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HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

BULLETIN

OP THE

MUSEUM OF COMPARATIVE ZOOLOGY

AT

HARVARD COLLEGE, IN CAMBRIDGE

VOL. 115

CAMBRIDGE, MASS., U.S.A.
1956-1957

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

.0
0 I

CONTENTS

PAGE

No. 1. — Notes on American Earthworms of the Family
LuMBBiciDAE. ITI-VII. By G. E. Gates. August,
1956 1

No. 2. — A Quantitative Study of the Equidae of the
Thomas Farm Miocene. By Robert S. Bader. Au-
gust, 1956 . . . ^ 47

No. 3. — Aneuretiis simoni Emery, a Major Link in Ant
Evolution. By E. 0. Wilson, T. Eisner, G. C.
Wheeler and J. Wheeler. (3 plates.) August, 1956 79

No. 4. — The Types of Naiades (Mollusca: Unionidae)
IN the Museum of Comparative Zoology. By
Richard I. Johnson. September, 1956 ... 99

No. 5. — Pseudemys scripta callirostris from Vene-
zuela w^iTH A General Survey of the scripta
Series. By Ernest Williams. (3 plates.) Septem-
ber, 1956 143

No. 6. — Revision of the African Tortoises and Tltitles
of the suborder Cryptodira. By Arthur Lov-
eridge and Ernest E. Williams. (18 plates.) Feb-
ruary, 1957 161

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. 115, No. 1

NOTES ON AMERICAN EARTHWORMS OF THE
FAMILY LUMBRICIDAE. IIl-VII.

By G. E. Gates

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

August, 1956

Publications Issued by or in Connection

WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

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Vol. 3, no. 35 is current.

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1948 — Published in connection with the Museum. Publication terminated
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Bulletin of the Museum of Compqrative Zoology

AT HARVARD COLLEGE
Vol. 115, No. 1

NOTES ON AMERICAN EARTHWORMS OF THE
FAMILY LUMBRICIDAE. III-VII.

By G. E. Gates

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

August, 1956

No. 1 — Notes on American earthworms of the family
Lumbricidae. III-VII.

By G. E. Gates
III

BiMASTOs TUMiDus (Eiseii, 1874)

Allolobophora tumida Eisen 1874, Ofv. Vet. Ak. Forh., 31, p. 45. (Type

locality, Mt. Lelnuion, New York-Massachusetts. Type, one half of

anterior end sagittally sectioned by Smith, in U. S. Nat. Mus.)
Allolohophora gic.srlcri Ude 1895, Zool. Aiiz., IS, p. 339, Zeit. wiss. Zool.,

61, p. 127. (Type locality, Savannah, Georgia. Types, numerous, in

Hanover Provincial Mus.?)
Allolohophora (Bimastm) tumida + -4. (-B.) gieseleri, Hclodrilus (B.) t.

+ E. (B.) g., Michaelsen, 1900, Abh. Nat. Ver. Hamburg, 16, p. 16,

Das Tierreich, 10, p. 502.
Helodrilu^ gieseleri hempeli Smith 1915, Bull. Illinois Sta. Lab. Nat. Hist.,

10, p. 551. (Type locality, Havana, Illinois. Tj-pes in U. S. Nat. Mus.)
Helodrilus (Bimastus) tumidus + S. (B.) gieseleri f. tijpica + H. (B.)

gieseleri var. hempeli, Smith, 1917, Proc. U. S. Nat. Mus., 52, pp. 170-

172.

?
Bimastus duels Stephenson 1933, Proc. Zool. Soc. London, 1932, p, 939.

(Type locality, Durham, North Carolina. Holotj^pe in Brit. Mus.)

Tamalpais, Marin County, California, 0-0-1 (macerated). California Ac.

Sci.
New Orleans, Louisiana. Under logs in moist river bottom forest, Dec.

31-Jan. 1, 1931-32, 0-10. J. M. Valentine per Dr. G. E. Pickford.
Big Thicket, Texas, May 1, 1938, 0-0-1. Ottys Sanders.
Athens, Texas. Six miles to the north, April 3, 1948, 0-0-3. Ottys Sanders.
Paris, Texas. Eleven miles to the south, March 29, 1952, 0-0 1. Ottys

Sanders. (Three specimens, without label, from Mr. Sanders probably

also from Texas.)
North Dublin, Virginia. Sawdust, Little Walker Creek, June 25, 1955,

0-2-17. Prof. Walter Harman.
South Rushton, Louisiana. In and near ash of sawdust pile, April 2, 1955,

0-0-3. Prof. Walter Harman.
Lincoln Parish, Louisiana, March 10, 1955, 0-0-2. Bobby Wise per Prof.

Walter Harman.

2 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

External characteristics. Length, 26-40 mm. Diameter, 2-3.5
mm. Segments, 81-122 (ef. Table 1). Anal segment of worms
not recently amputated usnally longer than penultimate one and
Avith setae in middle of anterior half but with little or no indica-
tion of a dorsal pore or an intersegmental furrow that would
mark off setigerous portion into a metamere. Body approximat-
ing to four-sided posteriorly, the dorsum slightly rounded and
wider than the ventrum, one setal couple at each corner. Pig-
mentation (unrecognizable in alcoholics except at site of clitel-
lum in a Louisiana worm) red, in dorsum, reaching ventrally to
or nearly to B through most of axis, also present in ventrum of
some or all of first fourteen segments, lacking at intersegmental
furrows, in wide transverse stripes in clitellar segments (recently
preserved Virginia worms). Prostomium epilobous, tongue open
(all). Setae begin on ii on which all usually are present, paired
but not as closely as in some lumbricids, d about at mL anteriorly
but more dorsal posteriorly where CD a t rifle <AB, BC<.AA,
DD<^iC hn\y AA. Nephropores, recognizable only on a few
segments, just above B on xv (all worms on which pores of that
segment were recognizable), on clitellum irregularly and some-
times asymmetrically alternated between levels just above B and
well above D (Virginia). Possible sites on more anterior seg-
ments have been noted on several specimens after some softening
and removal of the cuticle ; at or above D on iii-vi, slightly above
B on vii-ix, above D on xi. First dorsal pore on 5/6 (all).

Female and male pores equatorial and at usual levels on xiv
and XV respectively. Male pore tumescences in median half of
BC, confined to xv but sometimes dislocating 14/15 and 15/16
slightly, occasionally reaching on one or both sides slightlj^ into
xvi.

Clitellum saddle-shaped, reaching ventrally nearly to B
(Louisiana) or to A, located as follows; xx-xxix (1, Louisiana),
xx-xxx (2, Texas), (xx-xxi?) xxii-xxix (2, Texas), (xxi?) xxii-
xxix (1, Texas), posterior part/xxi-anterior part/xxx (1, Vir-
ginia), (xxi?) xxii-xxx (1, Louisiana), xxii-xxix (14, Virginia
and 4, Louisiana), xxii-xxix and onto xxx (2, Virginia), xxii-
xxix and possibly xxx (2, Texas), xxii-xxx (3, Virginia, Texas
and California). Tubercula pubertatis lacking.

Genital tumescences lacking or unreeognizable.

GATES : AMERICAN LUMBRICIDAE 3

Internal anatomy. No septa thickly muscular. Longitudinal
muscle band at niD with pigment, scattered flecks of red pigment
present on (in or under?) peritoneum of dorsum anteriorly (10,
Virginia).

Calciferous gland extends well into x, the posterior portion of
the gut in that segment often with an appearance as of a pair of
vertical sacs. Gland of about the same calibre through xi-xii,
nneonstricted at insertion of 11/12. Intestinal origin in xv
(24). Gizzard in xvii-xviii (24). Typhlosole begins gradually in
leaion of xxi and in places may reach floor of gut, slightly
widened veutrally in an anterior portion and there with a single
median groove. The typhlosole ends abruptly as shown in the
table.

Subneural trunk unrecognizable (22), very narrow but in part
red and on ventral face of cord (2). Extra-oesophageal trunks
empty, apparently passing into dorsal trunk posteriorly in xii
(1, Virginia). Hearts present in vii-xi, those of vii empty in
two worms (Virginia). Nephridial ducts, at least behind xv,
pass into parietes at B. Vesicle, U-shaped.

Table 1

Typhlosole termination and segment number

in Bimasios tumidus

Typhlosole

Atyphlo-

ends in

solate

Number of

segment

segments

segments

Remarks

81

19

100

Posterior amputee

83

17

91

*

Posterior amputee ?

85

15

100

Posterior amputee ?

95

20

115

97

16

113

« .1

Fcisteiiur aiiiputees?

97

20

117

Louisiana

98

19

117

Posterior amputee ?

99

12

111

»

Posterior amputee ?

101

13

114

*

Posterior amputee ?

103

15

118

Louisiana

106

16

122

Louisiana

107

15

122

Louisiana

• Last segment with setae as indicated on page 1.
AU worms from Virs'inia cxceiit as otherwise indicated.

Segment numher of other wnims : St (posterior amputee). 107 (posterior
amputee), ins, ii:;. 120.

4 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Seminal vesicles rather small, two pairs, in xi-xii (24). Male
funnels crenellate (24). Male deferent ducts slender, shortly
looped just behind funnel septa, the loops crowded into a small,
leaf-shaped disc. The ducts of a side come into contact in xii
where they apparently unite. They disappear from sight at
anterior margin of atrial gland or are recognizable further pos-
teriorly on surface of the gland (24). The ducts must have
grown posteriorly to or nearly to eq/xv, in some of these worms,
before the developing atrial gland had penetrated through the
longitudinal musculature. Spermathecae lacking (24). Ovisacs
of about the same size as the ovaries and slightly lobed (24).

Atrial gland crossed in xv by an equatorial cleft, a second cleft
possibly at insertion of 15/16 as the gland seems to extend
slightly into xvi. No glands on parietes in BC of clitellar seg-
ments nor in association with setal follicles. No setal follicles
especially protuberant into coelom (no genital setae?) in four-
teen Virginia and Louisiana worms.

Abnormality. Spiral metamerism involving v-vi only (1,
Virginia). Metameric abnormality in preanal region (1, Vir-
ginia).

Life history. Slight iridescence was recognizable on each male
funnel in ten Virginia worms collected on the 25th and 27th of
June, as well as in four from Louisiana (March 10 and April 2).
Male deferent ducts also appeared to be slightly iridescent. No
spermatophores. Seminal vesicles of an aclitellate Virginia worm
were of the same small size as in clitellate specimens and had
small brown bodies near dorsal ends, but none of the usual ex-
ternal indications of postreproductive regression was recognized.
The clitellum of some Virginia and five Louisiana worms may
have been at maximal tumescence. Sparse sperm production ap-
parently is indicated by these worms, in which case uniparental
reproduction perhaps should be anticipated, possibly' partheno-
genesis.

Parasites. Cysts, in coelomic cavities of anterior half of body
of a Texas worm, are attached to parietes and septa by long
filaments.

Remarks. Condition of the author's worms (most of them
alcoholic) is fair to very poor; the softened body wall of the
California worm was alreadv broken through before removal

GATES : AMERICAN LU:mbRICIDAE 5

from the tube in which it arrived. Much better material was,
however, loaned by Prof. Ilarnian, a favor which is much ap-
preciated.

Spermathecal vestijres were not recognizable in the parietes
after removal of lon<i'itudinal musculature but broad transverse
bands of pigmentation became quite obvious.

Septa 6/7 or 7/8 to 11/12 or 14/15 are somewhat strengthened
but none is thickly muscular. The thickness seems to vary with
the contraction or relaxation of the worm at preservation as
well as with degree of maceration. Differences detectable in dis-
section, insofar as present material is concerned, are slight and
do not appear to be of any taxonomic value.

Function and phylogeny of the gland present on the parietes
over site of male pore and its tumescences in many lumbricids
are unknown. The characterization "atrial" is retained, for
want of a bettor term, but without any connotations as to func-
tion or evolutionary history.

The clitellum of one Texas worm was about as in tiimidus, of
two as in gieseleri, of another as in hempeJi. The dift'erences ap-
pear to be no greater than are now to be expected in normal intra-
specific variation.

B. duels, known only from the original description of the
type, was thought to be distinguished, presumably from tumidus
and gieseleri, by a longer clitellum of 11^ segments. The last
hearts of duds are in x, according to Stephenson, but hearts of
xi when empty may not be readily recognizable in ordinary
museum material. Number of chambers in the calciferous gland,
which Smith (1917) believed to be of taxonomic importance
(though he did not always mention them) is unknown.

The clitellum covers 8-11 segments in tumidus and another
half segment may be within the range of normal intraspecific
variation in Bimastos. Location of anterior boundary of the
clitellum, at or in front of 21/22, also characterizes tiimidus but
no other species of Bimastos. Coverage of xxxi-xxxii by the cli-
tellum now seems to be the only reason that can be cited in sup-
port of retention of duels.

B. longicinctus Smith and Gittens 1915 has a clitellum, in
Illinois, according to its authors, of 10-11 segments, on xxiii-xxxii
or xxxiii. Location in worms subsequently recorded by others
from Ohio and Arkansas without data as to variation presumably

6 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

was the same, as the organ is all that enables routine identifica-
tion of the species. Number of segments covered is about the
same as in tumidus, but the anterior margin is at 22/23 rather
than at 21/22 or anteriorly. The only other method available
for separating these two forms involves counting chambers or
partitions in the calciferous gland. There are about 60 of these
in longicinctus, about 40 in tumidus. The difference, according to
Smith (1917, p. 175), is sufficient to warrant specific distinction.
Although little is known as yet about variation in such a charac-
ter, requiring microtome sections, the number of partitions in
the calciferous gland of Eisenia rosea (Savigny, 1826) is 38-54.

IV

BiMASTOS PARVUS (Eisen, 1874)

Allolobophora parva Eisen 1874, Ofv. Vet. Ak. Forh., 31, p. 36. (Type

locality, Mt. Lebanon, New York-Massachusetts. Types in TJ. S. Nat.

Mus.)
Allolobopliora heddardi Michaelsen 1894, Zool. Jahrb. Syst., 8, p. 182.

(Type locality, Orlando, Florida. Type in Hamburg Mus.?)
Allolobophora (Bimastus) parva + A. (B.) beddardi, Hclodrilus (B.) p.

-f H. (B.) h., Michaelsen, 1900, Abb. Nat. Ver. Hamburg, 16, pp. 14-13,

Das Tierreieh, 10, p. 502.
Eelodrilus {Bimastus) parvm -f H. (B.) beddardi, Smith, 1917, Proe.

U. S. Nat. Mus., 52, p. 173.
Bimastus parvus -f B. beddardi, KobayasM, 1940, Sci. Eep. Tohoku Univ.,

15, pp. 297, 298.
Bimastus beddardi, Cemosvitov and Evans, 1947, Linnean Soc. London,

Synopses British Fauna, 6, p. 18.

Mexico. In soil around roots of plants from Guadalajara, Jalisco, in
baggage arriving at Nogales, May 10, 1949, 0-0-1.

Fish bait in soil from Juarez, Chihuahua, in passenger's baggage
arriving at El Paso, May 21, 1949, 0-3-2.

In soil with two plants in passenger's baggage arriving at Santa
Fe bridge, August 13, 1953, 0-0-1.

In soil with plants from Torreon, Coahuila, in passenger's baggage
arriving at Santa Fe bridge, August 13, 1953, 0-0-1.
Italy. In soil witli strawberry and rosemary plants in baggage on SS

Atlantic arriving at New York, March 22, 1950, 0-0-1.
.lapan. In 25 ll)s. soil with plants in baggage on SS Cavnlier ainivin.? at
San Pedro, April 23, 1954, 0 0-1.

GATES: AMERICAN LUMBRICIDAE 7

England. In soil with plants in baggage on SS Britannic arriving at

New York, April 17, 1949, 0-1-1.
Wales. In soil with plants in baggage on SS Britannic arriving at New
York, April 24, 1949, 0-0-3.

(Worms listed above were intercepted by officers of the Division of
FoT-oign Plant Quarantines, U. S. Department of Agriculture and
were available through courtesy of Dr. C. W. F. Muesebeck and Dr. P.
W. Oman.)
Texas (probably), 0-0-3. Ottys Sanders. (In an unlabelled tube along
with B. tumidus.)

External eharacterisfics. Length, 25-42 mm. Diameter, at
clitellum, 2-3 mm. Segments, 101 (1), 102 (1), 104 (1), 109 (2).
Pigmentation, whenever recognizal)le (formalin preservation),
red. Body rather four-sided posteriorly. Prostomium epiloboiis,
tongue open. Nephropores (-when recognizable) just above B
or above D. First dorsal pore at 5/6.

Female and male pores e(iuatorial. Male pore tumescences
confined to xv or extending well into xiv and xvi (10).

Clitellum saddle-shaped, on (xxiii?) xxiv-xxx (1), (xxiii?)
.\xiv-xxxi (2), (xxiii?) xxiv-xxxii (2), xxiv-xxx (2), xxiv-xxxii
(1), xxiv/2-xxxii/2 (3), xxv-xxxi (9). Tubercula pubertatis
lacking or possibly represented by quite indistinct areas on xxvi-
.Kxx (several).

Internal afiatomy. Calciferous gland, intestinal origin, gizzard
and typhlosole as in tuniiclus. The typhlosole has no ventral
groove anteriorly and, in a worm of 109 segments, ends in xcvi.
Hearts were recognizable only in vii-xi (1) or viii-xi.

Seminal vesicles quite small. Atrial glands markedly pro-
tuberant into coelom. No setal follicles especially protuberant
from parietes (genital setae lacking?).

Life Jiistory. Spermatozoal iridescence was not recognized on
male funnels and spermatophores were lacking externally
though the clitellum in some of the worms appeared to be at
maximal tumescence. Parthenogenesis and male sterility are sus-
]:)ected.

Abnormality and variation. Helicometameres were noted be-
hind clitellar region in two worms. Seminal vesicles were lack-
ing in ix but were present in x of two worms.

Remarks. Many worms lacked the posterior end and most
were poorly preserved. If pigment once was present in alcoholic

8 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

material it had become unrecognizable. Anterior and posterior
margins of clitellum, as in tumidus, occasionally may not be
distinct because of quite gradual decrease in height of epidermis.
Whether male deferent ducts are looped on posterior faces of
funnel septa Avas not determinable.

If the usual assumption as to American origin of B. parvus
is correct, this species is the only American lumbricid that has
become "peregrine". It had been recorded, usually without
data as to variation, from at least sixteen states, as well as from
Mexico, Guatemala, Brazil, Hawaii, Japan, Manchuria, China.
Tibet, India, Burma, the Malay Peninsula, Java, Germany and
the British Isles.

Two species recognized by Smith (1917), B. parvus and hed-
dardi, were distinguished from each other almost only by dif-
ferences in clitellar location and in intersetal spacings- that now
appear to be within limits of normal intraspecific variation.

B. parvus is distinguishable from tumidus, also with about
40 partitions or chambers in the calciferous gland, only by
clitellar characteristics. The organ tends to be shorter, usually
of 7-8 segments, though 9 have been found and 10 may be
possible. Situation of anterior margin at 23/24 or 24/25, even at
22/23 when anterior margin is less definite, provides a further
distinction but not, in the latter case, from longicinctus. To that
species one such specimen, with a clitellum also covering xxxi-
xxxii, would have been referred but for its foreign origin.

So much emphasis on the clitellum is required because of more
or less recent elimination in Bimastos of spermathecae, tubercula
pubertatis, perhaps also genital setae and certain glands. Species
usually differ most obviously from one another, in the Lumbrici-
dae as well as other earthworm families, in their reproductive
organs on which the classical taxonomy largely has been based.
Satisfactory substitutes for the characters that are no longer
available still have to be found, not only in the Lumbricidae but
also in other genera where similar evolutionary changes have
been or still are under way (cf. Gates, 1954, p. 237).

GATES: AMERICAN LUMBBICIDAE 9

ADDENDUM

EisENiA ROSEA (Savigiiy, 1826)

External characteristics, of several American alcoholic speci-
mens, are such as to permit identification as B. parvus. Just as
in that species, seminal vesicles of ix-x and all spermathecae are
lacking. However, enlarged setal follicles are protuberant, in
each worm, into coelomic cavities of one or more segments ;
a-h or c-f//x-xiii, a-b/xxx, xxxii. Such follicles, with genital setae,
are to be expected in those very segments in the exotic rosea
where their presence usually is indicated by genital tumescences,
but apparently not in any American species of Bimastos.

The calciferous glands of rosea were described by Smith
(1924) immediately after those of parvus. The only difference
mentioned is in width of lamellae and of chambers.

Tubercula pubertatis have been lost during evolution of an
A morph ( ''Bimastos stage") in species of several lumbricid
genera, and their absence in these athecal individuals of rosea
is not, then, surprising.

Follicles that are to produce genital setae become enlarged,
in this as well as other exotic species, during a juvenile stage
before development of tubercula and clitellum which appear
ontogenetically in that order. Tumescences develop around aper-
tures of GS follicles as the clitellum is differentiated. Suppres-
sion of GS tumescences in this American parthenogenetic strain
may be an early stage in an evolutionary development that will
result in elimination of the associated GS follicles and their
modified setae. Just such an early stage apparently has been
reached by certain species of Bimastos. In others, tumidus and
parvus, such evolutionary development may have been com-
pleted. Genital setae, presumably functioning only during
copulation, are of no use to animals that have become partheno-
genetic.

Bimastos palustris Moore 1895

Xew Jersey, 0-0-7. Harold Davies (1954).
External characteristics. Length, 18-38 mm. Diameter, 1 mm.
or slightly more at posterior end, ca. 2 mm. anteriorly, 2+ mm.
in clitellum, 3 mm. in xv-xvi. Segments, to 101 (cf. Table 2).

10 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Pigmentation red, in circular musculature of dorsum anteriorly,
lacking close to intersegmental furrows but present in ventrum
of first five or six segments, recognizable only 'after stripping off
longitudinal musculature, a light pink coloration all that is vis-
ible externally. Body squarish posteriorly and with setae at
corners. Prostomial tongue broad, short, open (7). Setae begin
on ii on which all are present (7), AB=, < or > CD, but dif-
ference slight, BC < AA < DD < IC. Nephropores certainly
recognizable on'y behind xv, irregularly and asymmetrically
alternate ])etween levels just above B and dorsal to D. First
dorsal pore on 5/6 (6), W/7 (1), a weak spot at mD on 4/5
in one worm clearlj^ not a definite pore.

Female pores diagonal to transverse slits (or in such slits?),
posterolateral to h (7), at most midwaj^ between eq/xv and
15/16. Male pores deeply invaginate, the usual transverse cleft
apparently slightly postequatorial in some but probably really
equatorial on all. Male pore tumescences conspicuous, maximal
height at or close to 16/17, sloping anteriorly down to the cleft,
scarcely noticeable anterior to eq/xv.

Clitellum annular (7), epidermis not as thick in AA as dor-
sally but intersegmental furrows unrecognizable (6), anterior
and posterior margins distinct and as much so ventrally as in
the dorsum (7), on xxiii-xxvii (1), xxiii-xxviii (6). No tubercula
pubertatis (7).

Genital tumescences include a-h setae, on xiii (7) and xvi (7).

Internal anatomij. Septa l:)aek to 14/15 somewhat more opaque
than posteriorly, 8/9 apparently the thickest, 7/8-9/10 less so,
all weak — merely pushing gut over to one side or the other
breaks the septa at parietal insertions, even anteriorly. Pigment
apparently lacking in longitudinal muscle band at mD and in
]ieritoneum.

Calciferous gland extends into x (7), the gut in that segmen*:
with an appearance as of two vertical sacs posteriorly. Constric-
tions at 10/11-11/12 usually slight or unrecognizable but in one
specimen so deep externally that the gland appears to be monili-
form as in Eisenia Idnnbergi (Michaelsen, 1894) and F. carolin-
ensis Michaelsen 1910. Constriction at or near 12/13 external or
internal or both and then different on opposite sides. Intestinal
origin in xv (7). Gizzard in xvii-xviii (7). Typhloscle begins
rather gradually behind the gizzard and is a simpli- lamella,

GATES : AMERICAN LUMBRICIDAE

11

slijihtlj' thicker ventrally, often reaching fioor of gut, ending
as shown in Table 2.

Snbncural trunk mostly unrecognizable but occasionally indi-
cated, on ventral face of cord, in two worms by an indistinct red
streak visible only in a segment or two. Extra-oesophageals, pre-
sumai)ly empty, unrecognizable. Hearts in vii-xi (7), those of
\ii em])ty and small (4), all empty (1). A vertical red column,
resembling a blood vessel and doubtless of blood, recognizable in
lateral Avail of calciferous gland on each side in one or more or
all of x-xii. Nephridial ducts, at least beliind xv, pass into
j)arietes at B.

Tabi>k 2

Typldosole termination and segment number
in Bimasfos lyahtsiris aiul B. sp.

Typhlosole

Atyphlo-

Number

ends in

solate

of

segnient

segments

segments

Remarks

47

8

55

Posterior amputee

48

4

52

Posterior amputee

62

6

68

Posterior amputee

69

8

77

Posterior amputee

83

10

93

Posterior amputee?

84

11

95

Posterior amputee?

87

14

101

B.

sp.

101+ ?

?

101

Posterior portion broken oiT
and lacking.

130

17

147

Seminal vesicles equisized and fairly large, in xi-xii (7).
Male funnels large, crenellate, reaching well up at sides of gut.
Male deferent ducts straight, those of a side coming into contact
in xii, passing up to apex of atrium where primary male pore
doubtless is located. Spermathecae lacking, rudiments unrecog-
nizable in parietes after removal of longitudinal musculature
(7). Ovisacs present (7). lobed, smaller than the ovaries. 0\\-

12 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

ducts seem to be unusually thick relative to size of worm and
obviously pass into parietes behind eq/xiv.

Atrium thumb-shaped, erect in xv, reaching well up toward
level of dorsal face of gut (7), basal portion distended pos-
teriorly and continued into xvi as a transversely placed reni-
form mass that bulges back 16/17 at parietes. No corresponding
extension on parietes anteriorly.

Follicles of ventral setae of xiii and xvi slightly thickened but
thin- walled, elongated, apices reaching into coelomic cavities just
above associated glands. Acinous glandular masses on parietes
median and posterior to each such pair of follicles.

Ah7iorniality. Male pore of right side and atrium in xvi, right
male tumescence in xvi-xvii. Basal gland of atrium in xvii. Geni-
tal tumescences of right side in xiii and xvii. Clitellum not as
thick ventrally in BB as in other worms and intersegmental
furrows faintly indicated across mV.

Habitats. Apparently restricted to banks that are kept moist
by running water (cf. Moore, 1895 and Davies, 1954).

Natural history. Slight iridescence is recognizable on male
funnels of two specimens with spermatophores, but not on funnels
of other worms.

Spermatophores were found by Moore from latter part of
February to December. Cocoons may then be deposited in any
season of the year that temperature permits. Although not now
anticipated in the summer, some sort of a diapause may be
expected in January or whenever the weather is too cold for
normal activity.

Biparental reproduction can be assumed, at present, as Moore
mentioned release from spermatophores of sperm presumably
normal.

Distrihtition. Pennsylvania, within thirty miles of Phila-
delphia. Northern New Jersey. Raleigh, North Carolina. The
latter record was accompanied by no information as to habitat,
number of specimens, structure, etc.

Remarks. Greater opacity of epidermis than usual in the
dorsum anteriorly, in vivo as well as after preservation, pre-
sumably is responsible for a previous assumption as to absence
of pigment (cf. Smith, 1917, p. 169).

Spermatophores are 2-2-|- mm. long, attached to xiv in DD.

Male pore tumescences in the types, so far as can be judged

GATES: AMERICAN LUMBRICIDAE 13

from the figure provided by Moore, seem to have been as marked
anterior to the cleft as behind it.

The calc'iferous gland appears to be constricted at or near
insertion of 12/13 as in various other American lumbricids both
exotic and endemic. The constriction may be external on one
side and internal on the other or internal all around when gut
is straiglit. These dift'erences probably are of little significance,
perhaps a matter of differential contraction at time of preserva-
tion. However, the constriction is usual and in dissections ap-
pears to provide a posterior boundary of the gland though calcif-
erous lanudlae, according to Smith (1924) frequently extend
into xiii.

Hearts tiid not pass in any of these specimens into gut wall
((•r. Moore, 1895) which confirms Smith's finding (1928, p. 354).
The vei'tical cords of blood mentioned above may have been
mistaken for hearts. Inability to trace the extra-oesophageal
trunks in the present specimens is unfortunate because of a
possibility that they may provide characters of taxonomic im-
portance in a group where such are much needed. However, it
can be suggested, after examination of the vessels in many speci-
mens of various other species that the extra-oesophageal trunks
do not join the ventral trunk in x (Moore, 1895) or the dorsal
trunk in x (Smith, 1928). Distention by blood of an anterior
portion of the extra-oesophageal and of a fairly large branch
that occasionally passes (in other species) to the dorsal trunk in
ix and/or x, with the portion of the extra-oesophageal behind
the branch empty and collapsed, may have been the condition
seen by Smith. A branch from the subneural arising in any of
segments xiv-x (again in other species) may pass to the extra-
oesophageal. AYhen that branch is distended with blood the
trunk at first appears to turn laterally and join the extra-oeso-
phageal but does again become recognizable anteriorly wherever
blood is present. The diificulties posed by temporarily empty
vessels in earthworms rarely have been appreciated even by
those who made special studies of the circulatory systems.

Microscopic examination of the setae in the enlarged ventral
follicles of xiii and xvi has not been possible. Some modification
certainly can be expected even though not recognized by Moore
nor mentioned by Smith.

14 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

B. palustris is distinguished from the previously erected
species, tumid'us and parvus, by the shorter clitellum of only
5-6 segments as well as its obvious annularity, posterior disloca-
tion of female pores, deep invagination into coelom of male
pores at apices of large atria, the enlarged setal follicles in xiii
and xvi, the epidermal tumescences and the coelomic glands
("postsetal", Moore) associated with those follicles. The folli-
cles, genital setae therein, glands and tumescences have not been
used in the classical taxonomy. Little is known, however, about
the variation of any of these characters in this species or of
the newer ones just mentioned in any lumbricid. Another dis-
tinction from tumidus and parvus may be in the method of
reproduction.

Closest to palustris of the Bimastos species now appears to be
heimburgeri Smith 1928, erected on a single specimen from
Illinois. Four individuals subsequently recorded from North
Carolina, in absence of any description, must be assumed to be
like the type. The species is distinguished from palustris by the
clitellum, of 8-|- segments on xxv-xxxiii(part) and saddle-
shaped, absence of genital setae and presumably also of the aci-
nous glands. The type was not "sexual" though clitellate, and
Smith suggested (1924, p. 856) that genital setae "may be
present at the time of sexual activity." This now seems unlikely.
Enlargement and protuberance into coelom of GS follicles is
obvious in lumbricids studied by the author before any indica-
tion of clitellar development is recognizable, and in thecal species
even before appearance of the tubereula pubertatis. If Smith's
statement re gonads of diminished size referred to the testes,
and if clitellar tumescence was maximal, the type may be par-
thenogenetic. Otherwise iridescence should have been recogniz-
able on the male funnels of the unsectioned half and in the sec-
tions of the other half sperm on the funnels should have been
obvious.

Bimastos sp.

Moonshine Dell, Giles County, Virginia, June 20, 1955, 0-0 2. Prof. Waltpr
Plarman.
External characteristics. Size, 71: x 4.5 (at xv) mm. Seg-
ments, 147. Pigmentation red, in dorsum but sparse behind
clitellum until near hind end, in ventrum of first seven seg-

GATES: AMERICAN LUMBRICIDAE 15

merits. Body approximating to transversely rectangular in cross
section posteriorly, M-itli setae at the corners. Setae begin on
ii on which all are present, rather closely paired, AB ca. = CD,
BC ca. = AA, DD < iC posteriorly. Prostomium epilobous,
tongue open (2 specimens). First dorsal pore on 5/6 (2).
Nephropores unrecognizable.

Clitellum annular, only slightly tumescent, intersegmental
furrows recognizable all around, on xxiii-xxxi (1), still quite
indistinct on the other but apparently indicated by a whitening
of the dorsum on xxv-xxx. Tubercula pubertatis lacking.

Female pores definitely equatorial on xiv, of about the same
size and shape as adjacent aperture of a GS follicle. Male pores
each in a deep equatorial cleft or slit at bottom of a depression
sloping down from 15/16 and 14/15, tumescences markedly pro-
tuberant, extendino- across all or nearly all of xiv and xvi,
equally developed anterior and posterior to eq/xv.

Genital tumescences unrecognizable (perhaps not yet de-
veloped) but a-h setae of xii-xviii retracted into parietes, follicle
apertures enlarged, slit- or comma-shaped, longitudinally placed.

hiternal anatomy. Septa 8/9-14/15 strengthened, 12/13 the
thickest. Pigment sparse in longitudinal muscle band at mD.
Peritoneal flecks very few.

The calciferous gland extends well into x and there is widened
so that there is an appearance as of two vertical sacs. A slight
constriction at insertion of 10/11, none at 11/12, internal at
12/13. Intestinal origin in xv (2). Gizzard in xvii-xviii (2).
Typhlosole begins in xxii, nearly 2 mm. high but rather thin
anteriorh- where it reaches floor of gut, and has a deep longi-
tudinal groove on ventral face, ending abruptly (cf. Table 2).

Subneural trunk recognizable only for a distance of one seg-
ment here and there where a little blood is present. Extra-oeso-
phageals empty but traceable (1) into dorsal trunk posteriorly
in xii. Hearts present in vii-xi, empty in vii (1) or with a little
blood but small. Nephridial ducts pass into parietes at B.

Seminal vesicles rather small, tough, in xi-xii. Male funnels
crenellate. Male deferent dacts without looping on funnel
septa. Ovaries and ovisacs as usual, spermathecae absent (2).

Atrium hemispheroiclal, the lumen reaching above level of
parietes, with glandular extensions into xvi and xiv. Follicles
of ventral setae of xii-xviii enlarged and conspicuously pro-

16 BI^LLETIN : MUSEUM OP COJNTPARATn'E ZOOLOGY

tuberant above parietes. Acinous glands apparently are associ-
ated with the follicles.

Remarks. Thickened septa, nephridia, GS follicles and acinous
glands are closely crowded and adherent to each other. Deter-
mination of relationships in such circumstances sometimes be-
comes possible after several months further treatment with
preservative.

Enlarged apertures of GS follicles presumably are common
external openings of the follicles and of the associated acinous
glands.

Atria are not as high as in palustris.

Spermatozoal iridescence on male funnels is slight. Such
iridescence has been noted in other lumbricid species during
postsexual regression, at onset of maturity, and when matura-
tion is sparse as in certain supposedly parthenogenetic forms.

These worms are to be compared with those of species having
the anterior margin of the clitellum at 22/23, palustris, longi-
cinctus and parvus (only occasionally). They cannot go in
palustris as at present understood though they do have GS
follicles in xiii and xvi and acinous glands associated therewith.
Presence of those follicles and glands presumably rules out
2)arvi(s in which they are lacking. Nothing is known as to GS
follicles and glands in longicinctiis in which the clitellum is sad-
dle-shaped though this latter difference may prove to be of no
importance.

BiMASTOs sp.

Whitetop Mountain, Viiginia, under bark of log at elevation of 4700 feet,
August 17, 1955, 0-0-1. Prof. Walter Harman.

External characteristics. Size, 16 x 2 mm. (through clitellum).
Segments, 61 (probable posterior amputee). Pigmentation red,
obvious externally. First dorsal pore at 5/6.

Female pores on xiv, further from B than usual, almost at
level of male pore cleft, only slightly if at all postequatorial.
Male pore clefts equatorial on xv. Male tumescences extend
well into xvi and xiv, 1-1/15 and 15/16 unrecognizable across
them, and include a-h of xv-xvi. Clitellum annular, on xxiv-
xxxii(xxxiii?), posterior margin indistinct. No tubercula.

Internal anatomy. As in previous species except as otherwise
mentioned.

GATES: AMERICAN LUMBRICIDAE 17

Typhlosole ends in liii but is much lower there than in the
preceding segment.

Male deferent ducts looped between funnel septa and parietes
but in a loosely zigzagged manner, not compacted into a ball or
disc. Oviducts pass into parietes anterior to equator of xiv and
are relatively thick. GS follicles, a-b of xv-xvi, markedly pro-
tuberant into coelomic cavities. No acinous glands associated
with those follicles.

No atria ; even atrial glands lacking.

Parasites. Five transparent cysts, each of about the same
size as the prostomium, are present in coelomic cavities of xi-xii.

Eemarks. No iridescence recognizable on male funnels.

Species with the anterior margin of the clitellum at 23/24 are
longicinctus and parvus. Presence of GS follicles in xv-xvi (that
are lacking in parvus) apparently obviates further consideration
of that species. Nothing is known as to presence or absence of
atrial glands in longicinctus.

VI.

Species of Eisenia

The distinctness and relationships of our endemic species of
Eisenia, which were characterized about as well as their Eu-
ropean congeners, have been misunderstood abroad. So little has
been known about variation in and distribution of the American
forms that the collections made by Dr. Pickford and Prof. Har-
man, in the hitherto uninvestigated mountains of North Caro-
lina, Tennessee and Virginia (Gates, 1955), though secured in
a probably unfavorable season, have joroved to be of considerable
importance. Material received since completion of the manu-
script of parts I-II of this series now has provided some of the
additional information that was w^anted for a discussion of
these relationships.

E. hortensis obviously is exotic in North America. A brief
description is included to provide hitherto unrecorded but now
necessary information. Notes on other exotics, E. foetida and
rosea, are provided for similar reasons.

The author's thanks are extended to Prof. Walter Harman,
Harold Davies, and Curator Fenner Chace, Jr., for the oppor-
tunity of studj-ing the various specimens they supplied.

18 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Genus ElSENIA Malm 1877

EiSENiA CAROLiNENSis Michaelsen 1910

Whitetop Mountain, Virginia, August 17, 1955. Under rocks at summit,
4-0-4. Under bark of log at elevation of 4700 feet, 0-0-1. Prof. W. A.
Harman.
Mountain Lake, A'irginia. Barn, Mountain Lake Hotel, June 20, 1955,
0-0-1. Moonshine Dell, June 20, 1955, 0-0-2. Castle Rock, July 8, 1955,
0-0-1. Prof. W. A. Harman. Mann's Bog, MLBS., August 4, 1955,
0-0-2. Doris Hatfield per Prof. W. A. Harman.

Segments, 133 (2 helicometameres in intestinal region), 136,
141, 144 (all juvenile), 141 (largest clitellate worm, 110 x 7 mm),
152. Pigmentation red, quite dense in juveniles and obvious ex-
ternally clear to hind end, also externally recognizable all the
way to the anal region in one clitellate worm but in others visi-
ble just in the preclitellar portion of the body and even there
sometimes indicated only Iw a faint pink. Pigment appears to be
sparse in dorsum of ix-xii, except near mD, of the clitellate worm
with most obvious red color. Clitellum whitish or light grayish
and, even though in formalin, about as in previous alcoholic
material.

Tubercula pubertatis as before, quite rudimentary but obvi-
ously tripartite on two late juveniles. Variation in location of
genital tumescences and clitellum is shown in the table.

The typhlosole ends in cv, but is smaller in last two or three
segments (worm of 141 segments), cxiv (152 segments).

Seminal vesicles of a juvenile are large, filling coelomic cavi-
ties except in ix, very soft and fragmenting on slight pressure
even after a month in formalin (preservation very good). Testes
discoidal. Iridescence lacking on male funnels.

Eemarks. The red coloration in clitellar epidermis previously
mentioned (Gates 1955, p. 8) must have been slowly developed
in the formalin. No trace of the coloration is as yet recognizable
on these recently preserved worms.

GATES: AMERICAN LUJIBRICIDAE

19

Table 3

Variation in external characteristic's of E. ('(iroiiju'usis Itoui

Virginia

Genital tumescences,

including ah,

additional to those

Serial

on xxv-xxx, ioentcfl

Clitellum

number

in segments

on segments

Remarks

1

11

17 18 - 31

Juvenile. TP rud.

O

11

17 18 31

24-32/eq

3

11

17 21 24 31

24-31

■±

11

18 21 31

24 32

Sph.

5

16 17

eq/2431

()

16 17 21 31

(231)24-31

7

16 17 21 31

24-31

8

16 17 21 31

24-31

9

17 21 31

Juvenile. TP rud.

10

17? 24 31

Juvenile.

11

17 21 31

24-31

12

17 21 31

25-31

13

17 18 20 21 31

2432/eq

14

17 24 31

Juvenile.

15

21 31

2531

TI' iwl. Tiil)erculM i)ubprtatis ruilimoiitary. (Rudiments still iiiir('Co;,aii7,al)le in

other two jnveniles. )
eq Equator.

Sph Small spermatophorc median in -1 on riuht side of xxvii.

Genital tumescences on juveniles are, of course, still riidimeutary Inu

are definite except at the one site indicated by a " ?".

Two small knobs mark sites of invisible follicle apertures on each of

the conspicuous tumescences of clitellate worms.

EiSENiA LONNBERGi (Michaelseii*; 1894)

Pigment, in or associated witii circular muscle layer, obviously
is not red and that layer, after stripping off the longitudinal
musculature, has a dirty or sooty appearance somewhat sug-
gestive of an India-ink suspension. Externally, pigment ap-
pears to be lacking in the dorsum of viii-xii except close to mD,
but after removal of longitudinal musculature is recognizable
though more sparse than elsewhere. Peritoneal flecks appear to
he black and thev mav be numerous and large.

20 BULLETIN : MUSEUM OF COMPARATIYE ZOOLOGY

Clitelliim greyish (formalin preservation), on xxiii-xxx (2),
xxiv-xxx (1). Genital tumescences, including o-h, on xii (1),
xxiii-xxx (2), xxiv-xxx (1).

Typhlosole ends in xci (worm of 108 segments and like the
other two a posterior amputee).

Hearts of x-xi empty and slender (3). From the subneural
a large branch passes up on anterior face of 13/14, on each side,
to dorsal trunk. Extra-oesophageals again empty, traceable only
in one worm and then just into xii.

Male deferent ducts may be quite obviously looped just behind
funnel septa but loops are loose, not bound by connective tissue
into a disc or ball. GS follicles of xii surrounded on parietes
by annular glands. Glandular tissue is median to A on parietes
in xxiii or xxiv to xxx but is lateral to B only in xxvi-xxix (and
there TP glands?). GS follicles of xxiii or xxiv to xxx (as well
as of xii, in 1 specimen) conspicuously protuberant into coeloni,
those of xxvi-xxviii somewhat more so.

Remarks. These New Jersey worms (Davies, 1954, p. 6) had
not been in formalin long enough to permit development in
clitellum of the red coloration previously mentioned (Gates 1955,
p. 2). The dark coloration previously mentioned {ihicl., p. 6),
that often "develops" during preservation when formalin is in
contact with cork, obviously is not involved here.

Gut lumen in xi-xii may be only a vertical slit as shown by
Smith (1924, figs. 25, 28 ) or the lumen may be much larger, even
wider than the gut wall. Differences presumably are due to
physiological state of glands at time of fixation and/or to varia-
tion in method of preservation. Even with greatest width of
lumen and maximum narrowness of the wall the ealciferous
gland is obviously moniliform. The gland always appears, in
horizontal sections, to end without reaching into segment x.

Relationships of glandular masses on parietes in clitellar
region still remain to be determined ; distinguishing setal glands,
possible peritoneal blisters elevated by accumulations of some
kind of corpuscles, and compacted masses of coelomic coagulmn,
from each other and from possible TP glands not permitted by
material that has been available. If TP glands are lacking,
lonnhergi will have one more character in common with caro-
lincnsis as well as E. foctida (Savigny, 1826). Presence of TP
glands will be shared with horfensis.

GATES: AMERICAN LUMBRICIDAE 21

EiSENiA FOETiDA (Savigny, 1826)

External cl)a)'actcnstics. Spermatheeal ])ores appear to be
slightly more lateral to inD than in American species. Male pores
each at lateral end of a sliuht cleft and at or slightly lateral to
niB(\ Tumescences slight, restricted to xv and to median half
of BC.

Clitellum of 6-8 segments, usually beginning at 24/25 or 25/26
and ending at 31/32 or 32/33. Tubercula puliertatis extending
across three or four segments, on xxviii-xxx (most frequently),
xxviii-xxxi, or bet^veen equators of those segments.

Genital tumescences often quite distinct, in the elitellar region
as obvious as in the macedonian form of E. rosea or indicated
only by deep retraction of genital setae, each tumescence around
a setal couple. Any pair of ix-xii or each couple of one of those
segments may be involved but there is considerable variation
as well as asymmetry. \''entral couples of xxvi-xxxii usually
are included in timiescences, those of xxv, xxiv and xxxiii less
often, in one of xvii-xxiii much more rarely.

Infernal anatomy. No septa thickly muscular. Calciferous
gland begins behind insertion of 10/11, much wider in xii, with
a slight external constriction at 11/12 and an internal constric-
tion at or just behind 12/13 on one or both sides. No trace of
sacs in x. ({izzard variable, layer with brilliant muscular sheen
in horizontal section ends at or slightly behind insertion of 17/18
or is continued though with decreasing thickness to or nearly
to 18/19. (Jut narrowed in xix and valvular at 19/20.

Dorsal blood vessel empty and unrecognizable anterior to 5/6.
Subneural trunk adherent to cord. Extra-oesophageals turn up
in ix, pass through 9/10 and into dorsal trunk on posterior face
of 9/10, a posterior continuation of the trunks behind the up-
ward bend recognizable in none of the specimens. Hearts in vii-
xi onl.y (20), no branches from dorsal trunk in vi (20), hearts of
x-xi usually smaller than those of ix. Nephridia appear to be
vesiculate, shape of bladder doubtful (J-shaped?). The duct,
very short and slender, passes into parietes at B. Necks of fun-
nels sometimes are long enough to reach into contact with septum
next in front (contracted material). Brain and circumpharyn-
geal eorami.ssures left in iii by a transverse section exactly along
3/4.

22 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Holandric. Seminal vesicles, four pairs, in ix-xii (20), one
vesicle of xii occasionally extending into xiii and markedly con-
stricted by 12/13. Male deferent ducts rather thick entally,
just behind funnel septa irregularly and variously looped, con-
nective tissue binding loops together delicate and easily rup-
tured, the looped portion ("epididymis") sometimes reaching
a length of 2 mm. Ovaries, ovisacs and oviducts as usual, except
that an extra ovary (with large ova distally) is present though
not associated with a female funnel or ovisac in two of ten Maine
worms. Spermatheeae free, in ix-x (20), a short duct portion
often recognizable in coelom.

Atrial and TP glands lacking (20). GS follicles (those in
tumescences) protuberant into coelom but often bent laterally
(a-h) or mesialh' (c-d) against parietes. The "postsetal glands"
unrecognizable (field preservation) but some slight parietal
blistering around GS follicles usually is recognizable.

A half-moou-shaped slight elevation with base at D extends,
on parietes in viii or ix-xii, well toward mD and coincides with
an obvious external white area in which the epidermis is
thickened. The parietal elevation appears to be little more than
an accumulation of flocculent matter (corpuscles?) underneath
elevated peritoneum. Protuberance, like the epidermal thicken-
ing, is most marked when pigment associated with circular
muscle layer has been completely lysed, less marked when lysis
is as yet only partial.

Reproduction. Male funnels, in a large number of clitellate
worms, always have a brilliant spermatozoal iridescence. A
similar iridescence usually is recognizable in spermathecal am-
pullae. Spermatophores often are present externally. Reproduc-
tion presumably is sexual and probably almost always biparental.

Remarks. The above characterization, based on material from
ten states, supplements a previous description (Gates, 1943, p.
95) published after side by side comparison of "wild" worms
and those being sold by earthworm culturists as "hybrids."

Nephropores, as before, could not be identified with certainty
though possible sites on some segments may be slightly more
lateral to B than usual.

GATES: AMERICAN LUMBRICIDAE 23

EiSENiA HORTENSis (Michaelseii, 1890)

Uibaiia, Illinois, April 25, 1923, 00-3. J. L. Hyatt per F. Smith (U. S. Nat.
Mus. No. 26205).

External characteristics. Se<i-nieiits, 85-103. Pi<iment unrecog-
nizable externally (alcoholic preservation). Prostomium epilo-
bous (3), tongue open. Setae begin on ii, on which all are pres-
ent, paired, AB ca.=CD apparently < BC, d rank about at niL
anteriorly, somewhat more dorsal posteriorly where c rank seems
to be at niL. Xephropores small, first recognizable on iii, located
on left side of one specimen as follows : iii-iv/Z>, v/well above D,
vi/nearer mD than to D, vii-ix/D, x-xi/slightly above B, xii-
xiii/D, xiv-xvi/above B, xvii-xviii/D, xix-xx/above B, xxi-
xxii D, xxiii/above B, xxiv/D. Both pores of xiv are just above
B (1) or at Z> (2), both pores of xv-xvi just above B (3). First
dorsal pore at 5/6 (3).

Spermathecal pores minute, each on a tiny tumescence close
to mD, on 9/10-10/11. Female pores minute, just above B,
equatorial on xiv. Male pores minute, about the same size as
nephropores and smaller than the female pores, each laterally
in a slight equatorial cleft (almost unrecognizable in one worm
with greatest protuberance of tumescences), on xv in median
half of BC. Tumescences markedly conspicuous, confined to
median half of BC, longitudinally elliptical, bulging 14/15 an-
teriorly and interrupting 15/16 to reach well into xvi.

Clitellum saddle-shaped, unrecognizable below B, on xxviii-
xxxii (2), xxviii-xxxiii (1). Tubercula pubertatis on xxx-xxxi
at least, apparently reaching onto xxix and to xxxii/eq (2),
longitudinal bands of translucence without any indication of
elevation (2), each band on median face of a markedly pro-
tuberant and rather lamelliform ridge highest on xxx-xxxi and
gradually disappearing on xxix and xxxii (1).

Genital tumescences indistinct, setae retracted out of sight,
a-b/x (1), a-h and c-d/x (1), c-d/xi (1). Unrecognizable pos-
teriorly.

Internal anatomy. Septa, none thickly muscular. Pigmenta-
tion still unrecognizable after stripping off: longitudinal muscula-
ture but circular layer not white as is the other.

Gut wide in x, with wall thin to 10/11, no trace of sacs. Calci-
ferous gland in xi-xiii, not quite reacliing 10/11, no special

24 BFTLLETIX : MUSEUM OF COMPARATIVE ZOOLOGY

constrictions recognized. Intestinal origin in xv (3). Gizzard in
xvii (3), the layer with brilliant muscular sheen obviously ex-
tending in horizontal sections only slightly behind 17/18. Gut
narrowed and valvular at 19/20 (3). Typhlosole begins in
region of xxi-xxiii and is simply lamelliform, ending in Ixxvi
(worm with 85 segments), Ixxxix (100 segments), xc (103 seg-
ments).

Dorsal blood vessel recognizable into v, with two pairs of
vessels to gut in each segment from xii posteriorly, one pair
anteriorly in each of xi-x, none in ix. Extra-oesophageals and
subneural trunk unrecognizable. Hearts in vii-xi (3). Nephridia
vesiculate, the vesicle usually collapsed but when distended
elongately ellipsoidal and extending on the parietes from A or
B to beyond D or almost halfway to mD. Ducts not certainly
recognizable, prol;ably very short and passing into parietes at
B. Brain and circumpharyngeal commissures left in iii, once
well anteriorly, by a transverse section exactly along 3/4.
Pharynx roof in iv with a median groove that shortly divides so
that lumen is Y-shaped in transverse section.

Testes in x-xi (3), male funnels medium-sized and crenellate.
Male deferent ducts straight, Avithout sinuosities or looping (no
epididymis), uniting posteriorly in xii, passing posteriorly
about at mBC and into anterior margin of atrial gland laterally.
Seminal vesicles in ix, xi and xii, those in ix the smallest. Ovaries,
ovisacs, oviducts as usual. Spermathecae in ix-x (3), ducts prac-
tically confined to parietes.

Atrial glands low, finely acinous, tough, longer than wide,
in median half of BC, apparently reaching into xvi. TP glands
on each side, a longitudinal tough mass with an acinous appear-
ance like that of the atrial gland, in xxx-xxxi, possibly extending
slightly into xxix and xxxii, in the worm with ridges almost
concealed from view in a deep groove.

GS follicles not especially thickened but protuberant above
general level of parietes, anteriorly as indicated by the tu-
mescences, posteriorly b of xxx-xxxi (3), xxix (2), xxxii (1). The
a follicles of xxix-xxxii appear to be slightly enlarged and
slightly protuberant into coelom.

Reprochfction. Two spermatophores present in AB in presetal
half of xxviii (1) have an opaque central portion with slight
iridescence. Spermatozoal iridescence on male funnels and in

GATES: AMERICAN lATMBRTCTOAE 25

distended spermathecae brilliant. Biparental reproduction is
anticipated.

Remarks. These worms had been preserved in near-maximal
relaxation and jruts are only partially filbnl. They are too soft,
though not macerated, for convenient study of external charac-
teristics. Inability to recognize genital tumescences posteriorly
probably results in part from the method of preservation. Ne-
phropores, however, in several regions are visible under high
power of dissecting binocular as definite openings. Blood in
distended hearts is perfectly white and this bleaching is of no
assistance while looking for the subneural trunk oi- in learning
relationships of the extra-oesophageals. Blister-like parietal
elevations, around a and b setae of xvi (with slight indications
of lobing), and extending uninterruptedly through clitellar
segments exactly in AB, may be artifacts ( ?).

The clitellum of the only other specimen seen (Gates, 1948,
p. 96) was on xxvii-xxxiii. More anterior extension, across xxv
or even xxiv-xxv, apparently indicated by Smith (1917, p. 166),
may have been lacking in his own material.

Asymmetry in iiitrasegmental location of nephropores was
recognized or suspected only very rarely and may be less fre-
(juent than in some si)ecies though condition of material did not
permit enough observations for a more definite characterization.

Anterior termination of calciferous gland behind insertion
of 10/11 in hortensis, previously recorded by Smith (1924),
certainly appears to be more than sufficient justification for
specific distinction from worms in which the gland extends well
into X.

The taxon Jiortensis. originally a forma of Allolohophora .suh-
rubicinida, eventually became a variety of vcncia in Eisenia.
then a species in Eisenia (Omodeo, 1952, p. 188), and lastly a
species in Dendrohaena (Omodeo, 1955, p. 6). The transfer to
Dendrobaena, along with roieta, seems unlikely to assist in
resolving any of the confusion so prevalent in lumbricid tax-
onomy.

Variation greater than usual in a species seems to have been
imputed to hortemsis, which may be in need of revision and more
accurate definition. The species Avill have to be considered as
characterized above, in the discussion (p. 29), since European
material has been unavailable and examination of the types has
been impossible.

26 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

EiSENiA ROSEA (Savignv, 1826)

This si)eeies is polymorphic, various formae that have been
named being distinguished ahiiost entirely by differences in the
reproductive system. Number of chromosomes already recorded
(cf. Muldal, Omodeo) ; 54 = 3n, 72 = 4n, 90 = 5n, 108 = 6n,
160 to 174 = lOn-6 to 20. Ploid}^ varies within the formae. Very
common, probably throughout most of North America, possibly
more so than any other parthenogenetic species. Most of the
material available from a dozen states appears to be referable
to forma macedonica which has, according to Muldal, 53 chromo-
somes = 3n-l.

Calciferous glands, as previously noted by Smith (1924, p. 32,
fig. 44), extend well into x. No variation with regard to that
character was found during examination of a considerable num-
ber of specimens. Differences in appearance and size of "sacs"
as well as of other portions of the gland are recognizable but
presumably are expressions of physiological state and/or variety
of fixation and preservation. Hearts are present in vi.

Atrial glands are lacking but TP glands are present, except in
more advanced stages of morph evolution in which spermathecae
and some of the seminal vesicles also may be absent.

Transfer of rosea to AUobophora, proposed by Pop (1941)
and approved by Omodeo (1955), may prove to be no more
acceptable than retention in Eiscvia where the species may as
well remain, in hope of avoiding frequent changes in name of
a widely distributed w'orm, pending the drastic revision of the
entire family Lumbricidae that is so much needed.

DISCUSSION

The close relationship of the American species of Eisenia to
each other is shown by many characters to which little or no
attention has been given in the past. Among such are: segment
number, maxima — so far as known at present — being 142 in
Idnnhergi, 152 in carolincnsis ; lysis of pigment, at maturity, in
dorsum of viii or ix-xii ; altei'iiation of nephropores between two
widely separated levels and the frequency of intrasegmental
asymmetry in position; size and shape of the typhlosole as well
as its length, termination being anterior to cxvi; abortion of
hearts or commissures in v but presence of such vessels in vi ;

GATES : AMERICAN' LUMBRICIDAE 27

emergence of spermathecae, during their development, into
eoelomic cavities so far that ducts are not confined to parietes ;
the large battery of GS follicles which includes the a-h setae of
xxv-xxx, and frequently of additional segments as well (taxo-
nomic value of characters of genital setae as yet uninvestigated).
Close pairing of spermathecal pores near mD, in these species,
though iH^rhaps not always in the Lumbricidae, absence of
seminal vesicles in x, etc., provide further evidence of close
relationship.

Taxonomically important differences between carolinensis and
ldnnhcrf)i, respectively, are : pigment, red or black ( ? certainly
not red). First dorsal pore located at 5/6 or 4/5. Spermathecae
two or three pairs, with pores on 9/10-10/11 or 8/9-10/11.
Clitellum of 7-10 or 7-8 segments, ending at 31/32-32/33 or at
30/31. Tubercula pubertatis, on xxvii-xxix or xxvi-xxviii. An
"epididymis" present or lacking. Spermathecae, preseptal or
postseptal. GS glands lacking in coelom or present on parietes.
Another difference of some importance, if established, will be
absence or presence on parietes of TP glands. Male tumescences
of XV usually are larger in carolinensis but shape, size and seg-
mental extent, which occasionall.y have been cited as evidence
for specific distinctness in closely related forms (Kobayashi,
1940, p. 289), may have little or no such value. Additional
differences may be anticipated, perhaps in the genital setae, as
sexual setae in other families of earthworms have been thought
to have considerable taxonomic value (cf. Pickford, 1937), as
well as in other systems as yet unstudied.

Difference in pigmentation of two such closely related species
is of especial interest as it demonstrates the futility of attempt-
ing to divide the Lumbricidae (Pop, 1941, p. 508), on the basis of
that character, into subfamilies. Even generic definition in
terms of pigmentation is now impossible, at least in ease of
Eisenia.

Endemicity and distinctness of American species of Eisenia,
of which there now appears to be only two, have been questioned
in several recent contributions ("maldistinte," Omodeo, 1955,
p. 50). "In the eastern part of the U. S. A., a few 'endemic'
species are found, but," according to Muldal (1952, p. 56), "it
seems as if there were no really old species there, and that these

28 BULLETIX : MUSEUM OF COMPARATIVE ZOOLOGY

forms are recently introduced and rapidh' changed from their
European progenitors.''

The European species Avith which relationships first have to
be considered are E. foetida, sul)moniana and nordcnskioldi,
none of which has been characterized adequately for present
needs. E. focfida, the only one available on this continent, ac-
cordingly has to be taken as a typical representative of the
genus of which it presumably is also the type species.

Close relationships of foetida to both American species of
Eisenia are indeed suggested by many characters, but to caro-
linensis more especially because of pigmentation, number of
spermathecae, and location of spermathecal pores. Differences
from carolinensis include : the shorter clitellum of 6-8 segments ;
location of anterior boundary of the clitellum at 24/25 or at some
more posterior level back to 26/27 ; location, usually, of tubercula
pubertatis on xxviii-xxx or xxxi ; abortion of hearts in vi ; reten-
tion of seminal vesicles in x. Physiological differences may be
even more important, for foetida, in spite of rather definite re-
strictions to certain sorts of special habitats, has been able to
avail itself of many of the opportunities for colonization in
remote areas that have been provided, accidentally or inten-
tionally, by man.

More important, however, and di.-spruving the supposed lack
of distinctness, are differences in the ealciferous glands. These
organs, in foetida, according to Smith (1924-, p. 28, fig. 20),
begin only in xi well behind the insertion on the gut of septum
10/11. No variation in that character has now been found, in
many specimens examined for this particular purpose. The
glands in the two American species do appear, in dissections
(I-II), to be restricted to xi-xii, but microtome sections (Smith,
1924, p. 28, fig. 25) showed that the ealciferous lamellae, though
gradually narrowed, extend through all of x in lonnhereji. The
same lack of variation with respect to anterior boundary of
the gland, as in foetida, can be confidently expected in both
American species though the assumption that the boundary is
the same in each requires confirmation for carolinensis.

Differences just mentioned certainly are of considerable im-
portance and may prove to be of value at subgeneric or even
generic level. Resemblances that can justify inclusion of the

GATES: AMERICAX LUMBRICIDAE 29

three species in the same genus are only : greater thickness of
the gland in xii, and absence of any sort of sacs in x.

Presence of calciferous lanielhie between 9/10 and 10/11 is
tentatively assumed to be primitive, and the anterior narrowing
is thouglit to be evidence that evolutionary reduction of the
portion of the gland in x has been under way. Elimination of
calciferous lamellae in x has been (•()mj)leted in fo( tida.

The calciferous gland in hortensis begins, just as in foetida,
in xi. That character, together with location near mD of sperma-
thecal pores, abortion of hearts or vascular commissures in v-vi,
and other similarities, appear to justify retention of hortensis
in the same group as foetida, unless, of course, presence of atrial
and of TP glands in hortensis is to be considered of sufficient
importance for generic distinction. Such importance at present
seems unlikely as those glands sometimes are eliminated, in the
Lumbricidae, within infrasubspecific taxa.

The calciferous gland in E. rosea not only extends well into
X but is there responsible for widening of the gut and for forma-
tion of a pair of "sacs." This certainly is adequate justification
for exclusion from any genus of which foetida is the type.

The calciferous gland in Dendrohaena octaedra is, at least in
l)art, monilit'orm, having a deep external constriction between
the portions in xi and xii. This produces a marked resemblance
to the glands of Umnhergi and Caroline nsis. If sacs are lacking
in X, as now appeai-s to be the case, relationships of octaedra to
other species of Dendrohaena must be reconsidered. If the
thickening of gut wall in the posterior quarter of x ])roves to be
calciferous, relationships of octaedra thereby indicated will be
closer to American than to European species of Eisenia.

Characters provided by calciferous glands have not l)eeii
used in lumbricid taxonomy hitherto, except at species level and
then, only the number of chaml)ers or partitions in the gland.
Such a character now seems unlikely to have much importance
as Omodeo (various papers) already has found intraspecific
variation that nearly reaches known limits. Other and more
readily recognized characters obviously are available. Neglect
of, or inadequate attention to, the calciferous glands in the much
larger family, the Megascolecidae, was in part at least responsible
for the unnatural and morphologically heterogeneous "genera"
on which Stephenson (cf. 1930) based his thesis of convergence.

30 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

That can be concisely but fairly summarized as a belief that an
extant genus evolved, in different parts of the world, from two
or more genera all still extant.

Lumbricid genera still are defined (cf. Stephenson, 1930;
Cernosvitov and Evans, 1947; Pop, 1941) mostly by genital
characters. One such, for example, is number of seminal vesicles,
organs that may undergo all of the standard phylogenetic re-
ductions as well as j^et others within limits of a single infra-
subspecific taxon. All other structures of the same system, ex-
cepting only the ovaries of xiii, also are subject to change at
species or lower level. In large sections of various earthworm
families the genital organs provide most of the macroscopic
characters by which species are distinguished and defined. This
remarkable evolutionary plasticity (cf. Gates, 1956 b-e) of the
reproductive system necessitates considerable caution in use of
its structures for taxonomic purposes at genus or higher level.

Pigmentation is one of the four somatic characters and that
by which Pop (1941) would subdivide the Lumbricidae. The
procedure proposed would place the two closely related Ameri-
can species, as already noted, in different subfamilies. Gizzard
characters, at least as usually stated, no longer are diagnostic
even for Eiseniella. The prostomium may always be tanylobous
in Lumhricus but the same shape is found occasionally in all but
one of the other genera. Setae have provided the fourth charac-
ter, the value of which is shown by the following quotations
from generic definitions (Cernosvitov and Evans, 1947) : "Setae
more or less closely paired," Allolohophora ; "closely or widely
paired (or distant)," Bimastos; "mostly widely paired or dis-
tant, rarely closely paired," Dendrohaena ; "closely to widely
paired or distant," Eisenia; closely paired (seldom widely
paired)," Eophila; "(mostly distant) rarely closely paired,"
Octolasinm. Such obviously undiagnostic characters, in Pop's
system (1941), are second in importance only to pigmentation.
Even with more recent changes suggested by Pop and Omodeo,
the lumbricid genera appear to be only congeries of species asso-
ciated because of relatively unimportant or superficial converg-
ences.

As one rather indirect result of a study of evolution of repro-
ductive organ polymorphism (Gates, 1956c) it is now possible
to suggest, but only very tentatively because of ignorance of so

GATES: AMERICAN LUMBBICIDAE 31

much of earthworm anatomy, some of the directions in which
evolution may have proceeded in the Lumhricidae, with refer-
ence especially to the species that have been considered above.

From a hypothetical protomeg:adrilid to be characterized else-
where, an ancestral protolumbricid can be derived by the follow-
ing changes: Acquisition of dorsal pores at each intersegmental
level. Dislocation laterally of spermathecal pores from region
of AB to CD. DiflPerentiation of tubercula pubertatis within
the clitellum and in association therewith of coelomic glands
(TP) of unknown function. Elimination of any oesophageal
gizzard. Extension of oesophagus into xv. Union at median
margins of calciferous lamellae of x-xiii or xiv to i)roduce an
intramural gland opening anteriorly into gut lumen. Develop-
ment of an intestinal gizzard in xvii and behind that of a typhlo-
sole. Differentiation of a subneural trunk and, median to tlie
hearts, of extra-oesophageal trunks. Elongation of male deferent
ducts to open to exterior in BC at equator of xv. Invagination
into coelom of xv of atria (with glands thereon) through which
male ducts opened to exterior. Slight elongation of oviducts to
open at equator of xiv just lateral to B. Specialization of setae
in anterior segments for genital functions. Other characteristics
that can be assumed are : presence of lateral hearts or commis-
sures in v-xi. Penetration of nephridial ducts into parietes to
open externally at or near B. Presence of glands of unknown
function in association with follicles of genital setae, and presence
of four pairs of seminal vesicles in ix-xii. The si)ermathecal
battery alread\' had been enlarged by addition of two pairs with
pores at 9/10-10/11, or shortly was so augmented.

A more recent ancestor of most species hitherto included in
Eisenia had continued dislocation of spermathecal pores toward
mD and regardless of positions of setae, begun reduction of
calciferous lamellae of segment x, aborted hearts or commissures
of V, acquired a tendency for nephridial ducts during embryonic
development and, after parietal penetration, to grow laterally
before passing on to the epidermis, eliminated the original pair
of spermathecae that had opened at 8/9, aborted atria and even
the atrial glands. If that ancestor had a clitellum of six or
seven segments with anterior margin at 23/24 little change addi-
tional to acquisition of the peculiar dark pigment, and possibly
elimination of TP glands, is required to give Jonvhergi. No evi-

32 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

dence has been found of any tendency to eliminate the remain-
ing; pair of ancestral spermathecae. On the contrary, such varia-
tion as has already been recognized seems indicative of increase
in size of battery by addition of a further pair opening at
11/12, a change that already has been made in other evolutionary
lines.

In evolving from its common ancestor with Idnnhergi, caro-
linensis has acquired a different kind of pigment, eliminated
the last of the original pairs of spermathecae, eliminated (or
reduced) the CIS glands as well as the TP glands if the latter
had not been lost previously, and shifted the tubercula into
xxvii-xxix.

Tubercula, in xxvi-xxviii in Idnnhergi, xxvii-xxix in carolinen-
sis, xxviii-xxx or less often xxviii-xxxi in foctida, xxix-xxxi in
rosea, xxx-xxxi in hortensis, xxxi-xxxiii in octaedra, provide an
interesting sequence. Though the series may be indicative of the
way evolution has proceeded it does not, in this case, evidence
phylogenetic relationships. Transfer of rosea from Eise7iia to
some other genus, as already mentioned, is necessary. American
endemics may not be congeneric with the European forms to
which they seem to be most closely related. The transatlantic
species, having lost calciferous lamellae in x, and the hearts of
vi, are more advanced but organs that have been left out of
consideration in the past are suggestive of long divergent
lineages. Thus, hortensis, which has retained atrial glands after
loss of atria as well as the coelomic TP glands, now appears to
be older than foctida, in which both kinds of glands have been
lost. D. octaedra, which has retained atrial and TP glands,
except in advanced stages of infrasubspecific morph evolution,
then would be older than hortensis as spermathecae still must
be developed at levels determined by location of a particular
rank of setae. Female pores, like the spermathecal pores, are
more dorsal than in many lumbricids but because of shift in the
b rank in association with which the oviducts seem obliged to
acquire their external openings.

Abortion of hearts in x-xi of octaedra, except for vestiges in
a rare individual, has been completed. Presence of hearts in xii
has been recorded in an occasional lumbricid species but all such
instances require confirmation because of known misidentifica-
tions of large vessels connecting extra-oesophageal and dorsal

GATES: AMERICAN LUMBRICIDAE 33

trunks. Behind xi, in the Lumbrieidae as in the Oenerodrilidae,
hearts ])robably have lieen developed rarely, if ever. A snpra-
oesopha^eal trunk seems never to have been developed in the
Lumbrieidae. Latero-oesoi)hageal and oesophageal hearts are
therefore impossible ; all are lateral.

The g-eog'raphical ranges of our native forms are as yet little
known, earthworms having been collected mostly in and around
municipal centers, farmyards, golf courses, etc., where intro-
duced exotics are most likely to be found. Considerable approxi-
mation to extermination of endemics in com})etition with the
intruders certainly seems to have been implied by Smith (1928).
llis conclusions, according to one of his graduate students
(verbal communication) who secured many of the Illinois speci-
mens mentioned in that contribution, were largely based on ma-
terial from urban areas and banks of streams contaminated by
sewage and industrial wastes. The influence of man, on the
competition he has been responsible for initiating, certainly
ought to be investigated in other sorts of habitats. Meanwhile,
species of Eisenia apparently are to be sought in often neglected
niches by stream banks and swamps, in forests and especially
perhaps in the mountains of the Appalachian chain. The dis-
tributional ranges as now known, except for \evy recent coloni-
zation in the extreme southwest of New England, extend from
New Jersey into Louisiana and are south of the limit of glacia-
tion. Paucity of endemics of Eisenia in the vast area of North
America may, then, be due, in part, to extinction of other species
during the last glacial period. A similar paucity of species in
Eurasia, where several different phylogenetic lineages (instead
of one) now seem to be required, likewise may have resulted,
in part, from ice age extermination.

A score or more of lumbricids, mostly of European origin,
have been carrie^l to almost every part of the world and to many
places very many times, since the days of Henry the Navigator.
Some (cf. Ciates, 1954, p. 255) have become established, even
widely distributed, in remote regions where they would not
have arrived unaided. Intentional introductions of several of
the most successful of the colonizing species have, however, often
failed and the characterization of "cosmopolitan" (cf. Cer-
nosvitov and Evans, 1947; Muldal, 1952) is unwarranted, prob-
ably for any lumbricid. Only thirteen individuals, all of a

34 BTLLETIX: MUSEUM OF COMPARATIVE ZOOLOGY

single and possibly American species, were secured during a
four year survey (Gates, 1945, 1947, 1956a) of the earthworm
fauna of the Allahabad sector of the Gangetic valley and an
adjacent area to the south from which more than 50,000 speci-
mens were identified. Furthermore, a thirty -year study of the
earthworms of southeast Asia showed that in spite of frequent
introductions, intentional and accidental, lumbricids have not
been able to nuxintain themselves where climate is tropical. Only
in hill resorts at elevations of more than four thousand feet, in
peninsular- India, Burma and the Malay Peninsula, has there
been some successful colonization. To find lumbricids elsewhere,
it is necessary to go well above the tropics, even north of the
Gangetic and Brahmaputra valleys into the eastern Himalayas.
There, and again so far as is now known, in and around centers
of European influence (Gates, 1951), endemics probably have
l)een replaced by exotics some of which, however, are megascole-
cid. Indigenous lumbricids can be expected only in the western
Himalayas and the far uorthAvestern corner of India. Whatever
the nature of the restriction, even the most successful of the
colonizing lumbricids now appear to be limited to temperate
zone types of climate. The less adaptable American endemics,
in past ages, may then have been similarly prevented from estab-
lishing themselves in tropical portions of the continent.

►SUMMARY

American species of Eisenia are closely related, obviously
■'good," and clearly distinguishable from their European con-
geners. The latter, in accordance with newlj- i)roposed evolu-
tionary sequences based on characters hitherto unrecorded or
neglected, now appear to be more advanced. Paucity of species
on the American continent is attributed to inability to colonize
in tropical climates, and to ice-age extinctions.

Lumbricid genera of the classical system now appear to be
mostly congeries of si)ecies associated because of relatively recent
and superficial convergences acquired at species or, sometimes,
even lower levels. Genera must be redefined and by reference to
more conservative organs less liable to rapid change during in-
frasubspecific evolution.

GATES : AMERICAN LUMBRICIDAE 35

vri

BiMASTOs ZETEKi Siuith aiid Gittins 1915

Highlands, North Carolina. In loaf mould and saudy soil (pH ca. 5.0)

near path loading up to Whitosido Mountain, at altitude of ca. 4500 ft..

August 21, 1932, 2 0 1. Dr. G. K. Pickford. In moderately damp, loamy

.soil by path to Primeval Forest, at altitude of m. 3900 ft., July 24,

1931, 01-0. J. M. Valentine per Dr. G. E. Pickford. In pocket of sandy

black soil (])H 5.0) under dead leaves by stream no;ir path to Primeval

Forest, July 25, 1931, 0-0-1. Dr. G. E. Pickford.
Linville, North Carolina. In earth (j)!! between 5.0 and 6.0) and leaf

mould by stream in ravine at 4000 ft., on road to Blowing Rock, July

31, 1931, 0 0-4. Dr. G. E. Pickford.
Indian (iap (now Newfound (iap;, Tennessee. In loamy soil (pH 5.0)

among rotten wood by stream at ca. 4500 ft., near road and ca. 500 ft.

below gap (Great Smoky Mountains), July 30, 1931, 0 0 9. Dr. G. E.

Pickford.
.Mt. Carmel, Connecticut. Jn rotten log, April 14, 1946, 0-0-1. Dr. G. E.

Pickford.
Whitetop Mountain, Virginia. Under bark of log at elevation of 4700 feet,

August 17, 1955, ?-0 1. Prof. Walter Harman. (15 small .juveniles, or

some of them, may be of this species.)
Mountain Lake, Virginia. Leaf mould at reservoir of Biol. Sta., June 20,

1955, 0 0-3. Sawdust pile. Castle Rock, July, 1955, 16-0 0. Castle

Rock; August 12, 1955, 24-0 0, August 2, 1955, 0-1-0. Prof. Walter

Harman.
Culver, Indiana, June 19, 1914, 0 0-2. II. V. Heimburger. (U. S. Nat. Mus.

No. 26,214. FLxamined by courtesy of Dr. Fenner Chace.)

External characteristics. Length of worms not obviously am-
jmtees, 73-99 mm. (Carolina, Tennessee), 95-100 mm. (Indiana,
relaxed?), to 135 mm. (Virginia, not relaxed). Diameter, 4 mm.
(Indiana), o-S mm. (others). Segments, 12G-154 (cf. Table 5),
135 (Indiana). Pigmentation unrecognizable in old (1914-1946)
specimens and in formalin as well as in alcohol preservation.
A red coloi", in old formalin material, usually is present in epi-
dermis of the region between xv and the elitellum. The color
may l)e lacking on xvi-xvii, indistinct or slight and then equa-
torial in the dorsum only. The colored band becomes wider
posteriorly until intersegmental furrows are nearly reached and
continuous around the ventrum, finally becoming indistinguish-
able from the elitellum. Pigmentation red, in recently preserved
Virginia worms, in circular muscle layer and at both ends of

36 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

the body also in an outer portion of the longitudinal muscula-
ture. The color gradually disappears ventrally in BC, except in
first and last few segments where the ventrum is red, and may
be sparse except at mD in dorsum of probable cliteUar region
in largest juveniles. The body tends to be four-sided posteriorly
though the dorsum, slightly wider than the ventrum, remains
slightly convex. Prostomium epilobous, tongue open or closed
(3 specimens, but adventitious?). Setae begin on ii on which all
usually are piesent, one pair at each angle of the body posteriorly
where CD < or ca. == AB, BC < AA < DD < JC. Nephro-
pores first recognizable on iii, usually close to or above D on
anterior segments, posteriorly lateral to B (between levels of
male and female pores) or in DD, sometimes well above Z),
without regular alternation and often at different levels on op-
posite sides of a segment. Both situations, with or without sym-
metry, have been found on xiv-xvi. Pores of six consecutive
segments occasionally may be at one level on one side in the
intestinal region but in each worm the pores on opposite sides of
the same segments alternate between the two positions. First
dorsal pore on 4/5 (1), ?5/6 (3), 5/6 (59), 7/8? (1).

Female pores equatorial, closer to 1) than a is, on xiv. Male
pores each in a small, shallow, transversely slit-like depression in
median half of BC, equatorial on xv.

Clitellum red (old formalin specimens) or wliite (old alcoholic-
and recent formalin material), saddle-shaped or annular, located
as shown in Table 4. Tubercula pubertatis lacking (all).

Genital tumescences lacking or so slightly developed as to be
almost unrecognizable. The o-h setae of certain segments are
retracted, the margin of eacli follicle aperture tumescent, the
annulus so small as to be easily overlooked. Genital tumescences,
as so indicated, are on xvi (16 Carolina and Tennessee worms, 1
from Whitetop Mt., Virginia, possibly also in several small
juveniles from the latter locality), on xxvii (6), xxvii-xxviii
(11), xxvi-xxviii (1), all from Mountain Lake, Virginia. Male
pore tumescences are small, scarcely or only slightly protuberant,
confined to xv, sometimes (old formalin material) indicated
merely by an epidermal translucence that does not reach either
14/15 or 15/16.

Internal anatomy. Septa 7/8-9/10 and 12/13-14/15 (especially
13/14) rather generally are muscular and thicker than 10/11-

GATES: AMERICAN LUMBRICIDAE 37

11/12, 10/11 the thinner, but none is, compared to size of body,
very much thickened. Long^itudinal muscle band at mD of freshly
preserved material has a pink color. Peritoneal pigment flecks
are lacking. Epidermis at parietal incisions of recent formalin
material perfectly' white as is circular musculature just undei--
neath intersegmental furrows.

Calciferous gland extends well into x, often reaching nearly
to 9/10 but only apparently, usually much wider in x, and then
with an appearance as of a pair of vertical sacs. Constrictions
at 10/11 and 11/12, when recognizable, are slight, but that in
region of insertion of 12/13 and usually internal on both sides
is marked. The gut lumen through the gland (x-xii) may be only
a vertical cleft or wider and then variously so, sometimes wider
than wall of gut. Oesophagus narrowed through xiii-xiv. In-
testinal origin in xv (25). Gizzard in xvii-xviii (25), the layer
with brilliant muscular sheen obviously extending somewhat
behind insertion of 17/18 in horizontal sections but not reaching
that of 18/19. Gut narrowed in xix and valvular at 19/20.
Typhlosole, beginning gradually or more abruptly in region of
xxii-xxvi, 1^-2 mm. high, reaching to or nearly to floor of gut,
shape as in Eisenia Idnnbergi Michaelsen 1894, often with a
deep longitudinal groove on ventral face of largest portion (an-
teriorly), ending abruptly or otherwise as shown in Table 4.

Subneural trunk usually large and filled with blood (occa-
sionally (juite unrecognizable), adherent to cord, continued for-
ward to subpharyngeal ganglia. Extra-oesophageal trunks turn
upward posteriorly in xii to dorsal trunk, the vertical portion
often distended with blood and looking much like a heart. A
slender )K)sterior continuation behind the upward curve is
shortly recognizable only rarely. A branch of the extra-oesopha-
geal, as large as the vertical portion in xii, passes upward in
ix (several worms), through 9/10 and to the dorsal trunk just
in front of the calciferous gland. From the subneural a-large
branch passes out in xii, xiii or xiv, on one or both sides, to
join the extra-oesophageal in xii just as the latter turns up.
The whole subneural trunk sometimes appears to turn out to one
side to the extra-oesophageal but a slender continuation usually
is recognizable further forward on the cord. The dorsal trunk
gives off four large branches to the calciferous gland in x, two
anteriorly, the others under the hearts. The trunk is empty

38 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

anterior to 6/7 (all), quite small, unrecognizable in front of
5/6 though probably not ending there. Hearts or commissures
were not found in v-vi. Hearts of vii often rather small and
empty. Last hearts in xi (25). Nephridia have been definitely
identified only from iii posteriorly (present in ii?). Nephridial
ducts pass into parietes at B. Brain, the posterior margin
deeply incised, left in iii by a transverse section exactly along
3/4. Median, longitudinal groove in roof of pharynx in iv in
cross section Y-shaped.

Seminal vesicles two pairs, in xi-xii (25), not rudimentary
but rather small to quite small, in dorsal half of segment or be-
tween upper face of gut and dorsal parietes. Male funnels crenel-
late, small, often only two to three times the size of the nephridial
funnels just beneath them, somewhat larger in the Indiana worm,
with some opacity (usually) or translucent. Male deferent ducts
slender, just behind funnel septa straight, sinuous, zigzag looped
or loops (various) held together by connective tissue in a small
flat disc, visible on parietes (southern worms) into xv. Sperma-
thecae lacking (25) and rudiments unrecognizable in parietes
after stripping oif longitudinal musculature. Oviducts pass into
parietes anterior to eq/xiv. Ovisacs, always present (25), may
be of about the same size as the ovaries and slightly lobed.

Atrial gland nearl}^ circular, flattened against parietes, crossed
l)y an equatorial cleft, another cleft at insertion of 16/17, the
gland extending into xvi (19).

Follicles of the ventral setae in xv-xxxviii usually appear to
be slightly larger than those of the lateral setae in the same
segments. None of them, in the Indiana w^orm, appear to be
especially enlarged. Special enlargement in other specimens is
associated with the external modifications already mentioned.
(xS follicles of xvi are surrounded on the parietes by an acinous
glandular mass or the lobules may be present only on posterior
and median sides. No such glands are recognizable in posterior
segments.

Juveniles. Atrial glands are lacking in worms as large as 110
X 8, 125 X 6 nnn., even in one 135 mm. long, but without any
recognizable epidermal tumescence in presumed clitellar region.
GS follicles are, however, protuberant into coelom and modifica-
tions around the follicle apertures are recognizable. Associated
glands are not yet recognizable in the coelom. Enlargement of

GATES: AMERICAN LUMBRICIDAE 39

setal follicles may be recognizable in much smaller juveniles on
some of which rudiments of male pores and their tumescences
are not yet detectable.

Abnormality. Metamerism is abnormal in one or more regions
of a posterior portion of the body in two worms.

Regeneration. An anal seg^nent had been reconstituted in each
of the posterior amputees. No other regeneration had taken place
though amputation in some of the cases certainly had not been
recent. Posterior regeneration, as in Lumhricus terrestris L.,
may not be possible in ordinary conditions.

Parasites. Large, thin-walled cysts with a watery content
(2-6 per segment in coelomic cavities of x-xi) were present in
seven Carolina-Tennessee worms in one of which each of the
unusually large seminal vesicles contained 2-4 more of the cysts.
Further cysts of the same sort were present in the coelom of the
last fifteen segments of one worm, along with much smaller
opaque spheroidal bodies. The latter were scattered through the
postclitellar portion of the body of several worms but were
especially numerous in the last fcAV segments of posterior am-
putees along with ' * brown ' ' bodies, some of which were white.

Life history. Clitellar tumescence had become unrecognizable
in a July worm but the site of the clitellum was indicated by the
usual postreproductive discoloration. Full clitellar tumescence
may not have been attained by several July worms in which
intersegmental furrows still were faintly visible on the dorsum
or in which acinous glands around GS follicles still were very
small. The clitellum of the April and some July specimens was
at or near maximal tumescence. Three spermatophores on one of
the Carolina-Tennessee worms are firmly attached to xxviii just
lateral to B and in CD and to xxv at m\'. Each is a transparent
flattened, nearly square disc containing centrally a rather small
ellipsoidal to ovoidal wiiite body.

The breeding season, judging from clitellar development,
spermatophores, etc., would seem to include April in Connecti-
cut, June in \'irginia, July in the mountains of Tennessee and
North Carolina. During June, in Indiana, cocoons were found
according to Heimburger (1915, p. 285) and worms Avere at
"height of sexual maturity." Material that would provide
evidence as to a fall breeding season has not been available.

40 BULLETIX : MUSEUM OF COMPARATIVE ZOOLOGY

No spermatozoal iridescence was recognized on male funnels
of any worm including one from Indiana, nor in the opaque
central bodies of the spermatophores. Male funnels are not of
normal size and development in some of the worms. Seminal
vesicles are of about the same small size in some worms with
maximal clitellar tumescence as in the postsexual aclitellate and
in juveniles. The largest seminal vesicles are merely mem-
branous sacs filled with cysts. The worms, if deductions as to
breeding season are correct, are male sterile in which case re-
production must l)e parthenogenetic.

Remark.s. The red color of the epidermis from xv-xxvi, like
that of the clitellum, presumably is a formalin-developed artifact
but must indicate, because of constant localization, some chemi-
cal difference from other regions. The difference probably i:>
established at maturity and the change must be reversible during
postreproductive regression as the color was lacking in the
regressed aclitellate worm. The brown of the "purplish-brown"
coloration mentioned by Smith (1917, p. 175) may have been
a product of postreproductive metabolic changes.

The clitellum in this species, according to the original de-
scription, covers 11 or 111 segments, xxvii-xxxvii or eq/xxvii-
xxxviii. Only xxvii-xxxvii is mentioned by Smith in his subse-
quent characterization of the species (1917) though he had "con-
siderable numbers" of worms from three states. No further data
have been provided subsequently by authors who recorded zeteki
from New York, Ohio, Indiana, Michigan, Missouri, Arkansas,
Louisiana and North Carolina. In absence of such data it should
l)e assumed that the clitellum was on xxvii-xxxviii in material
from all of those localities since location of that organ has been
about the only character ust^d for routine identification of species
of Bimastos. However, no single individual herein considered
has a clitellum exactly on xxvii-xxxvii. Anterior and posterior
margins both appear to be rather indistinct often, due to a
gradual rather tiian the usual abrupt decrease in epidermal
thickness. At present it seems that the clitellum may cover nine
to fourteen segments, beginning at or behind 25/26 and ending
between 35/36 and 39/40. Location of anterior margin at or
behind 25/26 seems to be about the only character now available
to distinguish zefeki from Jongicincfits Smith and Gittins 1915.

GS follicles have not been recorded hitherto. The}' may have

GATES: AMERICAN LUMBRIC'IDAE 41

been lacking in Illinois and Indiana worms. If so, two other
strains may be recognizable, one with follicles in xvi (or xv-xvi)
only and associated with acinous glands, another having the
follicles more posteriorly, in the region of xxvi-xxviii, but no
glands. Strain ii (Carolina to Virginia), with acinous glands
such as are present in the sexual paliistris, may be the most
primitive. Strain iii (Mountain Lake, Virginia) without acinous
glands, then would be more advanced and i (Indiana, Illinois?)
the most advanced. The latter is the one nearest to the southern
limit of the recent glaciation.

Parthenogenesis is suspected in two other species of Bi)itasi<>s
(cf. III-V), tumidus and paribus, and in the latter may in part
have been responsible for colonization of various extra-American
regions, but has not been similarly advantageous to zeteki.

Spermatophores must have been developed, in the ancestry of
zeteki, before spermathecae were eliminated. They still can be
produced and i)resumably are attached to the surface of the
body during a futile copulatory act (pseudocopulation). The
male sterility could have become established in this form only
after modification in oogenesis to permit parthenogenesis.

42

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Table 4
Clitellar characteristics in Biniastos zetehi

Clitelluni

Epidermis

on

Tumes-

red on

segments

Shape

cence

Locality

segments

1.

26-35 (?)^^

?

0

Highlands

0

2.

26-35

A

medium

Highlands

f

3.

26-35

A

maximal 1

Linville

17-25

4.

26-35

A

marked

Indian Gap

16-25

5.

26-35

A

marked

Indian Gap

16-25

6.

26-35(36f)

S

medimu

Indian Gai)

24-25 + VC

7.

26-35(36?)

A

maximal?

Indian Gap

(17 18?) 19 2.-,

8.

26-35(36?)

A

marked

Indian Gap

16-25

9.

26-36

A

marked

Linville

17 25

10.

26-36

A

maximal ?

Linville

17-25

11.

26-36

A

marked

Indian Gap

17-25

12.

26-36(37?)

S

medium

Indian Gap

0 VC

13.

eq/26-36/eq

A

marked

Indian Gap

17 25

U.

eq/2636/eq

A

marked

Highlands

17-25

15.

(26-27)28-

37(38-39)

S

marked

Mt. Carmel

?

16.

27-35(361)
26-35

A

S

maximal
maximal ?

Indian Gap
Whitetop Mt.

17-25

17.

18.

(26?)27-37(38?)

s

maximal ?

Virginia

19.

(26?)27-38/eq

s

maximal?

Virginia

20.

(27?)28-37

s

maximal ?

Virginia

21.

27-38/eq

s

maximal ?

Indiana

22.

28-38/eq

s

maximal ?

Indiana

•Site of clitelluni indieati'd by pustreproiluctive discoloration.

A. Annular. S. Saddle-shaped. eq. Equator.

VC. Epidermis iu .-l.-t as red as clitellum.

(86?) Epidermis of dorsum hi xxxvi red, slightly thickened, decreasingly so
anteroposteriorly.

(26-27) Epidermis red, imieasin^'ly thickened anteroposteriorly in xxvi-xxvii
but furrow 20/27 still visible.

Dorsal pores are occluded except in an early staj?e of clitellar development. Inter-
segmental furrows are obliterated as tumescence becomes marked but even
when clitellum is annular still are slightly indicated in BB or AA according
to degree of tumescence. Setae and nei)hropores retained.

Thickening of epidermis in AA or BB in worms with a supposed saddle-shape of
clitellum, might have been recognizable in sections.

GATES : AMERICAN- LUMBRICIDAE

4:^

Table 5
Typhlosole termination and segment number in Bimastos zeteki.

Typhlosole Atyphlo- Number
ends in solato of

segment segments segments

Locality

Remarks

76

18

94

Highlands

amputee

80

16

96

Linville

amputee

84

16

100

Indian Gap

amputee

85

22

107

Highlands amputee? juvenile

87

20

107

Indian Gap

amputee

89

11

100

Whitetop Mt.

amputee?

96

33

129

Highlands

97

24

• 99+

Indian Gap Bi

•oken off posterior
end lacking

97

29

126

Indian Gap

amputee?

98

28

126

Linville

100(±l-2)

40

140(±l-2)

Linville

101

30

131

Indian Gap

103

45

148

Linville

104

35

139

Indian Gap

104

37

141

Mt. Lake, Virginia

104

42

146

Indian Gap

104+

?

104+

Indian Gap Broken off posterior

end lacking

103-106

43-46

149

Mt. Lake, Virginia

106

38

144

Highlands

juvenile

106

45

151

Mt. Lake, Virginia

97-107

15-25

122

Mt. Lake, Virginia

104108

20-24

128

Mt. Lake, Virginia

108

44

152

Indian Gap

109

29

138

Highlands

amputee 1

109

33

142 ■

Mt. Carmel, Conn.

110

44

154

Mt. Lake, Virginia

(±1-2) Anterior end which had been broken off in front of clitellum was lacking.
103-106 Typhlosole terminates very gradually, rather than abruptly, exact end
of the organ uncertain.
Pofeterior amputation f)f long standing may be unrecognizable; because of re-
arganization of last segments. Number of segmients in "complete specimens",
according to Smith (1917, p. 175), usually 130. The maximum he mentioned, 142.

44 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

DISCUSSION
The genus Bimastos was erected for a single American species,
palustris Moore 1895, which lacks tubercula pubertatis, seminal
vesicles in ix-x, as well as spermathecae. The genus shortly
became (cf. Michaelsen, 1900) a taxonomic waste-basket into
which similarly defective forms, regardless of morphology and
geographical distribution, were thrown. One of those so mis-
placed is AUolohopliora tenuis Eisen 1874 which occasionally has
one to three, more or less rudimentary spermathecae as well as
more or less indistinct markings assumed to be tubercula. Be-
cause of a "tendency" to develop spermathecae, Smith (1917,
p. 178) suggested that an admittedly confused taxonomy could
be somewhat simplified by tranfer of tenuis to Dendrohaena
Eisen 1874 which has spermathecae. Actually, most specimens
of tenuis, as long suspected (cf. Gates, 1953, p. 531, footnote),
have been of an A morph of D. ruhida (Savigny, 1826), the
individuals with more or less rudimentary spermathecae being
"intermediates" in which elimination of spermathecae has not
yet been completed (cf. Gates, 195 ?c). D. ruhida obviously is
exotic in America. Other Eurasiatic A morphs that have been
erroneously referred to Bimastos may prove to be capable of
similar association with a normal H morph, when intermediates
have been found if not before. The finding of H morphs of the
various American species of Bimastos cannot now be anticipated
as the only other endemic lumbricids are distinguishable at
generic level.

EEFEEENCES

Cernosvitov, L. and A. V. Evan.s

1947. Lumbricidae. Linnean Soc. London, Synopses British Famin,
6: 1-36.

Davies, H.

1954. A preliminary list of the earthworms of northern New Jersey
with notes. Breviora, 26: 1-13.

Gates, G. E.

1943. On some American and Oriental earthworms. Ohio Jour. Sei.,

43: 87-116.
1945. The earthworms of Allahabad. Proc. Nat. Acad. Sci. India., 15:

45-56.

GATES: AMERICAX LUMBRICIDAE 45

1947. Earthworms of the Allahabad sector of the Gangetic plain.
Proc. Nat. Acad. Sci. India (B), 17: 117-128.

1951. On the earthworms of Saharanpur, Dehra Dun, and some Hima-
layan hill stations. Proc. Nat. Acad. Sci. India (B), 21: 16-22.

1953. On the earthwojms of the Arnold Arboretum, Boston. Bull.

Mus. Comp. Zool., Harvard, 107: 500 534.
195-4. Exotic earthworms of the United States. Bull. Mus. Comp. Zool..

Harvard, 111: 219-258.
1955. Notes on American earthworms of the family Lumbricidae. I II.

Breviora, 48: 1-12.
1956a. Earthworms of the region south of the Allahabad sector of the

Gangetic plain. Proc. Nat. Acad. Sci. India (B), (in press).
1956b. The earthworms of Burma. VII. The genus Euiypiioeus. With

notes on several Indian species. Eec. Indian Mus. Calcutta, (in

press).
1956c. Reproductive organ polymorphism in earthworms of the oriental

megascolecine genus Phrretima Kinberg 1867. Evolution, (in

press).

Heimburger, H. V.

1915. Notes on Indiana earthworms. Proc. Indiana Acad. Sci., 1914:
281-285.

KOBAYASHI, S.

1940. Terrestrial Oligocliaeta from Manchoukuo. Sci. Rep. Tohoku
Univ., (4) 15: 2G1-315, figs. 1-15.

MiCHAELSEN, W.

1900. Oligochaeta. Das Tierreich, 10.

Moore, H. F.

1895. On the structure of Bimastos palustris, a new Oligochaete.
Jour. Morph., 10: 473-496, pis. 26-27.

MULDAIi, S.

1952. The chromosomes of the earthworms. I. The evolution of
polyploidy. Heredity, 6: 55-76.

Omodeo, p.

1952. Cariologia dei Lumbricidae. Caryologia, 4: 173-275.

1955a. Lombrichi cavernicoli di Grecia e Turehia raccolti dal Dr. K.

Lindberg. Ann. 1st. Zool. Univ. Napoli, 7: 1-16.
1955b. Aspetti biogeografici della speciazione. Boll. Zool., 21 (2): 156.

46 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

PiCKFOED, G. E.

1937. A monograph of the acanthodriline earthworms of South Africa.
Cambridge, England.

Pop, V.

1941. Zur Phylogenie und Systematic der Lumbriciden. Zool. Jahrb.
Syst., 74: 486-527.

Smith, F.

1917. North American earthworms of the family Lumbricidae in the

collections of the United States National Museum. Proc. U. S.

Nat. Mus., 52: 157-182.
1924. The ealciferous glands of Lumbricidae and Diplocardia. Illinois

Biol. Mon., 9 (1): 1-76.
1928. An account of changes in the earthworm fauna of Illinois and

a description of one new species. Bull. Illinois Nat. Hist. Survey,

17: 347-362.

Stephenson, J.

1930. The Oligochaeta. Oxford.

Publications Issued by or in Connection

WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

Bulletin (octavo) 1863 — The current volume is Vol. 115.

Breviora (octavo) 1952 — No. 57 is current.

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

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

Occasional Papers of the Department of Mollusks (octavo) 1945 —
Vol. 1, no. 18 is current.

Proceedings of the New England Zoological Club (octavo) 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.

The continuing publications are issued at irregular intervals in numbers
which may be purchased separately. Prices and lists may be obtained on
application to the Director of the Museum of Comparative Zoology,
Cambridge 38, Massachusetts.

Of the Peters "Check List of Birds of the World," volumes 1-3 are out
of print; volumes 4 and 6 may be obtained from the Harvard University
Press ; volumes 5 and 7 are sold by the Museum, and future volumes will be
published under Museum auspices.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. 115, No. 2

A QUANTITATIVE STUDY OF THE EQUIDAE OP
THE THOMAS FARM MIOCENE

By Robert S. Bader

University of Florida

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

August, 1956

No. 2 — A Ouanlitiil/vc Study oj the Equ'idae of the Thomas
Farm Miocene

Bv Robert S. Bader

INTRODUCTION

The most important terrestrial vertebrate locality in the Ter-
tiary of eastern North America is the Thomas Farm of north
central Florida. More terrestrial mammalian species have been
described from this Gilchrist County Miocene locality than from
any other Tertiary deposit east of the High Plains. A general
review by Romer (1948) includes a complete bibliography of the
work to that date. Since 1948. additional herpetological remains
have been described by Tihen (1951), Vanzolini (1952), Williams
(1953), Coin and Auft'enberg (1955), and Auffeuberg (1956).
A new bird genus has been described by Brodkorb (1954).

The most abundant species occurring at this site are included
in the Equidae, especially in the genus Parahippiis. These as-
sume primary importance not only because of their frequency
of occurrence but also because of the critical stage of equid evolu-
tion which they represent. This most interesting and crucial
phase in the history of the family was marked by an increase
in gross size, in hypsodonty and in the intricacy of the dental
pattern, as well as by the appearance of cement on the teeth.
Coincident with, and inferentially in consequence of, these wide-
spread morphological changes there occurred an equally mo-
mentous ecological shift from a browsing to a gi'azing habitat.
Suljsequent to the last detailed discussion of the horses (White,
1942), the accruement of material has been considerable (pri-
marily through the efforts of the Museum of Comparative Zool-
ogy ; recently, collections have been made for the University of
Florida by the author). Because of this fact and since the large
collection is amenable to statistical analysis, a quantitative study
was undertaken in the hope of shedding new light on the relation-
ships of the several species previously described.

Simpson (1932) recognized 3 species from the earlier Miocene

50 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

deposits of Florida: ArchaeoMppus nanus Simpson from the
Thomas Farm, Parahippus leonensis Sellards (1916) type from
a well on the Griscom Plantation near Tallahassee and referred
specimens from Thomas Farm, and Anchitherium clarencei Simp-
son, type from Midwaj^ also near Tallahassee, and referred mate-
rial from Thomas Farm. White (1942), with a much more exten-
sive series at his disposal than was available to Simpson ten years
earlier, recognized seven species from Thomas Farm, viz., Mio-
hippus sp., Parahippus hlackhcrgi, Parahippus harhouri, Para-
hippus leonensis, Merychippus gunteri, Anchitherium clarencei,
and Merychippus westoni. The last named species, found near
Newberry, (Alachua County), Florida, and described by Simpson
(1930), was not identified with certainty by White and none of
the more recent material can definitely be assigned to it. A lower
jaw was referred to A. clarencei by White in addition to the ma-
terial described earlier from this locality by Simpson (1932).
Although there is no doubt that this large, conservative form did
occur at Thomas Farm, more recent collecting has failed to reveal
any new material of it.^

The great bulk of the equid material is composed of repre-
sentatives of the four remaining species recognized by White,
and it is principally with these that the present study is con-
cerned. P. hlackhcrgi was originally defined on the basis of ma-
terial from the Garvin Gulley, Texas, and placed in Miohippus
by its author. Hay. It was assigned to Archaeohippus by Stirton
(1940) in his general review of the family. As mentioned above,
Simpson described a new species, A. nanus, from the Thomas
Farm material available to him in 1932, but White (1942)
referred to this form as P. hlackhcrgi, placing A. nanus in syn-
onymy. P. harhouri was described by White from the Thomas
Farm, and M. gunteri by Simpson (1930) from Midway. White
considered these four species to represent a phyletic series in the
following manner: P. hlackhcrgi ))))) > P. harhouri )))» > P.
leonensis );» > 31. gunteri. This evolutionary sequence, spanning
the morphological range between a very primitive Parahippus
and a very primitive Merychippus, in his view presumably oc-
curred in situ in the Thomas Farm area.

1 A first niedlau phalanx has been found in the cla.v below the boulder bar on
the north face of the quarry since this was written.

Tablk

1.

Parahipjjm hJaclherc/i

Permanent

uppers

23

lowers

23

Total

46

p.

hai

-houri,

P. Iconensis, Mer

■ychippih<i

ffimteri

uppers

61

lowers

82

Total

143

Grand Total

189

BADER : EQUIDAE OF THOMAS F^ARM 51

ANALYSIS

A tabulation ui' the total imnibcf of associated dentitious meas-
ured is given in Table I. The total sample is seen to include over
200 specimens, many of which were, of course, incomplete.
Numerous isolated teeth also occur in the collections; however,

Deciduous
2
1
3

6

9

15

18

Table 1. A tabulation of the total number of associated dentitions measured
in the study.

since P''and M- often could not be distinguished with certainty
from P ' and ]\I '. res{)ectively, the.y were not included in the
statistics given beloAV. Length, width, and height measurements
were taken on all associated cheek teeth. Certain measures or
counts (e.g., dimensions of the protoeone, metaconid-metasty-
lid and crochet and number of plications on the metaloph) were
attempted and then abandoned. It was felt that statistics of such
measures (and estimates of parameters derived therefrom) would
add little to the classical procedures of description due to the in-
consistencies introduced by the variation of these dimensions with
wear, the nonhomologous nature of certain measures in different
wear stages (e.g., width of protoeone), and the inability to repeat
measures within satisfactory limits. All but the measuring error
could be eliminated by treating only specimens exhibiting little
or no wear. Such an analysis, however, would be limited to only
a small percentage of the total and would thus largely nullify the
most attractive feature, from an evolutionary point of view,
afforded by the Thomas Farm material, i.e., the opportunity to
study evolving populations.

The use of tooth length as a valid measure perhaps requires
some defense since it is known to be influenced by wear and for

52 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

that reason it has not been regarded as reliable, at least by some
students of the Bquidae, in the sort of detailed comparison that
is attempted here. Correlations and jjartial correlations of
length, width, and height were calculated for various members
of the dental series in both the larger and smaller population.
The r values ranged from near-]- .7 to almost 0, many of which
were not significant. Part of the correlation of height and length
must be attributable to the heritability of the dimensions in-
volved, i.e., in unworn teeth, height and length would be expected
to show a positive correlation since they are probably different
phenot^'pie expressions of the same general genetic complex (in
addition to which there may be factors for each trait). Width,
not affected by wear as measured on these specimens, is a third
manifestation of the same complex. Partial correlations of length
and height (width held constant) almost invariably exhibited a
lower value than the simple correlations between length and
height. In other words, some of the correlation between height
and length is caused \)y factors also common to width. These must
be factors affecting general dental size (or perhaps gross size?)
and cannot be attributed to tlie effects of age (wear). The con-
vergence in many of the teeth is most striking a few millimeters
above the enamel base, a point not reached by wear in almost all
of the species studied. It may be assumed, from such evidence,
that dental length may be used, at least in these forms, with
relative impunity. Data obtained from coronal length follow
essentially the same pattern as that of the widths and is thus
susceptible to the same inter]-)retation.

The measurements and their symbols are as follows :

Length of total cheek series (TCS) — the total length of the

cheek series taken at the grinding surface.
Length of the cheek tooth (LP-, etc.) — the maximum length
of the ectoloph (or ectolophid) measured at the grinding
surface.
Width of cheek tooth (WP-, etc.) — Uppers: the maximum
distance from the mesostyle to tlie lingual border of the
protocone measured at the enamel base. Lowers : the dis-
tance from the protoconid to a point essentially midway
between the paralophid and the metaconid, measured at
the base and normal to the long axis of the tooth.

BADER : EQUIDAE OF THOMAS FARM

53

20_

15 _

I0_

LP

<7>

<T

<7)

«•

<r>

^

0)

■t

<J>

«•

en

•»

0)

«r

0)

*

(7>

<t

<T>

O

O

z

-

pj

CJ

to

lO

f

fl-

m

m

<c

u>

r-

f^

CO

20_

15 .

WP

I0_

5_

en <n

o

CJ

20.

15 _

LP,

I0_

5 _

en

<7>

O

d —

<n

to

en

FiGUEE 1

54

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

20_

15 _

LM'

I0_

5-

6

en ^ en
6 - -

CO

en

(M

en o)
to to

^

'T

If)

en

^0 _

15 _

LM,

10 _

5 _

nr

20_

a>

<<■

en

<«•

en

^

en

T

en

*

en

f

<T>

^

0)

t

<j>

t

CO

en

(D

o

o

,

I

CVJ

CVJ

to

to

^

t

lO

in

u>

(0

N

15 _

I0_

5 _

LP,

T L

CD

^

en

^

en

'I-

<T> rt <n 't

en

*

en

*

en

*

en

CD

CD

en

o

o

-

— CVJ ^j fO
Figure 2

to

<r

*

in

m

u>

u>

BADER : EQUIDAE OF THOMAS FARM

55

20-

■

W

M,

15-

10-

5_

n

_,

CO

<r>
00

o

o>
o

*

o>

1;

0>

20-

I5_

I0_

5-

LM^

00

0>

<J>

i~n

d

o) «r o>
d - -

M

cvi

in

20-

I5_

WM'

I0_

J — r

«^

0)

*

01

*

o>

*

0)

f o ^ a>

*

a>

t

<»

*

0)

*

0)

c

0)

<\J

CJ

(2

(O

1-

t

m m <0 <c

Figure 3

c-

r-

oo

oo

o>

<T)

o

o

CO

CM

C\J

Figures 13. Frequency distributions of typical dental dimensions for the
entire sample, including Parahippus hlacTcbergi, Parahippus barhouri,
Parahippus leonensis, and Merychippus gunteri. Class midpoints are
plotted on the abscissae, frequencies on the ordinates.

56 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Height of the cheek tooth (HP-, etc.) — Uppers: from the
tip of the mesostyle to the base of the enamel. Lowers :
from the tip of the entocouid to the enamel base.

Frequency distributions were plotted for the length and for
the width of both the upper and lower cheek teeth for the com-
bined sample, a total of twenty-four distributions. Nine of these
have been selected as typical and are presented in Figures 1 to 3.
These distributions demonstrate fundamentally the same pattern
irrespective of the dimension or tooth selected, i.e., there is a
larger, more numerous population distinctly separable from a
population smaller with respect to both dimensions and number
of individuals. Only minor deviations from a normal distribution
are in evidence, with little or no suggestion of bi- or trimodality
in the larger group. The approximation to a normal curve is
much closer for the larger assemblage, as would be expected from
the relative sample sizes. An upper dentition (MCZ 3742) falls
at about the 'Sa limit for several characters and is usually almost
a milliinetei- smaller than the next larger individual (Figs. 1
to 3). This specimen was referred to P. harhouri by White.
Frequency distributions for two postcranial elements are giA-en
in Figure 4. The sample size is very large in both, numbering
338 for the caleanea and 552 for the phalanges. The same basic
pattern as that of the dental elements is discernible here. The
extremes of the.se variates in the two populations overlap slightly
as seen from the grouped distributions ; hoAvever, the ungrouped
data show a hiatus of about 1 mm. in the caleanea distribution
and of 0.3 mm. in that of the phalanges. The observed ranges,
then, do not overlap, although the standard ranges do to a limited
extent. Limited samples of several other postcranial elements
were also available. Lisofar as conclusions can be drawn from
these small samples (N<30), they appear to demonstrate basi-
cally the same features as those of the more numerous elements.

Scatter diagrams for several of the pairs of variates Avere
plotted and three, selected as representative, are presented in
Figure 5. On the basis of these ratios, the group is clearlj- sepa-
rable into only tAA-o populations. The slope and position of re-
gression lines is nearly the same in the tAvo populations.

Sample size (N), mean (x), obserA^ed range (O.K.), standard
range (S. R. [S. D.]), standard deviation (a), and coefficient
of A'ariation (V) for the dental dimensions of the tAvo pupu^^ations

BADER : EQUIDAE OF THOMAS FARM

57

are given in Tables 2 and 3. There is but slight overlap in the
standard ranges for a few traits and none in the observed ranges.
A most pertinent point relative to the present discussion is the
magnitude of the variation in the respective populations. That
the smaller population represents a single species is the unani-
mous opinion of all who have studied the group, a judgment with
which the present findings are in agreement. The variation in
this "good" species represents, therefore, the best available
standard of comparison for the larger grouj). Although the
material referred to P. leonensis alone, has been characterized
as unusually variable, the variation in the odontometric traits
of the total larger population from Thomas Farm (P. harbouri,

60-

50_

40-

30,

20.

CALCANEA

. — n

ro_

60-
SO-
40_
30_
20-
I0_

FIRST MEDIAN PHALANX (MANUS)

tbz.

— wojcsjWfticsjevj

Figure 4. Frequency distributions of two posteranial elements for the entire
sample. Class midpoints are plotted on the abscissae, frequencies on
the ordinates.

58 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

II

10-

9_

O

7

6.

FIG. 5A

Pa

_i I I I I I I I I I I

10 II 12 13 14 IS 16 17 . 18 19 20

LENGTH
Figure SA

IS
17

:

FIG.

5B

P'

•

•

•

• •

• • •

« •

•

16

-

•

*

• •

•
•
•

IS

.

•

I
1-
O 14

UJ

_l

-

•

13

-

12

r

•

•
•
•

• •
• •

1 1

•

• •

1

1

1 1

I

1 1 1

'

13 14 15 16 17 18 19 20 21 22 23

WIDTH

Figure 5B

BADER : EQIHDAE OF THOMAS FARM 59

P. leonensis and M. guntcri) is quite comparable to that of P.
hlackheryi. The mean V for tlie maxillary teeth is 5.06 in /'.
hlackhcrgi compared to 4.98 in tlie larger population. The figures
for the mandibular teeth are 5.64 and 5.88 respectively. The V's
in each group are at or near the mode for homologous traits in
both fossil and Recent maunnalian species. Patently, on the basis
of quantitative considerations alone, the total sample cannot be
divided into more than two populations.

The maximum unworn height of the M- in P. hlackhergi is
aI)out 10 mm. and approximately 16 mm. in the larger group.
Seasonal deposition of the remains of animals which reproduce
seasonally should result in a multi-modal distribution of a charac-
ter which is highly correlated with age, e.g., the crown height of
teeth. Matthew (1924) divided a large sample of Merychippus

17

- FIG.

50

M^^

•

• •
• • • • •• •

• • •

16

15

. • • •

• •
• • • •

• • •

• * •
•

•

14

.

X

ZI3

_l

•

•

12
1 1

•
•

•

• • •

•
*

•

•

10

"

-J 1,

' '

1

L 1111

12 13 14 IS 16 17 18 19 20 21 22

WIDTH

Figure 5C

Figure 5A, B, C. Scatter diagrams of coroTial length and width for selected
teeth. The total sample is included.

60 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

primus from the Snake Creek horizon into several distinct age
groups based on eruption and wear of the molars. Kurten
(1953) found a striking multi-modality in the distribution of
molar height in several artiodaetyls from the Pontian fauna of

Table 2

Varlate

N

X

O.R

•

S.R.

(S.D.)

(•

V

TCS

8

62.60 ♦ .97

58.6 -

66.2

53.9

- 71.7

2.71*1

.69

1*.1*9 1 1.09

1^2

15

12.59 1 .lii

11.6 -

13.U

10.8

- 2h.3

.51*1

.10

l*.33l .78

LP3

17

11.1»1* + .10

10.5 -

12.1

10.1

- 12.8

.U3 1

.07

3.83 1 .65

IPU

11*

11.33 1 .13

10.I4 -

12.0

9.8

- 12.9

.1*7 +

.09

U.25 1 .78

Ull

16

11.61 + .13

10.7 -

12.3

9.9

- 13.3

.53 i

.09

U.63 1 .81

I«2

17

10.87 + .10

10.3 -

11.8

9.5

- 12.2

.Ul ♦

.07

3.83 1 .65

IM^

13

9.19 + .22

7.3 -

10.2

6.6

- n.8

.79 1

.15

8.72 1 1.68

WP2

15

12.2li + .16

11.0 -

13.0

10.2

- Ua.3

.62 +

.11

5.161 .93

Wp3

15

13.95 + .19

12.3 -

2h.9

11.6

- 16.3

.73 1

.13

5.29+ .95

wp'*

m

Uj.65 + .20

13.0 -

15.7

120

- 17.1

.71* 1

.11*

5.11 1 .95

WM^

13

Ui.Ol + .15

13.1 -

lit.8

12.3

- 15.7

.521

.10

3.83 1 .73

WM^

18

13.l4l< 1 .17

12.1 -

lli.6

11.1

- 15.7

.711

.12

5.32 1 .88

vim3

13

10.99 1 .a

9.0 -

11.8

8.5

- 13.5

.761

.15

7.01* 1 1.36

TCS

9

6U.68 +1.02

60.5 -

68.6

5U.8

- 7U.6

3.05 1

.72

1*.85 1 l.U

1^2

13

U.Oli + .16

10.3 -

11.8

9.2

- 12.9

.57 1

.11

5.28 1 1.01

""3

17

10.61i + .15

9.6 -

ll.U

8.6

- 12.7

.62 1

.u

5.91 1 1.00

^h

21

10. 5U + .13

9.8 -

11.8

8.3

- 12.8

.60 +

.09

5.77 1 .88

%

20

10.13 + .13

9.2 -

10.9

7.9

- 12.U

.60 1

.10

6.031 -91*

Ul2

16

9.99 + .13

9.2 -

U.l

8.3

- U.7

.521

.09

5.25 1 .91

1^3

12

10.80 ♦ .10

10.3 -

11.6

9.7

- 11.9

.31*1

.07

3.20 1 .6h

VJPg

13

5.73 1 .13

5.1 -

6.U

U.2

- 7.2

.1*61

.09

8.27 1 1.58

WP3

17

7.39 i .11

6.7 -

8.5

5.9

- 8.9

.U7 1

.08

6.1*7 1 1.09

WP||

a

8.37 1 .10

7.7 -

9.U

6.9

- 9.9

.U61

.07

5.50 1 .81*

\m^

21

7.96 + .09

7.1 -

8.9

6.6

- 9.3

.1*11

.06

5.15 1 .79

WI2

17

7.57 i .10

6.9 -

8.3

6.2

- 8.9

.1*2 1

.07

5.61 1 .95

WM3

13

6.50 ♦ .U

5.9 -

7.1

5.3

- 7.7

.381

.07

5.97 1 i.:il*

Table 2. Dental statistics of the smaller population, Parahippus blackbergi.
Symbols explained in the text.

BADER: EQUIDAE OF THOMAS FARM 61

China. It might be hsshhuh] that a simihir (listributiou could be
recoti'uized in the Thomas Farm material since deposition is defi-
nitely of a seasonal nature now and probably was in the middle
and late Tertiary. A study of both the limited samples of the

Tahle 3

Variate

X

0.

R.

S.R. (£

;.D.)

a—

V

TCS

21

92.35 1

.81

82.3

- 98.U

80.3 -

loU.U

3.73 i .58

U.09 ♦ .62

LP2

33

19.56 +

.17

16.8

- 21.1

16.1, -

22.7

.98 + .12

5.01* + .62

U>3

U2

16.58 +

.12

U4.IJ

- 18.0

lU.l -

19.1

.78 + .09

1*.7U 1 .51

uh

U6

16.65 1

.11

U;.6

- 18.1

IU.3 -

19.0

.73 1 .08

li.l*l + .U6

nf

h2

15.52+

.lij

12.9

- 16.9

12.7 -

18.U

.88 ♦ .10

5.72 ♦ .62

Ii-l2

U9

15.58 1

.U

13.5

- 16.8

13.2 -

18.0

.7li + .08

1.79 i .U8

I^l3

U6

lli.h9 i

.09

13.3

- 15.7

12.5 -

16.5

.61 + .06

U.2U i .lil*

Vjp2

3U

18.37 1

.20

lh.9

- 20.5

IU.6 -

22.2

1.18 + .11*

6.1*6 + .78

wp3

38

20.21* +

.20

16.8

- 22.5

16.3 -

21*.2

1.21 + .11;

6.02 + .69

^^

U3

20.60 +

.16

17.3

- 22.6

16.8 -

2li.U

1.18 + .13

5.76 + .62

VJi?-

U2

19.25 1

.13

16.9

- 20.8

16.5 -

22.0

.81* + .09

u.ia 1 .U8

vm2

U9

19.53 1

.13

17.0

- 21.2

16.6 -

22.5

.90 + .09

I1.62 + .1*6

m^

U3

18.01 +

.12

16.2

- 19.9

15.U -

20.6

.80 + .09

Ii.l*7 + .1*8

TCS

18

92.16 +

,99

82.2

- 97.7

78.5 -

105.8

l*.22 + .70

U.6!* + .76

1^2

30

16.16 +

.20

13.5

- 18.7

12.6 -

19.7

1.09 + .11*

6.70 + .87

IP3

52

15.12 +

.U

13.5

- 16.7

12.6 -

17.7

.78 + .08

5.15 1 .50

^U

61

15.00 +

.09

13.U

- 16.7

12.7 -

17.3

.72 + .06

1*.79 + .1*3

1%

60

li;.39 +

.12

12.2

- 15.9

11.3 -

17. U

.91* 1 .09

6.58 ♦ .60

ut.

60

li;.75l

.11*

12.5

- 17.2

11.1 -

18. U

1.12 + .10

7.62 + .69

^2

U8

I8.1;5i

.12

16.7

- 20.9

15.8 -

21.1

.81 ♦ .08

U.UO + .U5

WP2

31

9,06 +

.13

7.1

- 10.3

6.7 -

1J..5

.7U + .09

8.27 +1.01*

W3

57

12.09 +

.09

10.3

- 13.3

9.8 -

Ui.li

.70 * .07

5.79 1 .5U

^u

59

12.78 +

.10

10.8

- 11*. 3

10.3 -

15.3

.78 + .07

6.15 + .56

WMi

61

U.81 +

.09

10.2

- 13.2

9.6 -

m.o

.67 + .06

5.69 + .51

m^

58

U.13 +

.08

9.7

- 12.1

9.2 -

13.1

.60 + .06

5.U0 + .50

TOI3

U3

9.77 ♦

.08

8.5

- 10.7

8.1 -

u.l,

.51 1 .05

5.2U 1 .56

TnhlcS. Dental statistics of the total progressive population {Paialiip})iis
barlxviri, rarahippus h-onensis, and Merychippus r/nnteri of White).

62 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

younger age group and the total frequency distributions of crown
height fails, however, to bring this expectation to realization.
The distinctness of the age groups may have been obliterated due
to an increase in mean height of the crown during the interval
of deposition at Thomas Farm.

Table 4

O.R.

.78 - .79

.8U - 1.02

1.00 - 1.07

1.03 - 1.3U

.9U - 1.03

1.05 - 1.39

.li6 - .50

.52 - .61

.68 - .73

.70 - .89

.6U - .67

.73 - .98

Table 4. A comparison of the width/length ratios of the deciduous and
permanent premolars in the total progressive population.

A small sample of milk molars was available in the larger
population. The width/length ratio is more characteristic of the
deciduous set than the linear dimensions and is compared with
that of the permanent dentition in Table 4. This ratio appears
to be of value in the assignment of isolated teeth to the proper
set. The means are distinctive with but modest overlap in the
standard ranges. As is common in horses, the milk molars are
both longer and narrower than their successors. The variation
in the ratio is notably greater in the permanent series.

Although this investigation is concerned chiefly with measur-
able traits, valid species, may, of course, exhibit no significant
differences in such characters and yet be separable on non-
metrical or qualitative grounds. Careful attention, therefore,
was directed to the several characters which have been con-
sidered diagnostic of the four species. These include principallj' :

N

1

r

dp2

3

.78

e2

32

.97 +

.007

dp3

5

1.03 1

.011

p3

37

1.22 +

.010

dP^

3

.99

P^

U3

1.2li +

.011

dPg

5

.U8 +

.007

^2

28

.56 +

.005

dP3

6

.70 +

.008

^3

U7

.80 +

.006

"^U

7

.66 +

.005

\

5U

.85 +

.006

S.R.

(S.D.)

0 —

.81

- 1.07

.OliO ♦

.005

.95

- 1.11

.02U +

.008

1.02

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.061 ♦

.007

1.01

- I.li7

.071 +

.008

.U3

- .53

.015 *

.005

.1*7

- .65

.029 ♦

.oou

.6U

- .76

.020 +

.006

.68

- .92

.038 ♦

.OOlt

.62

- .70

.012 +

.003

.71

- .99

.OUh±

.OOli

BADER : EQUIDAE OF THOMAS FARM 63

amount of cement, degree of hypsodonty, prominence of cingnla,
distinctness of metaconid and metastylid, development of plica-
tions (largely from the metaloph) and the degree of nnion of
the various lophs. Detailed descriptions and figures of the pro-
posed species were presented by Simpson (11);}2) and White
(1942) and need not be repeated here. White empliasized the
considerable variation and intergradation in these characters,
which make for difficulty in species identification. Additional
collecting has served to demonstrate interg:radation at even more
subtle levels, though the range of morphological expression re-
mains about as previously known. The manifestation of the
crochet in the total sam])le varies from essentially a Miohippus
to a Merychippus stage, i.e., from complete absence to full union
with the protoconule at an early stage of wear. Cement is absent
or scantily present in the more primitive species while the larger
group most often has a moderately heavy coat, though the varia-
tion is extensive. In the more progressive forms the metaloph
plications may be several or almost totally wanting. The dis-
tinctness of the metaconid and metastylid, though varying iu
these forms, offers no opportunity for taxonomic separation. In
short, detailed study of these and other traits has revealed no
valid basis for distinguishing more than two populations within
the total assemblage. In general, within the larger population,
the linear dimensional traits and the degree of hypsodonty tend
to be positively correlated with the amount of cement, complexity
of the plications and the extensiveness of crest development,
though the association is by no means complete.

DISCUSSION

The results of this detailed investigation substantiate the more
superficial impression gained from field work, i.e., that specimens
are assignable without equivocation to a larger or smaller popu-
lation, but only with great difficulty (and, then, with little
enough assurance) allocated to one of the described species of
the larger group. P. barhouri was separated from P. leonensis
primarily because of its smaller size and modest crochet develop-
ment. With one exception, the specimens referred to the former
are clearly small members of the latter, as evidenced by the
want of an appropriate mode in the frequency distribution, the
similarity in the width/length ratios as seen in the scatter dia-

64 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

grams, the failure of the incliisiou of these specimens to do
violence to the ma»'nitude of the V of P. leonensis, and the almost
perfect intergradation demonstrated by crochet development.
One specimen (M.C.Z. 3742), alluded to in the previous section,
is, on strictly statistical grounds, very improbably a member of
the larger population. It cannot be absolutely excluded on this
basis, however, and since there was probably some evolution
toward larger size in the population, its separation would be
even more unjustified.

The difficulty attendant upon the consistent separation of P.
leonensis and i¥. (junteri was recognized by both Simpson and
White. White (1942, p. 28) stated, "The transition between the
two species is well represented by complete dentitions. For-
tunately it is still possible to set up an arbitrary rule which
will distinguish the conservative members of this species [M.
gunteri] from the progressive members of the preceding one.
This rule is: — The crochet nuist have joined the protoconule
on M^"- by the time wear has exposed the principal cusps
on M'^ before the specimen can be placed in M. (junteri. At
this time both species have a height of tooth at the mesostyle
on M- of 10 mm. When the teeth of the progressive specimens
of P. leonensis are worn so that M- has a height of 7.5 mm. at
the mesostyle they display most of the characters of M. gunteri.'^
This criterion is, admittedly, completely arbitrary and leaves
many of the specimens (in the middle and later wear stages)
indeterminal)le. It is much to be preferred to group these pro-
gressive forms witii the other larger specimens for much the same
reasons as indicated for P. barbouri (only a very small percentage
have been found which satisfy White's rule). Again, no evidence
of segregation within the lineage can be discerned.

It happens that individuals which are here referred to one spe-
cies Avere placed by White not only in different species and genera
but in different subfamilies (employing the arrangement of Simp-
son, 1945). The diiferenee in the two viewpoints is, of course,
not nearly so great as would be inferred from the alternative
taxonomic schemes; i-ather, it is largely a reflection of the ex-
cellent documentation of horse evolution. Now, it is clear that a
few of these specimens so assigned to /'. leonensis exhibit Mery-
rhippus qualities and would be properly referred to that genus
if found in isolation though the modal expression of the popula-

BADER: EQUIDAE OF THOMAS FARM 65

tion to wliieli they belong' remains j)arahippine in nature. In
making taxonomic decisions the total situation must, of course, be
evaluated. A specific sej^aration of these specimens would imply
that two populations (between which interbreeding was absent
or minimal) eoexisted at the same time and in the same general
area, an hyi)othesis for which thiM-e is no adequate justification
on the basis of the known evidence. Traits uniformly present in
a species do not arise de novo but spread gradually throughout
the population until they become fixed. A contemporaneous inter-
breeding assemblage, only a part of which possesses a given
character or whose members vary in the degree of character
expression, should not be split into different species simply
because the character in question is highly diagnostic of closely
related taxa. The development of the crochet is an excellent
case in point. The overwhelming majority of equid genera are
characterized either by complete absence of this crest or by its
full union with the protoconule, the primary exceptions including
a fcAV species of Miohippus and Arckaeohippus and the species
of Parahippus. The elapsed time from its inception to ultimate
expression must be in the order of 5 million years as a minimum.
Thus, while it is of taxonomic importance before and after this
development period, its diagnostic value during the transition
is greatly reduced (although not altogether eliminated) and
its arbitrary use completely unjustified. It cannot be seriously
doubted that in the later stages of the time represented by the
deposits at Thomas Farm, some members of an essentially con-
temporaneous population exhibited the typical ParaJiippus mode
of crochet development while others possessed the Merychippus
type, a not unexpected situation if the two did, indeed, have an
ancestral-descendant relationship. Simpson (1944) has discussed
certain theoretical implications of the incorporation of this
"mutant" in equid populations.

A similar example concerns the smaller population, P. bluck-
bcrgi. White (1942) assigned a few isolated upper teeth to Mio-
hippus sp., primarily on the basis of the incomplete union of tlie
ectoloph and metaloph. Since the dimensions of these teeth are
close to the means of their homologues in the P. hlackbergi sample,
it is most proljable that they are deviants of the same population,
though failure of ectoloph-metaloph union, if taken at face value,
would certainlv exclude them from that genus. There remains the

66 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

possibility that such deviations may occasionally be due to non-
heritable factors, producing essentially a phenocopy. This "lump-
ing" procedure facilitates the study of the dynamics of eyolying'
populations while detracting in no wise from the biological mean-
ingfulness of a group so lung as the deviants are fully discussed in
the species descriptions. The inferential determination of popula-
tional characteristics, not those of individuals as such, is the
proper object of taxonomic inquiry, but this is a concept too
often ignored even in modern paleontological work (see Simpson
1943). It offers, however, the best hope of bringing some order
out of the chaos in which taxonomic paleontology ( at least in
many areas) finds itself today.

There remains one other taxonomic possibility to consider, i.e.,
the use of subspecific names for the diff'erent phases of evolu-
tion in the larger group. This has been rejected due to the lack
of detailed stratigrai)hic records and independent criteria with
which the names might be associated, e.g., different formations
or members. The dilemma posed by the vertical species (or sub-
species) concept is well known. The differences in the relation-
ships of contemporaneous as opposed to vertical popidations are
several and profound. From a strictly taxonomic viewpoint, in-
creased knowledge enhances the understanding of the former
while tending to reduce the latter to the absurd. Contrariwise,
the untenable consequences of continued extension of vertical
unit limits has been pointed out by Simpson (1932) and others.
Fortunately for the jn-esent Liunean procedures in paleontology,
a continuous record of the type seen at Thomas P^arm is usually
of limited vertical extent. P. Iconensis (present usage) probably
exhibits a somewhat greater range of variability in non-dimen-
sional traits than any species of the genus heretofore defined. A
revision of the genus would, however, almost certainly result in
the placing of many names in synonymy, effecting an increase
in the variation of the valid species. As indicated in the previous
section, the dimensional variation is quite within the usual range.

The major outlines of the relationships of these Thomas Farm
horses emerge with considerable clarity. That there were, at any
one time, but two distinct, non-interbreeding populations seems
reasonably certain. These two populations, though manifestly
closel}^ related, present little or no evidence of intergradation.
If the smaller species is, indeed, the progenitor of the larger

BADER: EQUIDAE OF THOMAS FARM 67

(which is probable), the evidence must be sought in the deeper
levels of the Thomas Farm ''dig" or elsewhere in deposits of
somewhat earlier age. It may be assumed that some evolutionary
change occurred in both populations in situ, and to a greater
degree in the larger population. Although detailed stratigraphic
records were not kept on the earlier material collected from
the site, it was noted by White (1942) that several ''species/'
including Merychippus gunteri (in his sense) were found only
in the initial phases of the excavation. Teeth falling into this
category, according to White's rule, have more recently been
found at deeper horizons, but are extremely rare. The dimen-
sional means of all of these more advanced teeth are as a rule
greater than those of the remainder of the population, though
the difference is not statistically significant. Dimensions of teeth
and postcranial elements taken in the past two years from the
deepest portion of the pit (now about 20 feet below the surface)
are generally below their respective means for the larger popula-
tion as a whole. Simpson (1930) described M. gunteri from
Midway, also referring several specimens from Quincy to it. The
fauna from these localities is much less perfectly known than that
of Thomas Farm, practically nothing having been added to our
knowledge since Simpson's report. Though intergradation with
the more advanced specimens of Thomas Farm is apparent, the
typical development of these Midway and Quincy horses is
somewhat more advanced, exhibiting essentially merychippine
characters. That this group is derived from the larger popula-
tion at Thomas Farm, as Simpson tentatively suggested, is now
established beyond any reasonable doul^t. If the interpretation
presented here is correct, the absence of "P. harhouri" and
"P. leonensis" fi-om the Midway-Quincy deposits is satisfacto-
rily explained. The occurrence of P. hlackhergi throughout the
Thomas Farm deposit and probably at Midway also (Simpson,
1932) demonstrates its coexistence, in a relatively unchanged
state, with its larger, more progressive descendant. It has been
found in the later Miocene of the AVest although not reported
above the Garvin Clulley horizon in Texas. These interpretations
are diagrammatically presented in Figure 6.

In these Miocene equids, factors controlling such characters
as quantity of cement, hypsodonty, and complexity of the enamel
pattern appear to be moving through the population at a fairly

68

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

rapid rate, undoubtedly under selective control, as the grazing
niche of the plains is more fully exploited. Simpson (1944)°
suggested that the shift from a browsing to a grazing adaptive
peak partook of characteristics Mhich he termed quantum evolu-
tion. If such an evolutionary mode existed (its most convincing

MIDWAY-OUINCY

THOMAS FARM

1
1
1

1

BLACKBERGI

1

1
1

1

LEONENSIS 1 GUNTERI

PARAHIPPUS

MERYCHIPPUS

Figure 6

Figure (3. A sclu'inatic presentation of the relationships of the horses of
Thomas Farm and Midway-Quincy, late lowei' and early middle Miocene
respectively (see text).

support has come and, of necessity, must cuntinue to come, from
negative evidence), the larger population at Thomas Farm is in
the post-quantum phase, moving fairly rapidly up the slope of
the grazing adaptive peak. The smaller population remains
under the influence of the selective forces for browsing. The
transition between them would entail passage through the in-
adaptive ''valley." It should be extremely interesting to see
what sort of intergradation between tlie two, if any, is revealed
by continued excavation at the ''dig."

The relationships of selection, A^ariation, population size, and
rate of change are quite complex. The horse sequence at this
site oifers a rather limited opportunity for evaluation of these
interrelationships due to the restricted time interval involved.
Certain features may, however, be briefly considered. It may be

[iADER : EQUIDAE OF THOMAS FARM 69

inferred that there is a strong linear component to the selection
on the larger group while that on the smaller population was
primarily centripetal in nature. The coefficients of variation
given in Tables 2 and 8, closely similar in the two species, have
a vertical as well as a horizontal component. It may be assumed
with some confidence that the variation exhibited by contempo-
raneous populations was lower in P. honensis, since it was chang-
ing at a more rapid rate. Tf so. the relationship of quantitative
variation and evolutionary rate would appear to be similar to
that shown in the oreodonts over a longer temporal interval
(Bader, 1955). In the Thomas Farm horses, however, population
size is positively correlated with rate of change, the opposite
being true in the oreodonts.^

The proper Linnean designation of the two populations pre-
.sents a problem. The smaller population was described as Archaeo-
hippiis )wnus by Simpson (1982) and was referred to Parahip-
piis [Miohippus] hJackbergi Hay 1924 by White. Adequate
comparative material has not been available during this investi-
gation to make a meaningful choice between these tAvo genera,
the relationships of which are far from clear. However, if
Archaeohippus represents a natural assemblage, the smaller
species should be assigned to it. The valid application of the
specific name nanus as opposed to hlacJihcrgi hinges on the ques-
tion of conspecificity between the Florida and Texas forms.
Quinn (1955 and personal communication) found little overlap in
length-width measurements of small samj)les from the two areas.
It is hoped that more extensive comparisons can be made in the
near future. Pending further studies, I prefer to follow White.
The larger population was included in Parahippus by Sellards,
Simpson and White; however, Quinn (1955), in the most exten-
sive revision of the post-Oligocene equids of the past several
years, placed the previously described species of this population
in Hippodon. a genus redefined by him after its description by
Leidy in 1854 and subsequent rejection. Parahippus, a genus
originally described from milk molars, has not been rigorously
defined and may (along with Menjchippus) represent a struc-
tural stage attained independently by several phyla, as Quinn

1 It is assumed, in tlie absence of evidence to tlie contrary, tiiat the sample
sizes of the several species offer a reliable indication of the relatixc abnnflain-c
of the populations in nature.

70 BFLLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

infers. However, the distinction between Hippodon and Para-
hip pus remains nebulous and it is to be doubted whether the
resurrection of the former will effect a greater understanding
of the relationships of early and middle Miocene Equidae if it
is merely to be substituted for the latter.^ The problem of the
correct designation of the Thomas Farm species can be solved
only through a study far outside the confines of the present
investigation. A revision of the entire assemblage of Miocene
Equidae is clearly in order. Until such revision (a truly monu-
mental task) is undertaken, it is preferred to tentatively retain
the names used by White, P. hlackhergi and P. leoncnsis (the
oldest available name for the larger species ).-

A sequence very similar to that in Florida is seen in the Texas
Gulf Coastal Plain, species conspecific (or nearly so) with the
Florida Miocene equids appearing in progressively later strata.
Several possible relationships of the faunas of the two areas
suggest themselves : both areas could have been populated by
successive immigrations from the north ; Florida may have been
the principal site of evolution with repeated emigrations to
Texas (or vice versa) ; the two populations could have evolved
in parallel fashion from a common ancestry ; or one continuous
progressive population may have occupied the Gulf Coastal
region and evolved essentially as a unit to the Merijchippus stage,
with the continued survival of the primitive /*. blachbergi, the
progenitor of the sequence. The last hypothesis is the most
tenable. The intergradation, particularly in Florida, is too com-

1 This is not tlic iilaee to attciiipt a full review nl' Uuinn's iiiterestinjr study,
thouffh it iiia.v Ix' noted that in some ways his apinoat-h is almost diametrically
opposed to that used heic : witness, tor ex;inuile, the reeoj;nition ot seventeen
contemporaneous horse speeies (and el(>ven genera) in the latest tipper Miocene
(LaPara Creek) from a limited area of the Texas <iulf Coastal Plain. Quinn cor-
rectly points out that ai>pIicatiou of the Gause principle must he made with cau-
tion wdien dealin.t: with a thanatocoenose rather than a l>iucoenose. This does not
imply, however, that the principle may he completely ignored or that it may not
judiciously l)e applied at an order of magnitude level, particularly when dealing'
with a restricted time interval. A situation even remotely approaching that de-
I)icted for these Texas forms is an extreme rarity among the larger modern mam-
mals. On the other hand. Quinn has properly stressed the importance of the
lower teeth, heretofore largely ignored in classificatory studies.

- Mr. Bryan Patterson has pointed out that the smaller species, when compared
with the larger form, has longer lateral metapodials relative to the median
metapodials, more slender proximal phalanges and s somewhat different incidence
of the various ligamentary scars. These pedal disparities, in addition to the differ-
ences in the dentitions, are cogent reasons for placing the forms in different
genera. However, the prohleni of the correct generic assignation remains, as
outlined above.

BADER : EQUIDAE OF THOMAS FARM 71

plete to invoke an immioration hypothesis. Parallel evolution is
possible, thoiig:h rendered improbable by the great similarity of
the two series. Whether there oeeurred but one progressive
population at any one time is, of course, difficult to demonstrate.
The hiatus in material from the area between Texas and Florida,
is complete and likely to remain so. A more intensive comparison
of the terminal members of the distribution (Texas and Florida)
should aid in the interpretation. Quinn's suggestion (1955) that
the Garvin GuUey is slightly older than the Thomas Farm was
based on the relationship of the equids and certain other genera,
such as Synthetoceras. This has not been definitely established
but may be true and is tentatively accepted here pending more
detailed comparisons. The relation of the two sections and their
included horse faunas as now known is presented in Table 5
(based on Simpson, 1930, 1932 ; White, 1942 ; Stenzel, Turner and
Hesse. 1944; Quinn, 1955; and the present study. The horse
terminology is my own).

GEOLOGICAL PROBLEMS

Several problems of a geological nature have been posed by
the ' ' dig, ' ' most of which are but incompletely solved. Simpson
(1932) suggested that the materials were deposited in a sink
hole, a common feature in this part of Florida; White (1942)
considered the lack of sorting of the material in the boulder bar
and the lenticular nature of the clays to be indicative of stream
action. Cooke (1945) supported the sink hole hypothesis, point-
ing out that the tremendous aecumidation of bone demands some
sort of catchment basin such as would be provided by a sink.
Although articulated skeletoiLs have not been found, a gentle
water action is indicated by the unworn state of most of the
material and the intact condition of some delicate structures
(e.g., the mandibular coronoid process of the small cervids and
the distal end of the horse ulna). Recently an articulated eal-
caneum and astragalus of a rhinoceros were found by S. J. Olsen
and a few other articulated elements are known. W. Auffenberg
(personal communication) ha^s pointed out the similarity of the
deposits at the "dig" and those in recent sinks. Intensive screen-
ing of material by Auffenberg has revealed but a single fish
spine, whereas such spines occur in some quantity in most of the

72

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

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BADER: EQUIDAE OF THOMAS FARM 73

undoubted stream deposits he has worked. E. C. IMrkle has
suggested the possibility that the l)()ulder bar may be the remnant
of a solution joint in the Oeala limestone, both of whieh trend
NW-SE. It may be doubted thai a stream on this low-lying land
mass would be rejuvenated to an extent suftieient to trans])ort the
large boulders of the boulder bar, as proposed by AVhite. Al-
though incontestable evidence is yet to be forthcoming, the facts
seem best accommodated bj^ the sink theory. The alternatives are
not necessarily mutually exclusive ; the sink may have been fed
b^' small streams, as are existing sinks in the area today. The
mode of tleposition has obvious biological implications. Since
the material accumulated in a sink is usually from a more circum-
scribed area than that in a stream bed, the number of niches
"sampled" would be smaller, and, in consequence, the number
of closely related species to be expected in the deposit would be
reduced.

White stated that Florida was an island at this time ; Cooke
(1945) suggested a narrow peninsula. Vernon (1951), while dis-
cussing other aspects of the deposit, did not choose between the
alternatives. The critical evidence lies in the Okefenokee trough
area to the north and appears, at ]iresent, to be inconclusive.
The great similarity of the Texas and Florida horses as well as
that of many other components of the two faunas indicates that
the isolation of Plorida must have been, at best, very ephemeral.
White pictures the horses trapped on the island, from which they
finally emigrated in the early middle Miocene, giving i-ise to the
later, more advanced species of Merychippus in the West.
That Meriji'hippHs has a polyphyletic origin has been advocated
by several authors (e.g., Simpson, 1932, and Stirton, 1940) ; if
this view is correct, the western development of the genus may
have been relatively independent of that in Florida.

The time interval represented at the Thomas Farm was placed
by White between the deposition of the Suwannee Limestone
(upper Oligocene) and the Hawthorne formation (middle Mio-
cene). Invertebrates derived from the Suwannee have been iden-
tified in the deposit ; that this is the lower limit seems adequately
established. Romer (1948), quoting Simpson (1932), indicated
that the site was post-Tampan since the echinoid, Cassidulus
(jouldii, found here, was confined in Florida to the Tampa Lime-
stone. The species is, however, characteristic of the Suwannee

74 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

as now recognized (Cooke and Mansfield, 1936). The surface sand
and a superficial sandstone layer were identified by White, on
strictl}' lithological grounds, as belonging to or derived from the
Hawthorne. It was, therefore, assumed that the encroachment
of this sea concluded the period of fresh water deposition. Ver-
non (1951), in a detailed study of Levy and Citrus counties
(adjoining Gilchrist County), concluded that neither the Tampa
nor Hawthorne seas covered the area in question, both being
limited to the flanks of the Ocala uplift. He failed to recognize
Hawthorne deposits Avithin about 25 miles of the "dig," evidently
rejecting AVhite's interpretation of the sands. The Thomas Fai'm
deposit Avas assigned by Vernon to the Alachua formation, the
terrestrial facies of the Tampa and Hawthorne formations, and
it was further suggested that the site included deposits sjmnning
the entire Miocene. If the Hawthorne is not present, the defini-
tive upper limit of the interval of deposition is removed. There
is no paleontological reason to believe, howevei-, that the fauna
extends beyond the early middle Miocene at the latest. The fauna
as a whole is best assigned to the late lower Miocene as emphasized
by both White (1942) and Romer (1948). The previous alloca-
tion to the lower Miocene caused some concern since Merychip-
piis first occurs in the Great I^lains area in the Hemingfordian.
The removal of MerycliippKS from the species list, while not alter-
ing the morphological situation, does lend greater emphasis to the
late lower Miocene age of the deposit.

DENTAL CORRELATION

The most compreheiLsive investigation of dental correlations
in mannnals is to be fovind in the interesting and important work
of Kurten (1953). The following salient facts were brought
out in Kurten 's study: the r value of the linear dimensions of
the various members of a dental series is negatively correlated
with the distance between them (the rule of neighborhood of
Karl Pearson) ; the correlation is higher between occluding teeth
than between adjacent teeth and is very similar in a given area
of the dentition indicating identity of the morphogenetic
"fields" in both jaws; the correlation between the terminal mem-
bers of a series is often greater than expected from the rule of
neighborhood, suggesting a recurvature of the field on itself ;

BADER : EQUIDAE OF THOMAS FARM 75

and the correlation between functional homologous right and
left teeth is very high, considerably lower where non-functional.
Coefficients of correlation (/■) for dental width were calculated
for both species and are presented in Table 6. Lengths were not
included since the etfect of wear, even though mo<:Iest, would
add still another variable to an already complex situation and
since the labor of calculating from incommensurate sample sizes
is considerable. The correlations of the lengths of the uppers in
P. hlackhcrgi were, however, calculated and found, in essence,

Table 6
(a)

N- =

= 9 to 14

N =

= 8 to 19

p3

p4

Ml

iM2

M"'

P.J

P4

Ml

M2

M:i

p-

.79

.87

.67 (

:.45)

(.51;

P2

.77

.85

(.47) (

:.52)

.79

p3

.9.5

.08

.72

(.52)

P.i

.91

.73

.08

.67

p4

.81

.71

(.48)

Pi

.79

.77

.82

Ml

.59

(.17)

Ml

.89

.OS

M2

.76

M2

.79

(b)

N =

25 to 38

X =

18 to 48

p.-!

P-t

Ml

M2

MS

Ps

P4

M,

M2

Ms

p2

.89

.93

.77

.71

.75

P2

.79

.81

.82

.73

.76

p3

.97

.87

.78

.78

P3

.94

.84

.84

.78

P^

.S8

.81

.82

P4

.90

.92

.82

Ml

.92

.80

Ml

.92

.69

M2

.80

M2

.77

Table ti.

The e.

^efficient;

s of

correlation

of tO(

)t}i width in

I P. hlaclehercii

(a-

and P. leoneiiftis, present usage (b). Figures in parentheses are not
significant. All correlations are positive.

to follow the same pattern as that of the widths. Some of
Kurten's generalizations are substantiated by the present study
(he did not include data from horses). In general, the rule of
neighborhood is confirmed, though not without exceptions.
The correlation of P^ and P| is invariably higher than that of
the former with P4. The correlation of the last molar follows
almost a random pattern, being greatest with the last premolar
in three of the four examples. The low correlations of M^ in
P. blackbergi maj- be spurious due to the small sample. There is
some tendency for the correlatioii of the third molar with more
anterior members of the series to be greater than that of the
second molar with the same members but this is not as striking

76 BT'LLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

as ill some of Kurten's data which led him to suggest a reeurva-
ture of the tiehi. The disparity in correlations between the two
species is, for the most part, insignificant.

Although the dental battery has been considered a highly inte-
grated complex by most students of fossil and Recent mammals,
correlations within this series are usually lower than those be-
tween the difterent skull elements or between varioiLs postcranial
structures (Olson and Miller, 1951, and unpublished data). The
nature of the morphogenetic field is, as stressed by Kurten and
previous workers, certainly very complex. In dental traits as Avitli
other quantitative, polygenic characters there remains the strong
possibility that the character selected for study, e.g., width of
tooth, is not a manifestation of the primary action of any genetic
determinant but rather a secondary result or "by-product" of
the action of a very large number of determinants. Although
there is a pressing need for some understanding of the heredity
of the dental and osteological characters eommonly employed by
the paleontologist, solutions to problems in this area will not be
readily available in the foreseeable future.

ACKNOWLEDGMENT

1 would like to acknowledge my indebtedness to Dr. A. 8.
Komer, JMuseum of Comparative Zoology, Harvard University,
and to Dr. Herman Gunter, Florida State Geological Survey,
for generously loaning material in their cure for study. One
specimen was l)urrowed from the U. S. National Museum, AVash-
iiigton, D. C. Although my interpretation of the facts is some-
what at variance with those of Dr. Theodore E. White, I am
deeply indebted to his earlier studies of the fauna. The many
courtesies extended by Mr. Stanley J. Olseii, Florida State
Geological Surve.v, were greatly appreciated. Funds for field
work were supplied by the Department of Biology and the
P^lorida State Museum, T'niversity of Florida. Thanks are due to
Dr. B. C. Olson and Dr. R. G. Johnson, I'niversity of Chicago,
for making space and equipment available during the course of
this investigation. ]\Ir. Bryan Patterson, IVIuseum of Compara-
tive Zoology, Harvard University, critically examined the manu-
script and suggested several changes which were incorporated
in the text ; however, I am solely responsible for the opinions
expressed herein. . .

BADER : EQI'IDAE OF THOMAS FARM 77

LITERATURE CITED

AUFFENBERO, W.

1956. Remarks on some Miocene anurans from Florida, with a de-
scription of a new species of Ilyla. Breviora, No. 52: 1-11.

Bader, R. S.

1955. Yariai)ility and evolutionary rate in the oreodonts. Evolution,
9: 119-140.

Brodkorb, p.

1954. A ehaehalaca from the Miocene of Florida. Wilson Bulletin,
66: 180-183.

Cooke, C. W.

1945. Geology of Florida. Bull. Florida Geol. Surv., 29: 1-339.
and W. C. Mansfield

1936. Suwannee limestone of Florida (abstract). Proc. Geol. Soc.
Amer., (1935): 71-72.

GoiN", C. J. and W. Auffexberg

1955. The fossil salamanders of the Family Sirenidae. Bull. Mus.
Comp. Zool., 113: 497-514.

KURTEN, B.

1953. On the variation and population dynamics of fossil and recent
mammal populations. Acta Zool. Fennica, 76: 1-122.

]\Iatthe\v, W. 1).

1924. Third contribution to the Snake Creek fauna. Bull. Amer.
Mus. Nat. Hist., 50: 59 210.

Olson, E. C. and R. L. Miller

1951. A mathematical model applied to a study of the evolution of a
species. Evolution, 5: '325-338.

QriNN, J. H.

1955. Miocene Equidae of the Texas Gulf Costal Plain. Bur. of Econ.
Geol. Univ. Texas, Publ. 5516: 1-102.

R<3MER, A. S.

1948. The fossil mammals of Thomas Farm, Gilchrist County, Florida.
Jour. Florida Acad. Sci., 10: 1-11.

Sellards, E. H.

1916. Fossil vertebrates from Florida: a new Miocene fauna; new
Pliocene species ; the Pleistocene faiuia. Ann. Rep. Florida State
Geol. Surv., 8: 77-119.

78 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Simpson, G. G.

1930. Tertiary laud mammals of Florida. Bull. Amer. Mus. Nat.
Hist., 59: 149-211.

1932. Miocene land mammals from Florida. Bull. Florida Geol. Surv.,
10: 7-41.

1943. Criteria for genera, species and suljspeeies in zoology and
paleontology. Ann. New York Acad. Sci., 44: 145-178.

1944. Tempo and mode in evolution. Columbia University Press, New
York.

1945. The principles of classification and a classification of mammals.
Bull. Amer. Mus. Nat. Hist., 85; 1-350.

Stenzel, H. B., F. E. Turner, and C. J. Hesse

1944. Brackish and non-marine Miocene in southeastern Texas. Bull.
Amer. Assoc. Petrol. Geol., 28: 977-1011.

Stirton, R. a.

1940. Phylogeny of North American Equidae. Univ. Calif. Publ.,
Bull. Dept. Geol. Sci., 25: 165-198.

TlHEN, S. A.

1951. Anuran remains from the Miocene of Florida, with the descrip-
tion of a new species of Bufo. Copeia (3): 230-235.

Vanzolini, p. E.

1952. Fossil snakes and lizards from the lower Miocene of Florida.
Jour. Paleont., 26: 452-457.

Vernon, E. O.

1951. Geology of Citrus and Levy Counties, Florida. Bull. Florida
Geol. Surv., 33: 1-256.

White, T. E.

1942. The lower Miocene mammal fauna of Florida. Bull. Mus.
Comp. Zool., 92: 1-49.

Williams, E. E.

1953. A new fossil tortoise from the Thomas Farm Miocene of
Florida. Bull. Mus. Comp. Zool., 107: 537-554.

Bulletin of the Museum of Comparative Zoology
AT H A K \^ A K D COLLEGE
Vol. 115, No. 3

ANEURETUS SIMONI EMERY, A MAJOR LINK IN

ANT EVOLUTION

by E. O. Wilson, T. Eisner, G. C. Wheeler and

J. Wheeler

With Three Plates

CAMBRIDGE, MASS., U. S. A.
PRINTED FOR T H E MUSE U M

August, 1956

Publications Issued by or in Connection

WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

Bulletin (octavo) 1863 — The current volume is Vol. 115.

Breviora (octavo) 1952 — No. 57 is current.

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

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

Occasional Papers of the Department of Mollusks (octavo) 1945 —
Vol. 1, no. 18 is current.

Proceedings of the New England Zoological Club (octavo) 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.

The continuing publications are issued at irregular intervals in numbers
which may be purchased separately. Prices and lists may be obtained on
application to the Director of the Museum of Comparative Zoology,
Cambridge 38, Massachusetts.

Of the Peters "Check List of Birds of the World," volumes 1-3 are out
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Biilletm of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 115, No. 3

ANEURETUS 8IM0NI EMERY, A MAJOR LINK IN

ANT EVOLUTION

by E. 0. Wilson, T. Eisner, G. C. Wheeler and

J. Wheeler

With Three Plates

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

August, 1956

No. 3 — Aneitretus simoni Emery, a Major Link in Ant

Evolution

By E. 0. Wtlson/ T. Eisner,^ G. C. Wheeler,''' and J. Wheeler-

INTRODUCTION

To students of ants, the genera of the tribe Aneuretini
[Aneuretus, Protaneuretus, Paraneuretus, and Mianeuretus)
have always been of primary interest because of their supposed
intermediate phylogenetie position between the most primitive
known ants (Myrmeeiinae, Ponerinae) and the advanced sub-
family Doliehoderinae. Pew tribes of ants have been more
deserving of careful study, and yet few have been so little known
or so inaccessible to myrmecologists. Aneuretus simoni,^ the only
living species of the tribe, is known only from Ceylon and has
been reported from only three collections made there. Until
recently the total number of specimens in the museums of the
world numbered no more than five or six. Published descriptions
have contained only sketches of the gross features of external
morphology, and no information whatsoever has been available
concerning the internal anatomy, ecology, or behavior of this ant.

During a recent visit to Ceylon one of us (Wilson) made a
special attempt to find Aneuretua simoni and study it in the
field. This attempt was successful to an unexpected degree.
Not only were large series of A. simoni collected, including all
brood stages and adult castes, but a strong beginning was made
in the study of the ecology and behavior of the species. Upon
his return to the United States in September, 1955, Wilson
turned over part of his collection of Aneuretus to the Wheelers,
who are currently engaged in comparative anatomical studies
of ant larvae, and to Eisner, who at that time was in the process
of completing a comparative anatomical study of the ant pro-
ventriculus. The present paper is the product of the cooperative
effort of the four authors, each of whom has concentrated on

1 Biological Laboratories, Harvard University, Cambridge, Mass.

2 Department of Biology, University of North Daljota, Grand Forks, N. D.

3 Equals Aneuretus biitteli Forel, 1913, Zool. Jahrb. (Abt. Syst.), 36: 87, fig.
Al, queen. NEM' SYNONYMY. This species was based on an isolated queen,
which we have now definitely associated with the A. simoni worker. Types of
both species have been recently examined by Wilson in the Museum d'Histoire
Xaturelle. Geneva.

82 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

those aspects of special interest to himself. Specifically, Eisner
and Wilson have collaborated in the section on adult morphology,
the Wheelers have contributed the section on larval morphology,
and Wilson has contributed the sections on distribution, ecology,
and phylogeny.

ADULT MORPHOLOGY

Worker. (PI. 1, figs. E, F, H, I ; PI. 2, fig. B; text-figs. 1, 2).
A surprising discovery, made when the first nests of Arieuretus
simoni were opened, was that the worker caste of this species is
dimorphic. As sho'\\ai in the size-frequency histogram of Figure
1, this is true complete dimorphism, with intermediates lacking.
While the distribution given represents only a single colony, it
is probable that if all the colonies collected were compounded,
two completely discrete size-distributions would still be obtained.
As shown in Plate 1, the major worker differs from the minor
primarily in its proportionately larger and broader head, and
in its relatively shorter, stouter propodeal spines. In the queen,
it will be noted, both of these allometric trends are carried a bit
further. The head is wider with respect to its length than in the
worker castes, and the propodeal spines are reduced to mere
angles.

The following description is based on the minor worker. Eyes
small, containing only about 30 ommatidia. Antennae 12-
segmented, with the segments of the flagellum increasing distally
both in length and width. Clypeus broad and flat, lacking a
median carina, the anterior border strongly emarginate in full-
face view. Mandible as shown in Plate 2B. Mandibular denti-
tion reminiscent of Bolichoderus (s. str.), relatively constant
except in the following two characters: (1) the shapes of all the
individual teeth vary somewhat, and (2) the number of the
reduced median teeth (located between the three apical and
three major basal teeth) varies from four to eight. Maxillary
palp 3-segmented, the terminal segment showing a variable
median constriction or local cuticular thickening, which may
well indicate a phylogenetically recent fusion site. Labial palp
4-segmented.

Alitrunk as figured. Propodeal spines spike-like, tapering
abruptly near the tips to form sharp points; seen from above,

WILSON AND OTHERS : ANEURETUS SIMONI EMERY 88

they diverge from one another stron*i'ly. Metasternal gland bulla
prominent, its openings supplied with a small number of coarse
guard hairs originating from both the dorsal and ventral lips.
Anterior peduncle of petiole long and slender, somewhat
narrowed toward its anterior attachment. The petiolar node
well differentiated from the anterior i)eduncle by dorsal and
lateral swellings. Seen from above, the node proper is nearly
twice as broad as long. Posteriorly, it narrows abruptly to the
point of attachment to the gaster.

2B -

ACC. NO. 1288

e
e

0.26 0.44

Pronotol width

Fig. 1. Size-frequencj' histogram of the entire worker population of a
single colony of Ancurctiis simoni from Katnapura, Ceylon, showing worker
dimorphism.

The gaster lacks an}' form of intersegmental constriction what-
soever. The sting is Avell-developed, sclerotized, and exsertile.

Integument relatively thin and collapsible, resembling in this
respect that of the higher dolichoderines and formicines. Color
of living and fresh alcoholic material varies within most nest
series from light yellow (in callows) to medium yellowish orange.
Body lightly shagreened overall, subopaque to feebly shining;
the dorsal and most of the lateral surfaces of the propodeum are
in addition transversely rugulose.

Pilosity generally sparse. Appendages covered with abundant
short, oblique pubescence, but lacking longer hairs except on the
coxae and tarsal claws. Body supplied with scattered, relatively
long, pointed erect hairs distributed as shown in Plate 1 ; body
pubescence appressed and sparse.

Proventriculus as shown in Text-figure 2. Because of its ex-
tremely small size and flaccid framework this organ is easily
deformed during preparatory handling, and it accordingly varies

84

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Fig. 2. Worker proventriculus of Aiieuretus simoni. A, cuticular frame-
work with musculature removed. B, longitudinal section through interplicary
plate and portal quadrant, showing musculature. Bl., bulb ; c. m., circular
muscles ; cp., cupola ; Cr., crop ; interpl., interplicary plate ; /. in. 1 and
I. m. 3, longitudinal muscles; M. G., midgut; pL, plica; pt., portal; qd.,
cupolar quadrant; st. vlv., stomadaeal valve. Further explanation in the
text.

WILSON AND OTHERS : ANEURETUS SIMONI EMERY 85

from one preparation to the next; for this reason, the habitus
portrayal of Text-figure 2 must be considered semidiagrammatie.
The terminology of the following description is the same as that
adopted in tlie forthcoming monograph on the formicid pro-
ventriculus by Eisner. The cuticular framework consists of a
thin-walled, flask-shaped bulb (hi.), opening anteriorly into the
crop (Tr. ) through a relatively wide cruciform portal (pt.).
The walls of the bulb are four broad, sclerotized, inwardly
curved interplicary plates (interpl.), joined basally and merging
into a slender, tubular stomodaeal valve (st. vlv.). The thin and
flexible cuticle of the valve, after extending into the midgut,
folds back u))()n itself so as to constitute a compound tube. The
interplicary plates of the bulb are separated from one another
by relatively narrow longitudinal strips of thin, flexible cuticle,
the pleats or plicae (pL), and extend anteriorly into the lumen
of the crop as four apically rounded lobes or quadrants (qd.),
which collectively constitute the cupola (cp.). Actually, the
cruciform shape of the portal is determined by the presence of
the cupolar quadrants. The bulb is surrounded by a strongly
developed layer of circular muscles (cm, Text-fig. 2B). The
longitudinal muscles are arranged in two distinct groups (1. m. I
and /. m. 3). The more strongly developed of these muscles
(/,. m. 1) originate on the interplicary plates of the bulb, extend
over the plates and insert anteriorly on the quadrants of the
cupola. The second group {I. m. 3) consists of a few scattered
fibers that extend from the crop to the midgut outside the circu-
lar muscles. Some of these fibers terminate at the base of the
proventriculus. where they insert on the cuticle at the point of
origin of the stomodaeal valve.

The glandular system has not yet been studied, but it is note-
worthy that in the field the workers defending their nests were
never observed to produce anal droplets, nor did they produce
the "Tapinoma-odor," resembling rancid butter or coconuts,
which is supposed to characterize most Doliehoderinae.

Queen. (Plate 1.) The queen is much larger in size than either
of the worker subcastes and differs further in its proportionately
broader head and greatly reduced propodeal spines. Tn addition,
it is much darker, being overall dark l)rown. It is essentially
similar to the workers in petiolar structure, mandibular denti-
tion, pilosity. sculpturing, etc.

86

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

The wing venation (of both the queen and the male) is rela-
tively generalized and similar to that of the most generalized
true dolichoderine, Dolichoderus s. sir., as typified by D. aftela-
hoides (Fabr.). The latter species, as pointed out by Brown and
Nutting (1950) in their important review of formicid wing vena-
tion, possesses all of the venational elements of the primitive
ponerine pattern except for the first radial cross-vein. Aneuretus
simoni shows two significant further advances. First, Mf2 is
completely contracted, so that the ends of Rs2 and the cross-vein

Fig. 3. Larva of Aneuretus simoni (immature). A, head in anterior
view, X 130. B-D, three types of body hairs, X 546. E, larva in side view,
X 43. F, left mandible in posterior view, X 325. G, left mandible in
anterior view, X 325. H, left antenna in lateral view, X 546.

m-cu are contiguous. Second, Rs4 has contracted to bring about
the alignment of the cross-veins 2r and r-m.

Male. Plates i and 2 show almost all of the important features
of external morphology of this caste. Especially noteworthy are
the conservation of the "myrmecioid" petiolar structure shown
also by the worker and queen, the relatively generalized structure

WILSON AND OTHERS : ANEURETUS SIMONI EMERY 87

of the mandible, and the generalized structure of the genitalia,
especially the volsella. One perhaps significant advanced feature
is the absence of serration on the ventral edge of the penis valve.

LARVAL MORPHOLOGY

Immature larva. (Text-fig. ;}.) Body length (through spira-
cles) ]. 1-2.1 nun. Straight length 0.8-1.6 mm. Somewhat sig-
moidal in side x'lew ; thorax forming a stout neck Avhich is bent
ventrally at 90° ; abdomen rather stout, diameter greatest at
abdominal somite 1\'; abdominal soinites IX and X small and
directed posterodorsally, the tenth forming a terminal naked
knob. Anus posterior. Leg and wing rudiments present. Spira-
cles small, decreasing in diameter posteriorly. No spinules seen
on the integument. Body hairs short, moderately numerous, uni-
formly distributed, of three types: (1) 0.027-0.045 mm. long,
slightly curved or flexuous, with a few minute denticles, without
alveolus and articular membrane, the most numerous type; (2)
0.045-0.072 mm. long, longest and most numerous anteriorly,
slightly curved or flexuous, with a few minute denticles, with
alveolus and articular membrane; (3) minute (about 0.001 mm.
long), few, widely scattered. Head moderately large, feebly
cordate ; cranium transversely subelliptical, with the occipital
border impressed and the corners rounded. Mouth parts large.
Antennae small, each with three sensilla, each of which bears a
moderately long spinule. Head hairs moderately numerous, uni-
formly distributed, slightly cui'ved or flexuous, long (0.027-
0.072 mm.), with numerous minute denticles on the distal 2/3.
Labrum short and broad (breadth 2.8 times the length), bilobed
due to a rather deep impression of the ventral border; anterior
surface of each lobe with one minute hair and three sensilla;
ventral border witli two sensilla near the middle and with a
few minute spinules; posterior surface of each lobe with one
large and three minute sensilla; posterior surface spinulose, the
spinules minute and in numerous short arcuate rows, the rows
radiating from each dorsolateral corner. Mandibles moderately
large, heavily sclerotized and subtriangular in anterior view,
longer than broad; slightly curved posteriorly; apex forming a
rather long tooth which is slightly curved medially, two rather
large subapical teeth on the medial border; anterior surface

88 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

with uumerous short longitiidiual ridges (rows of exceedinglj'
minute spinules ? ) ; posterior surface with several acute denticles
on the distal half and numerous ridges on the basal half, some
of the ridges bearing a row of minute spinules. Maxillae lobose,
with the apex a small conoid directed medially, with minute
spinules in short rows which are arranged in longer rows en-
circling the galea; palp a frustum bearing four apical (two
encapsulated and two with a spinule each) and one lateral
(with a spinule) sensilla ; galea a tall peg with two apical sensilla.
Labium hemispheroidal and protruding ; with all surfaces spinu-
lose; the spinules more numerous and in more numerous rows
on the anterior surface, and larger, fewer, and in short rows or
isolated on the posterior surface ; palp a slight elevation with
five sensilla, three encapsulated and two bearing a spinule each.
Opening of sericteries wide and salient. Hypopharynx spinu-
lose, the spinules so long that those in adjacent rows overlap; the
rows numerous and radiating from each dorsolateral angle.

The larva of Aneuretus shows only one dolichoderine character

the short, broad labrum and even this is not distinctive.

The one and only character that affiliates it with the Dolicho-
derinae is, strangely enough, one which is not characteristic of
the subfamily at all, namely the terminal knob projecting pos-
teriorly or posterodorsally. Now, a terminal knob occurs in the
larvae of several genera in other subfamilies, but there it pro-
jects posteroventrally, ventrally, or anteroventrally. Only in the
most advanced dolichoderine genera Engramma, Tapinoma and
Technomyrmcx have we previously found it projecting poster-
iorly or posterodorsally. (See G. C. and J. Wheeler, 1951.)

In marked contrast are the many non-dolichoderine characters
of Aneureius larvae. These are given in the accompanying table.
It will be seen that all the non-dolichoderine characters of Aneu-
i-ehis except 3 and 4 are more generalized than their dolicho-
derine counterparts. Thus we find Aneureius standing out con-
spicuously as a generalized ant larva when considered in refer-
ence to the Dolichoderinae.

WILSON AND OTHERS : ANEURETUS SIMONI EMERY 89

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iJU BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

PHYLOGENETIC POSITION AND TAXONOMIC STATUS

OP THE ANBURETINES

Like so many archetypal or anuectant forms, Aneuretus simoni
by itself shows a remarkable mixture of primitive and specialized
characters. In some of these characters the species seems to
have remained close to the mainstream of dolichoderine evolu-
tion; in others, it has moved off on a branch of its own. Before
commenting on the totality of Aneuretus morphology and at-
tempting to place the genus phylogenetically, let us review
briefly the principal aneuretine characters and attempt to assess
them with respect to their evolutionary significance.

Female mainlihnlar dentition. In the form and placement of
the basal teeth, Aneuretiis closely resembles the primitive true
dolichoderines, especially Dolichoderus s. sir. At the same time,
the set of reduced, semicircular median teeth represents a special-
ization not shared with any of the living dolichoderines.

Integument. The adult integument is relatively thin and
collapsible, as in the higher dolichoderines and formicines.

Clypeus. In shape and proportionate size, the clypeus re-
sembles the Dolichoderinae-Formicinae more closely than the
Myrmeciinae-Ponerinae. The prominently emarginate anterior
border is highly characteristic of the Dolichoderinae.

Palpal segmentation. The reduction of the number of segments
in the maxillary palp is a specialization peculiar to Aneuretus;
it is not shared by Protaneuretus or any primitive dolichoderine
or formicine genus.

Petiole structure. The .1 neuretus petiole, with its long anterior
I)eduncle and low, rounded node, is undoubtedly of a very primi-
tive type. It is in fact most similar to that of Notlwmyrmecia,
probably the most primitive of all the known living ant genera.
There is nothing comparable to it anywhere in the Dolichoderinae
or Formicinae.

Gaster. The absence of any constriction behind the postpetio-
lar segment constitutes an important difference from most un-
specialized Ponerinae. and makes it unlikely that the aneuretines
could have been derived from any of the known ponerine tribes.
Nothomyrm.ecia, on the other hand, also lacks any form of gastric
constriction.

WILgOX AND OTHERS : ANEVRETUS SlMONl EMERY 91

Sting. The well-developed, exsertile sting of Aneiirctus is a
primitive character not shared by any of the Dolielioderinae or
Formicinae.

Wing venation. In tliis character, Aneuretus is generalized
with respect to the Dolichoderinae and Formicinae as a whole,
liiit is more advanced than Dolichodents, having undergone re-
ductions in two important venational elements (Mf2, Rs-4).

Male genitalia. Until a much-needed survey of formicid male
genitalia is available, the most that can be said about Aneuretus
is that it is unspecialized, having undergone no impoi-tant reduc-
tions or modifications beyond the loss of serration on the ventral
penial valve blades. By contrast, some of the higher dolicho-
(lerines have undergone such changes as partial reduction of
the volsellae and complete loss of the pygostyles.

ProrcntiicuJus. The proventriculus of Aneuretus simoni
conforms dost ly to the primitive formicid type, as seen in the
Myrmeciinae, Pseudomyrmicinae, Ponerinae, and Cerapachyi-
nae, which in tui-n does not differ much from the generalized
aculeate i>attern. Specifically, there is a mobile portal associated
with tiexit)le i^licae in the bulb, with no suggestion of the portal
gi-owth and reinforcement through plical sclerotization seen in
the higher Dolichoderinae and Formicinae. At the same time,
there is apparently lacking in Aneuretus a group of longitudinal
muscles (/. in. 2) which is characteristic of the lower ant groups.
This lack of /. rn. 2 is surprising in view of the otherwise primi-
tive structure of the Aneuretus proventriculus, and may repre-
sent an important evolutionary' step toward the proventricular
type of the higher Dolichoderinae and Formicinae.

Odur. It was established in the field that Aneuretus simoni
adults lack the " Tapi)K)ma-()(lor" which is supposed to character-
ize the Dolichoderinae.

Larval morphology. The larva of Aneuretus simoni has onlj-
one character that definitely affiliates it with the Dolichoderinae :
the posterodorsally projecting terminal knob, which is character-
istic only of several of the most specialized dolichoderine genera.
Otherwise, larval morphology is so generalized as to set Aneu-
retus off from the Dolichoderinae as a group. Phylogenetically
the larva could have been derived from that of the Myrmeciinae,
but so could the larvae of most other primitive ant groups. More
immediately, it can 1)e linked to the lai'vae of the more primitive

92 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

formicine genera such as Melophorus, which it resembles in num-
ber and shape of body and head hairs, size of mouth parts, shape
and sclerotization of mandibles, and size and shape of maxillary
palp and galea. Melophorus in turn resembles very closely the
larvae of some of the Ponerinae, especially Ectatomma (see
Wheeler and Wheeler, 1953). At the present time, however, the
interrelationships of these last two genera are poorly understood,
and it is not known to what extent larval morphology by itself
can be used as a phylogenetic standard.

Summing up the above information, it is clear that the aneure-
tines, as exemplified by Aneuretus simoni, are a group of para-
mount significance in the evolution of the ants. Their relation-
ship to the Dolichoderinae is undeniable. Some characters in
both adult and larval morphology are shared exclusively with
the Dolichoderinae. But in other characters, the aneuretines are
so generalized as to seem to place them closer to the Myrmeciinae
and Ponerinae than to the Dolichoderinae. There is evidence to
suggest, and apparently none to deny, that the aneuretines repre-
sent the direct ancestors of the Dolichoderinae, and perhaps
also of the Formicinae. At the same time it appears, on the
important basis of external abdominal anatomy, that the aneure-
tines are more closely related to NotJiomyrmecia, the living
"archetypal" myrmeciine ant of Australia, than to any other
primitive ant group. On the basis of these two major considera-
tions, future phylogenists may feel justified in deriving the Doli-
choderinae and Formicinae ultimately from a NotJwmyrmecia-
like stock through the aneuretines.

In the past, the aneuretines have been conventionally con-
sidered a tribe of the Dolichoderinae. The totality of anatomical
evidence we have presented in this paper should now make it
clear that such a taxonomic placement is awkward, since an in-
clusion of the aneuretines in the Dolichoderinae would make
that subfamily heterogeneous out of all proportion to the re-
mainder of the ant subfamilies, and its diagnosis would then
have to depend on relatively minor characters in cephalic struc-
ture. We consider that the best course is to elevate the aneure-
tines to subfamilv rank, coordinate with the Dolichoderinae.

WILSON AND OTHERS : A^^EURETUS SIMONl EMERY 93

Subfamily ANBURETINAE Emery

Aneuretini Emery (tribe). 1912, Gen. Ins.. 187: (5. Type: AneureUis Emery;
monobasic.

Aneuretini, Wlu-eler, I'JU, Sclu-ift. Phys.-okon. Ges. KiJnigsbeif^, 55: 71.
(Description ;ind inclusion of the Baltic amber genera rrotancurctits
and Paraneitretus) .

Aneuretini, Carpenter, 1930, Bull. Mus. Comp. Zool. Harv., 70: 39. (De-
scription and inclusion of the genus Miaywiircfus from the Florissant
shales).

Aneuretinae (subfamily), Clark,^ 1951, Formicidae of Australia, Genus
Myrmecia (Commonwealth Sci. Indust. Rt-s. Org:., Australia), p. 16.

Diagnosis. (1) Clypeus broad, flat, and anteriorly emarginate,
as in the Dolichoderinae.

(2) Petiole with an anterior peduncle at least as long as the
basal portion of the node.

(3) Gaster completely lacking constrictions behind the first
postpetiolar segment.

(4) Sting well-developed, sclerotized, and exsertile.

(5) Larva primitive in form, with well developed neck, large,
sclerotized mandibles, and paxilliform maxillary palps and
galeae.

( 6 ) Pupae enclosed in cocoons ; larvae with large, salient seric-
tery-openings.

(7) Adult integument relatively thin and collapsible.

(8) Proventriculus generalized, with simple, mobile portal.
Pro ventricular muscle group I. m. 2 seemingly lacking.

(9) Fore-wing venation relatively generalized, but with the
following specializations: first radial cross-vein lacking, Mf2
and Rs4 completely contracted.

Note on internal classification. The known genera of the sub-
family include the nominate genus, with the single species de-
scribed herein, and the extinct Protaneuretus, Paraneitretus,
and Mianeurctus. Together, the.se four appear to constitute a
single tribal entity, to be known as Aneuretini.

1 While we lue in agreement witli Clark's elevation of the Aneuretini from
tribal to subfainilial rank within the Formicidae, we consider most of his other
subfamilial crcatious unwarranted, cs.. Nothomyrnieciinae, Eusphinctinae, Disco-

thyrinac. and Odontiiinachinai'.

94 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

DISTRIBUTION AND ECOLOGY

The present known distribution of Aneurefus simoni is shown
in Figure 4. At present this species is known only from Ceylon.
Here it is apparently more abundant in the southern part of
the range, where it is associated with the rather limited rain-
forest area. Around Kandy and Peradeniya, the two localities
from which the species had previously been recorded by other
authors, Wilson searched in vain for it during a period of five
days (July 10-14), before moving on into the more fruitful Rat-
napura area.

The least disturbed forest within the Kandy-Peradeniya area
is that of the Udawaddatekele Sanctuary, located just north of
Kandy and representing the remains of the old royal gardens of
the Kandy kings. This forest is relatively lush, with many large
evergreen trees, but it is noticeably drier than the forests of
the Ratnapura area and does not have a rainforest aspect. The
impression that an insect collector gets here is that the ant
fauna is diverse, but that the total population (of individuals)
is rather sparse. Scarcity of individuals seems to be due in part
to the scarcity of rotting wood on the ground, brought about in
turn by the incessant activity of Singhalese wood-gatherers from
Kandy. The few nesting sites left for wood-dwelling ants are
further diminished by the dense population of termites, which
on Ceylon are extremely abundant and more than able to hold
their own in competition with the ants.

The following general observations were made with respect to
the ant fauna of the Udawaddatekele Sanctuary. A great many,
perhaps most, of the myrmicines, dolichoderines, and formicines
nest on the ground in pieces of rotting wood, and forage both
on the gromid and up onto low vegetation. Ponerines nest
primarily in pieces of rotting wood on the ground also, and con-
fine their foraging to the ground. A high-arboreal fauna is not
well developed, either in species or individuals. Judging from
the number of individuals seen both in and away from nests,
the dominant genera on the ground appear to be Ponera and
Pheidole, followed by M]jrmicaria, SoJenopsis (Diplorhoptrum),
and Euponera {Mesoponera) , in that order. Also abundant, but
considerably less so than the above genera, are Leptogenys (s.

WILSON AND OTHERS : ANEURETUS SIMONI EMERY

95

lat.), ParatrecJmia (Nylanderia), and Pseudolasius. Ainonjz' the
low veg'etation foragers, Tecknomynnex leads, followed by Tctra-
morium (with Xiphomyrme.r), and then rolyrhachis and Cmv-
ponotus in that order. Croiiatogastcr forages o)i the ground
and up into the trees, but is rather scarce in this locality.
While not a single specimen of Aneuretus simoni was taken

Colombo

Pig. -i. Map of the suutlieni liulf ut Ceylon, showmy the localities wheif
Aneurcius simoni has been eollet-ted. The port of Coloiiilio is included
only for reference.

from the Kandy-Peradeniya forests, where it was most expected,
it proved to be one of the dominant ants not far to the south, in
the area between Ratnapura and Adam's Peak. At Ratnapttra
it was found to be extremely abundant in second-growth forest

96 bullp:tin .- museum op comparative zoology

around the city reservoir, and in similar "scrub" forest just
two miles east of the city. It was also discovered in the extensive
tract of rainforest that lies northeast of the villap:e of Gilimale
and near the foot of Adam's Peak.

The (Hlimale forest was made the site of further intensive col-
leetino- in order to gain a picture of the natural environment of
Aneureiu^. Although far from being- primar^' over most of its
range, this forest is one of the least disturbed in Ceylon, and
is probably not greatly ditferent in elementary composition from
the virgin rainforest of the nearby Sinharaja district.^ In at
least one spot remote from the main trails, the forest was found
to approach what appeared to be a primary condition and ex-
hibited several aspects of the "normal" tropical rainforest : trees
forming several stories and a closed canopy, undergrowth
sparse, and leaf litter and mold thin. Hotting wood is generally
more abundant on the ground than at Kandy, but the total ant
population is only slightly denser. The order of genus-dominance
in the ant fauna is quite similar to that at Kandy, with the
striking introduction of Aneurctus among the most prevalent
components. On the ground, Ponera and Phcidole appeared to be
the most abundant, as at Kandy. These are followed by Myrmi-
caria, AiieuretHs, Paratrechina (Nijlanderia), Eu ponera {Meso-
ponera), and Euponera {Bracliy ponera) , in that order. Among
the genera foraging on low vegetation, Techtiomyrmex is bj' far
the most abundant, followed by Polyrhachis and Camponotus.
Tetramorium occurs, but is far less common than at Kandy.
Crematogasfcr is also present but relatively scarce.

In the Oilimale forest, colonies of Aneuretus simoni were most
abundant at the edge of clearings (see Plate 3). This, coupled
with the fact that the species is also successful in the scrubby
forests around Ratnapura, may suggest that it is not naturally
a dweller of the deep rainforests, but is adapted to clearings and
more open forest types.

Some consideration should be given as to why Aneureius
shnoni has been able to survive, and indeed flourish locally, in
Ceylon, w^hile in other parts of the world the aneuretines have
faded to extinction. The general problems of extinction and
range delimitation are certainly among the most complicated
and least understood in biology, and to attempt to supply an

1 Fiir ;i description of the Sinharaja fori'st, sfc Baiter (1938).

WILSON AND OTHERS : ANEURETUS SIMONI EMERY 97

answer in this particular case with any tone of resoluteness would
be unwarranted. Nevertheless, it may be of help to future stu-
dents to mention a special condition of the Ceylon environment
which appears to favor the survival of Aiiciirehi.s. This is the
relatively weak development of the two ant genera Pheidole and
Crematognster. These two groups between them tend to occupy
both the preferred nest site and food niche of Aneuretus (so far
as these are known), and in many other tropical areas of the
world they are overwhelming dominants of the ant fauna. On
Ceylon, Pheidole is, to be sure, (|uite abundant, but relatively
much less so than in comparable forests studied (by Wilson)
in tropical Mexico and Melanesia, and it is represented by a
smaller number of species. Crematogaster is very sparsely repre-
sented in both species and individuals.

It is also important to emphasize in this connection that since
Aneuretus simoni is both ground-dwelling and primarily ground-
foraging, its principal competition probably comes not from
the dolichoderines, which are the phylogenetic successors of the
aneuretines, but rather from the ponerines and myrmicines.

During the course of the field work in 1955 a total of 20 nests
of Aneiiretits siynuni were collected. The number of individuals
in each ranged from only two to over one hundred. Only two of
the nests yielded dealate queens. One of these had, in addition
to the queen, 94 minor workers, one major worker, and a quantity
of eggs and small-to-medium larvae. The other contained what
appeared to be an incipient colony, with eight relatively small
minor workers and a (piantity of all stages of brood in addition
to the queen. Two other ne-sts contained alate queens, and two
contained males. Eight contained major w-orkers, but there were
never more than two of these to a nest. Nearly all of the twenty
ne.sts contained brood of various stages.

Eighteen of the 20 nests were in pieces of rotting wood on the
ground. The pieces ranged in size from small twigs up to fallen
tree-limbs three inches in diameter. In most cases the wood had
decomposed to the extent that it could be easily crumbled in
the hands. In two instances the Aneuretus were found beneath
rather large rotting logs on the ground. No nests at all were
found under rocks or in the open soil away from w^ood.

98 BULLETIN : MUSEUAI OP COMPARATIVE ZOOLOGY

Numerous workers were found foraging during the daytime,
during both overcast and sunny weather. They seemed to prefer
shady places, and very likely were nocturnal also, although no
field observations could be made during the night to check this
supposition/ The great majority of workers encountered away
from the nest were running over the surface of the ground; a
few were taken on herbaceous plants and shrubs at heights of up
to four feet.

Only a few observations were made on food habits. As might
be expected of a primitive ant, Aneuretus is predaceous. On two
occasions in the field, workers were found carrying entomobryid
collembolans. In an artificial nest, workers of a colony accepted
entomobryids readily and on one occasion captured about twenty
of these insects in the course of one morning. Most significantly,
the larvae were given the entomobryids directly, in the primitive
ponerine fashion, and they fed on them separately from the
workers. AYorkers in the artificial nest also accepted sugar water
readily, and at each feeding this substance was distributed
rapidly around the colony until all the adult individuals had
full, distended gasters.

Adult trophallaxis was observed on several occasions. The
donor opened its mandibles wide while the acceptor fed with
closed mandibles from the donor's lower mouthparts. On one
occasion a worker was seen donating to tw^o of its sisters simul-
taneously. The occurrence of adult-larval trophallaxis was not
investigated, although it is perhaps significant in this respect
that callow workers in the artificial nest seemed extremely atten-
tive of the brood, remaining close to it at all times.

In life the Aneureius workers have the deceptive appearance
of small yellow Pheidole minor workers, although to the ex-
perienced observer their general behavior is very un-myrmicine.
They almost always keep their long petiole folded up against
the posterior face of the propodeum, with the gaster correspond-
ingly partly elevated. In foraging, they move at about an average
gait' for ants of comparable size; very roughly speaking, they
are slower than most dolichoderines, such as Technoniyrniex
alhipes, but faster than most ponerines, such as Ponera coarctata.
Like many of the higher dolichoderines, they are much given to

1 The workers of a colonv nmintaiiu'd for a short tiuK- iu an artificial nest
forajrf'd from the brood (•h:rnil..T into the uncovered food cliamber during both
Uic day and night.

WILSON AN'D OTHERS : ANEURETVS SlMONl EMERY 99

halting abruptly at intervals, standing motionless for a moment,
and then abruptly starting off again, often in another direction.
When they are disturbed away from the nest, this l)ehavior is
intensified, with the workers dashing for a short distance, then
halting and remaining perfectly still for a while, then dashing
off again; under these conditions they show themselves capable
of considerable speed and agility. AVhen a nest is first broken
open, the minor workers swarm out aggressively and run over
any alien object offered them, l)ut they do not attempt to bite or
sting. Major Avorkers seem to be more timid, and do not venture
out in this fashion. The aggressiveness of the minor workers is
usually shortdived, and they soon retreat back into the undis-
turbed part of the nest, leaving unprotected any brood which
may happen to be in the exposed chambers. Later, tliey venture
out singly to retrieve the brood.

LITERATUEE CITED

Baker, J. E.

1938. Rainforest in Ceylon. Kew Bull., (1938) : 9 16.

Brown, W. L. and W. L. Nutting

1950. Wing venation and phylogenj- of the Formicidae (Hymenop
tera). Trans. Amer. Ent. Soc, 75: 113-132, 2 pis.

Wheeler, G. C. and J. Wheeler

1951. The ant larvae of the sultfaniily Doliehoderinae. Proe-. Ent.
Soe. Wash., 53: 169-210, 4 pis., 1 text-fig.

1953. The ;mt larvae of the subfamily Formicinae. Ann. Ent. Sot.
Amer.. 46: 126-171, 175-217, 6 pis., 2 text-figs.

PLATES

PLATE 1

A, Wings of queen. B, Wings of male. C, G, Queen. D, Male. E, E,
Worker ninjor. F, I, Worker minor.

wiiiiiiimiinrawu

PLATE 1

PLATE

A, Male mandible. B, Worker mandible. C, Terminus of male abdomen,
showing intact genitalia. D-H, Components of male genitalia isolated:
D, subgenital plate (sternite IX); E, paramere; F, volsella; G. penis
valve; H, tergites IX and X with pygostyles.

PLATE 2

PLATE 3

1. Rainforest near Gilimale, Ceylon; looking from a hill at the outskirts
of Gilimale over the forest canopy toward the cloud — enveloped base of
Adam's Peak in the distance.

2. Trail in the interior of the Gilimale forest, at a spot where Aneuretvs
simoni was one of the dominant ants.

©

;*&*

PLATx^. 3

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. 115, No. 4

THE TYPES OF NAIADES (MOLLUSCA: UNIONIDAE)
IN THE MUSEUM OF COMPARATIVE ZOOLOGY

By Richard I. Johnson

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

September, 1956

Publications Issued by or in Connection

WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

Bulletin (octavo) 1863 — The current volume is Vol. 115.

Breviora (octavo) 1952 — No. 61 is current.

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

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

Occasional Papers of the Department of Mollusks (octavo) 1945 —
Vol. 1, no. 18 is current.

Proceedings of the New England Zoological Club (octavo) 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.

The continuing publications are issued at irregular intervals in niunbers
which may be purchased separately. Prices and lists may be obtained on
application to the Director of the Museum of Comparative Zoology,
Cambridge 38, Massachusetts.

Of the Peters "Check List of Birds of the World," volumes 1-3 are out
of print; volumes 4 and 6 may be obtained from the Harvard University
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Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. 115, No. 4

THE TYPES OP NAIADES (MOLLUSCA: UNIONIDAE)
IN THE MUSEUM OP COMPARATIVE ZOOLOGY

By Richard I. Johnson

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

September, 1956

No. 4 — The Tyjjes of Naiades (MoUusca: Unionidae)
in the Museum of Comparative Zoology

Bv IvTCHAHD I. .lOIINSOX

INTRODUCTION

The collectiou of Naiades of the Museum of Comparative
Zoology had its beginuing in 1863 when Louis Agassiz purchased
the collection of J. G. Anthony of Cincinnati, Ohio. Anthony
was primarily interested in North American fresh-water shells
and, through voluminous correspondence and patience, he gath-
ered together a very extensive number of authentic contemporary
specimens. Like many pre-Darwinian naturalists, Anthony was
akin to a stamp collector in his eagerness to possess accurate
examples of each described species. Through his zeal, he laid
the foundation, as the following list will illustrate, of a large
and accurately labeled collection of type material. In his search
for authentic specimens, Anthony anticipated the present day
emphasis on the importance of types.

Agassiz not only purchased Anthony's shells, but induced him
to become the museum's first Curator of Mollusks. As curator,
Anthony continued to expand the collection. Anthony's chief
rival as a collector of fresh-water shells was the rich and socially
prominent Pbiladelphian, Isaac Lea, who devoted his very long
life to the study of the Unionidae. While many of Anthony's
own species have fallen into synonymy, he seemingly correctly
recognized Lea as an inveterate splitter who described the slight-
est variation as a new species^ and as an egotist who attempted
to attach his name to every species that he shifted from the
genus under which it was first assigned to a genus of his own
making. It appears that the two rivals began cordially to dislike
one another early in their careers, though there was a certain
cold cooperation between them throughout their lives. The only
primary types of Isaac Lea in the Museum of Comparative
Zoology are those of species which Anthony sent to Lea, which
proved to be new. Though Lea did not send Anthony any of
his types, he did identify much of Anthony's material. These
idiotypes and metatj'pes are included in this list because Lea
was prone to base a species on a single example, and it was

]i}'2 lUir.LETIX: MITSEIJM OF COMI'ARATIVE ZOOLOGY

thuujilit that these subsequent identifications might be of interest,
since they are what Lea thought he had described.

Lea described many species sent to him by C. M. Wheatley, and
these shells are now in I'^nion College, Schenectady, New York.
Wheatley sent Anthony paratypes of many of these species.
Among- other people who sent Anthony types of Unionidae were :
W. H. Benson, W. Theobold, W. B. R. H. Dunker, G. Von Dem
Busch, Largilliert, and T. A. Conrad.

The next important acquisition was purchased in 1898 by
Alexander A?rassiz from K. E. ('all. It consisted of land and
iresh-water mollusks of the United States, and had formed the
basis for Call's monographs of the Unionidae of Indiana and
Arkansas and was rich in geographic series.. In this series of
several thousand lots are some types from Gerard Troost, though
the main portion of Troost 's collection went to the Rijksmuseum
in Leiden, Holland.

It was a quarter of a century later before another large series
of Unionidae was obtained. In 1926, AV. J. Clench became
Curator of Mollusks and he donated his collection to the museum.
It contained the results of his own field efforts and duplicates
from Bryant Walker of Detroit, Michigan. In 1928 and 1929,
AV. J. Clench and P. Okkelberg made collecting trips to Ken-
tucky, Tennessee, Alabama, Georgia, and Florida. In 1933 W. J.
Clench and H. Vander Schalie collected in Virginia, Tennessee,
Alabama, and Georgia. The shells that were gathered on these
trips were divided between the L^niversity of Michigan and the
Museum of Comparative Zoology.

Prom 1914-1930 the Boston Society of Natural History parted
with all of its Unionidae that were not from New England,
and recently even the New England material has come to the
^luseum of Comparative Zoology. A number of lots from this
collection, founded in 1880, have proved to be type material.

In 1930 Joseph Bequaert ijresented to the museum a series of
African L^nionidae and Mutelidae which contained tj^pes of most
of the species he and H. A. Pilsbry described in 1927.

In 1946 while in China with the United States Navy, R. T.
Abbott was able to purchase for the museum a remarkable set
of types from the Musee Heude in Shanghai. These specimens
formed the basis of R. P. Heude 's work on the mollusks of the
Province of Nanking.

.JOHNSON : TYPES OF NAIADES IN MCZ lOo

In 1953-54 W. .1. Clench and K. I). Turner in collaboration
with the University of Florida extensiveh^ surveyed northern
Florida and the adjacent areas of Geor<2:ia and Alabama, coverinf:'
the rivers from the Escambia to the Suwannee. As a result of
this work the Museum now lias the largest collection of ITnionidae
from this area extant.

i'robably the largrest and finest assemblage of Unionidae in
the world is that of the JNIuseum of Zoology of the University of
Michigan at Ann Arbor, Michigan. Certainly its series of North
American forms is unrivaled. Its basis is the great collection of
Bryant Walker which itself consisted of many other famous ag-
gregations including those of J. LcAvis, E. R. Showalter, and L. S.
Frierson. The latter collection (see Frierson, L. S., 1927, p. 5)
was made up from the labors of other well known malacologists.

Probably ranking next in size is the collection in the Museum
of Comparative Zoology, which in addition to containing the
Unionidae described herein, has been painstakingly added to by
W. J. Clench through his efforts in the field and by world-wide
exchange. The geographic series is being constantly increased
through the devoted interest of H. D. Athearn and many others.

The United States National Museum at Washington, District
of Columbia, while lacking the extensive geographic series of
the above-mentioned institutions, doubtless has the largest num-
ber of types of Naiades in the world, containing as it does the
life work of Isaac Lea and numerous lesser collections such as
that of B. II. Wright.

The Academy of Natural Sciences of Philadelphia has a rela-
tively small series, but contains many of the types of T. Say,
T. A. Conrad and the so-called Pulson-Rafinesque types.

These are the most important collections in the United States,
but there are also A-aluable ones in the Chicago Natural History
^luseiun, Chicago Academy of Sciences, and American Museum
of Natural History in New York Citv.

The Senckenberg Museum in Frankfurt am Main, Germany,
has a large and important collection of Naiades, especially from
the Far East, which was greatly enhanced in value through
years of labor by Fritz Haas. The Berlin, Brussels, Paris, and
liritish museums all contain important types, as do many of the
smaller continental museums at Rouen, France, and Geneva,

104 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Switzerland. In general, the European collections have smaller
series and fewer records. This observation is based on the
author's rather hasty visits to most of the above-mentioned
museums.

As few of the earlier authors designated holotypes, we have
been content to call the types syntypes or cotypes. In cases
where one specimen has been well figured or we have been able
to locate the measured specimen, we have regarded these as
either holotypes or lectotypes, and considered any other speci-
mens in the type lot as paratypes. An allotype is a paratype of
the opposite sex to the holotype. Idiotypes are merely speci-
mens identified by the author of a species, and metatypes are
topotypes (from the original locality) identified by the author.
The importance of these last two categories can be questioned,
but in the absence of primary types, which can not be located
or have been destroyed, or in cases where the species was based
on a unique specimen, they have some value.

BIBLIOGRAPHY

Clench, W. J., 1936. The collection of mollusks. [in] Notes concerning
the history and contents of the Museum of Comparative Zoology. Har-
vard College, Cambridge, Massachusetts: 69-72.

Frierson, L. S., 1927. A classified and annotated check list of the North
American Naiades. Waco, Texas: 5-8.

Lea, Isaac, 1834-1874. Observations on the Genus Unio. Philadelphia, 1-13.

Simpson, C. T., 1900. Sj-nopsis of the Naiades, or pearly freshwater mus-
sels. Proc. United States Nat. Mus., 22: 504-515.

Turner, E. D., 1946. John Gould Anthony, with a bibliography and
catalogue of his species. Occasional Papers on Mollusks, Camliridge,
Massachusetts, 1 (8): 81-94.

ACKNOWLEDGMENTS

The author is especially indebted to Drs. W. J. Clench and
R. D. Turner for their ever willing help and encouragement,
which has meant time out from their own busy schedules of
research. Thanks are also extended to Mr. H. D. Athearn for
his willing aid in certain curatorial tasks connected with this
work.

JOHNSON : TYPES OF NAIADES IN MCZ 105

A List of the Types of Recent Naiades in the Museum of

Comparative Zoology in 1956, with their Original

References and Type Localities

The following list is arranged alphabetically by species, giving
the original author, the year and place of publication of its
original description, the type locality and collector (in paren-
thesis), the kind of type material — paratype or idiotype, etc.
— and finally the MCZ number and the source from which it
came to this museum. In the case of the species described by
Isaac Lea, we have only given the original reference whereby he
fixed the date of publication, and the later reference to the
particular volume of the "Observations on the Genus Unio" in
which Lea reprinted all of his descriptions and plates which
originally appeared in other journals. All locality data, in
brackets, contained in this report are additions to already pub-
lished records and have been obtained from original labels or
other sources.

Dr. J. Bequaert has kindly afforded notes on the present
status of certain species which he described with Dr. H. A.
Pilsbry in 1927.

The initials PANSP refer to the Proceedings of the Academy
of Natural Sciences of Philadelphia, JANSP to the Journal of
the same society, Tr. APS to the Transactions of the American
Philosophical Society, Pr. APS to the Proceedings of the same
society, Obs. to Observations on the Genus Unio, and MCZ to
Museum of Comparative Zoology.

ablatus [sio] Lea, Unio: 1863, PANSP, p. 193; 1867, Obs. 11, p. 17, pi. 4,
fig. 10 [name changed to ohlatus] (Long Creek, Gaston County, North
Carolina, C. M. Wheatley; North Carolina, J. G. Anthony) 2 Paratypes
MCZ 150806 ex J. G. Anthony.

alortivus Heude, Unio: 1883, Conch, Fleuv. Nanking, Part 8, pi. 63, fig.
124 (certains torrents ou ruisseaux du Ts' ing-yang Men, province dii
Ngan-houe [China]) 41/2 Paratypes MCZ 167255 ex Musee Heude.

acutissimus Lea, Unio: 1831, Tr. APS, 6. p. 89, pi. 10, fig. 18; 1834, Obs.
1. p. 99, pi. 10, fig. 18 (Alabama Eiver, Judge Tait) 4 Idiotypes MCZ
178887.

106 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

aethiopiformis 'von Ihering' Simpson, Unio: 1914, Desc. Cat. Naiades,
Detroit, Michigan, 3, p. 1312 [nomen nudum] 2 Cotypes MCZ 19236,
ex S. Putzeys coll'n.

affinis Lea, Unio: 1852, Pr. APS, 5, p. 251; 1852, Obs., 5. p. 27, pi. 19,
fig. 26 (Alexandria, Louisiana, T. Hale) 1 Idiotype MCZ 178885.

aheneus Lea, Unio: 1843, Desc. 12 Unionea [no pagination] ; 1848, Obs., 4.
p. 38, pi. 41, fig. 9 (Black Creek, Florida, S. B. Buckley), 1 Idiotype
MCZ 178713.

anceps Deshayes, Unio: 1874, Nouvelles Archives du Museum, Paris, France,
10, p. 127, pi. 6, figs. 9-12 (siir les rivages du Mekong, Cambodia
dans le sable fin, Jullien) 1 Cotype MCZ 187536 ex G. P. Deshayes.

andrewsii Marsh, Quadrula: 1902, Nautilus, 15, p. 115; 1902, Nautilus, 16,
p. 8, pi. 1, upper two figs, as Q. andrewsae (Holston Eiver, Tennessee,
Mrs. Geo. Andrews) 1 Paratype MCZ 134060 ex B. Walker coll'n.

appressus Lea, Unio: 1871, PANSP, p. 189; 1874, Obs., 13, p. 16, pi. 3, fig.
p. 8, pi. 1, upper two figs, as Q. audreivsae (Holston River, Tennessee.
Holston River, Miss Law and C. M. ^Mieatley) 1 Paratype MCZ
37102 from Miss Law in the E. R. Mayo coll'n. ex Boston Soc. Nat. Hist.

approximus Lea, Umio : 1845, Pr. APS, 4, p. 164; 1848, Obs., 4, p. 48, pi. 5,
fig. 13 (Red River at Alexandria, Louisiana, J. Hale) 2 Idiotypes MCZ
178900.

aquiliLS Lea, Unio: 1857, PANSP, p. 172; 1858, Obs., 6, p. 92, pi. 20,
fig. 72 (Flint River, near Macon, Georgia, J. C. Plant; Chattahoochee
River at Eoswell, Georgia, N. A. Pratt, Jr.) 3 Idiotypes MCZ 124884.

arcus Conrad, Unio: 1834, Amer. Jour. Sci., 25, p. 340, pi. 1, fig. 8 (Alabama
River) 1 Idiotype MCZ 178767.

argenteus Lea, Unio: 1841, Pr. APS, 2. p. 82; 1842, Obs., 3. p. 80, pi. 25,
fig. 57 (Holston River, East Tennessee, Dr. Troost and Mr. S. M.
Edgar) 4 Idiotypes MCZ 178813.

annathwaitensis B. H. Wright, Unio gibhosus : 1898, Nautilus, 11, p. 123
(A branch of the ^outh Fork of the Cumberland River at Armathwaite,
Fentress County, Tennessee) 1 ParatATe MCZ 20167 from B. H.
Wright ex L. S. Frierson.

ascia 'Benson' Hanley, Vnio: 1856, Cat. Recent Bivalve Shells, p. 385,
pi. 23, fig. 20 (Penang [Straits Settlements]). Listed by Simpson,
1914, Desc. Cat. Naiades, Detroit, Michigan, 3, p. 1192 as indeter-
minate. Lectotype, here selected, MCZ 17046, pi. 2, fig. 2, ex W. H.
Benson.

JOHNSON : TYPES OF NAIADES IN MCZ 307

askewi Marsh, Unio: 1896, Nautilus, 10, p- 91; 1897, Nautilus, 10. pi. 1,
figs. 3, 4 (Village Creek, Ilardin County, Texas, A. G. Wetherby ;
Sabine River, Texas, II. G. Askew) 2 Paratypes MCZ 30555 from the
latter locality from \V. A. Marsh, ex B. Walker coU'n.

asperatus Lea, Unio: 1861. PANSP, p. 61; 1862, Obs., 8. p. 72, pi. 7, fig.
218 (Alabama River, Claiborne, Alabama, Judge Tait ; Coosa River,
Alabama, E. R. Showalter) 1 Idiotype MCZ 178830.

auroreu^ Heude, Vnio: 1883, Conch. Fleuv. Nanking, Part 8 ,pl. 57, iig.
106 (A I'origine des torrents Ning-Kouo hien, Kien-te hien [Cliina])
4 Paratypes MCZ 167254 ex Musee Heude.

amtrali^ Simpson, Lampsilis: 1900, PANSP, p. 75, pi. 2, fig. 2, (Little
Patsaliga Creek, Southeastern Alabama) 1 Paratype MCZ 20154 ex
L. S. Frierson.

nveriUii B. H. Wright, Vnio: 1888, PANSP, p. 115, pi. 3, fig. 3 (Lake
Ashby, Volusia County, Jlorida) 16 Paratypes MCZ 134058 and 190123
from B. H. Wright ex B. Walker eoll'n.

balonnensis Conrad, Unio: 1850, PANSP, p. 10; 1854, JANSP, 2. p. 295,
pi. 26, fig. 3 ([Balonne River, Queensland, Australia 1") 1 Paratype
MCZ 72078 ex Acad. Nat. Sci. Philadelphia.

bardwelli Clench, Eyiid.clla (H)/ii(IeUa) : 1934, .Jour, of Couch., 20, ]). 89,
pi. 2, figs. 1-5 (Lower Reaches of the Glenelg River, Kimberly Division,
Western Australia, Beiesfoid BardwelH Holotvpe :\tCZ 41999; 12
Paratypes MCZ 42000.

barnesuinun Lea, Unio: 1838, Tr. APS, 6, p. 31, pi. 10, fig. 26; 1838,
Obs., 2, p. 31, pi. 10, fig. 26 (Cumberland River, Tennessee, Prof.
Troost) Ilolotype MCZ 5001 from Troost, ex R. E. Call eoll'n. Lcii
based this species on one specimen in the Troost collection. This is pre-
sumably that specimen, though it is slightly smaller than the measure-
ments given by Lea. 3 Idiotypes MCZ 178819.

hcaverrnsis Lea, Unio: 1868, PANSP, p. 161; 1869, Obs., 12. p. 57,
1)1. 44, fig. 109 (Beaver Creek and Long Creek, North Carolina, C. M.
Wheatley; and Carter's Creek, Geo. J. Postell) 2 Paratypes MCZ
178769 ex C. M. Wheatley. Localities not separated.

bMiia Morelet, Anodonta: 1866, Rev. et Mag. ZooL, (2) 18. p. 167 (Lacu
Toui-Sap, Cambodia) Lectotype, here selected, MCZ 175610, PI. 1, fig. 1,
ex A. Morelet. This is the first time that this species has been figured.

hcrthae Ortmann, Biplodon : 1921, Mem. Carnegie Mus. Pittsburgh, Pa., 8,
p. .128, pi. 38, figs. 1-4, [gills], pi. 46, fig. 6 (Rio Jacuhy, Caehoeira, Rio
Grande do Sul, Brazil, .1. D. Hasoman) 1 Cotype MCZ 100974 ex
Carnegie Museum.

108 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

bighyenais Lea, Unio: 1841, Pr. APS 2. p. 30; 1842, Obs., 3. p. 75, pi. 22, fig.
51 (Big Bigby Creek, Maury County, Tennessee, T. R. Button) 3
Idiotypes MCZ 178816.

hirgei Baker, Anodontoides : 1923, Nautilus, 36. p. 123; 1928, Bull. Univ.
Wisconsin, serial 1527, p. 180, pi. 68, figs. 1-4 (Sturgeon Bay, Door
County, below railroad bridge, Wisconsin) 4 Paratypes MCZ 89212.

hi.sseiianu.s Lea, [>?«;.• 1867, PANSP, p. 81; 1869, Obs., 12. p. 37, pi. 37,
fig. 90, (Bissel's Pond, Charlotte, North Carolina, C. M. Wheatley) 4
Paratypes MCZ 178690 and 202939 ex C. M. Wheatley.

blandianus Lea, Unio: 1856, PANSP, p. 263; 1858, Obs., 6. p. 65, pi.
11, fig. 47 (Othcalooga Creek, Gordon County, Georgia, Bishop Elliott)

2 Idiotypes MCZ 178786.

blandingianus Lea, Unio: 1934, Tr. APS, 5. p. 101, pi. 15, fig. 44; 1834,
Obs., 1. p. 213, pi. 15, fig. 44 (St. John's River, ? [sic] Florida, Dr.
Blanding) 2 Paratypes MCZ 178689, ex Dr. Blanding.

bolli Call, Unio: 1881, Araer. Natur., 15. p. 390 (Colorado River [Austin]
Texas) Lectotype, here selected, MCZ 5017, PI. 1, fig. 2. 15 ParatjTpes
MCZ 5016.

borealis A. F. Gray, Unio : 1882, Tr. Ottawa Field Nat. Club, No. 3, p. 53,

3 figs. (Ottawa River at Duck Island; Leaniy's Lake, near Hull,
Quebec, Canada) Lectotype, here selected, MCZ 51470 from the former
locality ex A. P. Gray coll 'n.

honrnianus Lea, Unio: 1840, Pr. APS, 1. p. 288; 1842, Obs., 3, p. 51, pi. 15,
fig. 28 (Scioto River near Chillicothe, Ohio, A. Bourne) 1 probable
Paratype MCZ 178833. The original label reads, "(unique) Genuine
rec. from the original finder Henry Moores [sic] who sent it to Lea for
description. ' '

boiil-inianus Lea, Unio: 1840, Pr. APS, 1- p. 288; 1842, Obs., 3. p. 46,
pi. 13, fig. 22 (Chattahoochee River, Columbus, Georgia, Dr. Boykin)
3 Idiotypes MCZ 141087.

breviculus Call, Unio: 1887, Pr. U. S. Nat. Mus., 10, p. 499, pi. 28 (Cur-
rant River, Shannon County, Missouri, and in Jack's Fork and Big
Creek, tributaries to it.) Figured Holotype MCZ 5020, from Jack's
Pork ; figured AllotjT)e MCZ 5023. In addition there are extensive lots
of paratypes from each locality. Ex R. E. Call coll'n.

bucMeyi Lea, Unio: 1843, Desc. 12 Uniones [no pagination]; 1848, Obs., 4,
p. 34, pi. 39, fig. 2 (Lake George and Lake Monroe, Florida, S. B.
Buckley) 3 Idiotypes MCZ 178723.

JOHNSON : TYPES OF NAIADES IN MCZ 109

buddianus Lea, Unio: 1843, Dese. 12 Uniones [no pagination] ; 1848, Obs., 4,
p. 35, pi. 40, fig. 5 (Lake George and Lake Monroe, Florida, S. B.
Buckley) 1 Idiotype MCZ 178726.

burlcensis Lea, U'tiio : 1859, PANSP, p. 112; 1859, Obs., 7. p. 33, pi. 27,
fig. 95 (Buckhead Creek, Burke County, Georgia, Bishop Elliot) 5
Idiotypes MCZ 150810.

hursa-pastoris B. H. Wright, i'uio: 1896, Nautilus, 9. p. 133, pi. 3 (Powell
River, Virginia) 3 Idiotypes MCZ 30574 from the Clinch River,
Tennessee.

huxtoni B. H. Wright, V7iio: 1897, Nautilus, 11. p. 55; figured by Simpson,
1900, PANSP, p. 80, pi. 1, fig. 6 (Lakelets of Monroe County, Florida)

2 Paratypes MCZ 20173 from B. H. Wright ex L. S. Frierson.

vacruUus Lea, Unio: 1831, Tr. APS, 4, p. 95, pi. 13, fig. 25; 1834, Obs. 1.
p. 105, pi. 13, fig. 25 (River Hoogly, Hindostan [India], G. W. Blakic)

3 Idiotypes MCZ 175565.

calif orniensis Lea, Anodonta : 1852, Tr. APS, 10. p. 286, pi. 25, fig. 47 ; 1852,
Obs., 5, p. 42, pi. 25, fig. 47 (Rio Colorado, California, J. L. Le Conte)
3 Paratypes MCZ 161905 ex I. Lea.

caliginosm Lea, Vnio : 1845, Pr. APS, 4. p. 165 ; 1848, Obs., 4. p. 53, pi. 7,
fig. 21 (Red River, at Alexandria, Louisiana, J. Hale) 2 Idiotypes
MCZ 178894.

capsaeformis Lea, Unio: 1834, Tr. APS, 6. p. 31, pi. 2, fig. 4; 1834, Obs., 1,
p. 143, pi. 2, fig. 4 (Cumberland River [Tennessee], W. Cooper) Holo-
type MCZ 178570 ex W. Cooper; 3 Idiotypes MCZ 178568.

cariosiis Say, Unio: 1817, Nicholson's Encyclopedia, 2 [no pagination]
pi. 3, fig. 2 (Delaware and Schuylkill rivers; [Susquehanna River]
Wilkesbarre [Pa.]) Neoholotype MCZ 178839, Neoallotype MCZ
151644, selected by Johnson 1947, Occ. Papers on MoUusks, 1, p. 148, pi.
19. figs. 1, 2. The type locality was limited to the Schuylkill River, near
Philadelphia, Pa.

castaneus Lea, Unio: 1831, Tr. APS, 4. p. 91, pi. 11, fig. 27; 1834, Obs., 1.
p. 91, pi. 11, fig. 27 (Alabama River [Alabama], Judge Tait) 3 Idio-
types MCZ 178925.

(Yf

Uiirbrnsis Lea, Unio: See under wlieatleiji Lea, Unio.

celtiformis Heude, Unio: 1874, Jour, de Conch., 22. p. 113; 1875, Conch.
Fleuv. Nanking, Part 1, pi. 1, fig. 4 (Kiang-Si, dans la riviere dc
Fou-tcheou [China]) 1 Paratype MCZ 167253 ex Musee Heude.

1 lU BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

cerinas Conrad, Unio: 1838, Monography Family Unionidae, Part 11, p.
95, pi. 52 (Probably inhabits the waters of Louisiana not far from
New Orleans) 1 Idiotype MCZ 178781.

chattanoogaensis Lea, Unio: 1858, PANSP, p. 166; 1859, Obs., 7. p. 27,
pi. 25, fig. 90 (Chattanooga, Tennessee, T. Stewardson; Etowah Eiver,
G. White; Coosawattec :ind Oostenaula Elvers, Bishop Elliott) 6 Idio-
types MCZ 178791.

chinensis Lea, Unio: 1868, PANSP, p. 150; 1869, Obs., 12. p. 85, pi. 53,
fig. 138 (Hong Kong, China) Holotype MCZ 175567 from Yale College,
New Haven, Conn, ex J. G. Anthony.

clncvnnatiensis Lea. Unio: 1840, PANSP, p. 285; 1842, Obs., 3. p. 32,
pi. 8, fig. 4 (Oliio River at Cincinnati [Ohio], T. G. Lea) 3 Idiotypes
MCZ 23604.

daihornesiii Le:i, Unio: 1838, Tr. AP8, 6. p. 105, pi. 24, fig. 115; 1838, Obs.,
2. p. 105, pi. 24, fig. 115 (Alabama River, near Claiborne [Alabama],
Judge Tait) 1 IdiotjTe MCZ 146969.

rlLntoiicnsis Winipson, Ptjichobrnnclius : 1900, PANSP, p. 79, pi. 5, fig. 3
(Archie's Fork of Little Red River, near Clinton, Arkansas) 11 Para-
types MCZ 192371 from C. T. Simpson ex B. Walker coll'n.

compactus Lea, Unio: 1859. PANSP, p. 154; 1859, Obs., 7. p. 36, pi. 28,
fig. 98 (Etawah River, Georgia, Bishop Elliott and Rev. G. White; and
Connasauga River, Bishop Elliot) 2 Idiotypes MCZ 178905.

co7icestator Lea, Unio: 1857, PANSP, p. 31; 1858, Obs., 6, p. 66, pi. 12, fig.
48 (Creeks, near Columbus, Georgia, Bishop Elliott and Dr. Lewis)
5 Idiotypes MCZ 178926.

confertH.s Leu. Unio: 1834, Tr. APS, 5, p. 103, pi. 16, fig. 47; 1834,
Obs., 1, p. 215, pi. 16, fig. 47 (Santee Canal, South Carolina, Prof.
Ravenel) 2 Idiotypes MCZ 178875.

connasaugainsis Lea, Margaritana : 1857, PANSP, p. 135; 1859, Obs., 7,
p. 47, pi. 32, fig. 113 (Connasauga River, one of the head waters of
the Alabama River, in Gilmer County, Georgia, Bishop Elliott; Ten-
nessee, J. G. Anthony) 2 Paratypes MCZ 146985 ex .T. G. Antiiony.

nniradicus Lea, Unio: 1834, Tr. APS, 5. p. 63, pi. 0 fig. 23; 1834, Obs., 1,
p. 175, pi. 9, fig. 2.^. (H.nl. . . ., Prof. Troo^t) 3 Idiotypes MCZ 178931
from Virginia.

cnnf<picui!ti Lea, Unio: 1872, PANSP, p. 156; 1874, Obs., 13. p. 38, pi. 11,
fig. 31 (Yadkin River, near Salisbury, North Carolina, C. M. Wheatlcy)
2 Paratypes MCZ 178878 ex C. M. Wheatley.

-lOlINSOX : TVinOS OK NAIADES IN MCZ 111

cor Couiad, i'nio: 1834, New Fresh Water Shells of tlie United Slates, p.
128, pi. 3, fig. 3 (Inhabits Elk and Flint Rivers, Alabama) Holotype
MCZ 178792 purehased by J. G. Anthony. Ftierson figured the "type"
of this species (1916, Nautilus, 29, pi. 3) and claimed later (1927,
Check list North American Naiades, Waco, Texas, p. 59) that the shell
figured by Conrad was not the one described, whereupon he selected the
"described" shell as the tj^pe. In his first article Frierson mentions
that Dr. Pilsbry thought that Conrad's figure of the shell was a
slightly older shell, but of the same species. In either case, the name
must lest on the figured Holotype which is here reproduced, PI. 2, fig.
3, as the original figure is pooi'.

crcbi i.slriata Anthony, Mnnoamdylaca : 1865, Anier. Jour. Conch., 1, p. 2Uo,
pi. 18, fig. 1 (Pegu, British Burma) Holotype MCZ 161872, 30 Para-
types MCZ 178935.

cnspata Gould, Unio: 1843, Pr. Boston Soc. Nat. Hist., 1, p.^141 (Tavoy,
British Burma, Rev. F. Mason) 3 Idiotypes MCZ 175564.

crocatu.s Lea, Unio: 1841, Pr. APS, 2. p. 31; 1842, Obs., 3. p. 76, pi. 22, fig.
52, (Savannah River, Georgia, T. B. Button) 1 Idiotype MCZ 146988.

cumberlandictis Lea, Unio: 1838, Tr. APS, 6, p. 25, pi. 7. fig. 19;
1838, Obs., 2. p. 25, pi. 7, fig. 19 (Cumberland River, Tennessee. Prof.
• Troost) 1 Idiotype MCZ 178928; 3 Tdiotypes MCZ 178899 from the
Clinch River, Tennessee.

camingii Lea, Monocondylaea: 1850 [1851] Pr. Zool. Soc. London, p. 199
[as Anodonta oumingii] ; 1860, Obs., 7. p. 53, pi. 33, fig. 114 (Malacca
[Straits Settlements], H. Cuming) 2 Paratypes MCZ 175582 ex Tl.
Cuming.

cimeolus Lea, Unio: 1840, Pr. APS, 1, p. 286; 1842, Obs., 3, p. 31, pi. 7, fig.
3 (Holston River, Tennessee, S. M. Edgar) 7 Idiotypes MCZ 178799.

cunningliami B. H. Wright, Unio: 1883, PANSP, p. 58, pi. 1, figs. 1-4 (Lakes
of Sumter County, Florida) 3 Cotypes MCZ 178700 ex B. H. Wright.

eurtus Lea, Unio: 1859, PANSP, p. 112; 1862, Obs., 8. p. 107, pi. 18, fig.
253 (Tombigbee River, Columbus, Mississippi, Wm. Spillman) 1 Idio-
type MCZ 178812 from Selma, Alabama.

dacUjlus Lea, Unio: 1840, Pr. APS, 1, p. 287; 1842, Obs., 3, p. 34, pi. 9, fig. 7
(Cany Fork River, Tennessee, S. M. Edgar) 1 Idiotype MCZ 178910.

dakola Frierson, Anodonta: 1910, Nautilus, 23, p. 113, pi. 10 (Ulvers
Point, Clear Lake, Deuel County, South Dakota, W. H. Over) 1 Para-
type MCZ 134118 ex B. Walker coll'n.

112 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

dalli B. H. Wright, U7uo: 1888 PANSP, p. 119, pi. 4, fig. 1 (Lake Beres-
ford, Volusia County, Florida) 5 Idiotypes MCZ 178704.

dariensis Lea, Vnio: 1842, Tr. APS, 8. p. 246, pL 26, fig. 61; 1842, Obs., 3.
p. 84, pL 26, fig. 61 (Near Darien, Georgia, J. H. Couper) 2 Idiotypes
MCZ 28330.

datus Lea, Unio : 1868, PANSP, p. 161; 1869, Obs., 12, p. .59, pi. 44, fig. 3
(Paw Creek, Beaver County and Long Creek, North Carolina, C. M.
Wheatley) 2 Paratypes MCZ 178710 ex C. M. Wheatley. Localities not
separated.

decisa Lea, Unio: 1831, Tr. APS, 4. p. 92, pi. 12, fig. 23; 1834, Obs., 1,
p. 102, pL 12, fig. 23. (Alabama Eiver, Judge Tait) 1 Idiotype MCZ
178797.

delpliinus Gruner, Unio: 1841, Archiv fiir Natur., I, p. 276, pi. 11, figs. 1,
la-e. (isn songi flumine Malacca [Straits Settlements]) 2 Cotypes
MCZ 186371 ex C. B. Adams eoH'n. Apparently G. Yon dem Busch
of Bremen sent specimens of this species to C. B. Adams and Isaac
Lee with the locality "New Holland?". It is with this locality and
under Gruner 's name that Lea described the shell the year following
Gruner (1842, Tr. APS, 8, p. 218, pi. 17, fig. 35), apparently unaware
that Gruner had already published the name. After Lea's paper went
to press. Von dem Busch sent Lea the correct locality. Thus, the Adams
specimens of delpliinus Gruner were those distributed by Gruner through
Von dem Busch prior to publication.

depygis Conrad, Unio: 1866, Amer. Jour. Conch., 2. p. 107, pi. 10, fig. 1
(Harpeth Eiver, Tennessee W. H. DeCamp) Type lost, see Johnson,
1948, Nautilus, 62, p. 102.

de-viatus Anthony, Vnio: 1865, Anier. Jour. Conch., 1, p. 156, pi. 12, fig. 2
(Tennessee) Holotype MCZ 161859.

diazensis S. H. Wright, Unio: 1897, Nautilus, 11, p. 5 (Lake Diaz, Volusia
County, Florida) 2 Cotypes MCZ 37314 from B. H. Wright from the
E. W. Eoper coll'n., ex Boston Soc. Nat. Hist.

dignatus Lea, Unio: 1863, PANSP, p. 189; 1867, Obs., 11. p. 55, pi. 17,
fig. 48 (Eiver Tigris at Bagdad [Iran], C. M. Wheatley) 2 Paratypes
MCZ 161975 ex C. M. Wheatley.

dimotus Lea, Unio: see under sumatrensis Lea.

discoidea Lea, Unio: 1834, Tr. APS, 5, p. 75, pi. 11, fig. 33; 1834, Obs., 1,
p. 187, pi. 11, fig. 33 (China) 1 Idiotype MCZ 175574.

JOHNSON : TYPES OP NAIADES IN MCZ 113

discrepans Lea, Vnio: 1860, PANSP, p. 92; 1860, Obs., 8. p. 22, pi. 55,
fig. 165 (North Alabama, Prof. Tuomey) 2 Idiotypes MCZ 178918.

dispar Lea, iiiio: 1860, PANSP, p. 305; 1860, Obs., 8. p. 9, pi. 51, fig. 153
(Columbus, Georgia, Bishop Elliott and 0. Ilallenbeck) 2 Idiotypes
MCZ 178932.

distaTis Anthony, Vnio: 1865, Amer. Jour. Conch., 1. p. 156, pi. 13, fig. 2
(Ohio) Holotype MCZ 146978.

dolabelloides Lea, Vnio: 1840 PANSP, p. 288; 1842, Obs., 3. p. 53, pi. 15,
fig. 31 (Holston Eiver, Tennessee, Prof. Troost) 1 Paratype MCZ
25254 from G. Troost ex E. E. Call coll'n.

dolabraeformis Lea, Vnio: 1838, Tr. APS, 6, p. 103, pi. 24, fig. 118; 1838,
Obs., 2. p. 103, pi. 24, fig. 113 (Altamaha River, Liberty County,
Georgia, Lewis Leconte; Altamaha River, near Darien, Georgia, Prof.
Ravenel) 1 Idiotype MCZ 28283.

dorei B. H. Wright, Unio: 1888, PANSP, p. 115, pi. 3, fig. 1 (Lake Monroe,
Florida) 4 Paratypes MCZ 178721 ex B. H. Wright.

downiei Lea, Vnio: 1858, PANSP, p. 166; 1859, Obs., 7. p. 28, pi. 25,
fig. 91 (Buck Lake, a bayou of Satilla River, Wayne County, Georgia,
T. C. Downie) 2 Paratypes MCZ 178716, received prior to publication.

edgariumis luecL, Vnio: 1840, Pr. APS, L p. 288; 1842, Obs., 3, p. 52, pi.
15, fig. 30 (Holston River, Tennessee, Prof. Troost and Dr. Currey;
Tennessee River, at Florence, Alabama, Mr. Button) 5 Idiotypes MCZ
178821 from the Sequatchee River, Tennessee.

cUiottii Lea, Margaritana : 1858, PANSP, p. 138; 1859. Obs., 7, p. 44, p!.
31, fig. 108 (Chattahoochee River near Columbus, Georgia) 1 Idiotype
MCZ 146990.

cllipticum Conrad, Pseudodon: 1865, Aijier. Jour. Conch., 1, p, 352, pi. 25,
fig. 1 (Cambodia, C. M. Wheatley) Holotype MCZ 175575 ex C. M.
Wheatley.

excavatus Lea, Vnio: 1857, PANSP, p. 32; 1858, Obs., 6, p. 71, pi. 13, fig.
52 (Othcalooga Creek, Gordon County, Georgia, Bishop Elliott; Etowah,
Georgia, Rev. G. White; Claiborne, Alabama, Judge Tait) 3 Idiotypes
MCZ 23602.

exigum Lea, Vnio: 1840, Pr. APS, 1, p. 287; 1842, Obs., 3. p. 29, pi. 7,
fig. 1 (Chattahoochee River, Columbus, Georgia, Dr. Boykin) 3 Idio-
types MCZ 178912.

fabula Lea, Margaritana: 1838, Tr. APS, 6. p. 44, pi. 13, fig. 39; 1838,
Obs., 2, p. 44, pi. 13, fig. 39 (Cumberland River, Tennessee, Prof.
Troost) 5 Idiotypes MCZ 146482.

114 BULLETIN : M[J8EUM OP COMPARATIVE ZOOLOGY

falcata Gould, Alasmidonta : 1850, Pr. Boston Soc. Nat. Hist., 3, p. 294;
1852, U. S. Expl. Exped., 12, p. 433, figs. 545-545b (Wallawalla,
Spokane, Wahlamet, etc., J. Drayton; Oregon and Upper Sacramento
Eiver, California, Dr. Pickering) 1/3 Cotype MCZ 155763, labeled
"Oregon District" ex Smithsonian Institution; 3 Cotypes MCZ 28296
labeled "California" ex A. A. Gould.

falcatus Higgins, Mycetopns : 1868, Pr. Zool. Soc. London, p. 179, pi. 14, fig.
6 (Forest streams near Chyavetas, Upper Amazons [Brazil], E. Bart-
lett) 1 Cotype MCZ 74218 ex E. F. Geale. This specimen agrees with
the plate and may well be the one figured.

fallax Lea, Unio : 1857, PANSP, p. 32 ; 1858, Obs., 6. p. 79, pi. 15, fig. 59
(Streams near Columbus, Georgia, Bishop Elliott; and French Broad
River, Tennessee, Joseph Clark) 2 Idiotypes MCZ 178869.

fannyae Johnson, Hyridella: 1948, Nautilus, 62. p. 47, pi. 3, fig. 1 (Marco
River, Merauke, South Dutch New Guinea, P. T. L. Putnam) Holotype
MCZ 160663.

feminalis Gould, Anodon: 1850, Pr. Boston Soc. Nat. Hist., 3. p. 293; 1852
U. S. Expl. Exped., 12. p. 436, fig. 547-547b (Wallawalla,, Columbia
River, Oregon, Dr. Pickering) 2 Cotypes MCZ 37294 ex Boston Soc.
Nat. Hist.

f rrrussaciana Lea, Anodonta: 1834, Tr. APS, 5. p. 45, pi. 6, fig. 15; 1834,
Obs., 1, p. 157, pi. 6, fig. 15 (Ohio River, near Cincinnati, Ohio, T. G.
Lea) 13 Paratypes MCZ 178770 ex T. G. Lea.

fimhriafa Frierson, Lampsilis: 1907, Nautilus, 21, p. 86, pi. 12, two upper
figures and lower left hand one (Valles River, Mexico, A. A. Hinkley)
3 Cotypes MCZ 167699 from A. A. Hinkley ex B. Walker coll'n.

fisherianus Lea, U'nio : 1838, Tr. APS, 6. p. 8, pi. 4, fig. 8; 1838, Obs., 2,
p. 8, pi. 4, fig. 8 (Head of Chester River, Maryland, T. Fisher) 2
Idiotypes MCZ 178567.

flava Pilsbry and Bequaert, Aspatharia : 1927, Bull. Amer. Mus. Nat. Hist.,
15. p. 418, pi. 43, figs. 3-3a; pi. 44, figs. 2-2a (Medje, Belgian Congo,
Lang and Chapin) 1 Paratype MCZ 142360. First published as a
distinct species, Dr. Bequaert now regards this as a variety of Aspatlia-
ria trapezia v. Martens. ' ' This form is close to typical trapezia and
may prove to be only a synonym." (Bequaert)

flava Anthony, Anodon: 1865, Amer. Jour. Conch., 1. p. 160, pi. 14, fig. 3
(Michigan) Holotype MCZ 161866.

JOHNSON : TYPES OF NAIADES IN MCZ 115

flavidens Benson, Unio: 1862, Ann. Mag. Nat. Hist., (3) 10. p. 188; [was
figured without name] 1862, Gleanings in Science, 1, pi. 7, fig. 1 (Ganges
River at Bhitoura, between Cawnpore and Allahabad [India]) 3 Para-
types MCZ 175559 ex W. H. Benson.

florentina Lea, Unio: 1857, PANSP, p. 83; 1862, Obs., 8. p. 68, pi. 5, fig.
213 (Florence, Alabama, G. White; Cumberland River, Tennessee, Prof.
Troost) 2 Idiotj-pes MCZ 178929.

fluclci Walker, Anod&ntites : 1924, Nautilus, 38, p. 53, pi. 1, figs. 3, 4
(Wanks River or Rio Coco, Nicaragua, W. IT. Fluek) 3 Paratypes
MCZ 167989 ex W. H. Fluck.

fiucM Morrison, Diplodon: 1943, Nautilus, 57, p. 14, pi. 5, fig. 5-9 (Orinoco
River above Munduapo, Venezuela, W. H. Fluck) 4 Paratypes MCZ
167994 ex W. H. Fluck.

flucki Bartsch, N ephronaias : 1906, Pr. U. S. Nat. Mus., 30. p. 393, pi. 27,
figs. 2, 3, pi. 38, figs. 2, 3, pi. 39, figs. 1, 3 (Wounta River, north west
of Kukallaya, Nicaragua, W. H. Fluek) 2 Paratypes MCZ 167491 ex
W. H. Fluck.

folliculatus Lea, Unio : 1838, Tr. APS, 6, p. 38, pi. 11, fig. 33 ; 1838, Obs., 2,
p. 38, pi. 11, fig. 33 (Savannah River [Georgia], Major Leconte) 1
IdiotjTpe MCZ 178702 from South Carolina.

foremnnmnus Lea, Unio: 1842, Pr. APS, 2. p. 224; 1842, Obs., 3, p. 85,
pi. 27, fig. 64 (Coosa River, Alabama, Dr. Brumby) 1 Idiotype MCZ
178753.

forsheyi Lea, Unio : 1859, PANSP, p. 155 ; 1859, Obs., 8, p. 39, pi. 60, fig.
182 (Fayette County, Texas, Prof. Forshey; Alabama, J. G. Anthony)
8 Paratypes MCZ 178785 ex J. G. Anthony.

friersoni B. H. Wright, Unio: 1896, Nautilus, 9, p, 134, pi. 3 (Bayou Pierre,
an arm of the Red River in Desoto Parish, Louisiana) 5 Paratypes MCZ
165697 ex L. S. Frierson.

fryanus B. H. Wright, Uiiio : 1888, PANSP, p. 113, pi. 2, fig. 1 (Lake
Ashby, Volusia County, Florida) 4 Paratypes MCZ 37202 from B. H.
Wright, from the E. W. Roper coll'n. ex Boston Soe. Nat. Hist.

fucatus Lea, Unio: 1860, PANSP, p. 92; 1860, Obs., 8, p. 35, pi. 59, fig.
178 (North Alabama, Prof. Tuomey; Tuscumbia [Alabama], L. B.
Thornton) 1 Idiotype MCZ 178898, from Murfreesboro, Tennessee.

fulgidus Lea, Unio: 1845, PANSP, p. 164; 1848, Obs., 4. p. 47, pi. 4, fig. 10
(Alexandria, Louisiana, J. Hale) 1 Idiotype MCZ 178784.

116 BULLETIN : jMUSEI'M OF COMPARATIVE ZOOLOGY

fii.sciitiis Lcti, Unio: 1843, Desc. 12 Ilniones [uo pagiiiiitioii] ; 18-18, Obs., 4.
p. 35, pi. 40, fig. 4 (Black Creek, Florida, S. B. Buckley) 3 Idiotypes
MCZ 178725.

l/anina Pilsbry, Lamptiili'i mrlalliru : 1909 [1910], PANSP, p. 539, pi. 27, fig.
6 (Ganina River, 3 miles southwest of San Dieguito [Mexico], A. A.
Hinkley) 15 Cotypes MCZ 189371 from A. A. Hinkley ex B. Walker
coll 'n.

r/u.'^foncmii: Lea, Unio: 1863, PANSP, p. 191; 1867, Obs., 11, p. 24, pi. 6,
fig. 18 (Mine Creek, Gaston County, North Carolina, C. M. Wheatley)

1 Paratype MCZ 178773 ex C. M. Wheatley.

(/rnerosvs Gould, Vnio: 1847, Pr. Boston Soc. Nat. Hist., 2. p. 220 (Newvillo,
Tavoy [Burmah], Mrs. Vinton) Vn Cot^X' MCZ 161900 from A. A.
Gould ex .1. G. Anthony coll 'n.

(Kiimetricus Lea, Unio: 1832, Tr. APS, 5. p. 38, pi. 4, fig. 10; 1834, Obs., 1,
p. 150, pi. 4, fig. 10 (Bayou Teche, Louisiana, W. M. Stewart) 2 Idio-
types MCZ 178697.

grnihii Lea. Unio: 1857, PANSP, p. 85; 1862, Ob.«., 8, p. 61, pi. 2, fig. 204
Catawba River, Gaston County, North Carolina, Dr. Genth and C. M.
Wheatley; Deep River, Gulf, North Carolina, Prof. Emmons) 2 Para-
types MCZ 178895 ex C. M. Wheatley.

(jcsnerii Lea, Anodonta : 1858, PANSP, p. 139; 1859, Obs.. 7, p. 49, pi. 31,
fig. 109 (Uphaupee [Uphapeej Creek, Macon County, Alabama, Wm.
Gesner) 2 Paratypes MCZ 152882 ex Wm. Gesner; 1 Idiotype MCZ
152883.

gibber Lea, Unio: 1838, Tr. APS, 6, p. 34, pi. 10, fig. 30; 1838, Obs., 2,
p. 35, pi. 10, fig. 30 (Caney Fork River, Tennessee, Prof. Troost) 3
^[etatypes MCZ 178804.

ijibhum Benson, Anodonto : 1855, .lour. Asiatic Soc. Bengal, 34, p. 135;
(Tangtse Kaing, China) 3 Cotypes MCZ 177314 from W. H. Benson ex
C. B. Adams coll 'n.

glaber Lea, Unio: 1838, Tr. APS, 6, p. 34, pi. 10, fig. 29; 1838, Obs., 2,
p. 34, pi. 10, fig. 29 (Holston River, Tennessee, Prof. Troost) 4 Idio-
tj-pes MCZ 178907.

gladiolus Heude, Unio: 1877, Conch. Fleuv. Nanking, Part 2, pi. 16, fig. 35
(Rivieres de Ning-Kouo-fou et de Kouang-te-tcheou; lac Tai [China!)

2 Idiotypes MCZ 167248 from Hia-in-fou, China ex Musee Heude.

ghindulosa Anthony, Aiiodon: 1865, Amer. Jour. Conch., 1, p. 163. jd. 16,
fig. 3 (Michigan) Holotype MCZ 161869.

JOHNSON : TYPES OF NAIADES IN MCZ 117

(louldianus 'Ward' Jay, Unio: 1839, Jaj''8 Catalogue (third edit.), p. 24
[nomen iiudutn] 1 Cotype MCZ 143238 ex Dr. Ward.

gracilcntus Lea, I'nio: 1857, PANSP, p. 85; 1862, Obs., 8, p. 62, pi. 3, fig.
205 (Catawba River, Gaston County, North Carolina, C. M. Wheatley)
3 Idiotypes MCZ 178701.

gracilis Lea, Anodonta: 1850 [1851] Pr. Zool. See. London, p. 197; 1860,
Obs., 7. p. 57, pi. 34, fig. 119 (Dingle, Isle of Panay [Philippine Islands],
H. Cuming) 2 Paratypes MCZ 175581 ex H. Cuming.

granulatus Lea, Uiiio: 1861, PANSP, p. 60; 1867, Obs., 11. p. 52, pi. 16, fig.
46 (Big Prairie Creek, Alabama, E. R. Showalter) 1 Paratype MCZ
178915 ex E. R. Showalter; 2 Idiotypes MCZ 178896.

grateloupiana Gassies, Anodonta: 1849, Moll. Agenais, p. 193, pi. 2, figs.
13-15, pi. 3, figs. 1-3, pi. 4, fig. 2 (la Garonne dans les nasses tranquilles
et a fond vaseux; conmiune a Beauregard avant les travaux du Canal,
trouvee aussi au bas de Rouquet par M. Ailkes; a Boe, dans les nasses)
1 Cotype MCZ 175636 ex J. P. S. de Grateloupe.

gravidits Lea, Uyiio: 1856, PANSP, p. 93; 1857, Obs., 6. p. 12, pi. 24, fig. 5
(Siam) 4 Idiotj-pes MCZ 175553.

gundl-achi Dunker, Unio: 1858, Malakozoologische Blatter, 5, p. 228 (in
flumine Taco Taco dicto insulae Cubanae, J. Gundlaeh) 1 Cotype MCZ
94849 ex W. Dunker.

haglicri Frierson, Unio: (Pleurobenia) : 1900, Nautilus, 13, p. 109, pi. 2
(North River, Alabama, near Tynes) 2 Cotypes MCZ 134119 ex B.
Walker coU'n.

hallenbeckii Lea, Anodonta: 1858, PANSP, 2. p. 139; 1859, Obs., 7. p. 50,
pi. 32, fig. 112 (Uphaupee [Uphapee] Creek, Macon County, Georgia, W,
Gesner). This is an error and should read Macon County, Alabama. 1
Paratype MCZ 150174 ex W. Gesner with the label: "A rare new
species. All I have are of the same imperfect character having been
carried over land by negroes for forty miles. ' '

hallenhecMi Lea, Unio: 1859, PANSP, p. 170; 1860, Obs., 8. p. 10, pi. 51,
fig. 154 (Flat Rock Creek and Four Mile Creek, near Columbus, Georgia,
G. Hallenbeek and Bishop Elliott) 5 Idiotypes MCZ 178728.

hanleyamiis Lea, Unio: 1852, Tr. APS, 10, p. 279, pi. 23, fig. 37; 1852, Obs.,
5, p. 35, pi. 23, fig. 37 (Coosawattee River, Murray County, Georgia,
Dr. Boykin) 4 Metat\T.es MCZ 178802.

harperi B. H. Wright, Unio: 1899, Nautilus, 13, p. G (Altamaha, Suwannee
and Flint Rivers) 4 Paratypes MCZ 186969 from Suwannee River,
Florida, from B. H. Wright ex B. Walker coH'ii.

118 BULLETIN : MirSEUM OF COMPARATIVE ZOOLOGY

Imitii B. II. Wright, U)iio {Kllii)(io) icchbianu.s: 1934, Nautilus, 47. p. 95,
]>1. 10, figs. 3-4 (Lake Consuolo, Citrus County, Florida, near Floral
City) 2 Paratypes MCZ 95947 ex B. IT. Wriglit.

liartmaHtanum Lea, Unio: 1860. PANSP, p. 370; 1862, Obs., 8, p. 77, pi. 8,
fig. 222 (Coosa River, Wetunipka, Alabama, E. R. Showalter) 2 Para-
typos MCZ 178827 ex E. K. Showalter.

hartwrightii B. H. Wright, I'liio: 1S96, Nautilus, 9. p. 121, pi. 2, figs. 4-6
(Lake Beresford, Florida) 2 Cotypes MCZ 167687 ex B. Walker coll'n.

Iiuttvmani Ortmann, IJiplodon : 1921, Mem. Carnegie Mus. Pittsburgh, Pa., 8.
p. 478, pi. 34, figs. 1-4; [gills] pi. 47, fig. 5; [glochidium] text fig. 4a
(p. 469) (Rio Guapore, near Eio San Siniao, Matto Grosso, Brazil, J.
D. Haseman) 1 Cotype MCZ 47040 ex Carnegie Museum.

hasethurstianiis Lea, Unio: 1858, PANSP, p. 166; 1859, Obs., 7, p. 29,
pi. 26, fig. 92 (Satilla River, Camden County, Georgia, T. C. Downie)

3 Idiot jT<?s MCZ 178724.

h< hi talus Conrad, L'nio: 1854, PAX8P, 2, p. 296, pi. 26, fig. 5 (Missouri)
1 Idiotype MCZ 178782.

hinUeyl B. IT. Wright, Vnio: 18S6, PANSP, p. 117, pi. 4. fig. 3 (Lake
Monroe, Florida) 1 Idiotype MCZ 30299.

hippopoetis Lea, Unio: 1845, Pr. APS, 4. p. 163; 1848, Obs., 4, p. 41, pi. 1,
fig. 1 (Lake Erie, Dr. Budd) 1 Metatype MCZ 176968.

}u>lmhergi Doello-Jurado, Mi/celopoda : 1923, Anal. Mus. Nac. Hist. Nat.
Buenos Aires, Argentina, 31, p. 518, pi. 1 (Rio Santa Lucia, provincia
de Corrientes, como a 35 kilometios de su desemboeadura en el Eio
Parana, S. Aquirre and J. W. Gez) 1 Paratype MCZ 111722 ex Mnseo
Nacional de Historia Natural de Buenos Aires.

hopeionensis Lea, Unio: 1838. Tr. APS, 6, p. 29, pi. 11, fig. 24; 1838. Obs., 2,
p. 29, pi. 9, fig. 24 ([Altamaha River] Hopeton, near Uarien [Georgia]
procured by Prof. Shepard from his friend J. H. Cowi^er [Couper])

4 Paratypes MCZ 165691 ex J. II. Couper; 2 Paratj-pes MCZ 155567
ex Prof. Shepard.

hydianu-s Lea, Unio: 183s, Tr. APS, 8, p. 14, pi. 6, fig. 14; 1838, Obs., 2,
p. 14, pi. 6, fig. 14 (Teehe River Louisiana, W. M. Stewart; vicinity of
New Orleans [Louisiana], ]\Ir. Barabino) 5 Idiotypes MCZ 28263.

ihi'rint/i Clessen, Anodontn : 1882, Malakozoologische BUitter [n. s.], 5. ]>.
191, pi. 4, fig. 5 (Taquara [Rio Grande do Sul] del niundo nova,
Brasiliae) 1 Paratyi.e MCZ 194094 from H. Von Ihering ex S. Putzeys
coll'n.

.lOllXSOX: 'I'VI'KS Ol' NAIADES IN MCZ 119

ihcringi B. IJ. Wriglit, L nio : 1898, Nautilus, 12. p. 93; figured 1900, Simp-
son, PANSP, p. 79, pi. 4, fis. 5 (Sau Saba River, .Menard County.
Texas) 2 Paratyi>es MCZ 20ir)2 ex I.. S. Fricrson.

iiiibricata Authouy, Anixlon: ISti.l, Anier. .lour. Contdi., 1, \k 159. pi. 14, tiy.

1 (Camp Lake. .Mieliit^an) Ilolotype MCZ 161860.

hniilicala Say, Anuiluitta : IS29, New Jlarmouy [Indiana] Disseminator, 2,
No. 22, p. 340 (Pond in Danvers, Massachusetts, Dr. T. W. Harris)
Neoliolotype MCZ 176769, selected by Jolinson, 1946, Occ. Papers on
Mollusks, I, p. 112, pi. 16, figs. 1, 2. The type locality was changed
to the Agawam River (Outlet of Halfway Pond), Plymouth, Massa-
chusetts.

iniprcsaa Anthony, Alanmodon : 1865, Amer. Jour. Conch., 1. p. 157, pi. 12,
(ig. 4 (Tennessee) Ilolotype MCZ 150666.

incrassatas Lea, Vnlo: 1840, Pr. APS, 1, p. 286; 1842 Obs., 3. p. 55, pi. 16,
fig. 34 (Chattahoochee Rivei', near Columbus, Georgia, Dr. Boykin) 4
Metaty^es MCZ 178771.

meptus Lea, rnio: 1852, Tr. APS, 10. p. 261, pL 15, fig. 12; 1852, Obs., 5,
p. 17, pi. 15, fig. 12 (Abbeville District, South Carolina, .T. P. Barratt)

2 Idiotypes MCZ 178694.

hnflata Lea, Si/mpJnjnota : 1831, Tr. APS, 4. p. 99, pi. 14, tig. 28; 1834, Obs.,
1. p. 169, pi. 14, fig. 28 (Alabama River [Alabama], Judge Tait) 2
Idiotypes MCZ 178754.

ininimta Anthony, Aiwdon: 1866, Amer. Jour. Conch., 2, p. 145, pi. 7, fig. 1
(Slawson's Lake, Michigan) Ilolotype MCZ 161876.

inoniatus Lea, Unio: 1856, PANSP, p. 93; 1857, Obs., 6, p. 13, pi.
25, fig. 6 (Slam, S. R. House) 1 Tdiotype MCZ 175554.

insolidiis Lea, inio: 1872, PANSP, p. 159; 1874, Obs., 13. p. 44, pi. 13,
fig. 37 (Abbeville District, South Carolina, Dr. Barratt; Fredericks-
liurg, Virginia, Dr. Emmons; Irwin's Creek, Mecklenberg Co., North
Clarolina, C. M. Wheatley) 1 Paratype MCZ 189369 ex C. M. Wheatley.

Inapecta Iredale, Aparcthyria : 1843, Australian Zoologist, 10, p. 191, pi. 2
(Holmes Creek, 12 mi. N. E. of Dar%vin, No. Territory, Australia, M. M.
Ward) 1 Paratype MCZ 196597 ex Australian Museum.

iiiterruptuti Lea, Unio: 1838, Tr. APS, 6. p. 15, pi. 6, fig. 15; 1838, Obs., 2,
p. 15, pi. 6, fig. 15 (Ilarpeth River, Tennessee, Prof. Troost) 2 Para-
types MCZ 5477 from I'rof. Troost ex R. E. Call coll 'n. 1 Idiotype
MCZ 178923.

120 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

interventus Lea, Vnio: 1861, PANSP, p. 60; 1862, Obs., 8. p. 88, pi. 11,
fig. 233 (Cahawba River, Alabama, E. R. Showalter) 1 Para type MCZ
178796 ex E. R. Showalter; 1 Idiotype MCZ 178790 from Tennessee.

iridella Pilsbry and Frierson, Lampsilis: 1907, Nautilus, 21. pi. 12, [2
lower right-hand figures] ; 1908, Nautilus, 22. p. 81 (Valles, Mexico,
A. A. Hinkley) 3 paratypes MCZ 100405 from A. A. Hinkley ex
Mus. Nat. Hist., Univ. Illinois, Urbana, 111.

iridineus Heude, Mycetopus: 1874, Coneh, Fleuv. Nanking, Part 1, pi. 8,
fig. 19 (les rivieres de Tai-ping-fou et de Fou-tcheou-fou [China]) 1
Paratype MCZ 167108 ex Musee Heude.

irisans Anthony, Anodon: 1865, Anier. Jour. Coneh., 1. p. 163, pi. 16, fig. 2
(Bostwick's Lake [Kent County] Michigan) Holotj-pe MCZ 161870.

irwinensis Lea, Unio: 1872, PANSP, p. 159; 1874, Obs., 13. p. 48, pi. 15,
fig. 42 (Irwin's Creek, North Carolina, C. M. Wheatley) 4 Paratypes
MCZ 178787 ex C. M. Wheatley.

jacTcsoni Marshall, Dipindnn : 1928, Pr. U. S. Nat. Mus., 74. p. 1, pi. 4, figs.
1-3 (Areas, Province of Minas Geraes, Bi-azil, R. W. Jackson) 2 Para-
types MCZ 52367 ex R. W. Jackson.

javanus Lea, Vnio: 1840, Pr. APS, 1. p. 285; 1842, Obs., 3. p. 58, pi. 18,
fig. 36 (Java, Von dem Busch) 3 Idiotypes MCZ 175568.

jayensis Lea, Vnio: 1838, Tr. APS, 6. p. 28, pi. 9, fig. 23; 1838, Obs., 2,
p. 28, pi. 9, fig. 23 (Florida, J. C. Jay) Va Metatype MCZ 178720.

jejunus Lea, Vnio: 1838, Tr. APS, 6. p. 9, pi. 4, fig. 9; 1838, Obs., 2. p. 9.
pi. 4, fig. 9 (Roanoke, also near Camden, South Carolina, Dr. Blanding)
2 Idiotypes MCZ 178719.

jetscliiiii Westerlund, Margaritana margaritifera: 1890, Fauna Palaarctis-
chen Region, Part 7, p. 185 (Oester-Schlesien [Juppel River] b.
Weidenau [Austria], R. Jetschin) 2 Cotypes MCZ 183476 from R.
Jetschin ex D. Thaanum coll'n.

jewettii Lea, Vnio: 1867, PANSP, p. 81; 1869, Obs., 12. p. 36, pi. 37, fig.
89 (Sink of Noonan's Lake, Florida, Col. E. Jewett) 2 Paratypes MCZ
178695 ex E. Jewett.

jonesh Vander Schalie, Lampsilis : 1934, Nautilus, 47. p. 125, pi. 15 (Pea
River, at Priston's Mill, Dale County, Alabama, J. A. Burke) 2 Par;i
types MCZ 98802 ex Mus. Zool. Univ. Michigan, Ann Arbor, Mich.

Iceinerianus Lea, Vnio: 1852, PANSP, p. 251; 1852, Obs., 5. p. 37, pi. 23.
fig. 40 (Coosawattee River, Murray County, Georgia, Dr. Boykin) 2
Idiotypes MCZ 178828.

.JOHNSON : TYl'KS OK NAIADES IN MCZ 121

l-inffu B. II. Wright, Unto: 1900, NautihiH, 13. p. 138 (A branch of the
Flint River, Baker County, Georgia) 2 Cotypes MCZ 201.56 from B. il.
Wright ex L. S. Frierson.

LirLiandianiift S. II. Wright, Unio: 1897, Nautilus, 10, p. 136 (Ochlookonce
River, Leon Co., Florida) 6 Paratypes MCZ 189802 and 186840 from
B. H. Wright ex B. Walkei- coll 'n.

lirtlandiantts Lea, Unio: 1834, Tr. APS, 5, p. 98, pi. 14, fig. 41; 1834, Obs.,

I, p. 210, pi. 14. tig. 41 (Mahoning, Ohio, J. P. Kirtland) 2 Idiotypes
MCZ 178738.

l-uUethenni^ Lea, Un.io: 1864, PANSP, p. 285; 1869, Obs., 12. p. 11, pi. 29,
fig. 65 (Near Mardin, in a stream from Kulleth falling into the Tigris
River [Iraq], C. :\I. Wheatley) 1 Paratype MCZ 152884 ex C. M.
Wheatley.

hnnarrki(m.m Lea, Unio: 1852, Tr. APS, 10, p. 266, pi. 17, fig. 20; 1852,
Obs., 5, p. 22, pi. 17, fig. 20 (Caddo River, Arkansas, Prof. Powell;
Washitta River, near the Hot Springs, Arkansas, Dr. Hale) 2 Idiotypes
MCZ 846.-1.

lanmiensis Frierson, Quadnila: 1901, Nautilus, 15, p. 75, pi. 4 (Lauana and
Banita Creeks, near Nacogdoches, Texas, Messrs. Askew, Strode, and
Frierson) 1 Paratype MCZ 69757 ox Dr. Strode.

lanceolatm Lea, Unio: 1828, Tr. APS, 3, p. 266, pi. 3, fig. 2; 1834, Obs., 1,
p. S, pi. 3, fig. 2 (Tar River at Tarborough [North Carolina], I. Lea)
3 Idiotypes MCZ 17892(i.

laiif/i Pilsbry and Bequaert, Mutcia: 1927, Bull. Amer. Mus. Nat. Hist.,
53, p. 432, pi. 39, figs. 3, 3a- b (Zambi, in the bed of an arm of the
Congo, Belgian Congo, Lang and Bequaert) 6 Paratypes MCZ 41068.
"Is a synonym of Mutela nilotim angustata Sowerby." (Bequaert).

laosensin Lea, Unio: 1866, PANSP, 4, p. 63, pi. 21, fig. 61; 1867, Obs., 11.
p. 67, pi. 21, fig. 61 (Laos Mountains, Cambodia, Mons. Mouhot, \)uy

II. Cuming) 2 Paratypes MCZ 33982 ex C. M. Wheatley. Lea refers
to these specimens.

Jargillierti Philippi, Unio: 1847, Zeit, fiir Mai., 4, p. 94 (Yucatan [Mexico],
Largilliert) Lectotype MCZ 155569, selectt-d by Johnson, 1951, Nautilus,
64, p. 79, pi. 5, fig. 2. The type locality was restricted to San Geroninu),
Campechc, Mexico. From Largilliert ex C. B. Adams coll 'n. 1 ParatyiH'
MCZ 184531.

lato-marginata Lea, Anodonia: 1834, Tr. APS, 5, p. 76, pi. 12, fig. 34; 1834,
Obs., 1. p. 188, pi. 12, fig. 34 (River Parana [Argentina], Dr. Burrough)
1 Idiotype MCZ 175657.

122 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

latus Lea, Unio: 1859, PANSP, p. 171; 1860, Obs., 8. p. 16, pi. 53, fig. 159
(Savannah Eiver, near Savannah, Georgia, G. White and Major Le
Conte) 3 Idiotypes MCZ 150808.

laiui Lea, Unio: 1871 PANSP, p. 189; 1874, Obs., 13. p. 12, pi. 2, fig. 4
(Tennessee Eiver, Tuscumbia, Alabama, B. Pybas ; Tennessee Eiver,
Ur. Edgar; Holston Eiver, A. E. Law and Prof. Cope) 2 Paratypes
MCZ 37300 from A. E. Law ex J. Lewis.

lecontianus Lea, Unio: 1838, Tr. APS, 6. p. 40, pi. 12, fig, 35; 1838, Obs., 2,
p. 40, pi. 12, fig. 35 (Conoochee Eiver, Georgia, Major Le Conte) 1 Idio-
type MCZ 178731.

lehmanii S. H. Wright, Unio: 1897, Nautilus, 10, p. 138; figured by Simp-
son 1900, PANSP, p. 80, pi. 4. fig. 9 (St. Mary's Eiver, Florida) 8
Idiotypes MCZ 178711.

lenior Lea, Unio : see imdei' lenis.

lenis Lea, Unio: 1840, Pr. APS, 1, p. 286; pi-eoccupied, changed to lenior,
1842, Obs., 3, p. 42, pi. 12, fig. 18 (Stone's Eiver, Tennessee, S. M.
Edgar) 5 Idiotypes MCZ 178901.

leprosiis Miles, Unio: 1861, Eeport on the Zoology of Michigan for 1859-60,
Lansing, Michigan, p. 30 (Huron Eiver, Livingston County, JNIichigan^
1 Cotype MCZ 151653 from M. Miles ex Peabodv Museum, Salem, Mass.

lesleyi Lea, Unio: 1860, PANSP, p. 360; 1860, Obs., 8, p. 34, pi. 58, fig.
177 (Kentucky, J. Lesley; [near Murfreesboro] Tennessee, J. G.
Anthony) 3 Paratypes MCZ 178815 ex J. G. Anthony.

lesueurianus Lea, U7iio: 1840, Pr. APS, 1. p. 286; 1842, Obs., 3, p. 33.
pi. 8, fig. 6 (Cany Fork and Holston Eiver, Tennessee, S. M. Edgar)
"Two specimens only of this species were brought by Mr. Edgar"
(Lea). 1 Para type MCZ 5293 from Edgar to Troost ex E. E. Call coll'n.

Icivisii Lea, Anodonta: 1857, PANSP, p. 84; 1862, Obs., 8. p. 44, pi. 62, fig.
187 (Erie Canal and Mohawk Eiver, Herkimer County, N. Y., James
Lewis; Genesee Eiver, below Eocliester, N. Y., C. Dewey) 2 Paratypes
MCZ 161904 ex J. Lewis.

lewisii Lea, Unio: 1861, PANSP, p. 40; 1862, Obs., 8. p. 75, pi. 8, fig. 220
(Coosa Eiver, Alabama, E. E. Showalter) 1 Paratype MCZ 178835 ex
E. R. Showalter; 2 Idiotypes MCZ 178823.

lineata Heude, Anodon: 1879, Conch. Fleuv. Nanking, Part 5, pi. 36, fig.
70 (Ou-yuen-hien [Cliina]) 1 Paratype MCZ 167247 ex Mus6e Heude.

lineatus Lea, Unio: 1840, PANSP, p. 287; 1842, Obs., 3. p. 44, pi. 12, fig.
20 (Chattahoochee River, Columljus, Georgia, Dr. Boykin) 1 Idiotype
MCZ 3 46976.

JOHNSON : TYPES OF NAIADES IN MCZ 123

lobensis Frierson, Parreyaia: 1913, Nautilus, 27. p. 85, pi- 5 (Lolio River,
Kauierun, Africa, George Schwab) llolotype MCZ 21164.

luc^i Deshayes, Anodonta: 1847, Hist. Nat. Moll. Algeria, pi. 108, ligs.
1-2 (J.a Calle, Algeria) 3 Cotypes MCZ loOCKU ex G. B. Deshayes.

liigcns Drouet and Chaper, Unio: 1892, Mem. See. Zool. France, 5. p. 147,
1)1. r>, figs. 1-3 (le Kapoeas (Seniitau) a Borneo) 2 Paratypes MCZ
174097 from Chaper ex S. Putzeys coll 'n.

lugubris Lea, Vnio: 1834, Tr. APS, 6. p. 30, pi. 9, fig. 25; 1838, Obs., 2,
p. 30, pi. 9, fig. 25 (Hopeton, near Darien, Georgia, Prof. Shepard) 2
Paratypes MCZ 37044 from J. H. Couper who gave them to Prof
Shepard, who, in turn, sent them to Lea for description, ex Boston
Soc. Nat. Hist.; 2 Idiotypes MCZ 178712.

latulentuA- Gould, L' nio : 1850, Pr. Boston Soc. Nat. Hist., 3. p. 295; 1852,
U. S. Expl. Exped., 12. p. 428, fig. 542 a-c (New Zealand, Drayton) 12
Paratypes MCZ 155765 from the Smithsonian Institution ex C. B.
Adiims coll 'n.

I lion a. Lea, Unio: 1865, PANSP, p. 89; 1869, Obs., 12, p. 19, pi. 32, fig.
74 (East Tennessee, S. S. Lyon) 1 Idiotype MCZ 178814.

macer Lea, Inio: 1857, PANSP, p. 8G ; 1863, Obs., 9. p. 24, pi. 29, fig. 271
(Roanoke River, Weldon, North Carolina, Prof. Emmons) 1 Idiotype
MCZ 150813.

)mu.^t,i.s Lea, Unio: 1841, Pr. APS, 2. p. 82; 1842, Obs., 3. p. 82, pi. 26, fig.
60 (French Broad River, East Tennessee, Prof. Troost and S. M.
Edgar) 3 Paratypes MCZ 5596 from Prof. Troost ex R. E. Call coll'n;
2 Idiotypes MCZ 178917.

marceiis Benson, Vnio flavidens: 1862, Ann. Mag. Nat. Hist., (3) 10, p. 188;
figured liy Ilanley and Theobald, 1876, Conch. Indica, p. 19, [d. 42, figs.

4 6 (Berhampooter River, Assam [India], Col. Jenkins) 1 Cotype MCZ
1 75550 ex W. H. Benson.

mardinexsi^ Lea, Unio: 1864, PANSP, p. 286; 1869, Obs., 12, p. 12, pi. 30.
fig. 66 (near Mardin, in a stream falling into the Tigris River, Asin
[Trail], C. M. Wheatley) 2 Cotypes MCZ 152886 ex C. M. Wheatley.

martoisi Ihcring, Castalina : 1893. Arehiv fiir Natur., 59, p. 81, pi. 3, iig.

5 (Rio Camaquam, Rio Grande do Sol [Brazil]) 8 Parntypes MCZ
192368 from TL von Ihering ex S. Putzeys coll'n.

mcnielii Anthony, Anodon : 1866, Amer. Jour. Conch., 2, p. 144, pi. 6,
fig. 1 ([Sand Lake] :Michigan) Ilolotype MCZ 150644.

124 JJULLETIX : MUSEUM OF COMPARATIVE ZOOLOGY

mearufii Simpson, Lampsilis: 1900, PANSP, p. 75, pi. 1, fig. i (vicinity of
Fort Clarke, Kinney County, Texas, E. A. Mearns) 4 Paratypes MCZ
87962 ex E. A. Mearns.

medelUnus Lea, Unio: 1830, Tr. APS, 6. p. 39, pi. 12, fig. 34; 1838, Obs., 2,
p. 39, pi. 12, fig. 34 (Eiver Medellin, near Vera Cniz [Mexico], Dr.
Currough) 2 Paratypes ^\VZ 178881 ex Dr. Burrougli.

medius Lea, Unio: 1861, PANSP, p. 40; 1862, Obs., 8. p. 82, pi. 10, fig.
227 (near Coosa River, Alabama, E. B. Showalter) 3 Paratypes MCZ
178822, mentioned by Lea as being in the J. G. Anthony coll'n.

menManus Lea, Unio: 1830, Tr. APS, 5, p. 76, pi. 19, fig. 59; 1838, Obs., 2,
]). 76, pi. 19, fig. 59 (Harpeth River, Tennessee, Prof. Troost) 2 IdJo-
types MCZ 178908.

mercdithii Lea, Unio: 1858, PANSP, p. 40; 1862, Obs., 8. p. 69, pi. 5, fig.
214 (Tennessee River, Tuscumlda, Alabama, L. B. Thornton) 4 Para
types MCZ 178807 ex L. B. Thornton; 2 Idiotypes MCZ 178803.

/,/^r(^v Lea, Unio: 1852, Tr. APS, 10. p. 260, pi. 15, fig. 10; 1852, Obs., 5,
p. 16, pi. 15, fig. 10 (Abbeville District, South Carolina, J. P. Bar-
ratt) 2 Idiotypes MCZ 178768 from Georgia.

niicauA Anthony, Amxloii : 1S65, .\mcr. .loiii. Coiicli.. 1, ]>. 162, jil. 16, fig.
1 (Texas) Holotype MCZ 187290.

mican.s Lea, Unio: 1857, PANSP, p. 85; 1862, Obs., 8. p. 63, pi. 3, fig. 207
(Catawba River, Gaston County, North Carolina, C. M. Wheatley and
Dr. Genth; Deep River, Gulf, North Carolina, Prof. Emmons) 2 Idio-
types MCZ 178718.

niicropteni.s Morelet, Unio: 1866, Jour, de Conch., 14, p. 63; 1875, Series
Cunchyliologiciue, Part 4, p. 349, pi. 15, fig. 6 (riviere Battambang,
Cambodje [Cambodia], M. Le Mesle) 3 Paratypes MCZ 17045 ex A.
Morelet.

mitohelli Simpson, Unio: 1896 {in Dall] Pr. U. S. Nat. Mus., 18. p. 5; 1896,
ihid. 19. p. 371, pi. 32, figs. 13 (Guadaloupe River, Victoria County,
Texas, J. D. Mitchell) 2 Paratypes MCZ 165695 from J. D. Mitchell ex
Boston Soc. Nat. Hist.

nworesianua Lea, Unio: 1857, PANSP, p. 83; 1867, Obs., 11. p. 43, pi. 114,
fig. 37 (Tuscumlna, Alabama, L. B. Thornton) 2 Paratypes MCZ 25621
from Tliornton ex II. E. Call coirn.; 2 Idiotypes MCZ 178795 from
Tennessee.

mordetMnus Heude, Unio: 1877, Couch. Fleuv. Nanking, Part 8, pi. 60, fig.
117 (La riviere de Kouang-to-tcheou [China]) 1 Paratype MCZ 167115
ex Mnsee Heude.

JOHNSON : TYPES OF NAIADES IN MCZ 125

mosnlensifi Lea, Unio: 1863, I'ANSP, p. 190; 1867, Obs., 11. p. 56, pi. 17, fi^.
49 (River Tigris at Mosul [Iraq], C. M. Wheatley) 2 Paratypes MCZ
161976 ex C. M. Wheatley.

■mucidus Morelet, Vnio: 1845, Moll, dii Portugal, p. HI, pl- 1-1, fit,'. 3
(rivieres do nord et notaninient dans la Tain6ga, le Cavado et la Lima
[Portugal] ) -2 Paratypes MOZ 152880 ex A. Morelet.

wundu.s \Aia, Unio: 1857, PANSP, p. 83; 1867, Obs., 11. p. ,44, pl. 14, tig.
38 (Tusc-unil.ia, Alabama, L. B. Thornton) 5 Idiotypes MCZ 178805.

murchisoniamis Lea, Vnio: 1834, Tr. APS, 5. p. 33, pl. 3, fig. 6; 1834, Obs.,
1, II. 145, pl. 3, fig. 6 (China, Mrs. Murchison) 4 Idiotypes MCZ
175569 from India.

ii(i)iki>i(j<'ii.si.'i lleude, Monocoiulylaca : 1874, Jour, de Conch., 22, p. 116; 1875,
(bneh. Fleuv. Nanking, Part 1, pl. 4, fig. 9 (riviere de Nanking [China] )
21/2 Paratypes MCZ 167240 ex Musee Ileude.

),a}nancnsi.s Conrad, Unio: 1852, PANSP, p. 10; 1854, JANSP, 2. p. 296.
pl. 26, fig. 3 ([Nepean Eiver, New South Wales, Australia]) 1 Paratype
MCZ 72077 from T. A. Conrad ex Acad. Nat. Sei. Philadelphia.

jmshvilUanius Lea, Unio: 1834, Tr. APS, 5. p. 100, pl. 14, fig. 43; 1834,
Obs., 1. p. 212, pl. 14, fig. 43 (Cumberland Eiver [Tennessee], Prof.
Troost; Ohio [River] at Louisville [Kentm-ky], Dr. Fitch) 2 Paratypes
MCZ 163S0 from Prof. Troost ex R. E. Call coll'n; 3 Idiotypes MCZ
178902.

7i€gatus Lea, Unio: 1861, PANSP, p. 59; Obs., 8. p. 80, pl. 9, fig. 225 (Big
Prairie Creek, Ala))ama, E. E. Showalter; Columbus, Mississippi, W.
SpiUmau) 4 Paratypes MCZ 130562 ex E. R. Showalter; 2 Paratypes
MCZ 5665 ex \V. Spillman; 2 Idiotypes MCZ 17877(5 from Alabama.

neialerii Lea, Unio: 1858, PANSP, p. 165; 1859, Obs., 7. p. 30, pl. 26, hg.
93 (Flint Eiver at Lanier, Georgia, H. M. Neisler; at Macon [County]
Georgia, J. C. Plant) 2 Paratypes MCZ 189796 ex H. M. Neisler.

Lanier was al)Out 10 miles north of Oglethorpe, Macon County,
Georgia. It does not appear on modern maps. It is not to be confused
with the town of the same name in Bryan County. Both of the type
lots probably came from the same source.

nicaraguae Philippi, Anodonta: 1848, Zeit. fiir Mai., 5. p. 130 (Nicaragua.
Largilliert) 1 Paratype MCZ 184530 from Largilliert ex C. B. Adams
was selected by Johnson (1951, Nautilus, 64, p. 81, pl. 5, fig. 1) as
lectotype. This selection was, hoAvever, invalid as a lectotype for this
species had been previously selected by Haas (1930, Senckenbergiana,
12. p. 322, fig. 1); see, Johnson (1951, Nautilus, 65. p. 32); also 1
Paratype MCZ 17071 from Largilliert ex J. G. Anthony.

126 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

nitens Lea, Unio: 1840, Pr. APS, 1. p. 288; 1842, Obs., 3. p. 43, pi. 12, lij?.
19 (Long Creek, Cocke County, Tennessee, S. M. Edgar) o I'aratypes
MCZ 5666, mentioned by Lea as being in the G. Troost coll'n.. ex R. E.
Call coU'n.

iintatits- Lea, I nio: 1838, Tr. APS, 6. p. 28, pi. 8, fig. 22; 1838, Obs., 2, p. 28,
pi. 8, fig. 22 (Cumberland River [Tennessee], Prof. Troost) 1 Idiotype
MCZ 178911.

nuhilus Lea, Unio: 1868, PANSP, p. 161; 1869, Obs., 12. p. 58, pi. 44,
fig. 110 (Paw Creek, Menklenburg County, North Carolina, C. M.
Wheatley) 1 Paratype MCZ 178730 ex C. M. Wheatley.

nu.e Lea, Vnio: 1852, Tr. APS, 10. p. 283, pi. 24, fig. 43; 1852, Obs., 5,
p. 39, pi. 24, fig. 43 (Alabama River [Alabama], C. M. Wheatley;
3 Idiotypes MCZ 178794.

Hi/angensis Frierson, Parreysia: 1913, Nautilus, 27. p. 86, pi. 5 (Nyang
River, Kamerun, Africa, George Schwab) Ilolotype iICZ 21160.

ohesa Connolly, Cafferia caffra: 1925, Records Albany [Soutli \fiica] Mus.,
3, p. 263, pi. 11, figs. 1-4 (Bushman's River, Alicedale, Cape Provinee
[Union of South Africa] ) 4 Paratj^pes MCZ 52457 from M. Connolly.

nhesiis Lea, Unio: 1831, Tr. APS, 4. p. 96, pi. 113, fig. 26; 1834, Ob.s., 1,
p. 106, pi. 113, fig. 26 (York River, Virginia, [Georgia, teste Lea, Obs., 1,
p. 118] \V. Cooper) 2 Paratypes MCZ 165693, from W. Cooj-er ex
Boston Soc. Nat. Hist.; 1 Idiotype MCZ 17869S from Prairie Creik.
Florida.

ohlattiK (.s'(c] Lea, Vnio: see under alhatus.

nbtusiu^ Lea, Vnio: 1840, Pr. APS, I. p. 287; 1842, Obs., 3, p. 39, pi. 11, fig.
13 (Chattahoochee River, Columbus, Georgia, Dr. Boykin) 3 idiotypes
MCZ 146980.

iiccultus Lea, Vnio: 1843, Des. 12 Uniones [no pagination]; 1848, Oli.s., 4,
p. 37, pi. 41, fig. 7 (Black Creek and Lake Monroe, Florida, S. l;.
Buckley) 2 Idiotypes MCZ 178916.

nchracea Say, Vnio: 1817, Nicholson's Encyclopedia, 2. [no pagination],
1)1. 3, fig. 8 (Delaware and Schuylkill Rivers [Pennsylvania]) Neoholo-
tyi)e MCZ 178838 selected by Johnson (1947, Occ. Papers on MoUusks. 1,
p. 153, pi. 20, fig. 2). The type locality was restricted to the Schuylkill
River, near Philadelphia, Pa.

ogeecheensis Conrad, Vnio: 1849, PANSP, p. 153; 1850, JANSP, 1. p. 275,
pi. 37, figs. 3-4 ([Ogeechee River, Georgia], J. II. Couper) Lectotype,
here selected, MCZ 146971 is the specimen from which figure 3 was
drawn ex J. H. Couper.

JOHNSON : TYl'ES OF NAIADES IN .MCZ 127

oleivonis Heudo, Mycetopns: 1877, Conch. Flouv. Nanking, Part 3, pi. 22,
fig. 4(5 (La Iloal supcriore, department do Ing-tch'eou [China]) 2 Para-
typos MCZ 10711(1 ox Miisoo Ilcudc

iipalioia Anthony, Aiiodon: 18C5, Auier. Jour. Conch., 1, p. 159, pi. 1-1, fig. 2
(Shear's Lake [Kent County] ISlichigan) Hoiotype MCZ 1.50633.

()l)ali)iu.s Anthony, Unio: 1866, Amer. Jour. Conch., 2, p. 146, pi. 7, fig. 2
(Michigan) Holotyi)e MCZ 161893.

(>riiior( n.'^Ls Morrison, Castalia: 1943, Nautilus, 57. p. 14, pi. .5, figs. 1-4
(Orinoco River at Maipures, U. S. Colombia, W. 11. Fluck) 2 Paratypes
MCZ 167992 ex W. H. Fluek.

orphacnsis Lea, Vnio: 1864, PANSP, p. 285; 1869, Obs., 12, p. 10, pi. 29,
fig. 64 (Tigris River, near Hardin, Pashalic of Orpha, Asiatic Turkey
[Iraq]. C. M. Whoatley) 3 Paratypes MCZ 152885 ex C. M. Wheatley.

nilnunini Walker, M'uromyu : 1925, Oce. Papers Mus. Zool., Univ. Michigan,
No. 163, p. 1, pi. 1, figs. 1-6 (Green River, Mammoth Cave. Also
Sulphur Fork of Russell Creek, Adair County, Kentucky) Hoiotype
MCZ 167705; Allotype MCZ 167702; 1 Paratype MCZ 167703 all from
the former locality.

oiimanni Pilsbry and Bequaert, Pseuclospatha : 1927, Bull. Amer. Mus. Nat.
Hist., 53, p. 446, pi. 45, figs. 5, 7 (Lake Tanganyika: Bay of Toa in
5 m.; Tembwe; Moliro [Belgian Congo]) 1 Paratype MCZ 41070
from Tembwe. "Is a variety of Pse^idospatha ianganyicensis Smith."
(Bequaert).

ortmanni Frierson, Unio (Nephronaias) : 1913, Nautilus, 27, p. 14, pi. 2
(Conchin's River, near Quirigua, Guatemala, Atlantic Drainage, A. A.
Hinkley) 3 Cotypes MCZ 20916 ex A. A. Hinkley.

o.'<becU Philippi, Unio: 1845, Zeit, fur Mai. 1. p. 164; 1847, Abbild. und
Beschr. Conch., 3, p. 45, pi. 3, fig. 1 (Yang-tsokiang, China, Largilliert)
2 Paratypes :MCZ 187289 ex Largilliert.

oscai-i B. H. Wright, Unio: 1892, Nautilus, 5. p. 124; 1896, Nautilus, 9,
p. 122, pi. 2, figs. 1-3 (A creek near Lake Osceola, at Winter Park,
i^lorida) 15 Cotypes MCZ 178717 ex B. H. Wright.

ovata Simpson, Quadrvla: see under vestitiis Heude, Unio:

(Karhensi^ Call, Unio: 1887, Pr. U. S. Nat. Mus., 10. p. 498, pi. 27 (Currant
River, Shannon County, Missouri, and in Jack's Fork and Big Creek,
tributaries to it) Hoiotype MCZ 5707; Allotype MCZ 5705 both fiom
Jack's Fork, ex E. E. Call coll'n.

128 BULLETIN : MUSEUM OF COMPARATRTI ZOOLOGY

pallaryi Longstaff, Nodularia teretiscuJa var: 1914, Jour. Linn. Soc. Lon-
don, Zool., 32, p. 256 (near Mogran, Anglo-Egyptian Sudan [Also
listed from a dozen nearby localities], J. Longstaff) 1 Paratype MCZ
173051 ex T. Pain.

pallida Anthony, Anodon: 1865, Amer. Jour. Conch., 1, p. 162, pi. 15, fig. 3
(Michigan) Holotj^e MCZ 161871.

paludicohis Gould, Unio : 1845, Pr. Boston Soc. Nat. Hist., 2, p. 53 (Ever-
glades of IHorida, J. Bartlett) 1 Idiotnie MCZ 178705.

panacocnsis Yon dem Buseh, T'nio: 1843, [in Philippi] Abbild. und
Beschr. Conch., 1, p. 75, pi. 2 (Plumea Panaeo, prope Tampico, Mexico)
3 Paratypes MCZ 178882 ex Von dem Busch.

papyracea Anthony, Anodon: 1865, Amer. Jour. Conch., 1. p. 161, pi. 15,
fig. 2 (locality unknown [Potomac River, Virginia] ) Holotype MCZ
150656.

paramattensis Lea, Unio: 1862, PANSP, p. 176; 1867, Obs., 11, p. 64,
pi. 20, fig. 59 (Paramatta River, New South Wales, Smithsonian Institu
tion) 1/2 Paratype MCZ 17015 ex Smithsonian Institution.

parccdcntaia Haas, Nodularia: 1911, Nach. Deutschen Mai. Ctes., 43, p. 43;
1911, Conch, Cab., 9, Part 2, Section 2. p. 104, pi. 12a, figs. 3-4.
(Mikawa, Japan, Hirase) 1 Paratype MCZ 44650 ex S. Hirase.

parvulus Heude, Unio: 1878, Conch. Fleuv. Nanking, Part 4, pi. 32, fig. 65
(Petits ruisseaux souvent a sec, qui re<joivent les eaux des rizieres dans
les vallons du Kien-te sud (Ngan-houe) [Giina]) 1^^ Paratypes MCZ
167246 ex Musee Heude.

patsaligensis Simpson, Pleurohema : 1900, PANSP, p. 82, pi. 2, fig. 1 ;
(Little Patsaliga, southeast Alabama) 2 Paratji)es MCZ 20179 ex L. S.
Frierson; 2 Paratypes MCZ 210276 ex B. Walker coll'n.

paulus Lea, Unio: 1840, Pr. APS, 1. p. 287; 1842, Obs., 3. p. 51, pi. 15,
fig. 29 (Chattahoochee River, Columbus, Georgia, Dr. Boykin) 3 Idio-
tj-pes MCZ 178892.

peguensis Anthony, Monocondyloea [sio] : 1865, Amer. Jour. Conch., 1,
p. 205, pi. 18, fig. 2 (Pegu, British Burma) Holotype MCZ 161877; 22
Paratypes MCZ 161874.

peguensis Anthony, Unio: 1865, Amer. Jour. Conch., 1, p. 351, pi. 25, fig. 2
(Pegu, British Burma) Holotype MCZ 161875; 1 Paratjiie MCZ 161876.

pellis-laeerti Morelet, Unio: 1865, Jour, de Conch., 13, p. 22; 1875, Series
Conchologique, Part 4, p. 355, pi. 17, fig. o (Siam) 3 Paratypes MCZ
175551 ex A. Morelet.

.)()I1.\8U.\ : TVl'KS OK XAIAUKS I\ MCZ 129

},e)iicillatii,-i Lea, Cnio: 1857, PANSP, p. 171; 1859, Obs., 7, p. 21, pi. 23,
fi's. 85 (Chattahoochee [River] near Columbus, Georgia, Dr. Boykin ;
near Atlanta [Georgia], Bishop Elliott; Flint River, near Albany.
Georgia, Bishop Elliott and G. White) 3 Idiotypes MCZ 178930.

]),'icnarr((it>t.s Lea, Umio : 1857, PANSP, p. 85; 1862, Obs., 8, p. (53, pi.
;>, (is. l'(H; (Catawba River, Gaston County, North Carolina, C. M.
Wheatloy and Dr. Genth) 3 Idiotypes MCZ 152239.

jicniodo.sus Lea, Unio: 1845, Pr. APS, 4, p. 163; 1848, Obs., 4, p. 45,
pi. 3. fig. 8 (North Carolina, W. B. Budd) 2 Idiotypes MCZ 178831.

pertionatu.s Say, Unio: 1829, New Harmony [Indiana] Disseminator, 2, No.
20, p. 309 (Wabash River [Indiana]) 4 Idiotj^es MCZ 25763 from the
Cumberland River, Tennessee. Named by Say (see Call, 1898, Mollusea
vf Indiana, p. 475), from G. Troost ex R. E. Call coll'n.

lirr.siriatu.s Lea, U^iio: 1852, Tr. APS, 10. p. 255, pi. 12, lig. 3; 1852, Obs., 5,
p. 11. pi. 12, fig. 3 (Abbeville District, South Carolina, Dr. Barratt)
3 Idiotypes :^ICZ 178691.

pictus Lea, Unio: 1834, Tr. APS, 5. p. 73, pi. 11, fig. 32; 1834, Obs., L
p. 185, pi. 11, fig. 32 (Harpeth River, Tennessee, Prof. Troost). Figured
Holotype MCZ 5792 from G. Troost ex R. E. Call coll'n; 1 Idiotype
MCZ 146977.

inlaris Lea, Uiiio: 1840, Pr. APS. 1, p. 285; 1842, Obs., 3, p. 47, pi. 14,
fig. 24 (French Broad and Ilolston Rivers, East Tennessee, S. il.
Edgar) 1 Paratype MCZ 5783 from the French Broad River, mentioned
by Lea as being in the Troost coll'n., ex R. E. Call coll'n.; 3 Idiotypes
MCZ 178780.

pinei B. H. Wright, Unio: 1897, Nautilus, 11, p. 40; figured by Simpson,
1900, PANSP, p. 80, pi. 3, fig. 1 (An unnamed lake in Withlacoochee
River Region of Hernando County, Florida) 2 Paratyi^es MCZ 20153
ex L. S. Frierson.

pinkstonii B. H. Wright, Unio: 1897, Nautilus, 10. p. 136 (Tuscaloosa
[Tallapoosa] River, Macon County, Alabama) 2 Cotj-pes MCZ 189647 ex
B.Walker coll 'n.

phi n Hat ems Conrad, Unio: 1838, Monography Family Unionidae, Part 12, p.
103, pi. 57, fig. 1 (Black Water River, Virginia; Neuse River, North
Carolina; Yadkin River, North Carolina, Dr. Blanding) 2 Idiotypes
MCZ 124885 from Louisiana.

plid.sii Marsh, Unio: 1891, Xautilus, S, p. 2 (Little Red River, Arkansas)
5 Idiotypes MCZ 29988.

ISO BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

plebeius Adams, Unio compressus: 1842, Thompson's History of Vermont,
Part 1, p. 166 (Middlel)ury [Yormont]). There were four specimens
in the original lot. The largest one, whose measurements are given in
the original description is here selected as Lectotype MCZ 154361 ;
3 Para types MCZ 154366 ex C. B. Adams coll'n.

plectophorus Conrad, Unio: 1849, PANSP, p. 162 [as plectrophoriis] ; 1850.
JANSP, 1. p. 277, pi. 38, fig. 7 (Flint Kiver, Georgia, J. H. Couper)
3 Paratypes MCZ 178840 and 28339 ex J. II. Couper.

plieiferus Lea, Unio: 1838, Tr. APS, 5, p. 61, pi. 17, fig. 53; 1838, Obs., 2,
p. 61, pi. 17, fig. 53 (Mexico, Dr. Burrough) 7 Idiotypes MCZ 6584.

polymorphm B. H. Wright, Unio: 1899, Nautilus, 13, p. 42 (Spanish Creek,
Okefenokee Swamp, Charlton County, Georgia) 5 Cotypes MCZ 30290
ex B. Walker coll 'n.

polystictus Heude, Unio : 1877, Conch. Fleuv. Nanking, Part 2, pi. 12, fig. 25
(riviere Siang [Hounan Province, China]) 1 Paratype MCZ 167116 ex
]V[usee Heude.

popenoi Call, Unio : 1885, Bull. Washburn College, 1, p. 49, pi. 2 (Fall River,
Wilson County, Kansas; A'erdigris River, Kansas) Holotype MCZ 40.'!4
from the former locality ex R. E. Call coll'n.

postelUi Lea, Unio: 1858, PANSP, p. 165; 1859, Obs., 7. p. 32, pi. 26, fig. 94
(Randall's Creek, near Columbus, Georgia, Bishop Elliott; Carter's
Creek, Baldwin County, Georgia, Mr. Postell) 5 Idiotypes MCZ 124883.

powellii Lea, Unio: 1852, Pr. APS, 5. p. 252; 1852, Obs., 5. p. 26, pi. 19,
fig. 25 (Saline River, Arkansas, Prof. Powell) 1 Paratype MCZ 1788S4
ex Prof. Powell.

prattii Lea, Unio: 1858, PANSP, p. 166; 1859, Obs., 7, p. 24, pi. 24, figs.
88, 88a (Chattahoochee River at Roswell, Cobb County, Georgia, N. A.
Pratt, Jr. and J. Postell) 1 Idiotype MCZ 146989.

productus Conrad, Unio: 1836, Monography Family Uuionidae, Part 3, p.
31, pi. 14, fig. 1 (Savannah River at Augusta [Georgia], M. Phillips)
3 Idiotypes MCZ 178692 from the Shenandoah River, Jefferson County,
West Virginia ex Dr. W. D. Hartmann.

propinquus Lea, Unio: 1857, PANSP, p. 88; 1862, Obs., 8, p. 67, pi. 5, fif;.
212 (Florence, Alabama, G. White; Tuscimibia, Alabama, L. B. Thorn-
ton) 4 Paratypes MCZ 178566 ex L. B. Thornton.

proximo Lea, Unio: 1852, Pr. APS, 5, p. 252; 1852, Obs., 5, p. 27, pi. 20,
fig. 27 (Georgia, J. P. Barratt) 2 Idiotypes MCZ 146981.

JOHNSON : TYPES OF NAIADES IN MCZ 131

pugio, Benson, Unio: 1862, Ann. Mag. Nat. Hist., (3) 10. p. 193; 1870,
Hanley and Theobold, Conch. Indica, p. 5, pi. 19, fig. 7 (In rcgione Ava
[Burma 1, W. Thooljold) Theobold sent only one worn right valve of
this species to Benson for description. Our specimens are from Theo-
bold's original lot. 3 Paratypes MCZ 175560 ex W. Theobold.

pulcher Lea, Unio: 1838, Tr. APS, 6. p. 6, pi. 3, fig. 6; 1838, Obs., 2, p. 6,
pi. 3, fig. 6 (near Nashville, Tennessee, Prof. Troost) Prof. Troost lent
Lea one specimen of this species for description. We have Troost 's
original lot with the label, "U. pulcher Lea, Harpeth Eiver, Tennessee.
Mr. Lea has I think improperly made from the varieties of this two
other species" (presumably U. menl'ianus and U. interruptus) . We
are unable to tell which specimen was the one figured. 6 Cotypes MCZ
5824 ex E. E. Call coll 'n.

puUatus Lea, Unio: 1856, PANSP, p. 262; 1858, Obs., 6. p. 57, pi. 8, fig. 39
(Creeks near Columbus, Georgia, Bishop Elliott) 1 Idiotype MCZ
178715.

puh'inulus Lea, Unio: 1845, Pr. APS, 4, p. 164; 1848, Obs., 4. p. 55, pi. 8,
fig. 24 (Tuscaloosa and Coosawattee River, Murray County, Alabama,
P,. W. Budd) 3 Idiotypes MCZ 178820.

punctatus Lea, Unio: 1865, PANSP, p. 89; 1869, Obs., 12. p. 21, pi. 32, fig.
76 (Caney Fork, Tennesseee, S. M. Edgar; Tnscumbia, Alabama, B.
Pybas) 1 Idiotype :MCZ 178897 from Tennessee.

pasilla Simpson, Quadnda granifera: 1900, Pr. U. S. Nat. Mus., 22. p. 795
(Green River, at Mammoth Cave, Kentucky) Lectotype, here selected,
MCZ 20176, PI. 2, fig. 4. This is the first time that this species has been
figured. Ex li. S. Frierson.

pusillus Lea, Unio: 1840, PANSP, p. 286; 1842, Obs., 3. p. 58, pi. 18, fig. 36
(Ogechee [Ogeechee] River, Georgia, Major Le Conte) 2 Idiotypes
MCZ 178766.

pyhasii Lea, Unio: 1858, PANSP, p. 40; 1862, Obs., 8. p. 71, pi. 6, fig. 216
(Tennessee River, Tuscumbia, Alabama, B. Pybas) 3 Idiotypes MCZ
178904.

pi/riformis Lea, Unio: 1857, PANSP, p. 31; 1858, Obs., 6, p. 69, pi. 12, fig.
50 (near Columbus, Georgia, Bishop Elliott) 2 Idiotypes MCZ 178808.

radians Lea, Unio: 1857, p. 32; 1859, Obs., 7. p. 19, pi. 23, fig. 84 (Othca-
looga Creek, Gordon County, Georgia, Bishop Elliott) 1 Idiotype MCZ
178906.

ratlulomts Drouet and Cliaper, Unio: 1892, Mem. Soc. Zool. France, 5, p.
150, pi. 5, figs. 10-12 (le Sebroeang a Borneo) 2 Paratypes MCZ 174096
ex S. Putzeys coll 'n.

132 BULIjETIX : MUSEIM OK CU.M I'AKATIVE ZOOLOOV

rat-cnsi.s Le;i, Unio: 1859, PANSP, p. 17J ; IHtJd, OIjs., 8. j). i:), i>l. o'i, tij;-. loG
(Chattahoochee [Rh'cr] near Columbus aiul Rae's Ci-eek, Georgia,
Bishop Elliott) 2 Idiotypcs .MCZ 124S81.

lavcncimiiHs Lea, Unio: 1834, Tr. APS, 5, p. 32, pi. 3, lig. o; 1834, Obs., 1,
p. 144, pi. 3, fig. 5 (French Broad River, tributary to the Tennessee,
near Asheville, North Carolina, Prof. Ravenel) 2 Idiotypes MCZ 17880G
from Alabama.

racinfellu^s Morelet, Unio: 1849, Testacea Novissima Insulae Cubaiiae, 1,
p. 29 (in Lacu Yzabal, reipublicae Guatemalensis) 1 Idiotj-pe MCZ
17870(5.

rcdusHs B. H. Wright, Unio: 1898, Nautilus, 11. p. Ill (Ockloeknee [Och-
locknee] River, Leon Co., Florida) 25 Paratypes MCZ 167685 from B. II.
Wright ex B. Walker coll 'n.

reevcianns Lea, Unio: 1852, Tr. APS, 10, p. 272, pi. 20, lig. 28; 1852, Obs., 5,
p. 28, pi. 20, fig. 28 (Alexandria, Louisiana, J. Hale and Prof. Powell)

1 Paratype MCZ 178918 ex Prof. Powell with tlie label ' ' T. Ecrveionu.s
Lea, Arkansas [sic] Lea so decided from this specimen and another,"

2 Idiotypes MCZ 178879.

!( sapinatus Von Martens, PN^'ududun.: 1902, Nach. Deutsch. Mai. Gesell., 34,
p. 131 (Than-Moi, Tonkin [French Indo-China], Fruhstorfer) Lecto-
type MCZ 167691 selected by Johnson, 1948, Nautilus, 62. p. 50, pi. 3,
'ig- 3; Vn Paratype MCZ 167672 ex B. Walker coll 'n. For a discussion
of this tjT3e designation see: Baker, II. B. 1950, Nautilus, 63. p. 128;
Johnson, R. I., 1950, Nautilus, 63, p. 125; Zilch, A., 1950, Archiv fiir
Molluskenkunde, 79, p. 87.

rhoinbic.a Anthony, Alasmidon: 1865, Anier. Jour. Conch., 1, p. 158, lA. 12,
fig. 5 (Michigan) Holotype MCZ 50296.

ricliardsoni Von Martens, Anodonta: 1900, Biologia Centrali Americana,
Moll., p. 529, pi. 41, figs. 1, la, 2, 2a (Rio Ameca, state of Jalisco,
Mexico, Richardson), 3 Cotypes MCZ 6808 ex F. D. Godman.

riograndensis Von Ihering, Anodonta: 1890, Arch. Naturgesch., 1, p. 154,
(Rio Grande do Sul, Brazil) 3 Cotypes MCZ 192370 ex S. Putzeys colFn.

rosaceiis Conrad, Unio: 1850, JANSP, 1, p. 275, pi. 37, fig. 5 (Savannah
River [Georgia], J. H. Couper) Holotype MCZ 178779 ex J. H. Couper.

ro.^ea Heude, A)iodo)i : 1881, Condi. Fleuv. Nanking, Part 7, pi. 50, fig. 93
(les environs de Fou-tcheou (Fou-kien) [China]) % Paratype MCZ
167249 ex Musee Heude.

JOHNSON : TYPES OF NAIADES IN MCZ 133

rostrum Lea, Umio : 1861, PANSP, p. 391; 1863, Obs., 9. p. 23, pi. 29,
fig. 270 (Davidson County, North Carolina, Dr. Genth) 1 Idiotype MCZ
178698.

rotula Pilsbry and Bequaert, Caelatura (Laevirostris ?) : 1927, Bull. Amer.
Mus. Nat. Hist., 53. p. 405, pi. 33, figs. 5 5a (Congo River at Leopold-
ville [Belgian Congo]) 2 Paratypes MCZ 86183. "Is a s^nionjan of
Caelatura (Laevirostris) hotirguignati de Roehebrune. " (Bequaert).

rugatu^ Menke, Unio: 1828, SjTiopsis Molluscorum, p. 90 (In Germaniao
rivis; in principatus Pyrmontani fluvio Emmcr, in principatus Sc-liaum-
burgo-Lippiaci rivo Aue, et in rivo Aue prope Hildesiam) 1 Cotype
MCZ 152876 from Germany, from C. Menke ex W. Dunker.

rutilan-s Lea, Vnio: 1856, PANSP, p. 262; 1858, Obs., 6. p. 59, pi. 9, fig. 41
(Othealooga Creek, Gordon County, and Columbus, Georgia, Bishop
Elliott) 3 Metatypes MCZ 178933 from Columbus, Georgia.

saccellus Drouet and Chaper, Vnio: 1892, Mem. See. Zool. France, 5. p. 148,
pi. 5, figs. 4-6 (le Sebroeang a Borneo) 1 Paratype MCZ 174095 from
Chaper ex 8. Putzeys coll 'n.

sacculus Anthony, Unio: 1865, Amer. Jour. Conch., I. p. 157, pi. 12, fig. 3
(Tennessee) Holotype MCZ 161898.

sagittiformis Lea, Unio: 1852, Tr. APS., 10, p. 277, pi. 22, fig. 35; 1852,
Obs., 5, p. 33, pi. 22, fig. 35 (Oconee River, near Athens, Georgia,
Major Le Conte) 8 Idiotypes MCZ 178687.

salehrosus Lea, Unio: 1859, PANSP, p. 170; 1860, Obs., 8, p. 14, pi. 52,
fig. 157 (Flat Rock Creek [and] Bull Creek [Georgia], G. Hallenbeck ;
Chattahoochee River, near Columbus, Georgia, Bishop Elliott) 3 Idio-
types MCZ 150812.

sanctorunijohanium B. H. Wright, U7iio (Elliptio) : 1933, Nautilus, 47. p. 17,
pi. 1 (Lake Druid, near Floral City, Florida) 2 Paratypes MCZ 81972
ex B. H. Wright.

satnr Lea, Unio: 1852, Pr. APS., 10. p. 252; 1852, Obs., 5. p. 21, pi. 17,
fig. 19 (Alexandria, Louisiana, J. Hale; Lake Calcashue, near New
Orleans [Louisiana], J. G. Anthony) 1 Paratype MCZ 53862 with the
label, "Amite River, Mississippi, Mr. Lea drew up his description of
the species partly from my specimen." (J. G. Anthony.)

sarannahensis Lea, Unio: 1857, PANSP, p. 169; 1858, Obs., 6, p. 81, pi.
16, fig. 61 (Savannah River, also Brantley's Mill, Washington County,
Georgia, G. White; Santee Canal, South Carolina, Dr. Ravenel; and
Sugar Creek, Mecklenburg County, North Carolina, C. M. Wheatley) 1
Paratype MCZ 178729 ex C. M. Wheatley.

lo4 BULLETIX : MUSKIM OF COMPARATIVE ZOOLOGY

.•icluidei Marshall. Anodanlitcn : 1934, Jour. Washington Acad. Sci., 24. p.
80, figs. 4 6 (Tubicuary River, at Aro.ja, southorn Paraguay, F. Sehadej
1 Taratype MCZ 62479 ex F. Sehade.

.schicerzcnhaclii 'Parreyss' Bourguigiiat, I'liio: ISuti, Rev. et Mag. de Zool.,
(2) 8. p. 75, pi. 8, figs. 1-5 (Brousse, Anatolic, [Turkey] ) 4 Cotypes MCZ
177313 from L. Parreyss ex C. B. Adams coll'n.

scitulus Lea, Unio: 1860, PANSP, p. 93; 1860, Obs., 8, p. 24, pi. 55, fig. 167
(Tuscunibia, Alabama, L. B. Tliornton ) 1 Idiotype MCZ 178913.

dcobinatus Lea, Umo: 1856, PANSP, p. 93; 1857. Obs., 6, p. 19, pi. 26, fig.
13 (Siani, T. R. Ingalls and 8. R. House) 3 Para types MCZ 175555
ox I. Lea.

scittiim Sowerby, Unio: 1868, Conchologia Iconiea, 16, pi. 94, fig. 510
(Tcnasserim [Burma] Benson, teste Thcobold) 1 Parat.vpe MCZ
175548 ex W. Tlieobold.

xccundus Heude, Psc^^(lo^lon : 1877, Conch. Fleuv. Nanking, Part 3, pi. 18,
fig. 38 (La riviere Hoai [China] dans la partie inoyenne de laquelle
il abonde) 2 Paratypes MCZ 167242 ex Musee Heude.

semlfjranosus Von dem Busch, Unio: 1845 [in Philippi] Abbild. und Besehr.
Conch., 1. p. 19, pi. 1, figs. 1-3 (Flumen prope Tampico, Mexico) 2
Paratypes MCZ 152861 ex 0. Yon dem Busch.

semirasa Pilsbry, Lampsilis : 1909 [1910], PANSP, p. 534, pi. 26, figs. 7, 8,
9 (Ganina River, 3 miles southwest of San Diequito . . . also in Valles
River, tvro miles above Mecos [Mexico], A. A. Hinkley) 20 Cotypes
MCZ 189370 labeled, "Mexico," from A. A. Hinkley ex B. Walker
coll'n.

shepardiamis Lea, Unio: 1834, Tr. APS, 5, p. 95, pi. 13, fig. 38; 1834, Obs., I.
pi. 13, fig. 38 (Hopeton, near Darien, Georgia, Prof. Shepard) 3
Paratypes MCZ 155568 ex Prof. Shepard.

showalterii Lea, Anodonto : 1860, PANSP, p. 307; 1863, Obs., 9, p. 37, pi. 33,
fig. 284 (Coosa River, Wetumpka, Alabama, E. R. Showalter) 1 Para-
type MCZ 178772 ex E. R. Showalter.

showalterii Lea, Unio: 1860, PANSP, p. 307; 1862, Obs., 8, p. 77, pi. 8,
fig. 223 (Coosa River, Wetumpka, Alabama, E. R. Showalter) 3 Para-
types MCZ 178920 mentioned by Lea as beLiig in the J. G. Anthony
coll'n. 1 Paratype MCZ 178818 ex E. R. Showalter.

shurtleffianm Lea, Unio: 1856, PANSP, p. 94; 1857, Obs., 6. p. 22, pi. 27,
fig. 17 (Sina River, India, Major Le Conte; Ahmednugger, India, S.
Shurtleff) 1 Paratype MCZ 175566 ex S. Shurtleff.

.lolI.VSOX: TYl'KS OK XAIADES IK M<Z 135

sif/nata Pilsbry, Lampsdls: 1909, PANSP, p. 536, pi. 25, figs. 5, 6; pi. 26,
figs. 5, 6 (Valles River, noiir a snuill falls about 3 miles below Valles,
Mexieo, A. A. Hinliley) 10 Colypos MCZ 192367 ex A. A. Hinkley.

srmiUimu.s Urtiiianu, Diplodori: 1921, Mem. Carnegie Mus. [Pittsburgh, Pa.],
8. p. 495, pi. 35, figs. 3-6; [gills] pi. 45, fig. 2; [gloehidia] text fig. 4c
(p. 469) (Rio Nhuiuliaquara, Morretes, Paraaxa, Bra/.il, J. D. Ilaseman)
2 Cotypes MCZ 47050 ex Carnegie Musenin.

Hmplex Lea, Unio: 1845, Pr. APS, 10. p. 163; 1848, Obs., 4. p. 50, pi. 5,
fig. 15 (Black Warrior River, Alabama, B. W. Budd) 1 Tdiotype MCZ
178834.

sitnus Lea, Vnio: 1838, Tr. APS, 6, p. 26, pi. 8, fig. 20; 1838, Obs., 2. p. 26,
1)1. 8, fig. 20 (Cuiiibeiland Kiver. Tennessee, Prof. Troost) 5 Idiotypes
MCZ 178903.

sinalueusis Crosse and Fischer, Anodonia f/lauca: 1883, Jour, de Conch., 31,
p. 219 (Rio Presidio in provineia Sinoloa dicta, Reipublicae Mexicanae,
A. Forrer) 1 Cotypc MCZ 87540 from A. Forrer, ex Boston See. Nat.
Hist.

."linuosa Morrison, Guianadetunu : 1943, Nautilus, 57, p. 46, pi. 8 (Cuyuni
River, opposite Kartabo Point, near its junction with the Mazaruui
River, Essequibo District, Britisli Cuiana) 1 Paratype MCZ 144796
ex U. S. Nat. Mus.

KomemU 'Moores' Call, Jnodonta: 1880, American Naturalist, 14. p. 530
[Monday Creek, Carbon Hill, Hocking County, Ohio] 3 Cotypes MCZ
158090 [nonun iiadu))i\ ex R. E. Call coll'n.

sfliverbiunus Lea, Unio: 1833, Tr. APS, 4. p. 68, pi. 10, fig. 28; 1834, Obs., 1,
p. 180, pi. 10, fig. 28 (Tennessee, G. B. Sowerby) 4 Idiotypes MCZ
178886 from Caney Fork River, Tennessee.

spadictius Lea, I'niu: 1857, PANSP, p. 86; 1862, Obs., 8. p. 59, pi. 1, fig.
201 (Deep River, Gulf, North Carolina, Prof. Emmons; and mountain
streams. North Carolina, .T. Clark) 3 Idiotyi>es MCZ 178714.

spatalatus Lea, Unio: 1845, Pr. APS, 4, p. 164; 1848, Obs., 4, p. 54, pi. 8,
fig. 22 (Rock River, Wisconsin, Capt. Maryatt and B. W. Budd) 5
Idiotypes MCZ 17S835 from Michigan.

spillmanii Lea, Margaritana: 1858, PANSP, p. 138; 1862, Obs,, 8, p. 109,
id. 17, fig. 252 (Tombigbee Rivei', near Columbus, Mississippi, W.
Spillman) 1 Paratype MCZ 6395 from Spilhuan ex R. E. Call coll'n.

1 36 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

spinosus Lea, Unio: 1836, Dese. of New Sp. TJnio [colored figs.] ; 1838, Obs.,
2. p. 57, pi. 16, fig. 50 (Altamaha [River] Hopeton, near Darieii,
(jeorgia, J. H. Couper ; Altamaha [River] Liberty County, Georgia, L.
Le Conte) 5 Paratypes MCZ 87951 from J. H. Couper ex Boston Soc.
Nat. Hist.

spinosus Simpson, Shistodesmns : see under vestitus var. b Heude, Unio.

spissus Lea, Unio: 1859, PANSP, p. 112; 1859, Obs., 7, p. 26, pi. 25, fig. 89
(Satilla River, Wayne County. Georgia, T. C. Do^sniie) 4 Meta types
MCZ 178707.

splendidm Lea, Lampsilis : 1838, Tr. APS, 6. p. 70, pi. 19, fig. 61; 1838, Obs.,
2. p. 70, pi. 19, fig. 61 (Altamaha River, near Darien, Georgia, J. H.
Cowper [sic] ) 4 Paratypes MCZ 165692 from J. H. Couper, ex Boston
Soe. Nat. Hist.

squameus Lea, Unio: 1861, PANSP, p. 391; 1863, Obs., 9. p. 22, pi. 28,
fig. 269 (North Carolina, J. G. Anthony) 1 Paratype MCZ 124889
ex J. G. Anthony.

stagnalis Conrad, Unio: 1849, PANSP, p. 152; 1850, JANSP, L p. 275,
pi. 37, fig. 2 (Lihabits mill ponds; Ogeechee River, Georgia, .T. H.
Couper) Holotype MCZ 178778 ex J. 11. Couper.

stanleyvillensis Pilsbry and Bequaert, Caelatura: 1927, Bull. Amer. Mus.
Nat. Hist., 53, p. 403, pi. 32, figs. 3, 3a, 4; pi. 33, fig. 3. (Congo River
at Stanleyville [Belgian Congo], Lang, Chapin, and J. Bequaert) 1
Paratype MCZ 142354. " Is a synonym of Caelatura (Zairia) elegans
de Rochebrune. ' ' (Bequaert.)

stapes Lea, Uriio: 1831, Tr. APS, 4. p. 77, pi. 7, fig. 8; 1834, Obs., 1. p. 87,
pi. 7, fig. 8 (Alabama River [Alabama], Judge Tait) 2 Idiotypes MCZ
178789.

strwardsonii Lea, Unio: 1852, Tr. APS, 10, p. 278, pi. 23, fig. 36; 1852, Obs.,
5, p. 34, pi. 23, fig. 36 (Chattanooga [sio'] River, Tennessee, T. Steward-
son) 6 Idiotypes MCZ 178877.

straitissimus Anthony, Unio: 1865, Amer. Jour. Conch., 1, p. 155, pi. 12,
fig. 1 ([Clinch River] Tennessee) Holotype MCZ 17356.

striatus Lea, Unio: 1840, Pr. APS, 1, p. 287; 1842, Obs., 3, p. 41, pi. 12,
fig. 16 (Chattahoochee River, Columbus. Georgia, Dr. Boykin) 4 Idio-
types MCZ 178800.

.strodeanus B. H. Wright, Unio: 1898, Nautilus, 12. p. 5 (Escambia River, W.
Florida) 6 Paratypes MCZ 210277 from B. H. Wright ex B. Walker
coll 'n.

.lOllXSOX: TYl'KS OF XAIADES IX MCZ 137

strumosm Lea, Vnio: 1872, PANSP, p. 158; 1874, Obs., 13. p. iil, pi. 19,
fig. 00 (Yadkin Kiver, North Carolina, C. M. Wheatloy ) 1 I'aratype
MCZ 30403 ex C. M. Wheailey.

tinbaiiguldta Anthony, Anodon: 1865, Amor. Jour. Conch., 1. p. 158, pi. 13,
fig. 1 ([Shears Lal<c', Kent County | Michigan) llolotype MCZ 150645.

t:iihri<is,s<i Lea, Anodonta : 1850 [1851], Pr. Zool. iSoc. London, p. 198; 1860,
Ohs.. 7, p. 54, pi. 33, tig. 115 (Laguna de Bai, Luzon. Philippines, II.
CiiMiing) '2 Paratypes ]MCZ 175579 ex TI. Cuming.

subcrocem Conrad, Unio: 1854, JANSP, 2. p. 297, pi. 27, fig. 1 (One of the
tri))utaries to Canadinn River, Arkansas, Dr. Woodhou.se) 1 Idiotype
MCZ 1 78693.

subfUipsis Lea, Vnio: 1856, PANSP, p. 262; 1858, Obs., 6. p. 62, pi. 10,
fig. 44 (Creeks near Colunil)us, Georgia^ Bishop Elliott) 2 Idiotypes
MCZ 178934.

f~iibflavu.^ Lea, Unio: 1857, PANSP, p. 170; 1858, Obs., 6. p. 90, pi. 19,
fig. 70 (Walnut Creek, above Macon, Georgia, Bishop Elliott and J. C.
Plant) 2 Tdiotypes MCZ 72523.

sub(/il)bom Anthony, Anodon: 1866, Amer. Jour. Conch., 2, p. 144, pi. 6,
fig. 2 (Bhick Lake, Michigan) llolotype MCZ 161854.

mtb-inflaia Antliouy, Anodon: 1865, Amer. Jour. Conch., 1, p. 160, pi. 15,
fig. 1 (Michigan) Holotype MCZ 161863.

sKblatu.^ Lea, Vnio: 1857, PANSP, p. 169; 1858, Obs., 6. p. 82, pi. 16, fig.
62 (Ucliee Bar, liclow Columbus, Geoi'gia, Bishop Elliott) 2 Idiotypes
MCZ 124882.

auccineus Heude, Mycetopus: 1885, Conch. Fleuv. Nanking, Part 9, pi. 70,
fig. 134 (Les rivieres de Lu teheou fou et de Tong-tch'eng vers leur
embouchure dans le lac Tch'ao, province de Ngan-houe [China]) 1
Paratype MCZ 167109 ex Musee Heude.

sumatnnsia Lea, Vnio: 1859, PANSP, p. 153; 1860, Obs., 7. p. 57, pi. 34,
fig. 118 (Sumatra, H. Cuming) 2 Paratypes MCZ 175563 ex H. Cuming.
Tlie name was changed to dimotus l»,v Lea 1870, because sumatrensis was
preoccupied for a Vnio Ijy Dunker.

superbus Lea, Vnio: 1843, Desc. 12 Uniones [no pagination]; 1848, Obs., 4,
p. 39, pi. 42, fig. 11 (New Holland [Australia], J. C. Jay) 2 Idiotypes
MC^ 175551 from Siam.

auperiorensis Marsh, Vnio: 1897, Nautilus, 10, p. 103, pi. 1, figs. 1. 2, 5
(Michipicoton River, upper shore of Lake Superior, Canada, J. H.
Ferris) 1 Cotype MCZ 20169 from J. H. Ferris ex L. S. Frierson.

138 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

suttoni B. H. Wright, Vnio: 1897, Nautilus, 11. p. 56 (Laie near Candkr,
Marion County, Florida, B. Sutton) 2 Paratypes MCZ 20172 from
B. H. Wright ex L. S. Frierson.

siimmetricus Lea, Vnio: 1845, Pr. APS, 4, p. 164; 1848, Obs., 4. p. 47,
])1. 4, fig. 11 (Red River, at Alexandria, Louisiana, J. Hale; 2 Idiotypes
MCZ 178732.

taitianm Lea, Vnio: 1834, Tr. APS, 5. p. 39, pi. 4, fig. 11; 1834, Obs., 1,
]). 151, 1)1. 4, fig. 11 (Alabama River [Alabama], Judge Tnit) 1
Idiotype MCZ 178811.

tennesseensis Frierson, Strophitus undulatws: 1927, Check list N. American
Naiades, Waco, Texas, p. 22 (Headwaters of the Tennessee River j.
Lectotype, here selected, MCZ 16690, PI. 2, fig. 1. The original descrip-
tion is here repeated, "A form found in the headwaters of the Tennes-
see River, very thin, much compressed, widely biangul.nted behind,
yellow inside and out," ex L. S. Frierson.

hniii^ Lea, Anodonta : 1850 [1851] Pr. Zool. Soc. London, p. 198; 1860,
Obs., 7, p. 55, pi. 33, fig. 116 (Sual, Luzon, Philippines, H. Cuming)
4 Paratypes MCZ 175578 ex H. Cuming.

tcnuiseulus Frieison, Vnio: 1911, Nautilus, 25. p. 29, pi. 1, figs. 4-6 (Reedy
Lake, Polk County. Florida) 3 Cotypes MCZ 20180 ex L. S. Frierson.

trtricm Lea, Vnio: 1857, PANSP, p. 170; 1859, Obs., 7, p. 13, pi. 22, fig.
78 (Flint River, near Alliaiiy, Georgia, Bishop Elliott) 2 Idiotypes
MCZ 178722.

/f./(/.sr/(A/.s- Lea, Vnio: 1860, JANSP, 4. p. 359, pi. 61, fig. 184; 1860, Obs., 8,
p. 41, pi. (il, fig. 184 ( Dewitt County, Texas, W. Xewcomb) 4 Paratypes
MCZ 178921 ex W. Newconib.

Ouianumi Sehlesch, Anodonta piscinalis: 1907, Ann. Soc. Roy. Mai. Belgique,
(2) 42. p. 161 (in Zwei Teichenhinter "Gl. Carlsberg" in valby bei
Kopenhagen [Denmark]) 9 Cotypes MCZ 192022 ex D. Thaanum eoll 'n.

thornton.it Lea, Vnio: 1857, PANSP, p. 83; 1867, Obs., IL p. 42, pi. 14,
fig. 36 (Tuscumbia, .Alabama, L. P.. Thornton) 8 Paratypes MCZ
178809 ex L. B. Tliornton.

tinheri B. H. Wright, Vnio: 1899, Nautilus, 13, p. 7 (Tonibigbee River
[Moscow] Alabam;! ) 2 Paratypes MCZ 20165 ex L. S. Frierson.

t(iiiit)i[/htaniini Frierson, Fh urobema : 1908, Nautilus, 22, p. 27, pi. 3, figs. 3, 4
(Tombigbee River, Deiuopolis, Marengo County, Alabama; Columbus.
MississipiH ) 2 Idiotypes MCZ 178824 from the Alabama River, Selnia.,
Alabama.

.lOlINSON : TYPES OV NAIADES IN MCZ 139

tnwhiiibrcn.six Lon, Margaritana : 1858, PANSP, p. 138; 1862, Oba., 8. p.
Ill, pi. 18, fig. 2.15 (Toiuhigbee Kivor, near Columbus, Mississippi, W.
.Spillmnu) 1 Paiatype MCZ 1520.37 ex W. Spillman.

tuitivus Lea, Vnio: 1840, Pr. APS, 1, p. 287; 1842, Oba., 3. p. 42, pi. 12,
fig. 17 (Chattahoochee River, Columbus, Georgia, Dr. Boykin) .3 Idio
types MCZ 178688.

iriangulata Lea, Margaritana: 1858, PANSP, p. 138; 1859, Obs., 7, p. 46,
111. 32, fig. Ill (Upper Chnttnhooi-hee, Georgia, Bishop Elliott; Colum-
bus, Georgia, Dr. Boykiu and J. Postell ; Polato County, Georgia, G.
White; Sawney's Creek, South Carolina, Dr. Blandiug) 3 Paratypes
MCZ 165696 from Columbus, Georgia, from the E. E. Mayo eoH 'n ex
Boston Soc. Xat. Hist.; 2 Idiotypes MCZ 150661.

tridavus Heude, JJnio: 1877, Conch. Fleuv. Nanking, Part 2, pi. 10, figs. 21,
21a (La riviere Siang (prov. de IIou-Nan) [China]) 1 Paratype MCZ
167111 ex Musee Ileude.

tricmholuj^ Benson, Unio: 1862, Ann. Mag. Nat. Hist., (3) 10. p. 190,
(Ihuiiina Kamgunga, prope Moradabad [India]) 3 Cotypes MCZ 175558
ex W. H. Benson.

friiimphans B. H. Wright, U-nio: 1898, Nautilus, 11. p. 101 (Coosa River, St.
Clair County, Alabama) 2 Paratypes MCZ 139625 ex B. Walker coirn.

trochelianus Lea, Unio: 1852, Tr. APS, 10. p. 280, pi. 23, fig. 39; 1852,
Obs., 5. p. 36, pi. 23, fig. 39 (Coosawatte River, Murray County, Georgia,
Dr. Boykin) 1 Idiotype MCZ 178810.

troinpi. Drouct and Chaper, Unio: 1892, Mem. Soe. Zool. France, 5. p. 153, pi.
6, figs. 8-10 (le Kapoeas (Semitau) a Borneo) 1 Cotype MCZ 174098
ex S. Putzeys coll 'n.

troostensis Lea, Unio: 1834, Tr. APS, 5. p. 71, pi. 10, fig. 30; 1834, Obs., 1.
p. 183, pi. 10, fig. 30 (Cumberland River [Tennessee], Prof. Troost) 3
Idiotypes MCZ 178922.

truncata B. H. Wright, Margaritaim marginata: 1898, Nautilus, 11. p. 124
(It occurs in eastern as well as western waters and into Virginia and
Tennessee) 4 Cotypes MCZ 152253 from Greyson County, Virginia
ex B. H. Wright.

tuberosm Lea, Unio: 1840, Pr. APS, 1. p. 286; 1842, Obs., 3, p. 48, pi, 14,
fig. 25 (Cany Fork and Cumberland Rivers, Middle Tennessee, S. M.
Edgar) 2V^ Paratypes MCZ 6205 from the Cumberland River, men-
tioned by Lea as being in the Troost coll'n., ex R. E. Call coll'n.

140 BULLETIN- : MUSEUM OF COMPARATIVE ZOOLOGY

tumidus Morelet, Monocondyhis : 1866, Jour, de Coneh., 14, p. 62 (In torren-
tibus montanis Cambodiae) ; 1875, Series Conehologique Part 4, p. 337,
pi. 16, fig. 1; type figured. Jour, de Conch., 40, pi. 3, figs. 42-44. 3 Para-
types MCZ 175576 ex A. Morelet.

tuomeyi Lea, Unio: 1852, Tr. APS, 10. p. 256, pi. 13, fig. 4; 1852, Obs., 5,
p. 12, pi. 13, fig. 4 (Abbeville District, South Carolina, J. P. Barratt)
1 Idiotype MCZ 178699.

tiircicus 'Parreiss' lsiG'\ Kuster, Unio: 1862, Conch. Cabinet, (2) 9. Part
2, p. 267, pi. 90, figs. 3-4 (Kleinasien [Turkey] ) 2 Cotypes MCZ 152877
from L. Pprreyss ex C. B. Adams coll 'n.

turgidulus Lea, Unio: 1858, PANSP, p. 40; 1862, Obs., 8. p. 66, pi. 5, fig.
211 (Cumberland Eiver, Prof. Troost and T. C. Downie; Florence, Ala-
bama, G. White) 3 Idiotypes MCZ 178565.

tiirgidus Lea, Unio: 1838, Tr. APS, 6. p. 11, pi. 5, fig. 11; 1838, Obs., 2.
p. 11, pi. 5, fig. 11 (near New Orleans [Louisiana], Mr. Barabino) 2
Idiotypes MCZ 178832.

umbrans Lea, Unio: 1857, PANSP, p. 104; 1858, Obs., 6, p. 72, pi. 13, fig.
53 (Othcalooga Creek, Gordon County, Georgia, Bishop Elliott) 2
Idiotypes MCZ 178874.

urnbrosa Lea, Unio: 1856, PANSP, p. 95; 1857, Obs., 6, p. 31, pi. 30, fig. 26
(Mendellin Eiver [near Vera Cruz] Mexico, Dr. Burrough) 1 Paratype
MCZ 178872 from Dr. Burrough, mentioned by Lea as being in the
J. G. Anthony coll 'n.

undivaga Pilsbry, Unio: 1909, PANSP, p. 536, pi. 26, figs. 1-4; pi. 27,
fig. 5 (Valles Eiver, two miles above Mecos in a muddy bank just above
the rapids, A. A. Hinkley) 3 Cotj-pes MCZ 30633 from A. A. Hinkley ex
B. Walker coll 'n.

unicolor Lea, Unio: 1845, Tr. APS, 4, p. 163; 1848, Obs., 4, p. 48, pi. 4,
fig. 12 (Tuscaloosa, Alabama, W. B. Budd) 6 Idiotypes MCZ 178909
from the Pearl Eiver, Alabama.

unicostatus B. H. Wright, Unio: 1899, Nautilus, 13. p. 69 (Spring Creek,
Decatur County Georgia) 2 Paratypes MCZ 20150 ex L. S. Frierson.

upsoni Marsh, Unio: 1880 "Eead before the Mercer County, Illinois Scien-
tific and Historical Society March 2, 1880. Mr. Marsh had copies of
the description published and thinks that the species was described on
the records. It was not figured," Simpson, 1900, Pr. U. S. Nat. Mus.,
22, p. 539; Marsh, 1887 Conch. Exch., 1. p. 51 (Kishaeaukee Eiver,
Winnebago County, Illinois) 1 Cotype MCZ 5551 from W. A. Marsh
ex E. E. Call coll 'n.

■lOIINSON : TYPES OF NAIADES l.\ MCZ 141

rnniuemoisL^ Leu, Zhiio: 1838, Tr. APS, 6. p. 30, pi. 11, fig. 31; 1838, Obs.,
2, p. 36, pi. 11, fig. 31 (Cumljerland River, Tennessee, Prof. Troost) 1
Tdiotype MCZ 178888.

rrii list lis Lea, U-nio : 1838, Tr. APS, 6, p. 4, pL 2, fi}--. 4; 1838, Obs., 2.
p. 4, pi. 4, fig. 4 (Potosi. Missouri, .John Pony; Cincinnati, Dliio, T. (J.
Lea) 5 Idiotypes MCZ 178880.

rents Lea, Uiiio: 1860, PANSP, p. 140; 1862, Obs., 8, p. 87, pi. 11, fig. 231
(Cahawba River, Perry County, Alabama, E. R. Showalter) 1 Paratype
MCZ 178798 mentioned by Lea as being in the J. G. Anthony t-oll'n.

(•(.v/i7«N var. a Ileude, L'nio: 1883, Conch. Fleuv. Nanking, I'art 8, pi. 58, fig.
112 (La riviere de Ning-kouo fou, celle de T'sing-yang Men (Ngan-
houe) [China]) 1V> Paratypes MCZ 167241 ex Musee Heude. This
name was preoccupied by Lea for a Vnio, therefore Simpson, 1900 Pr.
U. S. Nat. Mus., 22, p. 801 changed it to Quadnda ovata.

VI. si il IIS v;ir. h Heude, I'lua: 1883, Conch. Fleuv. Nanking, Part 8, \t\. ')9,
fig. 115 (La riviere de Ning-kouo fou, celle de T'sing-yang hieii
(Ngan-houe) [China]) l^/o Paratypes MCZ 167245 ex Musee Ileude.
This name was preoccupied by Lea for a Unio, therefore Simpson, 1900,
Pr. U. S. Nat. Mus., 22, p. 804 changed it to Shistodcsmus spinnsus.

liUosa B. 11. Wright, Unio: 1898, Nautilus, 12, p. 32 (Suwannee River,
Suwannee Co., Florida) 1 Paratype MCZ 186836 ex B. II. Wright.

virescens Lea, Unio: 1858, PANSP, p. 40; 1860, Obs., 8, p. 23, pi. 55, fig.
166 (Tennessee River at Tuscumbia, Alabama, B. Pybas) 6 Paratypes
MCZ 178829 ex Pybas and Thornton, mentioned by Lea as being in
lioth collections.

vondenbushiana [sic] Lea, Margaritana : 1840, Pr. APS, 1. p. 288; 1842,
Obs., 3. p. 60, pi. 18, fig. 39 (Java, G. Von dem Busch) 2 Paratypes
MCZ 175596 ex G. Von dem Busch.

ivalkeri B. H. Wright, Unio: 1897, Nautilus, 11, p. 91; figured by Simpson.
1900, PANSP, p. 77, pi. 1, fig. 5 (Suwannee River, Madison County,
Florida) 2 Paratn^es MCZ 20178 ex L. S. Frierson.

icalpolei Hanlej', Monocondylaea : 1871, Pr. Zool. Soc. London, p. 587
(Sarawak, Borneo) Lectotype MCZ 175577 selected by Johnson, 1948,
Nautilus, 62. p. 49, pi. 3, fig. 2 ex R. F. Geale.

ualtoni B. H. Wright, Unio: 1888, PANSP, p. 114, pi. 2, fig. 3 (Lake Wood-
ruff, Volusia County, Florida) 7 Paratypes MCZ 167701 from B. II.
Wright ex B. Walker coll 'n.

tcardiana Lea, Anodorita: 1838, Tr. APS, 6. p. 46, pi. 14, fig. 42; 1838, Obs.,
2, p. 46, pi. 14, fig. 42 (Near Chilieothe, Ohio, J. C. Ward) 1 Paratype
MCZ 37229 ex J. C. Ward.

142 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

wardii Lea, Unio: 1861, PANSP, p. 392; 1863, Obs., 9, p. 9, pi. 24, fig. 257
(Walhonding River, Ohio, J. C. Ward; Wassepinicon River, Iowa, Dr.
Foreman ; Coal River, Logan County, Virginia, Dr. Hartmann and Mr.
Anthony) 1 Paratype MCZ 178774 from Dr. Hartmann ex J. G.
Anthony. ' ' Tlie specimen f lom Mr. Anthony is marked ' Tennessee, '
but Dr. Hartnianu, on seeing this specimen, thinks he recognizes it
as the specimen which he gave Mr. Anthony from the same habitat
(Coal River, Virginia) with his own." (Lea).

n-atersoni Tomlin, Anodonta: 1923, Jour, of Conch., 17, p. 68, (Lake Beschik,
Salonika [Greece]) 2 Paratypes MCZ 78226 ex J. R. Le B. Tomlin.

inhbian.us B. H. Wright, Unio (Ellipfio) : 1934, Nautilus, 47, p. 94, pi.
10, figs. 1-2 (Lake Consuelo, Citrus County, Florida, near Floral City)
2 Paratypes MCZ 95548 ex B. H. Wright.

websterii B. H. Wright, Unio: 1888, PANSP, p. 113, pi. 2, fig. 2 (Lake
Woodruff, Volusia County, Florida) 1 Paratype MCZ 178765 ex B. II.
Wright.

ivheatleyi Lea, Anodorita: 1852, Tr. APS, 10, p. 287, pi. 26, fig. 49; 1852,
Obs., 5, p. 43, pi. 26, fig. 49 (Para [Brazil] South America, C. M.
Wheatley) 3 Paratypes MCZ 192020 ex C. M. Wheatley.

wheatleyi Lea, Monocondylaea: 1862, PANSP, p. 176; 1863, Obs., 10, p. 35,
pi. 50, fig. 307 (River Tigris, Assyria [Iraq], C. M. Wheatley) 1 Para-
type MCZ 178755 ex C. M. Wheatley.

icheatleyi Lea, Unio: 1857, PANSP, p. 85; 1860, Obs., 8, p. 58, pi. 1, fig.
200. [The name was herein changed by Lea to Unio catawhcnsis, wheat-
leyi being preoccupied.] (Catawba River, Gaston County, North Carolina,
C. M. Wheatley) 1 Idiotype MCZ 20166 from Ashley's Creek, Mecklen-
berg County, North Carolina. From C. M. Wheatley ex L. S. Frierson.

luheeleri Ortmann and Walker, Arhansia: 1912, Nautilus, 25, p. 98, pi. 8
(Old River, Arkadelphia, Arkansas, H. E. Wheeler) 1 Paratype MCZ
135712 from H. E. Wheeler ex Boston Soc. Nat. Hist.; 1 Paratype
MCZ 23319 from H. E. Wheeler ex L. S. Frierson.

looodwardimius Lea, Unio: 1859, JANSP, 4. p. 199, pi. 23, fig. 82; 1859,
Obs., 7, p. 17, pi. 23, fig. 82; [anatomy] pi. 29, fig. 103 (Etowah and
Connasauga Rivers, Cass County, Georgia, Bishop Elliott and G. White)
5 Idiotypes MCZ 192019.

zeiglerianm Lea, Unio : 1838, Tr. APS, 6. p. 32, pi. 10, fig. 27 ; 1838, Obs., 2,
p. 32, pi. 10, fig. 27 (Cumberland River, Tennessee, Prof. Troost) 2
IdiotyiDes MCZ 178893.

^onatm Heude, Unio: 1883, Conch. Fleuv. Nanking, Part 8, pi. 61, fig. 120
(La riviere de Ning-kouo fou, au sud de la ville [China]) 1 Paratype
MCZ 167112 ex Musee Heude.

PLATES

PLATE 1

Fig. 1. Anodonia hellua Morelet. Laeu Toui-Sap, Cambodia. Lectotype
MCZ 175610. Natural size.

Fig. 2. Unio holli Call. Colorado River [Austin] Texas. Lectotype MCZ
5017. Natural size.

PLATE 1

PLATE 2

Fig. 1. Strophitus undulatus tennesseensis Frierson. Headwaters of the
Tennessee River. Leetotype MCZ 16690. Natural size.

Fig. 2. Unio ascia 'Benson' Hanley. Penang [Straits Settlements].
Leetotype MCZ 17046. Natural size.

Fig. 3. Unio cor Conrad. Elk and Flint Rivers, Alabama. Holotype MCZ
178792. Natural size.

Fig. 4. Quadrula granifera puaUla Simpson. Green River at Mammoth
Cave, Kentucky. Leetotype MCZ 20176. Natural size.

PLATE 2

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. 115, No. 5

PSEUDEIilYS SCRIPT A CALLIBOSTBIS FROM

VENEZUELA WITH A GENERAL SURVEY

OF THE SCRIPT A SERIES

By Ernest Williams

With Three Plates

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

September, 1956

Publications Issued by or in Connection

WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

Bulletin (octavo) 1863 — The current volume is Vol. 115.

Breviora (octavo) 1952 — No. 61 is current.

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

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

Occasional Papers of the Department of Mollusks (octavo) 1945 —
Vol. 1, no. 18 is current.

Proceedings of the New England Zoological Club (octavo) 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.

The continuing publications are issued at irregular intervals in numbers
which may be purchased separately. Prices and lists may be obtained on
application to the Director of the Museum of Comparative Zoology,
Cambridge 38, Massachusetts.

Of the Peters "Check List of Birds of the World," volumes 1-3 are out
of print; volumes 4 and 6 may be obtained from the Harvard University
Press ; volumes 5 and 7 are sold by the Museum, and future volumes will be
published under Museum auspices.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. 115, No. 5

PSEUDEMYS SCRIPTA CALLIROSTRIS FROM

VENEZUELA WITH A GENERAL SURVEY

OF THE SCRIPTA SERIES

Bv Ernest Williams

With Three Plates

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

September, 1950

No. 5 — PsEUDEMYs scRiPTA CALLiROSTRis from Venezuela with
a general surviy of the scripta series

By Ernest Williams

In 1947 Benjamin Shreve reported a juvenile Pseudemys from
the Falcon district of Venezuela, assigning it to the scripta
group but not further identifying it. Re-examination of this
specimen has revealed that it is referable to callirostris Gray
and is the most easterly record to date of this form.

In the course of identifying this individual I have looked into
the scripta series as a whole and I will therefore present a few
comments here on the overall picture. This portion of the discus-
sion is intended to be suggestive rather than definitive. A rather
full record in regard to callirostris is included since there is no
adequate account in English.

TAXONOMIC HISTORY

John Edward Gray in 1855 described Emys callirostris —
"The Beautiful Beaked Emys" — from a single specimen,
stuffed, half-grown, presented by the Haslar Hospital with the
locality ''America." Gray furnished in his plate Xllb an ex-
cellent figure of dorsal and ventral views of the type showing
the "symmetrical rings" on the chin which were the most dis-
tinctive feature of the form, and the occasion of the specific name.

In his description of the type. Gray recognized the close rela-
tionship of this form to "Eiiiys ornata" and in 1863 (again in
1870) placed callirostris in the genus Callichelys of which ornata
was the type.

Giinther (1885) was apparently the first to suggest that cal-
lirostris may have come from the "West coast of Central Amer-
ica." Cope (1887) followed, listing Pseudemys callirostris from
Central America. Boulenger (1889) did not suggest any locality
for callirostris but, though listing it as a variety, thought it was
perhaps only an individual variation of Chrysemys or^iata.

The second specimen of callirostris was described by Sieben-
rock (1907) who regarded it as representative of a distinct sub-
species of Chrysemys ornata but doubted the ostensible locality
— "Florida" — of his specimen since he had received it from a

146 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

dealer. He abstracted his 1907 information in his 1909 "Synop-
sis."

The next notice of caUirostris was as a full species in the list
of Mertens and Miiller {in Hans Rust, 1934). The reason for
this treatment was not revealed until Miiller 's paper of 1940.

In the interval, however, Carr (1938) (in a paper primarily
concerned with indicating intergradation between Pseudemys
scripta elegans, then called troosti, and populations of the north-
east coast of Mexico which he called cataspila) mentioned cal-
lirostris rather parenthetically. He wrote: "Gray's description
of the form (1855) is based on a single specimen and the char-
acters by which he distinguishes it all fall within the range of
individual variation in cataspila. I am inclined to agree with
Boulenger that Gray's specimen was an aberrant cataspila.'^
Carr was apparently unaware of Siebenrock's second specimen,
although he cites the 1909 Synopsis.

In 1940, in a paper which (because of World War II and be-
cause of its rather obscure place of publication) has not been
generally available to American herpetologists, Lorenz Miiller
redescribed caUirostris from a series of 10 specimens in the
Miinchener Zoologische Staatssammlung.

Of Miiller 's seven specimens with good locality data, three
came from Barranquilla, Colombia, and four were collected by
llellmich at Jesus del Rio in a tributary of the Rio Magdalena.
Miiller published excellent photographs of the young and half
grown and full descriptions of these and of an adult, comparing
them pictorially and in the text with comparable stages of
(>r)iata.

In 1950 Smith and Taylor cited caUirostris in their checklist
as a possible synonym of Pseudemys scripta cataspila (and in-
judiciously restricted the ty])e locality to Alvarado, Vera Cruz,
Mexico ) .

In 1952 Mertens published a discussion, with photographs, of
tAvo more living specimens from the vicinity of Barranquilla.
He mentioned also another probable record of the form (Stuve.
1907^) from the neighborhood of Soplaviento on the lower
Magdalena.

1 Wochenschr. Aquar. Terr. 4, 228-229 (n^'t .seen).

WILLIAMS: PSEL^DEMYS SCRIPTA CALLIROSTHIS 147

Mertens and WCi-niuth (1955) list Pseudemys callirostris as a
full species with the distribution "Northern South America
(Colombia)."

C^HARACTERS

The distinctive characters of callirostris as compared witli
ornaia liave been stated by Gray (1855), Siebenrock (1!)09), and
Mertens (1952). Lorenz Midler's discussion, though more lengthy
and based on more material than any other, does not state what
characters are diagnostic but merely alternates detailed descrip-
tions of individual specimens of callirostris and ornata. I sum-
mariz(^ in Table 1 the characters found to be valuable.

TABLE 1

callirustri.s onmta

1. a broad supratempoial stripe or 1. a narrow supratemporal stripe,
spot cinnabar red in life, usually yellowish or reddish in life, typi-
rounded off anteriorly and w(dl eally entering the orbit,
separated from the orbit. '2. rather narrow stripes on the

2. a variable number of ocelli on underside of the snout, the max-
the underside of the snout and illary area and tlie mandilile.
on the maxillary area. Ocelli The mandilde typically with three
(usually 3) on the mandible. very elongate spots which may
Less regular ocelli posteriorly on either round off posteriorly or
the throat. may connect in varial)le fashion

with one or more of the stri]U's
on the throat.

Siebenrock could tind no characters of the shell to distinguish
callirostris from ornata, nor can I. Various statements made by
]\Iiiller implying shell differences require confirmation. He sug-
gests, for example, that in the adult state the dorsal ocellar pat-
tern is less clearly shown in callirostris than in ornaia and that
the shell is thus more "eiutiinig." He finds differences also iu
shell shaj^e, plastral i)roi)ortions, etc.

The ocellar as opposed to striped pattern of chin and snout
appears to be a ([uite constant diff'erence between all known
calliroslris (15+) and every oniafa seeu by uie ' nr figured in

1 All spcciiufiis of onidtn in tlic (•(illcitiuns of the .Viiit-rican Muscuiii of .Natural
History, the Chicajro Xatural History Museum, the Philadelphia Academy of
Naturai Sciences and the United States National Museum have been examined as
well as all ornata in the Miiseiim of Comparative Zoology.

148 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

the literature. Indeed, in this respect caUirostris differs from
all other members of the script a series. (There is some approach
in this character however in P. s. gaigeae Hartweg in which the
lateral stripes are shortened to long- ovals.)

The supratemporal spot seems not to be constant to the same
degree. Midler mentions that in one of his specimens of caUi-
rostris this spot was well separated from the orbit on one side,
but narrowed forward and approached the orbit very closely on
the other. Further, a series of specimens from Costa Rica ( Coen
Dispensary, MCZ 19351-3) show supratemporal spots not enter-
ing the eye, much as in callirostris. This is surprising since all
the Panamanian specimens which I have seen (and an Antioquia.
Colombia specimen) have supratemporal stripes entering the eye
quite as in Ilonduran or Mexican ornata.

There appears to be, however, much parallelism within scripta
in regard to the development of a supratemporal spot. Thus
Pseud cmijs scripta gaigeae is essentially only a regional variety
in which just such a supratemporal spot as is found in callirostris
or the Costa Rica individuals is especially well defined. Pseud-
emys scripta elegans also approaches cedlirostris in this regard
though in that form the supratemporal spot regularly enters the
orbit. In each of these cases also the supratemporal spot is red
whereas the similarly placed stripe of neighboring forms is
yellow.

As compared with all previously illustrated Colombian cal-
lirostris the Venezuelan specimen (Plate 2) appears to differ by
the narrowness of the central figure on the plastron. Probably,
however, there is high individual variability in this character.
There is certainly extreme variability in this regard within
ornata and except, perhaps, in very local populations there
appears to be no geographical pattern to the variation.

DISTRIBUTION

Callirostris has previously been reported from only three local-
ities in Colombia: (1) the vicinity of Barranquilla ; (2) Jesus
del Rio (Departamento Bolivar) between Calamar and Magan-
gue; (3) near Soplaviento. In addition. Brother Niceforo Maria
informs me that two additional localities are represented in the
collectious of the Instituto de la Salle: the Canal del Dique

WILLIAMS : PSEUDEMYS SCRIPTA CALLIROSTKIS 149

(Bolivar), not far from Barranquilla, and the borders of the
Cienaga del Guajaro (Atlantico), again near Barranquilla. The
United States National Museum also has a specimen from
Cartagena (USNM 61676).^

The new record in the Falcon district of Venezuela at La
Penita and Maravita Springs, Cerro Chicliiriviche extends the
known range of caUirostris hundreds of miles to the east. That
an extension of this magnitude is possible, points, as does the
general inadequacy of our knowledge of caUirostris, to the ex-
treme imperfection of our information regarding the turtles of
South America.

There is no ground for assuming that the eastern boundary
of caUirostris has been discovered. Between the Venezuelan
record and Pseudeniys dorhigni of South Brasil and Argentina
there is an immense gap which, though it may be in great part
real, will bear investigation.

As regards the western boundary of caUirostris there is no
exact knowledge either. From the Rio Chepo on the west coast
of Panama the Museum of Comparative Zoology has a typical
specimen of ornata (MCZ 7868). The American Museum has a
number of ornata from various localities in the Canal Zone. They
are again essentially typical. The Chicago Natural History Mu-
seum has the most easterly specimen (CNHM 63902) referable
to ornata known to me — from Turbo, Gulf of Uraba, Antioquia,
Colombia. This individual has a hroad supratemporal stripe
entering the eye and an extraordinarily reduced plastral pattern.
This single individual is not typical of, but neither is it outside
the variability of, ornata. The broad supratemporal stripe prob-
ably betrays some influence of caUirostris, but only further
knowledge of the Uraba Turbo population can show whether
there is true intergradation.

" r-'

AFFINITIES

Certainly, in most characters caUirostris is extremely close to
ornata. It is closer to this than to any other form. From this
lK)int of view Siebenrock was justified in regarding it as a sub-
species of ornata. However, it seems advisable to view this form

1 Dr. F. Mfrtciii is making morp fxti^nsivp ooUpctions of rnlliroxtria in Colombia
upon which he will report.

150 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

in a larger context, in terms of the entire series related to P.
scripta to which it certainly belongs.

1 have not attempted to review the scripta series in a thorough
fashion ; to do so will recinire many more specimens from many
more localities than have been available to me. But in order to
make clear the place of caUirostris in the scripta series I have
examined juveniles of every member of this widely distributed
series, except nfbuJosa and Jiiltoni, and adults of most forms
including both itchulusa and hilfoni.

In this series the most striking differences are those in pattern,
and these are most clearly seen in juveniles. Fully adult animals
of either sex may show patterns much more obscurely, and in
adult males the phenomenon which is commonly called "melan-
ism" typically results in the complete disruption of the juvenile
pattern.

I state the geographic variation in pattern in terms of four
conspicuous elements only — the dorsal carapace pattern, the
pattern of the sides of the head, of the chin, and of the plastron.

1. The scripta series begins on the east coast of North Amei--
ica with the populations called scripta. These populations ex-
tending from southeastern Virginia to northern Florida have a
carapace typically with a transverse stripe on each costal though
occasionally the last costals may bear ocelli, a head pattern with
a very consi)icuous vertical yellow spot behind the eye (the spot
usually sha])ed like a distorted triangle), a chin which is striped
longitudinally, and a plastral pattern varying from immaculate
yellow to ocelli or smudges on each plastral scute.

2. To the west the scripta populations intergrade with popu-
lations given the name of elegans, occupying the Mississippi
Valley from Illinois to the (lulf, ranging westward into Iowa.
Kansas, Missouri, Oklahoma and Texas, eastward into Alabama.
Tennessee, Kentucky and Ohio. These populations centering on
the Mississippi Valley resemble scripta in the transversely lined
carapace. They differ strikinuly in the presence of a longitud'na]
red temporal spot behind the eye. The chin is longitudinally
striped, while the plastral pattern is typically one of open ocelli
in juveniles or smudges in adults, one on each plastral scute.

3. In the upper courses of the Cumberland and Tennessee
rivers, populations formerly regarded as not distinct from ele-
qans have recently been redescribed under the name troosti.

WILFjIAMS : PSEUDEMYS SCRIPTA CAI.LIROSTiaS

151

These aiiiiimls have transverse earapaee stripiuj^,' a iianoir
longitudinal yellow head spot, chin with broad stripes, and on the
plastron small ocelli or sniud»i'es or no dark pigmentation.

4. In portions of Chihuahua, Coahuila, l)uran<?o and extreme
western Texas, there is a population ( /'. s. (jagcne) which in
typical specimens is very distinctive hut whieh apparently inter-
<j;:rades broadly and complexly to the east with clcfjans and to the
south with the form sometimes called rataspild out wliieli I
would unite with ornata. (inifjcac has a reticulate carapace ])at-
tern with frequently some ocelli present at least posteriorly, a
hlack-hordcred oval red spot behind the eye and well sej)arated
from it, a chin striped medially but with the lateral stripes
shortened into elon<;ate ovals that are almost ocelli, and a plastral
pattern of broken lines and spots mostly extendino; laterally

• ornota
X callirostns

Fig. 1. Ma)) of the southenmiost localities (Panama and Colombia) for
Pimudenn/s .scripta ornata and the kno-v\-n localities for Pscudeiiuifi scripta
callirostrls (Colombia and Venezuela).

along the sulci. The plastral pattern may sometimes approach
that of script a and sometimes be more like that of ornata (see
below) but is never as well defined as in either of these.

1 MCZ 49Uo, a juvenile from Kuoxville, Tennessee, regarded as trooati on bal-
ance of characters and on locality, is hi«hl.v peculiar in several aspects of dorsal
pattern especiall.v in havinjr a rather oeellar light spot in each costal rather than
the transverse stripe. This specimen is tijriired in ventral view in Plate 1.

152

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

o. To the west of gaigeae in Mexico there is another popula-
tion in southern Sonora which has been described under the
name of hiltoui. The carapace is reticulate dorsally. There is
a lonjiitudinal light orange temporal spot behind the eye, not
black-bordered and not separated from the eve. The chin is
striped, and the plastral pattern is one of discrete paired, rather
rectangular, smudges arranged in two lines along the middle of
the i)lastron (Fig. 1). This population is not known to inter-
grade with gaigeae or ornata, but Pseud emys from intermediate
localities are still undiscovered.

Fig. 12. Plastnil pattorn in Pscudemy.s scripta liiltani type. (After Carr
1942.)

6. Across the Gulf of Baja California in the southern part of
the peninsula of that name is an isolated j^opulation named
nehulosa by Van Denburgh (1895). This form is described as
ihaving rather reticulate markings dorsally on the carapace, an
orange or yellow spot, not black-bordered, on the side of the
head, a striped chin, and (Fig. 2) a narrow smudged central
plastral pattern, not broken into discrete spots, much resembling
the plastral figure of Chrysemys picta marginata and with ob-
vious similarities also to the plastral pattern of ornata (see
below). A specimen (USNM 75854) from La Purisima, La Paz,
Lower California has a plastral figurt^ of broken smudges very

WILLIAMS : PSEUDEMYS SCHII'TA CAIJJROSTRIS

153

like those of llic type scries of Inlloiti, wliieli may tlius be syn-
onyinous witli luhulosa — tlu' earlier named of the two forms.

7. I (listiii<>nish in the remainin»' portions of Mexico and in
all oF Ceiiti-al America only a siii<ilc member of the scripta series
(1 omit /'. (ir<nji from discnssioii U)v the moment). This set of
populations is variable bnt not in any defined p-pocvraphical sense;
there is certainly no clcdv noi'th-south differentiation witlun it as
Carr (li).SS) sujugested.' 1 thei-efore reject cataspila (liinther
1885 (type locality "Mexico") in favor of ornata Gray 1831
(type locality Mazatlan, Sinaloa, Mexico). h\ thus synonymizinf;

Fig. 3. Plastral pattern in P.scinlcinns scripfa nehidosa type. (After A'aii
Donlmrgh 1895.)

cataspila, 1 adhere to tiie philoso])hy that no iufraspecific popu-
lation should be given a formal latin name unless it is sharply
set off by one or moi'c distinctive features that are constant or
nearly so within a "'eographic ai-ea the borders of which can be
rather sharply defined.

The plastral pattern is one of closed concentric lines sometimes
occupyin'.-' the center of the plastron or sometimes spread out

1 Carr (lil'lS) formud liis concept of mtn^vilii in part on the specimen made
the type of P. .v. o^igexie by Hartweg (lllMHa. T.tMith),

154 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

along the sutures in very complex fashion. There is perhaps a
tendency' to greater width and complexity of pattern in more
southerly localities but the range of variation is extreme; even
Costa Rica and Panama individuals may have very narrow
plastral figures. Some of a series from Rama River Nicaragua
(AMNH 12415-23) show especially wide and elaborate plastral
figures ; in others of the same series the figure is narrow. Typi-
cally from Mexico to Panama there are on the head in the tem-
poral region only narrow yellow lines entering the orbit, but in
our Costa Rica locality (Coen Dispensary) there is an expanded
supratemporal spot much as in clegans. The dorsal pattern is
more or less clearly ocellate ; whether the ocelli touch or do not
touch the posterior margins of the dorsal shields appears to be a
jnirely individual feature.

The impression which these notes give of greater variability
in or7iata than in the previously mentioned populations is prob-
ably erroneous. Because these populations approach callirostris
so closely both geographically and in their characters, I have
examined greater numbers of specimens and I am thus more
aware of variability. Ornata appears to be a set of populations
fully as well defined as any other. Intergradation to the north
with gaigcae and dcgans appears to be well established (Ham-
ilton, 1947).

8. CaUirosfris in Colombia and Venezuela has the dorsal pat-
tern ocellate like ornata. lias a plastral figure, whether narrow
or wide, of closed concentric lines as in ornata and differs from
most ornata including Panamanian populations in having a red-
dish supratemporal spot that does not enter the eye, and differs
from all other Pseudemijs in a fully ocellate pattern on the chin
and snout. Intergradation with oDiata in western Colombia is
inferred but not demonstrated.

9. No Pseudemfjs is yet known in eastern Venezuela or the
(luianas. The only South American population other than cal-
lirostris (and at the extreme northwest ornata) is that in the far
south in Rio Grande do Sul, Brasil, and in neighboring parts of
Argentina. This is again a member of the scripta series but is
usuallv considered a distinct species, P. dorhigni. Though sep-
arated by half a continent from caUirostrif< it shows some decided
resemblances to that form. The plastral figure is a broadened
intense ornaia-like figure quite similar to the plastral figures

WILLIAMS: PSEI'DEMYS SCRIPTA CALLIROSTHIS 155

ill Coloiiibiau callirostri.s. It is (listinctivc in the inteiLsity of the
black pigmentation rather tlian in any detail of pattern. The
snpratemporal stripe is enlarged and separated from the orbit
as in caJlirostris. The snont is stri]~>ed and there is a median
stripe forking posteriorly on tlie eliin bnt at the corners of the
lower jaw there is on each side an ocellns not significantly dif-
ferent though perhaps slightly h)nger than in the callirostris.
Even the dorsal i)attern of the carapace is sometimes ocellate as
in ornata or in caJliro.stris, but here the similarity is verbal rather
than exact. The ocelhis in ornata and callirostris is a light ring
enclosing a darker central spot. In dorhigni the ocellus when
present is a transversely elongate ring of black surrounding a
very light (in alcohol, white) center. In other specimens the
ocellus is drawn out into a transverse streaked pattern.

Pscudoujis dorhif/iii is the terminal UKmilier of the continental
scripta series. There is, however, a subsidiary series in the West
Indian islands. 1 have briefly surveyed this series also and dis-
tinguish within it three subgroups witli regard to plastral i)at-
terns.

1. Pseiidemys decussata from Cuba, 7*. s. stejnegeri from
Puerto Rico, P. s. vicina from Hispaniola, though differing each
in a minor but characteristic way, all resemble to an amazing
degree some specimens of ornata, in particular those from Rama
River in Nicaragua (AMNH 12415-28) already cited.

2. 1 have seen only a few specimens of young P. terrapen,
none of which is fully similar to the deeussata-steJ7iegeri sub-
series. The single ^ICZ juvenile is quite unique, as compared
with all Psrudemj/s that I have examined, in its broad unified
black plastral marking. Slightly older specimens at the United
States National Museum are more ornata-Yike (probably falling
within the variability of <n-i\ata i but distinctly less complex in
pattern than deciissata and stejnegeri. Even the solid pattern
of the MCZ juvenile, however, seems best interpreted as a modi-
fied ornafa pattern.

;j. (^uite isolated in plastral pattern is P. dccorata (said to
occur sympatrically with /'. .s. vicina). In this, there are small
light-centered spots on the plastron like those of elegans but
without the regular arrangement characteristic of elegans. This
condition is again probably derivable from the ornata pattern
and ])robably from the more complex varieties of this by break-

156 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

down and diminution of pigment. An occasional specimen of
dccussata approaches decorata by the fragmentation and simpli-
fication of the plastral pattern but without achieving the scat-
tered ocelli of decorata. Some specimens of decorata have the
plastral ocelli elongate and smudged so as to approach, but at a
distance, the jilastral pattern of dccussata.

All three groups of the West Indian Pseudemys are otherwise
very similar and very like ornata. All have an ocellate carapace
pattern and striped heads and chins. All, however, tend to have
the head striping fainter than in continental forms and all tend
to lose at least dorsal pattern early.

It is possible that I have here somewhat oversimplified the
picture. Certainly there is much to be done in clarifying the
variation within as well as the intergradation between named
forms. There are manv general statements in the literature re-
garding intermediates between one oi' another subspecies usually
with remarks on greater similarity to one than to the other.
Such statements cannot be independently evaluated until there
is more ]:)recise information on the characters on which these
judgments are made. I have not myself seen the intergrading
populations in any of the several cases.

A special problem still remaining is that of Pseudemys grayi
Bocourt.^ The United States National Museum has several skulls
and shells belonging to this form. The slender skulls look very
different from those of ornata, but it is not clear to me how the
shells of grayi are to be separated from those of ornata. Carr
(1938) has suggested that P. grayi may represent only the males
of ornata, but if so the differences seem both very extreme and
highly localized. A biological proI)lem is evidently concealed
under the name P. grayi which merits more attention than it
has received .-

However, in spite of unresolved problems, an inspection of
scripta series as a whole does make it clear that there are several
grades of differentiation within it. Morphologically, the major
break lies between the populations of the United States, with

1 Pov the use of P. firaiii Bo('<iurt r.-ithiT lli.-iii /'. iti\i\tr<i < )'Shaut:liness.v see
Mortens and Werniuth ( 10.")."i') .

i! Th(^ heads of males of P. (Iccnrnta fri in Hispaniola and P. titcjncgcri from
Puerto Uioo are also strikingly narrow ; rliose of males of P. dreu.s:inta from ("nha
are not.

WILLIAMS: PSEITDEMYS SCRIPTA CALLIROSTRIS 157

their linear dorsal, and absent or uneoinplieated ventral, ])atterii,
and the populations to the South (Mexico, Central and South
America and tlie West Indies), with theii- ocellate dorsal and
typically "ornate" ventral pattern. Possibly the latter is the
more primitive. Certainly it is sug'ji'estive that the ventral pat-
tern of the ornafa .subseries is substantially identical with a
ventral pattern which occurs in the P. jloridana series {P. rubri-
ventris and /*. floridana) and also in the two genera closely
related to Paeudemys, Graptcmys and Chrysemys (C. picta belli,
C. picta 1)1(1 r(ji)iata and G. psendogeograplvica) .

Geographically situated at the major break within the scripta
series, the po])ulations called (joigcac, nebulosa and hiUo7ii are
in some regards intermediate but appear also to have characters
of their own. Gaigeae appears to bridge the gap biologically
(Hamilton 1947).

In the scripta subseries the differences between clegans and
scripta are sharp and constant within the typical areas of these
two named populations. 2' roost i as currently restricted (Burger
1952) is a population somewhat intermediate between scripta
and elegans.

In the ornata subseries there is a sharp single character dif-
ference between ornata and caUirostris and a more complex but
less obvious ditference between both these and dorbigni. Certain
of the West Indian forms {decussata and stejnegeri) differ less
from some ornata than ornata may differ within itself. Yet these
forms superficially so close to ornata show a probably more mean-
ingful affinity with their more differentiated island neighbors
tcrrapen and dccorata.^

The example of the insular forms suggests that caution may
be desirable in evaluating degrees of difference on the mainland.
While it is clearly possible that all the scripta series {P. decorata
and perhaps P. grayi apart ) is one species, it is equally clear
that the situation is not simjile and needs much further detailed

1 .Mcrtfiis anil Wcriuutli il'.>'>'> p. :!(;(>-:',C>T ) list all rhc ciirn-nl ly naiiHMl West
Indian forms as siibspec.es (if trnaixn. Wliile tliis iadicafes a relntidii amnnt;
these fcirnis which I believe to be real, I do not think it indicates correctly the
several grades of this relationship. I do not ret,'ard (nujuxln and plniiii as reco;:-
nizable. Fv}i>< and nuilomi may be neither native nor recoirnizablc. while graiiti
niav be retained suh jiidicc. 'I'entatively. I sujTKest the following ai'ran>:ement :
Pfieudcmiis dccnsNtitd drcitssntii (the sole Cnban form). I'si:idcin!if drcitsxata
riciiia (Ilisiianiohui), P-scndrmiis dcciisnata xtrjiK i/cri (Puerto Kican). I'sciidcmiix
dccii>'x<it(i f/riniti (Cayman). PKcitdcmjix terrain n (.lamaican). I'xriidnniix dcronitii
(Ilispanicdan) .

158

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Scripta subseries

troosti

elegans

/

scripta

gaigeae

nebulosa (hiltoni)

Ornota subseries

ntermediate

populations

ornata

decussata-
vicina-
sfejnegeri

terrapen
decorata

callirostris

dorbigni

Fig. -4. Diagram to show groupings and approximate geographic position
ind aftinities in the .srripia series.

WILLIAMS: PSEUDEMYS SCRIPTA CALLIROSTKIS 159

study. For my i)ai-t 1 am eonteiit to modify the current iiomeii-
I'laturo of mainland forms to a minimal extent, indieatinf>- by
the new eombination /*. sciiiita (■(illiroslris that callirostris differs
only in minor though eonstant ways from its nearest neiji'hbor in
the scripta series, P. scripia onxihi, that these differences seem
less than those between |)oi)\dations known to interfjrade, and
that though there is no present critical evidence of intergrada-
tion there are also no counter arguments to the hypothesis of
subspecies status.

AcknowkHigments : I am indebted to the authorities at the
United National Museum, the Philadelphia Academy of Natural
Sciences, the Chicago Natural History Museum and the Ameri-
can Museum of Natural History for the privilege of examining
their collections. I wish to thank also Mr. Benjamin Shreve,
Mr. Arthur Loveridge and Mr. Walter Auff'enberg for reading
the manuscrijit and making suggestions.

REFERENCES

BOULENGER, G. A.

1S89. Catalogue of the ehelonians, rhyiiehocephalians and croeodiles in
the Britisli Museum (Natural Tlistory). T.ondon, 311 pp.

BlRGER, W. L.

1952. A neglected subspecies of the turtle P.'^cudemys ncHpta. Jour.
Tenn. Acad. Sci., 27: 75-79.

Carr, a. F.

1938. Notes on the Pscudemt/.s .scripld complex. Herpetologica, 1: 131-

135.
HUl'. a new P.^^cHdemi/.s from Stiudra, Mexico. Amer. Mus. Novitates,

No. 1181: 1-2.
UI5-J. Handbook of turtles. The turtles of the United States, Canada

and Baja California. Ithaca, N. Y. 542 pp.

Cope, E. D.

1887. Catalogue of the batrachians and reptiles of Central America
and Mexico. Bull. U. S. Nat. Mus., 32: 1-98.

Cray, J. E.

1855. Catalogue of shield reptiles in the collection of the British
Museum, Part I. Testudinata. London, 79 pp.. 42 plates.

160 BULLETIN' : MTSEITM OF COMPARATIVE ZOOLOGY

GllNTHER, A.

1885. Chelonia, in Keptilia and Batracliia. Biologia Centrali - Ameri
cana, pp. ]-18.

Hamilton, R. D.

1947. Tho range of I'.sfiuli mi/s si-npla (iai(ieac. Copeia, 1947: (55-66.

Haktweg, N.

1939a. A lu'w American Pscaili nii/s. On-. Papers Mns. Zool., Univ.

Michigan, No. 397: 14.
19391). Further notes on the Psctalcmiiit scripla complex. Copeia, 1939:

55.

Mertens, R.

1952. Pseudemys calliroatris, eine Schnuick-Hchildkrote aus Kolumbien.
Aquar-Terrar. Zeitschr., Jahrg. 5: 184-185.

Mertens, R. and H. Wermuth

1955. Die rezenten Sehildkroten, KrokodiK^ und Briickenechsen. Zool.
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MULLER, L.

1940. Ul)er P.scndem!/.s call i runt ri't (Gray). Iti Tier und Unnvelt Sud-
anierika. Ibero-americanische Studien, 13: 108-126.

RrsT, H.

1934. Systematische Liste der lebenden Sehildkroten. Blat. Aquar.
Terrar., 45: 42-45, 59-69.

Shreve, B.

1947. On Venezuelan reptiles and amphibians collected by T)v. H. P.
Kugler. Bull. Mus. Comp. Zool., 99: 519-537.

SlEBENROCK, F.

1907. Chri/semys ornaia caUiro.stri.'< (Tray, eine selbstandige Unterart.

Zool. Anz., 31: 790-792.
1909. Synopsis der rezenten Sehildkroten mit Beriicksichtigung der in

historischer Zeit ausgestorbeuen Arten. Zool. .Tahrb., Suppl., 10:

427-618.

Smith, H. and E. H. Taylor

1950. An annotated checklist and key to the reptiles of Mexico exclu-
sive of the snakes. Bull. U. S. Nat. Mut., 199: 1-253.

Van Denburgh, J.

1895. A review of the herpetology of lower California. Part I. Rep-
tiles. Proe. Calif. Acad. Sei., Sec. 2, 5: 77-162.

PLATES

Plate 1. Ventral views of turtles of the scrlpta series. Upper left:
Pseudemys scripta elegans M.C.Z. 12638, Aniston, Alabama. Upper right:
P. s. troosti M.C.Z. 4905, Knoxville, Tennessee. Lower center: P. s. gaigeae
M.C.Z. 4551, San Pedro, Mexico.

PLATE 1

Plate 2. Ventral views of turtles of the scripta series. Upper left : P. .'*.
ornata M.C.Z. 19352, Coen Dispensary, Costa Rica. Upper right: P. dor-
bigni M.C.Z. 33502, Porto Alegre, Brasil. Lower center: P. -s. canirn.<<tri.s
M.C.Z. 49055, Cerro Ohichiriviche. Silva district. Falcon State, Venezuela.

PLATE 2

Plate 3. Wntial views of turtks (jf tlie scripta series. Upper left: P.
(Iccu.ssafa (h'ni.-i.'iiiUi. M.C.Z. 34179, twelve miles from Soledad, Cuba. Upper
ri^lit: P. fcrrapt'u M.C.Z. 52399, Rio Cobre Canal along Spanishtown Road,
.lamaica. Lower center: P. drmrdtd M.C.Z. 36855, Thoniazean. Plaiti.

PLA TE 3

BuUetin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. 115, No. 6

REVISION OF THE AFRICAN TORTOISES AND
TURTLES OF THE SUBORDER CRYPTODIRA

By Arthur Loveridge and Ernest E. Williams

With Eighteen Plates

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

February. 1957

Publications Issued by or in Connection

WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HAHVAKD COLLEGE

Bulletin (octavo) 1863 — The current volume is Vol. 115.

Breviora (octavo) 1952 — No. 61 is current.

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

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

Occasional Papers of the Department of Mollusks (octavo) 1945 —
Vol. 1, no. 20 is current.

Proceedings of the New England Zoological Club (octavo^ 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.

The continuing publications are issued at irregular intervals in numbers
which may be purchased separately. Prices and lists may be obtained on
application to the Director of the Museum of Comparative Zoology,
Cambridge 38, Massachusetts.

Of the Peters "Check List of Birds of the World," volumes 1-3 are out
of print; volumes 4 and 6 may be obtained from the Harvard University
Press; volumes 5 and 7 are sold by the Museum, and future volumes will be
published under Museum auspices.

Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE

Vol.11?i.No.R

REVISION OF THE AFRICAN TORTOISES AND
TURTLES OF THE SUBORDER CRYPTODIRA

By Arthur Loveridge and Ernest E. Williams

With Eighteen Plateb

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

February, 1957

No. 6 — Revision of the African Tortoises and Turtlen
of the Suborder Cryptodira

By Arthur Loveridqe and Ernest E. Williams^

CONTENTS

PaOI!

Introduction 163

List of the Cryptodira in Africa 175

List of Keys and Tables 177

Systematic Discussion 180

Bibliography of African Cryptodira 503

Index to All Scientific Names and Sjiionyms 542

INTRODUCTION

The last comprehensive treatises to embrace all African
testudinates were the "Catalogue of the Chelonians, Rhyncho-
cephalians, and Crocodiles in the British Museum," by G. A.
Boulenger (1889a), and "Synopsis der rezenten Schildkroten
mit Beriicksichtigung der in historischer Zeit ausgestorbenen
Arten," by F. Siebenrock ( 1909a ).^ The present contribution is
an attempt to bring our knowledge up to date by summarizing
all the available information of importance regarding African
Cryptodira. This completes the coverage of African Testudinata.
as the Pleurodira have already been dealt with by one of us.^

Material. We have collaborated in completely rewriting de-
tailed descriptions of the external characteristics of each of the

1 These names are listed alphabetically, or according to the authors' respective
antiquity, without further significance. The entire osteologifal and paleontologi-
•-■al accounts, all major discussions, and the .selection of illustrations are the
work of my industrious colleague. Dr. E. E. Williams, whose unrivalled knowledge
of the Testudinata is well known. Without his erudition and drive, it is safe to
say that this monograph would never have been completed in so comprehensive a
form. A.Iv.

2 Though published on October 31, 1955, the checklist of the turtles of the world
by Drs. Mertens and Werinuth, unfortunately 'lid not reach us until the manu-
script of this paper had been submitted for publication. It was thus too late for
citation under genera and species and should be consulted for the numerous
differences.

3 Loveridge, A., 1941, Bull. Mus. Comp. Zool., 88: 465-524. One of a series of
revisioiiary studies of various families of African reptiles published between 1938
and 1947.

164 BimLETTN : MUSET^M OF COMPARATIVE ZOOLOGY

32 species or forms here recognized. In doing so we have been
fortunate in having available in the Museum of Comparative
Zoology African material of all 32 forms with the exception
of one marine species, together with many paratypical or topo-
typical examples of them or their synonyms.

In order to examine other types, and in search of additional
information, one of us (E. E. W.), as a Guggenheim Fellow,
spent many months studying testudinates in most of the princi-
pal museums of Europe (Leiden; London; Munich; Paris;
Prankfurt-am-Main) and eastern North America (Chicago; New
York; Philadelphia; Washington). Upon his return, the
curators of all these and many other institutions generously
loaned for further comparative study much critical material —
often of a bulky or fragile nature necessitating careful packing.

When such loaned material is referred to, we have indicated
the institution by one of the following abbreviations :

A.M.N.H., American Museum of Natural Historj-, Xew York.

B.M., British Museum (Natural History), London.

CM., Carnegie Museum, Pittsburgh.

C.N.H.M., Chicago Natural History Museum, Chicago.

G.M., Museum d'Histoire naturelle, Geneva.

H. M., Zoologisches Museum, Hamburg.

I.F.A.N., Institut Frangais d'Afrique Noire, Dakar.

L.M., Rijksmuseum van Natuurlijke Historie, Leiden.

M.M., Zoologische Sammlung des Bayerischen Staates, Munich.

P.M., Museum National d 'Histoire naturelle, Paris.

N.R., Naturhistoriska Riksmuseum, Stockholm.

S.M., Senckenbergische Naturforschende Gesellschaft, Frankfurt a. ?it.

T.M., Transvaal Museum, Pretoria.

U.S.N.M., United States National Museum, Washington.

V.M., Naturhistorisches Museum, Vienna.

W.M., Naturwissenschaftliche Sammlung, Neues Museum, Wiesbaden.

Y.P.M., Yale Peabody Museum, New Haven, Connecticut.

Z.M.U., Zoologisches Museum der Universitat, Berlin.

Acknowledgements. We herewith welcome this opportunity
to express our appreciation for the cooperation of colleagues
both here and abroad, especially of those who, with unfailing
courtesy, have answered endless questions of a seemingly trivial
nature that involved careful examinations of specimens in their
charge. We have made exceptionally heavy demands on the

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 165

British and Transvaal Museums, whose collections of African
Cryptodira ai'e particularly rich.

For lending specimens and/or answering queries we desire
to express our grateful thanks to:

V. Aellen (Geneva Museum)

J. C. Battersby (British Museum)

G. Belloc (Institute Oceanographique Monaco)

C. M. Bogert (American Museum of Natural History)
R. D. Brongersma (Leiden Museum)

P. Chabanaud (Paris Museum)

D. M. Cochran (United States National Museum)
J. Eiselt (Vienna Museum)

V. F. FitzSimons (Transvaal Museum)
J. T. Gregory (Peabody Museum, Yale)
J. Guibe (Paris Museum)
F. Heineck (Wiesbaden Museum)
W. Hellmich (Munich Museum)

A. C. Hoffman (Bloemfontein Museum)
R. F. Inger (Chicago Museum)

W. Lanz (Geneva Museum)

R. F. Lawrence (Natal Museum)

M. E. Malan (Stellenbosch University)

R. Mertens (Senckenberg Museum)

B. L. Mitchell (Nyasaland Tsetse Control)

E. Mohr (Hamburg Museum)

C. F. Moysey (Devonshire, England)

F. Neubar (Wiesbaden Museum)
H. W. Parker (British Museum)
J. A. Peters (Brown University)

N. B. Richmond (Carnegie Museum, Pittsburgh)

H. Rendahl (Swedish Museum)

C. A. du Toit (Stellenbosch University)

A. Villiers (I.F.A.N. Museum, Dakar)

H. Wermuth (Zoologische Museum, Berlin)

J. G. Williams (Coryndon Museum, Nairobi)

G. F. de Witte (I.R.S.N. de Belgique)

We also wish to express our appreciation to Mr. Thomas
Andersen of Kidugallo, Tanganyika Territory, for endeavoring
to obtain for us certain specimens of Malacochersus ; to W. H.
Archer of East London, South Africa, for writing us about
Geochelone ; to G. S. Cansdale of London, England, for answer-

166 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

ing questions regarding Gold Coast Kinixys; to C. Jacot-Guillar-
mod of Basutoland, respecting the occurrence in that country of
Psammohates; to our colleague Dr. Ernst Mayr for advice on
various taxonomic enigmas; to T. S. Jones, Esq., of Sierra Leone,
for securing specimens that established the presence of Kinixys
h. nogueyi in that country, and not least to J. M. Savage of
University of Southern California for his kindness in reading
over, and commenting on, this manuscript.

We welcome this opportunity to acknowledge our indebtedness
to the following artists whose numerous drawings add much to
the usefulness of this revision: Mr. S. B. McDowell of New
York, Miss E. R. Turlington of London, Miss P. M. Washer of
the Museum of Comparative Zoology. We are also grateful to
Messrs. Peter Green of London, England, and Prank White of
Cambridge, Massachusetts, for their photographic achievements,
and to the Trustees of the British Museum (Natural History)
for permission to reproduce various photographs of specimens
in that institution.

Taxonomy. Though, as a result of collaboration, the scope of
this revision is greater than that of its predecessors, changes in
nomenclature are remarkably few, for we have endeavored to
maintain a conservative attitude throughout. The only fresh
names proposed are : Aldahrachelys, new subgenus for the giant
tortoises of Aldabra, etc., Pseudotestudo, new subgenus for the
dwarf tortoise of Egypt and Libya. No new species or races have
been erected, though in certain instances {Malacochersus, Cyclo-
derma) the possibility of their existence has been noted.

At the specific and infraspecific level emphasis has been rather
on the suppression of names, never lightly or on theoretical
grounds, but only after a careful examination of the available
material failed to support their claims to specific or racial dis-
tinction. In some cases (such as Kinixys helliana) the accumula-
tion of additional tortoises may produce valid reasons for the
revival of forms we have not recognized. But to sustain these
names will require more comprehensive and careful studies than
those on the basis of which they were proposed. It cannot be
too strongly emphasized that testudinates should never be de-
scribed on single characters and but rarely on a single specimen.
Conclusions based on series that are limited to less than ten

LOVEBIDQE AND WILLIAMS: AFRICAN CRYPTODIRA 167

individuals should be viewed with suspicion, for it is highly
probable that the apparently constant characters displayed by
so small a sample will prove to be unstable when a larger series
is available.

The status of one species, Cyclanorhis elegans Gray, founded
on a juvenile, has been clarified by synonymizing with it oligoty-
lus Siebenrock, founded upon an adult.

At the generic level the changes recommended,^ though not
minor, are infrequent. Thus, after a careful survey which in-
cluded consideration of extra-limital forms, we have concurred
with present South African usage in recognizing the large tor-
toises of Africa as generieally separable from Testudo, sensv
stricto, as are also the members of the geometricus group and
angulata (for details cf. pp. 218ff). Another change in the con-
cept of a genus has necessitated our removing the North Ameri-
can pond-turtle species hlandingii to the genus Emydoidea, re-
taining in Emys only the species orhicidaris of Europe and
Africa.

At every point in our taxonomic study we have been faced
with the fact that in all characters the amount of variation ex-
hibited by members of this order is frequently astonishing and
always disconcerting. In our descriptions of external characters
we have endeavored to record every major variation encoun-
tered. The number of such observed variations, it should be
noted, is apt to bear a direct ratio to the number of specimens
studied. Thus, where relatively large series were available to
us, as in the case of Geochelone p. bahcocki, Malacochersus tor-
nieri or Kinixys h. helliana, our earlier crude observations were
constantly subjected to qualifying comment. If we have used
such words as "always" or "never" anywhere in the descrip-
tions, it has been in a moment of unguarded enthusiasm.

This high degree of variability extends to osteology also. There
are no grounds for assuming that internal characters are more
constant than external ones. The only certain test of stability
for both is the empirical one of examining a large series. Here
a practical difficulty arises, as series of skeletons are rarely avail-
able. Since, with good reason, generic definitions are largely

1 See also footnote on p. 220.

168 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

based on osteological characters, the lack of skeletal series has
been a real handicap.

In this predicament our method has been to rely upon deter-
mination of the range of variation in a test species, Geochelone
pardalis hahcocki, for which numbers of skulls and skeletons
were available. Whatever species, judged by even a few speci-
mens, lay wholly outside the variation of this form in several
major skeletal characters (e.g. maxillary ridging, pygal number,
neural shape, etc.) could, we felt, be safely regarded as only
remotely related and hence possibly generically distinct. How-
ever, we have not been content with this estimate of morphologi-
cal distance only, but wherever possible have endeavored to
trace lineages in the fossil record. We have accepted paleonto-
logical evidence as constituting additional osteological material,
and where the separation of lineages proved to be of great
antiquity — as, for example, going back to the Eocene — ac-
cepted it also as modest warranty of generic distinction.

While availing ourselves of published descriptions and illus-
trations, our first assessment of the degree of osteological varia-
tion in any given species or genus has been based on specimens
we ourselves have examined. Thus of the 27 African terrestrial
or freshwater species dealt with in this study, we have seen
skeletal material of all except three, viz. Psammohates geometri-
cus, Homopus signatus and H. femoralis. Of certain other
species relatively little material (less than three skulls or skele-
tons) has been available : only single skulls of Geochelone sulcata,
Psammohates oculifer, Homopus boulengeri and Cycloderma
auhryi. On the other hand, our understanding of the African
species and our ideas as to the variational range within their
subfamilies and families have benefited by the examination of
much extralimital material including, for example, both skulls
and skeletons of most of the species and every species-group of
the Testudininae.

Categories at every taxonomic level need to be studied against
a background of the next higher category if they are to be fully
understood. Thus no subspecies can be properly evaluated
without a reasonably adequate knowledge of the entire species
of which it forms a part. No species can be appreciated without
a substantial acquaintance with the whole genus in which it is

LOVERIDQE AND WILLIAMS: AFRICAN CBYPTODIRA 169

included. No genus can be satisfactorily defined except in terms
of full information of at least the section of the subfamily to
which it is assigned. Consequently we have not hesitated to
explore — for their background value — extralimital forms and
problems whenever these seemed likely to shed significant light
on their African counterparts. It is on this account that family
or subfamily phylogenies have been included, together with
discussions of non-African genera or species, and even defini-
tions of certain extralimital forms.

Despite all our care, our definitions, whether specific, generic
or familial, are weakened, at least in appearance, by the fantastic
variability on which ^Ye have commented above. Our key to the
external characters of families mentions lack of hornj^ scutes in
a solitary sea turtle. On another occasion one of us (Williams,
1954, Breviora, No. 29) has called attention to variation in a
"family character" of an African pleurodire species. Such radi-
cal variations cannot in honesty be omitted from the record, but
it is important to recognize that they do not actually diminish
the value as phyletic units of the categories whose "definitions"
they disturb. However, they do show how necessary it is to use
the greatest care and the most broadly based knowledge in deal-
ing with the vexing problem of the recognition of natural
groups. Some aspects of these matters have been discussed in
very percipient fashion by Zangerl and Turnbull (1955, Field-
iana: Zoology, 37:366 if.)

At the outset, the idea occurred to us to include under each
species a synopsis of any Problems remaining, or gaps in the
life-history requiring to be filled. However, we soon realized
that such synopses would be unavoidably repetitious and much
too long. Perhaps the most fully documented form is the Eastern
Leopard Tortoise {Gcochelone pardalis babcocki), but no Afri-
can cryptodiran can be said to be well known. One has only to
refer to the various headings given under each species or race
to realize how very sketchy is our knowledge of their respective
ecologies, dietetic preferences, reproductive habits, geographic
variation and range.

In the case of Anatomy the information was in general so
scanty that this heading was also omitted throughout, though
references to anatomical papers are retained in the citations for

170

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

each species. Only once has the anatomy of an African species
been studied adequately, and this was done on non-African ma-
terial. We refer to the very old but classic work of Bo j anus
(1819-1821) on Emys orbicularis. Less meritorious, but possibly
based on African specimens, at least in part, is that of Thomson
(1932) on "the Tortoise" (i.e. "Testudo graeca" = liermanni,
and "Testudo ihera" = graeca). Other anatomical discussions
involving African turtles either mention African forms inci-
dentally or in a more general context (e.g. sundry papers by
Siebenrock), or else record anatomical observations of no great
scope.

/-Sf^^

/h V

e^

\ P

/ abd

hypo '

\ f

P

xiphT^

Van

B

Fig. 1. Clemmys caspica leprosa: diagram of shell to show nomenclature
of horny scutes and bony plates, the latter 's abbreviations underlined
in the figure and italicized below. A, Carapace: c l-4 = costals; m 2,
10 = second and tenth marginals; n l-7 = neurals; nu = nuchal scute;
nu = nuchal bone ; p 2, 9 = second and ninth peripherals ; py = pygal ;
spy = suprapygal ; v 2 = second vertebral. B, Plastron ; abd = abdominal ;
an = anal; ent = entoplastron ; epi = epiplastron; f = femoral; g = gular;
h = humeral ; hyo = hyoplastron ; hypo = hypoplastron ; p = pectoral ;
xiphi = xiphiplastron.

♦ (P. Washer del.)

LOVERIDQE AND WILLIAMS: AFRICAN CRYPTODIRA

171

Terminology. We have introduced no new terras or new
usages to the nomenclature of either bones or horny shields. For
the dorsal shell our usage is that of George Baur in his several
papers, differing from that of Boulenger only by distinguishing
the rib-containing bony plates as "pleurals," and the bones
surrounding the shell as "peripherals." Our nomenclature for
the dorsal shell is contrasted with that of other authors in
Table I (see also Fig. 1).

Fortunatel}^ there has never been any confusion with regard
to the names of either the bones or horny shields of the ventral
shell. To express length-relationship between the various plastral
shields we have introduced a new "plastral formula" in which
the shields are listed in order from longest to shortest. This
formulation is not a substitute for precise studies of the length
ratios between plastral shields, but, despite variability, it tends
to be characteristic of species, genera, and occasionally of even

Table 1

Nomenclature of the parts of a turtle shell

HORNY SHIELDS

Our usage

Boulenger 1889

Hay 1908

Carr 1952

nuchal

nuchal

nuchal

precentral

vertebral

vertebral

vertebral

central

supracaudal

supracaudal

supracaudal

postcentral

costal

costal

costal

lateral

marginal

marginal

marginal

marginal

BONES

Our usage

Boulenger 1889

Hay 1908

Carr 1952

nuchal

nuchal

nuchal

proneuraP

neural

neural

neural

neural

suprapygal

pygal

suprapygal

epipygal

pygal

pygal

pygal

pygal

pleural

costal

costal

pleural

peripheral

marginal

peripheral

peripheral

1 Though, bv the invention of this and other noveUies. Carr intended his termi-
Qolotry to reduce confusion, here it is delinitely at fault. The very similar term
"preneural" has long been in use for an element that is immeiliately posterior to
the nuchal of autliors in certain Asiatic and fossil trionychids and some other
fossils.

172 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

higher units. Thus a plastral formula ^ with Abd>h> (all other
elements), or with Abd> (all other elements) turns out to be
very characteristic of testudinines.

As for head shields in the testudinines, we have adopted the
terms used by Miss Procter for Malacochersiis tornieri. In this
species, as also in the members of the genus Testndo, the dorsal
aspect of the head possesses the most elaborate regular scutella-
tion to be found in tortoises. The terms employed are "supra-
nasals, " " prefrontals ' ' and ' ' f rontals. ' ' With regard to the still
more elaborate head scutellation of the marine turtles, we follow
Deraniyagala (1939 :192 :fig. 76) except in employing the term
" supratemporals " for what he calls "temporals," as being more
appropriate.

Folklore. Probably fcAv reptiles figure so frequently in African
folk tales as do tortoises. All manner of superstitions are
entertained concerning them, and references to such beliefs con-
stantly crop up in books of travel by the earlier explorers. As
we came across relatively few examples in our search of zoologi-
cal papers we have omitted the subject altogether, rather than
treat it inadequately. The matter merits the attention of some
anthropologist who, preferably in collaboration with a zoologist,
would scan the literature and publish a classified synopsis of
these tales and beliefs.

Citations. Following the name of each species, when used as
a heading, citations to it or its synonyms are given in an ab-
breviated form which can be amplified by reference to the
Bibliography on p. 503. Almost 550 papers (1758-1955) in which
we have found references to African Cryptodira are listed.
Omitted from the bibliography are papers involving some single
generic or specific description for which an adequate citation
has already been given in the synonymy. Also omitted are cita-
tions to non-African species or races mentioned in the text ;
these are given in parentheses or as footnotes.

Attention is directed to the Synopsis Methodica (a folding
chart or table in which binomials are employed) at the end of
the first volume of Lacepede, 1788 "Histoire naturelle des Quad-
rupedes ovipares et des Serpens," a work frequently rejected by
systematists since only popular names are employed in the text.

1 See Pig. 1 for significance of abbreviations.

LOVEBIDGE AND Wn^LIAWS : AFRICAN CRYPTODIBA 178

An extraordinary volume that cannot be taken seriously,
is that of Rochebrune, 1884a, "Faune de la Senegambie. Rep-
tiles. " In it the author lists as occurring in Senegal such impossi-
bilities as Testudo marginata, T. geometrica, Homopus signatus,
H. areolatus and Cycloderma frenatum. Despite the fact that
definite Senegambian localities are furnished for them, only four,
or at most five, of the eighteen land and freshwater turtles
listed by Rochebrune actually occur in Senegal.

A question arises with regard to the priority of J. E. Gray's
contributions cited as 1831b and 1831c. The latter — Synopsis
Reptilium — contains many original descriptions and was clearly
intended to be published first. Unfortunately, the evidence sug-
gests that 1831b — Synopsis of the Species of the Class Reptilia
— which appears as an appendix to Edward Griffith's transla-
tion of Cuvier's Animal Kingdom, came out first. In Gray's
own bibliography the date is given as 1830; this is possibly
the date of its completion, an advance copy, or an earlier edition
than the usually accepted one of 1831. This synopsis in Grif-
fith makes only passing reference to, or gives the scantiest de-
scriptions of, species more fully dealt with in 1831c. For this
reason, whenever a new species is involved, we have cited 1831c
in advance of 1831b. In no case does it affect priority of
nomenclature.

In listing a reference to some form, we do not intend to infer
that the entire synonymy of the author cited necessarily applies
to the species or race to which it is assigned.

Localities. It will be noted that localities are listed alphabeti-
cally under their respective countries. The latter, beginning with
Morocco, are arranged clockwise around the continent. The only
exception to this is in the case of trionychids from north of the
equator, whose ranges are given from the Nile west to Senegal.
Generally speaking, we have adopted the orthography of the
government administering the area, though a few exceptions
such as Algiers (instead of Alger), and Tangiers (in lieu of
Tanger), have been allowed to stand. Where the current spelling
of a place name differs from that used in the original record
given in the literature, the rejected spelling is given in paren-
thesis after the preferred one.

174 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

A locality preceded by an asterisk implies that a specimen
from the place in question has been studied by us ; usually it is in
the Museum of Comparative Zoology. If it is in some other in-
stitution, however, the key letters of the museum where it is
preserved follow the locality in parenthesis. In many instances
such asterisk-bearing localities constitute fresh records of oc-
currence for the species and will not be found in the literature.

A locality that appears in quotation marks is one that has been
taken from the literature but which we have failed to find on any
map. In some instances, at least, the name may have been
misspelled or misprinted.

Unfortunately, all localities appearing in the literature can-
not be accepted. The handsome and ornate shells of tortoises
have attracted the attention of mankind from earliest times. This
is especially true in South Africa where primitive tribes were
accustomed to wear the shells of the smaller species as ornaments,
or used them as scoops or receptacles, especially for huchii
ointment (cf. Psammohaies geometricus under the heading
Enemies). Doubtless they were used in barter and, passing from
one itinerant African to another, were transported far from their
place of origin. In due course some were seen and purchased
by European travellers, especially those bent on acquiring ob-
jects of anthropological interest. Naturally the shells were
labeled as coming from the place where purchased.

In recent times car drivers are likely to stop and pick up any
small tortoises encountered wandering on the road. After being
transported great distances the reptiles may escape, be liberated,
or handed over to whoever happens to put the driver up for the
night. All localities that appear questionable are either men-
tioned in footnotes or at the end of the section dealing with
the Range of the species in question.

For the convenience of those who wish to see at a glance
what species are currently known to occur in a particular coun-
try, a chart is provided (p. 179). In general the name of the
country only is given, but the Union of South Africa covers so
vast an area that its component divisions (e.g. Transvaal, Natal,
etc.) have been cited. Similarly, though not invariably, French
Equatorial and West Africa have been listed under their major
territories (e.g. Gabon, Senegal, etc.) when this appeared advis-
able.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 17^

List of the CRYPTODIRA in Africa'

Pagf.

Order TESTUDINATA

Suborder CRYPTODIRA 180

Family TESTUDINIDAE 181

Subfamily [PLATYSTERNINAE | 182

Subfamily EMYDINAE 183

Genus [Emydoidea Gray] 189

Genus Clemmys Ritgen 191

C. caspica leprosa (Schweigger) 192

Genus Emys Dumeril 201

E. orbicularis (Linnaeus) 202

Subfamily TESTUDININAE 209

Genus Geochelone Fitzinger 221

Subgenus Aldabrachclys new 225

Subgenus Geochelone Fitzinger 225

G. sidc^ita (Miller) 230

G. pardalis babcocki (Loveridge) 235

G. pardalis pardalis (Bell) 251

Genus Testudo Linnaeus 254

Subgenus Testudo Linnaeus 261

T. graeca graeca Linnaeus 261

Subgenus Pseudotestudo new 276

T. Tcleinvianni Lortet 276

Genus Malacochersv^ Lindholm 283

M. tornieri (Siebenrock) 286

1 Names in square brackets are those of non-African groups which had to be
defined or discussed for taxonomic reasons.

176 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Genus Psammobates Fitzinger 294

P. oculifer (Kuhl) 315

P. geometricus (Linnaeus) 319

P. tentorius verroxii (Smith) 324

P. tentorius trimeni (Boulenger) 333

P. tentorius tentorius (Bell) 336

Genus Chersina Lindholm ;ji2

C. angulata (Schweigger) 345

Genus Eomopus Dumeril and Bibron 352

H. signatus (SchoepfE) 359

H. houlengeri Duerden 362

H. femoralis Boulenger 365

E. areolatus (Thunberg) 367

Genus Kinixys Bell 374

K. belliana ielliana Gray 384

K. belliana nogueyi (Lataste) 'A9C<

K. homeana Bell 4U0

K. erosa (Schweigger) 404

Family TRIONYCHIDAE 412

Genus Trionyx Geoffrey 420

T. triunguis (Forskal) 423

Genus Cyclanorbis Gray 436

C. elegans Gray 438

C. senegalensis (Dumeril and Bibron) 443

C. sp 4.'0

Genus Cycloderma Peters 451

C. aubryi (Dumeril) 453

LOVERIDGE AND WILLIAMS : AFRICAN" CRYPTODIRA

177

C. frenatum Peters 459

Family CIIELONIIDAE 472

Genus Chelonia Brongniart 472

C. mydas (Linnaeus) 474

Genus Eretmochelys Fitzinger 484

E. imbricata (Linnaeus) 485

Genus Caretta Kafinesque 489

C. caretta (Linnaeus) 490

Genus Lepidochelys Fitzinger 494

L. olivaoea olivacea (Eschscholtz) 495

Family DEEMOCHELYIDAE 498

Genus DermocJielys Blainville 499

D. coriacea (Linnaeus) 499

LIST OF KEYS AND TABLES

Key to the Families of Cryptodira in Africa 180

" " ' ' Subfamilies of Testudinidae in Africa 182

" " " Emydinae in Africa 190

" " " Forms of Clemmys caspica 192

" " " Genera of Testudininae in Africa 220

" " " Species and Subspecies of Geochelone in Africa 229

" " " Subgenera of Testudo 258

" " " Circum-Mediterranean Species of Testudo 260

" " " Species and Subspecies of Psammobates 314

" " " Species of Eomopns 358

" " " Species and Subspecies of Kinixys 383

" " " Trionychidae of Africa 419

" " " Marine Turtles Breeding in Africa 471

178 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Page

Table 1. Nomenclature of the Parts of a Turtle Shell 171

Table 2. Recorded Distribution of African Cryptodira 179

Table 3. Characters of the Subgenera of Geochelone 223

Table 4. Characters of the African Species and Subspecies of Geochelone 228
Table 5. Chronological List of the Specific and Subspecific Names Pro-
posed in Psammobatcs 301

Table 6. Resemblances of Chersinn to other Genera 343

Table 7. Characters of the Species of Kinixys 38;^

Tables. Characters of the Two Sections of the Family Trionychidae . . 414

Table 9. Characters of the African Species of Trionychidae 419

Table 10. Marine Turtles Caught on the Senegal Coast 469

Recorded distribution |
of African Cryptodira -

around that contiuent ^

E

5

S3

y a-

0 0,

w 0

5 .s 'n

11 -si

^ S g S 0

a 1: tic jtf ^
^ ^ 0 0 -o

u -- — -3 0
0 0 w g ~

:^ u H H S

•= i S i
S £ ^ ~

Sill

c- a ca aJ

-O -Q -Q Lil

0 o 0 0

a c £ s
a E i E
^ S § ?

a, 0. cl, X

c

til

-s « s

S .3 -
s g 1

iei

7,

|i 1 s-g. ». .2 i 1

i ^ S -2 ° i 1 1 1 .J

II 1 & & & & 1 i s
i i g :§ 1 :s :i ■§ 1 -5

2 S

E S3 S

1 g a £ :• 3 s
S g ^ 1 1 s 5

i I 3 S :^ S 3
0 u y w J 0 e

Bio de Oro

•

Spanish ]\Iorocco •

French Morocco •

•

•

Algeria •

*

• •

Tunisia •

•

• «

Libya (Tripoli etc.)

• •

Egypt

•

•

* • ? • •

Sudan

•

•

• • • •

Eritrea

•

• •

• •

Ethiopia

•

•

• •

•Frt^nch Sonialiland

British Soroaliland

•

Somalia

•

• •

Sokotra Island

• • • •

Uganda Protectorate

•

* • •

Kenva Colony

• •

•

• • • ?

Tanganyika Territory

• «

•Pemba Island

Zanzibar Island

• •

Mafia Island

Mozambique

•

• * • •

Nyasaland

•

•

Northern Rhodesia

•

7

Southern Rhodesia

.

Bechuanaland Protectorate

•

•

Union of South Africa

(Transvaal)

•

*

(Natal)

•

(Orange Free State)

« •

•

'• Basutoland

(Cape Province)

• 7

South-West Africa

• •

• •

. ?

?

a •

•St. Helena Island

Ascension Island

• • »

Angola

•

• •

•

Cal.inda

• •

• •

Belgian Congo

• • • •

• • • •

•Belgian Ruanda-Urundi

French Equatorial Africa

• • •

• • * » • • «

Rio Muni

» »

•Annobom Island

Sao Thom6 Island

• •

Principe Island

• •

•Fernando Po Island

French Cameroon

• •

British Cameroon

• • •

•

Nigeria

4

• • •

c

Dahomey

•

Togo

• • •

• •

Gold Coast

• ft • «

• • • •

Ivory Coast

« a

•

Liberia

• • «

a • m •

Sierra Leone

• • «

•

French Guinea

•

?

Portuguese Guinea

» •

»

Gambia

•

m • m

*

Senegal

•

m A

• ft • « » •

French West Africa

• 9

• *

Cape Verde Islands

• • •

Canary Islands

• •

Madeira Islands

«

'

' N« ramrdfi koo»o tfl lu

Table 2

180 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

SYSTEMATIC DISCUSSION
Suborder CRYPTODIRA

1870. Cryptodvra Cope (part), Proc. Amer. Assoc, Adv. Sci., 19:235.

Definition. Skull with temporal region frequently emarginate
behind ; parietal usually not in contact with squamosal ; nasals
rarely, and lacrimals never, present; prefrontals almost always
joining vomer; epipterygoids usually present; pterygoids with-
out lateral rolled-up expansions, always excluding basisphenoid
from quadrate ; quadrate never enclosing eustachian tube ; artic-
ular region of mandible concave; splenial rudimentary or ab-
sent; dentaries always united.

Head withdrawn by a vertical flexure of the neck ; atlas rarely
fused to odontoid; cervical vertebrae with rudimentary trans-
verse processes situated anteriorly ; posterior cervical spines low ;
cervical postzygapophyses widely separated; central cervical
articulations well developed, posteriorly broad and (usually)
double ; sacral ribs well developed ; pelvis never ankylosed to
carapace or plastron; caudal vertebrae usually procoelous.

Shell primitively complete (except for the always absent meso-
plastra), variably reduced or its elements lost in specialized
forms; horny shields present or absent; if absent, an epithecal
component of the bony shell greatly developed.

Key to the Families of Cryptodira in Africa

(Based on external characters)

1. Limbs not modified as flippers, each with 3, 4 or 5 claws 2

Limbs modified as flippers, with 0, 1 or 2 claws 3

2. Carapace with horny shields; feet with 4 or 5 claws. . TESTUDINIDAE

(p. 181)
Carapace without horny shields ; feet wtih 3 claws . . TEIONYCHIDAE

(p. 412)

3. Carapace with large horny shields ;i flippers with 1 or

2 claws CHELONIIDAE

(p. 472)
Carapace without large horny shields ; flippers clawless

DEEMOCHELYIDAE
(p. 498)

1 Shields were absent on a single aberrant adult Chelonia mpdas captured at
Karaiiuva Island ( 13. xi. 19:^8) where it was examined, while still alive, by
Deraniyagala (1939: 227).

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 181

Family TESTUDINIDAE

1825. Testudinidae Gray, Ann. Philos. (2), 10:210.

Definition. Cryptodirous testudinates, semi- or fully aquatic
or terrestrial in habit. Horny shields always present; costal
scutes almost always 4 pairs; marginals exclusive of the nuchal
and supracaudal, usually 11 pairs; inframarginals complete or
reduced to axillary and inguinal scutes; plastron with 6 pairs
of scutes; normally no intergulars.

Skull without nasal bones; prefrontals always in contact dor-
sally, with descending processes that may be scarcely or wddely
separated inferiorly; parietal never meeting squamosal; either
postorbital or quadratojugal sometimes absent; temporal region
posteriorly emarginate or not; quadrate enclosing or not enclos-
ing stapes; post-otic antrum well developed; upper jaw fre-
quently with one or more ridges on its triturating surfaces ;
vomer always present, separating palatines ; mandible W'ith well-
developed coronoid bone.

Neck vertebrae usually with 2 biconvex centra, typically the
eighth centrum doubly convex in front; coracoids with median
borders narrow or markedly widened ; tuberosities of humerus
widely separated to closely approximated; trochanteric fossa of
femur tending to be reduced; phalanges with condyles; claws
normally 4 or 5.

Carapace without epithecal component, united to plastron by
suture or ligament; sometimes either carapace or plastron with
more or less of a hinge ; nuchal without well-developed costiform
processes ; typically peripheral and pleural bones solidly united ;
neural bones variably shaped ; pygals usually 3 ; plastron never
cruciform, usually without fontanelles in adults (except in
Malacochersus) ; entoplastron always present; buttresses very
strong to absent.

Range. "World-wide except for the x\ustralian region.

Remarks. The conception of the TESTUDINIDAE to which
we have adhered is that of Williams (1950), in which the family
includes the Platysterninae together with the Emydinae and
Testudininae. Though only two of these subfamilies occur in
Africa, the family definition has been phrased so as to embrace
the genus Platysternon and the definitions of Emydinae and
Testudininae drafted so as to afford contrasts with that Asiatic

182 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

genus. In order to clarify the statement of some of the charac-
ters cited, it has seemed advisable to present, in addition to the
diagnoses of the African subfamilies, one of the Platysterninae
also.

Subfamily PLATYSTERNINAE

1950. Platysterninae Williams, Bull. Amer. Mus. Nat. Hist., 94:513, 556.

Definition. Aquatic terrapins. Skin of head undivided ; scales
on f orelimb neither spurlike nor with dermal ossifications ; small
spurlike tubercles on hinder side of thigh ; scent glands present ;
bursae anales present.

Skull with prefrontals whose descending processes are closely
approximated inferiorly; frontal excluded from orbit; postor-
bital enormously developed, largely covering temporal region;
temporal region posteriorly but very slightly emarginate; tem-
poral arcade very solidly constructed; quadratojugal large, in
contact with the maxilla; jugal not entering orbit, surrounded
by other bones; quadrate not enclosing stapes; surangular
largely covered \)y dentary, only narrowly exposed laterall3^

Coracoids with narrow medial borders ; tuberosities of humerus
widely separated; trochanteric fossa of femur widely open;
epipodials moderate; second and third digits always have 3
phalanges.

Carapace very low; rib heads stout, well developed; neural
bones rather quadrate; inframarginal series of scutes complete.

Range. Southeastern Asia.

Key to the Subfamilies of Testudinidae in Africa

Skin of head smooth and undivided ; digits fully webbed,
the second and third with 3 phalanges. Range:

ponds and streams of northwest Africa

EMYDINAE
(p. 183)
Skin of head divided into larger or smaller shields;
digits not webbed, the second and third with only 2
phalanges. Range : all of Africa in suitable areas . .

TESTUDININAE
(p. 209)

LOVERIDQE AND WILLIAMS : AFRICAN CRYPTODIRA 183

Subfamily EMYDINAE

1909. Emydinae Sicbenroek, Zool. Jahrb. Syst., Suppl., 10:451.

Definition. Testudinid terrapins of a<inatic to fully terrestrial
habits. Skin of head entirely smooth or posteriorly divided into
small shields ; scales on f orelimb sometimes bandlike, never spur-
like or with osteoderms; spurlike tubercles on liinder side of
thig'h absent or very small; scent glands present; bursae anales
present, occasionally rudimentary.^

Skull with prefrontals whose descending processes are typi-
cally very closely approximated inferiorly, at most only moder-
atel}" separated ; frontal often entering orbit, sometimes entering
temporal fenestra ; postorbital typically well developed, never
absent; temporal region posteriorly always emarginate; tem-
poral arcade sometimes interrupted by reduction or absence of
the quadratojugal ; quadratojugal tending to be reduced or ab-
sent; quadratojugal and maxilla rarely in contact; quadrate
usually not enclosing stapes ; surangular largely covered by den-
tary, only narrowly exposed laterally.

Coracoids with narrow medial borders ; tuberosities of humerus
usually widely separated ; trochanteric fossa of femur usually
widely open, except in the more terrestrial forms where it tends
to be narrowed or reduced ; epipodials moderately elongate ;
second and third digits always with 3 phalanges, except on the
forefeet of the American genus Terrapene, where there are only
2, as in tortoises.

Carapace usually low arched ; rib heads typically well devel-
oped, sometimes much elongated; neural bones primitively hex-
agonal, short-sided anteriorly, sometimes short-sided posteriorly,
rarely octagonal, never vestigial ; of the inframarginal series only
the axillaries and inguinals normally present.

Range. North and South America; Europe; Asia; North
Africa. Absent from tropical and South Africa and the Aus-
tralian region.

Fossil record. Throughout Cenozoic of North America, Europe
and Asia; first reported in the Pliocene of Africa; not known
fossil in South America.

1 r.ursap anales were not found in Cyclctnys (— Pymdea) mouhoti by J. Ander-
son, 187«, .Tour. Linn. Soc, 12: 434-444.

184 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Remarks. Only two emydine genera occur in Africa. A study
of the African testudinid fauna is, tlierefore, not an appropriate
place for a revision of emydine genera. Nevertheless we have be-
come involved in the problem of their definition and grouping
since the distribution of one of the African genera (Eniys), as
currently understood, includes a species (orMcularis) from
North Africa, Europe, and neighbouring portions of Asia, and
a second {Mandingii) in eastern North- America. An investiga-
tion of the evidence for this peculiar distribution has convinced
one of us (E.E.W.) that the two species are not congeneric.

The basic reasons for this decision were in fact stated by Baur
in 1889 (Amer. Nat. 23: 1099-1100), and will become evident to
anyone who studies skeletons of Deirochelys reticularia, "Emys"
hlandingii and Emys orbicularis.

A decision of this sort involves a sorting out of convergent and
phylogenetically meaningful characters. It is best done in a
frame of reference that includes a conception of the relationships
of all emydine genera. Figure 2 is an attempt to provide this
appropriate background. The scheme given there is tentative ;
it is not a phylogeny since it neglects the temporal dimension
and omits all fossil forms (disregarded for this subfamily, as
they are poorly known and worse analysed). However, the dia-
gram, wherever it indicates the direct derivation of one genus
from another does predict that the actual ancestral fossil wiU
"key" out very near the living genus which has been given the
more central and thus "primitive" position.

In Figure 2 we have postulated an unknown (presumably Asi-
atic) ancestral emydine, inferring as the characters of this hypo-
thetical animal those which would most closely approach the
characters found in related and primitive families (estimating
primitiveness from the characters held in common by the older
fossil families). On this basis we consider that the ancestral
emydine had:

1. a carapace with surface sculpture ;

2. a skull with a strong temporal arcade ;

3. moderate axillary and inguinal buttresses;

4. maxillae with rather wide triturating surfaces but no
secondary palate ;

5. quadrate not enclosing the stapes;

Cuora ^^

0=,i,(3/i,or^-v^

Cyclemys '^^^
0.-1,(3/1/0) >.

Terrapene ^^ ^ Geoemyda As,NA3A

e=, 1,(1/0) 9.1,(3/1)

^-•'•"'°' ^/5/77/77;rE^As.Af.NA^
0,1,(3/1/0}

Annamemys ^'^ \

6,1.(3)

/ proto-\

Tesfudininae .^^J emyoinel
Eu,As>:\f.NA3A V_-X

Hieremys ^

9.1,(1)

.Hi+--I'-I

Deirochelys ^^

/ 1,(1/0)

Emydoidea ^^

Notochelys^^
0.-1.(1)

Chine mys^

0,11,(3/0

Geoclemys As

11.(3)

Morenia ^*

-w-.d)

Pseudemys ^^^^

^.(1/0)

Chrysemys "^

,1' (0)

Groptemys ^^
i/il,(i)

J,»

Kachuga ^

Bafagur^^

.0)

— , 1,(0)

-^^,^ strong buttresses 0 enfoplastron crossed by humeropectorol sulcus

-^ -w^ moderate buttresses ii wide triturating surfoces, unridged

"semi box" tortoise i narrow triturating surfaces , unridged

= "full box" tortoise ,1- weakly ridged surfaces 'Tstrongiy ridged surfaces

j^ very wide triturating surfaces with one or more

ridges.

Fig. 2. Dendrogram of emydine relationships. With the exception of Emy-
doidea, the names are those of currently accepted genera. Numerals in
parentheses refer to the number of keels on the carapace. NA, SA, Eu.
Af , As = abbreviations for the continents on which these genera occur.

(P. Washer del.)

186

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

6. the entoplastron anterior to the humeropectoral sulcus;

7. moderate maxillary triturating ridges;

8. three keels on the rather flat shell.

The listing of these characters is somewhat in the order of our
confidence in them. Pseudemys in the Western Hemisphere, and
Ocadia in the Eastern, are living forms possessing most of these
characters and so in our estimation closest to the postulated an-
cestral em^'dine. From the vicinity of these two central types
there radiate, according to this view, three major phyletic lines,
each with certain characteristic trends, and each with their sub-
sidiary radiations.

Fig:. 3. Condition of the prefrontal descending process. A. typical eniydine
(Clemntys in.'iCHlpfa) ; B. typical tostudinine (Gopherus agassizii).

(S. McDowell del.)

The land tortoises, we believe, form a unified series diverging
at a very earl}'^ date from the emydine stock. They appear to be
united by certain parallel trends in the scalation of the head,
thighs, and lower limbs; by the universal extreme weakness or
absence of vertebral keeling ; by the downgrowths of the f rontals
underarching the olfactory tract; and by a wide separation of
the ventral processes of the prefrontals (in contrast to the usual
median approximation of these processes in emydines, see Fig.
3). Further discussion of this lineage is given below (pp.
211-219).

Within the emydines, after the separation of the tortoise line,
one major group appears to be primarily North American. This

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 187

group — Fseudemys and its relatives — never has a tricarinate
carapace. One subgroup (Chrysemys) loses the surface sculp-
ture of the shell and weakens the ridging of the maxillary trit-
urating surfaces; another subgroup Malaclemys and Graptemys
acquires a tuberculate condition of the median keel and a greater
or lesser development of a secondary palate, while the subgroup
Deirochelys-Emydoidca narrows the triturating surfaces, loses
the maxillary ridging, elongates the head and neck, strengthens
the extensor muscles of the neck with a corresponding bowing
out of the carapace ribs at their point of attachment to the col-
umn and, finally, in Emydoidea itself achieves a plastral hinge.

The second major division of the emydines is primarily Asiatic
and primitively tricarinate. This series subdivides immediatelj'
into several lines.

One lineage is that of the ''river turtles" or "diving turtles"
of Asia. In all, the skulls have a very characteristic habitus with
a rather angular, somewhat elongate, shape and a tendency to
an upturned snout ; all have an extraordinarily developed sec-
ondary palate with strong maxillary ridging. Typical members
of the group have extreme development of the shell buttresses,
with a resulting internal partitioning of the shell far more ex-
treme than in any other emydines — with the possible exception
of Annamemys. Of the entire group only Callagur shows any
trace of lateral keels on the carapace. In skull structure and gen-
eral habitus Morenia is an obvious member of this lineage, but it
has diverged radically in having the buttresses very poorly
developed.

A second lineage within the major group is that of Orlitia and
Siehenrockiella, a small and relatively primitive group, lacking
the evident specializations of the other lines and sharing minor
liabitus features.

A third lineage comprises the group of Asiatic genera (united
by Boulenger as " Damonia") which are strongly tricarinate
with reduced buttresses and wide unridged triturating surfaces.

The most successful of the primarily Asiatic lineages is that
to which Ocadia and Clemmys belong. This series has invaded
Europe, North Africa; North America and (as Geoemyda) even
South America.

188 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

This lineage has as its conspicuous specialization the combina-
tion of narrowing of the triturating surfaces of the skull with
a bringing forward of the humeropectoral sulcus of the plastron
so that it lies across the entoplastron well in front of the latter 's
posterior border. This series begins with Ocadia, a form with
moderate buttressing, and proceeds in Annamemys to a second-
ary extreme development of buttresses. However, on four
separate occasions (Emys, Terrapene, Notochclys and Cyclemys-
Cuora) the same series has given rise to forms with hinged
plastra and consequent loss of buttresses.

According to this interpretation Emys orhicularis is a close
relative and, in fact, derivative of the genus Clemmys, while the
species hlandingii, here referred to the genus Emydoidea Gray,
is considered very remote from orbicularis and Clemmys, being,
instead, a close relative and derivative of the genus Deirochelys.

There are undoubted resemblances between hlandingii and
orbicularis and equally undoubted resemblances between hlan-
dingii and Deirochelys reticularia. One or the other of these sets
of resemblances must be convergent.

The resemblances between orhicularis and hlandingii are four
in number :

1. coloration of the carapace ;

2. the plastral hinge ;

3. narrow maxillary triturating surfaces;

4. quadrate tending to enclose the stapes.

The last two are resemblances also to Deirochelys. In the face
of any contradictory evidence the color character would not be
regarded seriously by anyone, especially as the plastral colora-
tion is quite different in the two species. The plastral hinge and
the enclosure of the stapes are phenomena which have repeatedly
occurred; the hinge has an evident adaptive value and the en-
closed stapes may similarly convey some selective advantage.

The resemblance between hlandingii and reticularia are gen-
erally less easy to verbalize consisting of characters in the general
habitus of sliell and skeleton. However, certain of these lend
themselves to ready statement :

1. Both have an elongate head and neck, this elongation
being more extreme than in any cryptodirous turtles ex-
cept the trionychids.

LOVERroGE AND WILLIAMS: AFRICAN CRYPTODIRA 189

2. Both have extremely narrow maxillary triturating sur-
faces with a very similar, geutlj^ arching, external contour.

3. Both have the orbits wholly exposed dorsally with corre-
spondingly narrow interorbital space.

4. Both have the dorsal rib heads very slender and greatly
arched, thus accommodating very powerful extensor
muscles of the neck.

Dealing with these characters in the same order we offer the
following comments :

1. Apparently no other emydines exhibit any tendency to
elongation of head and neck, i.e., unlike plastral hinging,
this is not a repeated trend.

2. This character exemplifies one of the many striking sim-
ilarities in skull shape that are greater and more detailed
than any to be expected except in cases of direct relation-
ship.

3. In this orbital character it might be noted that blandingii
and orbicidaris are poles apart. This character also serves
to separate broad groups within the Emydinae, the
Clemmys series tending to have the orbits covered, the
Pseudemys group tending to have them exposed dorsally.

4. The slender arched ribs are undoubtedly correlated func-
tionally with the elongate neck, but the similarity in
detail between hlandingii and reticularia is far greater
than between these and such a form as Chelydra in which
powerful cervical extensors also occur.

In order to further clarify some of the modifications in our
definition of Emys, made necessary by our new restricted concept
of the genus, we append a definition of Emydoidea.

Genus Emydoidea Gray

1870e. Emydoidea Gray, Suppl. Cat. Shield Kept. Brit. Mus., part 1, p. 19.

Type: Cistiida blandingii Holbrook (by nionotypy).
1929. Neoemys Lindholm, Zool. Anz., 81:282 nom. nov. for Emydoidea

Gray, considered unavailable on nomenclatorial grounds.
Definition. Beak never hooked, inesially notched; skin on
hinder part of head smooth, undivided ; back of thighs without
tubercles ; digits fully webbed.

190

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Skull with orbits fully exposed in dorsal view ; triturating sur-
face of maxilla narrow, without ridges; anterior palatine fora-
mina small ; posterior palatine foramina large ; temporal arcade
complete ; quadrate enclosing stapes.

Carapace smooth, never with vermiculate sculpture; neurals
hexagonal, short-sided in front, broader than long; buttresses
absent; rib heads slender and strongly arched to accommodate
the strongly developed extensors of the long neck.

Plastron united to the carapace by ligament, a more or less
distinctly developed hinge between hyo- and hypoplastra: ento-
plastron posteriorly touching, though usually not crossed by, the
humeropectoral sulcus.

w

Fig. 4. Plastral shapes of juvenile African Emydiuae. A, Clemmys caspica
leprosa (M.C.Z. 53798), B, Emys orbicularis (M.C.Z. 1896).

(P. Washer del.)

Range. A limited area in eastern North America.
Fossil record. Known only from the Pleistocene of North
America a little west of its present range.

Key to the Emydinae in Africa

Plastron never hinged ; anals pointed, their median suture always shorter
than the interabdominal suture (cf. Fig. 4a) . .Clemmys caspica leprosa

(Schweigger) (p. 192)

LOVERIDGE AND WILLIAMS: AFRICAN CBYPTODIRA 191

Plastron hinged in adults; anals rounded, their median suture ahvajs longer

than the interabdominal suture (cf. Fig. 4b) Emys orbicularis

(Linnaeus) (p. 202)

Genus ClEMMYS Ritgen

1828. Clrm)iiy.<i Ritgen, Nova Acta Acad. Leopold. -Carol., 14:272. Type:

Tcstudo puvctnta Schocpff = T. r/uttata Schneider (designation by

Stejneger 19(t7, Herpetology of Japan, p. 492).
1832. Chelopus Eafinesque, Atlantic Jour., 1:64. Tj-pe: Testudo guttata

Schneider (designation by Stejneger).
1857. Naiirmjis Agassiz, Contr. Nat. Hist. U.S. ,1:442. Type: Testudo

guttata Schneider (by monotypy).
1857. Calemys Agassiz, Contr. Nat. Hist. U.S., 1:44.3. Tj^e: Testudo

■muhlenbergii Schoepff (by monotypy).
18.57. Glyptemys Agassiz, Contr. Nat. Hist. U.S., 1:443. Type: Testudo

insculpta Leconte (by monotypy).
1869d. Mauremys Gray, Proc. Zool. Soc. London; 500. Type: Emys fuligi-

nosa Gray (by original designation).
1870e. Geodemmys Gray {not Geoclemmys Gray: 1855), Suppl. Cat. Shield

Rept., part 1, p. 26. Type: Testudo guttata Schneider (designation

by Stejneger.)
1870e. Sacalia Gray, Suppl. Cat. Shield Rept., part 1, p. 35. Type: Cistuda

healeiGray (by monotypy).
1870e. Emmenia Gray, Suppl. Cat. Shield Rept., part 1, p. 38. Type: Emys

grayi Gunthev =: Testudo caspica Schweigger (by monotypy).
1870e. Eryma Gray, Suppl. Cat. Shield Rept., part 1, p. 44. Type: Emys

laticeps Gt&j ^ Emys leprosa Schweigger (by monotypy).
1919. Melanemys Shufeldt, Aquatic Life (Philadelphia), p. 155. Type:

Testudo guttata Schneider (designation by Dunn, 1920. Copeia,

p. 8).
Definition. Beak never hooked, not or mesially notched; skin
on hinder part of head smooth, undivided ; back of thighs Avithout
tubercles ; digits fully webbed.

Skull with orbits largely concealed in dorsal view ; triturating
surface of maxilla narrow, without ridges ; anterior palatine fora-
mina small; posterior palatine foramina large; temporal arcade
complete ; quadrate not enclosing stapes.

Carapace smooth or with concentric grooves, never with ver-
miculate sculpture; neurals hexagonal, short-sided in front, as
long as, or longer than, broad; buttresses short, extending only
to the outer margins of the pleurals (costal bones) ; rib heads
short, not slender or strongly arched.

192 BULLETIN : MUSEUM OF COMPAKATIVE ZOOLOGY

Plastron -united to the carapace by suture, without a hinge;
entoplastron crossed by the humeropectoral sulcus well in ad-
vance of its posterior border.

Bange. North America, North Africa, Europe and Asia.

Fossil record. Cenozoic of North America, Europe, Asia. The
earliest occurrence in Africa is in the Pliocene of Algeria.

Key to the Forms of Clemmys caspica^

Plastral pattern of both young and adults predominantly dark brown or
black, the bridge always dark with at most some small yellow spots or
streaks. Eange : South Dalmatia ; Greece ; Ionian Islands ; Crete^ ;
Cyprus; Asia Minor (only in west and south) ; Syria c. rivulata

Plastral pattern of both young and adults consisting of a dark brown or
black blotch covering most of each shield but leaving a yellow margin,
at least laterally; bridge predominantly yellow with only the sutures
between the shields dark. Range: Asiatic Eussia; Asia Minor (middle
and east) ; Iraq and Iran c. caspica

Plastral pattern in young consisting of a dark brown or black blotch occupy-
ing the entire medial area but leaving a wide yellow margin laterally;
bridge yellow with two large dark brown blotches usually coalescing;
in adults all markings become obsolescent. Eange: Iberian Peninsula
and northwest Africa (Morocco to Tunisia) south to the central Sahara,
west to the Gambia (i.e. about 13°N) c. leprosa

Clemmys caspica leprosa (Schweigger)

1812. Emys leprosa Schweigger, Konigsberger Arch. Naturw. Math., 1:298,

339 ; No locality.
1814. Schweigger, 29 (reprint of 1812).

1862b. Strauch, 18.

1894. Oliver, 101.

1896b. Oliver, 118.

1899. Doumergue, 247, pi. i.

1903. Mayet, 10.

1904. Chaignon, 2.
1907. Barbier, 73.
1919. Leblanc, 177.

1824. Emys marmorea Spix, Animalia nova Species testud. ranarum Brasil

. . ., p. 13, pi. x; Xo locality.
1831c. Gray, 25.

1 We are without material of C'. v. cretica Mertens (1946, Senekenbergiana, 27:
115) of Crete, which appears to differ but slightly from c. rivulata.

LOVERroGE AND WILLIAMS: AFRICAN CRYPTODIRA 193

1835. Dum^ril and Bibron, 248.

1829. Clemmys Sigris Mk-hahelles, Isis von Oken, col. 1295; Spain.

1831. Terrapene sigris Bonaparte, 87 (translation of last).

1831c. Emys vulgaris Gray, Synopsis Eeptilium, p. 24, pi. iv: Southern

Europe.
1831b. Gray, 9 (Spain).

1841. Schlegel, 108.

1860. Tristram, 405 (Sahara).

1833. Les Emydes Eozet, 231.

1835. Emys sigriz Dumeril and Bibron, 240.
1850. Guichenot, 2.

1854. Eichwald, 415.
1887. Lortet, 19, pi. vii.

1897. Bateman, 54.

1836. Emys luiaria Bell (part: not of Lacep^de), text and col. pis.
1844. Emys caspica Gray (part: not of Gmelin), 19.

1855. Gray, 22.
1867a. Steindachner, 5.

1854. Emys laticeps Gray, Proc. Zool. Soc. London, 1852, p. 134: Gambia

Kiver, West Africa (M. Castang coll.).

1855. Gray, 23, pi. ix.

1857. Tortue aquatique Labouysse, 83.

1860c. Emys fuliginosus Gray, Proc. Zool. Soc. London, p. 232, pi. xxx:

North Africa?

1862a. Clemmys laticeps Strauch, 32.
1865. Strauch, 75.

1884a. Eochebrune, 18.

1862a. Clemmys marmorea Strauch, 32.
1865. Strauch, 75.

1862a. Clemmys leprosa Strauch, 122.
1865. Strauch, 74.

1880c. Vaillant, 33, 88.

1889a. Boulenger, 105.

1889b. Boulenger, 306.

1890. Strauch, 69.

1891c. Boulenger, 96, 106.

1892. Anderson, 11.

1892. ■ Konig, 15.

1894. Werner, 76.

1895. Konig, 404.

1896. Escherich, 278.

1898. Jeude, 7. (Clemnys)

1900. Boettger and Tornier, 64.

1901. Gadow, 357, fig. 80 (skull).

194

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1908. Kerville, 96.

1908. Zulueta, 451.

1909. Siebenrock, 481.

1911. Lampe, 144.

1912. KoUman, 101, figs. 1, 3, 4. (Clemnys)
1912b. Pellegrin, 262,

1912b. Werner, 408.

1913. Pellegrin (1912), 420.

1914. Pellegrin, 347.
1914b. Werner, 334.
1916e. Chabanaud, 233.
1917c. Cliabanaud, 105.
1918. Seurat, 23.
1920. Mourgue, 233.
1925. Flower, 922.

1926a. Pellegrin (1925), 315.

1926e. Pellegrin, 121.

1927. Fejerv^ry, 517.

1927a. Pellegrin, 261.

1929. Dollfus, 112,

1929. Flower, 17.
1929b. Werner, 12, 15, 21.

1930. Seurat, 182.
1931c. Werner, 275.
1932. Ghigi, 208,

1934. Mosauer, 51.

1935. Laurent, P., 345.
1935. Hediger, 3.

1937. Hediger, 187, 191.

1938. Angel and Lhote, 376.
1938. Gorham and Ivy, 181.
1947. Braestrup, 5.

1950c. ViUiers, 343.

1950. Williams, 550.

1951. Aellen, 168, 195.

1952. Stemmler-Morath, 217, photos 1-2.
1953e. Girons, 76, 79.

1954. Noel-Hume, 90, 112.

1865. Clemmys fuliginosa Strauch, 76.

1869d. Maureviys laniaria Gray, Proc. Zool. Soc. London, p. 499, pi. xxxvii:

No locality (ex Zool. Gardens from a dealer).
1869d. Mauremys fuliginosa Gray, 500.

1870e. Gray, 35.

1872. Sclater, P. L., 603 (synonymizes with Zeproso).

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 195

1873b. Gray, 35.

1869e. Emys flavipes Gray, Proc. Zool. Soc. London, p. 643 ; No locality.

1869e. Emys fraseri Gray, Proc. Zool. Soc. London, p. 643 : North Africa

(ex Mr. Fraser).
1870e. Gray, 36.

1873b. Gray, 35.

1873d. Gray, 146.

1870e. Emys laniarw, Gray, 37,
1870e. Eryma laticeps Gray, 45.
1873b. Gray, 36.

1873b. Emys lamaria (sio) Gray, 36.
1874. Emys caspia var. leprosa Boettger, 126.
1882. Clemmys caspica sigris Bedriaga (1881), 340.
1883a. Clemmys caspia var. leprosa Boettger, 131.
1886. Clemmys Caspica Parenti and Picaglia (not of Gmelin), 86.
1927. Clemmyde lepi-euse Seurat, 81.

Synonymy. Schweigger (1812) cites Schoepff as the author,
having taken the name from an unpublished manuscript of the
latter. He had seen the type in the Paris Museum.

Common Names. Leprous Terrapin (preferred) ; Marbled Ter-
rapin (Gray: 1831c); Mud Tortoise (Tristram: I860); Iberian
AVater-Tortoise (Gadow: 1901); Spanish Terrapin (Flower:
1929); Fakroun-el-ma (Arabic: Doumergue: 1899).

Illustrations. This terrapin is well represented in the fine
colored plate of Lortet (1887 : pi. vii) .

Description. Beak distinctly notched; edge of jaws not or but
finely denticulated; mandibular width at symphysis less than
(young) or almost equal to (adults) the horizontal diameter of
the orbit; skin on upper surface of head undivided; forelimb
anteriorly covered with smooth, flattened shields of very diverse
size of which about 5 (3 along outer margin and usually 2 in-
ternal to them) greatly enlarged; digits webbed to the 5 claws;
hind foot strongly webbed, claws 4; tail long.

Carapace rather strongly depressed, anterior and posterior
margins not expanded, not or but feebly reverted, not serrated,
not notched — at most but slightly incurved — in nuchal region ;
nuchal moderate, Iroader than long,^ wider posteriorly ; vertebral
keel distinct in young ^ but scarcely disting-uishable in some

1 Longitudinally divided in an Algerian terrapin (Strauch : 1890).

2 Up to individuals of about 120 or 130 mm. according to Doumergue (1899:
51) who has additional notes on growth changes.

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adults; vertebrals 5, rarely 6/ broader than long, broader than
the eostals (young) or variable (adults) ; costals 4, costal (lat-
eral) keels present but interrupted and indistinct; marginals
(exclusive of nuchal and supracaudals) 11 ; supracaudal divided.
Front lobe of plastron anteriorly truncate, not or but slightly
produced, not notched, slightly concave in S $ , flat or convex
in $ 9 ; gulars paired ; pectorals wide ; axillary 1, moderate ;
inguinal 1, moderate, not in contact with, femoral ; hind lobe
broadly or deeply notched, anals pointed (cf. Fig. 4 for contrast
with Emys).

Fig. 5. Skull of Clemmys caspica leprosa (M.C.Z. 2210). Condylobasal
length 39 mm. c/i =: angular ; ari = articular ; co = coronoid ; d =: den-
tary; / = frontal; j = jugal; m = maxillary; pa = parietal; pal =:
palatine; pm = premaxillary; po = postorbital; prf = prefrontal; pt =
pterygoid; q = quadrate; qj = quadratojugal; so = supraoecipital ; sq
= squamosal; sur =surangular; v = vomer.

(S. McDowell del.)

Plastral formula: Abd > or = (p > or = or < f ) > (an >
or = or < g) > h.

Color. Carapace of young pale or dark olive to olive brown,
uniform, or each costal shield with a black-bordered yellowish
spot or streak; the vertebrals with a narrow or broad median
streak ; marginals uniform or each marbled with j^ellow to a
variable degree, frequently edged with yellow laterally. When
any pattern is present each shield is apt to be narrowly edged
with black, becoming more or less uniform in adults.

1 Six in a specimen from Ain Sefra (M.C.Z. 27343).

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 197

Plastron of young yellow with the central area dark brown
(with or without a median light line), and dark brown blotches
usually coalescing on t)ie bridge ; marginals ventrally exhibit
dark brown blotches at the sutures. Both plastron and marginals
more or less uniform without definite pattern in adults.

Head olive to olive gray, its sides streaked or vermiculated
with yellow (reddish orange), between eye and tympanum a small
round yellow (orange) spot usually present, around the tym-
panum a more or less well-defined ring of yellow (orange) ; sides
of neck olive gray with 4 or 5 longitudinal yellow (orange)
streaks; throat streaked or vermiculated with yellow (orange) ;
limbs olive streaked or vermiculated with yellow (orange).

In life, judging by the detailed description given by Bedriaga
(1882), the coloration would appear to be even more striking than
suggested by our description. However, according to Mosauer
(1934), considerable color variation occurs between juveniles
from different localities such as Gabes and Gafsa. In old indi-
viduals all bright markings tend to disappear. For further color
changes due to age, see Doumergue (1899: 52).

Breeding. Mating takes place either on land or in water,
usually on the surface though sometimes a mated pair will descend
to the bottom where they may remain immobile for several hours
(Lortet : 1887). The artificial conditions of captivity that render
hibernation unnecessary, upset the breeding behavior so that
eventually mating may take place at any time of the year. In
his eagerness to secure a mate the S is apt to bite at the collarlike
folds of skin within which the 9 withdraws her head. His re-
peated attentions result in sores and swellings that in due course
prevent the 9 from wiping her eyes with the back of a forefoot —
a practice common to many species. Ultimately the eyes fester
and the terrapin becomes almost blind, gives up feeding, and
leaves the water in a condition that is very difficult to cure
(Gadow: 1901).

Early May, according to Lortet (1887) is the usual month for
laying. Doumergue (1899), however, records that between the
evening of September 2 and the following morning a large Al-
gerian 9 laid 9 white eggs ranging in size from 21 x 34.5 to
21 X 38 mm. Usually 6 to 8 are deposited in shallow excava-
tions in mud, sand, or between the roots of a tree. Twenty-five

198 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

days after the eggs have been laid, hatching takes place (Lortet :
1887). In Tunisian oases, towards the end of May, hatched-out
eggshells were encountered in small holes, chiefly between the
roots of trees, bordering the waterholes. No hatchlings were
seen, however (Mosauer: 1934),

Upon hatching, the young immediately hasten to the water
where few remain near the surface or venture to bask on the
rocks; instead they dive to the bottom where they hide in the
mud. There, by means of a net, large numbers can be captured
with comparative ease (Lortet: 1887). The carapaces of many
young measured on April 13, 1898, were 30 mm. ; by mid- August
they had increased to 45 mm. (Doumergue: 1899).

Longevity. Despite their dispensing with seasonal rest as a
result of congenial climatic conditions, most of Gadow's many
captives lived ' ' with undiminished appetites for more than twelve
years" (Gadow: 1901). A terrapin in the Giza Zoological Gar-
dens lived for 5 years, 1 month, and 10 daj^s; another in the
London Zoo only survived 4 years, 6 months, twelve days
(Flower: 1925b).

Diet. Hatchling terrapins are eaten by old ones (Doumergue),
though the principal food of this species consists of frog's, toads,
tadpoles, fish worms, aquatic insects and their larvae (Lortet).
Twenty terrapins were observed feeding on the corpses of horses
that had fallen into the water (Rozet: 1833). In the absence of
animal food these essentially carnivorous reptiles will eat algae
and aquatic plants (Labouysse: 1857), lettuce, legumes and
scraps of bread (Doumergue) . By seizing bait and getting hooked
they constitute a nuisance to fishermen (Labouysse).

Clearly Lortet errs when he says that the prey is always de-
voured under water, for Stemmler-Morath (1952) often found
some feeding on Arab excrement on the far side of an earthen
wall at least a yard from the nearest water. Another half-dozen
were found in a dry ditch used by natives as a toilet. They were
in good health and were said to have lived there as long as a local
farmer could remember.

Parasites. The name leprosa refers to the leprous appearance
of the carapace resulting from attacks by freshwater algae, to
which this terrapin is especially prone owing to its manner of
life. When the mud-encrusted reptile emerges from its slimy

LOVERIDGE AND WILLIAMS : AFRICAN CBYPTODIRA 199

pool to bask in the hot sun, its horny shields tend to become
brittle and flake off. Entering through the cracks, the algae
flourish in the Malpighian layer and even in the underlying bone,
which becomes gangrenous in places. Terrapin inhabiting perma-
nent lakes or watercourses are not subject to attack and remain
as clean as other species (Gadow: 1901). Stemmler-Morath
(1952) remarks that these green and brown algae may attain a
length of as much as 50 mm., and were present on all the speci-
mens caught by him. Seurat (1918) has recorded three species
of nematodes {Camallanus fnicrocephalus ; Falcaustra lambdi-
ensis, and Spiroxys contortus) from Algerian terrapin.

Enemies. Hundreds were on sale in the fish market at Algiers
(Strauch: 1865). As previously mentioned, small terrapin are
eaten by the adults (Doumergue: 1899). At Rabat one terrapin
was recovered from the stomach of a heron (Pellegrin : 1926a).

Defense. Apart from seeking safety by burying themselves in
the muddy bottom of their habitat, the chief defense of these
terrapin would seem to be their odoriferousness. This derives
from a large pair of inguinal glands that open just behind the
plastral bridge. Freshly caught leprosa emit a powerful stench,
but captive specimens cease to do so after becoming accustomed
to being handled ( Gadow : 1901).

Apparently the odor may be influenced by diet, for at times
it reeks of fish and an excess of animal food increases its intensity
(Werner: 1912b). Doumergue (1899) claims that the smell is
scarcely noticeable in winter and early spring, but we reject his
deduction that the odoriferousness of these terrapins derives from
the mud in which they live, and tends to disappear when they
are kept in clear water.

Temperament. Excessively w^ary, scrambling into the water
when anyone approaches to within ten or five meters. Such be-
havior makes their capture difficult. Why they should be so
nervous is not readily explained, for Moroccans do not molest
them and no other potential enemy is in evidence (Stemmler-
Morath: 1952). This opinion is certainly inapplicable to Alger-
ian Arabs if Strauch 's statement (1865) holds good today (cf.
Enemies).

Aestivation and Hibernation. During the hot summer months
(August-September), as ponds and watercourses shrink in the

200 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

torrid heat, these terrapin crowd into the remaining pools. There
their numbers contribute to the daily increasing foulness of the
stagnant water until everything edible is consumed. Then the
reptiles leave the pool to seek shelter among the rocks where they
bury themselves until aroused by the return of rainy weather.

During the winter months (November-March) hibernation
occurs but is intermittent (Lortet: Doumergue : Aellen). These
terrapin neither aestivate nor hibernate in captivity when tem-
perature conditions are congenial and a current of warm water is
kept circulating through their tanks. However, Gadow's terra-
pins showed a fondness for the hot-water pipes against which
dozens of them would huddle until their shields, and even the
plastral bones, suffered from the excessive heat. To prevent re-
currence of such injuries it was found necessary to keep the rep-
tiles away from the pipes by screening. Some leprosa in an
English garden successfully passed the winter under a heap of
moss and rubbish ; others remained in the mud beneath the ice in
a deep concrete pond where they survived several very severe
winters to emerge each spring in perfect health (Gadow: 1901).

Habitat. Many early travellers — from Rozet (1833) onwards
— record the astonishing abundance of leprosa in Algeria, where
scarcely a pool, stream or river is without some terrapins. Werner
(1894), however, considered it less common in east Algeria than
in the west. In Morocco they are present in almost incredible
numbers in all bodies of water except the most temporary of
desert streams ( Stemmler-Morath : 1952). The latter writer
found terrapins living in the swift streams of the High Atlas,
an observation that appears to qualify Werner's statement that
leprosa prefers level country. He also found half-a-dozen indi-
viduals living in a ditch some distance from water, as is recorded
under Diet.

Localities. Spanish Morocco: *Larache (U.S.N.M.) ; Tangier
(Tanger). French Morocco: Ain el Auda; Behalil near Sefrou;
Berguent; Bin el Ouidan (Ouidane) ; Casablanca ; Da'iet el Roumi ;
Fes (Fez) ; Mogador; *Oudjda; Oued Akrech (Akreuch), south
of Rabat; Oued Berkine; Oued Fes; Oued Ifrane, middle Atlas;
Oued Imoughoud (Tmoughout), south of Taza; Oued Ksib near
Mogador ; Oued Liboud ; Oued Sebou ; Oued Sous ; Oued Tensif t ;
Oued Tiflet ; Oued Yguem ( Yquem) ; Oued Za ; Ouezzane ; Rabat ;

LOVERIDGE AND WDjLIAMS : AFRICAN CRYPTODIRA 201

Sehoul; *Sidi Yahia; Sous; Taourirt; Tetouan (Tetuan). Al-
geria: Ain el Bahir ; Ai'n el Had jar ; Ain Sefra ; Ain Temoucheut ;
*Algiers (Alger); Arba; Arlal; Aumale; Batna; Bedeau;
*Biskra (U.S.N.M.) ; Biskra to Constantine; *Bone; Bordja
Saada; Boudsareah; *Bou Saada; Cheliff (Chelif) ; Constantine;
Der Kaid Embarek (Embareck M'Toughi) ; El Khreider; Gery-
ville; Hamman Salahine near Biskra; Lake Fetzara (Tetzara;
Tetzura) ; Macta ; Medea ; Oran ; Ouargla ; Oued Andalouses at
Bredeah ; Oued Asseila ( ? Asselar) ; Oued Baeeaura ; Oued el
Biodli ; Oued el Harrach ; Oued Saf saf at Tlemcen ; Oued Saida
at Ain el Hadjar ; Oued Sebaou near Tizi Ouzou ; Oued Sig (Zig) ;
Oued Tafna at Sebdou; Oued Tlelat; Perregaux; Rhadames;
Sahara; Touggourt; Ziban Oases. Tunisia: Ain Draham ; Ain
Hameraia; Bir Mcherga to Zaghouan; Djerba Id.; Douirat
(Duirat) ; El Hamma (Hamman) near Tozeur; Gabes; Gafsa;
Ischkeul ; Kairouan (Kairwan) ; Maxula Rades; Oued Bagra;
Oued Debbane; Oued el Amor; Oued el Mahdi; Oued Gued-
ouiaris; Oued Leben; Oued ]\Iiliani (Milane) ; Oued Oum Mela;
Oued Rzella ; Oued Siliana; Soliman; Tabarea; Tozeur (Touzla;
Tozzer) ; Zaghouan. Libya: Murzuch (Murzouk) in Fezzan;
Tripolitania. French West Africa: Adrar des Iforas; Agades
(Agadez), Air region; Fort Gouraud or Idjil (probably Koudja
d'Idjil), Mauretania. Gambia: Gambia River (Gray: 1853).

Range. Iberian Peninsula and north Africa (Morocco to Libya)
south to the central Sahara, west to the Gambia (i.e. about
13°N.). We reject the record of Porto Novo, Dahomey (Cha-
banaud: 1917c). ^

Genus EmyS Dumeril

180fi. Emys Dumeril (part), Zool. analytique, p. 76. Type: Testudo lutaria

Linnaeus = T. orbicularis Linnaeus (designation by Fitzinger,

1843, Syst. Kept., Part 1, p. 29).
1814. Eydrone Eafinesque, Specchio Sci. (Palermo), 2, p. 66. Type: T.

orhicnlaris Linnaeus (by present designation). 2
1844. Lutremys Gray, Cat. Tortoises, Crocodiles, Amphisbaenians in Brit.

Mus., p. 31. Type: Testudo europaea Schneider (by monotypy).

1 This record should be discarded. Dr. Chabanaud (1917c :105) who identified
the two specimens, informs us (22.xii.5."')) thnt they were subseciiiently destroyed
with other material identified at that time. Furthermore nothing concerning the
itinerary of the donor (Waterlot) can be found in the Museum records at Paris.

2 The included species were T. orbicularis, lutaria and geometrica.

202 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Definition. Beak never hooked, mesially notched; skin on
hinder part of head smooth, undivided ; back of thighs without
tubercles ; digits fully webbed.

Skull with orbits wholly concealed in dorsal view ; triturating
surface of maxilla narrow, without ridges ; anterior palatine
foramina small; posterior palatine foramina large; temporal ar-
cade complete ; quadrate barely enclosing stapes.

Carapace smooth or with concentric grooves, never with ver-
miculate sculpture; neurals hexagonal, short-sided in front, as
long or longer than broad ; buttresses absent ; rib heads stout, not
slender or strongly arched.

Plastron united to the carapace by ligament, a more or less
distinctly developed hinge between hyo- and hypoplastra ; ento-
plastron crossed by the humeropectoral sulcus well in advance of
its posterior border.

Range. North Africa, Europe and contiguous parts of Asia.

Fossil record. Most fossil species assigned to this genus do not
really belong to it. Perhaps the only certain records are those
from the Pleistocene of Europe.

Emys ORBICULARIS (Linnacus)^

1758. Testudo orbicularis Linnaeus, Syst. Nat., ed. 10, 1. p. 198: Southern

Europe.
1766. Linnaeus, 351.

1758. Testudo lutaria Linnaeus (part), Syst. Nat., ed. 10, 1. p. 198:

' ' Italia, Oriente. ' '
1783. Testudo Earopaea Schneider, Naturg. Schildkroten, p. 323: most

• countries in Europe.

1819. Bojanus, 1-178, figs. 1-201 (anatomy).

1788. La Jauue Lacepede, Hist. Nat. Quad. Ovip. Serpens, 1. p. 135, and

Testudo flava at end of same volume in Synopsis methodica, a table

in -which binomials are employed: Europe (other localities in

error ) .
1790. Testudo meleagris Shaw, Nat. Misc., 4, pi. cxliv: "America" (in

error).
1792. Testudo pulchella Schoepff, Naturg. Schildkroten, p. 134, pi. xxvi (ed.

2, 1801, p. 113, col. pi. xxvi) : No locality.

1 In conformity with our practice we have included all known synonyms : in this
instance howeve'r, our orbicularis material is too inadequate for us to express an
opinion as to whether any of them are possibly recognizable geographical races.
We have omitted '-Emys antiquorum Valenciennes," listed as a synonym by Gray
(1855 :41) as no such name was found in the reference he gives.

LOVERIDGE AND WILLIAMS : AFRICAN" CRYPTODIRA 203

1802. Testudo rotunda Sonnini and Latreille, Hist. Nat. Rept., 1., p. 107;
2. p. 282: No locality (Sometimes attributed to LacepMe (1788)
who, however, iises only the French equivalent in conjunction with
orbicularis) .

1833. Cistuda hellenica Bibron and Bory de Saint Vincent, Reptiles et
Poissons, in Bory de Saint Vincent Exped. Sci. Moree, 3: Zool. 1,
p. 61: Nisi Plain, Pamisus Basin, Messina, Greece.

1833. Emys hellenica Valenciennes (for plate only) in Bory, op. cit., Atlas,
pi. viii, figs. 2-2a.

1833. Emys iberica Valenciennes (for plate only) in Bory, p. 61, op. oit.,
Atlas, pi. Lx, fig. 1.

1836. Emys Eofinanni Fitzingcr, Ann. Wiener Mus., 1. p. 123: n.n. to com-
bine hellenica with orbicularis.

1850. Cistudo Europaea Guichenot, 2.

1860. Tristram, 405 ("Sahara" applied erroneously).

1899. Doumergue, 252.

1904. Chaignon, 3.

1925a. Seurat, 150.

1851. Emys lutaria Bianconi, 71.
1862a. Strauch (part), 101.
1865. Strauch, 49.

1897. Bateman, 50, fig. 34 (habits in captivity).

1854. Emys europaea Eiehwald, 416.

1877. Bruhl, pi. xxxiii, figs. 8, 14; pi. xxxiv, figs. 4, 9.

1880. Bruhl, pi. Ixix, figs. 3-4.

1915. Rawitz, 671, pi. xlix, figs. 76-78.

1855. Lutremys europaea Gray, 1, 40 (Lutremys was proposed in 1844).
1887. Lortet, 15, pi. vi (did not encounter any when in Africa).
1862b. Cistudo lutaria Strauch, 17.

1867. Lallement (not seen).

1894. Oliver, 101.
1889a. Emys orbicularis Boulenger, 112.

1891c. Boulenger, 96, 105.

1896. Doumergue, 477.

1897. Siebenrock, 247, pi. iii, fig. 14.
1901. Gadow, 351, fig. 79,

1907. Johnson, 13, 69, photo.

1908. Kerville, 96.
1909a. Siebenrock, 486.

1912b. Werner, 412, fig. (not used).

1913. Pellegrin (1912), 420.

1920. Mourgue, 233.

204 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1925b. Flower, 922.

1927a. Pellegriii, 261.

1929. DoUfus, 112.

1930. Seurat, 179.
1937a. Flower, 4.

1938. Gorham and Ivy, 181.

1951. Aellen, 169, 195.

1954. Noel-Hume, 94, 112.

1903. Emys orbicularis hellenica Kovatscheff, Verb. Zool.-Bot. Ges. Wien,

53, 171 (revives name as subsp.).
1916. Emys orbicularis aralensis Nikolsky, Faune Eussie, Eept., 1, p. 24:
Lake Aral (presumably Aral Sor, just north of the Caspian Sea,
Astrakhan, U.S.S.E.).
1934a. Emys orbicularis orbicularis Mertens and Miiller in Eust, 7.

Another citation of Emys hot aria will be found under Clcmmys
caspica leprosa. An extensive bibliography of orhicularis is fur-
nished by Boulenger (1889a), and another by Siebenrock (1909).
After careful checking we have omitted most of them from the
above list as they deal exclusively wdth European or Asiatic
material ; a selected few have been included on account of their
taxonomic importance or because of the observations that they
contain.

Names. European Terrapin (preferred) ; European Pond Tor-
toise (Flower: 1929); Speckled Terrapin (Shaw: 1790); Mud
Tortoise (Bateman: 1897).

Illustrations. This terrapin has often been well figured, though
usually from European specimens as in the colored plate of Lortet
(1887:pl.vi).

Descriptio7i. Beak distinctly notched ; edge of jaws smooth ;
mandibular width at symphysis less than (young) or subequal
to (adults) the horizontal diameter of the orbit; skin on upper
surface of head undivided ; forelimb anteriorly covered with
smooth, flattened shields of very diverse size, of which none is
conspicuously enlarged; digits webbed to the 5 claws; hind foot
strongly webbed, claws 4 ; tail long.^

Carapace moderately depressed, anterior and posterior margins
not expanded, not reverted, not serrated, not notched in nuchal
region; nuchal moderate to small, longer than broad; dorsal
shields concentrically striated except in aged specimens; verte-

1 Alleged • a?:e anrl sexual distinctions displayed by tail length that are men-
tioned by Boulenger (lS89a :113) are not disrermble even in our limited material.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 205

bral keel distinct in young but scarcely distinguishable in adults;
vertebrals 5, broader than long, broader than the costals (young)
or narrower than the costals (adults) ; costals 4, with indistinct,
interrupted keels (young) or without trace of keels (adults) ;
marginals (exclusive of nuchal and supracaudals) 11 ; supra-
caudal divided.

Front lobe of plastron anteriorly truncate, not produced, not
notched, possibly slightly concave in S $ , flat or convex in $ 9 ,
hinged at humeropectoral suture ; gulars paired, rarely two
pairs ;'^ pectorals wide; axillary 1,"^ moderate; inguinals variable,
often indistinct, not in contact with femoral ; hind lobe very
broadly or not notched ; anals truncate or rounded, not pointed
(cf. Fig. 4, for contrast with Clemmys) .

Fig. 6. Skull of Emys orbicularis (A.M.N.H. 73604). Half grown. Condy-
lobasal length 28 mm. Certain features of this specimen are due to its
comparative youth. (S. McDowell del.)

Plastral formula: An > (g, p, abd subequal) > f > h.

Color. Carapace of young, dark brown or black, uniform, or
with more or less numerous yellowish dots or radiating lines.
Adults similar.

Plastron of young, dark brown or black, the outer side of each
plastral shield and underside of each marginal with a large

1 Two pairs in an Algerian specimen (U.S.N'.M. 10986).

2 Occasionally 2, as on right side of a Tiflis terrapin (M.C.Z. 5309).

206

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yellow spot. Plastron of adult nearly entirely blackish brown,
brown and yellow, or yellow, each shield more or less narrowly
edged with black. Adults similar.

Head above, dark brown or black, uniform, or spotted or ver-
miculated with yellow; sides of head and throat spotted or ver-

Fig. 7.

Emys orbicularis (M.C.Z. 31976) Internal view of plastron, x 2/3.

(P. Washer del.)

miculated with yellow, the latter sometimes predominating on
the throat; limbs and tail dark brown or blackish, more or less
sparsely spotted with yellow, the tail streaked with yellow,

"We are unable to confirm Boulenger's statement (1889a) that
the color of the spots has a sexual significance, i.e. yellow in 9 ,
pale brown in $ . Gadow (1901) states that both the color and

LOVERIDGE AND WILLIAMS : AFRICAJST CRTPTODIRA 207

shape of the carapace alter with age.

Size. Carapace length of largest S (M.C.Z. 5187) from Algiers,
128 mm., breadth 94 mm., height 48 mm.; carapace length of
largest $ (M.C.Z. 5189) from Algiers, 138 mm., breadth 96 mm.,
height 62 mm. Both far surpassed by an unsexed European
record of 190 mm. (Boulenger: 1889a). A juvenile (M.C.Z.
3482) from southern Europe has a carapace length of 30 mm.,
breadth 24 mm., height 14 mm.

Sexual dimorphism. Both tail length and spotting were be-
lieved by Boulenger (1889a) to be indications of sex, but we
have been unable to confirm his findings with the limited material
at our disposal.

Breeding. On warm spring nights during the pairing season
these terrapins emit short piping calls until they find a mate,
after which the couple swim about together.

When gravid, the 9 , having selected a suitable spot free of
vegetation, where the soil is firm, prepares the site by ejaculations
of fluid from the bladder and anal water-sacs. Then, holding her
body rigid while stiffening the tail, she uses the latter as a borer,
pushing it into the ground. Following this preparatory loosening
of the soil, it is scooped out and heaped around the periphery of
a hole that may be as much as five inches in depth, i.e. as far
down as her hind feet can reach. The feet are also employed to
separate and spread the eggs which are deposited in a single layer
on the bottom of the pit.

After about 10 eggs have been laid, the loose soil is returned
to the hole and tamped down by allowing the plastron to fall
upon it as, time and again, the terrapin raises herself to her full
height and then drops on the site. After some additional stamp-
ing over the area, the terrapin roughens the surface with her
claws before leaving the place for good.

The incubation period varies in relation to locality and tem-
perature, embryos sometimes hibernating within the egg. In a
garden at Kiev, U.S.S.R., some eggs did not hatch until 11
months after they were laid. For some obscure reason, hatch-
lings of orhicularis are more difficult to rear than are those of
other terrapin. (The foregoing remarks probably refer to
European specimens and are taken from Gadow: 1901.)

Growth. One hatchling, reared in captivity, which wintered
beneath moss in an unheated room in England, took 4 years to

208 BULLETIN : MUSEUM OF COMPAEATIVE ZOOLOGY

attain a carapace length of 50 mm. A 25 mm. terrapin grew to
134 mm. in carapace length during 11 years, at the end of which
time it weighed 491 grams (about 1 lb.). Another, over a period
of 8 years, increased in carapace length from 110 to 132 mm.,
and from 83 to 106 mm. in breadth. This particular terrapin,
however, did not hibernate as it was kept in a greenhouse
(Gadow: 1901).

Longevity. Perhaps the best authenticated record is 27
years, 11 months, 17 days, in the Jardin des Plantes (Vail-
lant:1892:223i) cited by Flower (1925b) who, however, (1937a)
refers also to a $ , already adult in 1868, which bred regularly
for 60 years in a French garden until her death on February
27, 1928. This, and a record of 120 years, are taken from
Rollinat (1934:110-). Probably all these instances of longevity
are based on French, rather than African, terrapins.

Diet. In captivity young European Terrapins will readily take
flies, tiny worms or tadpoles ; larger individuals eat insects, frogs,
fish and even raw meat. Though normally feeding in water,
tame terrapins will come out on land to be fed if sufficiently
hungry. Usually, however, the prey must be seen in motion
before a terrapin will attack it.

Frogs are stalked as they sit on a floating leaf. Rising slowly
from below, the terrapin thrusts its nostrils and eyes above the
surface close to the frog and waits motionless. After a while it
may sink to rise again with its snout actually touching the
unsuspecting frog's toes which, after smelling at them, it seizes
with a sudden sidewise biting motion. While maintaining its
hold the terrapin employs its sharp foreclaws to tear the living
prey to pieces. This occupies considerable time for only the
intestines and scraped-off flesh are devoured.

Fish too are stalked, the terrapin moving slowly along the
bottom as it cautiously approaches its prey. Then, with a few
gentle movements of its fullj^-extended webbed feet, the reptile
rises almost imperceptibly and, gaping widely, grabs at the
fish's belly. The bones are picked clean as the skeleton sinks
to the bottom, but the air-bladder floats away on the surface
to serve as an indication of the presence of a terrapin in the
pond (Gadow :1901).

1 Vaillant, L., 1892, Arch. Mus. Paris (3). 4, pp. 221-253.

2 Rollinat, R., 1934, La Vie des Reptiles de la France Central (Paris).

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 209

Defense. The European Terrapin lacks a defensive odor like
that which is so characteristic of Clemmys (Gadow:1901).

Hibernation. In Europe this terrapin buries itself in mud and
does not reappear until spring is well advanced (Gadow:1901).

Migration. At times, extended migrations take place, either
under the stimulus of a food shortage or because the occupied
pool is in process of drying up (Gadow:1901).

Habits. European Terrapins should not be kept in an aquar-
ium unless there is growing vegetation and facilities to land,
stones to bask on and bark or moss under which they can retire
when so inclined (Bateman: 1897; Gadow:1901).

Habitat. The statement as to the abundance of this terrapin
in Algeria, made by Guichenot (1850), is due to confusion with
Clemmys, according to Boulenger (1891c). The same author
points out that its reported occurrence in the "Sahara" by
Tristram (1860) is due to a misapplication of this term to the
high plateau of southern Algeria. The frequency with which
Etnys occurs in Tunisian streams in the absence of Clemmys, or
vice versa, is remarked upon by Chaignon (1904) ; however, both
apparently coexist in a dozen localities.

Localities. French Morocco: Oued Ifrane; Oued Nkhol. Al-
geria: * Algiers; Bone; La Calle; Lake Fetzara; Oued Harrach;
Oued Sebaou (as AVed Sebaon) ; "Sahara" (in error). Tunisia:
A'in Draham; Cap Bon streams; Krombalea to Soliman; Oued
Belli ; Oued Bezirk ; Oued el Amor ; Oued el Kebir ; Oued Melah
(as Mala); Oued Sidi Saad, 3 km. south of Cebala; Tabarca
(as Taborca).

Range. Northwest Africa (Morocco to Tunisia) ; central and
south Europe; southwestern Asia.

Formerly the range v.-as much more extensive. Post-glacial
remains have been found in Sweden, Denmark, the Netherlands
and in the peat of England (Norfolk and Cambridgeshire).
According to Bateman (1897) specimens that have escaped
from captivity survive English winters, having been recaptured
in a healthy condition years later.

Subfamily TESTUDININAE

1909. Testudininae Siebenrock, Zool. Jahrb. Syst., Suppl., 10. p. 508.

Definition. Testudinid tortoises of strictly terrestrial habit.
Skin of head divided into larger or smaller shields; scales on

210

BULLETIN : MUSEUM OF COMPAKATIVE ZOOLOGY

forelimb more or less enlarged, often with dermal ossifications ;
hinder side of thigh often with large spurlike tubercles : scent
glands absent ; bnrsae anales absent.

Skull with prefrontals whose descending processes are more or
less widely separated inf eriorly ; frontal entering or excluded
from orbit; postorbital tending to be reduced, rarely absent;

Fig. 8. A dendrogram of testudinine relationships. Names in capitals
are those of full genera. Names underlined are those of subgenera as recog-
nized in this revision. With each subgenus or genus is given its range and
the number of Recent species, if any. The symbol t indicates an extinct
form.

(P. Washer del.)

temporal region always emarginate posteriorly; temporal arcade
rarely interrupted by absence of the postorbital ; quadratojugal
typically well developed, never absent, never in contact with

LOVERIDGE AND WILLIAMS: AFRICAN CBYPTODTRA 211

maxilla; jugal entering orbit; quadrate usually enclosing stapes;
surangular not covered by dentary, often extensively exposed
laterally.

Coracoids with median borders markedly widened ; tuberosities
of humerus tending to be approximated; trochanteric fossa of
femur restricted by union of trochanters ; epipodials very short ;
no digit with more than 2 phalanges, except in an extinct genus.

Carapace usually arched, high ; rib heads often vestigial ; neu-
ral bones primitively hexagonal, tending to become alternately
quadrangular and octagonal, sometimes vestigial; of the infra-
marginal series only the axillaries and inguinals normally pres-
ent.

Range. World-wide except for Australian region.

Retnarks. The separation of the land tortoises into generic
groups is not easy. Like the emydines they are a closely knit
assemblage in which there has been much parallel evolution.
Even more than in the emydines, it is difficult to distinguish
between those characters that are merely convergent and those
that may indicate natural divisions.

In a revision of African forms, however, the problem is a
pressing and immediate one since nowhere else in the world do
the tortoises achieve the diversity or the numbers of species that
they have in Africa.

We have made a beginning by assembling the recognizable
forms into species groups. Estimation of relationships at this
level rests upon relatively secure foundations. With almost equal
confidence we may proceed to the level of groups that may be
termed subgenera, but above this uncertainties increase. In
Figure 8 we have attempted a diagram of relationships between
species-assemblages that are at least subgenera, and to which
the available Latin names have been attached. We have included
the evidence of fossil forms where these have been sufficiently
well analyzed to be usable. Fortunateh^ in the testudinines, in
contrast to the emydines, the fossil record approaches adequacy,
at least in some periods and places.

In preparing Figure 8 we have considered the following as
the characters which the primitive testudinine must have had.

212 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

A B __ C

FIGURE 9

LOVERIDGE AND WILLIAMS: AFRICAN CBYPTODIEA 213

Osteological characters

1 . Interval betweeii ventral processes of the prefrontals only
moderately widened.

2. Maxilla with triturating surface ridged, but premaxilla
unridged.

3. Anterior palatine foramina small, concealed.

4. Temporal arcade strong.

5. Prootic well exposed dorsally.

6. Quadrate not enclosing stapes.

7. Surangular subequal in height to prearticular.

8. Fourth cervical centrum biconvex.

9. Anterior neurals hexagonal.

10. Suprapygal one, anterior to the vertebral-supracaudal
sulcus.

11. Entoplastron anterior to humeropectoral sulcus.

12. Digital phalangeal formula 2, 3, 3, 3, 3.

External characters

1. At least prefrontal and frontal head scutes present.

2. Scales on f orelimb numerous, not greatly enlarged.

3. Large femoral tubercles present.

4. Claws on forelimb 5, on hind limb 4.

5. Tail claw absent.

6. Carapace moderately convex.

7. Neither carapace nor plastron hinged.

8. Nuchal present.

9. Vertebrals not conical.

10. Vertebral keel very weak, lateral keels absent.

11. Supracaudal divided.

Fig. 9. Forelimb scalatiou in certain testudinids. A, Clemmys caspica
leprosa (M.C.Z. 1894) (x 2) ; B, Geochelone pardalis labcocU (M.C.Z.
50304) (x 1) ; C, Emys orbicularis (M.C.Z. 5189) (x IVa); D, Testudo her-
manai (M.C.Z. 3063) (x IVa) ; E, Testudo Meinmanni (M.C.Z. 5081) (x 2) ;
F, Testudo graeca graeca (M.C.Z. 1497) (x IVa) ; G, Malacochersus tornieri
(X.R. Stockholm), Njoro, (x 1); E, Chersina angulata (M.C.Z. 3998)
(x 1); I, Malacochersus tornieri (M.C.Z. 30003), Mangasini, (x 1); J,
Psammohates t. trimeni (M.C.Z. 42227) (x 1) ; K, Psammobates t. ten-
torius (M.C.Z. 21332) (x 1).

(P. Washer del.)

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BULLETIN : JiUSEUM OF COMPARATIVE ZOOLOGY

12. Submarginal scute absent.

13. Gular area but slifi'htly j^rodnced or thickened.

14. Gulars paired.

15. Anal notch moderate.

Most of these characters are primitive emydine and are in-
ferred to be also primitive testndinine on the well-grounded
theory that the Emydinae are ancestral to the Testudininae. In
several instances where there is a good fossil record for a tortoise
lineage, it is possible to observe a shift in these characters from

A

Fig. 10. Interprefrontal space in African testudinids I. A, Clemmys
caspica leprosa (M.C.Z. 2210) ; B, Emys orbicularis (A.M.N.H. 74604) ;
C, Geochclone pardalis habcocM (A.M.N.H. 7203); D, Testudo graeca
gra.eca (M.C.Z. 4485) ; E, Testudo Ideinmanni (Yale Mus. 662) ; F, Mala-
cochersiis iornieri (A.M.N.H. 45081); G, Chersina- angulata (A.M.N.H.

50725).

(P. Washer del.)

IvOVERID(iE A\n WII,T.TA:">!P : APRirAN CRYPTOniRA

215

the earlier to the later meiubers, e.g. from the emydiiie hexagonal
neural pattern to the advanced testudinine octagonal cum
(juadrilateral condition.

Fig. 11. Iiiteipiefioutal space in African testudinids II. H, Psammohatcs
t. ientorius (M.C.Z. 3465) ; I, Psammobates t. verroxii (M.C.Z. 21330) ; J.
Ilomopus Jjoulengcri (A.M.N.H. 7107); K, Homopus areolatus (A.M.N.H.
17792); /., Kinixys erosa (A.M.N.H. (39727); M, Kinixxjs Iwmeana (A.M.
X.H. 50725); N, Kinixys h. hellmva (A.M.N.H. 10029).

(P. Washer del.)

All the osteological characters listed are clearly primitive
emydine, with the possible exception of the small, concealed,
anterior palatine foramina. The latter feature is considered

216 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

primitive for testudinines because of its more frequent occur-
rence in forms regarded as primitive on other grounds. Similarly
we treat as primitive for testudinines certain kinds of scutella-
tion unknown in emydines, e.g. complex head scutellation with
supranasals, prefrontals and frontals.

No living tortoise exhibits the whole complex of characters^
here considered primitive. Rather, the members of the Testu-
dininae display various combinations or tnosaics of primitive
and advanced characters, characteristic for each subgroup or
lineage .2

Superficially considered, the tortoises give a great overall im-
pression of homogeneity — far greater than that provided by
the emydines. But when all the evidence is considered and par-
ticularly that of the rather good fossil record, it is apparent
that this homogeneity is primarily one of trends and only sec-
ondarily of realized conditions. The records seem to indicate
there has been no single sequence, but rather a number of starts
from a single point of departure.

The development of advanced characters in the separate lines
has been quite independent; the characters have been neither
consistently synchronous in appearance nor consistently consecu-
tive nor in any evident way correlated. Each lineage appears to
be characterized by the timing of initiation of the several ad-
vanced characters, and some lineages indeed are still primitive
in certain aspects today, some in others. A discussion of the
paleontological evidence supporting these conclusions is not
pertinent here and will be presented separately by one of us
(EEW). It is however with these considerations in mind that
we have framed our diagrams of relationship and based our
decisions as to generic rank.

1 In our generic definitions we have used many, but not all 27, of the listed
characters, or their advanced alternatives. Certain of them have proved too
variable within groups, or even within species, to be of taxonomic value. We
have also preferred osteological to external characters.

2 Anyone desiring to check these statements is referred to our numerous figures.
Figures 10 and 11 show the condition of the interval between the ventral proc-
esses of the prefrontals in most of the African Testudinidae. Figures 19 and 22
furnish examples of head scutellation. Figures 9, 39 and 44 illustrate types of
limb scutellation. Figure 12 shows the suprapygal area. Under each genus will
be found additional illustrations of, at least, major skull and shell characters.
With regard to the latter a warning is necessary. There has been neither the
opportunity nor the desire to figure "typical" specimens. Both age and in-
dividual variations have been drawn when present in the specimen depicted, and
attention has been directed only to the more striking peculiarities or abnor-
malities'.

LOVEBIDGE AND WILLIAMS: AFRICAN CRYPTODIRA

217

Fig. 12. Pygal patterns in testudinids (diagrammatic). A, Clemmys
caspica leprosa (Pygal pattern differs within the genus Clemmys as cur-
rently recognized) ; B, Geochelone pardalis babcocki; C, Emys orbicularis ;
D, Testudo graeca graeca ; E, Malacochersus tornieri; F, Psammobates ten-
torius tentorius ; G, Chersina angulata ; H. Uomopiis signatus ; I, Kinixys
erosa; J, Aeinixys planicauOa ; K, Pyxis arachnoides.

(P. Washer del.)

218 BULI^ETIN : MUSEUM OF COMPARATIVE ZOOLOGY

In summary, we have come to the following eonclnsions :

The assemblage that has the Eocene subgenus ^Hadrianus as
its base is a natural grouping and one that appears to us to
extend back to the very base of the tortoise line. We have
adopted the generic name Geochelone for the whole of this
rather homogeneous group.

•\Stylemys^ and Gopherus share a "unique specialization — the
median premaxillary ridge — and thus clearly belong together
but their differences (i.e. in phalangeal formula) point to a very
ancient separation at practically the emydine level (see below,
p. 255 ff ) , and we therefore regard them as two quite distinct
genera.

Testudo in the strict sense (with 2\ graeca Linnaeus as the
type) and Pseudotestudo new subgenus (T. hleinmanni Lortet
type) seem aijpropriately bracketed together by certain very
singular specializations (prootic completely concealed l)y the
parietal,- and plastron posteriorly hinged in one or both sexes),
hut Pseudotestudo, which has not been separated from Testudo
previously, shows some amazing ditferences in skull structure
(see p. 259 and Figs. 20, 24). There are, fortunately, transitional
forms in this series and the unity of the genus appears satisfac-
torily established. The record demonstrates that this lineage is
as ancient as that of Geochelone {yTestttdo comptoni of the
Eocene of England, see pp. 353 ff below).

For the groups so far mentioned there has been, happily, a
fossil record. For the diverse forms of the Ethiopian region there
is no such aid. We have proceeded here on the strict ground of
degree of morphological difference, influenced, however, by the
extra-African evidence that only modest morphological difference
may mean very ancient divergence.

We shall argue below that Acinixys, Pyxis;^ Psammohates,
Chersina, Bomopus and Kinixys are related. The individual

1 As a matter of conventiou throughout this paprr ovor.v pxtiurt l'imhis or
species is preceded by the symbol t.

2 This character was first noticed hy Baur (1802), who was also the first to
point out the '•Testudo leithii" (i.e. k'leinmnnni) characters that have led us to
sei)arate it as a subgenus.

3 Pi/ J" is Bell, 182.5, Is used rather tlian Belleiiina Williams. in.'iO, I't.'i'i. in as
nuuh'as the supposedly preoccupying name Pyxis Chemnitz 1784 is not binomial,
arul the next use of the name (Humphreys in the "Musenm Calonniauiim" 1T!«T)
has now been outlawed by ruling of the International Comuiis.sion on Nomen-
clature (Decision 31).

IjOVERIDOE and WIIJJA.MS: AFRICAN CRVPTOniRA

2 in

specializations of each, however, are so marked that we regard
each as constituting a separate genus. Malacochersus we have
found difficult to place, its relationships quite obscure ; we sug-
gest that it has branched off at or very near the base of the
testudinine series, and we have therefore retained it as a
genus also.

Shown on the relationshi]) diagram for the sake of complete-
ness, are two fossil forms of uncertain position. Though im-
perfectly known, on present evidence they appear to be quite

Fig. lu. Tliiid and loiulli eostals of Klnixya for comparison with those
of Ilomopun. A, Kinixya h. helUana (M.C.Z. 40008); B, Homopus boulen-
iirri (M.C.Z. 42231).

(P. Washer del.)

isolated. A few other fossil genera — jAchilemys Hay, jCheiro-
gaster Bergounioux, ^Sinohadrianus Ping — listed in Williams
(1952), are impossible to place at this time. Further comment
on the African genera will be found under each genus.

220 BULLETIN : MUSEUM OF COMPAKATIVE ZOOLOGY

Key to the Genera of Testudininae in Africa^

1. Carapace hinged posteriorly in adults, usually between 7th and 8th mar-

ginals; outer side of fourth costal markedly wider than outer side of
third (in all ag.es) (cf. Fig. 13). Eange: Africa from 17° north to

Bechuanaland and Natal Kinixys Bell (p. 374)

Carapace without hinge ; outer side of fourth costal subeqiial to, or
smaller than, outer side of third . . . ; 2.

2. Gular single, strongly projecting. Eange: South Africa

Chersina Gray (p. 342)

Gulars paired 3.

3. Carapace very flat, flexible, fenestrated; neural bones reduced; plastron

with a great fontanelle at all ages. Range: arid areas of Kenya and

Tanganyika Malacochersus Lindholm (p. 283)

Carapace more or less convex, rigid, solid; neural bones unreduced;
plastron with a fontanelle only in hatchlings 4.

4. Maxilla uuridged; anterior neurals hexagonal or quadrilateral 5.

Maxilla ridged (except in Tcstudo Meinmannl of Egypt) ; anterior
neurals octagonal and quadrilateral 6.

1 Some divergence of opinion occurs with regard to the advisability of breaking
up Testudo into several genera — first attempted by Fitzinger (1835) and more
recently bv Hewitt (l'J'^1 : 1933b). Uninfluenced by either of these authors, my
colleague, "Dr. E. B. Williams, is fully convinced by his own studies that certain
structural differences indicate so early a divergence among the several groups as
to justify their recognition as full genera. lie considers their numerous similari-
ties, when not due to common ancestry, are attributable to parallel evolution or
due to convergence. Assumptions as to which characters reliect close affinity, and
which are to be discarded as subsequent convergences, seem somewhat speculative
to me in view of the fact that the fossil history of five of the seven suggested
genera is completely unknown.

More impressed than my colleague by the many similarities (which may be
seen by comparing the several generic descriptions), I take a more pragmatic
view of the purposes of nomenclature. A multiplicity of genera that are based
on single characters, or aggregations of characters that in themselves may be
subject to variation, and sometimes difBcult to observe or evaluate, tends to
nomenclatorial confusion. For an instructive example of the extent to which
genera-making can burden posterity, one has only to turn to the 28 synonyms of
Trionyx (cf. p. 420).

Fragmentation of Testudo offers the advantage of separating into groups those
species whose characters are believed to indicate close affinities. On the other
hand, this result can be achieved by treating such groups as subgenera, an
arrangement that has the additional advantage of emphasizing their over-all
relationship, though to this my colleague's views naturally prevent him from
subscribing (cf. Fig. S). To me, as a working herpetologist, the desirability of
maintaining as stable a nomenclature as possible outweighs the advantages of
emphasizing a new theory of phylogeny (that may not find general acceptance or
have to be set aside when more is known of the fossil history involved) by
raising the several groups to full generic status. In the conservative view,
Gcochelouc, Punmmobaten and Chersina would be retained as subgenera of
Testudo.

Possibly this attitude is a mistaken one, so for the purposes of this revision
I defer to a current trend in herpetology. This is done in the confident belief
that- — as increasing knowledge reveals the disadvantages of a multiplicity of
genera that are difficult to define — in due course there will be a return to the
larger units as is even now occurring in South African ornithology following an
extended period of nomenclatorial chaos. A.L.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 221

5. Carapace somewhat depressod ; vertebral shields never conical ; each gular

usually broader than long (cf. Fig. 14) ; anterior palatine foramen
large, conspicuous; maxillary bone not entering roof of palate. Range:

South Africa Tlomo'pus Dumeril and Bibron (p. 352)

Carapace rather convex; vertebral shields sometimes conical; each gular
usually as long or longer than broad; anterior palatine foramina
small, concealed; maxillary bone entering roof of palate. Eange:
South Africa rsammohates Fitzinger (p. 294)

6. Supranasal scales present; nuchal scale present; femoral tubercles absent

or only 1 ; suprapygal single or divided transversely ; posterior lobe
suprapygals 2, a larger anterior one embracing a smaller posterior one
of plastron (in life) more or less clearly movable in one or both sexes;
prootic usually concealed dorsally by the parietals. Range: North

Africa Testudo Linnaeus (p. 254)

Supranasal scales absent; nuchal scale absent; femoral tubercles 2 or 3;
suprapygals 2, a larger anterior one embracing a smaller posterior one
(cf. Fig. 12B) ; posterior lobe of plastron never movable; prootic
usually well exposed dorsally. Range: Tropical and South Africa. . . .

Geoohelone Fitzinger (p. 221)

Genus GrEOCHELONE Fitzinger

1835. Geochelone Fitzinger, Ann. Wiener Mus., 1: pp. Ill, 112, 122. Type:

Testudo stellata Schweigger := T. elegans SchoepfP (designation by

Fitzinger: 1843).
1872c. Centrochelys Gray, Appendix Cat. Shield Eept. Brit. Mus., p. 5.

Type: Testudo sulcata MMler (by monotypy).
1873b. Stigmochelys Gray, Hand-List Shield Rept. Brit. Mus., p. 5. Type:

Testudo pardalis Bell (by monotypy).
1933b. Megachersine Hewitt, Ann. Natal Mus., 7. p. 257. Genotype: Testudo

pardalis Bell (by original designation).
Definition. Skull with triturating surface of maxilla strongly
ridged; median premaxillary ridge absent; maxillary not enter-
ing roof of palate; anterior palatine foramina small, concealed
in ventral view; prootic typically well exposed dorsally and
anteriorly; quadrate usually enclosing stapes; surangular sub-
equal in height to prearticular ; neck with second, third or
fourth centrum biconvex.

Carapace never hinged; typically the anterior neurals alter-
nately octagonal and quadrilateral; outer side of third costal
scute about as long as, or longer than, that of the fourth; no
submarginal scute ; suprapygals 2, the anterior larger, bif urcat-

222

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

ing posteriorly to embrace the smaller posterior element, which
(in post-Eoeene forms) is crossed near its middle by the sulcus
between the fifth vertebral and the supracaudal.

Plastron not hinged ; gular region more or less thickened and
produced ; gulars single or paired, longer than broad.

Range. Galapagos Islands; South America; Africa; Mada-
gascar and other islands of the Indian Ocean (including Cey-
lon) ; southeastern Asia (including India).

A

B

Fig. 14. Typical shape of gular scutes in Psammohates for comparison
with those of Eomopus. A, Psammohates t. verroxii (IM.C.Z. 42222) ; B,
Homopus areolatns (M.C.Z. 17524).

(P. Washer del.)

Fossil record. Well represented in the Tertiary and Pleistocene
of North America and the Tertiary of Europe ; known from the
Oligocene and Pliocene of Asia ; from the Oligocene and Miocene
of Africa ; recorded in South America only since the Miocene,
and in the West Indies from the Pleistocene.

Remarks. The genus Geochelone tends to large size. In cara-
pace length some fossil and Recent species may exceed a meter
in length and none attains to less than 250 mm.

Characteristic of this genus is the fact that the shell takes on
certain specializations early in its phyletic history. Earliest of
these are a thickened, produced gular region and the peculiar
pygal pattern with the first suprapj'gal embracing a smaller
second one, both already present in jHadrianus of the Eocene.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA

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224 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

The alternate octagonal-quadrilateral neural pattern arises a
little later, but even in fHadrianus one of the neurals may some-
times be octagonal and several of them are often hexagonal short-
sided behind. The latter conditions are intermediate, indicating
the establishment of a trend which, at least in North America,
culminated in the perfected pattern in the Oligocene. An octag-
onal-cum -quadrilateral pattern for the anterior neurals is now
very characteristic of the genus, though by no means invariable.

Other shell specializations occur irregularly within the genus.
The nuchal scute is lost in many species. The supracaudal is
usually single in post-Eocene subgenera with the exception of
Manouria. The pectoral scutes tend to become narrowed, mod-
erately so in the African species, to an extreme degree in the
typical members of the fossil American subgenus ^Resperotestu-
do. In Cylindraspis, and independently in Asterochelys yniphora,
the gulars have fused to form a single scute, while in the sub-
genus ■\Megalochelys the gular region becomes not merely pro-
duced but strongly bifurcate. Variation in height of shell occurs
also (contrast O. radiata and G. emys) as well as striking dif-
ferences within a species (cf. G. pardalis). The humeropectoral
sulcus crosses the entoplastron well anteriorly in the subgenus
Indotestudo, in which this feature was established early {jG.
kaiseni^ and t(r. nana^ of the Oligocene of Mongolia). The
same character occurs irregularly in a number of fossil species
elsewhere, e.g. jG. uintensis^ and ■\G. costaricensis^ of North
America.

In contrast to the shell, the skull in most of the subgenera of
Geochelone is remarkably primitive (for a testudinine) and sur-
prisingly uniform. However, there is a tendency, several times
repeated, for a downgrowth of the opisthotic to more or less con-
ceal the postotic fenestra in posterior view. Apart from this the
only striking modification occurs in the Seychelles-Aldabra
tortoises and ^G. grandidieri of Madagascar. In these the ex-
ternal naris is extraordinarily extended dorsoventrally in eon-
junction with a distortion of the whole front of the skull, giving

1 C. W. Gilmore, 1931, BuU. Am. Mus. Nat. Hist.. 59. pp. 213-257.

2 C. W. Gilmore, 1915, Mem. Carnegie Mus., 7, pp. 101-161.

a A. Segura, 1944, Escuela de Farmacia, Guatemala, 6, Xos. 73-74, pp. 9-29 ;
Nos. 75-76, pp. 16-25 ; Nos. 77-78, pp. 13-24.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 225

the latter a bulbous appearance. Since this modification appears
to be quite as distinctive as any of the features, or combinations
of features, that distinguish subgenera of Geochelone, and since
the minor lineage involved seems to be quite a distinct one, we
here propose a new subgenus for gigantea, sumeiri and -fgrandi'
dieri, to be called :

AldABRACHELYS new subgenus
Type species : Testiido gigantea Schweigger

Definition. External naris higher than wide ; quadrate enclos-
ing stapes or not ; postotic fenestra not concealed in posterior
view; nuchal present or absent; typically the anterior neurals
alternately octagonal and quadrilateral; first dorsal vertebra
short and stout ; supracaudal divided or undivided ; gulars paired
but not divergent; entoplastron not crossed by the humero-
pectoral sulcus.

Range. Aldabra and Madagascar.

The continental African species 0/ Geochelone belong to the
nominate subgenus, defined as follows :

Subgenus GeOCHELONE Fitzinger

1835. Geochelone Fitzinger, Ann. "Wiener Mus., 1. pp. 108, 112, 122: as a
subgenus. Type species: Testudo stellata Schweigger = T. elegans
Schoepff.

Definition. External naris not higher than wide ; quadrate en-
closing stapes ; postotic fenestra not concealed in posterior view ;
nuchal absent; anterior neurals alternately octagonal and
quadrilateral ; first dorsal vertebra short and stout ; supracaudal
undivided; gulars paired but not divergent; entoplastron not
crossed by humeropectoral sulcus.

Distinguishing marks. The African members of this subgenus
are easily distinguished from other African tortoises. They are
the only ones with 2 or 3 subconical tubercles on the hinder side
of each thigh, and are by far the largest in adult size.

Range. Africa ; Ceylon ; India and Burma.

Fossil record. Not known fossil with any certainty except in
the Tertiary of Europe. Probably the fossil giant tortoises of

226

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

the islands of Malta, Minorca and Teneriffe belong to this sub-
genus.

Remarks. The living members of the subgenus divide into two
species groups, an Asian one consisting of G. elegans and G.
platynota, and an African one comprised of G. sulcata and

Ondurman O

Geochelone sulcata
G. pordalis babcocki

Dongola o

Jebel Telga O

V.

Bahr el Ghazal

Province Bahr el Jebel
Bor ^
Oondokoro V
--V. Mongalla #
"..,--•- (

Lado Enclave

EqooJor

Fig. 15. Locality records of Geochelone in East Africa north of the

Equator.

(P. Washer del.)

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 227

G. pardalis. The African species are much the larger and differ
also in not possessing the tail claw which is present in the
Asian forms. Both species groups appear to have been present
in Europe during the Tertiary.

The large tortoises of Africa present a relatively simple pic-
ture. The largest and most northerly species, G. sulcata, is con-
fined to a rather arid belt extending from Senegal to Eritrea,
consequently inhabiting mostly the Sudanese subprovince of
the Savannah Province of botanists, but also present in the
northern portion of the Northeastern subprovince. G. pardalis
babcocM, relatively uniform in all its characters, occurs south-
wards from the southern Sudan and Ethiopia throughout most
non-forested areas of the eastern half of the continent, west-
wards into Angola and the northern half of Southwest Africa.
Known from very few specimens is G. p. pardalis, which oc-
cupies, or did occupy (it is perhaps now extinct), a very limited
region in western South Africa.

Though available evidence shows G. sulcata to be allopatric to
G. pardalis they have always been separated specifically, a view
we see no reason to reverse. G. sulcata differs more from the two
pardalis forms than do the latter from each other. The frontal
scute characteristic of sulcata is lacking in all the pardalis we
have seen. While the pale color of sidcata is approached by
individual pardalis, especially those from arid regions that have
lost most of their black pigmentation, no pardalis is known to us
in which the black flecking is entirely absent. More significant
is the striking difference in convexity displayed by the much
larger sidcata when contrasted with its nearest neighbor, par-
dalis hahcocki.

The situation in regard to the two subspecies of G. pardalis
is sufficiently confused to require some extended discussion.
The subspecies hahcocki was separated from the typical race by
one of us (Loveridge :1935c) when he discovered that two speci-
mens from Southwest Africa in the collection of the Museum
of Comparative Zoology did not correspond to the form known
to him in East Africa. Photographs of these two Southwest
African tortoises are presented here for the first time and con-
trasted with East African material. The markings of the adults

228

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Table 4
Characters of the African species and subspecies of Geochelone

Character

sulcata

p. babcoelci

p. pardalis

frontal scute

usually
present
and large

absent

absent

convexity of
carapace

flattened
dorsally

very convex

flattened dorsally

shape of
gular region

enlarged
scales on
forelinib

color of
adult

color of
hatchling

maximum size
recorded

somewhat
produced
and notched

not or but slightly
produced, not or
but slightly
notched

slightly produced,
shallowly notched

juxtaposed

usually not
juxtaposed

not juxtaposed

uniform horn
color

yellow, richly
variegated with
black; or rarely
mostly yellow

yellow to greenish
variegated with
black

pale yellow,
the dorsal
shields edged
with brown;
plastron pure
yellowish white

light brown, the
dorsal shields
broadly edged
with black and
typically contain-
ing a red-bordered
areola within
which is a single
spot centrally
located and
usually united to
the posterior
areolae, or, in the
extreme north,
the central spot
flanked by two
lateral onesi,
plastron as Ln
p. pardalis.

dull yellow, the
dorsal shields
narrowly edged
with black and
containing a red-
bordered areola
within which is a
pair of spots
separated by a
light vertebral
line: plastron
dull yellow, su-
tures of the
shields more or
less margined
with black.

762 mm.

585 mm.

432 mm. (type)

1 As in a specimen from Mon^alla, Sudan, photographed for us by Dr. J. Eiseli
of the Vienna Museum; also an Ethiopian tortoise figured by Vaillant (1904a) as
calcarata. In some Kenya hatchlings the areolar spots may be altogether absent.

LOVERIDGE AND WILLIAMS : AF^RICAN CRYPTODIRA 229

are probably indistinguishable, but in juveniles the marked
flattening of the carapace is usually correlated with a color pat-
tern so distinctive that the nomen nudum higuttata^ was pro-
posed in a footnote by Cuvier (1829:10; Dumeril and Bibron:
1835).

Although twenty years have elapsed since the description of
p. hahcocki, it is still difficult to clearly define the ranges of the
two forms because of the paucity of p. pardalis material with
good locality data. A number of specimens at the British Mu-
seum (measured at the time hahcocki was described) appear to
belong to p. pardalis, but are labeled only "South Africa." In
addition to these the only specimens definitely known to us are
the following : Bell 's type of pardalis with the locality ' ' Cape of
Good Hope"; the adult shell (M.C.Z. 16713) labeled as from
Kolmanskop (Dr. W. Beetz coll.) ; a hatchling (M.C.Z. 22473)
from Aroab (W. S. Brooks coll.) ; the young specimen figured
by Duerden (1907b :fig. 7) and Hewitt (1937e:pl. i, fig. 2; pi. ii,
figs. 1-2). 2

We have also hesitantly assigned to the nominate race the
Archer tortoise from Keetmanshoop because that locality is so
near to Aroab, and, because of their proximity to the type locality
of pardalis, the Delalande specimens from Oliphants River, Cape
Province. How^ever, one Delalande tortoise in the Paris Museum,
labeled as the type of higuttata Cuvier, with locality "Cap,"
is an intergrade — being as convex as p. hahcocki but with the
juvenile pattern of p. pardalis.

For further remarks on this subject see the footnote on p. 250.

Key to the Species and Suhspecies of Geochelone in Africa

1. Carapace horn-color or brownish, uniform; frontal shield usually present,

large. Range: Eritrea and Ethiopia west to Senegal

sulcata (Miller) (p, 230)

Carapace yellow or olive more or less richly variegated with black; frontal

shield absent 2

2. Carapace distinctly convex, its height being included in its length (in a

straight line) from 1.61 to 2.11 times; vertebral shields of young with
a central black spot. Range: Anglo-Egyptian Sudan and Ethiopia

1 Erroneously cited as T. bipunctata by Gray (lS31c :12).

2 Thanks to Dr. Robert Mertens, we have a photograph of two additional speci-
mens from Kehoboth, south of Wiudhoels, Southwest Africa.

230

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

south to Natal, west through Cape Province to Southwest Africa (where
it meets with the typical form, but is dominant north of 27° S.) and

southern Angola p. habcocTci (Loveridge) (p. 235)

Carapace distinctly flattened dorsally, its height being included in its
length (in a straight line) from 2.02 to 2.62 times; vertebral shields
of young with a pair of lateral black spots separated hy a light longi-
tudinal line. Range: southern Southwest Africa (in former times pos-
sibly extending south to the Cape Peninsula, as the type locality was
said to be the Cape of Good Hope) p. pardalis (Bell) (p. 251)

Geochelone sulcata (Miller)^

1780.

Tcstudo sulcuta Miller, Icones animalium et plantarura (Various

Subjects of Natural History . . .), pi. xxvi: "India occidentali"

(in error).

1789.

Gmelin, 1045.

1792.

Schoepff, p. 156 (German ed.).

1796.

Shaw, in Miller (vide supra), 53, pi. xxvi.

1800.

Bechstein, in Lacepede, 346.

1801.

Schoepff, p. 135.

1802b.

Daudin, 313.

1802.

Shaw, 39,

1812.

Schweigger, 323, 452.

1814.

Schweigger, 53, 55 (reprint of 1812).

1831c.

Gray, 11.

1835.

Dumeril and Bibron (part), 74, pi. xlii, figs. 1-la.

1844.

Gray, 7.

1845b.

Riippell, 297.

1851.

Guiehenot, 189.

1855.

Gray, 9.

1860.

Dumeril, 162.

1862a.

Straueh, 78.

1865.

Strauch, 22.

1870a.

Steindachner, 326.

1872b.

Gray, 4, in Sowerby and Lear, pis. xvii-xviii.

1884a.

Eochebrmie, 13 (ignored).

1890.

Strauch, 52.

1905.

Barbier, 83, pi. iii.

1912.

Siebenrock, 214, fig. 2.

1924a.

Werner, 263, figs. — .

1928.

Flower, 654, figs. 1-2.

iTestudo Schopfl Fitzinger {nom. nud.), 1826, sometimes referred to the
synonymy of sulcata, was subsequently synonymized by Fitzin^er with Kinixys
eroga.

LOVERIDGE AND Wll.l.lAMS: AFRICAN CRYPTODIRA 231

1929. FloYsrer, 32.

1937ii. Flower, 9.

17S4. T<studo calcarala yt'liiu'iiU'i-, Haiuiiil. Verniisc-lit. Aljliandl. Zuol.
]). 317, pi.— (based on Miller's figure of sulcata).

1835. lliippell, 4.

1880c. Yaillant, 33, 88, pi. xxvi, fig. 3.

1889a. Boulenger, 159.

1891. Prato (not seen).

1893. Trinien, 79.

190-ta. Vaillant (part), 186 (fig. of juv. is p. laicocTci).

1906. Kanmierer, 48, fig. 7.

1906a. Siebenrock, 821.

1908c'. Kammerer, 769, figs. 6 7.

1908. Werner (1907), 1826.

1909a. Siebenrock, 521.

1911a,. Masi, 34.

1914a. Pellegrin, 122, pi. i, fig. 2.

1925b. Flower, 924.

1928b. Scortecci, 334.

1930a. Scort«cci, 215.

1950. Williams, 550.

1820. Cher sine cnlcarata Merrem, 32.

1831c. Testudo radiata var. senegalensis Gray, Syn. Eept., p. 4; Senegal.

1836. Geochelone sulcata Fitzinger, 122.
1855. Geochelone senegalensis Fitzinger, 251.
1869c. Peltastes ? sulcatus Gray, 173.

1870c. Gray, 656.

1870e. Gray, 12.

1873b. Centrochelys (Peltastes) sulcatus Gray, 13.

1873e. Gray, 725.

Further citations of ''sulcata" and " calcarata" will be
found under Geockelane p. babcocki.

Synonymy . The use of the name sulcata has been dealt with
at some length by Siebenrock (1912:214). As the copy of
Miller's work in the library of the Museum of Comparative
Zoology lacks the pertinent part of the text, and plate xxvi bears
no name, we appealed to H. W. Parker for help in verifying
Siebenrock 's conclusions. Dr. Parker replied that though plate
xxvi in the British Museum copy is also unnamed, there is a list
of plates which states:

232

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

''Tabula XXVI

Fig. A. Testudo sulcata. Lin. Syst. Nat. Classis III. Orclo I.
Genus CXIX. Spec. Habitat in India occidentali.

B. Pars inferior.

C. Caput magnitudine naturali."

Thus no doubt remains as to sulcata being the correct name to
apply to this species. Furthermore, Parker points out that plate
xxvi appeared in Part 5, which, according to Sherborn and
Iredale (1921, Ibis:302'309) was published apparently in 1779
or 1780.

Common names. Spurred Tortoise (preferred) ; Grooved Tor-
toise (Flower) ; alu gatta (Arabic in Kordofan, fide Fitzinger,
who translates it as "father of covers") ; abu gefne (Arabic in

Fig. 16. Skull of Geochelone sulcata (B.M. 1904.7.28.1).

(S. McDowell del.)

Dongola: Riippell) ; gafot (at Massaua: Riippell).

Illustrations. For most people Miller's fine plate (1780) and
its reproductions will be unavailable, but Sowerby and Lear's
(1872) (our Plate 1) are equally good. Werner (1924a) and
Flower (1928) have figured the young.

Description.^ Beak weakly hooked, edge of jaws strongly
dentate; prefrontal large, divided longitudinally; frontal usu-
ally large, rarely broken up; remaining upper head shields
small, irregular; forelimb anteriorly with large unequal, juxta-

1 Based chiefly on the head, limbs and shell of an adult $ (M.C.Z. 11975) from
Katsina Emirate ; together with notes on British Museum material as of 1953,
two without data in the A.M.N.H., and one without data in the C.N.H.M.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 233

posed or imbricate scutes, forming 3-6 longitudinal and 6-7
transverse series from elbow to outer claw; claws 5; posterior
side of thigh with 2-3 large conical tubercles; heel with large,
conical, spurlike, bony tubercles; claws 4; tail without terminal
clawlike tubercle.

Carapace flattened dorsally, sides descending abruptly, deeply
notched in nuchal region, anterior and posterior margins re-
verted and serrated, not more than twice as long as deep ; dorsal
shields concentrically grooved ; nuchal absent ; vertebrals 5, not
convex, the second, third and fifth much broader than long,
broader than the costals; costals 4, not forming an angle with
the marginals ; marginals 11 ; supracaudal undivided.

Front lobe of plastron somewhat produced and bifid; gulars
paired ; pectorals very narrow, their anterior border usually
straight, widening abruptly towards the axillary notch ; axillaries
2; outer moderate to small, inner minute; inguinals 2, outer
large to moderate, inner small, meeting femoral; hind lobe
deeply notched posteriorly, angular or crescentic.

Plastral formula : Abd>h>f >g>or=p>an.

Color. Carapace of juvenile (50 mm.^) pale yellow, the shields
with narrow brown sulci. Plastron yellowish white.

Carapace of adult brownish to horn color, uniform. Plastron
yellowish, uniform.

Size. Carapace length of ? (M.C.Z. 19975) from Katsina
Emirate, 670 mm., breadth 470 mm., height 270 mm. Apparently
exceeded by that of a specimen mentioned below.

The carapace of a young $ hatched at Giza Zoo on 5.x. 10,
was about 50 mm. long by 45 mm. broad at time of hatch-
ing. On 26.iii.28 this tortoise measured 762 mm. (30 inches) in
length and weighed 184 lbs. (Flower :1928). The 800 mm.
carapace from Eguei mentioned by Pellegrin (1914a) may have
been measured over the curve.

Longevity. Two tortoises hatched at Giza Zoo on 5.x. 10, lived
for 19 years, 9 months, 26 days, and 20 years, 7 months, 22 days
respectively, but were possibly surpassed by an adult $ measur-
ing "28 inches across the shell" which had been in captivity
"over 38 years" when presented to the London Zoo on 3.x. 31. It
died at Whipsnade on 19.xii.35 when reputedly over 42 years
old (Flower :1937a).

iCf. Flower (1928:654).

234

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Enemies. The flesh of a Spurred Tortoise is "never eaten"
by the natives, according to Rtippell (]835). However, his
informants may have been Moslems.

Habitat. In the Sudan, according to Werner (1908:1826)
sulcata extends no further south than Kordofan, being separated
by the swamps of the Bahr el Gebel region from the range of p.
bahcocki, still known only from points south of Bor on the White
Nile — i.e. about 5° N.

Fig. 17. Skull of Geochelone pdrdalis habcocli (A.M.N.H. 7703). Con
dylobasal length 81 mm.

(S. McDowell del.)

Localities. Sudan: Dongola ;'* Jebel Teiga, 30 miles west of
Jebel Medob (B.M. specimen seen by E.E.W.) ; Kordofan; Om-
durman (bought in market by Werner :1924a). Eritrea: Chena-
fena; between Cheren (Cliire) and Mareb (Marembe) River;'
Godolefassi; Massaua Coast. Ethiopia: Shoa (Chona) near
Awash (Heonacha) River;' Takkaze [River]. French Equa-
torial Africa: Eguei. Nigeria: *Katsina Emirate. Senegal:
Dagana; near Dakar; Saint Louis. French ^yest Africa: Maure-
tania; Sudan and Niger Territories {fide Villiers in litt.).

Range. Trans-Africa from Eritrea west to Senegal in thp
vicinity of 15° N.

1 Mons. Paul Chabanand infnrnis us that Lefebvre left no map of his itinprary
that would enabln us to fix his Incalirios with precision. Lefebvrp's orthography,
combined with some inisiirinting. renders the identification of his place names
with cnn-eutly recoirnized localities somewhat difficult. Thus "Chona", a mis-
print for Choua, = Choa or Shoa.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIHA

235

Erroneously reported from Antilles (Merrem) ; Buenos Aires
(A. Dunieril) ; Cape of Good Hope (A. Dunieril) ; Giza, Egypt
(Werner) ; Nyasaland (Johnston) ; Patagonia (Duraeril and
Bibron; Port Natal (A. Smith); South Africa (Boulenger).
The South American records are due to confusion with Geoche-
lone chilensis (see Siebenrock :1912).

Geochelone pardalis babcocki (Loveridge)

1831c.

Testudo pardalis Gray (parti), 12, pi.

iii, fig. 3

1835.

Dumeril and Bibron, 71.

,

1835.

Temminck and Schlegel, 73.

1844.

Gray, 7.

1851.

Dumeril and Dumeril, 4.

1854a.

Peters, 215.

1855.

Gray, 9.

1862a.

Straueh, 77.

1863b.

Gray, 382.

1865.

Straueh, 22.

I866I1.

Peters, 887.

1867a.

Bocage, 217.

1869a.

Peters, 11.

1870e.

Gray, 6.

1872c.

Gray, 4.

1873b.

Gray, 5.

1878.

Fisk, 790.

1878a.

Peters, 202.

1880e.

Vaillant, 8, 33, 88.

1882a.

Peters, 2.

1884a.

Roehebrune, 11 (but in error).

1886.

Vaillant, 137.

■

I88y.

Boettger, 281.

1889a.

Boiilenger, 160.

1890.

Straueh, 52.

1893a.

Boettger, 9.

1893b.

Boettger, 113, 122.

1893.

Trimen, 79.

1894c.

Giinther, 85.

1895a.

Boeage, 3.

1896a.

Bocage, 97.

1896a.

Boulenger, 546.

1 Many of the following references might equally well be considered as '(part)'
for they' mention Bell's type, though their material is obviously O. p. babcocki.

236

BULLETIN : MUSEUM OP

COM]

1896c.

Boulenger, 15.

1896.

Tornier, 3.

1897g.

Boulenger, 277.

1897.

Johnston, 361.

1897.

Meek, 184.

1897.

Tornier, 63.

1898.

Jeude, 3.

1898.

Johnston, 361.

1898.

Sclater, W. L., 96.

1898.

Tornier, 282, fig. 1.

1899.

Ansorge, 288.

1899a.

Mocquard, 218.

1899.

Siebenrock, 566, pi. iii, figs.

34-36

1900.

Peel, 333.

1900b.

Tornier, 582.

1901.

Lampe, 191.

1902d.

Boulenger, 445.

1902.

Scherer, 255.

1905.

Neumann, 389.

1905c.

Tornier, 366.

1906a.

Duerden (not seen).

1906.

Kammerer, 48.

1906a.

Siebenrock, 823, fig. 3.

1907a.

Boulenger, 6.

1907a.

Duerden, 9.

1907.

Lonnberg, 1.

1908.

Chubb, 220.

1908a.

Kammerer, 741, figs. 3-5.

1908b.

Mocquard, 557.

1908.

Werner (1907), 1826.

1909a.

Chubb, 592.

1909b.

Chubb, 34.

1909a.

Siebenrock, 522.

1910.

Siebenrock, 700, 716.

1910a.

Werner, 300.

1911.

Griflani, 3.

1911.

Lampe, 146.

1911.

Lonnberg, 7.

1911a.

Masi, 29.

1911d.

Stemfeld, 49.

1912b.

Boulenger, 329.

1912c.

Stemfeld, 198, 200.

1912b.

Werner, 433.

LOVERIDQE AND WILLIAMS: AFRICAN CRYPTODIRA

237

1913c.

Nieden, 54.

1913d.

Werner, 40.

1914a.

Hewitt, 244.

1915.

Werner, 330.

1916.

Calabresi, 42.

1919.

Sclimidt, 600.

1921d.

Loveridge, 50.

1922.

Falk, 163, fig. — .

1923g.

Loveridge, 925, 931.

1924b.

Loveridge, 2.

1925b.

Flower, 925.

1927.

Calabresi, 37.

1928.

Cott, 952.

1928d.

Loveridge, 49.

1929.

Flower, 32.

1929h.

Loveridge, 14.

1929.

Eose, 188.

1931.

Hewitt, 499.

1932b.

Parker, 340.

1933h.

Loveridge, 206.

1933a.

Power, 214.

1934a.

Mortens and Miiller in Eust, 9.

1934.

Pitman, 307.

1935b.

FitzSimons, 303.

1937.

Buxton, 101.

1937a.

Flower, 9, 37.

1937b.

Monard, 146,

1937.

Euckes, 103.

1939b.

FitzSimons, 18.

1940a.

Scortecci, 126.

1943.

Scortecci, 270, 282.

1944.

Leaiey, 396, figs. — .

1946.

Bally (1945), 163.

1946.

CairncToss, 395, pi. i.

1946.

Mitchell, 20.

1947.

Stevenson-Hamilton, 319.

1950.

Williams, 551, 552.

1952.

Bally, 236.

1954.

Battersby, 248

1954.

Noel-Hume, 75, pis. i-iv.

1893a.

Homopus signatus Boettger (part), 8 (juv. Shell ex Great Namaqua

land).

1885.

Testudo sulcata Smets (not of Miller: 1780), 8.

238 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1904. Testudo calcarata Vaillant (part; not of Schneider), 186, figs, (are

clearly of a juvenile p. babcocJci) .
1926. Tortoise, Bowker, 37.
1942. Hull, 125.

1948. Cooper, 11.

1933b. Megachersine pardalis Hewitt, 257.
1935c. Testudo pardalis hahcocki Loveridge, Bull. Mus. Comp. Zool., 79,

p. 4: Mount Debasien, Karamojo, tJganda.
1936h. Loveridge, 18.

1936J. Loveridge, 220, pi. i, figs. 1-2.

1937f. Loveridge, 492, 495.

1937d. Mertens, 2.

1947g. Loveridge, 136.

1949b. Loveridge, 19.

1953e. Loveridge, 158.

1955a. Mertens, 35, pi. iii, figs. 9-10 ; pi. xxii, figs, 129-134.

1955b. Mertens, 52.

1937a. Geochelone pardalis Hewitt, 789, pi. x, fig. 6.
1937e. Hewitt, 3 and G; pis. i, figs. 1, 3, 4; ivA, fig. 4; xxvii, fig. 4.

1948a. Archer, 74, figs.

1948b. Archer (part), 75, figs, (omit Keetnianshoop).

1950. Rose, 338, figs. 203-205.

1937b. Testudo pardalis pardalis Mertens (not of Bell), 5.
1955. Hellmich, 14, 15 (of reprint).

Common namefi. Eastern Leopard Tortoise; Mouutain Tor-
toise. The first name, used by Gray (1855:9), is preferable, for
the species occurs on the plains as well as on mountains, where
it may be more frequently encountered today possibly as a result
of its having been killed off on the lowlands. Akuma''- (Karamo-
jong) ; anassi (at Cape Delgado; Peters) ; herg-schilpad (Afri-
kaans: Bowker); camba (at Tete :Peters) ; chepkoikochet (Kip-
sigis and Nandi:Hull); ekudu (Samia:Hull) ; ekutu (Kitosh:
Hull); enkuru (Kisii :Hull) ; fulahomha (at Capangombe :Bo-
cage) ; fiirgohi^ (Swahili) ; ikiidu'^ (Gishu) ; kavnba'^ (Nyungwe
at Tete); likudu'^ : (Maragoli; Tereki) ; liikutu (Bunyore and
Hanga:Hull); malugangi'^ (Gogo) ; nifudi'^ (Ganda) ; nguru'^
(Taita:Peters) ; opuk (Luo:Hull); prokoch (Suk:Hull).

1 Fide Loveridge. Mitchell gives furu and kamba for tortoises but no evidence
that Geochelone occurs at Dowa or Fort Maguire.

LOVEBIDGE AND WILLIAMS: AFRICAN CRYPTODIRA

2;i9

Description} Beak scarcely to strongly hooked, edge of jaws
strongly dentate ; prefrontal large, frequently single but usually
divided longitudinally ; frontal broken up ; remaining upper head

Fig. 18. Internal view of plastron of Geochelone pardalis babcocTci (M.C.
Z. 40004) (x 1/3).

(P. Washer del.)

shields small, irregular ; forelimb anteriorly with large, unequal,
usually scattered or rarely juxtaposed, generally imbricate

1 Based entirely on material in the M.C.Z. unless otherwise stated.

240 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

scutes, forming 3-4 longitudinal and 7-9 transverse series from
elbow to outer claw ; claws 5 ; hinder side of thigh with 2 or more
very large, rarely small, conical tubercles; heel with large, coni-
cal, spurlike tubercles; claws 4; tail without terminal clawlike
tubercle.

Carapace very convex, sides descending abruptly, deeply
notched in nuchal region except in the very young, anterior
margins not, and posterior margins only sometimes, expanded,
reverted, and more or less strongly serrated ; dorsal shields con-
centrically striated, sometimes swollen, subconical or convex;
nuchal absent ; vertebrals 5, rarely 6,^ more or less convex, first
as broad as, or broader than, long, the rest broader than long,
broader than the costals; costals 4, rarely 5, not forming an
angle with the marginals; marginals 10, 11 or 12;- supracaudal
undivided, somewhat incurved.

Plastron with front lobe not or but slightly produced and not
or but weakly notched; gulars paired; pectorals very narrow,
their anterior border usually straight, widening abruptly to-
wards the axillary notch; axillaries 2, one large, the other mi-
nute; inguinal moderate, normally in contact with the femoral,
rarely separated f hind lobe deeply or slightly notched pos-
teriorly, usually angular, occasionally crescentic,'* an interanal
scute sometimes present.

Plastral formula : Abd> h>or=g>or<f >or=p>or<an.

Color'. Carapace of hatchling'^ dull yellow, a broad black
border (sometimes mottled with yellow) surrounds each ver-
tebral and costal shield; within this border the ground color is
light brown; typically each areola has a reddish brown border
of moderate width within which there is a single central red-
brown spot that usually coalesces posteriorly with the surround-
ing areolar border; upper and lateral edges of marginals and
supracaudal bordered with black within which the areolae are
edged with reddish brown. Plastral shields yellow, more or less
mottled and edged with blackish brown. Head and limbs yellow-
ish brown, uniform.

1 In a juvenile from Artu only, fide Tornier (1905c).

2 In a specimen (A.M.N. H. 50739) from Blanket! River, Tanganyika.

3 Separated only in a specimen mentioned by Hewitt (1931).

4 In the holotype $ (M.C.Z. 40003) for example.

5 Chiefly based on a series of specimens (M.C.Z. 42162-7) from Waterberg.
Southwest Africa. For illustrations of juveniles from this area see Mertens

( 1955a :pl. xxii, figs. 129-134).

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIEA 241

A juvenile from Ethiopia, referred to "calcarata" {=sulcata)
by Vaillant (1904a) only differs from the Waterberg series in
having lateral areolar spots alongside the central ones of the
second and third vertebrals. Moreover Tornier (1905c :366) finds
a 74 mm. "pardalis" {=babcocki) from "Artu," Ethiopia,
differs but little from one taken in Ilereroland, Southv^est Africa.
A Mongalla juvenile in the Vienna Museum, photographed for
us by Dr. J. Eiselt, is very similar to these Ethiopian ones.

Two juveniles from the Ithanga Hills (M.C.Z. 31985) and
Southern Guaso Nyiro (U.S.N.M. 41685), measuring 51 and 40
ram. respectively, have yellow carapaces whose shields are devoid
of dark spots (except for an azj'^gous spot on the fourth vertebral
of the Guaso Nyiro tortoise). Both the dorsal and marginal
shields of the latter have an inner dark border, while in the
Ithanga Hills juvenile these borders are deep black and cover
the sutures, coalescing with the corresponding borders of the
adjacent shields.

Carapace of adults yellow with black markings. These are
zonary in the young (whose areolae are usually brown) but
i-adially arranged or broken in adults, where they may be so
dense as to almost obscure the yellow ground color. Plastrons
are invariably yellowish, either uniform or with black radiating
streaks and scattered spots.

A Sudanese specimen in the British Museum, accompanied by
full MS notes by Flower, is very light in color with very few
black flecks. Another from the same locality is more nearly
typical. According to Neumann (1905:389) Somali specimens
are paler than those from Tanganyika, and Siebenrock (1906a:
823) suggests that tortoises from north of the equator tend to be
lighter than those from the south, the coloring of j^oung Sudanese
bahcocki approaching that of sulcata. Actually the carapace of
a subadult Northern Guaso Nyiro specimen (M.C.Z. 7990) is
heavily variegated with black except on the areolae.

The literature teems with descriptions of individual coloring.
The pattern is discussed at some length by Duerden (1907b :74).

Size. Carapace length of a 5 from Albany, 585 mm. (i.e. 23
inches in a straight line) ; 916 mm. (36 inches^) over the cara-

1 Possibly it was these figures that gave I'ise to Stevenson-Hamilton's (1047 :
•319) misleading statement that leopard tortoises attain a maximum length of
"between 2 and 3 feet" (i.e. about 609 to 914 mm.).

242 BULLETIN : MUSIiUM 01'' COMPARxVTIVE ZOOLOGY

pace; girth 1116 mm. (44 inches) (Hewitt :1937e:4). Masi
records (1911a :29) an unsexed Ethiopian specimen as 450 mm.
Carapace length of a ^ from Arusha, 340 mm., height 196
mm.; that of the holotype 9 from Mount Debasien was only
364 mm., height 207 mm., being surpassed bj' a $ from Mount
Mbololo with a length of 385 mm., and a height of 208 mm.

A table giving the length, breadth and height of twenty-three
tortoises from Southwest Africa is given by Mertens ( 1955a :35).
Length of claws on hind foot, 30 mm. (Cairncross:1946).

Weight. A 2 from near Aberdeen, weighing about 18 lbs.
when captured in 1929, has had its annual growth and fluctuat-
ing weight recorded by Cairncross (1946:395) from 3.vii.38 to
7.xii.41 when it attained about 500 mm. {19%q inches) in
length, and weighed 45 lbs. Mrs. Leakey (1944:396) gives the
weight of a 5 as 18 lbs., and those of two 9 5 as 37 and 45
lbs. respectively. Hewitt (1937e:4) mentions a 3 of 50 lbs.,
while Archer (1948a :75) gives 69 lbs. as the maximum but
states that some hahcocki remain at about 10 lbs. while others
continue to grow until they reach 30 lbs. Both the heaviest
records appear to be those of specimens in the Albany Museum,
Grahamstown.

Sexual dimorphism. While some individuals are readily sexed,
others are not. Certain characters mentioned by FitzSimons
(1985b :304), though doubtless satisfactory for his material,
appear quite inadequate when applied to an extensive series.
Thus there are exceptions to the general rule that the lower
sides of the carapace are almost vertical in $ $ and bulging for
the accommodation of eggs in adult 9 9 , though it remains a
good average character. Again, the notch on the posterior lobe
of the plastron is frequently more angular in S $ , though in
one old S there is scarcely any angle whatever, the sides being
almost in a single plane ; in the 9 9 it is widely angular, even
semicircular. The shape of the supracaudal, often thought to be
diagnostic of sex, is not so in this species. The number of times
the breadth of the carapace is contained in the length is largely
an age character, ranging from 1.01 to 1.45 times in the young;
1.49 to 1.55 times in our 6 adult S 3 ; 1.45 to 1.68 times in the
10 oldest 9 9 . We are reduced to the following characters,
therefore, of which the relative tail length is the surest guide.

LOVERIDGE AND WILLIAMS: AFRICAN' CRYPTODIKA 243

Posterior third of plastron slightly concave in adults; tail
long $

Posterior third of plastron as flat as the rest in all ages;
tail short 2

Breeding. When two $ $ and a 9 shared an enclosure at
the East London Museum, Archer (1948a :75) observed that
the $ i would push and butt one another until one of the
combatants was overturned. Then, unless in a depression, the
vanquished tortoise would right itself and hurry away. The
victorious $ then pursued the 2 , which seemingly evinced no
interest, bumping against her shell and walking around her —
sometimes for hours at a stretch according to P. J. Styrdom (in
Hewitt :1937e :6) — until pairing took place.

In East Africa, Bally (1952:236), who had a single $ con-
fined with two 2 9 (he has satisfied us as to sexing), states that
it was the 9 9 who butted each other's flanks until one of the
contestants was overthrown. Furthermore, so Bally informs
us (by mail), the larger 9, apparently annoyed at the S
getting in her way, rammed him also! This seems to refute
Hewitt's (1937e:6) statement that 9 pardalis do not butt with
the front of their shells.

Mrs. B. E. Leakey (1944:396) records that mating was a com-
mon occurrence among her captive tortoises. It began with
the $ following one of the two 9 9 around and around, oc-
casionally for several hours, until he was accorded the oppor-
tunity to mount. On doing so he would extend his long neck and
utter a husky cry that carried for a considerable distance.
Hewitt (1937e:6) describes this call as ''a peculiar grunting

noise."

Nesting. For 2 or 3 days prior to laying, the expectant tor-
toise walks restlessly about the enclosure, pausing now and
again to scratch the ground, while in each eye a frothy white
foam^ gradually accumulates (Cairncross:1946 :395).

Apparently Duerden (1906a) was one of the first to record the
nesting of this species, but a more detailed account is that given
bj' Loveridge (1923g:926) of the nesting at Kilosa of a captive
tortoise from Pwaga. Briefly it may be said that the reptile
was first observed to be digging at 5.15 P.M. on May 21. Despite

1 Probably au individual characteristic only.

244 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

the extreme hardness of the soil, a circular hole about 4 inches
in diameter by 2 inches in depth had been dug already. Excava-
tion was accomplished by the insertion of a hind foot which
was scraped round and then brought up with a very small
quantity of soil precariously balanced upon it; even so, at the
last moment, much of the dirt fell back into the hole. The loss
would have been even greater had not the tortoise from time
to time micturated copiously so that the fluid, besides softening
the hard ground, caked and bound the powdery soil. The hind
feet were employed alternately with brief pauses for rest be-
tween the exchange. On reaching a depth of 4 inches the tortoise
undercut the sides to form a gallery round the periphery, then
continued excavating until a depth of 5 inches was reached.

Darkness fell at 6.30 P.M. At 7.12 P.M. the first egg was
laid, its descent being retarded by an enveloping column of
viscous fluid. At 7.24.10 P.M. the seventh and last egg was de-
posited. As each egg was laid the tortoise put do"«Ti a foot and
felt about for the egg which she then pushed beneath the under-
cut. At 7.26.10 P.M. the mother began breaking down the sides
of the hole after which she proceeded to fill it in with all the
recently excavated soil. Not once during the period of approxi-
mately three hours that she had expended on the task did the
parent face about to examine the progress of her labours, or
inspect the eggs which she had buried without ever seeing.

So impressed was Loveridge by the mechanical nature of
the entire proceeding that he lifted the tortoise off the nesting
site to some hard ground a couple of feet away. There she
continued to trample without intermission until returned to the
nest site. After a while she began raising herself on all four
limbs as high as they would permit, then, suddenly relaxing, she
dropped upon the site so that the disturbed soil was flattened
by her plastron ; this process of pounding the soil was continued
for a long time. The following morning she was found to be
resting on the site and, though the enclosure was a large one,
she returned to sleep upon it that evening and for many subse-
quent nights. On June 12 Loveridge noted that not only did
she sleep on the site but micturated and defecated on it fre-
quently, if not daily. Other observers have reported that the

LOVERIDQE AND WILLIAMS: AFRICAN CRYPTODIRA 245

parent tortoise may be so exhausted by her efforts that she can
scarcely drag herself away from the site.

Evidently nest holes vary considerably in size, for Archer's
had a surface diameter of 6 inches and a depth of 8 inches.
Cairncross says that his was 7 inches across at the surface, about
9 inches at the bottom, while the total depth was 9 inches. Mrs.
Leakey's flask-shaped cavity was about a foot in depth, but had
to accommodate an exceptionally large number of eggs, 30 in
all. When numerous, the eggs are arranged in layers that are
separated by earth {fide Bowker :1926). The number produced
at one laying, therefore, ranges from 5 (Cairncross :1946 :395)
to 30 (Leakey :1944:396). Photographs of a nesthole being dug,
and of one with eggs in situ, have been published by Cairncross
{loc. cit., pi. i), one of whose tortoises laid a total of 52 eggs
(in 6 installments) during the summer of 1939-40. Their gross
weight was 357.48 grams (about 6 lbs.).

Both in appearance and size the egg of a Leopard Tortoise
rather resembles a pingpong ball but the porcelain-like shell is
very brittle. Occasionally, as with poultry, a double, elongate
egg with median constriction (figured by Archer) may be laid.
Perhaps a tendency of this kind was exhibited by the Sudanese
egg mentioned by Siebenroek (1906a :823) as measuring about

35 or 37 x 43 mm., unless the 43 was a misprint for 33 mm.
Loveridge (1936j:220) removed 8 hard-shelled eggs measuring

36 X 38 mm. from a 2 taken on Mt. Mbololo on April 17, there-
fore larger than the 7 measuring 32.5 x 35 mm. laid by a Pwaga
tortoise on May 21, and smaller than 2, measuring 40 x 40 mm.
and 38 x 40 mm., laid by a captive on August 5 and 25 respec-
tively.

Eggs laid very early in the South African spring may hatch
in 8 months provided that warm weather continues on into late
autumn, but autumn laid clutches may take as much as 18
months to hatch if the following summer is cold (Archer :1948a :
75) . Eggs laid in a Grahamstown garden on February 7, hatched
on April 23 of the following year, i.e. they took 14 months,
whereas two clutches laid in the South African autumn took
only 11 and 12 months respectively (Bowker :1926 :37). In
South Africa, a presumably captive Leopard Tortoise laid 3

246 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

eggs on the ground at Grahamstown (Hewitt: 1937a :6), be-
tween July 26 and 28, mid-winter.

Mrs. Leakey (in Hit.), after removing eggs from the nest,
packed them in sand in a gasoline can which she placed on a
cement hot-water tank. Nine months later, sometime between
December 14 and 21, there emerged a single hatchling weighing
half an ounce. As no others appeared during the succeeding
3 weeks the rest of the clutch were examined. Two eggs only
held young tortoises, both of which were dead.

As eggs left in situ rarely hatched, Cairncross (1946:396)
removed many clutches to an incubator where, except for a 10
minute cooling off period each morning, a temperature ranging
from 88° to 92° was maintained for 6 months. The results were
disappointing as the majority of eggs proved infertile. For
further details consult his paper.

In one of BoAvker's nests (1926:37) the central clay plug
subsided slightly, permitting the hatchlings to squeeze their way
out into the sunlight between plug and wall. Normally the
young, whose shells are flexible at time of hatching, claw their
own wa}^ to the surface. Being well supplied with nourishment
from the yolk they require no food for several days, despite a
display of considerable activity.

Growth. Rate of growth is affected by availability of food
coupled with congenial temperatures, being accelerated during
warm weather and retarded by cold. Good and bad seasons are
reflected in the size of seasonal growth-rings on the shields of
the carapace. In immature tortoises these rings furnish an indi-
cation of age, but becoming blurred with wear are no longer a
trustworthy guide when maturity is reached. Consequently,
statements regarding growth which are based on data derived
from one or two tortoises, vary widely. P. J. Strydom is quoted
by Hewitt (1937e:4) as saying that in South Africa only a
few inches in length are gained during the first decade, making
a total of 7 inches after 17 years. Archer's (1948a :75) detailed
figures seem to support this. He furnished a growth chart of
one of many tortoises (No. 64) studied at the East London
Museum.

Fisk (1878 :790) reports that two fully-grown ''pan/a? ks" were
brought to Clanwilliam in 1846, and there remained without

LOVERIDGE AND WILLIAMS: AFRICAN CBYPTODIRA 247

progeny until 1857. In that year 2 young hatched from eggs
laid by the 9 and Fisk assumes this to imply that the parents
had only then become adult! In 1877 the parents died within
3 months of each other, at which time, though 20 years old, the
young ones were but two-thirds the size of their parents. The
young were then deposited in the London Zoological Gardens.

Longevity. According to Flower (1937a :9) the record for
known longevity is furnished by the above-mentioned tortoises
which lived in captivity for over 30 years (1846-1877).

Diet. Principally grass, though they also feed on various
succulents, crassulas, spekboom, thistles, prickly pear, pumpkins,
beans and water melons (Hewitt :1937e :5) ; sweet potato tops
or an entire, hard, white cabbage may be consumed at one meal
(Leakey :1944:396) ; verdure of ground nuts, lettuce, and mois-
tened bread (Loveridge :1933h :206) ; grapes (Rose :1950 :339).

Bally (1946; 1952) observed a captive Ethiopian specimen
take up and swallow a meatless chicken bone twice as long as
its own head. Some days later the reptile was found nosing and
pushing around a much larger bone. When Bally smashed this
bone the tortoise seized the jagged splinters, one of which was
an inch and a half long, and swallowed them one after another
until all were gone. Thereafter he supplemented the reptile's
vegetarian diet with a daily ration of bone splinters upon which
it throve. For several years a companion tortoise from Lake
Chala showed no interest in bones; then, in 1947, the Chala
reptile also started eating chicken bones and doubled its weight
in a year. It almost doubled its weight each succeeding year as
the Ethiopian tortoise had been doing.

Bally tabulates the annual growth and his figures show that
the Chala tortoise was still about doubling its weight each year
in 1950, at which time the Ethiopian reptile's development was
slowing down. Bally wondered whether the bone diet was re-
sponsible for the Ethiopian specimen's increase in size and
weight during the j^ears that the Chala tortoise remained almost
static. (AVe reject the suggestion that Ethiopian tortoises pos-
sibly grow larger. There may be some confusion with sulcata.)
A possible source of calcium for feral tortoises is to be found
in the excrement of hyenas and other carnivores for captive
specimens have been known to eat the dry dung of dogs as well
as owl eastings.

248 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

After eating, though it may have been on fleshy Cissus leaves
or lettuce, Bally 's tortoises drank regularly and eagerly. Though
drinking copiously in summer, leopard tortoises can go for long
periods without water (Hewitt :1937e :5).

Even when feeding daily on rich food a lapse of from 2 to 3
weeks occurred between defecations. Bally (1952:236) noted.
The volume of such excrement was correspondingly large, fre-
quently exceeding a tenth of the total weight of the depositor.

Parasites. Ticks (Aynbly omnia exornatum and A. nuttalli, not
A. marmoreum as previously reported in error) were present on
turtles from many localities.

Enemies. The eggs are dug up and devoured by dogs (Lea-
key), jackals, meerkats, muishonds. Young tortoises are attacked
by crows and ground-hornbills, larger ones by ratels (Hewitt),
seemingly by hyenas also; certainly by soldier ants (Loveridge).
Nyamwezi tribesmen were seen eating them by Ansorge (1899:
288). Scherer (1902:255), when in the Mkomasi Valley, en-
countered a couple of Africans carrying a pair of poles to which
was attached a huge tortoise about 500 mm. in length that the
men said would provide them with a good meal. One of us
(Loveridge) was told that in the Iringa Highlands a really
large tortoise was considered to be worth 2 goats! According
to FitzSimons (1935b :304) Bushmen prize these reptiles both
for their flesh and shell. The latter, after removal of the plas-
tron, is used for ladling or holding water, while Dr. R. Poch
states that the shields are made into snuffboxes by the Kalahari
Bushmen.

At Magugu, Cooper (1948:11) found a tortoise, apparently
of this species, lying on its back and unable to right itself.
Surrounding spoor suggested that the reptile had been over-
turned by a buffalo's nose, after which the ungulate had pawed
up the ground in the vicinity.

That many tortoises perish in the annual grass fires is sug-
gested by Ansorge (1899 :288) who not infrequently encountered
their calcined remains. Many shells and skeletal remains, but no
living tortoises, were found at Tsotsoroga by FitzSimons (1935b :
304) who concluded that the reptiles had perished during the
extensive flooding of the flats a few years before.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 249

Aestivation and Hibernation. In view of the admitted lan-
guage difficulties, the alleged burying of this species during dry
and cold seasons in Southwest Africa — as reported to Falk
(1922:163) by Bushmen, should be received with reserve pend-
ing confirmation.

Migration. Peel (1900:333) reports seeing in Somaliland "a
whole army of these monsters migrating" across the desert from
one patch of grass to another.

Habitat. Leopard Tortoises inhabit the coastal plain and up-
land savanna, excluding areas of primary forest, in rather
sandy, thornbush steppe and on kopjes. Such stone-strewn hills
should have scattered scrub to provide shelter from the noonday
sun. One tortoise, who skeleton was found between two sloping
rocks by Loveridge, had presumably perished through inability
to escape from its smooth-walled prison. The seeming predilec-
tion of this species for mountain masses (on Mt. Elgon it has
been taken between 9,000 and 10,000 feet) may indicate that in
such a habitat it is less liable to molestation from man and the
larger carnivores. According to Kammerer (1908a :741) low
temperatures are resisted better by Leopard Tortoises than by the
giant Spurred Tortoise.

Localities. Sudan: Bahr el Ghazal Province (Werner :1924a) ;
Bahr el Jebel (Gebel) ; Bor; Gondokoro; Lado Enclave; Mon-
galla; (the Omdurman market record of Kammerer is rejected).
Ethiopia: Ado-Shebeli (Adoshebai) Valley; Araro (Arero) ;
"Arruena" ( ?Arenda) ; Artu; Arussi Gallaland; Boorgha coun-
try; Caschei; Dabas (Daba-as) ; "Ennia Land" ( ? En) ; "Far-
re" (? Erre) ; Javello ; Lake Abaya; Lake Zwai; "Lasman";
Neghelli; Shoa; Sibi ; South of Harar (Harrar) ; "So-Omadu"
(? Soddu). British Somaliland: Hargeisa; Nogal Valley;
Toyo Plain {ca. 9°10'N.: 44°45'E.). Somalia: "Bambasse"i
between Berdale (Bardale) and El Condul (Condut) ;
Ircut (Pozzi-Ircudt) ; Jet; "Sahaieroi."^ Uganda: *Mount
Debasien. Kenya Colony: Athi Plains; *Guaso Nyiro; *Ithanga
Hills; Kabete; Kaimosi; Kedoug Valley; Kikuyu Escarpment
south of Lake Naivasha ; ' ' Kipopotue, Ukamba ' ' ; Lekiundu
River, Southern Guaso Nyiro; *Meru River Plains; "Mgana"
(Betton coll. A locality unknown to L. Leakey and J. G. Wil-

1 A Calabresi (1916) report on Stefani and Poli material; many misspellings.

250 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

liams) ; Mount Elgon; Mount Kenya; *Mouut Mbololo ; Mtito
Andei ; Murri ; Nairobi ; Ndi ; Ndogo, Lake Baringo ; Southern
Guaso Nyiro; Turkana; *Wema, Tana River; *Ziwani Swamp,
Northeast of Kilimanjaro (CM.)- Tanganyika Territory:
*Arusha ; Blanketi River ; Busisi ; Dodoma ; Irazo ; Kahama
(Leiden Mus.) ; *Kilosa; Kondoa Irangi; Korogwe; Lake Eyasi;
Lake Jipe ; Lindi ; Luguo ; Magugu ;^ Masai Nyika and Steppe ;
Mbonoa; Mdjengo's; Mkaramo, near Mombo, Pangani River
(1896) ; Mkomasi River; Moshi ; Mtali 's ; Ruvu River, 33 km. s.e.
of Moshi ; Mwanza ; Namanga ; Ngare na Nyuki ; Ngoma ; Nzinga ;
*Pwaga ; Ruvs'ira ; Sagayo ; Sanga ; *Saranda ; Shinyanga ; Sim-
bo; *Singida; Tabora; Tambali ; *Tukuyu; Ukerewe LI.; Ulugu ;
Usambara ; Wembere ; Zengeragusu. Mozambique : Cape Del-
gado Ids.; *Kasumbadedza ; Sena; Tete. Nyasaland: (occurs
south of the Mwanza River and west of the Shire in both Chik-
wawa and Port Herald districts, according to Mitchell : in litt.) ;
Tengani. Northern Rhodesia: Luangwa (as Loangwa) River;
Petauke. kSouthern Rhodesia: *Birchenough Bridge, Sa1)i River;
near Gwamayaya River; Mount Darwin. Bechimnaland Pro-
tectorate: Gemsbok ; Kalahari; Magalapsi; Makarikari ; Maun;
Serowe; Shaleshonto; Shorobe; Tsotsoroga Pan, Mababe Flats.
Transvaal: Mannamead; Naauwpoort ; Pietersburg. Natal: Dur-
ban. Cape Province :^ Aberdeen ; Albany ; Alexandria district ;
Algoa Bay ; Alice ; Bathurst district ; Brakkloof ; Colesberg ; Fort
Beaufort district ; Grahamstown ; Hamburg ; Hogsback ; Kei-
.skama; Malmesbury; Middleburg; Middleton; Modder River;
Mortimer Road ; Nonesis ; Orange River Station ; Port Elizabeth ;
Queenstown ; Saint Clair ; Somerset East district ; Tarkastad ;
Tottabi near Alicedale ; Uitenhage district; Vryburg; Warren-
ton ; Willowmore. Southwest Africa: *Etosha Pan-near ; Gibeon ;
G reat Namacpialand ; Karibib ; Landsberg Farm, Luderitz dis-
trict; Neudamm Farm; Ogosongomingo ; Aus (Oas) ; Oka-
handja ; Omaruru ; Omatjenne ; Rietmond near Gibeon ; Swakop-
mund; *Tses (U.S.N.M.) ; *AVaterberg; Windhoek (Windhuk).-

1 Unidentified "land tortoise" overturned by hulTalo (Cooper :1948) . Magugn,
near Mbugwe, slightl.v south of Lake Manyara.

2 Through the courtesy of Mons. J. Guibe we liave been able to examine one of
Dolalantle's voung tortdises from the "Cap." It is unfortunate that no mor<'
precise locality is known. In pattern it is a p. pardalis of the •'biguttata" type.
but in pioiiortions is clearly ref<>ral)le to /). habcovl;i tor its height of 4r( mm. is
included in its length of 83 mm., 1.S4 times. It must therefore be considered an
intermediate.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 251

Angola: Benguela — inland from; Capangombe — some distance
from ; Cubal ; Dombodola ; Forte Rocadas ; Mossamedcs — in-
terior of ; Mupa.

Range. Sudan and Ethiopia south to Natal, west through Cape
Province to Southwest Africa, where it meets with the typical
form, but is dominant north of 27°S. and in southern Angola.
It does not occur in northern Angola {fide Monard :1937b).

Erroneously reported from Madagascar (1873) and Sene-
gambia (1884a).

Geochelone pardalis pardalis (Bell)

1828a. Tcstudo Pardalis Bell, Zool, Jour., 3. p. 420, pi. xxv of Suppl.:

"Promont. Bonae Spei, " i.e. Cape of Good Hope.
18;U1). Gray, 4.

1831c'. Gray (part), 12 (oinit figs, as these are of p. babcoohi).

1835. Dumeril and Bibron (part), 71 (color of young, etc.).

1836. Bell, text and col. pis.
1849. Smith, A., App., 1.
]8o]. Dumeril and Dumeril, 4.
1860. Fitzinger, 411,

1872b. Gray, 3, in Sowerby and Lear, pis. ix-x.

1892. ?Muller, F., 214.

1907b. Duerden, 74, fig., pi. vii, fig. 7 (showing juv. markings).

1829. Tcstudo biguttata Cuvier, Eegne Animal, ed. 2, 2, p. 10, footnote

(nonien nudum) : No locality.
1831l'. Tcstudo hipunctata Gray, 12 (erroneously attributed to Cuvier ms8.
and cited with reference to the description of a juvenile in the
Paris Museum).
1831b. Gray, 4 (erroneously attributed to Cuvier in Regne Animal).

1831c. Test ado armuta Boie niss. Gray, 12 (listed as a synonym).
1831b. Gray, 4.

1835. Geochelone pardalis Fitzinger, 211.
1937e. Hewitt, pi. i, fig. 2 ; pi. ii, figs. 1-2.

1948b. Archer (part), p. 77 (Keetmanshoop specimen).

1935c. Testudo pardalis pardalis Lioveridge, 4: (key).
] 955a. Mertens, 35.

Most citations of "pardalis," and one of "pardalis pardalis,"
will be found listed under p. halcocki, with which race the typi-
cal form has been confused for almost a century.

Common names. Southwestern Leopard Tortoise (preferred) ;
Mountain Tortoise.

252 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Illustrations. Excellent colored plates of the $ holotype (side
and plastral views) appeared in Bell's (1836) rare Monograph
of the Testudinata, and were later reproduced by Sowerby and
Lear (1872b) . The distinctive coloration of the young p. pardalis
has been figured by Duerden (1907b :fig. 7, based on a specimen
beginning to assume the adult coloration), and by Hewitt
(1937e:pl. 1, fig. 2, a hatchling closely resembling the Aroab
specimen in the Museum of Comparative Zoology).

Description} Beak moderately hooked, edge of jaws not
strongly dentate; prefrontal large, divided longitudinally; re-
maining upper head shields small, irregular ; forelimb anteriorly
with a few large, unequal, widely separated, non-imbricate
scutes, forming 3-4 longitudinal and 7 transverse series from
elbow to outer claw ; claws 5 ; hinder side of thigh with 2 large
conical tubercles; heel with large, flat tubercles; claws 4; tail
without terminal clawlike tubercle.

Carapace flattened dorsally, sides nearly vertical (at least in
(5 adult), sloping gradually in young, deeply notched in nuchal
region, in adults the anterior margin not, and lateroposterior
margin only slightly, expanded and slightly serrated, its greatest
height included more than twice (2.02 to 2.62 times) in its
total length; dorsal shields concentrically striated except in
young, their areolae slightly raised ; nuchal absent ; vertebrals 5,
more or less (in young) convex, all broader than long, broader
than the costals; costals 4, not forming an angle with the mar-
ginals; marginals 11; supracaudal undivided.

Front lobe of plastron anteriorly truncate in young, slightly
produced and shallowly notched in adults ; gulars paired ; pec-
torals very narrow, their anterior border slanting in young,
more or less straight, widening abruptly towards the axillary
notch ; axillaries 2, one small, the other minute ; inguinal small
in contact with the femoral ; hind lobe deeply notched posteriorly,
almost crescentic.

Plastral formula: Abd>h>or=g>or<f>or=p>or<an.

Color. That of a hatchling from Aroab (M.C.Z. 22473) is
strikingly different from juveniles of G. p. habcocU, viz. carapace
dull yellow, black being confined to the narrow sulci surrounding

1 Cased on original description of 9 holotype, the shell of an adult $ (M.C.Z.
1.6713) and an aleoholic juvenile (M.C.Z. 22473), together with notes on British
Museum material as of 1953.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 253

each vertebral and costal shield; these are edged with yellow,
each areola with a reddish brown border, those on the vertebrals
flankinf>' a pale cream, broad, median band; upper and lateral
edges of marginals and supracaudal bordered with reddish
brown. Head and limbs peppered with black. The juvenile tor-
toise figured by Seba (1735 :pl. Ixxixtfig. 3), which Bell sug-
gested might be the young of pardalis, is more probably a young
P. geometricus of the luteola phase (see below).

The coloring of the adult, as described by Bell, is substantially
similar to that of our specimen, also of the eastern race — p.
hahcocki. Carapace light yellow; shields with black, somewhat
radiating, markings. Plastron dirty yellow, each shield with
black spots, dashes, or radiating streaks extending inwards from
the margins. Head, neck, feet and tail, dull brown with some
admixture of dull yellow.

Size. Carapace length of type $ , 432 mm., breadth 280 mm.,
height 213 mm.; carapace length of $ (M.C.Z. 16713), 302 mm.,
breadth 210 mm., height 147 mm. ; carapace length of juv.
(Aroab), 55 mm., breadth 42 mm., height 21 mm.

Enemies. Possibly the food value of this large tortoise may
have led to its extermination on the Cape for today its range
appears to be restricted to a small area in the southern part
of Southwest Africa.

Habits. Bell (1828a; 421) states that since the beginning of
summer the t\i:>e had been living in his small orchard, feeding
heartily on the grass which it plucked with movements similar to
those "of a goose." The strength of an adult Leopard Tortoise
is surprising for once it has braced its feet it is next to impossible
to pull the reptile backwards. After its death, Bell (1836) dis-
sected the type and found it held an estimated 200 ova ranging
in size from those no larger than a pin's head to others equalling
a pigeon's egg.

Localities. Cape Province: ? now extinct: Cape of Good
Hope 1 (Type) ; Olifants River (Delalande coll. ? this subspe-

1 Walter Rose informs us (6.V.54) that, during thirty years of rambling in the
vicinity of Cape Town, he has never encountered a leopard tortoise — except
I'scapees. He believes it formerly occurred 45 miles northwest of Cape Town In
the Malmesbury district where one meets with arid conditions resembling those
of the Karroo.

254 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

cies). Southwest Africa: *Aroab; Keetmanshoop^ *Kolman-
skop ;- liehoboth.''

Range. Southwest Africa possibly as far north as Rehoboth,-^
more certainly south of a line from Kolmanskop (near Luderitz
Bay) to Keetmanshoop, soutli (in former times presumably) to
the Cape.

Erroneously reported from Angola (Monard: 1937b).

Genus TesTUDO Linnaeus

1758. Tcstudo Linnaeus, Syst. Nat., ed. 10, 1, p. 197. Type: Testudo graeca

Linnaeus (designation by Fitzinger).
1820. Chersine Merrem, Vers. Syst. Amphib., pp. 12, 29. Type: T. graeca

Linnaeus (designation by Lindhobn).
1830. Chersn^ Wagler, Natur. Syst. Amphib., p. 138. Type: Tcstudo mar-

ghiata Schoepff {hj monotypy).
1869c. PcUastcs Gray (not of Eossi: 1807), Proo. Zool. Soc. London, pp.

167, 171. Type: T. graeca Linnaeus (designation by Lindholm).
1870e. ChersineUa Gray, Suppl. Cat. Shield Eept. Brit. Mus., part 1, p. 8.

Type: T. graeca Linnaeus (designation by Lindhobn).
1870e. Testudinella Gray (not of Bory: 1826), Suppl. Cat. Shield Eept.

Brit. Mus., part. 1, p. 12. Type: T. liorsfieldi Gray (by monotypy).
1872c. Peltonia Gray, App. Cat. Shield Eept. Brit. Mus., part 1, p. 4. Nom.

nov. for Peltastes Gray (preoccupied).
1916. Medaestiu Wussow, Wochenschr. Aquar.-Terrar. Kunde, 13, p. 170.

Type: T. graeca Linnaeus (designation by Mertens).
Definition. Skull with triturating surface of maxilla moder-
ately or weakly ridged or without ridging ; median premaxillary
ridge absent ; maxillary not entering roof of palate ; anterior
palatine foramina small, concealed or large and conspicuous ;
prootic typically concealed dorsally and anteriorly iy parietal :
quadrate enclosing stapes or not; surangular subequal in height
to prearticular ; neck with third or fourth centrum biconvex.

1 This Keetmanshoop record is assumed solely on geographical grounds, being
taken from Archer (194Sl3;T7) who lists it with many others as Oeochelone
pardalia.

2 It is only right to mention that in the Vienna Museum there is a specimen
allegedly from Swakopmund (Poch coll.) whose breadth is contained in its length
2.10 times, and another of 2.11 times from Aus where Poch collected many G. p.
habcocki. We propose regarding these two individuals as intermediates between
the two races which meet in the Luderitz Bay region.

3 The Rehoboth record is based on a photograph of two young specimens kindly
sent to us by Dr. Robert Mertens in a letter ot 18.V.56.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 255

Carapace never hinged ; typically the anterior neurals alter-
nately octagonal and quadrilateral ; outer side of third costal
scute about as long as, or longer than, that of fourth ; subraargi-
nal scute absent ; frequently a single suprapygal, if two, they are
typicallj' separated by a straight transverse suture.

Phistron with ])osterior lobe more or less hinged in one or both
sexes; gular region but little thickened or produced; gulars
paired, longer than broad.

Range. Northern Africa, southern Europe, and western Asia.

Fossil record. Extensively known from the Cenozoic of
P^urope, and since the Pliocene from North Africa.

Ronorks. The genus Testudo includes forms of moderate to
small size, the living species attaining as much as 280 mm. in
some cases, as little as 120 mm. in one instance,

Testudo graeca, though technically the type of Testudo, is not
typical of the Boulengerian grouping which has gone under this
generic name. In our belief, as indicated in the diagram of rela-
tionshij^s above (Fig. 8), Testudo graeca and its immediate rela-
tives are not closer to the species pardaHs or clegans or denticu-
lata or gigantea or emys or radiata than are the species of
Gopherus or of Homopus, forms which have long been put in
distinct genera. Though conventionally the graeca group, at
best, has been one subgenus among many subgenera, an inspec-
tion of the characters bj' which the subgenera that we unite under
Geochelone are separated, and a comparison of these with the
peculiarities of the graeca group, makes it plain that different
orders of magnitude of divergence are involved.

^Ye have here suggested the separation of two larger genera
Geochelone and Testudo — as w-ell as the smaller units Mala-
cochersus, Psammohates and also Chersina, the latter all Ethi-
opian endemics. We have considered another possibility, that,
while tlic Ethiopian endemics might still be separated, Geoche-
lone and Testudo might be regarded as subgenera within a larger
concept of Testudo. This would have the apparent merit of re-
moving from the Boulengerian genus Testudo only the Ethiopian
forms that have never fulfilled the Boulengerian definition of
the genus But it appears to us that this suggestion founders
when the ( ropherus-Stylemys lineage is taken into account. The
Gopher us-Stylemys line has quite as much in common character-

256 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

wise with Testudo and Geochelone as these have with each other.
Recent Gophcrus has the maxillary ridging, the concealed ante-
rior palatine foramina and the octagonal and quadrilateral neu-
rals that occur, though not quite invariably, in the other two
groupings. The sole conspicuous distinction is the median pre-
maxillary ridge, seemingly a triviality. Considering only Recent
material, as Boulenger did, it was entirely logical for him to in-
clude Gopher us in Testudo.

But the evidence now available from the fossil genus fStylemys,
which has the gopheride premaxillary ridge and is clearly a close
cousin of Gopherus, transforms the entire picture and reveals
that the roots of the gopheride lineage go back to forms with
many emydine characters. Thus it has been demonstrated (Case
1936 1; Williams :1952) that the North American Oligocene spe-
cies iStylemys nehrascensis had neurals with the primitive emy-
dine shape (hexagonal, short sided in front), and in both fore-
and hind feet the emydine phalangeal formula of 2,3,3,3,3.

Thus it is necessary to face the fact that many of the
characters common to modern land tortoises — for example, Geo-
chelone, Testudo, Gopherus and Fyxis share the octagonal and
quadrilateral neural pattern — have heen acquired in parallel.
Formerly this fact so impressed one of us (Williams :1950; 1952)
that he questioned the naturalness of the subfamily Testudininae.
Further investigation and consideration has convinced him that
all the land tortoises do spring from a common root within the
Emydinae. He now believes that the major lines within the land
tortoises had their start in the beginning of the Tertiary with an
ancestral form that in terms of its characters was then only a
slightly divergent emydine and only prospectively a testudinine.
It appears to him that as a correlative of this view, it is desirable
to recognize at least one genus for each of the major lineages that
go back to the postulated ancestral testudinine.

It is on the basis of this interpretation that we venture to
break up the Boulengerian genus Testudo and restrict its con-
cept to the graeca group. Previously (p. 218), we have mentioned
two distinctive characters of this genus — the posterior plastral
hinge (present in one or both sexes), and the concealed prootic.
A third, heretofore neglected, character which we have found

1 Case, E. C, 1936, Contrib. Mus. Paleont. Univ. Michigan, 5, pp. 69-73.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA

257

useful enough to employ in the key (p. 221), is the presence of
distinct supranasal scales (Fig. 19) otherwise occurring only in
Malacochersus. Perhaps this implies affinity between these two
genera. However, we are inclined to believe that a rather elabor-

B

Fig. 19. Head sealation in certain testudinines (cf. Fig. 22). A, Testudo
hermanni juv. (M.C.Z. 3063); B, Malacochersus tornieri (after Procter:
1922b) ; C, Testudo g. graeca juv. (M.C.Z. 1497) ; B, Geochelone p. bab-
cocU juv. (M.C.Z. 40005); E, Kinixys erosa juv. (M.C.Z. 12583).

(P. Washer del.)

ate head squamation was primitive for tortoises, and that break-
down of this condition has occurred to a greater or lesser extent
in many lines. In any event, and in spite of a certain amount of
variation (Fig. 22), this character proves a valuable external
indicator for the genus.

Within the restricted genus Testudo there are groupings at
various levels. We recognize two subgenera, both African, which
may be defined as follows :

258 BULLETIN: MUSEUM OP" rOMPARATIVK ZOOLOGY

Key to the Suhgetiera of Testudo

Maxillary with weak to moderate ridges ; quadrate enclosing stapes. Range :

Northwest Africa ; southern Europe ; western Asia

Testiulo Linnaeus (p. 261")
Maxillarj- without ridges; quadrate not enclosing stapes. Range: North

east Africa (Tripoli and Egypt only)

rscudotcstudo new subgenus (p. 276)

The subgenus Testudo sensii stricto is here interpreted to in-
clude only gracca, marginata, hoi'sfieldi and hermanni. Of
these hermanni appears to be the most primitive, possibly the
most primitive in the entire genus. Both its external and internal
characters testify to this. Externally the forelimb has relatively
small and numerous scales (in as many as ten longitudinal rows)
the larger ones mostly toward tlie outer side, and the supracaudal
is divided. The presence of a tail claw may also be primitive for
the genus. Internally the skull shows relatively strong ridging
on the maxilla and the anterior palatine foramina are fully con-
cealed. The trachea {fide Siel^enroek) is unspecialized in not be-
ing excessively long and coiled as in the two other members of
the subgenus in which this character has been studied. In all
these respects graeca is more specialized than herynanni; while
marginata, as Siebenrock (1910) has shown, is extremely close
to graeca. The two form a transition to the very specialized
dwarf species kleinmanni, graeca in particular being close enough
to suggest that kleinmanni was ultimately derived from it.

Testudo gracca has the scales of the forelimb enlarged and
forming 4 to 6 longitudinal rows, a trend continued by klein-
manni in which there are usually only 3 in longitudinal series.
In both graeca and kleinmanni the supracaudal is usually single,
and the tail claw is rudimentary or absent in both species. As
compared with hermanni the maxillary ridging is reduced in
graeca, absent in kleinmanni. The anterior palatine foramina,
more exposed in graeca than in hermanni, are fully exposed in
kleinmanni. In possessing a long and coiled trachea, graeca is
more specialized than kleinmanni; however, it is in the largest
individuals of graeca and marginata that this condition is most
pronounced (Siebenrock :1910). In one respect kleinmanni dif-
fers from other members of Testudo in a way not approached by
graeca, i.e., in having the quadrate open behind, not enclosing

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 259

the stapes. In size the two species differ radically, graeca being
one of the largest members of the genus and kleinmanni decidedly
the smallest. However, this disparity is bridged by an allegedly
dwarf subspecies, if valid, T. graeca floweri of southern Palestine
(cf. Flower :1933:745).

TesUido kleinmanni, despite the numerous ways in which it is
approached by graeca, from which in all probability it is directly
derived, is still a species rather sharply set apart. It might he
said to represent only the extreme of trends within graeca, but
the differences are abrupt and very striking. The end result is a
species which, in many of the characters considered of taxonomic
value by Boulenger (and for the most part so considered hy us),
parallels forms that are almost certainly not closely related. In
the absence of maxillary ridges it parallels Pyxis, Psammohates,
Chersina, Homopus and Kinixys, and parallels as well the last
two in its fully exposed anterior palatine foramina. In the quad-
rate failing to enclose the stapes it displays an emydine character
(probably secondary) elsewhere found in testudinines only in
Pyxis arachnoides,^ and Oeochelone sumeiri (type specimen ex-
amined at the Tring Museum).

If as we suspect it is a derivation of graeca, Testudo klein-
manni must surely have been at one time only a geographical
form of graeca. It is still allopatric to that species, dividing the
present range of graeca in two. This singular fact of distributioii
is indeed our only clue, for in Africa there is no fossil evidence
on which to found an opinion on the circumstances of klein-
manni's origin, and the evidence from Europe has not been suffi-
ciently analyzed.

Testudo graeca of North Africa and southern Spain is probably
racially distinguishable from the populations of eastern Europe
and Asia. However, Mertens (1946d) who separated it as g.
(jraeca, found only minor differences in shell proportions between
the typical form from North Africa and Ms g. ibera ranging fron)
the Balkans to Persia. Certainly the populations of eastern
Europe and Syria do not differ greatly from those in Africa. If
one may deduce from this that the separation of the African and
east European races has not been of long standing, it is natural

1 In Pyxig the condition is quite different and not primitive. Tlie slsuU structure
(jf this genus and of Acinixys is to be described in a separate paper by one of
us (E.E.W.).

260 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

to infer that formerly a continuous population extended across
southern Europe whence the invasion of western North Africa
occurred via Spain and not across Egypt. In this view the
southern Spanish population of g. graeca would be a relict
one, persisting after the general extinction of the species in west-
ern Europe. This might be due to an invasion by hermanni which
now occupies much of the region separating the two ranges of
graeca in southern Europe.

If this hypothesis is correct, then Testudo kleinmanni is not a
peculiar form that has differentiated in the middle of the geo-
graphic range of its parent species. Instead, at the time of its
origin, it was the most westerly form of the graeca complex on
whose periphery it developed. If in southern Palestine there is
really a dwarf race (T. g. floweri Bodenheimer : 1935^) resem-
bling kleinmanni in coloration — and a single shell ~ in the Brit-
ish Museum seems to support this view — the case for our
hypothesis respecting the mode of origin of kleinmanni will be
strengthened.

Key to the Circum-M edit err anean Species of Testudo

1. Forelimb anteriorly with 7-10 small scales across it; hinder side of thigh

without enlarged tubercle; tail terminating in a. distinct claw, always

present; supracaudal usually divided. Eange: southern Europe^

JwrTtianni Schneider

Forelimb anteriorly with 3-6 larger scales across it; hinder side of thigh

with or without a large tubercle; tail claw present or absent in all

three species; supracaudal usually undivided 2

2. Forelimb anteriorly with 3-6 scales across it; hinder side of thigh with

a large tubercle. Eange: western North Africa and southern Spain

(with races in extreme southeastern Europe and western Asia)

g. graeca Linnaeus (p. 261 )
Forelimb anteriorly with 3-5 scales across it; hinder side of thigh with-
out enlarged tubercle 3

1 Bodenheimer, F. S., 193o, "Animal Life in Palestine" {fide Mertens :1946cl).
We have also found the name referred to in M6m. Inst. Egypt, 1937, 33, p. 71.

2 While the carapace of this specimen (examined by us) has the predominantly
vellow hue of kleinmanni, the plastron was much blacker than in that species ;
also the pectoral shields were larger than the femorals, the reverse of the klein-
manni condition. While having the appearance of an adult, the length of its
carapace is only 114 mm., tliat of its plastron 97 mm.

3 As Lortet (1887:6) distinctly states that he never collected this species in
Syria, the specimen from Antakia, Syria, allegedly collected by Lortet, and re-
corded as "graeca," i.e. hermanni by Boulenger (1889a :177) requires confirmation.

LOVERIDGE AND WILLIAMS: AFRICAN CBYPTODIRA 261

3. Forelimb anteriorly with 5, rarely 4, scales across it; supracaudal and
posterior marginals spread out; coloration predominantly black; size
moderately large, maximum length of carapace about 280 mm. Range:

Greece only marginata Schoepff

Forelimb anteriorly with 3, rarely 4, scales across it; supracaudal only
spread out; coloration predominantly yellow; size small, maximum
length of carapace not exceeding 127 mm. Range: eastern North Africa
(Tripoli and Egypt) Meinmanni Lortet (p. 276)

Subgenus TeSTUDO Linnffius

1835. Testudo Fitzinger, Ann. Wiener Mus., 1. pp. 108, 113, 122: Type
species: Testudo graeca Linnaeus.

Definition. Supracaudal divided or single. Maxillary weakly
to moderately ridged ; anterior palatine foramina at least partly
concealed in ventral view ; quadrate enclosing stapes. Plastron of
adults with a more or less movable posterior lobe in one or both
sexes. Tail claw present or absent.

Range. Northern Africa, southern Europe, western Asia.

Testudo graeca graeca Linnaeus

1758. Testudo graeca Linnaeus, Syst. Nat., ed. 10, 1, p. 198: Africa.

1774. Miiller, S., 44.

1789. Poiret, 358.

1804. Blumenbach, 66 (illus. no.).

1828. Boiy de St. Vincent (part), 73.

1833. Rozet, 232.

1835. Temminck and Schlegel (part), 70.

1860. Tristram, 405.

1867a. Steindachner, 4.

1872b. Gray, 4, in Sowerby and Lear, pi. xvi (but omit "Asia Minor").

1880c. Vaillant, 8, 33, 88, pis. xxv; xxvii, fig. 5.

1886. Fade, 316.

1893. Fade, 368.

1924a. Flower, 921.

1925. Flower, 929.

1926. Flower, 133.
1929. Flower, 28.

1932. ^Thomson, 362, pis. — .

2G2 BULLETIN : MUSEUM OF COMPABATIVE ZOOLOGY

1933. Flower, 744.

1934a. Mertens and Miiller in Rust, 9 .

1936b. Mertens, 15, figs. 2-3.

1937a. Flower, 6-9, 36.

1937. llediger, 187, 191.

1946(1. Mertens, 112.

1948. Astre, 60.

1949. Moyle, 582.

1950. Williams, 550.

1951. Aellen, 167, 195.

1952. Williams, 555, 556.
1954. Noel-IIume, 20, figs. 5-6.

1766. Testudo pusilla Linnaeus (part: not of 1758), 353.

1802. Shaw (not of Linnaeus: 1758), 53.

1862a. Strauch, 67.

18621). Strauch, 14.

1865. Strauch, 14.

1874. Boettger, 123.
1880c. Vaillant, 8, 26, 88.
1886. Vaillant, 135.

1890. Strauch, 45 (but omit Asiatic localities).

1898. Jeude, 2.

1835. Testudo Mauritanica Dumeril and Bibron (part), Erpet. Gen., 2,

p. 44: Mauritanica; Algeria (omit coasts of Caspian Sea, etc.).

1850. Guichenot, 1.

1854. Eversmann, 443.

1857. Jan, 35.

1875. Kereado, p. XXXV {Tistudo Mauritanica) .
1894. Oliver, 102.

1896 b. Oliver, 118.

1897. Bateman, 48.
1904. Chaignon, 2.
1903. Barbier, 78, pi. iii.
1907. LeCerf, 22.

1912. KoUnian, 103 footnote.

1919. Leblanc, 178, figs. 1, 5, 6.

1920. Mourgue, 233.

1836. Testudo inarginata Gervais (not of Schoepff), 309.
1841. Schlegel, 138.

1 848. Gervais, 204.

1884n. Rochebrune (erroneous report of oecun-ence in Seuegambia) 15.

1898. Jeude (erroneous listing of skeletal material from Cape) 2.
1836. Testudo ibera Gervais (not of Pallas), 309.

LOVERIDGE AND WILLIAMS: AI-RICAN CRVI'TODIRA

2()3

1848.
1851 1)4.
1851.
1877.
1 8,S8ii,.

i88r>h.

1887.

1889m.

1891c.

1892.

1892.

lS92a.

18921).

1893a.

1894.

1895.

1895c.

1896.

189<).

1 S9i».

190(1.

1901.

1901.

1903.

lyuGc.

1908.

1908.

1909a.

I909u.

1911.

1912a.

1912b.

1913.

19131).

1919.

1922h.

1922.

1923a.

1924a..

1924.

1926a.

1927n.

1928.

(k-ivais, 204.

Blanehard, 72, pis. i-ii (.skull, carapace, etc.).

Kichwald, 414.

rameiano, 97, pi. iv.

lioL'tlstT, 130.

Hoettger, 472.

Lortet (part), 3, pi. i.

Boulenger, 176.

Bouleuger, 96, 104.

Anderson, 11.

Koenig, 15.

Werner, 354.

Werner, 269.

Boettger, 11.

Werner, 75.

Koenig, 403.

Werner, 127.

Anderson, 111.

Escherich, 278.

Doumergue, 243.

Boettger and Tornier, 64.

Gadow, 369.

Lampe, 193.

Mayet, 10.

Siebenrock, 847.

Kerville, 96.

Zulueta, 451.

Siebenrock, 540.

Werner, 597, 628.

Lampe, 147.

Pellegrin, 256.

Werner, 448.

Ghigi, 266, 283.

Siebenrock (part), 24, figs. 10-19.

Leblanc, 177.

Procter, 493, figs, la, lib.

Zavattari, 15.

Calabresi, 7.

Flower, 920.

Zavattari, 15.

PeUegrin (1925), 315.

Pellegrin, 261.

Ilediger, 408.

264 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1929. Lindholm, 295.

1929b. Werner, 15, 21.

1929. Zavattari, 86.

1930a. Zavattari, 263.

1931e. Werner, 275.

1932. Thomson, 363, pis. — .

1935. Hediger, 3.

1935. Laurent, P., 345.

1937. Zavattari, 530.

1944b. Kehl, 166.

1948. Astre, 60.

1953. Burton, 34, fig., photo.

1836. Testudo Whitei Bennett, in White, Nat. Hist. Selborne, p. 361, fig. :

No locality given, but type in British Museum.
1841. Testudo graeca var. mauritaniea Schlegel, 106.
1857. tortue terrestre Labouysse, 86.
1870e. Peltastes marginatum var. whitei Gray, 11.
1873c. Chersinella graeoa Gray, 725, pi. Ix, fig. 4 (skull: African).
1880d. Testudo campanulata Peters (not of Strauch)!, 305.
1881a. Peters, 365.

1928, tortue d'Algerie Bailly-Maitre, 111.
1946d. Testudo graeca graeoa Mertens, 112.

Further citations of "graeca," "maurit aniens," "marginata,"
and "campanulata," will be found under Testudo kleinmanni.
Owing to the composite and frequently involved nature of many
of the early references to "graeca," we have preferred to omit
them unless they definitely mentioned a North African locality.

Synonymy. The name graeca (according to Statins Miiller:
1774, as quoted by Siebenrock :1913 :26) was given to this tortoise
by Linnaeus on account of its carapacial pattern resembling
Grecian mosaic work. It was entirely without reference to its
place of origin which Linnaeus quite definitely stated to be
Africa, after Edwards {circa 1743-8: Nat. Hist. Birds, part iv).
In this work the life-sized colored plate carries this caption :

1 Strauch (1862b, Mem. Acad. Imp. Sci. St.-P6tersbourg, (7), 4, No. 7, p. 14)
lists campanulata as occurring in Algeria though not seen by him. His descrip-
tion is of marginata. Straiicli attributes the authorship of campanulata to Wal-
baum (1782, Chelonogr. Beschr. Schildkroten, pp. 80, 124), but Walbaum's name
— ''Lorica varietatis tabulatae testudinis campanulata" — is not binomial and
consequently invalid ; furthermore, it apparently does not refer to an African
species.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 265

"No. 204. The African Land-Tortoise.
Loc. Santa Cruz in West Barbary."

i.e., the old Spanisli fort of Santa Cruz near Oran, Algeria {fide
Strauch :1862a) , and the reptile figured is quite identifiable.

Strauch (1862a :67) points out that graeca Linnaeus, 1758, and
piosilla Linnaeus, 1758, are not specifically the same, the latter
being a composite; but pusilla Linnaeus (in part), 1766, and
pusilla Shaw, 1802, were both based on Edward's specimen and
are consequently synonymous with graeca Linnaeus, 1758.

The first to point out that graeca was the correct name to apply
to the African species, was Siebenrock (1913, Ann. k.k. Naturhist.
Ilofmus. Wien., 196-197) but because ihera Pallas 1831 had long
been employed for this species, he preferred to continue use of
the latter name. Subsequently Flower (1925b; 929) pointed out
that the type of ihera came from Iberia in Transcaucasia (cf.
Pallas, 1811, Zoogeographia Kosso-Asiatica, 3, p. 18, pi. ii, figs.
2-3). Specimens (M.C.Z. 5301-2) from Tiflis, Transcaucasia,
suggest that this is a race of graeca, recognizable by its broad and
low-vaulted carapace. T. g. ihera has been considered by Mer-
tens (19-46d:112) as one of several races inhabiting Asia Minor,
their range separated from that of T. g. graeca by Egypt.

Tcstudo graeca hettai Lataste, based on an old specimen of un-
known origin, having a divided supracaudal and a caudal spur,
hut lacking a conical tubercle on the hinder side of the thigh,
appears to be a straight synonym of liermanni Gmelin, 1789, ex-
cept for its allegedly 4-clawed forefeet and 3-clawed hind feet.

Common names. Western Spur-thighed Tortoise (preferred) ;
Mosaic Tortoise (Miiller :1774) ; Moorish Tortoise (Bateman:
1897); Spur-thighed Mediterranean Land-Tortoise (Flower:
1925) ; Fakroun (Tunisian Arabic: Mosauer) ; makroona (Alge-
rian Arabic :Koenig) ; zolhafie (Arabic :Forskal).

Illustrations. The finest colored representations of this tortoise
are those to be found on plate xvi of Sowerby and Lear (1872b),
correctly labeled graeca; and on plate i of Lortet (1887) Avhere it
is miscalled ihera.

Description. Beak weakly hooked, tricuspid ; edge of jaws
not dentate; supranasals present, not meeting medially; pre-
frontal entire, rarely divided longitudinally;^ frontal large,

1 Occasionally divided longitudinally in "ihera" according to Boulenger (1889a).

2G(i

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

occasionally broken up ; remaining upper head shields small,
irregular ; forelimb anteriorly covered with large to very large,
unequal, juxtaposed, more or less imbricate, rounded (young)
to pointed (adult) scutes forming 3-G^ longitudinal and 4-7
transverse series from elbow to outer claw ; claws 5 f hinder side
of thigh with a very large fiat or subcorneal tubercle ; heel with or
without an indefinite spurlike tubercle; claws 4; tail without, or
with a poorly developed terminal clawlike tubercle.

Fig-. :20. ykiill of Testudu gratca yrutca (M.C.Z. 4485). Coiidylobasal
length 26 mm.

(S. McDowell del.)

Carapace convex, slightly depressed in young only, sides de-
scending abruptly, broadly notched in nuchal region, anterior
and posterior margins not expanded, not or but slightly reverted,
not or but slightly serrated ; dorsal shields concentrically .stri-
jited, not or but slightly swollen; nuchal moderately elongate,
often broadei" beliind, rarely divided longitudinally;"^ verte1>rals

J Thrt'p on left, four on ri^'ht of jin Alunrian spooiinrn i M.C.Z. 44S."i).

2 Flower remarks (l!(24:i)21) that of .'U tortoises of unknown iiarcnt;i^'i'
(though prpsuuiably of_Mefl! terra nean origin) hatched in (Jiza Zoolojiieal Uardeiis
between the years T>ii(-1!»2 . 12 had 4 elaws on tlieir f(M'efeet. 7 others bore ."i
elaws, and the ooiiiliiion in I he r(>niaining 12 was not note<l.

•"•Divided in a North African tortoise (Lnnii)e :19()1 ).

LOVERIDGE AND WILLIAMS: AFRICAN CRVI'TODIKA

2(i7

5, rarely 6,' the first as broad as or broader than long,- second
to fifth narrower or broader than the eostals; costals 4, rarel.y
7 or 8,^ not forming an angle with the marginals; marginals 11,
rarely 10; supracandal nndividod, rarely divided;-'' not project-
ing beyond the marginals, slightly incurved in 5 ,$ , occasionally
in $ ? .

Front lobe of plastron anteriorly truncate, not or but slightly

Fig. 21. Typical condition of prootic in dorsal view of skulls. A, LUoclu
lune p. />abfocL-(( A.M.N. H. 7203) showing prootic exposed; B, Testado y.
araeca (M.C.Z. 4485) showing prootic concealed.

l)roduei'd and not or but slightly notched; gulars paired; pec-
torals moderate, their anterior border usually curved and widen-
ing towards the axillary notch; axillary 1, small or moderate:
inguinals (apart from some accessory scales) 2, outer moderate.

1 Six in an abnormal livin? individual which had 7 costals on thp rjirht and s
on the Ifft (Mertens :l'j;i(ib :ligs. 1-2).

2 The third vertebral was always broader than or c(|iial to the tifth in yonn;:
(liza-bred tortoises, in older ones Ihe lliird was narrower than the fifth (Flower:
l'.r_>4a).

a Occasiiinnlly divided in voun- < i'uU' Kceniir :ls'.ii; i or adnlis i Donmerjrne :
1S99).

268 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

triangular, inner small, quadrate and in contact with the femo-
ral ; hind lobe broadly notched posteriorly, especially so in S $ ,
immovable in juveniles^ but in life more or less movable in adults
of both sexes.^

Plastral formula: Abd>(g. h. p. f. an: very variable).

Color. Carapace of young yellowish or pale olive, each shield
bordered with black and brown and usually bearing a central
black spot.

In adults yellowish, pale olive, brownish to reddish brown,
rarely uniform; the darker pattern as described for the young
but much more irregular with black predominating. Plastron
usually yellow or greenish yellow, occasionally light brown, most
or all of the shields bearing a blackish blotch. In 5 <J , accord-
ing to Doumergue (1899), the plastron has a tendency to be
more melanistic than in the 9 9 .

Size. Carapace length of a <^ (M.C.Z. 18161) from Bone
(which lived for over twenty years — 10.vii.l890 to 3.vi.l913 —
in the Loveridge garden), 145 mm., breadth 109 mm., height
73 mm.

Carapace length of a $ (M.C.Z. 1498) from Algeria, 192 mm.,
breadth 133 mm., height 100 mm., far surpassed by an Algerian
5 276 mm. (see below) mentioned by Flower (1945:452) who
points out that the largest examples of this species may not be
more than 35 years old, though attaining a much greater age.
Moreover, he says that very large individuals of this species
need not necessarily have come from the Balkan Peninsula or
Asia Minor, but may sometimes be of the typical North African
race.

Growth and Weight. The carapaces of two tortoises of un-
certain origin (purchased from a dealer in Norwich, England)
increased during a seven-year period from 7 to 9, and ly^ to
91/2 inches, respectively, when measured over the curve. Their
weight, recorded every six months from 1886-1892, showed a gain
of 11 and 12 oz., respectively. Occasional fluctuations in the
carefully tabulated figures were attributed to variations in the

1 Under 65 mm. according to Strauch (1862b :15) ; occasionally movable in
young (Koenig :1892).

2 Very slightly (Schlegel :1841) ; observable in the largest adults (Strauch:
1862b) ; sufficiently movable in $ J to facilitate enlarging the opening (Dou-
mergue :1899) ; movable in both sexes (Siebenrock :1906c).

LOVERIDGE AND WILLIAMS : AFRICAN CBYPTODIRA 269

climatic conditions, hot summers and cold winters being con-
sidered as the most favoi'able for growth by Sir Peter Eade
(1893) whose papers contain a wealth of detail.

Flower (1945:451) furnishes the annual weights and meas-
urements of two female Algerian graeca that were hatched in
September, 1905, from a batch of 14 eggs laid the previous May.
Of the 14 hatohlings, two were acquired by C. F. Moysey on
22.ii.06. The following day he weighed and measured them for
the first time and gave them the names of "Daimler" and "Pan-
hard." Their respective weights were then .008 (Daimler) and
.012 kgm. (Panhard), their carapacial lengths over the curve
47 and 49 mm. respectively. During the succeeding 39 years (in
Somersetshire and Devon gardens) a fairly regular rate of
growth (measurements were not taken from 1921 to 1937) was
maintained while, except on two occasions, each tortoise regis-
tered an increase in weight every year from 1906 to 1944. In
the latter year their respective weights were 1.814 and 3.969
kgm., their corresponding lengths over the curve 279 (11 inches)
and 365 mm. (14% inches). In a straight line the length of the
latter (Panhard) was 276 mm. (10% inches) on 22.V.44. Of its
weight Flower remarks that it was (almost) the same as that of
the heaviest graeca in the Giza Zoological Gardens, which he
recorded as 4 kilos (8 lbs. 13 oz.).

Following the exceptionally fine and dry summer of 1955,
which probably stimulated the tortoises' appetites, Daimler and
Panhard were weighed on September 1st for the fiftieth succes-
sive year of their captivity. At that time Daimler weighed 2.629
kgm., and measured 298 mm., having ceased growth in 1949 at
44 3^ears, while Panhard weighed 4.252 kgm., and measured 365
mm., having ceased growth in 1942 at 37 years. The weight of
the larger tortoise may well constitute a record for a North Afri-
can graeca. Certainly Mr. Moysey 's achievement in weighing
these animals monthly, except when hibernating, for half a cen-
tury is something for which all herpetologists should be grateful.
For the supplementary data contained in this paragraph we
personally are indebted to Mr. Moysey (in letter of 14.i.56).

Mr. Moysey suggests that the difference in size of his two
female tortoises is possibly attributable to the sharper-edged beak
of the larger. "With a single snap this reptile would sever a piece

270 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

of leaf, whereas the smaller specimen often made several abortive
attempts before succeeding in separating a piece for mastication
and swallowing.

Sexual dimorphism. Eighteen alleged secondary sexual char-
acters are discussed and illustrated by Camerano (1877 :pl, Iv).
Most, if not all, are highly questionable. Werner (1895c) states
that sex cannot be determined by plastron for he says it may be
concave, flat, or slightly convex in either sex.

Courtship. Burton (1953:34) quotes C. H. Lay as saying that
sometimes between mid-May and mid-June a thirty-year-old cap-
tive " ihera" male annually executes a series of rhythmic con-
volutions which leave a definite and regular pattern (cf. fig. in
Burton) on the bare soil over an area of about forty feet square.
The course taken is not so obvious in years when grass or weeds
cover the plot. If this is indeed a courtship performance, supple-
mentary to the oft-recorded butting described below, then further
information should be gathered Ijy persons resident in Africa.

Breeding. Possibly the fullest account of the breeding of a
Moroccan tortoise is furnished Ijy Gaston Astre (1948:60-64),
based on a young and an adult $ and an old $ that had lived in
his garden at Toulouse for twenty years. Owing, perhaps, to the
difference in age and size of the $ $ , no combats were observed
between them. However, each spring the adult $ assiduously
pursued the 5 . Then, rising on his four feet and withdrawing his
head, with the front of his carapace he would deliver a smart
blow- on the back of hers. This performance would be repeated
at the rate of one or two blows every two seconds. Should the 9
hurry away the $ Avould give chase and, passing, bring her to
a halt by biting at her head and forelimbs until she withdrew
them within her shell, or, more usually, he would seize her cara-
pace in his jaws and shake it with a range of as much as one or
two centimetres. During copulation, according to Bailly-Maitre
(1928), the male gapes widely and emits a muffled whistling cry.

From mid-May to mid- June the instincts of the 9 , rather than
necessity, caused her to excavate with her hind feet a number of
shallow troughs — about 34 mm. deep. These she soon aban-
doned, making no attempt to fill them in. Eventually, about the
10th to 15th of June, she would select a sunny spot with southern
aspect, most frequently at the foot of a wall where the soil was

LO\"F.RIDOK ANT) WTI.T.rA ^T^; : AFRICAN rRVT'TODIRA 271

sandy or loose and the ground likely to remain dry. There, usin^
her hind feet, she would dig a conical, somewhat bell-shaped, pit.
though its sides might be almost vertical if the earth happened
to be well packed, to a depth of 70 or 100 mm. and a surface
diameter of 120 to 340 mm.

On the l)ottom of this funnel she usually deposited 2, though
occasionally o, white, ellii)soidal eggs measuring 25 x ;^0 mm. and
weighing from \-i to 14 grams. Next she proceeded to cover them
to a depth of about 30 mm. wdth loose earth taken from that
heaped around the periphery of the excavation. The hole was
never entirely filled, for after the tortoise had smoothed over the
surface a slight concavity of from 5 to 10 mm. remained. Having
finished her task the tortoise would unconcernedly walk away.

On one occasion Astre noted the time expended on nest-making
and egg-laying. Digging commenced at 12 noon and was com-
pleted at 17.30 hours. Laying began at 17.30 and continued until
18 hours. Covering the eggs and filling in the hole with earth
took until 18.30, after which the tortoise made for the far end of
the garden, where she customarily spent the night beneath a
heap of dry vegetational debris, and went to sleep.

Though eggs were laid each summer for a period of twenty
years they never hatched owing to the Toulouse climate. Follow-
ing the unusually hot summer of 1947, however, when the average
temperature for July was 23.02° C, for August 23.81° C, with
a maximum of 42° on August 1st, and averaging 19.7°C. during
September, two eggs hatched. It may be assumed that the tem-
perature of the soil when directly exposed to the sun was pos-
sibly 20° higher than the shade temperatures cited above.

If the eggs benefited by the exceptionally warm summer, the
liatchlings were favored by the unusually mild winter that fol-
lowed. The average temperatures were 7.76°, 7.25° and 12° C.
respectively for the months of January, February and March,
1948. In April, when the young tortoises would be about 6
months old, they w^eighed 9 grams apiece and their carapaces
measured 30 mm. in length by 25 mm. in breadth.

In October, Bailly-]\Iaitre (1928) found a hatchling Algerian
tortoise in his Narbonne garden and, on examining the nest, a
freshly dead embryo in an egg. Astre mentions that in Septem-

272 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

ber, 1944, M. LeCompte discovered in his Gironde garden a small
tortoise that appeared to have been hatched there.

During the Tunisian rainy season in early spring, these tor-
toises are rarely in evidence unless the sun is powerful. When
this happens the reptiles begin to wander about in search of
each other (Chaignon:1904). On March 12th, in Algeria, tor-
toises were observed fighting and mating among the bushes (Dou-
mergue:1899).

In May, 3 ellipsoid eggs, each the size of a pigeon's, were pres-
ent in a Tunisian tortoise eaten by Chaignon (1904). An old $ ,
captured in the Macta (Mactra) region on 19. v. 1890, laid an egg
on June 25th ; on July 4th two more eggs were found, and laying
continued until 7 eggs had been deposited (over how long a
period is not stated). These eggs measured 28 x 36 mm. (Dou-
mergue :1899). In August an Algerian tortoise laid 2 eggs meas-
uring 24 X 33 mm. (Werner :1894), while others laid by a 202
mm. Tunisian reptile measured 24.5 to 31 x 31.5 to 38.5 mm.
(Gadeau de Kerville:1908). An egg in our possession (M.C.Z.
19003) measures 26 x 37 mm. The remarks by Kercado (1875)
are intentionally omitted as being based on untrustworthy hear-
say.

Longevity. The record is given by Flower (1925b and 1937a)
as being 102 or 125 years, depending on whether Archbishop
Laud's tortoise died in 1730 or 1753. This reptile, whose shell
was examined and identified by Flower at Lambeth Palace where
it is preserved, was at Fulham Palace in 1628 when Laud was
Bishop of London. He took it with him to Lambeth Palace when
he became Archbishop of Canterbury in 1633. Much supplemen-
tary information regarding this and other historical tortoises will
be found in the references given above.

Possibly more reliable records are those of 96 and 54 years
down to one of 22 years, 10 months and 23 days for a tortoise
(M.C.Z. 18161) already adult when brought back from Bone,
Algeria, on 10.vii.l890. The latter lived until 3.vii.l913 when
one of us (A.L.) chloroformed him because he was so weak and
moribund that flies w^ere buzzing around him and depositing their
eggs about his tail. Each winter during his life in a South Wales
garden, he had dug himself under the surface in a strawed-over
strawberry bed.

LOVERIDQE AND WILLIAMS: AFRICAN CRYPTODIRA 273

Diet. Pood is taken with a snapping movement or torn away,
being- swallowed with little or no mastication. When not fed,
Eade's two tortoises sought, not grass but clover, trefoil and
fleshy-leaved plants such as Echweria or Sedum. They displayed
a marked preference for yellow blossoms, consuming quantities
of buttercups, dandelions, French beans and lettuce. Both would
leave all other food for a green pea, a vegetable of which they
were so fond that they would follow anyone who would feed them
peas and clamber up his legs in their eagerness to reach them.
One would take sliced apple, the other not. While a neighbor's
tortoise fed on milk-soaked bread, Eade's two could not be
tempted with milk and never drank water (Eade:1886). Cap-
tive tortoises at St. Petersburg were fed cabbages and potatoes
(Eiehwald:1851).

Though essentially vegetarian, according to Doumergue (1899)
they will take insects, moUusks and worms during the Algerian
dry season ! Street vendors are apt to advertise these tortoises as
useful in ridding a house of cockroaches or a garden of slugs. As
a result many perish of starvation unless liberated in a garden
where they can help themselves to lettuce, and strawberries.

Parasites. Ticks (Hyalomnia aegyptiacum Linnaeus) from a
Rabat tortoise were identified as its synonym {Hyalomnia syria-
cum C. L. Koch) by Dr. R. Schweizer (Hediger :1935).

Enemies. Tortoise eggs are richer in albumen than those of
birds ; the flesh also is edible and tortoise broth is recommended
for the feeble (Doumergue :1899). The flesh is like that of a
chicken and many were eaten by the French soldiers, though not
by the Arabs and Moors (Rozet :1833), for the latter fear them,
imagining that they are evil spirits (Poiret:1789).

Formerly it was, and possibly still is a common sight in early
summer in the larger cities of Europe, especially maritime ones,
to see barrow loads of these helpless reptiles being offered for
sale by street vendors, or crawling aimlessly about in the window^s
of so-called pet shops.

Temperament. Of three purchased from a hawker on a Nor-
wich street by Eade (1886:316-322) one died, the others ap-
peared ill but recovered. They showed little evidence of hearing,
but their sight was acute and their sense of taste well developed,
enabling them to "discriminate instantly." As Lady Eade was

274 BULLETIN : MUSEUM OF COMrAKAl'l VK /DOLOdV

apt to indulf^e them with their favorite foods, upon her arrival
they would leave Sir Peter — even though he might be feeding
them at the time — and go to her to be fed. They ate only once
a day, if at all, during cool weather.

On the finest summer days, however, they were early astir and
by 6 or 7 A.^I. might be found nibbling at trefoil. Normally they
did not venture abroad until between 8 and 10 o'clock, and not
then if it happened to he raining or overcast. Should a cloud
obscure the sun when they were feeding, both would promptly
withdraw their heads. In thundery weather, even though there
might be hot and sunny intervals, the tortoises remained quiet
and indifferent to food. When the solar rays were insufficiently
hot tlie reptiles would tilt themselves against the south wall of
a greenhouse in order to enjoy the maximum warmth. When too
hot they would bury their heads beneath leaves while leaving the
(!arapace exposed.

They objected to being placed in a sleeping place other than
the one to which they customarily resorted. In cool weather,
however, Sir Peter habitually placed them under a mat in the
greenhouse ; next morning they would be at the door waiting to
be let out. When shut up in a greenhouse one tortoise was fre-
quently found mounted on the other's back or on an inverted
floM'erpot. Apparently both were c5 $ but they did not fight or
display any interest in eacli other's society; when one got in the
way its companion would attempt to clamber over it. Both
evinced a love of climbing; when one tried to ascend some sticks
that were resting against a wall, it slipped and was found sprawl-
ing on its back, three legs waving helplessly in the air while only
one hind limb rested on the ground.

If carried from some spot they liked, they would make straight
for it again as soon as released. When timed it was found that
one took a minute to walk 10 feet, while the other could do 20
feet in this period ; at this rate they would take between 4 and 5
hours to cover a mile. Their memories appear good, for immedi-
ately after emerging from a hibernation of 7 to 8 months' dura-
tion, they would set off for their old haunts as if only a day had
intervened.

Aestivatio7L and Hibernation. Both aestivation and hibernation
ai-e intermittent in Xorth Africa. During the Algerian summer

I.OVKKIDC.K AND WILLIAMS: AFRICAN CRYI'TODIKA Z/O

these tortoises uoniiully seek slielter I'l-om tlie sun b\- burying
themselves, and eonsecjuently are rarely encountered (Dou-
niergue :189f)), l)ut in July and August two were collected in the
early morning by Ilediger (1928).

They reappear in September only to vanish again at the ap-
proach of winter by concealing themselves in burrows or beneath
a large stone. Even in tiie depth of winter, however, when the
ground has been sufficiently heated by the sun, individual tor-
toises emerge for a time. In the hot coastal dunes and hills they
come out again in February (Doumergue :1899).

Habitat. In Algeria these tortoises are extremely connnoii
througiiout the region from Oran to the Petit Atlas (Rozet :1833),
where they occur among the dwarf palms {Chamacrops hiimilis)
beneath whose fronds they conceal themselves (Schlegel:18-41) .
They are numerous among the bushes {Zizyphvs jujuha) that
are scattered throughout the waste lands (Eversmann :1854),
where the.v are more plentiful among the dunes than on the
edges of the plains (Le Cerf :1907). Scarce, though widely dis-
tributed throughout the mountainous regions and all the high
plateaus, this species is apparently absent from the oases, its most
southerly records being Beguira ; El Aricha, and Geryville
(Doumergue :1 899). Absent south of El Kantara (Kantarah),
which is also the most southerly limit for the lizard genera La-
certa, Ophisops {Ophi<)p.s) and Fsammodromus (Werner :1894).

Common on the plains of Tunisia (Olivier :1896b).

Localities. Spanish Morocco: Tangier (Tanger). French
Morocco: Azemmour (i^biimor) ; Azrou ; *Beni Snassene ; Ber-
guent; Casablanca; Bar el Beida, Casablanca; El Marit, Casa-
blanca ; Pes ( Fez ) ; Karia ba Mohammed ; Maarif near
Casablanca; *Mamora (Marmorea) Forest; Mogador; Mogh-
ran, a confluent of Sebou west of Beht; Oued Akrech, south
of Rabat; Oued Mella north of Mogador; Ouezzane; Rabat
(Am el Anda and La California); *Sefrou; Sidi Amar;
*Sidi Yahia (U.S.N.M.) ; Souk el Arba; Taourirt. Algeria:
Aumale. Guyotville ; Bainen Poorest near Algiers, Batna ;
Beguira ; Biskra ; *Bone ; Boudsareah near Algiers ; Bou Saada ;
Constantine ; El Aricha ; Geryville ; La Calle ; Lambeze
(Lambesa); Macta; Maison Caree near Algiers; Medea (Me-
deah) ; Metidja near Algiers; Nemours; Palestro; on mountains

276 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

between Rosmarin and Thymian; Santa Cruz=Oran. Tunisia:
A'in Draliam; Bir Mcherga ; Djebel Aehkel ( ? Isehkeul) ; Djebel
Rsass (sic); Douirat (Duirat) ; Graiba; Kairouan (Kairwan)
Maxula Rades; Sidi bou Ali; Tunis; Utique ruins; Zarzis. Lihya
Barce (= El Merg) ; Bengasi; Bir (Bil) Milrha; Marmarica
Merg (= Barce) ; Tobruk (Tobruch) ; Tolmeta (Tolmetta)
Tripoli.

Range. Morocco to Cyrenaica ; Pityusen Islands and southern
Spain.

PSETJDOTESTUDO new subgenus
Type species : Testudo kleinmanni Lortet

Definition. Supracaudal usually single. Maxillary without
ridges ; anterior palatine foramina large, not at all concealed ;
quadrate not enclosing stapes. Plastron of adults with a clearly
movable posterior lobe in both sexes. Tail claw absent.

Range. Northeast Africa.

Testudo kleinmanni Lortet

1823. Testudo graeoa Lichtenstein (not of Ltanaeus), 91.

1835. Riippell, 4.

1880d. Peters, 305.

1881a. Peters, 365.

1835. Testudo marginata Dumeril and Bibron (part: not of Schoepff), 37.

1844. Gray, 9.

1855. Gray, 11.

1857. Jan, 35.

1855. fChersus mauritanicus Fitzinger (not of Dumeril and Bibron), 252

(Egypt).
1869a. Testudo leitJiii Giinther, Proc. Zool. Soc. London, p. 502, figs. 1-4:

"Sindh" (error) ; preoccupied by T. leithii Garter, 1852.
1869b. Giinther, 110.

1880c. Vaillant, 32, 88.

1889a. Boulenger, 175.

1890. Boulenger, Fauna British India. Kept, and Batr., p. 20.

1890. Strauch, 45.

1892. Baur, 156.

1893a. Boettger, 11.

1896. Anderson, 77, 95.

LOVERIDGB AND WILLIAMS: AFRICAN CEYPTODIRA 277

1898. Anderson, 28, col. pi. ii.

1901. Steindaehncr, 326.

1904. Andersson, 9.

1908b. Kammerer, 757.

1908c. Kammerer, 772, figs. 9-10.

1909a. Siebenrock, 540.

1909a. Werner, 596, 628.

1912b. Werner, 452.

1913. Ghigi, 283.

1923a. Calabresi, 7.

1924a. Flower, 920.

1925b. Flower, 928.

1926. Flower, 133.

1927. Vinciguerra, 328.
1929. Flower, 29.
1929. Zavattari, 86.
1930a. Zavattari, 263,

1931. Gestro and Vinciguerra, 538.

1933. Flower, 745, fig. la.

1934a. Mertens and Miiller in Eust, 9.

1937. Zavattari, 530.

1950. Williams, 551.

1870c. Peltastes leithii Gray, 657.

1870e. Gray, 11, fig. 6.

1873b. Gray, 11.

1883. Testudo Kleinmanni Lortet, Arch. Mus. Hist. Nat. Lyon, 3, p. 188:

Lower Egypt and environs of Alexandria, Egypt.
1887. Lortet, 11, pi. v.

1894. Testudo campanulata d'Aubusson (not of Strauch), for 1893, 230.

8yno7iyniy. Mons. Kleinmann, after whom the species is
named, was Director of the Credit lyonnais en Egy^te (Lortet:
1887 :13). Testudo leithii, named for Dr. A. H. Leith by Giinther
(1869a), is preoccupied by Testudo leithii Carter 1852 proposed
for a fossil turtle from Worli Hill, Bombay, India, which Wil-
liams (1953) ^ has shown to be that of a pelomedusid and assigned
to a new genus — Carteremys.

Common names. Egyptian Tortoise (suggested) ; Leith 's Tor-
toise (Flower) ; solhalfa or zihlifa (Innes in Anderson 1898).

Illustrations. Exceptionally fine colored figures of this species
(dorsal, lateral and ventral views) are given by Lortet (1887 :pl.
v) and Anderson (1898 :pl: ii).

1 Williams, E. E., 1953, Breviora (Mus. Comp. Zool.), No 13, p. 5.

278

BULLETIN : MUSEUJVJ OF COMPARATIVE ZOOLOGY

Description. Beak feebly or moderately hooked, tricuspid;
edge of jaws not dentate ; snpranasals present, sometimes meet-
ing mesially ; prefrontal entire, divided longitudinally, or broken
up ; frontal large or broken up ; remaining upper head shields
small, irregular ; forelimb anteriorly covered with a few, contig-
uous, extremely' large, imbricate, rounded or pointed scutes
forming 3, rarel}' 4, longitudinal and 4-7 transverse series from
elbow to outer claw ; claws 5, rarely 4 ;^ hinder side of thigli
without enlarged tubercles; heel with large, conical, spurlike
tubercle ; claws 4 ; tail with or without a terminal clawlike
tubercle.

Carapace convex, sliglitly depressed in young only, sides de-
scending abruptly, deeplj" or broadly notched in nuchal region,
anterior and posterior margins not expanded, not, or but slightly,
serrated ; dorsal shields concentrically striated, not or but slightly
swollen ; nuchal moderate or large, triangular, rectangular or ex-

Fig. 22. Head sealation in Tcatudo kleinmanni (x 1). A, Saluiu sped
men (M.C.Z. 54045); B, Saluni specimen (M.C.Z. 54044); C, Alexandria
specimen (M.C.Z. 5081), Cotype.

ceptionally elongate ;- vertebrals 5, rarely G or 7,^ the first as
In-oad as, or much broader than, long, smaller than the second to
tlfth whicli are narrower than, as broad as, or broader than, the
i-ostals;"' costals 4, rarely 3 or 5,^' not forming an angle with the
marginals; marginals 11 or 12, rarely 10;*^ supracaudal usually
undivided, less frequently divided, spread out like, but project-
ing angularly ])eyond, the marginals.

1 Four in a tortoise bred at Giza Zoo from El Arish parents, both of whom had
four claws only on each forefoot (Flower :19oo :747).

2 In a ^ from Kl Arish in which it was 6 mm. x '2..j broad mm. (Flower :19o?) :

747).

3 Six in a hatchling and also in a 95 mm. ^ from El Arish (Flower :]933 :747) ;

seven in a cotype lU.S.X.M.. 10979) seen by lO.K.W.

* In a 95 mm. Derna tortoise the third verteliral has given rise to a couple of
su]>plementary shields lying l>etween the ord and 4th cost.als (Calabresi: 192 'a i.

■''Three in one seen by Flower (1953:748): 5 on right side of a carapace
(Y.I>.M. t>7l.') seen bv us.

« Ten on one side of a cotyjie (U.S.X.M. 10979) and a specimen (B.M.
l!t22.o.2.'!.2), both seen by E.E.W. : 12 in a 95 mm. ^ from El Arish. while others
fioni tliis locality liad 11 on one side and 12 on the other (P'lower -AW.y.i :747).

I.OVKRIDGI': AM) WILLIAMS: AIKIC'A.V fRVrTOniRA

279

Front lobe of plastron anteriorly truiieate, not or bnt slif,'htly
produced, s(mi('\vliat notelied and eaeli lateral point itself very
slightly toothed; <iulars paired, rarely absent;^ IJeetorals moder-
ate, their anterior border usually widening towards the axillary
notch; axillary 1, moderate; inguinals 1 or 2, small oi- moderate,
in contaet with, or separated from, the femoral; hind lobe very
broadly notched posteriorly, sliglitly movable in adults of both
sexes but especially in the $ .

Fig. 23. Skull of Testudo Jdeinmanni (Yale Mus. 667). Coudylobasal
length 21 mm. The entrance of the parietal into the orbit is an individual
peculiarity.

(S. McDowell del.)

Flastral formula : Abd> (g, h, p, an, subequal) >f.

Color. Carapace greenish yellow to pale yellow, uniform, or
each shield more or less edged with black or brown. Plastron
greenish to j'ellow, each abdominal shield with a dark brown
to black, triangular or cuneiform patch, w^hieh is usually con-
spicuous and never entirely absent ; pectorals occasionally mar-
gined anteriorly with dark brown ; remaining plastral shields
uniform. Hatchlings are completely yellow, according to Lortet
(1887).

1 Absent in a Giza tortoise (U.S.N.M. 5.5758) in which the anterior lobe is ab-
normally sbortonpd thoiieh apparently not the result of an Injury.

280

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Head, neck, limbs, feet, nails and tail, yellow ; rarely a black
spot on crown of head posterior to the eyes; the buttonlike eyes
strikingly black, the irides in both sexes being shining black and
very conspicuous according to Flower (1933:748) who should
be consulted for further details on coloration. His paper, based
on 159 living tortoises, contains far more information about
"leithii" than all of the other citations combined.

A

B

Fig. 24. Testudo Meinmanni. A, Dorsal view of carapace (M.C.Z. 1499)
(x %); B, Internal view of plastron (Yale Mus. 667). Note hinging of
the posterior lobe (x %).

(P. Washer del.)

Size. Carapace length of adult 5 (5 is from 86 to 110 mm. ;
the last mentioned, coming from El Arish, is exceptional for
a $ . Carapace length of adult 5 ? is from 113 to 127 mm.,
the latter from Mersa Matruh (cf. Flower :1933 :746). Carapace
length of a $ (M.C.Z. 1499) 100 mm., breadth 72 mm., height
57 mm. Carapace length of a kleinmanni cotype, 100 mm., of
the leithii holotype 120 mm. ; of a juvenile kleinmanni cotype
(M.C.Z. 5081) 55 mm., breadth 45 mm., height 28 mm.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 281

Weight. Adults average 7 to 9 oz. (.198 to .255 kgm.), the
largest attaining 13 or 14 oz. (.35 to .4 kgm.), fide Flower
(1933:746).

Sexual dimorphism. Tail of $ short and pointed, that of
the S very long, yet exceeded by the fully extended penis which,
though normally pink, then turns to a rich purple (Flower:
1933:746).

Fig, 25. Testudo hermanni 3 (M.C.Z.). Internal view of plastron to
show absence of hinging.

(P. Washer del.)

Breeding. Mating at the Giza Zoological Gardens, Cairo, took
place in September and October (Flower). March and April
among captive tortoises (Lortet:1883), while a $ that Kam-
merer (1908c) kept for five years, repeatedlj^ copulated with
captive hermanni (miscalled graeca) but, possibly because thb
2 $ were subadult, they remained infertile. Flower describes*
the voice of the mating $ as sounding like the intermittent
winding-up of a metal spring.

282 BULLETIN : MUSEUM OF rOMl>ARATIVE ZOOLOGY

Eggs with a rosy blush and measuring 23 x 30 mm., were
deposited in May and abandoned on the surface, according to
Lortet (1887), who adds tliat when the temperature is sufficiently
high the young may emerge in 20 days after laying. Suck
hatchlings were completely yellow.

On two occasions eggs were laid at Giza in June, one laid on
the 17th measured 22.75 x 29 mm. No records were kept as to
when the majority of the eggs were laid, but 69 hatchings took
place as follows: September (6) ; October (37) ; November (17) ;
January (3) ; February (6).

Kammerer (1908b) somewhat vaguely refers to a dozen eggs
measuring 22 x 28 mm., some of which were laid on July 6 by a
tortoise having a carapace length of 104 mm. After 125 days^ all
allegedly hatched on November 8 but, refusing food, died within
a few weeks of emergence. The dimensions of these hatchlings,
stated to be from 18 to 19 mm., are at variance with those fur-
nished by Flower (1933:748) for one which Avas 33 mm. in
carapace length and 28 mm. in breadth.

The outline of Jdchrmanni hatchlings is quite different from
that of the more spherical graeca. Contrasting figures to illus-
trate this are furnished by Flower (1933:748).

Longevity. Twenty-one years and 25 days. Average longevity
of twenty tortoises still living in the Giza Zoo in 1925, was IG
years, 7 months, and 6 d-dys (FloAver :1925b).

Diet. Largely Saltwort {Salsola sp.) and Sea Lavender {Li-
nionium sp.) according to Lortet (1887:14).

Hiheniatioit. Active throughout the Egyptian winter though
more sensitive to low temperatures than either graeca or margi-
nata, according to Lortet (1887:14).

Habitat. In Egypt it is very restricted, being absent from the
deserts around Cairo, and ]) resent only in two districts which
are about 180 miles apart, viz. the northwestern deserts of the
Nile delta and region of Suez Canal where, as well as in northern
Sinai, it may be found among the bushes growing in the vicinity
of brackish lagoons.

Localities. Lihya: Barca; between Bir Sceferzen and Esc-
Seegga on the Giarabub to Tobruk road; Sirtica; Wadi Tessina
in Kufra Oasis. Egypt: *Alexandria; Bir el Abd ; Damietta; El
Arish (Arise) ; *Giza (Gizeh) ; Ismailia ; Kantara (Kantareh) ;

1 Coiiipiire with Lortet above I

LOVERIDGK AND WIT-I.IAMS: AFRIfAX CRYPTODIRA 283

Khabra Abu (Juzoar; Katia ; Lahfan ; Mariiit (Maryut) ; Mersa
Matruh; Port Said; Komani ; Salmaiia; *Salum ; Wadi el Amr;
Wadi Hareidhin.

Range. WestiMii Cyrenaica and northern P^<j:ypt (including
northern Sinai ).^

Genus MaLACOCHEESUS Lindholni

1929. Malacochersm Lindholni, Zool. Anz., 81. p. 285. Type: Testudo
lar7iicrl Siebonroc-k (by original designation).

Dcjinitiun. Skull with triturating surface of maxilla strongly
ridged; median premaxillary ridge absent; maxillary not enter-
ing roof of palate ; anterior palatine foramina small, concealed
in ventral vieAv ; prootic well exposed dorsally ; (juadrate enclos-
ing stapes; surangular sometimes markedly lower than pre-
articular, exposing the latter to view externally; neck with third
or fourth centrum biconvex.

Carapace never hinged, 'persistently fenestrated; neurals at-
tenuated and greatly reduced; outer side of third costal scute
about as long as, or longer than, that of fourth; submarginal
.scute absent; a single suprapygal.

Plastron never hinged, persistently fenestrated ; gular region
neither thickened nor produced; gulars paired, broader than
long.

Distinguislmig marhs. No other tortoise is so nearly flat or
has a shell so flexible that it may be compressed between finger
and thumb. Carapace length not known to exceed 177 mm.

Range. Apparently restricted to certain rocky regions of
Kenya and Tanganyika.

Fossil record. None.

Remarks. The relationships of Malacochersus are far from
clear. At first glance the shell seems anomalous and unique, yet
its fenestration, as has l)een made clear by Miss Procter (1922b),
is similar to that of some adult Gopherus polyphenius and to the
hatchling stages of every tortoise (ef. Procter: fig. 21b of a
young Testudo horsfieldi and the figures in Gray: 1873e). In
Malacochersus we find persisting into adult life a stage in shell

1 Tristram's sratpmeiit (1SS4. The Snrvpy of Western Palestine. The Fauna
and Flora of Palestine, l."!*!) that kIriniiKiiiiti ranges northwards in snitaide lo-
ealiti<'s lo the western side of the P.laok Sea (in the vicini'y of_TIehronl. Pales-
tine, was based on a niisidentifieation .is shown by Flower (IHo:'. :74.')).

284 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

ontogeny usually present only for a short time after hatching.
At least this is true of the dermal shell. There is, however, some
resorption of the endochondral ribs, but this also is paralleled
in other land tortoises, in which reduction of the ribs is a very
common condition. Apart from fenestration, the shell is rather
primitive, i.e. essentially emydine, in its single suprapygal, the
hardly developed gular lip and the broad transversely extended
gular scutes.

However, skull, vertebrae and limbs are thoroughly testudinine
and differ but little from those of Geochelone. Limbs and verte-
brae are in fact in no way distinctive, but the skull has certain
discernible but rather subtle peculiarities, and the mandible is
more conspicuously distinct in a dorsal emargination of the
surangular. This is not consistently present and while it is
tempting to ascribe such an emargination to a general weakness
of ossification in this form, such a suggestion is not borne out
by the very solidly built skull in which even the anterior palatine
foramina are not at all enlarged.

In external features M. tornieri is somewhat peculiar. The
transversely, rather than longitudinally, developed gulars recall
Homopus, while the head scutellation with its large supranasals
along with the prefrontals and frontal, is more elaborate, so
possibly more primitive, than that in any other form except
Testudo. Miss Procter would call an azygous scute, occasionally
occurring on the anterior plastron, an intergular, comparable to
that of primitive turtles ; it may be doubted that these anomalies
have any atavistic or phylogenetic significance. The flatness of
the shell is extraordinary and, as Dr. Procter comments, is un-
related to the fenestration but appears to be a special develop-
ment correlated with the reptile's habits. Miss Procter believed
the feet to be more supple at wrists and ankles and less club-
shaped than in most testudinines.

Because of the nature of the shell the paleontology of this form
is unlikely to become known soon. The best hope would appear
to be a skull, since that is rather solidly built, but even this
would have to be well preserved, and, as is well known, well-
preserved turtle skulls are paleontological rarities.

In the preceding diagram of relationships (see p. 210) we have
tentatively suggested Malacochersus as arising quite independ-

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA

285

ently from the ancestral stock of the testudinines. It would seem
that the characters possessed by Malacochcrsus, other than
specializations peculiar to itself, are primitive for the subfamily.
This opinion, while the best we can offer, is a guess, but on the
basis of the data available no other conclusion seems likelv.

P

Fig. 26. Distribution of Malacochersus tornieri in East Africa. The liol-
low circle indicates the type locality of procterae. (P. Washer del.)

The distribution of this peculiar genus, dependent on a
rocky habitat, is not merely limited but discontinuous if judged
by present records. The presence of such a shy and cryptozoic

286 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

creature may well have been overlooked, hovt^ever, and some of
the gaps in its range are certain to be closed when intervening
areas are carefully searched.

The southernmost record, Lindi, whence came the second
known specimen, is on the southeast coast of Tanganyika Terri-
tory almost 400 miles southeast of Dodoma, which appears to be
the chief center of distribution. From Dodoma the species ex-
tends west to Tabora and north to Busisi (the original type
locality of tornieri) at the southern end of Lake Victoria, and
east to Lake Eyasi which is due north of Dodoma.

The most northerly record for which a specimen is available,
Xjoro, Kenya Colony, is approximately 200 miles due north of
Lake Eyasi. Another record in Kenya is Malindi about 360
miles southeast of Njoro on the Kenya coast and as far from
Dodoma as it is from Njoro. But Malacochersus is to be looked
for still further north, for in 1914 the late R. B. Woosnam,
when asked by Loveridge whether undescribed reptiles were
still to be found in Kenya, replied that he had once come across
two strange soft-shelled land-tortoises when on safari in the
Northern Frontier District. Being on the march at the time,
and having nowhere else to put them, he had popped them into
a cooking pot being carried by one of the bearers. Later, on
enquiring for them, he found they had been turned loose by
his indignant cook. AVoosnam added that he had never seen any
similar tortoises during the many years he had spent in Africa.

Malacochersus tornieri (Siebenrock)

1896. Cinicys (sic) helliana Tornier (part: not of Gray), 2.

1903. Testudo tornieri Siebenrock, Anz. Akad. Wiss. Wien, 40, p. 18-5:

Busisi, s. end of Lake Victoria, Tanganyika Territory.
1903b. Siebenrock, 442, pi.

1904a. Siebenrock, 29, fig.

1909a. Siebenrock, 538.

1911. Lonnberg, 7.

1913c. Nieden, 54.

1922d. Loveridge, 522.

1923g. Loveridge, 931.

1924b. Loveridge, 2.

1924. Wettstein, 201, figs. 1-2.

1928d. Loveridge, 49.

1929. Flower, 32. .^^^

LOVERIDGE AND WILLIAMS: AFRICAN CRVl'TODIKA

•>S7

1933h. Loveridge, 207.

1936. Kanberg, 187, photo.
]93r)h. Loveridge, 19.
1936.i. Loveridge, 221.
1937f. Loveridge, 492, 49.").

1937. Ruckes, 103.

1940. Conant and Downes, 48.

1950. Williams, 550, 551.

1954. Noiil-Huine, 76 (numerous niisstaUMiu'utsX

1920i-. Tc.studo Loveridgii Boulengor, C. R. Acad. Sci. (I'misi. 17ii, p. 2C..T :

Dodoma, Ugogo, Tanganyika Territory.
1920d. Boulenger, 68.

1920. Boulenger, E. G., 190.

1921(1. Loveridge, 50.

19221). Procter, 483, figs, lb, 2-lla, 12-21a, pis. i iii.

1922d. Loveridge, 522.

1 92:5g. Loveridge, 928.

19241.. Loveridge, 2.

1924b. Martens, 71.

1924. Wettstein, 201, figs. 1-2.

1924(1. Witte, 48, 61.

19251). Flower, 925.

19221). Cinixys beUiana Procter (not of Gray), 515.
1923g. Test lido procterae Loveridgi\ Proc. Zool. Soe. London, p. 928, pis.

iii: Ikikuyu, s. of Gulwe, Tanganyika Territory.
ly24lj. Loveridge, 2.

1924b. Mertens, 72.

1924d. Witte, 48, 62.

1937f. Loveridge, 495.

1929. Testudo (Malacochemus) tornieri Lindholm, 285.
1934a. Malacochersus inniieri .Mertins niid JNliiller in Rut;!, 9.
1937d. Mertens, 3.

1939. Zangerl, 355, 399, pi. ii.

1942d. Mertens, 245-251, figs. 1-4.

1949. Conant and Hudson, 4.

1949b. Loveridge, 19.

1934a. Malacochersus procterae Mertens and Miiller in Rust, 9.

Further citations of "tornieri" will be found under Kinixys
h. helUana.

Common natnes. Soft-shelled Tortoise (Flower) ; Pancake
Tortoise (W. M. Mann).

Illustrations. Siebenrock (1903 :pi.-) furnishes a dorsal view
of the type of tornieri, together with detailed drawings of the

288

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

beak and scaling on the forelimb. Procter (1922 :pl. i) has a
colored plate of topotypieal loveridgii, which should be con-
trasted with that of procterae as shown by Loveridge (1923g:
pi. i). Siebenrock (1904a :fig. -) pictures the skeletal carapace
from below ; x-rays of $ and $ , besides numerous develop-
mental figures, will be found in Procter (1922b), Loveridge
(1923g:pl. ii), and Zangerl (1939 :pl. ii).

Types. The Berlin type, collected by Stuhlmann, was at first
referred to Kinixys by Tornier under the impression that the
paperlike consistency of its carapace was abnormal, perhaps a
case of arrested development. Later Siebenrock (1903a), discov-

Fig. 27. Skull of Malacochersus tornieri (A.M.N.H. 45081). Condylo-
basal length 30 mm. The lateral exposure of the prearticular is not constant.

(S. McDowell del.)

ering this Berlin specimen was not a Kinixys but an undescribed
species of Testudo, named it tornieri, though still attributing its
peculiar softness and flatness to pathological defects which had
retarded the development of the bony carapace and plastron;
alternatively he postulated faulty preservation. The receipt of
a second specimen from Lindi, enabled him (1904a :29) to am-
plify his original description and give a detailed account of the
fenestration of the shell.

When five specimens collected by one of us (A.L.) and labeled
"Testudo tornieri" in the field, were submitted to Boulenger, he

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 289

regarded them as separable and distinct. Included in the ship-
ment were other tortoises including a young Kinixys helliana
from another locality which Boulenger (1920c) unfortunately
assumed was the young of his loveridgn.

The disposition of Testudo procterae Loveridge, poses some-
thing of a problem. Its type locality — Ikikuyu (not to be con-
fused with the place of the same name on the outskirts of Do-
doma) south of Guhve (Igulwe) Station on the Central Rail-
way— is about 75 miles southeast of Dodoma, type locality of
loveridgii Boulenger (=tornieri).

The supposed structural differences offer but weak support
(cf. comments under Size) to the idea that the very distinctively
colored (cf. Loveridge :1923g :col. pi. i.), but still unique, type of
procterae in the British Museum is a distinct species. In the
absence of more material, therefore, we incline to the view that
procterae may be only a very striking color variant of tornieri.
This view, if correct, is paralleled by the somewhat similar case
of "Homopus" iergeri Lindholm, wdiich we regard as merely
a color variety of Psammohates tentorius verroxii (Smith). In
both instances pattern reduction has taken place; in hergeri
there has been a melanistic simplification ; in procterae brown
pigmentation has become dominant to the exclusion of all other
markings.

How^ever, in its elongate and large prefrontals with correla-
tively absent supranasals (cf. Plate 6), the head scalation of
procterae differs strikingly from, and is not approached by, any
of the many tornieri we have examined (cf. characteristic scutel-
lation of tornieri in Fig. 19) . Symmetrical as is the arrangement
in the holotype of procterae there remains the possibility that
it is an individual variation. Variation in head scalation, com-
mon though it is among tortoises, usually consists in a breakdown
of the normal arrangement, not in the appearance of an alterna-
tive pattern. In this instance, with only a single specimen avail-
able, it is impossible to evaluate the importance of this character.
It emphasizes the importance of securing further Malacochersus
from the type locality of procterae; we have attempted to have
such material obtained for us, but the attempt was unsuccessful.

Description. Beak weakly or moderately hooked, bi- or tri-
cuspid, edge of jaws minutely denticulate ; a pair of supranasals

290 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

in contact, very rarely separated; prefrontal entire or divided
longitudinally or some-\vhat broken up ; frontal larpe, small, or
broken up; remaining upj)er head shields small, irregular; fore-
limb along anterior edge with some moderately enlarged, more
or less imbricating scutes^ forming about 5-10- transverse series
from elbow to outer claw ; on inner aspect of forelimb are 2-4
prominent, enlarged, separated or juxtaposed, pointed scutes;
claws 5, occasionally 4 through accident ; hinder side of thigh
with a rosette of enlarged scales surrounding a somewhat larger
subconical tubercle ; heel sometimes with trace of a spurlike
tubercle; claws 4; tail without terminal clawlike tubercle.

Carapace very much flattened dorsally, flexible, its sides
either straight and parallel or obli(iue. perpendicular or steeply
sloping, scarcely notched in nucha! region, anterior margin flat
and weakly serrate, lateral margins sometimes reverted, gutter-
like, posterior margin flat, rarely reverted, slightly serrated;
dorsal shields concentrically striated, sometimes swollen;^ nuchal
somewhat broader than long in young, usually elongate in adults,
anteriorly indentate and projecting, usually broader posteriorly,
occasionally completely divided;'* vertebrals 5 rarely 4,'' 6,^ or
1,'' sometimes flat, very rarely divided longitudinally'* or trans-
versely,^ the contact between fourth and fifth frequently very
narrow% first vertebral slightly broader than long or as broad as
long and smaller than the second to fifth, which are much broader
than long, as broad as, or broader than, the costals ; costals 4,
rarely 5,^'^' sometimes forming an angle with the lateral marginals;
marginals 11, sometimes 12; supracaudal divided, sometimes
above only, rarely undivided, downwardly directed or outwardly
flared irrespective of sex.

1 Thosf enlargerl scutes -avc bntli mrrt> niuiieruiis ami .strongly iinbriratiii.;; in
the Xjoro tortoise (Stockholm Mus.).

2 Twelve on both sides of tlii> Njoro specimen (Stocklmlm Miis.).

3 Swollen in onl.v one Doiloma individual (M.C.Z. 24!)1'J).

* Divided in a 9 (M.C.Z. :j(M»09) from Mangasini, and in U.S.X.M. 72505, prob-
ably from Dodoma.

5 Four in a ^ cotype of luriridyii (cf. Procter :1922b :flj^. 4).

6 Six in a $ (M.C.Z. 2:3000) from Kondoa Irangi.

" Seven ir. I'.S.N.M. 7o9;54. proliably from Dodoma.

8 Fourth vertebral is longitudinally divided in U.S.X.M. 72497, proiiably from
Dodoma.

9 Fifth vertebral is transversely divided in U.S.X.M. 7::i94.3, probably from
Dodoma.

10 Five in U.S.X.M. 72522, 739:;4. etc.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIKA 291

Front lobe ol' plasiruii anteriorly truncate, more or less pro-
duced, openly notched ; gulars paired, as wide as, or wider than,
long; exceptionally an intergular ;^ pectorals moderately broad,
not or but slightly narrowed medially, their anterior border
usually straight, widening gradually toAvards the axillary notch ;
axillaries 2, rarely 3, small ; inguinals 2 to 4, outermost normally
triangular, the others transverse, the innermost in contact with
the femoral ; hind lobe more or less deeply notched posteriorly.

Plastral formula: Abd>h>p>or<f>or=g>or=an.2

Color of a hatchling (M.C.Z. 18167). Ground color of carapace
pale yellow; sutures between dorsals irregularly, but broadly,
bordered with dull brown though frequently interrupted at one
or more points on each shield ; vertebrals 2 to 4 have a median
brown spot, while the first and fifth vertebrals and each of the
eostaJs have a fainter, less regular, brown spot; dorsal edge of
marginals with a less well-defined brown border tending to ex-
tend downwards between each marginal, free edge of marginals
without markings. Plastron pale yellow, immaculate.

Color of juveniles. No two specimens are quite alike. Ground
color of carapace pale yellow ; brown borders covering the
sutures become black, and both these and the inner brown
borders tend to be interrupted to a varying extent by yelloAv
rays, the brown spots on the vertebrals and costals may, or may
not, persist. Plastron pale yellow, all the sutures, except those
between the gulars and anals, dark brown crossed by light rays,
while the gulars and anals may show a trace of such pigmenta-
tion. Plastron substantially as described below for adults.

Color of adults. Ground color of carapace pale yellow to
horn with variable markings so that scarcely two are alike ; dorsal
pattern almost always more or less distinctly rayed like mem-
bers of the Psammohaies geometricus group with the rays tend-
ing to be much less regular, center of areola yellow surrounded
by very irregular traces of brown while immediately outside
the areolar area is a narrow zone of yellow, beyond which to the
])eriphery of the shield is a broad black border broken by fine or
broad yellow rays, sometimes one color, sometimes the other pre-
dominating according as the light rays are broad or narrow,

1 Fiffnrpfi by Procter n92L'b :49.'i :fi^'. 7), also prusent in U.S.N.M. 72530 and
~2')47, probably from Doiloina.

2 Gular smaller than the anal in thf Xjoro tortoise (Stockholm Mus.).

292 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

heavy or faint. Plastral shields with yellow areolae which may
be smudged with brown; around the areola a broad zone of
black, broken to a greater or lesser extent by yellow rays, in
some cases, extends to the periphery of the shield, while in
others it is separated from the periphery by a narrower or wider
zone of yellow. In certain extreme cases the plastral shields ap-
pear almost black, in others mostly yellow. Very rarely a speci-
men may be almost uniform horn color both above and below. ^

Size. Carapace length of a large S (M.C.Z. 30008) 145 mm.,
breadth 102 mm., height 32 mm. ; the corresponding figures for
a S cotype of loveridgii (Brit. Mus.) being 140 mm., 100 mm.,
and 27 mm. Carapace length of largest ? (M.C.Z. 23024), 177
mm., breadth 131 mm., height 40 mm. ; the corresponding figures
for the 2 type of tornieri (Berlin Mus.) being 160 mm., 110
mm., and 35 mm., while those of the juvenile type of proderae
(Brit. Mus.) are 85 mm., 72 mm., and 40 mm.

When proderae was compared with a tornieri of exactly the
same carapace length, a difference in its relation to the breadth
(84.5%) and depth (477c) was noted, the same figures for the
tornieri being 90.5% and 26^0 respectively. However, at least
in the breadth/length ratio proderae is within the range of
Dodoma tornieri which ranged from 71 to 83.8% in seventeen
specimens measured by Procter, and from 69.1 to 94.2 7o in
twenty-five others measured by Loveridge.

Breeding. Courtship consisted in the $ impatiently snap-
ping at the limbs of the 9 as he followed her about, at times
clambering on her back and biting viciously at her head when-
ever she ventured to thrust it out. Two $ $ were so ill-tem-
pered as to seize the edges of the 9 9 carapaces in their jaws,
drag them along and eventually get beneath as they appar-
ently attempted to overturn them. One $ persistently, though
unsuccessfully, endeavored to mate with a 9 Kinixys h. helliana
which shared the enclosure. Pairing took place in January and
February, as early as 9 A.M. and as late as 4.30 P.M.

Females appeared gravid in April. A single elongate egg is
laid in July or August in East Africa, but Conant and Downs
(1940:48) record an egg weighing 33.5 gm., and measuring 28 x
48 mm. being laid in the Philadelphia Zoo on January 9, 1937.

1 As in U.S.N.M. 73945.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 293

Two eggs removed from the enclosure at Kilosa, where they
had been buried beneath a rockery, measured 26 x 42 mm. and
28 X 44 mm. The shells were extremely thin and brittle but
one (M.C.Z. 43834) was successfully drilled and blown. On
December 19, 1922, an embryo on the point of hatching was
removed from another egg and preserved (M.C.Z. 18167).

Longevity. The Soft-shelled Tortoise is very hardy in cap-
tivity. One captured December 8, 1918, and taken to England
in midwinter, lived in the Zoological Gardens, London, until
July 2, 1927, i.e. 8 years, 6 months and 24 days in captivity
(Flower, 1937, records only its stay at the zoo). Conant and
Hudson (1949) mention one that lived at Philadelphia Zoo for
7 years, 5 months and 6 days.

Diet. Apparently Soft-shelled Tortoises emerge to feed only
during the early morning hours. At Tabora one was found nib-
bling dry grass. In captivity they throve on lettuce and tender
cabbage leaves or, for long periods, were fed on bread soaked in
jam (Loveridge). Bread soaked in milk was taken, also clover,
dandelion and lettuce ; fruit only sparingly and they refused
bananas altogether. The species is exclusively vegetarian (Mer-
tens: 1942d).

Parasites. A tick (Amhlyoynma nuttalii'^) was on the type of
procterae when found. None of the extensive Dodoma series was
so att'ected but soon acquired ticks when transported to Kilosa.

Habits. At 9 A.M. some young were found basking on a rock
slab, though usually during daylight these tortoises spend their
time in rock crevices into which they climb. Their depressed
shells enable them to push their way farther into such retreats
than if they were convex. When surprised suddenly the reptiles
may be removed with ease, but if warned by a shadow falling
across the entrance, they inflate their lungs so that their shells
press upon the rock both above and below as they brace their legs
like struts. Under these conditions it frequently took as much
as an hour to work a single tortoise loose (Loveridge).

Eleven w-ere found beneath one flattish slab of rock in Janu-
ary, possibly aestivating as that and February are the hottest
months at Dodoma. In conformity with their structure, these
tortoises are far more active than their allies of the genus Geo-

1 OriginaUy identified as marmoreuin by S. Hirst; identification corrected by
J. C. Bequaert.

294 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

chelone, while their predilection for clambering up and falling off
rocks calls for rapid recover}^ in turning over should they land
on their backs. In captivity they displayed surprising agility
in climbing up vertical wire netting two feet in height, and many
escaped. At night shelter was sought beneath the rockery where
they would pile up on top of one another (Loveridge).

Mertens (1942d) found his captive specimens, though not noc-
turnal, spent most of the daylight hours in their hiding place.
Even when feeding they did not venture more than nine metres
from their retreat, which is definitely selected and constantly
used thereafter. To it the tortoise hastens when surprised in the
open, for M. tornieri, unlike most tortoises, does not, or only mo-
mentarily, withdraw its head inside its shell, preferring to make a
dash for its home. This it locates in a matter of minutes if taken
to a distant corner of the enclosure and then released. Also,
after being removed to Avinter quarters, when liberated in the
spring it readily located its ' ' home. ' '

Habitat. Small hills with rocky outcrops in arid thornbush or
savanna, at altitudes from al)out 100 to 4000 feet. The most un-
usual habitat record was that of a tortoise taken by an African
when "cleaning round his rice plants about a hundred yards
from" Allen Turner's camp at Mida Creek.

Localities. Kenya Colony: 10 miles south of Malindi at Mida
Creek; Mathews Range {fide J. G. Williams) ; Njoro. Tangan-
yika Territory: Busisi (Bussisia) ; Dodoma; Ikikuyu; Kibakwe;
Kidengi; Kikombo ; Kondoa Irangi ; Lindi; near Lake Eyasi (as
Njarassa) ; Mangasiui; Matete ; Mfilima; Tabora; Usandawi.

Range. Kenya Colony (in suitable terrain from Njoro east to
Malindi) south to Tanganyika Territory (from Busisi, Smith
Sound, Lake Victoria southeast through Ugogo to Lindi on the
southeast coast ) .

Genus PSAMMOBATES Fitzinger

18:^.1. Psammobates Fitzinger, Ann. Wiener Mus., 1. pp. 108, 113, 122.

Type: Testudo geometrica Linnaeus (designation by Fitzinger.

1843, Syst. Eept., part 1, p. 29).
193311. Chersinella Hewitt (not of Gray: 1870), Ann. Natal Mus., 7, p. 259.

Ty]>(^: Trsfiido fjeomptrica Linnaeus (by original designation).

LOVKRllX^K ANT) WILLIAMS: AI'UICAN' CRYI'TODTRA 205

Definition. Skull with tritiiratinp: surfaces of maxilla and pre-
nuLxilla without ri{l<iiii<i- ; inuxiliaiy participatiuji^ in roof of
palate; anterior palatine foramina small, eoneealecl in ventral
view; prootie narrowly exposed dorsally ; quadrate enclosing
stapes; snran<i;ular subecpial iu h('i<>lit to prearticidar ; neck with
third or fourth centrum biconvex.

Carapace never liinoed; normally the anterior neurals hex-
agonal; outer side of third costal scute about as long as, or longer
than, that of the fourth ; submarginal scute absent ; suprapygals
1 or 2, if 2 they are separated by a straight transverse suture.

Plastron not hinged ; gular region scarcely thickened and pro-
duced ; gulars paired, longer than broad.

Range. Bechuanaland Protectorate; Union of South Africa
(Orange Free State and Cape Province) ; Southwest xVfrica.

Fossil record. None.

Remarks. When Lindholm (1929:284, footnote) stated that
the genus Testudo {sensii Strauch, Boulenger and Siebenrock)
was unique, not only among reptiles and vertebrates but in the
entire Animal Kingdom, he was referring to the fact that it
contained some species (citing Geochelone gigantea) that in vol-
ume and weight were several hundred times greater than such
alleged congeneric species as "fiski" (i.e. Psammobates tentorius
verroxii). This is one of the tortoises of the geomctricus group,
none of whose members is known to exceed 150 nnn. in carapace
length.^ Re-examining the ease, we have found that in addition
to size the other characters of the geometrictis group require its
separation as a genus distinct from both Testudo and Geoche-
lone.

The geometricus group is emydine-like in maintaining as a
normal condition the hexagonal neural pattern that Geochelone,
Testudo and Gopher us — all three independently — abandoned
(except as an occasional variant) 30 or more million years ago.
In the general discussion of testudinine genera above (p. 218)
the geometricus group (as Psammobates) was assigned to the
larger assemblage of which Homopiis is a central member. The
character of hexagonal neural pattern has been one reason for
this assignment, but several other considerations have entered

1 We have not seen any larger than this. Imt larger specimens of geometricun
itself have heen reported (p. 323).

296 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

into this decision. A discussion of the problems involved seems
pertinent here.

Among African tortoises several groups of species occur which
are evidently endemic, as species groups, to this continent. Ne-
glecting Malacochersus which seems quite isolated, the rest have
several features in common. None of them has any close relative
or representative outside Africa or even in North Africa. All
have the neural bones hexagonal or vaguely quadrate or irregu-
lar, never with the regular octagonal and quadrilateral pattern
that is characteristic of all Recent tortoises outside the Ethio-
pian region. Chersina approaches the advanced pattern more
closely than any other of this group but still onl}' occasionally
and imperfectly. In all, the maxillary triturating surfaces are
without definite ridging. Chersina again is atypical but it
has only a very feeble ridge which is more easily felt than seen.
All except Homopus have the primitive single or transversely
divided suprapygal, not the geochelonide type. In addition all
are of only small to moderate size.

Against these general resemblances are to be set the striking
peculiarities which separate the African endemic species groups
not only from each other but from everything else. We evaluate
these peculiarities, as we have stated elsewhere (p. 218), as imply-
ing generic distinction. Yet it may be worth consideration that
despite these peculiarities there have been instances of taxonomic
confusion. For example, a color variant of a member of the
geometricus group was described as a species of Homopus {II.
bergeri Lindholm), and juvenile specimens of Kinixys with the
carapacial hinge still undeveloped have been twice confused with
Homopus (H. nogueyi Lataste and H. darUngi Boulenger).
Though inability to separate forms is not always indicative of
relationship, such errors at least make clear that the characters
used to separate generic groups are not conspicuous always or
at all ages. On our part, we believe that the resemblances of the
African endemics to one another are really indicative of close
relationships among them. We regard these endemics as a true
autochthonous African group.

At this point it is necessary to mention the two endemic Mala-
gasy tortoises — Acinixys and Pyxis. The skull of Acinixys has
maxillary triturating ridges and its carapace has hexagonal

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIKA 297

neurals. In Pyxis this situation is reversed, its skull lacking trit-
urating ridges while its carapace shows the octagonal and quad-
rilateral neural pattern ; in addition there is in Pyxis the obvious
specialization of a very peculiar plastral hinge. Both possess the
primitive pygal pattern, and in size they fit in the middle of the
continental African series. Thus while seeming to resemble the
continental African endemics, each of the Malagasy endemics
differs from that series in a single character — in each case a
different one. Note, however, that Chersina of continental
xifrica approaches each of these in precisely its aberrant charac-
ter — Acinixys in its maxillary ridge and Pyxis in its approxi-
mations of an octagonal and quadrilateral neural pattern.

We interpret this case, as we have others, in terms of a con-
ception of the evolution of tortoises as resulting from modifica-
tions in parallel occurring at very different tempos in the sepa-
rate lineages and sub-lineages. Thus for the most part the Ethio-
pian endemics (including now the Malagasy with the continental
African forms) have been slow to modify the primitive (emydine)
neural and pygal patterns at the same time that most of them
have gone the whole distance in the loss of maxillary ridging.
Acinixys has been slow to evolve both maxillary and neural ad-
vanced features, while Pyxis has been "progressive" in both
these details, but both Acinixys and Pyxis remain primitive in
the pygal pattern in which Homopus has gone on to an advanced
condition.

We see the Malagasy endemics therefore as part of one series
with the African endemics — part thus of a larger autochthonous
Ethiopian group. Recognition of such a group has a certain geo-
graphical tidiness, but thei-e is very much more to support this
conception than just this tidiness. We may argue from any one of
the resemblances shared by these Ethiopian forms. Thus the
primitive pygal pattern in an Ethiopian tortoise can be explained
in one of three ways: (1) as an emydine inheritance going back
to a separate emydine ancestor; (2) as an emydine inheritance
shared with other Ethiopian tortoises and going back to a com-
mon ancestor; (3) a secondary reversion to a primitive condition.
The likelihood of the first and third of these possibilities is much
diminished by the occurrence of several forms with the same
condition in one region, still further reduced by other resem-

29S HrLLEilX: iVIlTSliUW OF COMPARATIVE ZOOLOGY

hlances amono: the forms. Thus, by elimination, the second of
these sug-f?estions appears to be the most likely. On the basis of
primitive neural })attern an argument essentially identical may
be set up, and a very similar argument will apply for the un-
ridged maxilla. All of these taken together and considered along
with the subtler cross resemblances among the forms would seem
to strongly support the naturalness of our postulated group.

If this reasoning is valid and the Ethiopian endemics as a
group do have a common ancestor independently of the other
tortoises, then the geometric us group as one of this series cannot
l>e retained in Testudo, at least not in any conception of tortoise
genera short of an inclusive genus that would embrace the entire
subfamily. Below that level tw inclusive genus would he at once
definable and natural. We do not argue against the genus-sub-
lamily concept on grounds of merit, but to apply this viewpoint
in only one subfamily would be to create a genus whose dimen-
sions and variability are not consonant with those of the genera
of any other subfamily or family in the order.

The argument that Psammobates must be separated from Tes-
tudo, as typified by T. graeca, does not automatically imply its
distinctness from every other genus. "We maintain it as separate
because of its morphological distance from its closest relatives.
Thus in skull structure Psammobates shows one feature in which
it is distinctive within its lineage, and one that is altogether
unique.

(1) The space between the ventral processes of the prefrontals
is more restricted and, consequently, more emydine-like than in
any other testudinine (Fig. 11 //, /). Indeed the opening in
Psammobates approaches, but is still not quite as narrow as the
widest narial apertures in emydines, e.g. Hiercmys and Sieben-
rockiella. !~5ince at least a moderate restriction of the narial
aperture is probably })rimitive for all the Testudinidae (the emy-
dine condition occurs in Platijsternon) , in this feature Psammo-
bates is probably more primitive than any other living group of
testudinines.

(2) Behind the anterior palatine foramina of Psammobates
the maxillary bones send up into the roof of the palate more or
le.-ss sizable wings which, from each side, approach, but never join,
the vomer. This is decidedly a specialization.

LOVEBIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 2n9

In its shell structure, Psammobatcs displays a condition (in
certain subspecies in very exfi<>p:erated form) unique among the
African endemic genera — conically raised vertebrals. This con-
dition, together with a very similar color pattern, occurs in some
Geochelone, notably in G. clegans of Asia, but conical vertebrals
also occur in G. pardaJis habcocki and some South American tor-
toises. We consider this similarity convergent only, not more
significant tlian the superficial similarity of Emydoidca with
Emys, or the convergent resemblance of various agamid and
iguanid lizards.

Differences in skull and shell as great as these seems to merit
generic separation. Such recognition has already been accorded
the group by Hewitt and other students of South African tor-
toises. The name Chersinella Ciray, 1870, was at first employed
by Hewitt. Unhappily tliis is doubly invalid, first because its
genotype, as selected by Lindholm in 1929, is Testudo graeca
Linnaeus, and also because a much older name is available. The
correct name Psammobates Fitzinger, 1835, with T. gcometricn
Linnaeus as genotype, was subsequently adopted b}' Hewitt
(1937e). The gender of Psammobates being masculine, it com-
pels changes in termination of all the familiar names of the
geonietricus group.

General Survey of the Psammobatcs Species

Xo group of tortoises has fostered the making of so many
specific names as has Psammobates. For the three species that
we recognize there are 32 names available (see following list).
This plethora of names provides examples of nearly every kind
of taxonomic error. Names have been applied to adults and
young separately (geometrica and luteola; semiserrata and
ocidifera). Names have also been erected: (1) due to the mis-
reading of a type description (smithi Boulenger) ; (2) for strik-
ing color variants [bergeri Lindholm) ; (3) for striking scute
variants {strauchi Lidth de Jeude) ; (4) for minor scute variants
{boettgeri Siebenrock) ; (5) for ill-defined local populations
(various names of Hewitt) ; (6) for selected series within a
single local population (other names by Hewitt) ; (7) for inter-
grading populations (names by Plewitt).

300 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Most of these unnecessary names apply to a single species —
tentorius — individuals of which show an astonishing range of
pigmentary and morphological variation. The errors of such
earlier workers as Boulenger and Siebenrock resulted primarily
from a lack of adequate series with definite locality data, their
"new species" being based on one or very few specimens.

The extraordinary multiplication of names by Hewitt had a
very different foundation — a pliilosophy of naming which w' e
regard as unfortunate. Finding, w^hen dealing witli fairly large
local collections, that he was more or less able to recognize mem-
bers of these local series, Hewitt provided each such population
with a formal Latin name. When these populations proved less
homogeneous than usual, Hewitt proposed additional names for
the extreme types, e.g. the three names given variants within the
Hanover District population.

A list of names proposed for members of the genus Psammo-
bates (Table 5) is offered as a ready means of recognizing the
status assigned to each in this revision.

Grouping of Species

As suggested by Siebenrock (1904 and 1910), the species of
Psanimohates fall into two groups distinguished by the size of
the nuchal, the number of the axillaries, and the plastral pat-
tern. As is usual in testudinines every one of these characters is
subject to individual variation, so that occasional individuals
appear to transgress their group boundaries in one or another of
these characters.

The first species group within Psammohates consists of two
allopatric forms — geometricus and oculifer. Both differ from
the teiitorius complex in having a plastral pattern that covers the
entire plastral area. Typically it is an elaborate rayed pattern
in both geometricus and oculifer, but in the first it may sometimes
be simplified (see Hewitt :1933b, pi. 14, fig. 6), and in the second
it is sometimes indistinct. Both forms have only one axillary,
but in oculifer it is almost always fused with the humeral (Fig.
■SOB).

Normally the nuchal shield is well developed and longer than
broad in both species, more consistently in oculifer where it is
regularly large and subtriangular. In geometricus, while usually

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA

301

Table 5

Chronological List of Specific and Subspecific Names
Proposed for Members of the ^'geometricus Group,"
(i.e. Psammobates) , and their suggested Synonymy.

C. = Chersinella ; H. = Homopus; P. — Psammobates ; T. = Testudo

3.
4.
5.
6.
7.

9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.

T. geometrica Linnaeus, 1758

T. lu.teola Daudin, 1802

T. oculifera Kuhl, 1820

r. tentoria Bell, 1828

T. verroxii Smith, 1839

T. semiserrata Smith, 1839

T. geometrica var. tentoria (non Bell)

Gray^
T. geometrica var. nigriventris Gray,

1855
r. trimeni Boulenger, 1886
T. smithi Boulenger, 1886
T. fisM Boulenger, 1886
T. strauchi Lidth de Jeude, 1893
T. seimundi Boulenger, 1903
T. hoettgeri Siebenrock, 1904
H. hergeri Lindholm, 1906
T. oscarhoettgeri Lindholm, 1929
C. tentoria alhonica Hewitt, 1933
C. tentoria tentorioides Hewitt, 1933
C. tentoria piseatella Hewitt, 1933
C. tentoria subsulcata Hewitt, 1933
C. tentoria Tcaruica Hewitt, 1933
C. tentoi-ia duerdeni Hewitt, 1933
C. tentoria lativittata Hewitt, 1933
C. tentoria Tcaruella Hewitt, 1933
C. tentoria hexensis Hewitt, 1933
C. schonlandi Hewitt, 1934
C. fisJci cronwrighti Hewitt, 1934
C. fisM orangensis Hewitt, 1934
C. fisTci coleshergensis Hewitt, 1934
C. fisM grica Hewitt, 1934
C. fisM gricoides Hewitt, 1934
C. verroxii amasensis Hewitt, 1934
P. depressa FitzSimons, 1938

= P. geometricus
= P. geometricus
= P. oculifer
= P. t. tentoriu.<i
=: P. t. verroxii
= P. oculifer

= P. geometricus

= P. t. tentorius

z= p. t. trimeni

= P. t. verroxii

=^ P. t. verroxii

z= P. geometricus

= P. t. verroxii

=^ P. t. verroxii

= P. t. verroxii

= P. t. verroxii

= P. t. tentorius

■=■ P. t. tentorius X verroxii

= P. t. tentoritis

:= P. t. tentorius

= P. t. tentorius X verroxii

= P. t. tentorius X verroxii

=: P.t. tentorius

= P. t. tentorius

= P. t. trimeni

= P. t. verroxii

= P. t. verroxii

^ P. t. verroxii

= P. t. verroxii

= P. t. verroxii

= P. t. verroxii

= P.t. verroxii

= P. t. verroxii

1 Though not a new name, this synonym is included here because of its im-
portance, due to misuse at so early a date (1855).

:302 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

a moderate sized though slender scute, it is sometimes absent —
as in the type of st ranch i v. Lidth de Jeude and another speci-
men in Leiden, in such cases of al)sence the first marginals meet
medially, thus seeming to imply that the nuchal has been lost by
extreme attenuation.

The scalation of the forelimb diifers in the two species (PI.
lA, B), but in both there are only a few enlarged scales and many
small ones. This is in contrast to the situation in the tentorius
group where there are more enlarged scales in rather definite
rows, and fewer smaller ones.

Though P. oculifer has a very extensive range, it seems not to
extend south of the Orange River. Boulenger's record of this
species from "between Richmond and Victoria West'' (ques-
tioned by Hewitt :1934) was based on a misidentified te^itorius
verroxii} On the other hand, geometricus is, or was — it is now
very rare or extinct — confined to a few districts in the vicinity
of the Cape Peninsula. Though these are well-marked forms,
readily distinguishable on color, shape of nuchal, scalation of
forelimb, etc., in view" of their geographic separation it is diffi-
cult to evaluate the differences in terms of species versus sub-
species distinction. It is always possible to claim that these are
clinal extremes of a foi'merly intergradient population. How-
ever, since distance has been demonstrated on more than one
occasion to mean reproductive isolation, even when morphologi-
cal dift'erence is not great, and since the two forms are separable
on many characters and cannot intergrade in nature, we have
conservatively maintained them as full species. In modern sys-
tematic terms, however, they certainly belong to one superspecies.

In this connection a further point may be mentioned. North
of the Orange River oculifer occurs over part of the range of
tentorius verroxii. Differences beyond sympatric species are
commonly used as a measure of the specific distinctness of allied
allopatric forms. With this in mind a comparison reveals that
the populations of oculife7- that coexist with the far more variable
t. verroxii, resemble the latter more than do the more distant
t. tentorius and t. trimeni. Botli sympatric populations are rela-
tively dull colored and have large femoral tubercles ; neither have
high conical vertebrals. Usually oculifer has a very serrate

1 Specimen examined at the British Museum by E.E.W.

LOVERIDGK AXD WILLIAMS: AFRICAN CKVPTODIRA 30!?

inarfrin hut sometimes varies towards a non-serrate condition;
while t. verroxii is normalh^ not serrate, occasionally within the
range of oculifer it may be stron<>ly serrate (Power :1932b). Also
in some specimens of t. verroxii the nuchal shield may be rather
large, in size and shape approaching the nuchal of ocidifer. In
general the differences between th(>se sympatric species, while
perfectly clear-cut, are no stronger than those between geomctri-
cm and oculifer. On this criterion also these two last may be re-
garded as full species.

The second species group is interpreted by us to consist of a
single highly variable species — tentorius — of complex popula-
tion structure. Reference has frequently been made to individual
variability among testudinines but — measured by the variation
seen within a single small local population — probably no one
species exhibits greater variability than does tentorius. Two ex-
cellent studies of such variation in tentorius verroxii have
already been published. That of Power (1932b) on a series of
25 from, or near, Xiekerks IIo])e. just north of t!ie Orange River,
and another by Lorenz Miiller (1939a) on 45 individuals of the
same race from Carmel Rust, near Phillipstown. Much further
information on variation can be found in Hewitt 's several papers,
also in Duerden's (1907c) review of the '' gcomctrieus group."
The last two authors have seen far more material than anyone
else, and therefore should have had a better general view of both
individual and geographical variation, but we must add that
each of these authors has, in his own way, diminished the value
of his work for subsequent investigators.

Far more unfortunate than Hewitt's peculiar taxonomic con-
cepts are the unsystematic and erratic nature of his descriptions.
A great many individual specimens are described in some detail,
but the descriptions are not comparable, i.e. do not consistently
deal with the same characters; consequently they are not usable
by anyone lacking access to his material.

Duerdeii's fault springs from an opposite prejudice to that of
Hewitt — denigration of all local varieties — and with it a ten-
dency to suppress all locality data. Thus in Duerden's paper
there are many general statements about variation, but too often
these statements are not tied to specimens with specific localities.

Comments such as these on Hewitt's and Duerden's Avork

304 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

would be neither necessary nor appropriate if theirs were not
the major works on this whole group, done on the spot and with
abundant collections. Although one or both of us have seen much
material, including types, in the British Museum, in Leiden, in
Frankfurt, Lorenz Miiller's series in Munich, cotypes of several
of Hewitt's forms in the Museum of Comparative Zoology and
certain material generously loaned by the Transvaal Museum,
we have not seen all of this material at one time or in one place.
Nor have we seen the animals alive, and we are in no position to
solve by judicious collecting in critical localities any of the many
problems that have become evident to us. It remains an ines-
capable fact that these tortoises are forms of Southern Africa
which are not too profitably studied at a distance of many thou-
sand miles. The complex relationships will be satisfactorily re-
solved only when South African workers, utilizing all the
material available to them, undertake studies of variation as
meticulous as those of Power and Lorenz Miiller.

Owing to the distance separating us from the principal South
African collections, and because of our inability to explore the
intricacies of local variation, we have been compelled to take a
broad view of the tentorius complex. While not fully satisfied
with our interpretation, we present the following picture :

In the vicinity of Grahamstown, in southeastern Cape Prov-
ince, a Psanwiohates population occurs that is characterized by:

(1) high conical vertebrals;

(2) a strongly contrasting dorsal pattern of narrow yellow or
orange rays on a dark ground ; ^

(3) a plastral pattern consisting of a dark central figure with
sharply defined boundaries that are not broken up by lateral
rays, the whole surrounded by a broad yellow border ;

(4) the scalation of the forelimb from elbow to wrist, usually
with three longitudinal rows of enlarged scales, none of which is
conspicuously disparate in size ;

(5) hinder side of each thigh with a very large, sharp, sub-
conical tubercle.

In the northwest corner of Cape Province, including western
Little Namaqualancl near the mouth of the Orange River, and in

1 However, we figure a specimen (PI. lOD) in which the relative width of
blaclc and yellow rays is reversed. This lighter pattern is still very different
from anything that has been observed in other populations.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 305

nearby localities on the slopes behind the high plateau, there is
present a second population characterized by :

(1) high conical vertebrals;

(2) a strongly contrasting dorsal pattern of moderately wide
yellow or orange rays on a black ground ;

(3) a plastral pattern consisting of a dark central figure with
sharply defined boundaries broken through by numerous yellow
rays, the whole surrounded by a broad yellow border ;

(4) the sealation of the forelimb from elbow to wrist without
three longitudinal rows of enlarged scales; instead, either a
single scale greatly enlarged in relation to the others, or else a
single row of especially enlarged scales ;

(5) hinder side of each thigh without any very large, sharp,
subconical tubercle, though small inconspicuous tubercles may be
present.

On the plateau of eastern Little Namaqualand, just east of
the last population, is one that difi'ers from both the preceding
groups. Its characters are :

( 1 ) vertebrals quite flat, not in the least conical ;

(2) a dorsal pattern that is not so strongly contrasting, its
rays broader, its coloring duller;

(3) a plastral pattern consisting of a dark central figure, as in
the preceding groups, but paler and characterized by the indis-
tinctness of its boundary with the surrounding broad yellow
border ;

(4) the sealation of the forelimb from elbow to wrist consists
of relatively few scales variously enlarged ;

(5) hinder side of each thigh with a very large, sharp, sub-
conical tubercle.

These are the three peaks of character differentiation which are
to be found within the tentorius complex. All three populations
share certain characters :

(1) the plastral pattern never covers the entire plastral area,
which always has a broad clear border ;

(2) usually two axillaries present, sometimes more, rarely one;

(3) the nuchal shield is usually broader than long, frequently
minute, occasionally absent, in the latter event the first marginals
do not meet.

With these characters uniting them in a species group, what

306

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

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liOVERIDGE AND WILLIAMS : AFRICAN CRTPTODIRA 307

do the three populations represent — species, subspecies, or local
assemblages unworthy of nomenclatorial recognition?

So far as we are aware no intergradation occurs between the
Orange River mouth population and the one on the plateau to
the east of that area. In fact, representatives of both populations
labeled ' ' Steinkopf ' ' are present in the Museum of Comparative
Zoology collection. Hewitt (1934:308-309) has suggested that
the populations are probably allopatric; an opinion with which
we concur though apparently the evidence is not positive. How-
ever, whether the populations overlap or not is relatively unim-
portant. They are adjacent; they are strikingly different, and
they show no sign of interbreeding. In short they are behaving
like good species.

But these populations in northwestern and northeastern Little
Namaqualand are the only adjacent populations in the tentorius
complex that are sharply distinct. Everywhere else contiguous
populations of the tentorius complex merge into one another.
Less strongly differentiated populations cluster around the three
pealvs of difference already described and, wherever they meet,
coalesce by rather gradual transitions.

At this point, in order to facilitate ready reference to the con-
cepts involved, we propose applying the available scientific
names, using for this purpose topotypical ones that may ulti-
mately be synonymized. Thus trimeni Boulenger, whose concept
is substantially the same as ours, can be applied to the Orange
River mouth population in which the plastral pattern is broken
into sharpl}' distinct fragments.

For the moment schonlandi Hewitt may be applied to the
neighboring population in eastern Little Namaqualand. The pre-
cise locality of the S type is unknown, but a 9 paratype was
suspected to have come from O'okiep. The type locality may con-
fidentlj^ be restricted to O'okiep as the population of this and
surrounding localities is clearly what we understand by schon-

308

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

landi. Our conception is based on three paratypes from Stein-
kopf (M.C.Z. 4222-4).

The name tentoria Bell may he applied to the population cen-
tering around Grahamstown. Here again the type has imprecise
locality data, merely "South Africa," but we may with confi-
dence restrict the type local it}^ to Grahamstown, as the type
conforms satisfactorily to that population.

Atlantic Ocean ^^^^

Fig. 29. Distribution of vertebral height in Psammobates tentorius. Sub-
jective estimates of average height in the various populations.

(P. Washer del.)

It is now necessary to di.scuss the populations satellite to those
which constitute peaks of difference.

To the south of typical trimeni in northeastern Little Nama-
qualand is a population displaying the characters of trimeni in

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 309

much reduced intensity. The vertebrals instead of being conical
are relatively low. The foreliinb no longer i^ossesses any scale
that is strikingly larger than its fellows. The dorsal coloration
is duller, the rays wider. The plastral pattern is interrupted by
3'ellow rays but is no longer separated transversely. We have an
example of this sort from Bitterfontein (M.C.Z. 33451), and have
seen others from near Calvinia (T.M. 18258) and Nieuewerust
(T.M. 18257). Similuar specimens from Hex Iliver Valley were
named tentoria hexensis by Hewitt.

To the north and west the typical population of tentorius
grades into others that possess its characters in an attenuated
degree. The vertebrals are less conical ; in this character there is
a considerable range of variation but it centers about a different
and lower mode. The dorsal pattern becomes less sharply defined
and may have broader rays. The plastral pattern changes verj'
little but occasionally may be very faint, even though still sharply
bounded. Still farther north on the Karroo plateau the popula-
tions progressively lose tentorius characters and approximate
those of schonlandi. The vertebrals are only slightly and occa-
sionally raised. The dorsal pattern is very variable but usually
broad rayed. The plastral pattern is extraordinarily variable,
sometimes almost a faded, vaguely outlined, tentorius pattern, or
an indefinite rayed pattern, in others just an indistinct central
infuscation, or entirely absent.

It is for these populations to the north and northwest of ten-
torius that extreme variability has been recorded by Power and
Lorenz Miiller. Of them it is said that no one individual is like
another, and the amplitude of variation is indeed such that from
one locality Boulenger described two full species, while from the
same place Hewitt described still a third form ; all regarded by
us as strict synonyms. These plateau populations appear to be
really more variable than typical trimeni or typical tentorius,
their characters only attaining some degree of definition and con-
stancy where, as schonlandi Hewitt, their ranges abut on that of
trimeni.

The variable populations of the Karroo plateau extend north
of the Orange River into Great Namaqualand. Here a distinct
variant occurs. This is the melanistic unicolored variety named
hergeri by Lindholm. So far, only single specimens have been re-

310 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

corded, viz. the type from Gibeon; another from **Herreroland"
(Werner :1910a) ; a third from Upington in Gordonia district
(Siebenrock: 1910). Also referred to hergeri by Mertens (1955a:
37) is a specimen without exact locality identified by Sternfeld
(1911d:49), and one in his own collection obtained at Klein
Windhoek but believed to be an escaped captive not native to the
locality. We have also seen a British Museum example from
Barby, Tiraz Mountains, Southwest Africa.^

Since in each instance these hergeri individuals appear to differ
from surrounding- populations only in color, and their distribu-
tion so far as known is quite erratic, we have regarded hergeri
as merely a color phase occurring in, but nowhere characteriz-
ing, the northern populations of the plateau tortoises of this com-
plex. On the other hand, when Mertens (1955a :37) was in
Southwest Africa he was told that all the tortoises in one part of
Great Namaqualand were of a uniform red-brown color. Such a
report requires verification, for of the "hergeri" localities only
Gibeon and Barby are relativel}' near one another. Not only is
Upington remote from both, but it is separated from them by
localities (e.g. Keimoes, Lower Molopo River; Klein Karas
Mtns.) in which patterned tortoises occur.

Whether the Great Namaqualand populations allied to schon-
landi meet trimeni on the south, or whether they are separated
from the latter by the Orange River, is uncertain. From north
of the Orange the only records of trimeni worth attention are
those of Werner (1910a) and Mertens (1955a). Werner (1910a)
presents a photograph of the specimen from Keetmanshoop, but
this is too poor to permit of reappraisal. The descriptions and
photographs of Mertens (1955a) are more satisfactory and indi-
cate that trimeni may really coexist with scJionlandi-like forms
north of the Orange River.^

Summarizing the information so far presented, we postulate
the following :

1 The skull of this specimen has been prepared and confirms the assignment
to Psammobates on three grounds: (1) the characteristic maxillary projection
extends on to the roof of the palate ; (2) the ventral processes of the vomer are
approximated ; (3) the anterior palatine foramina are small and inconspicuous.

2 Dr. Mertens states (in a letter of 19.xii.55) that the plastral patterns of the
Southwest African specimens referred by him to trimeni (19r)5a:36), agree "sehr
genau" wih the pattern of the type of trimeni as figured by Boulenger.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 311

The northwest form (typical trimeni) grades southward into
a less distinctive population that approaches more closely the
other assemblages of the tentorius complex. The southeast form
(typical tentorius) grades in almost cliual fashion northward
and northwestward into populations again less and less like the
topotypes and increasingly like a third distinctive group. This
third population, nearly or entirely confined to the inland plateau
(whereas the other assemblages are more nearly coastal), is over
most of its range very variable (to a limited extent geographi-
cally, to a much greater extent individually) and is only sharply
distinct and relatively uniform where, as schonlandi, it meets the
northwest population.

Undiscussed so far is the southwest area that the southern pop-
ulation of trimeni, and that allied to schonlandi, approach from
the north, and which the populations of tentorius enter from the
east. This is the meeting place of all three of the major series of
assemblages of the tentorius complex, and adjacent to this meet-
ing place the representatives of all three have already lost every
distinctive character except plastral pattern. All are relatively
dull in color, without very convex vertebrals and with rather uni-
form forelimb scalation, i.e. three longitudinal rows from elbow
to wrist.

Through the courtesy of the Transvaal Museum we have been
able to examine a series of nine specimens from Matjesfontein,
which is in this critical area. All three types of plastral pattern,
as well as intermediates between them, occur in this small sample
(PI. 11).

This then is the picture which has emerged for us as a broad
and general view of the tentorius complex. It remains to be con-
sidered how this situation is to be interpreted biologically and
treated taxonomically.

Despite the lack of any evidence of introgression or intergra-
dation between typical trimeni and adjacent ''schonlandi," the
tentorius complex taken as a whole appears to be a reproductive
unit. By the indirect route of a chain of allopatric populations
which appear to interbreed freely at their points of contact, it
seems clear that even trimeni and ''schonlandi" must be able to
exchange genes. We may, therefore, consider that the whole com-
plex constitutes one species for which the name Psammohates

312 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

tentorius (Bell) is available. The question arises as to whether
this species should be subdivided ; if so, how minutely ?

The problem of species subdivision is only partly a biological
one. To describe the biological situation completely would re-
quire a far more elaborate analysis than that provided above and
a knowledge that no one now possesses. No series of names can
prove a satisfactory^ substitute for such a biological description.
If it is admitted that names cannot tell the full story, is there
any use or purpose in applying subspecific names?

We believe that the three well-differentiated centers or peaks
already described, around which other populations cluster, merit
recognition, especially in view of the fact that two of them behave
toward one another as if they had already reached the specific
level. By granting these populations names we are endeavoring
to indicate something both in regard to their past and to their
potential history. We believe that the typical populations of
trimeni, " schonlandi," and tentorius, represent the focal loci of
the tentorius complex during a previous fragmentation of its
range ; that during this former separation these three popula-
tions built up the sj^stem of features that now distinguish them
as peaks of difference within the complex. In terms of characters
they are even now almost at the specific level, but the satellite
populations surrounding them — showing as they do every ap-
parent evidence of intergradation — testify that reproductive
isolation has not been achieved. Still the process of differentia-
tion has gone so far that we may rather confidently assert that if
fragmentation and isolation again occurred, these populations
showing peaks of difference are the best candidates to pass the
species threshold. As true "incipient species," we believe they
may legitimately receive names.

Our subspecies are to be conceived of primarily in terms of
the peaks of difference — the three topotypical populations of
trimeni, " schonlandi," and tentorius. Simply for convenience in
demarcating subspecific boundaries, we attach to each of these
central populations its satellite groups, without granting any of
these separate nomenclatorial recognition. We agree with Hew-
itt that each of these satellite local populations may be recognized
by modal tendencies, though probably not by any constant
feature, but we also think that, without extreme artificiality, it
will never be possil)le to set limits to these populations. To ac-

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA

313

cept or erect names for these local groups would be to multiply
boundaries that, by reason of intergradation, are vague at every
level.

We have chosen to recognize only three somewhat arbitrary
ranges which are based on the convenient character of plastral
pattern. All populations possessed of a plastral pattern that is
sharply bounded, but indented or ])roken through by a series of

Fig. 30. Axillaries iu various species of Psammobatea. A, P. geometricus
(M.aZ. 32184) ; B, P. oculif.er (M.C.Z. 42158) ; C, P. t. verroxii (M.C.Z.
42224).

(P. Washer del.)

yellow rays, Ave assign to trimeni. All populations possessed of a
plastral pattern sharply bounded, but very little indented or
quite intact, we refer to teniorius. All populations with a plas-
tral pattern that is both indistinct and variable, we group under
the name verroxii A. Smith which antedates schonlandi Hewitt.
The population around Matjesfontein we cannot allocate since it
comes from an area of intergradation between all three forms.

314 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

We have felt that the complex geographical variation of these
forms was a phenomenon meriting rather full discussion and,
indeed, further investigation. It may be noted that during this
discussion we have arrived at substantially the same taxonomic
allocations proposed by Duerden in 1907. It is hoped that by
dealing with the problem of geographic variation more elabo-
rately than he did, the conclusions are placed on a firmer footing.

Key to the Species and Suhspecies of Psammobates

1. Plastral pattern dark brown or yellow brown, highly ornate, covering

entire plastral area; anterior and posterior margins of carapace tj'pi-
cally very sharply serrat«, even spinose; nuchal large, usually sub-
triangular; axillary single, normally fused with humeral; forelimb
with a few large scales on its anterior surface. Range: north of the
Orange River, viz. Southwest Africa; Bechuanaland ; Transvaal;

Orange Free State ; and Griqualand West in Cape Province

ocuUfei- (Kuhl) (p. 315)
Plastral pattern, if covering entire plastral area, black on yellow, other-
wise restricted to central area of plastron or absent; anterior and
posterior margins of carapace typically 7iot sharply serrate, never
spinose; nuchal, if present, smaller, either elongate or broad; axillaries
1 or 2, not united with the humeral; forelimb Avith variable scaling
anteriorly 2

2. Plastral pattern never absent, black on yellow; margins of the plastral

shields, even laterally and on the bridge, either wholly black or black
broken by yellow rays; nuchal very variable in size but typically
elongate, occasionally absent; axillary 1; forelimb anteriorly with a
few large scales which are not in contact but separated by smaller
ones. Range: extreme southwest Cape Province within a 100 mile
radius north and east of Cape Town, .geometricus (Linnaeus) (p. 319)
Plastral pattern sometimes absent, otherwise black or brown and yellow,
peripherally a more or less broad zone of yellow that is usually entirely
devoid of markings; nuchal very variable in size but typically broader
than long, often minute, rarely absent; axillaries 2 or 3, rarely 1;
forelimb anteriorly with large juxtaposed scales, not separated by
smaller ones 3

3. Plastral pattern, when present, without a well defined outline; dorsal

shields never strongly conical. Range: Orange Free State (possibly)
and northern Cape Province north through Southwest Africa, occurring
mostly at elevations above 3000 feet. . . .t. rerroxii (A. Smith) (p. 324)
Plastral pattern with a well defined outline; dorsal shields more or less
strongly conical ; occurring mostly at elevations below 3000 feet .... 4

LOVERIDGE AND WILLIAMS: AFRICAN CBYPTODIRA 315

4. Plastral pattern broken through by light rays and/or its outline indented
by a series of encroachments of the pale ground color. Range: Cape
Province (Little Namaqualand south to near Touw's Eiver) north to

Southwest Africa (Keetmanshoop, the only record)

t. trimeni (Boulengcr) (p. 333)
Plastral pattern without light rays crossing and with few or no indenta-
tions. Range: Cape Province (mostly south of 33° parallel) from
vicinity of Fish River west to beyond Uniondale and Willowmore

t. tentorius (Bell) (p. 336)

PSAMMOBATES OCULIFER (Kuhl)

1820.

Testudo oculifera Kuhl, Beitr. Zool. Anat., p. 77: Cape.

1831c.

Gray, 22.

1865.

Strauch, 61.

1889a.

Boulenger (part), 165 (omit specimen Richmond to Victoria

West).

1890.

Strauch, 54.

1893a.

Boettger, 10.

1897.

Siebenroek, 247.

1898.

Sclater, W. L., 97.

1898.

Siebenroek, 425.

1899.

Siebenroek, 566.

1901.

Lampe, 192.

1902a.

Werner, 341.

1904c.

Siebenroek, 307.

1907a.

Duerden, 9.

1907b.

Duerden, 76, pi. viii, fig. 12.

1907c.

Duerden, 196.

1909a.

Siebenroek, 524.

1910.

Siebenroek, 704.

1910a.

Werner, 301.

1911.

Lampe, 147.

1911b.

Stemfeld, 409.

1911d.

Sternfeld, 49, fig. 61.

1912b.

Werner, 434, pi. — .

1914a.

Hewitt, 247.

1929.

Flower, 31.

1931.

Hewitt, 499.

1931.

Power, 44.

1932b.

Power, 467.

1934a.

Mertens and MiiUer in Rust, 9.

1935b.

FitzSimons, 304.

316 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1937b. Mertens, 5.

1939a. Miiller, L., 129, pi. xi, fig. 7.

1950. WUliams, 550, 551.

1955a. Mertens, 34, pi. iii, figs. 7-8.

1839. Testudo semiserrata A. Smith, 111. Zool. S. Africa, Kept., pi. vi aa:

' ' between Latakoo and the Tropic of Capricorn. ' '
1844. Gray, 8.

1851. Dunieril and DumerU, 3.

1855. Gray, 9.

1880c. Vaillant, 25.

1886b. Boulenger, 542.

1887b. Boettger, 138.

1844. Emys oculifera Gray, 28.
1862a. Clemmys oculifera Strauch, 32.
1870c. Peltastes semiserratiis Gray, 656.
1870e. Gray, 9.

1872c. Gray, 5.

1873b. Gray, 10.

1893a. Homopus signatus Boettger (part), 8 (juv. ex. " Namaland " ) •
1894a. Boettger, 88.

1894. I'leck, 83.

1933b. Chersinella oculifera Hewitt, 263.
1934. Hewitt, 337.

1935a. FitzSimons, 520.

1937a. Psammohates oculifera (sic) Hewitt, 791, fig. 2.

1938. Testudo geometrica Schepers (not of Linnaeus), 535-555, figs. 1-5.

1939. Schepers, 451-495, 1-8.
1948a. Schepers, 9-11, figs. 1-4.
1948b. Schepers, 1-212, figs. 1-250.

A further citation of '' oculifera" will be found under Psam-
mohates tentorius verroxii.

Common names. Serrated Tortoise (preferred) ; Toothed Cape
Tortoise (Gray:1844) ; Kulil's Tortoise (Flower :1929).

Illustrations. Excellent figures (lateral and plastral views)
copied as our Plate 7 are furnished by Sir Andrew Smith
(1839 :pl. vi), who contrasts the nuchal and forelimb (figs, aa)
with those of geometricus (figs. bb). Siebenrock (1910) shows
the carapace and plastron of a hatchling. Sternfeld (1911d:fig.
61) provides a sketch of the buttock tubercle. Werner (1912b)
has a colored plate of both adult and juvenile.

Types. The principal discussions on the status of this species
and its synonym have been by Strauch (1865; 1890), Boulenger

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 317

(1889a), Duerden (1907c), and Siel)enrock ( 1940c ; 1910).

Description. Beak more or less strongly hooked, tricuspid,
edge of jaws more or less serrate ; i)refrontal divided longitudi-
iiall}' or broken up ; frontal broken up ; upper head shields small,
irregular ; forelimb anteriorly with a few unequal (one extremely
large) tubercular scutes (but area distal to the largest formed of
small scales), forming 2-4 longitudinal and 4-6 transverse series
from elbow to outer claw ; claws 5 ; hinder side of thigh with a
large, and occasionally a few subsidiary, subconical tubercles;
heel with or without ^ conical, spurlike tubercles ; tail in both
sexes devoid of tubercles on sides and without a terminal claw-
like tubercle.

Fig. 31. Skull of Psammobates ooulifer (A.M.N.H. 7094). Condylobasal
length 23 mm.

(S. McDowell del.)

Carapace convex, sides descending abruptly, shallowly notched
in nuchal region, anterior and posterior margins more or less
expanded, sometimes reverted, and strongly (rarely weakly)
serrated ; dorsal shields concentrically striated ; nuchal present,
usually large, subtriangular, more or less projecting into nuchal
notch; vertebrals 5, rarely 4, 6, or 7,^ more or less swollen,

1 Both present and absent in United States National Museum material.

2 Four in a tortoise from Kooa iflde Werner :1910a) ; 6 in Keetmanshoop
specimen; 7 in one (U.S.N.M. 63021) from Bechuanaland.

318 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

typically not conical, broader than long, about as broad as, or
broader than, or narrower than, the costals ; costals 4, not form-
ing an angle with the lateral marginals; marginals 11, rarely
10 or 12 ; supracaudal undivided, incurved in 5 S , downwardly
directed in 9 $ .

Front lobe of plastron anteriorly more or less produced and
openly notched ; gulars paired ; pectorals narrow where they
meet, sometimes even separated,^ their anterior border sloping
towards the axillary notch ; axillary 1, almost always fused with
the humeral; inguinal moderate, meeting femoral;- hind lobe
deeply notched posteriorly.

Plastral formula: Abd>h>(g, f, an)>p; g usually>an,
though sometimes=or<.

Color. Carapace bro^vnish yellow or horn color { $ ), with a
very variable pattern of dark brown or black rays of more or
less equal width, numbering 6-10 on the vertebrals and costals.
Plastron yellowish with light or dark brown (black, iide Gray:
1844) rays.

For lengthy discussion of ontogenetic changes in coloration see
Siebenrock (1910) ; for detailed individual variations Hewitt
(1933b) and FitzSimons (1935b).

Size. Carapace length of a 5 (T.M. 15999) from near Kim-
berly, 118 mm., breadth 82 mm., height 61 mm. Carapace length
of a 9 (T.M. 15998) taken with above, 133 mm., breadth 99 mm.,
height 70 mm. (FitzSimons :1935a).

Sexual dimorphism. Siebenrock (1910) states that in 5 5
the vertebrals are flatter than in the 9 9 , but our meagre ma-
terial scarcely bears this out. However, Siebenrock is correct
in stating that the 2 carapace is relatively higher than that of
the $ . Unlike most species, according to Hewitt (1933b), there
is no disparity in size between the sexes.

Breeding. On November 29, 1933, a pair were engaged in
courtship on the Kimberly to Schmidtsdrif t road, the $ scuttling
around the 5 , butting against her shell, and periodically emit-
ting short, low, grunting coughs. Though approached closelj'
by FitzSimons (1935a), the $ only ceased his activities when
actually picked up.

1 As in the type and a Waterberg tortoise (M.C.Z. 4215S).

2 Except in a Quickbom specimen mentioned by Hewitt (1931).

LOVERTDOE AND WILLIAMS : AFRICAN CRYPTODIRA 310

The egg of a Kimberly tortoise measured 31 x 39.5 mm
(Hewitt :1934).

Enemies. Jackals, ratels and the brown hyena will all break
open the plastron to get at these tortoises (R. D. Bradfield in
Hewitt :1934) ; Bataleur Eagles also prey upon them according
to the Hereros (Hewitt :1934).

Habitat. Frequents sandy plains and grassgrown fiats (Power :
1931; 1932b) ; grassland savannas of the inland plateaus accord-
ing to Bradfield (in Hewitt :1933b).

Localities. Bechuanaland: Gemsbok; *Gomodimo; *Kalahari;
Kamelslip; Kaotwe ; Kooa; Lake Ngami; Lehututu (as Lekutu)
to Kang; Makarikari; Mookane (Moocanc) ; Palapye Road;
*Serowe (U.S.N.M.) ; Vlei Topan. Transvaal: Jamestown; Lino-
kana. Orange Free State: Bothaville ; Modder River. Cape Prov-
ince: as "Cape" (type locality) ; Danielskuil (Daniel's Kuil) ;
*Kimberly; Kuruman ; "Latakoo to Tropic of Capricorn" (type
locality of semiserrata) ; Niekerkshoop (Niekerk's Hope) ;
Schmidtsdrift to Kimberly; Saint Clair; Douglas; Warrenton;
Witput Siding; Zoetvlei near Vryburg. Soiithwest Africa:
Aukeikas, 25 km. west of Windhoek; Aus (Anas; Oas) ; *Etosha
Pan-near; *Gaucha (U.S.N.M.) ; Gibeon; Gobabis; Grootf ontein ;
*Nama (U.S.N.M.) ; Ogosongomingo to Great Waterberg; Oma-
ruru ; Ondonga ; Quickborn Farm, Okahandja ; Rehoboth ; Riet-
mond ; Sandup ; Swakopmund; * Waterberg; Windhoek (Wind-
huk).

Range. Extreme western Transvaal and western Orange Free
State northwest through Bechuanaland and Southwest Africa
almost to the frontier of Angola.

Clanwilliara, we believe, was not the true source of the speci-
men but the donor's address cited by Duerden (1907a). It is
probably responsible for Little Namaqualand being included in
the range by FitzSimons (1935b :305) and Mertens (1955:35).

PSAMMOBATES GEOMETRICUS (LinuaCUs)

1734. Testudo minor, amboinensis Seba, Eerum jSTaturalium Thesauri . . .

2. p. 130, pi. Ixxx, fig. 8: "Amboina" (in error).
1758. Testudo geometrica Linnaeus, Syst. Nat., ed. 10, 1, p. 199: "Asia"

(in error).
1766. Linnaeus, 353.

320

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1784.

Daubenton, 628.

1788.

LacepSde, 157, pi. ix and Synopsis.

1789.

Bonnaterre, 24, pi. vi, fig. 1.

1792.

Schoepff, 55, pi. x.

1793.

Schoepff, 49, pi. x.

1797.

Shaw and Nodder, pi. ccevi.

1802b.

Daudin, 260, pi. xxv, fig. 1.

1802.

Shaw, 20, pi. ii.

1812.

Schweigger, 325, 424 (omit non-African localities)

1814.

Sehweigger, 56.

1828.

Bory de Saint Vincent, 73 (omit Asia occidentale)

1831.

Grifilth, 10, 54.

1831b.

Gray, 5.

1831c.

Gray, 12.

1835.

DumerU and Bibron, 57.

1835.

Temminck and Schlegel (part), 73.

1836.

Bell, text, col. pi.

1838.

Ouvier, pi. i.

1844.

Gray, 8.

1845b.

Eiippell, 297.

1849.

Smith, A., App., 1, pi. vi, bb only.

1855.

Gray (part), 8.

1857.

Jan, 35.

1862a.

Strauch, 74.

1865.

Strauch, 19.

1867a.

Steindachner, 3.

1872b.

Gray, 3, in Sowerby and Lear, pi. xiii.

1882a.

Miiller, F., 165.

1882a.

Peters, 3.

1884a.

Eochebrune, 12 (in error).

1886b.

Boulenger, 541.

1889a.

Boulenger, 162.

1890.

Strauch, 56.

1893a.

Boettger, 10.

1893.

Trimen, 79.

1896.

Lonnberg, 11.

1898.

Jeude, 4.

1898.

Johnston, 361.

1898.

Sclater, W. L., 96.

1901.

Lampe, 192.

1904c.

Siebenrock, 309 (key).

1907c.

Duerden, 195.

1909a.

Siebenrock, 524.

LO^'ERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 321

1910. Siebenroek, 702.

1911. Lampe, 146.
1911d. Sternfeld, 49.
1914a. Hewitt, 247.
1929. Flower, 30.
1929. Rose, 189.

1934a. Martens and Miiller in Rust, 9.

1938. Bernstein, 327 (det. doubtful).

1939a. Miiller, L., 129.

1954. Noel-Hume, 74.

1767. Tortue Knorr, 127, pi. iii, fig. 3 (clearly geometricus from this fine

colored figure).
1802b. Testudo luteola Daudin, Hist. Nat. Rept., 2. p. 277, pi. xxv, fig. 3:

(purchased in Dieppe) No locality.
1820. Chersine geometrica Merrem, 32.
1829. Gravenhorst, 20.

1843. Psammobates geometricus Fitzinger, 29.
1861. Fitzinger, 411.

1844. Testudo geometrica var. tentoria Gray (not of Bell), 8.
1855. Gray, 8.

1869c. Peltastes geographicus Gray (lapsus: fide Gray), 173.

1870c. Peltastes geometricus Gray, 655, pi. xiii.

1870e. Gray, 9.

1873b. Gray, 8.

1893. Testudo Strauchi Lidth de Jeude, Notes Leyden Mus., 15, p. 312,

pi. ix: Cape of Good Hope.
1907e. Duerden, 201.

1909a. Siebenroek, 523.

1934a. Mertens and Miiller in Rust, 9.

1933b. Chersinella geometrica Hewitt, 260, pi. xiv, figs. 1-7.
1934. Hewitt, 336.

1933b. Chersinella strauchi Hewitt, 262.

Further citations of "geometrica" will be found under ocu-
lifer, t. tentorius and Kinixys h. helliana.

Common Names. Geometric Tortoise (Lacepede) ; seerpootjies
{fide Peers in Hewitt :1934:336).

Illustrations. For a side and plastral view in color cf. Schoepff
(1792 :pl. x). A. Smith (1839 :pl. vi, bb) contrasts the forelimb
and nuchal with those of oculifer. Photographs of variants are
furnished by Hewitt (1933b :pl. xiv, figs. 1-7).

Types. Testudo luteola was synonymized with geometricus by
Gray (1831b), a disposition with which we are in accord. Lidth

322 BULLETIX : MUSEUM OF COMPARATIVE ZOOLOGY

de Jeude separated strauehi (still in Leiden Museum where it
has been studied by one of us — E.E.W.) l)ecause it lacked a
nuchal and on the proportions of its gulars, characters that are
demonstrably inconstant. Tt was synonymized with geometricus
by Duerden (1906), and there seems no justification for its re-
vival by Siebenrock (190r>; 1910) and Hewitt (1933b).

Description. Beak moderately or strongly hooked ; prefrontal
divided longitudinally or broken up ; frontal broken up ; upper
head shields small, irregular ; forelimb anteriorly covered with
a few large, unequal, scattered tubercular scutes, forming 6-7
longitudinal and 7-10 transverse series from elbow to outer claw ;
claws 5 ; hinder side of thigh without large conical tubercles ;
tail in both sexes devoid of tubercles on sides and without a
terminal clawlike tubercle.

Carapace very convex, sides descending abruptly, deeply
notched in nuchal region, posterior margins not or Ijut weakly
expanded, slightly reverted, and but rarerly serrated ; dorsal
shields concentrically striated ; nuchal present, tj-pically elon-
gate, occasionally triangular, sometimes minute, rarely absent,
when present more or less projecting into nuchal notch ; verte-
brals 5, rarely 4 or 6,^ sometimes swollen as truncate pyramids,
much broader than long, as broad as, or broader than, or nar-
rower than, the costals ; costals 4, rarely 5, usually not forming
an appreciable angle with the lateral marginals; marginals 11 or
12 ; supracaudal undivided, more or less incurved in both sexes.

Front lobe of plastron anteriorly truncate, not or but slightly
produced and A^ery shallowly notched ; gulars paired ; pectorals
broadly in contact, their anterior border sloping tow^ards the
axillary notch; axillary 1, small; inguinal moderate, meeting
femoral ; hind lobe deeply notched posteriorly.

Plastral formula : Abd > g > or < h > or = f > or==an > or=p ; f
usually > an.

Color. Carapace with yellow areolae from which radiate yel-
low rays (8-15 on vertebrals, 9-12 on costals, 2-4 on marginals)
separated hy black. Plastron yellow with more or less ill-defined
black rays and bands.

The juvenile coloring is substantially the same as that of the
adult, according to Siebenrock (1910) ; however, hatchlings in

1 Six in figure on plate of Shaw and Xodder (1797).

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 323

the Paris Museum seeu by E. E. AV. possess the luteola patteru
(cf. Plate 8A, B).

Size. Carapace length allegedly attains to 240 mm. (fide
Duerden :1907c) ; one of 135 mm. is mentioned by Strauch
(1862a) with a breadth of 95 mm., and height of 76 mm. Cara-
pace length of type 9 of strauchi was 140 mm.

If correctly identified, a specimen recorded by Daudin (1802b)
far surpasses all recent records. In French inches it is said to
have a length (possibly over the curve ?) of 10" 6"'=283.6 mm.

breadth 8" =208.5 mm-

height 3" 9"'=101.5 mm.

Breeding. Twelve to fifteen eggs are laid according to Bruyere
(Lacepede:1788).

Enemies. In all probability mankind is responsible for the
extermination, or near extermination, of this handsome little
species. "Writing of his journey into Caffraria in 1772, the
botanist C. P. Thunberg made the following observations.
' ' Instead of China- vessels and calabashes, poverty had taught them
[the Hottentots] to use the shells of the tortoises, which fre-
quent the bushes in the sandy plains, particularly the Testudo
minuta and geometrica" (1795:1:182).
''The Testudo Geometrica was known here (vicinity of the
Cape) by the appellation of the Syren (Syrentie). This land-
tortoise, which probably is the most beautiful of its kind, was
found very common in the sandy downs among the bushes.
The shells of such as were very small, and consequently the
most beautiful, were used for making snuff boxes" (1795:1:
243).
'•Some of the women . . . had a tortoise-shell hanging at their
backs, in which they preserved either their tobacco or bucku
(diosma). For want of clay tobacco-pipes they use wooden
ones" (1795:1:194).
''Some of them [Kaffirs] wore about their necks a necklace made
of small shells, called serpents slmlls (Cypraea moneta) strung
upon a string, and to this hung a tortoise-shell, for keeping
the bukku ointment in " ( 1795 :2:82) .
"Among their [Hottentots] ointments they mix the powder of a
strong smelling herb, which they call Bucku (a species of
Diosma, frequently the Pulehella), and which gives them so

324 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

disagreeable, so fetid, and at the same time so rank an
odour, that I sometimes could not bear the smell of the Hotten-
tots that drove my wagon ' ' ( 1795 :2:187 ) .

Localities. Cape Province : Cape Peninsula ; Ceres ; Darling ;
Eendekuil (as Eendekind) district; Klapmuts; Moorreesburg
district; Porterville district; Riebeeck Kasteel. (Williston —
as Amandelbom Mission on Zak River — we reject on the grounds
that it is too far west, and because other species recorded from
the mission lead us to suspect it was merely a collecting center
for shells brought in by pupils).

Range. Cape Province (Tulbagh, Paarl and Malmesbury dis-
tricts of the extreme southwest, in former times possibly extend-
ing to the Cape Peninsula).

Reported in error from Ascension Id. and Madagascar (Lace-
pede:1788); Mauritius (Gray:1855); Mozambique (Bianconi:
1851); Nyasaland (Johnston :1898) ; Great Namaqualand
(Sternf eld :1911b) and Senegambia (Rochebrune :1884a).

PSAMMOBATES TENTORTU'S VERROXII (Smith)

1839, Testudo verroxii A. Smith, 111. Zool. S. Africa, Eept., pi. viii: Xear
the sources of the Orange Eiver (probably in Cape Province,
rather than in Basutoland where it has not been taken. See re-
marks under Type).

1844. Gray, 9.

1855. Gray, 8.

1909a. Siebenrock, 525.

1910. Siebenrock, 710 (but Transvaal locality erroneous).

1910a. Werner, 302, pi. x, figs. 15a-b, 16a-b.

1911d. Sternfeld, 50, fig. 62.

1915. Werner, 330,

1934a. Mertens and Miiller in Eust, 9.

1939a. Muller, L., 129, pis. x-xi,

1870c. Peltastes verreauxii (editor's emendation) Gray, 656.

1870e. Peltastes verroxii Gray, 9.

1872c. Gray, 5,

1873b. Gray, 10,

1884a. Testudo verreauxii Eochebrune (in error), 13,

1886b. Boulenger, 541.

1887b. Boettger, 139.

1889. Boettger, 282.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA

325

1889a. Boulenger, 163.

1892. Muller, 214.

1893a. Boettger, 10.

1895. Oudemans, 323.

190-lc. Siebenrock, 313, pi. iv and v, fig. 5.

1907c. Duerden, 198,

1911. Lampe, 146.

1914a. Hewitt, 247.

1932b. Power, 466.

1933a. Power, 211.

1886b. Testudo smithi Boulenger, Proe. Zool. Soc. London, p. 542 : South

Africa.

1889. Boettger, 285.

1889a. Boulenger, 165, pi. iv.

1892. Miiller, 215.

1893a. Boettger, 10.

1894a. Boettger, 88.

1894. Fleck, 83.

1898. Sclater, W. L., 97.

1904c. Siebenrock, 318, pi. iii.

1904b. Tornier, 304, figs. 12, 14, 15.

1907c. Duerden, 200.

1909a. Siebenrock, 526.

1910. Siebenrock, 712.

1911. Lampe, 147.
19111). Sternfeld, 410.
1911d. Sternfeld, 50.
1915. Werner, 330.

1934a. Mertens and Muller in East, 9.

1886b. Test ado fisTci Boulenger, Proc. Zool. Soc. London, p. 542, col. pi.

Iviii: De Aar, near Hopetown, Cape Province.

1889a. Boulenger, 165.

1898. Sclater, W. L., 97.

1904c. Siebenrock, 322, pi. v, fig. 0.

1907a. Duerden, 10.

1907c. Duerden, 200.

1909a. Siebenrock, 527.

1910. Siebenrock, 716.

1929. Flower, 31.

1933a. Power, 214.

1934a. Mertens and Miiller in Rust, 9.

1939a. Muller, L., 129, pi. ii, fig. 1.
1889. Testudo tentoria Boettger (not of Bell), 284.

.'i26 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1893a. Boettger, 10.

1894a.. Boettger, 88.

189-4. Fleck, 83.

1903e. Boulenger, 217.

1889a. Tcstudo oculifera Boulenger (part: not of Kuhl), 166 (specimen ex

Eichmond to Victoria West).
1903e. Testudo Seimundi Boulenger, Ann. Mag. Nat. Hist. (7), 12, p. 216,

pi. xvii: Three miles east of Deelfontein, Cape Province.
1907c. Duerden, 201.

1909a. Siebenrock, 527.

1934a. Mertens and Miiller in Eust, 9.

1904b. Testudo boettgeri Siebenrock (not of Mojsiso\acs), Anz. Akad.

Wiss. Wien, 41, p. 194: Great Namaqualand, Southwest Africa.
1904c. Siebenrock, 310, pis. i-ii.

1907c. Duerden, 202.

1909a. Siebenrock, 525.

1911d. Sternfeld, 49.

1922a. Mertens, 168.

1906. Eomopus hergeri Lindhohn,! Jahrb. Ver. Nat. Wiesbaden, 59, p.

348: Gibeon, Southwest Africa.
1907b. TesUhdo tentoria var. fisMi Duerden (part), 88 (fig. is of tentorm

trimeni).
1909b. Testudo bergeri Siebenrock, 623.

1910. Siebenrock, 713, pis. ii and iv.
1910a. Werner, 304, pi. ix, figs. 14a-c.

1911. Lampe, 146.
1911d. Sternfeld, 50.

1929. Testudo oscarboettgeri Liudholm, Zool. Anz., 81, p. 295; n.n. for

boettgeri Siebenrock, preoccupied.
1934a. Mertens and Miiller in Rust, 9.

1934. Chersinella schonlandi Hewitt, Ann. Natal Mus., 7, p. 303, pi. xvi,

figs. 40-50 : Little Namaqualand, Cape Province.
1934. Chersinella fishi Hewitt, 309, pi. xvi, figs. 51-56.
1934. Chersinella fislci s.eimiindi Hewitt, 314, pi. xvi, fig. 57.
1934. Chersinella fislci cronwrighti Hewitt, Ann. Natal Mus., 7, \t. 317.

pi. xvi, fig. 58: Hanover, Cape Province.
1934. Chersinella fislci orangensis Hewitt, Ann. Natal Mus., 7, p. 319, pi.

xvi, figs. 59-60: Between Phillipstown and Petrusville District,

Cape Province.
1934. Chersinella fisl'i colesbei-gensis Hewitt, Ann. Natal Mus., 7, p. 321,

pi. xvi, figs. 61-62: Colesberg, Cape Province.

1 Krroneoiisly synonymizcd with Homopus boulcngeri by Siebenrock (lltOH : .")!.=)),
w lio later (11)101 correctly assigned it to the geometriciis groiip.

LOVEBIDGE AND WIJ.I.IAMS: AI-RICAN CBYPTODIRA 327

1934. ChcrffineUa fiski grica Hewitt, Ann. Natal Mus., 7. p. 323, pi. xvi,

figs. 63-66; pi. xvii, figs. 67-71: Marydale, Prioska District, Cape

I'rovince.
1934. Chersinella fi.sl:i gricoides Hewitt, Ann. Natal Mus., 7, p. 326, pi.

xvii, figs. 72-75: Nickerks Hope (= Niekerkshoop), Hay District,

Cape Province.
1934. Chersinella verroxii Hewitt, 328, pi. xvii, fig. 78.
1937a. FitzSimons, 260.

1934. Chersinella verroxii smithl Hewitt, 331, pi. xvii, figs. 76-77.
1934. Chersinella verroxii boettgeri Hewitt, 333.
1934. Chersinella verroxii amasensis Hewitt, Ann. Natal Mus., 7, p. 333:

Ukamas District, Cape Province.
1934. Chersinella verroxii bcrgeri Hewitt, 335.
1935a. Chersinella boettgeri FitzSimons, 520.
1935a. Chersinella fiskii ? FitzSimons, 520.
1937e. Psammobates fiski eolesbergensis Hewitt, 8, pi. iii, fig. 6.
1937c. Psavimobates fi-ski eronivrighti Hewitt, 9, pi. iii, fig. 5.
1938. Psammobates depressa FitzSimons, Ann. Transvaal Mus., 19, p. 154,

pi. ii, figs. 1-4; pi. iii, figs. 1-2: Aus, Southwest Africa.
1938. Psammobates fiskii FitzSimons, 155.
1946a. Psammobates fiskii fiskii FitzSimons, 354.
1955a. Testiido verroxii bergeri Mertens, 37, pi. iv, fig. 12.

Further citations of ''verreauxi" will be found under t. tento-
riiis, and "'fiski" under t. trimeni.

Common name. Northern Tent-Tortoise (would seem to be the
most descriptive).

Illustrations. Excellent figures (lateral and plastral views;
nuchal and forelimb) of the type have been furnished by Smith
(1839 :pl. viii). Siebenrock (1904c :pls. i-v) supplies good photo-
graphs of verroxii, smithi, fi,ski and boettgeri (type), and Hewitt
(1934 :pls. xvi-xvii) of the types of his numerous subspecies.
More recently Miiller (1939a :pls. x-xi) has photographed many
extremes of this highly variable form.

Types. The type of verroxii has been definitely identified by
FitzSimons (1937a :260) as a stuffed specimen, now minus eleven
shields, in the Roj'al Scottish Museum at Edinburgh,

As to the type locality of verroxii, Power states (1932b :466)
that he has systematically searched Sir Andrew Smith's works
from which ''it appears that he did a great deal of collecting
in that part of the country at present occupied by the divisions
of Bethulie, Smithfield, and Rouxville." Power concludes that

328 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

the type ''was taken somewhere north of Aliwal North, between
the Orange and Caledon Rivers ; that is to say, roughly 260 miles
east of Niekerk's Hope."

Dr. V. FitzSimons, writing us on 12.V.54, considers Power's
suggestion regarding the type locality as reasonable, adding that
he has never seen any verroxii in the eastern Orange Free State.
Mr. C. Jacot-Guillarmod, a keen naturalist whom we also con-
sulted, replied that the few tortoises seen by him in the Orange
Free State appeared to be referable to Homopns.

Writing on l.ix.54 from Mamathes, Basutoland, Mr. Jacot-
Guillarmod says : ' ' I looked up Andrew Smith 's itinerary and
find that the type locality of the tortoise cannot be far from
here. I should say just north of here round about Levibe, Fieks-
burg, or even east of here, perhaps along the Phuthiatsana River
as he went up this river to its source. This means that he passed
within four or five miles of Mamathes."

However, though "the sources of the Orange River" are in
Basutoland, where Mr. Jacot-Guillarmod has been residing for
the past 36 years, he tells us that during all that time he has
never seen a tortoise in the Protectorate. Even the leopard tor-
toise can no longer be found, and if verroxii ever did occur it
has presumably been exterminated for tortoises are highly prized
by the witch-doctors.

HcAvitt (1933b) believes that verroxii may be distinguished
from fi^ki by the presence of a zone of small scales on the anterior
aspect of the front foot immediately above the claws, instead
of the large scales which he says continue to the base of the
claws in fiski. We have been unable to verify whether this key
character holds.

In all probability^ Boulenger (1886b) was induced to describe
fiski by his overlooking Smith's (1839) mention of the presence
of a conical tubercle on the hinder side of the thigh in verroxii,
and also by the uncertainty as to the latter 's type locality (which
is possibly much nearer to that of fiski than has hitherto been
supposed). Boulenger never saw a specimen of verroxii and,
as recently as April 6, 1954, Mr. J. C. Battersby assures us that
there are no representatives of the species in the British Museum,
i.e. all their material is referred to fiski. When at the British
Museum in 1953, one of us (E. E. W.) studied the types of

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 329

fiski, sciniundi and smithi, and shortly afterwards at Sencken-
berg the type of hoett(jeri=^oscarboettgeri. Paratypes of schon-
landi (M.C.Z. 42222-4) have been available to us also.

Description. Beak weakly or strongly hooked, tricuspid, edge
of jaws finely dentate ; prefrontal and frontal broken up ; upper
head shields small, irregular ; f orelimb anteriorly covered with
a few extremely large (in some instances a single shield is ex-
ceptionally enlarged as in oculifer), unequal, juxtaposed, subim-
bricate scutes, forming 2-4 longitudinal and 5-9 transverse series

Fig. 32. Skull of Psammolates t. verroxii juv. (M.C.Z. 21330). Condylo-
basal length 15 mm. The youth of the specimen is reflected in the very
open sutures.

(S. McDowell del.)

from elbow to outer claw ; claws 5 ; hinder side of thigh with one
or more enlarged, conical tubercles -^ heel with a large spurlike
tubercle ; claws 4, rarely 5.

Carapace more or less convex, sides rounded, descending
abruptly, very shallowly notched in nuchal region, anterior and
posterior margins more or less expanded, sometimes reverted,
and not or but slightly serrated; dorsal shields concentrically
striated ; nuchal usually present, typically broader than long
or slightly longer than broad, rarely divided,^ frequently minute,
rarely absent ; vertebrals 5, rarely 4^ or 6,"* more or less swollen,

1 In stating "without" Boulenger (1880a) misread Smith's description.

2 In type of dcpressa FitzSimons.

3 Fide Power (1933a).

* In type of seimmidi as well as other specimens.

330 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

much broader than long, as broad as, broader than, or narrower
than, the eostals ; costaLs 4, rarely 5,^ usually not forming an
angle with the lateral marginals ; marginals 11 or 12 ; supracau-
dal undivided, incurved in $ S , inclined to spread out like
marginals in 9 9; tail in S S usually devoid of lateral tuber-
cles, if present, small, in 9 9 large lateral tubercles present,
both sexes without a terminal clawlike tubercle.

Front lobe of plastron usually anteriorly truncate, occasionally
somewhat produced,- openly notched ; gulars paired, rarely
subdivided ;^ pectorals more or less narrow where they meet,
their anterior border sloping more or less steeply towards the
axillary notch ; axillaries 2 or 3, rarely 1, one large and the
others, if present, small; inguinal moderate to large, sometimes
with 1 or 2 supplementary shields, meeting femoral ; hind lobe
more or less deeply notched posteriorly.

Plastral formula: Abd>h>an>or<g>or=f >or<p. Aiml
typically longer than g or f ; p usually shortest.

Color. Carapace light to brownish yellow, the yellow on each
dorsal shield usually subequal in width to the 5 or 6 black rays
of which the anterior and posterior pairs on vertebrals and
costals meet their fellows to form three series of ocelli. At times
the black rays may coalesce so that black predominates; mar-
ginals with 1 to 3 black rays. Plastron yellow, sometimes uni-
form, though normally its median portion is variegated with
dark brown.

The plastral pattern in the type of smitJii approaches that of
tentorius trimeni. Separation of stnithi and hoettgeri from
verroxii on the grounds that their light rays are yellow (instead'
of red and yellow as in lyerroxii) appears unjustifiable.

The type S of seimundi had a yellowish horn-colored carapace
from the centre of whose shields radiated short or interrupted,
blackish-brown striae ; the ])lastron was similarly colored with
mere traces of the dark radiating striae. The head showed a
few dark l)rown specks, otherwise the soft parts were yellow.

The type $ of H. hergeri, based on a native huchu-hox shell,
was described as having a carapace that was uniform bright
reddish horn-brown except for the bright yelloAvish areolae.

1 In a Ciipe (ovtoisc. fide Siclii'iirock.

- In type of scimiindi.

3 Fide Power ( 1".»;'>21)) . wlioin sec for ji very deluiled study of rcrrouii.

LOVERIDGE AND WII,[JA.MS: AFRICAN CRYPTODIRA 331

Plastron horn-yellow, its central portion washed with bright red-
dish brown and a few obscure brownish rays. However, the
specimen from Upiugton, Gordonia, reported on by Siebenrock
(1910), had a chestnut brown carapace, each shield edged with
darker of varying intensity while the center of the areolae
tended to be lighter. Plastron light green, its central portion
brownish. Head aud liml)s gray brown; outer scales on the
forelimbs beautifully tinted with light brown. A specimen from
Barby (now in the British Museum), which we have seen, cor-
responds ver}'- closely to Siebenrock 's description. The Herero-
land hergeri mentioned by Werner (1910a) had: Carapace red
brown, uniform. Plastron yellow. Femoral and anal region
brown. Sternfeld (1911b) concluded that hergeri was nothing
more than a color variant.

In the type $ of depressa the head is described as being dirty
3'ellow with more or less symmetrical blackish margins, viz. a
bar down the front of the snout, a spot on top of the snout, a
large spot above each orbit, and a crossbar on the back of the
head. For full description of the shell cf. FitzSimons (1938).

Size. The type S of verroxii was only 93.1 mm. ; the type S
of depressa, 109 mm., its width 82 mm., breadth 50 mm. Cara-
pace length of largest S (ex Niekerk's Hope), 117.5 mm.; that
of the largest 5 (ex Ukamas), 141 mm., width 102.5 mm.,
height 74 mm. {fide Hewitt :1934)

Power (1933a) supplies data to show that S S average
.smaller than $ 9 , which bears out the importance of limiting
comparisons to individuals of the same sex. Siebenrock 's (1910)
suggestion, accepted by Power, that there is a size and habitus
difference justifying the separation of fiski from verroxii, is not
supported by the available data.

Breeding. Females collected in April and May held only
unripe ova. In October there was a single enlarged ovum be-
sides several smaller ones. In December the oviduct contained
a single shelled egg, besides several smaller ova. From this
Power (1932b) deduces that a single egg is laid each year, and
that this does not hatch until the following spring.

Diet. In the intestines of tortoises from Niekerkshoop one
of the Crassulaeeae was most frequently present, ])esides frag-
ments of quartz. Eaeh of these formed the core of a pellet of

332 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

vegetable matter averag:ing about 12 x 18 mm. in size; such
pellets were surprisingly numerous, no fewer than twenty-six
being present in one tortoise (Power :1932b).

Feeds on the Mesemhryanthemum bushes beneath which these
tortoises shelter at Aus (FitzSimons, writing of hoettgeri:
1935a).

Parasites. Every tortoise in the large series collected at
Niekerkshoop by Power (1932b) was infested with spirillum
ticks {Ornithodoros moubata), especially about the neck where,
in some instances, they formed solid masses.

Thread worms of several species referable to the Oxyuridae,
were invariably present in the large intestine at its junction
with the small intestine.

Habitat. South of the Orange River verroxii was found on the
flats by Power (1932b), but at Niekerkshoop north of the river
verroxii was restricted to koppies and oculifer to the sandy
plains.

Localities. Orange Free State: "near the sources of the
Orange River" (for type of verroxii, which Power would place
here). Cape Province: Abiam, Clordonia district; Albany dis-
trict; Alweynsfontein ; Amandelbom Mission, Zak River; Beau-
fort AVest; Brandvlei; Britstown; Britstown to Victoria West;
Bros Pan ; Bros Pan to Wyksvlei ; Burghersdorp ; Colesberg ;
Concordia ; De Aar ; Deelfontein ; Gamoep ; Gamoep to Alweyns-
fontein (Alwijnsfontein) ; Graaf Reinet; *PIauover (U.S.N.M.) ;
Hopetown; Houmoed ; Keimoes; Leliefontein (Liliefontein) ;
Little Namaqualand; Marydale; Middleburg; Molopo River —
lower; Naauwpoort; Niekerkshoop (Niekerk's Hope); O'okiep;
Orange River Station; Philipstown to Petrusville; Plaatjiesfon-
tein near Dwaal ; Prieska ; Richmond ; Richmond to Victoria
West; Sea-cow River; Somerset East district; South Gordonia;
Springbok (Springbokfontein) ; Springfield — near; *Stein-
kopf; Ukamas; Upington; Van Wyksvlei; *Victoria West.
Southwest Africa: Augustfelde and Plateau Farms, Aus dis-
trict; Aus (Oas) ; *Barby, Tiraz Mountains (Brit. Mus.) ;
Bethany ; Gibeon ; Great Namaqualand ; Jakalswater ; Karas
Mtns. ; Keetmanshoop ; Keibib ; Keimoes ; Klein Karas Mtns. ;
Klein Windhoek ; Kubub ; Kuibis ; ; Narudas Sud ; Rehoboth.

Range. Possibly southern Orange Free State (for type only),

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 333

northern Cape Province (from Burghersdorp east to Williston
and northwest to O'okiep, northwards through) Southwest
Africa (Great Namaqualand to Rehoboth on the Tropic of Capri-
corn and the solitary 1895 record of Ouderaans for Jakalswater
near Swakopmund).

"We consider erroneous the records of verroxii and smithi from
the Blaauw Mountains, west of the Zoutpansberg range in the
northern Transvaal. Siebenrock (1910) himself thought the data
was questionable, being so remote from the general range of
the species.

PSAMMOBATES TENTORIUS TRIMENI (Boulcuger)
1886h. Trstudo trimrni Boulenger, Proc. Zool. Soc. London, p. 541, col. pi.

Ivii: Mouth of Orange Eiver, Little Namaqualand, Cape Province.
1889. Boettger, 283.

1889a. Boulenger, 163.

1893a. Boettger, 10.

1898. Sclater, W. L., 97.

1904c'. Siebenrock, 320.

1907a. Duerden, 10.

1907c. Duerden, 199.

1909a. Siebenrock, 526.

1910. Siebenrock, 715.

1910a. Werner, 303.

1911d. Sternfeld, 50, fig. 63.

1914a. Hewitt, 247.

1929. Flower, 30.

1934a. Mertens and Miiller in Eust, 9.

1955a. Mertens (part), 36, pi. iv, fig. 13.

1907b. Testudo tentoria var. fishii Duerden (part, not of Boulenger), pi.

viii, fig. 11.
1933b. Chersinella tentoria hexensis Hewitt, Ann. Natal Mus., 7. p. 286,

pi. XV, figs. 39-40: Hex River, Worcester District, Cape Province.
1933b. Chersinella trimeni Hewitt, 287, pi. xv, figs. 41-45.
1934. Hewitt, 337.

1935a. FitzSimons, 521.

1937a. Psammohates trimeni Hewitt, 791, fig. 1.
1938. FitzSimons, 155.

Common name. Western Tent-Tortoise.

Illustrations. That of Boulenger (1886b) is the best, but those
of Duerden (1907b) showing plastral view of alleged "fiskii"
(error), and Hewitt (1933b) illustrating variations, give a good
idea of this race.

384 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Types. The three cotypes in the British Museum have been
studied by one of us (E. E. W.) ; a fourth is preserved in the
South African Museum {fide Sclater :1898).

Description. Beak scarcely or distinctly hooked, bicuspid,
edge of jaws not or but weakly dentate; prefrontal divided lon-
gitudinally or broken up ; frontal broken up ; upper head shields
small, irregular ; forelimb anteriorly covered with a few ex-
tremely large, unequal, juxtaposed scutes, of which one is often
strikingly larger than the others, forming 1-3 longitudinal and
4-6 transverse series from elbow to outer claw ; claws 5 ; hinder
side of thigh wdth or without a never-very-large, flat or subconieal
tubercle ; heel without definite spurlike tubercle ; claws 4 ; tail in
$ $ devoid of lateral tubercles, which are present in 2 9, both
sexes without a terminal clawlike tubercle.

Carapace convex, sides rounded, shallowly to deeply notched
in nuchal region, anterior and posterior margins not expanded,
slightly reverted, and moderately serrated -^ dorsal shields con-
centrically striated ; nuchal moderate, small, minute, or rarely
absent ; vertebrals 5, more or less convex, typically conical,
broader than long, as broad as, or broader than, or narrower
than, the costals; costals 4, forming an angle with the lateral
marginals, which are convex and consequently separated from
the costals by a well-defined longitudinal groove; marginals 11
or 12 ; supracaudal undivided, incurved in $ $ , downwardh"
directed in 2 9 .

Front lobe of plastron anteriorly truncate, openlj- notched ;
gulars paired ; pectorals moderately broad, their anterior border
sloping slightly towards the axillaiy notch ; axillaries 1-2,
moderate ; inguinal moderate to large, sometimes with 1-2 sup-
plementary shields,- meeting femoral ; hind lobe deeply notched
posteriorly.

Plastral formula: Abd>h>or=(p, an)>or=g.

Color. Carapace yellow or orange (occasionally salmon red in
life, iide Hewitt), the light rays on each dorsal shield narrower
than the 4-8 black rays so that black predominates. Plastron yel-

1 In which respect we disagree with Duerden (1907c) -who considered this
species non-serrate.

2 Fide Duerden (1907c) ; we have also seen them with one such supplementary
shield.

LOVERIDGE AXD WII-LTAMS : AKKICAX CHVI'TODIIJ A 33o

low, its central portion occupied by a broad black figure either
indented Avith lighter or variegated with light rays.

Snout yelloAv with a median black line and similar lateral
ones, side of head with a longitudinal suborbital bar terminating
in a roundish postorbital spot ; crown of head black. Cf. Hewitt
( 1933b :290) for color in life of soft parts of Bitterfontein speci-
mens, and FitzSimons (1935a) for the juvenile coloring of a
Lekkersing tortoise that he assigns to trimeni.

Size. Carapace length of largest S (M.C.Z. 33451), 97 mm.,
breadth 75 mm., height 52 mm. Carapace length of largest 9
(T.M. 16002), 121 mm., breadth 95 mm., height 67 mm.

Sexual dimorphism. According to Hewitt (1933b :290), sup-
ported by our $ and 'i , $ $ lack the patch of enlarged scales
on side of tail which are present in the 9 9 , whose tails — as
usual — are shorter.

Hewitt also claims that the foreheads oi 6 i are more
protuberant, and that this is correlated with a broad, down-
wardly projecting beak that bears a w^ell-developed median, and
two lateral, ridges. He asserts that the beak, when pronounced
in 9 9, lacks the sharp ridges. Hewitt also thinks that the
lobes of the plastron converge less strongly in $ $ and that this
results in the gulars and apical distance between the anals being
broader than in the 9 9 .

Breeding. An egg, measuring 24.0 x 34.5 mm., was obtained
from a 9 taken near Bitterfontein (Hewitt :1933b).

Habits. During normal summer weather, according to B.
Peers (in Hewitt :1933b) Trimen's Tent-Tortoises emerge to feed
in the early morning and late afternoon. Upon the approach
oP unfavorable weather they seek shelter beneath some low-
lying bush or euphorbia where they soon bury themselves by
excavating the sand, first with the left forefoot, then with the
right. Once completely covered they remain dormant until
the arrival of cool weather or rain. Following a heavy down-
l)our great numbers of tortoises emerge simultaneously so that
many may be captured in a short time. While the rain is falling,
according to Peers, whose description, however, is none too
clear, the reptile raises its hind legs high so that the grooved
shell, aided by the forelimbs, conducts the water to the corners
of its mouth where it is eagerly taken in.

336 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Habitat. The sand veld (Peers).

Localities. Cape. Province — Little Namaqualand : Anenous —
near; Calvinia — near;^ *Bitterfontein — near; *Lekkersing ;^
Nieuwerust;^ Orange River Mouth ^ (type locality) ; *Steinkopf ;
Van Rhyn 's Pass — near, between Van Rhynsdorp and Nieuw-
houdtville. Southwest Africa — Great Namaqualand : August-
f elde and Plateau Farms, Aus district ; Keetmanshoop ; 70 km.
south of Walfish Bay (W. Triebner coll.).

Range. Extreme western Cape Province (i.e. Little Namaqua-
land) from Lambert's Bay (extending eastwards into Bushman-
land according to Peers,^ though we are unaware of any records
from that area) north to the Orange River, and even occurring
north of the river in Great Namaqualand (see discussion p. 310
above).

The record of trimeni from Kamelslip on the Nosop River,
Bechuanaland, was based on a huchu--pouch. carapace, i.e. a re-
ceptacle for the sweet-scented powder the natives obtain from
the hiicliu plant. As this is the only occurrence of trimeni in
Bechuanaland we reject the record, assuming the shell to have
been carried to Kamelslip by its peripatetic owner.

The Clanwilliam and Stellenbosch records mentioned by Sie-
benrock (1910) as taken from Duerden (1907a), were stated by
the latter to be only the home addresses of the donors.

PSAMMOBATES TENTORIUS-TENTORIUS (Bell)

1828a. Testudo tentoria Bell, Zool. Jour., 3. p. 420: Africa?

1836. Bell, text and col. pi. — .

1872b. Gray, 3, in Sowerby and Lear, pi. xiv.

1886b. Boulenger, 541.

1889. Boettger, 284.

1889a. Boulenger, 164.

1892. Miiller, F., 214.

1893. Trimen, 79.

1897. Siebenrock, 247, pi. i, fig. 4.

1898. Sclater, W. L., 97.
1904c. Siebenrock, 321.

1907a. Duerden, 10 (a few localities are referable to t. verroxii).

1907b. Duerden, 74, pi. vii, fig. 8 ; pi. viii, fig. 10.

1907c. Duerden, 197,

1 In Transvaal Museum, but seen by us.

2 Cited by Hewitt (1933b).

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 337

1909a. Siebenrock, 527 (some of Duerden's t. verroxii localities ac-

cepted).
1914a. Hewitt, 247.

1929. Flower, 31.

1934a. Mertens and Miiller in Rust, 9.

1937a. Flower, 9.

1939a. Muller, L., 129.

1954. Noel-Hume, 76.

1835. Testiido geometrical Temminck and Sclilegel (part: not of Bell), 73.
1855. Testiido geometrica var. nigriventris Gray, Cat. Shield Kept., Brit.

Mus., p. 8 : South Africa.
1870b. PeJtastes tentorius Gray, 9.
1870c. Gray, 656.

1873b. Gray, 9.

1898. Tcstudo verreauxii Sclater (not of Smith), 96 (Beaufort West).
1933b. Chersinella tent&ria Hewitt, 265, pi. xiv, figs. 8-9.
1933b. Chersinella tentoria alhanica Hewitt, Ann. Natal Mus., 7. p. 266,

pi. xiv, figs. 10-15 : Mayfair, Albany District, Cape Province.

1933b. Chersinella tentoria tentorioides Hewitt, Ann. Natal Mus., 7, p. 268,

pi. xiv, fig. 16: Bowden Hall, Middlebury District, Cape Province.

1933b. Chersinella tentoria piscatella Hewitt, Ann. Natal Mus., 7. p. 269,

pi. xiv, figs. 17-18 : Little Fish River, Somerset East District, Cape

Province.

1933b. Chersinella tentoria siibsnlcata Hewitt, Ann. Natal Mus., 7. p. 270:

Brighton Farm, near Steytlerville, Cape Province.
1934. Hewitt, 336.

1933b. Chersinella tentoria Tcaruica Hewitt, Arm. Natal Mus., 7. p. 272, pi.
xiv, figs. 19-24; pi. xv, figs. 25-28: Drogekloof Farm, near Klaar-
stroom, Prince Albert District, Cape Province.
1934. Hewitt, 336.

1933b. Chersinella tentoria duerdeni Hewitt, Ann. Natal Mus., 7, p. 279, pi.

XV, figs. 29-31 : Graaf Reinet, Cape Province.
1933b. Chersinella tentoria lativittata Hewitt, Ann. Natal Mus., 7, p. 281,

pi. XV, figs. 32-34: Willowmore, Cape Province.
1933b. Chersinella tentoria Tiaruella Hewitt, Ann. Natal Mus., 7. p. 283, pi.

XV, figs. 35-38: Uniondale, Cape Province.
1937a. Psammobates tentoria Hewitt, pi. x, fig. 5 (skull).
1937e. Psammohates tentoria alhanica Hewitt, 7, pi. iii, figs. 1-2; pi. xxvii,

fig. 9.
1937e. Psammobates tentoria tentorioides Hewitt, 8; also as "tent tortoise"

skeletal shell, pi. xxviii, fig. 1.
1937e. Psammobates tentoria pisoatella Hewitt, 8, pi. iii, fig. 3; pi. ivA,
figs. 2-3.

338 BULLETIN: iMUSEI'.M OF (O.MPARATIVK ZOOLOGV

1937e. Psammohates tcntoria duerdcni Hewitt, pi. ii, fig. 3.
1946a. Pitammohates tcntoria karruica (sic) FitzSinions, 353.
lOoO. Geometric Tortoise Roso, fig. 206.

Further citations of "tentoria," also tentoria hexensis, will be
found under t. verroxii and t. trimcni.

Common names. Southern Tent-Tortoise (preferred) ; South
African Starred Tortoise (Noel-Hume .-1954) ; knoppiesdop or
Inoppiesskilpad (Afrikaans, fide Hewitt :1937e).

Illustrations. Bell's (1836) fine picture of the type will not
be available to most workers, who w^ill be grateful to Hewitt
(1933b and 1934) for his photographic reproductions of most of
the outstanding variants.

Types. The type of Testudo tentoria Bell in the Oxford
]\Iuseum has been examined by E. E. W.

Description. Beak weakl}^ or strongly hooked, bi- or tri-
cuspid; edge of jaws more or less serrate; prefrontal and
frontal broken up ; upper head shields small, irregular ; forelimb
anteriorly covered with a few, extremely large, unequal, juxta-
posed scutes, forming 2-4 longitudinal and 4-9 transverse series
from elbow to outer claw^ ; claws 5 ; hinder side of thigh with or
without 1 or more, large, subeonical tubercles; heel without
definite spurlike tubercle ; claws 4 ; tail in both sexes devoid of
lateral tubercles and Avithout a terminal clawlike tubercle.

Carapace convex, sides rounded, broadly notched in nuchal
region, anterior margins slightly expanded, posterior some-
times reverted and slighth' serrated ; dorsal shields concentric-
ally striated; nuchal occasionally well developed, usually small,
minute, or even absent; vertebrals 5, rarely 6 or 7,^ usually
strongly conical- and, except for the first, usually broader than
long, narrower than the costals ; costals 4, occasionally 5, some-
times forming an angle wdth the lateral marginals; marginals
11-13 ; supracaudal undivided, incurved in $ S , downwardly
directed in 9 9 .

Front lobe of plastron anteriorlj^ truncate, openly notched;
gulars paired ; pectorals very variable, narrow or relatively
broad, their anterior border curving or sloping towards the

1 Fhlc Iltnvitt (193.3b :271, 275. 277) ; 7 in U.S.N.M. 16225, seen by E.E.W.

- riiH degree of convexity is higlily variable even in tortoises from the sann'
locality, says Duerden (190(C : 189).

LOVERTDGE AND WILLIAMS : AFRICAN CRYPTODIRA 339

axillary iioteh ; axillarics 2 or more, variable in size ; inguinal
moderate to large, sometimes with 1 or 2 supplementary shields,
meeting femoral ; hind lobe deeply notched posteriorly,

Plastral formula: Abd>h>or<g>or<an>or<f>p. Pec-
toral the shortest scute.

Color. Carapace yellow or orange, the light rays on each
dorsal shield usually narrower than, though sometimes subequal
to or broader than, the 8-14 black rays (8-12 on vertebrals, 12-14
on costals, 3-4 on marginals) so that black frequently predom-

Fig. 33. Skull of Psammohates t. tentorius (M.C.Z. 3465). Condylobasal
length 24 mm.

(S. McDowell del.)

inates. Plastron yellow, sometimes faintly tinged with orange,
visible only on the periphery as the central portion is occupied
by a broad brown figure.

The tail and adjacent soft skin are red to orange red (^dc
Hewitt :1933b).

Size. Carapace length of the type of tentorius, 108 mm. Cara-
pace length of largest 6 (ex Matjesfontein), 100 mm., breadth
74 mm., height 74 mm. (FitzSimons :1946a). Carapace length
of largest 5 (a paratype of t. karuella), 138 mm., breadth 107
mm., height 74 mm. (Hewitt :1933b).

Breeding. In September, 2 or 3 eggs are laid. Two oval eggs
of '"alhayiica" measured 21.3 x 27 mm. and 23.5 x 31 mm.
respectively. Two tortoises that hatched out in May measured

340

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

25.4 mm. (given as 1 inch) in length (E. du Toit etc. in Hewitt
1933b and 1937e).

HiDlllllfllll'lHlimill]

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 341

Longevity. Seven years, 5 months, and 6 days for a specimen
i-eceived at the Loudon Zoo in 1920 (Flower :1937a).

Diet. In a wild state the diet includes Eragrostis hergiana
(fide E. du Toit) lettuce and lucerne in captivity (Hewitt:
1937e).

Enemies. In certain areas ostriches, by devouring the young,
have almost exterminated this form (Hewitt :1937e).

Habits. In June not a single tortoise w^as seen at Drogekloof,
though the veld was traversed in every direction daily by Miss
1. Z. Oosthuizen; in September — though there had been no
rain and the veld was even drier — tortoises were encountered
almost every day (Hewitt :1933b).

Tent-Tortoises in captivity develop regular habits, each rep-
tile returning to its favorite corner of the pen after feeding or
sunning. Rather more sluggish and timid than other species, the
forelimbs, when withdrawn, completely conceal both the head
and all surrounding skin (Hewitt :1937e).

Habitat. In the Albany district tentorius is found among low
bushes in the karroid vegetation on either side of the Great
Fish River (Hewitt :1937e).

Localities. Cape Province (but note remarks under Range) :
Abraham's Kraal, Prince Albert district; *Adendorp; *Beaufort
West ; Bowdon Hall, Middleburg district ; Brandeston Farm,
Albany district; Brighton Farm, Steytlerville ; Calitzdorp (Cal-
itsdorp) ; Carlisle Bridge; Dikkop Flats between Grahamstown
and Port Elizabeth ; Drogelkloof Farm near Klaarstroom ; *Fish
River near Grahamstown; Graaff Reinet; * Grahamstown; Klip-
plaat ; Koup ; Let jesbosch, between Beaufort West and Frazer-
burg Road; Little Fish River; Matjesfontein ; Mayfair, Albany
district; Middleton, Carlisle district; Muiskraal; Nelspoort;
Oudtshoorn; Prince Albert; Steytlerville; LTniondale; *Upper
Karroo; *Warmbad Farm, Uuiondale (seen at U.S.N.M.) ; Wel-
bedacht Farm, Olifants River, Oudtshoorn district; Weston-
dale, Pearston district; Willowmore; Zwartruggens (Swart Rug-
gens), Aberdeen district.

Range. Southeast Cape Province, mostly south of the 32nd
parallel. Typicallj^ concentrated in the region from Grahams-
town to Uniondale. Apparently intergrading wdth Psammohates
tentorius verroxii at Adendorp ; Graff Reinet ; Middleburgh dis-

342 BULLETIN: MUSEUM OF COMPAEATIVE ZOOLOGY

trict (which is north of the 32nd parallel) ; Nelspoort and Beau-
fort West. At Matjesfontein and vicinity, intergrading with
both P. t. verroxii and P. t. trimeni may be expected. Errone-
ously recorded from Mauritius by Gray (1873b).

Genus CheRSINA Gray

1831c. Chersina Gray (not of Humphreys: 1797),i Syn. Kept., pp. 7, 14,

Type: Testudo angulata Sctmeigger (by monotypy).
1929. Gonioohersiis Lindholm, Zool. Anz., 81, p. 285. Type: Testudo anpu-

lata Schweigger (by original designation).
1931. Ncotestudo Hewitt, Ann. Natal Mus., 6. p. 504. Type: Testudo
angulata Scliweigger (by original designation).

Definition. Skull with triturating surface of maxilla without
ridging but a weak ridge on the horny sheath; median pre-
maxillary ridge absent ; maxillary not entering roof of palate ;
anterior palatine foramina moderately large, not concealed ;
prootie well exposed dorsally; quadrate enclosing stapes in
adults ; surangular subequal in height to prearticular ; neck with
second, third or fourth centrum biconvex.

Carapace never hinged ; normally the anterior neurals hexa-
gonal ; outer side of third costal scute about as long as, or longer
than, that of the fourth ; submarginal scute absent ; suprapygal 1.

Plastron not hinged; gular region greatly thickened and pro-
duced; gular shield single, much longer than broad.

Distinguishing marks. The only continental African tortoise
with a single gular scute.

Range. Union of South Africa (Cape Province only), possibly
extending into Southwest Africa.

Fossil record. None.

Remarks. We have separated the peculiar species angulatus
from the genus Testudo on osteological grounds. An external
character, the single gular, indeed suffices to set it apart from
all the tortoises of continental Africa, but this condition is
paralleled by the tortoises of Mauritius and Rodriguez and by
the Malagasy species ynipJiora, all of which we refer to the genus
Geochelone. From the Geochelone species with single gulars,
Chersina differs in the very extensive thickening of the gular
region, in which as in some other features it resembles Ki^iixys.

1 See footnote •': on page lilS.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA

343

Not all of its features are resemblances to Kinixys, however, and
its true affinities are doubtful.

To illustrate the problem ami our method of meeting it we
tabulate (Table 6) the characters of Chersina and cite for
each of them the genera presenting the closest similarity.

Table 6

Chersina

Genera showing greatest
resemblance

/

Comment

Maxilla with a very
feeble ridge or none

Quadrate closed bo-
hind stapes in ad-
ults

Anterior palatine
foramina moderate
to large

Neurals variable
approaching the oc
tagonal and quad-
rilateral pattern

tiiiprapygal single

Gular process
greatly tliickened
and extended an-
teroposteriorly

No ridge in. Kinixys, Eomopiis,
Pyxis, Psammobatcs ; reduced
or absent in Testiido

An advanced char-
acter in which C. is
intermediate

Kinixys, IIoviopus, Psammo- Au advanced char-

hat.es, Tcstudo s. str., Malaco- acter achieved in

chersus, most Geoehelone, and parallel by many

Gopherus and fStylemys genera

Test udo

An advanced char-
acter in which C. is
somewhat interme-
diate

Anterior marginals Kinixys
elongate

A specialized char-
acter shared only
with K.

Pyxis, Testudo, Geoehelone, Go-
pherus

An advanced char-
acter which C. ap-
pears to be ap-
proaching Inde-
pendently

Kinixys, Pyxis, Acinixys, Psam-
mobates, Testudo, Malacocher-

S118

A primitive char-
acter shared with
emydines

Kinixys

A specialized char-
acter shared only
with K.

344

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

From this tabulation it will be seen tbat except in neural
pattern (Kinixys is obstinately hexagonal) and in the single
gular, Chersina is either close to Kinixys or more primitive than
that genus. If it were to be synonymized with any genus it
would seem more reasonable to equate it with Kinixys than with
Testudo or GeocJielone. But the differences from Kinixys
seem to us to imply a long separate history, and we are not
certain that the relationship to Kinixys is as great as emphasis
upon two specialized characters would suggest. We prefer
therefore to regard this form as a full genus belonging to the
series of Ethiopian endemics.

Atlantic Ocean ^

Fig. 35. Distribution of Chersina angulaia in South Africa.

(P. Washer del.)

The distribution of this tortoise is wholly included within the
range of Psanimohates, possibly within that of Honiopus.

There appears to be no clearly demarcated geographical varia-
tion within angulafa; as to red on the plastron, see paragraph
on Color below (p. 348).

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA

345

Chersina angulata (Schweigger)

1795. Testudo pusilla Thunberg, (not Linnaeus, 1758), 3.

1812. Testudo angulata Schweiggor, Konigsberger Arch. Naturwiss. Math.,

1, pp. 321, 360, 443 : No locality.

1814. Schweigger, 52 (reprint of 1812).

1835. Dumeril and Bibron, 130 (not 150).

1835. Temminck and Schlegel ( ? part erosa), 72.

1836. Bell, text and col. pi. — .
1845b. Kuppell, 297.

1857, Jan, 35.

1872b. Gray, 5, in Sowerby and Lear, pi. xix.

1889. Boettger, 286.

1889a. Boulenger, 178.

1889. Vaillant, 166, pi. xv, figs. A-B.

1890. Strauch, 57.
1892. Miiller, F., 214.
1893a. Boettger, 12.
1898. Jeude, 5.
1898. Johnston, 361.
1898. Selater, W. L., 97.
1901. Lampe, 194.

1907a. Duerden, 9-10 (only donor's addresses).

1907b. Duerden, 73, pi. vi, fig. 4.

1909a. Siebenrock, 543.

1910a. Werner, 305.

1911d. Sternfeld, 50.

1912b. Werner, 434.

1915. Werner, 331.

1925b. Flower, 932.

1928. Cott, 952.

1929. Flower, 30.

1929. Rose, 188, 225, 231, fig. 125.

1935a. FitzSimons, 519.

1937a. Flower. 9.

1937. Ruckes, 103, pi. x, fig. 4.

1950. Williams, 551.

1812. Testudo tahulata Africana Schweigger (not of Hermann), Konigs-
berger Arch. Naturwiss. Math., p. 322: No locality.

1814. Schweigger, 54 (reprint of 1812).

1820. Chersine pusilla Merrem (not of Linnaeus: 1758), 38.

1946a. FitzSimons, 354.

346 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1828. Tcstudo Bellii Gray, Spieilegia Zoologica. p. 2, pi. iv, figs. 3 3a:

Cape of Good Hope.
1831c. Chersina angulata Gray, 15, pis. i-ii.

1831L.

Gray, 5.

1844.

Gray, 11.

1849.

Smith, A., App., 1.

1855.

Gray, 12.

I860.

Fitzinger, 411.

1862a.

Strauch, 23.

1865.

Strauch, 36.

1866.

Gray (part), 306.

1867a.

Stemdachner, 4.

1870e.

Gray (part), 13.

1873b.

(jray, 14.

1873c.

Gray, 726, pi. Ix, fig. 6.

187311.

Gray, 490.

1885.

Smets, 2 (locality erroneous).

1887h.

Boettger, 137.

1887a.

Miillcr, F., 296.

1931.

Neotestudo angulata Hewitt, 504.

1934a. Goniochersus angulatus Mertens and Miiller in Eust, 63.

1955a. Mertens, 34, pi. iii, fig. 14.

1937a. Chersine angulata Hewitt, 789, pi. x, fig. 7.

1937e. Hewitt, 6, pi. i, fig. 3 ; pi. iv, fig. 4 ; pi. xxvii, fig. 10.

1938. FitzSimons, 153.

1950. Rose, 325, 343, figs. 207-208.

1946. Testudo (Chersine) angulata Cairncross, 396.

Synonymy. Sehweigger (1812) took the name Tcstudo angu-
lata from a specimen labeled by Dumeril in the Paris Museum ;
authorship was later claimed by Dumeril and Bibron (1835:
130 and 138). T. tahulata africana Sclnveigger (not of Her-
mann) was also based on a Paris Museum tortoise. Strauch
( 1862a :(i7) has explained why Testudo pusilla Linnaeus (1758:
199) is not identical with pusilla Linnaeus (1766:353). The
latter, being preoccupied, cannot be used, as was done by Fitz-
Simons (1946a :354:). FitzSimons (1938) suggested that Cher-
siua Gray 1831, having been proposed by Humphreys for a gastro-
l^od in 1797, is not available, hence his use of Chersine Merrem
(,see however, footnote 3, p. 218).

Common names. Angulated Tortoise (Hewitt :1937e) ; Bow-
sprit Tortoise (Flower :1929) ; bont-skilpad ; duine-skilpad ;

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIKA

;]-(■

ploegskaar-skilpad ; rooi-skilpad ; skaar-skilpad (Afrikaans, all
Hewitt :1937e).

Illustrations. There are excellent figures in Gray (1831 :pls.
i-ii), and Bell (1836), the latter reproduced in Sowerby and
Lear (1872:pl. xix).

Description. Beak weakly or strongly hooked, bi- or tricuspid,
edge of jaws very weakly serrate; prefrontal divided longi-
tudinally, sometimes separated from the frontal by a small scale ;
frontal large, or occasionally divided longitudinally,^ flanked
by 4-5 small scales; upper head shields small, irregular; fore-
limb, more especially anteroventrally, with several rows of mod-
erately large, subequal, juxtaposed or slightly separated scutes
forming 2-3 longitudinal and 7-9 transverse series from elbow

Fig. 36. Skull of Chersina anaulata (B.M. 67-4-2-152). Condylobasal
length 28 mm.

(S. McDoweU del.)

(on the lower or inuer side of which is an isolated large scute)
to outer claw; on the upper or outer aspect of the wrist is a
semi-bracelet of large scutes; claws 5; hinder side of thigh
without large tubercles; heel with one or more enlarged flat
scales; tail without terminal clawlike tubercle.

Carapace elongate, moderately convex, sides descending
abruptly, deeply notched in nuchal region, anterior margins ex-
panded but neither reverted nor serrated, posterior margins not
or but moderately expanded, reverted, somewhat sinuate; dorsal

1 Diviflod in ;i Steinkopf spt>oim<>n. fide Werntr (1910a).

348 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

shields concentrically striated except in aged specimens, neither
swollen nor convex ; nuchal narrow, elongate, always small,
sometimes absent ;^ vertebrals 5, rarely 6 or 7,^ flat, the first about
as broad as long or longer than broad, the second, third and
fourth much broader than long, the fifth broader than long or
longer than broad, the third narrower than the third costals;
costals 4, rarely 5,^ usually not forming an angle with the
marginals; marginals 11, rarely 10;^ supracaudal undivided,
more or less incurved in 5 5 , downwardly directed in 2 2 .

Front lobe of plastron always strongly produced, its end
truncate or rounded ; gular entire f pectorals narrow where they
meet, their anterior border sloping steeply towards the axillary
notch in adults, less so in young; axillaries 1 or 2 (single in
young, double in adults) moderate; inguinal large, meeting
femoral ; hind lobe deeply notched posteriorly in $ $ , more
shallowly in ? 2 .

Plastral formula, adults: Abd>g>h>or<an>f>or==p; ju-
veniles: Abd>h>g>an>p>f.

Color. In both juvenile and adult the carapace is pale straw
or pale horn to olive, the areola of each vertebral and costal
shield usually darker and the periphery broadly margined with
black, these borders occasionally interrupted by lighter rays; a
broad-based, black, triangular blotch on the sutures between
marginals; supracaudal with a median, down-pointing, black
triangular mark. Sometimes, however, the pattern may be
broken up, indistinct.

Plastron pale straw to yellow ; from gular to anals extends a
broad, black blotch, uniform, variegated with lighter, or
broken up, its borders not sharply defined; lower aspect of
lateral marginals yellow, but those posterior to the inguinal tend
to display a black blotch at their anterior edge, and there may be
a similar spot on one of the anterior marginals.

1 Absent in a Richtersveld tortoise (M.C.Z. 42211).

2 Six in a Great Namaqualand buchiu pouch, fide Boettger (1SS9) ; 7 in a Stein-
knpf hatchling (M.C.Z. 42210).

3 Five in a Great Namaqualand biicliu pouch, fi.(le Boettger (1889).

4 Ten on right side of a Steinkopf hatchling (M.C.Z. 42210).

5 Semidivided on the lower surface of an adult ^ from Penrock, fide Hewitt
(1931; see also Hewitt :1937a :790). In Rose's figure (1950:343) the gular is
called a "subgular", while the humerals are miscalled "gulars" and the pectorals
"humerals."

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA

349

In life the plastron is said to be reddish by Bell (1836) but
the color disappears soon after death. However, Duerden

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350 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

(1907b :74), seemingly suggesting that it is a local characteristic,
I'emarks that angulaia from tlie western part of Cape Province
possess plastrons having a diffusely reddish ground color. Fitz-
Simons (1935a) states that the sides of the plastron are suf-
fused with reddish in adults, especiallj- so in $ $ from Van
Rhj'nsdorp, and later (1938) remarks on the bright rosj'- red
plastrons of Kamieskroon and Klipfontein tortoises.

Snout at tip, and hinder part of head, black; prefrontal and
frontal shields j-ellow (Hewitt :1937e). The coloring of the
soft parts as given by Gray (1866:306, etc.) was based on the
head and limbs of a Geoemyda piinctularia associated with the
shell of an angulata (fide Boulenger :1889a :178).

Size. Carapace length of largest $ , 264 mm., breadth 59 mm.
( Hewitt :1937e), length of another $ (M.C.Z. 3998), 183 mm.,
breadth 112 mm., height 80 mm. Carapace length of largest $
(T.M. 15996), 163 mm., breadth 104 mm., height 81 mm. (from
Soebartsfontein, FitzSimons :1935a) ; length of a Port Elizabeth
5 (M.C.Z. 9328) 153 mm., breadth 103 mm., height 68 mm.

Weight. The largest $ listed above weighed 5 lbs. (Hewitt:
1937e). See also Cairncross (1946:397) for tabulated increases
among hatchlings.

Sexual dimorphism. The rather spadelike projection of the
front lobe of the plastron is much more strongly produced in
i S (whose plastron is also somewhat concave in its posterior
third) than in $ 9 (whose tails are noticeably shorter than
those of the S S ). These sexual differences have been well
illustrated in black and white by Vaillant (1889 :pl. xv=our
Plate 12).

Breeding. In August, at Cape Town, the 2 digs a nest hole
of from 3 to 4 inches in depth; only a single egg is laid (Rose:
1950). However, two eggs (M.C.Z. 21662) were laid on IS.viii.
1925 at the M.C.Z. by a Port Elizabeth tortoise (M.C.Z. 22475).
Eggs, though usually oval, are sometimes spherical, rarely
pointed at one end. In size they range from 35 x 37 mm. to
34 X 42 mm. (Hewitt :1937e :pl. xxvii, fig. 10).

While the incubation period at Cape Town may range from
12 to 14 months (Rose:1950), Cairncross mentions a 30-grain
egg laid on 7.V.40 which, placed in an incubator on 3.viii.40,
hatched on 3.xi.40, i.e. in 6 months; however, the hatchling was

LOVERIDGE AXD WILLIAMS : AFRICAN CRYPTODIRA 351

malformed and died 14.X.41. For o-rowtli and weight records
of otlier hatchlings, see Cairncross (1946:397). The hatehling's
carapace is flat and strikingly different from the convex shell
of the adult (Hewitt :1937e).

Rose managed to protect the egg from rats and other predators
by disinterring it and then reburying it at the same depth of soil
within a wire basket sunk in the ground and securely covered
over by wire netting.

Longevity. Eleven years, nine months in Giza Zoo (Flower:
1925b; 1937a).

Diet. Eats gazanias (A. Rothmann) ; fond of young beans and
regarded as a nuisance by gardeners (Hewitt :1937e) ; a captive
specimen displayed a liking for meat (Rose:1950) but this
author's somewhat contradictory statement that true tortoises
are "normally strictly vegetarian" should have the second
adjective omitted.

Enemies. Unquestionably man. Writing from IMrs. Miiller's
farm, apparently a day's trek northeast of Cape Town, on Sep-
tember 11, 1773, Thunberg (1795:3) says: "Among the bushes
in the sands we frequently saw land-tortoises crawling, and the
young ladies in the house had ordered the slaves to bring sev-
eral of them home of various sizes for our repast. The Testudo
piisilla was the most common species here, and it was this which
was now laid upon the fire for our eating. I slipped into the
kitchen on purpose to see the mode of dressing it, and found that
the girls were cruel enough to lay the poor animal wide open
on the live coals, where, sprawling with its head and feet, it was
broiled alive, till at length it burst to pieces with the heat. The
eggs, which were in great number, and consisted of yolk only,
were the most luscious and desirable part of it."

The 21 young tortoises, with unbroken shells about tw^o inches
in diameter, recovered from the stomach of a 5 Secretary Bird
{Sagittarius serpentarius) killed on the Cape Flats by Levail-
lant (1824, Hist. Nat. Oiseaux d'Afrique, 1:70, pi. xxv), were
probably referable to ayigulata suggests Rose (1950:344).

Teynperament. Males spend much time in sparring with their
rivals (Hewitt :1937e). One S, by biting the heads of other
c$ ^ in the enclosure, induced them to withdraw into their shells.
whereupon he would thrust his nose or plastral projection be-

352 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

neath their shells and so overturn them. Occasionally; he would
achieve the same result by butting them head on, then walk away
leaving his opponents sprawling on their backs (Rose:1950).
Females do not respond readily to a male's advances, in fact
they usually endeavor to escape him by running away (Cairn-
cross : 1946).

Habitat. The drier coastal and central districts of Cape
Province as far eastwards as the Keiskama (Hewitt :1937a).
Still occurs on the Cape Flats, but is more often met with on
the less-frequented uplands towards Cape Point (Rose:1950).

Localities. Cape Province :^ Adelaide ; *Cape of Good Hope
Cape Flats; Cape Point; Cradock; Dassen Island (introduced)
Fish River Valley ; Grahamstown ; Graaff Reinet ; Kamieskroon
Keiskama Valley ; Kenkelbosch ; Klipfontein ; Malmesbury ; Mat-
jesfontein ; Mortimer ; Penrock near Grahamstown ; *Port Eliza-
beth ; Queenstown ; *Richtersveld ; Lekkersing ; Rhynheath ;
Schoombie ; Soebatsfontein ; *Steinkopf ; Stillbay ; Van Rhyns-
dorp ; Welbedacht; Worcester; Zoetendahl's Valley. Southwest
Africa:^ Bethany (based on buchu pouches collected by Schinz,
etc. Boettger: 1887b); Karibib (Shell :Mertens :1955a) ; Kuibis
(shell lacking anterior lobe of plastron, so possibly native pre-
served. Werner :1915) ; Orange River mouth (2 live 9 5, coll.
G. May) ; *Pomona (A.M.N.H., alcoholic) ; Rietmond near Gib-
eon (Lampe:1901).

Range. Cape Province west of 28° East longitude, possibly
extending northwards into Southwest Africa.

Erroneously recorded from West Africa and Madagascar by
Bell (1836) ; from Natal by Jan (1857) ; from Matabeleland and
Zambezi region by Smets (1885) ; and the East Indies by Bur-
meister (not seen).

Genus HOMOPUS Dumeril and Bibron

1835. Homopits Dumeril and Bibion, Erpet. Gen., 2, p. 145. Type: Testudo
areolata Thunberg (designation by Fitzinger: 1843: 29).

1835. Chersobius Fitzrnger, Ann. Wiener Mus., 1. pp. 108, 112, 122. Type:
Testudo signata Schoepff (by original designation).

1 In an annual report Duerden (1907a) lists angiilata from 14 donors, whose
addresses he gives, but these we omit as there is no way of telling whether the
specimens actually came from the places mentioned.

2 The records from Southwest Africa should be investigated, a statement with
which Mertens (1955a) concurs.

LOVERIDQE AND WILLIAMS : AFRICAN CRYPTODIRA 353

1931. Pseudomopus Hewitt, Ann. Natal Mus., 6, pp. 496, 498. Type:
Testudo signata Schoepff (by original designation).
Definition. Skull with triturating surfaces of maxilla and pre-
maxilla without ridges ; maxillary not entering roof of palate ;
anterior palatine foramina moderate to large, conspicuous ; proo-
tic narrowly exposed dorsally ; quadrate enclosing stapes; sur-
angular subequal in height to the prearticular ; neck with third
centrum biconvex.

Carapace never hinged; normally the anterior neurals hex-
agonal or quadrate ; outer side of third costal scute longer than
that of fourth ; submarginal scute absent ; suprapj^gals 2, the
anterior larger, bifurcating posteriorly to embrace the smaller
posterior element which is adjacent to, but not crossed by, the
sulcus between the fifth vertebral and the supracaudal.

Plastron not hinged ; gular region slightly thickened and pro-
duced; gular shields paired, usually broader than long.

Range. Union of South Africa (western Orange Free State
and Cape Province), possibly extending into Southwest Africa.
Fossil record. None.

Ilomopiis alone of the present day Ethiopian endemic tortoise
genera has been alleged to have a fossil record. In 1889 Lydekker
(Cat. Foss. Kept. Brit. Mus., part 3, p. 91 seq.) assigned jEmys
comptoni Bell, of the London Clay Eocene of England, and jEmys
scutella V. Meyer, from the Oeningen Upper Miocene of Switzer-
land, to Homopus. Bell as Lydekker points out, when describing
■]-E. comptoni, indicated resemblances to H. areolatus and H. sig-
natus and was probably only deterred from referring the species
to Homopus by the fact that the London Clay is a marine de-
posit.

Since Lydekker 's time little attention has been paid to these
fossils. Hewitt (1937a :792) justly remarked: "It is interesting
to note that flattened shells of similar form and size to Homopus
did occur so long ago in Europe, but the generic identity should
be considered very doubtful, as the characters emphasized are
quite insufficient."

Fortunately one of us (E.E.W.) has been able to examine ma-
terial of both of these fossil species — the unique type of jcomp-
toni in the British Museum and two specimens of f scutella in

354 BULLETIN : MUSEI'.M OF COMPARATTVE ZOOLOGY

the Tevlpi" Museum, Haarlem, llollautl. The Teyler Museum geu-
erously permitted the specimeus of ■\scuieUa to be borrowed and
further prepared. Only one side of each had been exposed; skill-
ful preparation by Mr. Arnold Lewis of the Museum of Com-
parative Zoology has now revealed the other side in each speci-
men, despite the extreme rottenness of the bone in places.

Whereas superficial examination of this material had seemed
to support, or at last did not refute, the assignment to Tlomopus.
the more careful study now possible reveals that neither species
can be referred to the South African genus. Instead, on totality
of characters, both appear referable to Tcstudo.

The type specimen of comptoni is defective in various critical
areas, and probably could not be placed with certainty were it
not that '\.scutclla, which it so mm-h resembles, furnishes a guide
to its assignment. The two species are considered to be Tcstudo
rather than Homopus on the following characters :

1. The neural pattern is advanced in both species; already
incipiently octagonal ciun quadrilateral anteriorly in Eocene
■'fcomptoni, and definitely so in Miocene ■'(.scnfcUa.

2. There is a single suprapygal. not a larger first suprapygal
embracing a second suprapygal as in Homopus and Geochelone.

3. The gulars of each are slightly longer than broad.

4. The single inguinal scute (in Eocene -'{coniptoni) is not in
contact with the femoral, i.e. is already more specialized than in
Homopus signatus or JI. boulengcri.

By itself none of these features is conclusive or diagnostic, but
neither does any of them nor any other character positively sup-
port relationship of either species to Homopus, while at least the
presence of the specialized neural pattern in iMiocene scutella
seems to afford a definite negative. There are no grounds for
assignment to any other Ethiopian endemic genus. The single
suprapygal is unknown in Geochelone, the size of which also ren-
ders relationship improbable. On the other hand, all characters
are consistent with Tcstudo scnsu stricto, which is also geo-
graphically probable. While, with some hesitation, we refer these
European fossils to Testudo, it is with full confidence that we
exclude them from the ancestry of Homopus. Thus Homopus,
like all other Ethiopian endemics, is at present quite unknown as
a fossil,

Remarks. Despite certain emydine-like features, Homopm

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 355

does not appear to l)e a primitive genus. Even the seemingly
emj'dine features — relativelj' depressed shell and transversely
developed gulars — are probably as completely secondary as,
according to onr view, are the unridged maxillae. The anterior
neural pattern has departed somewhat from the primitive hex-
agonal towards a nearly quadrate condition, while the pygal ar-
rangement in Homopn>i is an incipient stage of the advanced
pattern. It differs from the typical condition in Geochelone,^
Gophcrus and jiStylemys in that the sulcus between the first verte-
bral and the supracaudal does not cross the middle of the small
second supra])ygal ; instead it coincides with the suture between
supra pygal and pj'gal.

Hewitt (1931) proposed generically separating signal us and
boulengeri from Homopus, under the name of Pseudomopus, but
later discovered that Chersobins Fitzinger (1835) was available.
The differences which Hewitt points out are valid, but we do not
interpret them as deserving generic recognition. To do so would
imply a wider separation and less certainty as to the real affini-
ties of the separated species than, we feel, exists. We regard the
species of Homopus as forming a single closely-knit series within
which are recognizable two species groups : one embracing sig-
natus and boulengeri, the relatively primitive members of the
series, the other comprising the more specialized members —
femoralis and areolatus.

It must be admitted that the skulls of H. boulengeri and areo-
latus (Figs. 40, 41) differ appreciably in many ways, but we do
not feel that these differences exceed the variation that might be
expected within a genus or even within a subgenus. Further-
more, we have not seen skulls of signatus or femoralis and conse-
quently do not know whether skull structure would group the
species in the same way as do external characters such as num-
ber of claws and size of inguinals. No very striking skull differ-
ences between signatus and boulengeri, on the one hand, and
femoralis and areolatus, on the other, have been described by
Hewitt (1937a), but he mentions diff'erences in skull proportions
and other details between femoralis and areolatus. However, ac-
cording to him, there does exist in the carpus a species-group
difference — the distal carpals of signatus and boulengeri under-

1 All subgenera except Eocene ^Hadrianus.

356 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

going fusions, while in femoralis and arcolatus they always re-
main separate.

So far as known, H. signatus and H. houlengeri are allopatric,
signatus mainly occurring in northwest Cape Province with a
range closely similar to that of Psammohates tentorhis irimeni,
but apparently penetrating farther inland on the plateau (cor-
responding to climatic region 3a of Finch and Trewortha :
1943.^ H. houlengeri has its principal range in a long inland zone
of southern Cape Province, mostly at elevations between 2000
and 3000 feet (approximateh^ climatic region 4b of Finch and
Trewortha).-

Recent very surprising evidence (Mertens 1955a, see below p.
362 and p. 365) that both H. signatus and //. houlengeri occur in
southern Southwest Africa complicates this apparently simple
picture. The ranges of and interrelationships of the two forms
in this area are still obscure, however; this is a problem upon
which further work is urgently needed.

The nearest recorded points of approach for the two species
are Clanwdlliam Mtn. (for signatus) and Piquetberg (for houlen-
geri) which are about 50 miles apart. We have not seen either
of these specimens and cannot state whether there is any approx-
imation of characters in these neighboring populations. Else-
where, however, the two forms are so distinct that there seems
little ground for .suspecting intergradation. No other member of
the genus displays the characteristic freckled pattern of signatus,
while houlengeri appears to be devoid of pattern though very
variable as to shade.

Less obvious, though possibly more significant, is the fact that
the normal number of marginals in H. houlengeri is 12 pairs —
unique as a modal condition among testudinids — while signatus
lias the usual 11 pairs.

H. femoralis and areolatus are less distinct, but are said to
occur together at Halesowen in the Cradock District. This area
of sympatry is very limited as compared with the total range of
each species. H. areolatus is essentially coastal, extending from
Cape Town to Alexandria in southern Cape Province, mostly at
low elevations and entering higher ones (above 2000 feet) only
where it is said to coexist with femoralis (for the most part its

1 In Dudley Stamp : 1953.

2 Piquetberg itself is at very low elevation (ca. 500 ft.) but altitudes up to
3000 ft. from which houlengeri may have come, occur in the vicinity.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA

357

range coincides with climatic region 5 of Finch and Trewortha).
On the other hand, fcmoralLs is a creature of the eastern high
l)lateau (roughly climatic region 6 of Finch and Trewortha)
mostly occurring at elevations above 3000 feet.

The morphological characters separating these two species are
not very striking. They differ in the degree of hooking of the
beak, serration of the margin, development of the femoral
tubercle, etc., characters which may be entirely valid but which
it is easy to imagine being bridged by intermediates. Coloration
also differs, but not so impressively as between si gnat us and
houlengeri.

Atlantic Ocean

^ signatus
■ boulengeri
•^ femoralis
O areolotus

Fig. 38. Distribution of the species of Homopus in South Africa.

(P. Washer del.)

We have not seen the specimens from Cradock district identi-
fied as areolatus; it is somewhat astonishing to find areolatus in-
vading the highlands at well over 2000 feet, entering an unusual
ecology at just the point where its closest relative occurs! We
suggest that the alleged coexistence of femoralis and areolatus
in the district be re-investigated. Cradock specimens (seen by
one of us) in the British Museum are clearly femoralis but lack
the serrate margin mentioned hj Boulenger in his original de-

358

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

scription of the type. Thus, though Cradoek is the type locality
of femoralis, it is possible that intergradation between femoralis
and areolatus may occur in the area and just to the south of it.

These same two species — femoralis and areolatus — have also
been reported by FitzSimons (1946a :352-353) as occurring to-
gether at Welbedacht Farm, Oudtshoorn district. This locality
is over 200 miles from any other from which femoralis is known.
As the identification, apparently correct, is based on a single in-
dividual, the possibility' of the tortoise having been transported
by human agency is worth consideration.

Fig. 39. Forelimb scalation in Eomopus. A, H. signatus (M.C.Z. 21329) ;
B, E. boulengeri (M.C.Z. 42231); C, E. femoralis (M.C.Z. 17523); D, E.
areolatus (M.C.Z. 20967).

(P. Washer del.)

Key to the Species of Homopus^

1. Forelimb with 5 claws; inguinal shield single, in contact with the

femoral shield 2

Forelimb with 4 claws; inguinal shields 2-4, small or absent, only the
innermost in contact with the femoral shield 3

2. Carapace ivory white, yellow or yellowish green, heavily overlaid, spotted

or radiated with black ; plastron shaded or streaked with brown ;
posterior margin of carapace in adults usually reverted and serrated ;
marginals 11-12, usually 11; scutes across forelimb in 5-6 rows. Range:

1 For H. bergeri see Psanimohatcs t. verroxii, not a synonym of boulengeri as
WHS oncp siifrsi'xtf'fi-

For H. darlingi and //. iiogueyi spp Kinixys b. belUana and A', b. noguctii re-
spectively.

LOVERIDGE AND \VIL1.1A>'S: AFRICAX CRYPTODIRA ^59

Wostcrn Cape Province to southern Southwest Africa

.sifjnatus (Sehoepff) (p. 359)
Carapace light olive green, pale yellow, dark yellowish, reddish or
blackish brown, uniform or vertebrals edged with brown or black ;
plastron yellow, dirty green or dark olive, uniform or mesially brovm,
or each shield edged with brown ; posterior margin of carapace not
reverted, and not or but slightly serrated; marginals 11-13, usually 12;
scutes across forelirab in 3-5 rows. Eange: Karroo Plain of south-
central Cape Province, South Africa. Also Southwest Africa (see

discussion bnulengeri Duerden (p. 3G2)

3. Carapace olive or brown, uniform or each shield narrowly edged with
black; plastron yellow or greenish, unifonn or each shield blackish
brown anteriorly; posterior margin of carapace more or less reverted
and serrated; beak not or but weakly hooked; usually several small
scales above nostril; a pair of prefrontal shields; buttock with a very
large conical tubercle; heel with conical spurlike tubercle. Range:
Eastern Karroo of Cape Province north to western Orange Free State,

South Africa femoralis Boulenger (p. 365)

Carapace yellowish olive, olive or greenish, each shield reddish brown in
the centre, the margins edged with dark brown or black; plastron
yellowish, mesially brown, rarely uniform; posterior margin of cara-
pace sometimes reverted and not or but slightly serrated ; beak strongly
hooked; no small scales above nostril; prefrontal usually single or
semidivided posteriorly; buttock without a large conical tubercle, at
most a flat and inconspicuous one ; heel without a conical spurlike
tubercle. Range: Cape Province south of 32° S. latitude. South Africa
areolaUts (Thunberg) (p. 367)

HoMOPUs siGNATUs (Schoepff)

1782. Lorica testudinis signatae Walbaum, Chelonogr. SchUdkroten, pp. 71,

120, pi. (which bears an abbreviated form of the name, viz.

' ' Testud. Sign. var. ") : No locality mentioned.
1792. Testudo signata Schoepff, Naturg. Schildkroten, p. 141, pi. xxviii,

figs. 2-3; ed. 2, 1801, Hist. Testud., p. 120, pi. xxviii, figs. 2-3).

No locality.
1812. Schweigger, 319, 442.

1814. Schweigger, 50.

1831b. Gray, 5.

1831c. Gray, 13.

1836. Bell, text and col. pi.

1865. Strauch, 35.

1872b. Sowerby and Lear, pi. xx (omit text by Gray, cf. areolatua).

360

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1802b. Testudo cafra Daudin, Hist. Nat. Eept., 2. p. 291: Kaffraria, Cape

Province, South Africa.
1812. Schweigger, 318.

1814. Schweigger, 49.

1820. Chersine signata Merrem, 30.
1835. Homopus^ signatus Dumeril and Bibron, 152.
1849. Smith, A., App., 1.

1870c. Gray, 13 (error).

1872c. Gray (part), 6.

1873b. Gray (part), 15.

1873g. Gray, 320, pi. xii, fig. 1.

1884a. Rochebrune, 15 (in error).

1888d. Boulenger, 136.

1889a. Boulenger, 148.

1890f. Boulenger, 521.

1898. Sclater, W. L., 96.

1905h. Boulenger, 252.

1906c. Duerden, 408.

1907a. Duerden, 10.

1907b. Duerden, 69, fig., p]. vii, figs. 5-6.

1909a. Siebenrock, 514.

1910a. Werner, 299, pi. ix, figs. 13a-b.

1911d. Sternfeld, 48.

1929. Flower, 27.

1955a. Mertens, 34.

1835. Testudo Chersobius signata Fitzinger, 122.
1835. Testudo Chersobius cafra Fitzinger, 122.
1931. Pseudomopus signatus Hewitt, 498.
1934a. Mertens and Miiller in Rust, 8.

1935. Hewitt, 345.

1935. Pseudoviopus signatus peeisi Hewitt, Rec. Albany Mus., 4, p. 345.

pi. xxxvi: Near Van Rhynsdorp, Klaver District, Cape Province.
1937a. Chersobius signatus Hewitt, 791, fig. 3, pi. x, figs. 1-2.
1938. FitzSimons, 155.

1950. FitzSimons, 253.

1950. Rose, 338.

Further citations of "signatus" will be found under Geoche-
lone p. habcocki, Psammohates oculifer and H. areolatus.

I Homopus, without desiguatcd species, was first proposed liy Dnin^ril and
Bibron in 1S34. We have no information as to whether their 1S.".5 volume ap-
peared prior to Fitzinger (183.5) in which Chersobius was proposed as a subgenus.
However, as Fitzinger himself relinquislied Chrrsobius in favor of Homopus he
apparently conceded priority. The matter is discussed by Lindholm (1929 :2S4 ;
footnote 11).

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 361

Common Names. Speckled Tortoise (Gray :1844) ; Marked Tor-
toise (Gray: 1831b) ; Walbaum's Tortoise (Flower :1928).

Illustrations. Walbaum's black and white figures of carapace
and plastral views are clearly recognizable, Schoepff 's and Bell's
colored plates less characteristic.

Description. Beak not or but weakly hooked ; prefrontal small,
divided longitudinally or broken up ; frontal small or broken up ;
remaining upper head shields small, irregular; forelimb anteri-
orly covered with very large, subequal, juxtaposed, strongly im-
bricate scutes forming 5-6 longitudinal and 6-7 transverse series
from elbow to outer claw; claws 5; hinder side of thigh with a
very large conical tubercle ; claws 4.

Carapace hardly convex, flattened dorsally, sides sloping,
scarcely notched in nuchal region, anterior and posterior margins
somewhat expanded, reverted, and more or less strongly serrated ;
dorsal shields concentrically striated w^ith slightly impressed
areolae, not swollen, separated by deep grooves, a vertebral keel
in the young; nuchal moderate or small, as broad as or broader
than long, sometimes divided ;^ notched anteriorly and occasion-
ally posteriori}" : vertebrals 5, occasionally 6 or 7, the first about
as long as broad, the third to fifth as broad as or broader than
long, the fourth and fifth rather broadly in contact, the third
narrower than or equal to the third costal ; costals 4, sometimes
5, usually not forming an angle with the marginals; marginals
11-12, usually 11 ; supracaudal undivided.

Front lobe of plastron anteriorly truncate, projecting some-
what at corners, not or but slightly produced, not or but scarcely
notched; gulars paired; pectorals moderate, their anterior bor-
der straight ; axillary 1, moderate ; inguinal 1, small or moderate,
in contact with the femoral ; hind lobe broadly notched pos-
teriorly.

Plastral formula : Abd>h>an> (g, i, p, f, variable, subequal).

Color. In both juvenile and adult the carapace is ivory white,
yellow or yellowish green, heavily overlaid, spotted or radiated
with black. Plastron ivory white or yellow, more or less heavily
shaded with brown or radiating brown lines. Head and neck
yellowish, spotted with black above.

Discussed in detail, and extremes figured by Duerden (1907b).

1 Divided in one Steinkopf specimen (M.C.Z. 42218).

.162 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Size. Carapace length of largest 9 (T.M. 22108) 100 mm.,
breadth 71.5 mm., height 44 mm.

Habitat. Fairly numerous on granite koppies 16 miles from
Bitterfontein.

Localities. Cape Province: Bitterfontein; Clanwilliam Moun-
tain; Grootmist; Kaffraria ; Kamaggas; Klipfontein; *Little
Namaqualand ; O'okiep; Springbok (Springbokfontein) ; *Stein-
kopf ; Van Rhynsdorp. Southwest Africa: Keetmanshoop (Ket-
manshoop ; fide Werner :1910a :300) .^

Range. South Africa — western Cape Province (chiefly in
Little Namaqvialand), northwards into southern Southwest
Africa.

Erroneously reported from Great Namaqualand by Boettger
(1893a) through confusion with Psammohates oculifer, also from
Rehoboth (Fleck :1894; repeated by Sternfeld :1911b) ; see Mer-
tens ( 1955a :34). Also in error from Abyssinia and Mauritius
(Gray:1844), and Senegambia (Rochebrune :1884a).

HoMOPUS BOULENGERi Ducrdcn

lOOfic. JJomoims boulengeri Duerden, Eec. Albany Mus., 1. p. '406, pi. xi,
figs. 1, 2 and 5: Willowmore; Aberdeen; and Beaufort West Dis-
tricts, Cape Province, Union of South Africa.
1907a. Duerden, 10.

1907b. Duerden, 67, fig., pi. vi, fig. 1.

1909a. Siebenrock, 515.

1910. Siebenrock, 697, pis. i and iii.

1955a. Mertens, 33.

1934a. Pseudomopus boulengeri Mertens and Miiller in Bust, 8.
1935. Hewitt, 345.

1946a. FitzSimons, 353.

1950. FitzSimons, 253.

1950. Chersobiiis boulengeri Rose, 338.

Synonymy. Homopus hergeri Lindholm was erroneously syn-
onymized with H. boulengeri by Siebenrock (1909a :515) ; actu-
ally it is a synonym of Psammobates tentorius verroxii.

Common names. Donner-weer (local name, since it appears
after thunderstorms: Hewitt).

Illustrations. Duerden (1906c) supplies rather indistinct

I Mertf^ns (19.5oa) conflrms the presence of this species in Southwest Africa but
his specimens are without exact locality.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 363

photos of the plastral and lateral view of a cotype, also figure of
head; later (1907b) a dorso-lateral photograph. Siebenrock
(1910) also presents dorso-lateral and plastron photos, besides
clear pictures of the head, forearm and egg.

Description. Beak not or but moderately or strongly^ hooked,
bicuspid- or tricuspid;^ prefrontal small, divided longitudinally,
usually preceded by a small median and two lateral shields of
nearly the same size as the prefrontals; frontal broken up; re-
maining upper head shields small, irregular; forelimb an-
teriorly covered with extremely large, subequal, juxtaposed.

Fig. -iO. Skull of Eomopus houlengeri (A.M.N.H. 7109). Condylobasal
length L'2 mui.

(S. McDowell del.)

strongly imbricate scutes forming 3-5 longitudinal and 8 trans-
verse series from elbow to outer claw ; claws 5 ; hinder side of
thigh with, or rarely w'ithout, a large conical tubercle ; heel of
S surrounded by smaller tubercles but without definite spurlike
tubercle; claws 4, rarely 5; tail Avithout terminal clawlike
tubercle.

Carapace scarcely convex,' flattened dorsally, sides sloping,
scarcely notched in nuchal region, anterior and posterior margins
expanded, not or but slightly reverted, not or but slightly ser-

1 strongly fide Diierden.

^ In S.M. 49822 from Platpan Farm, near Ann, Southwest Africa.

364 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

rated; dorsal shields concentrically striated, the areolae not
impressed,^ not swollen, separated by deep grooves ; nuchal small,
as long as or slightly longer than broad; vertebrals 5, broader
or much broader than long, as broad as or narrower than the
costals, the fourth and fifth rather broadly in contact; costals
4, not forming an angle with the marginals; marginals 11-13,
usually 12; supracaudal undivided.

Front lobe of plastron anteriorly truncate, projecting some-
what at corners, not or but slightly produced, not or but slightly
notched ; gulars paired ; pectorals moderate, their anterior border
somewhat sloping, widening slightly towards the axillary notch ;
axillary 1, moderate; inguinal 1, moderate or large, in contact
Avith the femoral; hind lobe broadly notched posteriorly.-

Plastral formula : Abd>h> (g, p, f, and an, variable).

Color. Carapace dark reddish or yellowish brown, or light
olive green, uniform (in young) or the vertebral shields edged
with bright brown or black. Plastron yellow, dirty green or dark
olive, uniform or mesially brown, or each shield edged with
brown. Jaws brown, head and limbs light olive green. In life
the naked parts of skin are said to be bright yellow with minute
orange scales.

Size. Carapace length of Duerden's largest cotype 105 mm.
Carapace length of largest $ (T.M. 19177), 108 mm., breadth
74.5 mm., height 48.5 mm. Carapace length of largest 2 (T.M.
19838), 110 mm., breadth 80 mm., height 49.5 mm.

Sexual dhiwrphism. Plastron of S S deeply concave pos-
teriorly, not in 9 9; hind lobe of plastron deeply notched in
S S , more open in 9 9 (Duerden).

The relatively small size of the posterior aperture of the 9
shell opening — 6 mm. long by 15 mm. broad in one specimen
— in relation to the large size of the egg, is referred to by Sieben-
roek (1910) who suggests that during deposition of the egg
significant widening of the aperture must take place. This could
be achieved by a spontaneous mobility of the posterior lobe of
the plastron, which, he says, has been demonstrated to occur in
Pyxis arachnoides (cf. Siebenrock :1906b) and in Testudo s. sir.

Breeding. The egg referred to and figured by Siebenrock (vide

1 Moderately or deeply according to Duerden (1906c :406).

2 111 S.M. 49822 from Plateau Farm, near Aus, Southwest Africa.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 365

supra) measured 22 x 39 mm. The poles were strikingly pointed
and it was hard-shelled when removed from the oviduct. As
there were no other mature eggs, Siebenroek suggests that only-
one is produced at a laying.

Localities. Cape Province : Aberdeen district ; Beaufort West
district ; *Cape Province ; Hoek-onder-Berg Farm between Mon-
tague and Touws River; Matjesf ontein ; Miller Station, Klip-
plaats between Aberdeen and Willowmore; Pearston district;
Piquetberg; Willowmore; Zwart Ruggens, Aberdeen district.
Southwest Africa: near Aus.^

Eange. South Africa — restricted to the Karroo Plain area of
south central Cape Province ; and apparently Southwest Africa
near Aus.^

Though its range is not so restricted as Duerden (1906c)
assumed, he is apparently correct in saying that no other Homo-
pus occurs in its habitat.

HoMOPUS FEMORALis Boulcngcr

188Sg. Homopus femoraUs Boulenger, Proc. Zool. Soc. London, p. 251, pi.

xiv: Cradoek, Cape Province, Union of South Africa.
1889a. Boulenger, 147.

1890f. Boulenger, 521.

1890. Strauch, 58.

1898. Johnston, 361 (in error).

1898. Sclater, W. L., 96,

1906c, Duerden, 407, pi. xi, fig. 6.

1907a. Duerden, 10.

1907b. Duerden, 67, fig., pi. vi, fig. 2, pi. viii, fig. 9.

1909a. Siebenroek, 514.

1929. Flower, 27,

1931. Hewitt, 496, fig. 4a.

1934a. Mertens and Miiller in Eust, 8.

1937a. Hewitt, 788, 791, 795, pi. x, fig. 3.

1937e. Hewitt, 10, pi. ivA.

1946a. FitzSimons, 353.

1950. Kose, 336.

1 Mertens (1955a :33) received a living pair ($, 9) [one of which we have
examined] from Plateau Farm near Aus, Southwest Africa (H. Erni donor). This
.astonishing record is so far removed from the generally accepted range that we
specially asked Dr. Mertens about it. He replied that, though he did not
collect them himself, he has every confidence in the collector and, furthermore,
that during his trip he heard reports of the occurrence of a strongly-flattened,
rock-dwelling tortoise in the southern portion of Southwest Africa.

366 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Common names. Karroo Tortoise or Greater Padloper (Hew-
itt) ; Boulenger's Tortoise (Flower) ; Padlopertjie (Afrikaans,
A. C. Hoffman in litt.).

Illustrations. Boulenger's (1888g) figure of the type is in
color.

Description. Beak not or but weakly hooked, tricuspid, edge
of jaws serrate; a few small scales above the tubular nostrils;
prefrontal large, divided longitudinally; frontal large or broken
up ; remaining upper head shields small, irregular ; forelimb
anteriorly covered with a few extremely large, subequal, juxta-
posed, strongly imbricate, pointed scutes forming 3-4 longi-
tudinal and 7-8 transverse series from elbow to outer claw ; claws
4 ; hinder side of thigh with a very large conical tubercle ; heel
with large, conical, spurlike tubercles; claws 4; tail without
terminal clawlike tubercle.

Carapace somewhat convex, flattened dorsally, sides sloping,
not or but scarcely notched in nuchal region, anterior and pos-
terior margins somewhat expanded, more or less reverted, and
more or less strongly serrated ; dorsal shields concentrically
striated with slightly impressed areolae divided by a weak keel,
not swollen, convex ; nuchal elongate, subtriangular or slightly
longer than broad ; vertebrals 5, the first as long as or longer
than broad, the second to fifth broader than long, the fourth
and fifth somewhat narrowly in contact, vertebrals two and three
narrower than the costals; costals 4, not forming an angle with
the marginals ; marginals 11 ; supracaudal undivided.

Front lobe of plastron anteriorly truncate, not produced,
scarcely notched ; gulars paired ; pectorals verj- narrow, their
anterior border widening abruptly towards the axillary notch ;
axillary 1 ; inguinals 2 or 3, the outermost triangular, the others
transverse, the innermost in contact with the femoral ; hind lobe
broadly notched posteriorly.

Plastral formula: Abd>h>an>or<f>or=g>or=p. Pec-
toral is typically the shortest scute.

Color. Carapace olive or brown, pale or dark, with or without
a dull red or orange tinge, each shield narrowly edged with
black (in young) or uniform (in adult). Plastron yellowish or
greenish, each shield blackish brown anteriorly (in 3''0ung, but
sometimes persists), or uniform (adult). The shielded or tuber-

I.OVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA ;>67

culate soft parts yellow or pale brownish, the naked areas tinged
with salmon or orange. See also discussion in Duerden (1907b:
67).

Size. Carapace length of largest 6 , 133 mm., breadth 102 mm.
Carapace length of largest 9, 157 mm., breadtli 121 mm. (l)oth
Hewitt :1937e, but no localities mentioned).

Sexual dimorphism. The plastron is flat in both sexes, but
the S supracaudal is usually larger and directed downwards.

Breeding. The eggs are oval to nearly spherical, ranging from
25 X 29 mm. to 25 x 33 mm., and quite soft when deposited
( Hewitt :1937e:ll).

Diet. Bulhine caulescens is a favorite food (P. M. Niven in
Hewitt :1937e).

Enemies. See under areolatus.

Hahitat. Grassveld up to 5,500 feet.

Localities. Cape Province: Aliwal North; Barkly West; Coles-
berg; Cradoek ; Dordrecht; Halesowen; Indwe; Lady Grey;
Middleburg ; New Bethesda ; *Philipstown ; Sneeuwberg ; Storm-
berg; Warrenton; AVelbedacht Farm, Oudtshoorn district;^
Wodehouse=Dordrecht. Orange Free State: Bloemfontein-
(specimen in Senekenberg Museum seen by E. E. W.).

Range. South Africa — ea.stern Karroo of Cape Province
north to western Orange Free State.

Erroneously reported from British Central Africa by John-
ston (1898).'

HoMOPUS AREOLATUS (Thunbcrg)

1787. Testudo areolnin Thunberg, Kongl. Vetensk, Akad. Handl., 8, p. 181,

fig.: "India" (in error).

1792. Schneider, 267.

1792. Schoepff, 121, pi. xxiii.

1795. Schoepff, 104, pi. xxiii.

1 802b. Daudin, 287.

1812. Schweigger, 320, 443 (omit localities).

1814. Schweigger, 51.

1831b. Gray, 5.

1831e. Gray, 13.

1 See p. -'J-'tS for comments regarding this record.

2 Dr. A. C. Hoffman, Director of the National Museum in Bloemfontein, sees no
reason to question this record as femoralis is "very plentiful" and "occurs in
practically every district of the Free State." (in litt.)

368

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1835.

1836.

1862a.

1865.

1872b.

1880c.

1887b.

1890.

1890.

1788.

1802b.
1804.

1820.
1829.
1820.

1831c.

1855.

1835.

1844.

1845b.

1849.

1855.

1857.

1860.

1867a.

1873b.

1873c.

1884a.

1885.

1889a.

1890f.

1893a.

1895.

1898.

1898.

1898.

1903e.

1904b

Temminck and Schlegel, 73.

Bell, text and col. pi. — .

Strauch, 92.

Strauch, 35.

Gray, 5, in Sowerby and Lear, pi. xxi.
Vaillant, 34, 88.

Boettger, 140.
Miiller, F., 705.

Strauch (part), 58.

La Vermilion Lacepfede, Hist. nat. Quadrup. ovip. Serpents, 1, p.

166, and Testudo miniata in Synopsis Methodica, a table at end of

same volume ia which binomials are employed: Cape of Good Hope.

Testudo pusilla Daudia (not of Linnaeus), 299.

Testudo (Africana) Hermann, Observ. Zool. Animal. Spec, p. 218:

No locality.
Chersine areolata Merrem, 30.

Gravenhorst, 18.
Chersine tetradactyla Merrem, Vers, Syst. Amphib., p. 32: "India

orientali" (in error).
Testudo areolata var. pallida Gray, Synopsis EeptUium, p. 13 : South
Africa.

Gray, 12.
Homopus areolatus Dumeril and Bibron, 146.

Gray, 10.

Eiippell, 297.

Smith, A., App., 1.

Gray, 11.

Jan, 35 (as Homoeopus) .

Fitzinger, 411.

Steindachner, 4.

Gray, 15.

Gray, 726, pi. Ix, fig. 5.

Eochebrune, 16 (in error).

Smets, 10 (locality erroneous).

Boulenger, 147.

Boulenger, 521.

Boettger, 8.

Oudemans, 321.

Jeude, 5.

Johnston, 361 (in error).

Sclater, W. L., 96.

Boulenger, 217.

Tornier, 209, 303, fig. 11.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 369

1906c. Duerden, 407, pi. xi, fig. 4.

1907a. Duerden, 9.

1907b. Duerden, 68, fig. — , pi. vi, fig. 3.

1907J. Boulenger, 482.

1909a. Siebenrock, 513,

1910. Siebenrock, 695.

1925b. Flower, 924.

1928. Cott, 952.

1929. Flower, 26.

1929. Eose, 188, 231, fig. 124.

1931. Hewitt, 496, figs. 4b-4c.

1934a. Mertens and Miiller in Rust, 8.

1937a. Hewitt, 790, fig. 4, pi. x, fig. 4.

1937e. Hewitt, 9, pi. ii, fig. 4; pi. iv, fig. 2.

1946a. FitzSimons, 352.

1950. Rose, 336, fig. 202.

1950. Williams, 551.

1844. Homopus signatus Gray (not of Schoepff), 10.

1855. Gray, 11.

1872c. Gray (part), 6.

1873b. Gray (part), 15.

1872b. Testudo .ngnatus Gray (not of Schoepff), 5 (text: not pi.).

A further reference to "Testudo areolata" will be found under
Kinixys h. Ijelliana.

Synonymy. Testudo miniata, which Lacepede called "La Ver-
milion" in allusion to the reddish tinge of its protuberant snout,
was first referred to the synonymy of areolatus by Bell (1836).
Testudo africana Hermann is placed in the synonymy of areola-
tus by us on account of its depressed shell, short gulars, and
coloration. Testudo fasciata Daudin (1802b), heretofore gen-
erally regarded as a synonym of areolaiiis, we reject because the
type was stated to have had five claws on either forefoot. It is
also said to have come from Ceylon.

Commow wames. Parrot-beaked Tortoise (Hewitt :1937e) would
appear preferable to either Areolated Tortoise (Gray :1831b),
Thunberg's Tortoise (Flower :1929), or Grooved Tortoise (Rose:
1929), the latter having long been used for Geochelone sulcata.
Padloper in Afrikaans, on account of their tendency to keep
to paths or roads (C. J. Latrobe :1818).

Illustrations. Schoepff 's (1792) plates supply dorsal, plastral
and lateral views in color. Bell's (1836), reproduced in Sow-

■AIQ

BULLETIN: MUSEU.M OK COMPARATIVE ZOOLOGV

erby and Lear (1872), show the entire animal, including its
plastral aspect, in color.

Description. Beak strongly hooked, tricuspid, edge of jaws
anteriorly weakly serrate ; no small scales above tubular nostrils ;
prefrontal large, single or posteriorly semi- or wholly divided
longitudinally ; frontal moderate or broken up ; remaining upper
head shields small, irregular; forelimb anteriorly covered with
extremely large, subequal, juxtaposed, strongly imbricate,
pointed scutes forming 3-4 longitudinal and 6-8 transverse series
from elbow to outer claw ; claw^s 4 ; hinder side of thigh with or
without (irrespective of sex) a slightly enlarged, low and incon-

Fig. 41. Skull of Homopus areolatus (A.M.N.H. 17792). Condylobasal
length 21 mm.

(S. McDowell del.)

spicuous tubercle ; heel without definite spurlike tubercle ; claws
4 ; tail without terminal clawlike tubercle.

Carapace moderately convex, more or less flattened dorsall3^
sides sloping, scarcely notched in nuchal region, anterior margin
not or but slightly expanded, posterior margin not expanded,
sometimes reverted, and not or but scarcely serrated; dorsal
shields concentrically striated, with impressed areolae some-
times showing a longitudinal keel, more or less swollen, separated
bj' grooves ; nuchal variable, elongate, subtriangular or quadrate ;

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA '571

vertebrals 5, rareJy 4/ G- or 8,"' scarcely convex, the first and
fourth longer than broad or (rarely) as long as broad, the sec-
ond, third and fifth broader than long, the fourth and fifth
only narrowly in contact, the third as broad as, though
usually narrower than, the third costal ; costals 4, rarely 5,*
not forming an angle with the marginals; marginals 11,
rarely 10, 12^ or 18; supracaudal undivided, rarely divided."

Front lobe of plastron anteriorly truncate, not produced, not
or but scarcely notched ; gulars paired, small but broad ; pectorals
somewhat narrowed, their anterior border steeply sloping and
widening abruptly towards the axillary notch ; axillaries usually
1 or 2, sometimes o or 4, even 5,'^ the inner small, the outer
moderate; inguinals 3 or 4, rarely absent,^ or only the outermost
present, outermost triangular, the rest transverse, the innermost
in contact with the femoral ; hind lobe broadly notched pos-
teriorly,

Plastral formula: Usually Abd>h>an>(g, p, f subequal).
F>an in a Grahamstown specimen (M.C.Z. 17524). The femoral
suture is always smaller than that of the anal, according to
Hewitt (1931:498).

Color. Carapace yellowish olive, olive or greenish, each shield
reddish brown in the centre, the margins edged with dark brown
or black. Plastron yellowish, mesially brownish, rarely uniform.

According to Siel)enrock (1910:697) the $ S are much more
lightly colored than the 9 9 , especially the plastrons which in
$ S are mostly uniform, while those of the 9 9 are almost
always more or less darkly pigmented ; the young of the two
sexes are indistinguishable. Duerden (1907b :68) remarks that
tortoises from around Cape Town are usually darker than those
from eastern Cape Province. For detailed descriptions of the
color of live specimens see Bell (1836).

1 Four in a specimen mentioned by Oudemans (1895).

2 Six in a tortoise figured by Schoepff (1792 : pi. xxiii), also in one mentioned
by Bell fl836).

3 Eigbt in a specimen figured by Gottwald (pi. K, flg. 14) fide Strauch (1862a).
-t Five in one mentioned by Bell (18.36), and in two of Oudemans (1895).

5 Twelve in a specimen (U.S.N.M. 131846) without data.

6 Divided in one of Oudemans' (1895) series from Knysna and I'ort Elizabeth.
" Five occasionally according to Hewitt (1931 :499).

s Absent in a Knysna tortoise (M.C.Z. 20967) only, but Siebenrock (1910)
alleges that they are frequently absent.

372

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Size. Carapace length of largest S , presumably from Knysna,
96 mm. (Oudemans:1895) ; another $ (M.C.Z. 21457) from
Table Mountain, has a carapace length of 95 mm., breadth 67
mm., height 39.5 mm.

Carapace length of largest 2 , a Grahamstown specimen ( He w-
itt:1937e), 114 mm., breadth 86 mm.; another Grahamstown 9
(M.C.Z. 22474), has a carapace length of 107 mm., breadth 82
mm., and height 53 mm.

Fig. 42. Internal view of plastron of Eomopus areolatus (B.M. 64.11.20.
15), X 1.

(P. Washer del.")

Sexual dimorphism . Judging by the material available to us,
Siebenrock's (1910:697) conclusion regarding sexual dif-
ferences in the plastral coloring of adults (vide supra) is not
quite accurate. We are not in a position to verify other alleged

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 373

distinctions that he cites. Both Boulenger (1890f) and Oude-
mans (1895) present charts of secondary sexual characters, the
former stating that $ $ have larger heads, longer and more
pointed snouts, longer mandibular symphyses, and a larger elbow
scute. Oudemans failed to note any appreciable sexual differ-
ence in size of elbow scute, but concurs as to the validity of the
cranial characters cited by Boulenger if restricted to specimens
with a carapace length of over 90 mm. Oudemans failed to find
the horny terminal tubercle which Boulenger observed on the
tail of one of his specimens, but notes that the tails of $ $ are
somewhat longer. Oudemans comments on the absence of any
concavity in the plastrons of his S $ ; and in our material at
most only a very slight central depression is discernible, the
plastron being practically fiat in both sexes.

Breeding. Nest holes are about 3 inches deep. The eggs, from
2 to 5, possibly averaging 3 at a laying, are elongate ovals rang-
ing from 21 X 27 to 23 x 33 mm. (Hemtt :1937a ; pi. xxvii: fig.
8). Incubation takes 7 to 8 months, fide Rose (1950:337).

Longevity. Three years, 1 months, 10 days in the London Zoo,
but usually die within one year of arrival (Flower :1925b).

Diet. The only information is indirect: it is said not to be
destructive in gardens or on farms (Hewitt :1937e).

Parasites. A tick is present on one of the M.C.Z. series.

Enemies. Jackals prey on the young whose shells are easily
cracked. One "Homopus," together with 23 other reptiles,
was recovered from the stomach of a 5 Secretary Bird {Sagit-
tarius serpentarius) shot on the Cape Flats (Stark and Selatan,
Birds of South Africa, 3:405). Other birds of prey, according
to Kolbe as quoted by Lacepede and others, carry them up and
drop them on the rocks to break their shells. Crows, on the
other hand, turn them over and with their strong beaks attack
the softer parts, pecking out the edible portions (Latrobe :1818).

Defense. The head is incompletely protected when it and the
forelimbs are withdrawn inside the shell (Hewitt :1937e :10).

Temperament. Readily tamed according to Bell (1836) who
found they would take food from one hand while being held in
the other. A fairly bold and active tortoise that struggles vio-
lently with outstretched head and limbs when picked up (Hew-
itt :1937e :10).

374 BULLETIN' : MUSEUJNJ OF COMPARATIVE ZOOLOGY

Habitat. The only Homopus species occurring on the Cape
Peninsula (Rose :1929). A shade-loving tortoise inhabiting the
sour-grass areas from sea-level to the summit of the Zuurberg
(Hewitt :1937e:9).

Localities. Cape Province : Albany district ; Alexandria dis-
trict ; Bathurst district ; Cape Peninsula ; Deelfontein ;^ De ^Yet ;
*Dunghye Park, Caledon ; Garcia; Glen Epstine (as Erskine in
Hewitt :1937e), Cradock district; *Grahamstown (including
Carlisle Bridge; Mountain Drive and Gunfire Hill) ; Halesowen;
*Kenkelbosch (seen at U.S.N.M.) ; Knysna; Malmsbury; Morti-
mer ; Oudtshoorn ; Pigott Bridge ; Port Elizabeth ; Simons Bay
Mountains; *Table Mountain; *Tootabi near Alieedale ; Uiten-
hage ; Welbedacht Farm, Oudtshoorn district ; Zuurberg.

Range. Cape Province south of 32° S. lat. (except for Deel-
fontein record^), Union of South Africa.

Erroneously reported from India (Thunberg) ; Matabelelaud
and Zambezi region (Smets) ; British Central Africa (Jolin-
ston) ; Victoria Nyanza (Stranch and Siebenrock) ; Senogambia
(Rochebrune).

Genus KiNIXYS Bell

1827. Kinixys Bell, Trans. Liun. Soc. London, 15, p. 398. Type: caslanra

Bell = Testudo erosa Schweigger (by original designation).
1835. Cinothorax Fitzinger, Ann. Wiener Mus.,1. pp. 108, 111, 121. Type:

Kinixys belliana Gray (designation by Fitzinger: 1843).
1843. Cinixys Fitzinger, Syst. Eept., p. 29 (emendation of Kinixys Bell).
1873b. Kinothorax Gray, Hand-List Shield Eept. Brit. Mus., p. 16 (emenda-
tion of Cinothorax Fitzinger).
Definition. Skull with triturating surfaces of maxilla and pre-
maxilla without ridges; maxillary not entering roof of palate;
anterior palatine foramina large, conspicuous; prootic well ex-
posed dorsally; quadrate enclosing stapes; surangular subequal
in height to prearticular ; neck with third centrum biconvex.

Carapace of adult movable posteriorly, usually hinged be-
tween the fourth and fifth pleurals {costal bones) and seventh
and eighth peripherals {marginal bones) ; outer side of third
costal scute shorter than that of fourth; anterior neurals hexa-

1 TliP Dcrlfoiilein rcoonl niinircs investigation and verification.

IvOVERIDGE AND W1M.IA:\1S: AFRICAN CRYPTODIRA .!/•)

gonal; submarginal scutes present; suprapygals 1 or 2. if 2
typically separated by a straight transverse suture.

Plastron not hinged; gular region greatly thickened and
produced; gular shields paired, longer than broad.

Distinguishioig marks. The distinctive carapacial hinge suf-
ficiently differentiates adult members of this genus. Young
Kinixijs, prior to development of the hinge, may be recognized
by the relationship between third and fourth costal shields as
defined above (see Fig. 13), and also by the presence of the sub-
marginal scute, though this may be diificult to see.

Range. Africa from 17° North, south to Bechuanaland and
Natal. Madagascar (? introduced).

Fossil record. None.

Remarks. In view of the many remarkable characters distin-
guishing Kinixys — some from all, others from the majority of
tortoises — it is not surprising that the genus was separated at
an early date and its status never since questioned. However,
well differentiated though it is, Kinixys is not an isolated genus.
Below, we cite special resemblances to single genera among
the Ethiopian endemics, while in a previous section we have
argued for the phyletic unity of an Ethiopian group that in-
cludes Kinixys.

Adult members of the genus Kinixys are recognizable by the
unique hinging of the carapace, but as hinging is ontogenetically
a late phenomenon, and one that may sometimes fail to develop
both in Kinixys and Testudo {vide ififra), it is not fully satis-
factory for generic definition. Startling though the hinge of
Kinixys may be, hinging is a recurrent phenomenon among testu-
dinates, an adaptation to which their solid armor is especially
suited. It has occurred once in the Pelomedusidae, once in the
Kinosterninae, four times independently in the Emydinae, three
times independently in the Testudininae. In many of these in-
stances species with or without hinging are obviously closely
related.

The three examples of hinging in the Testudininae all differ,
not only from one another, but also from the type of hinging
that occurs repeatedly among emydines. In emydines the
plastron is hinged between the hyo- and hypoplastron at the
junction of pectoral and abdominal scutes. In Kinixys, as al-

376 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

ready mentioned, the hinge is earapacial. In Pyxis it occurs far
forward on the anterior lobe of the plastron and, as shown by
Siebenrock (1906b), tends to involve a break across the ento-
plastron. In Testudo, when present, the hinge is at the base of
the posterior lobe where the abdominal scutes contact the fem-
oral, and so involves the hypo- and xiphiplastron. In Testudo
sensu strict 0 it may be present in one sex of a species and absent
in the other, so that the presence or absence of a hinge cannot
be considered as automatically implying generic rank. It is not
on account of its hinge that we have retained Kinixys as a
genus, but because of the complex of characters that indicate a
long period of separation and divergence from all related lines.

The hinging of the dorsal shell in Kinixys involves the back-
bone also. The morphology of this peculiar condition has been
discussed in some detail by Siebenrock (1907). It is an entirely
secondary condition that gradually develops with age. The inter-
digitating sutures originally present between the seventh and
eighth marginals (besides the numerically corresponding peri-
pherals), together with the second and third costal shields (also
fourth and fifth pleurals), disappear with maturity as fibrous
cartilage intervenes between the now smooth edges of these ele-
ments. The hinge first forms peripherally and spreads inward;
as a final step the transverse suture between the fourth and fifth
neurals is loosened.^

There is some evidence that this ontogenetic process occa-
sionally fails to continue to completion. At least, various authors
(Miiller; Hewitt, etc.) have described apparently mature speci-
mens of K. &. helliana in which the dorsal hinge was so poorly
developed as to be immovable, or almost so.

Though the hinge is fully formed only in mature members of
the genus and is totally absent in hatchlings, a preparatory
modification in hatchlings is shown by the shapes of the third
and fourth costals : the third being narrowed distally and the
fourth correspondingly widened. Furthermore, the fourth costal,
contrary to the general rule among turtles, may be substantially
greater in area than the third.

This dorsal scute modification appears to be the most constant

1 According to Siebonrock (1907) the fifth noural is somewliat displaced, mov-
ing: forward above the fourth so as to permit of increased mobility. We have
been unable to conflrni this.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 377

external character by which juvenile Kinixys may be recognized.
The presence of the so-called "submarginals" of Siebenrock —
one or more supernumerary scutes internal to the ventral sur-
face of the second marginal — is also diagnostic of the genus, but
these vary much in size and are not always easily visible.

Osteologically, there are various features unconnected with
the hinge that make Kinixys a specially distinctive genus —
though some of these features are shared with one or another
related genus. One minor point is the position of the costomar-

Fig. 43. Kinixys erosa (Yale Mus. 5403) (x %). Internal view of left
anterior edge of carapace; sm = submarginal scale.

(P. Washer del.)

ginal sulcus with reference to the pleuroperipheral suture.
Whereas in tortoises generally there is complete, or almost com-
plete, congruence between these two landmarks, in Kinixys the
costomarginal sulcus is, at least on the posterior half of the shell,
well down on the peripherals. In this respect Pyxis and Acinixys
are somewhat like Kinixys, and the feature is probably an emy-
dine inheritance, as is also the case with the position of the
vertebral-supracaudal sulcus on the pygal in Kinixys (well
down on the pj'gal in K. erosa and homeana, higher in helliana).
Another peculiarity, shared with Chersina however, is

378 RTTLLKTIX : MUSEUM OF COMPARATIVE ZOOLOGY

the anteroposterior lengthening of the anterior peripherals, the
anteroposterior length of the second peripheral being greater
than the anteroposterior length of the first pleural. Also rather
unusual in a tortoise is the nearly complete absence of dift'erenti-
ation among the pleurals by alternate widening and narrowing.

In Kinixys tlie gular region of the plastron is strikingly modi-
fied by hypertrophy of the epiplastral lip. A tendency to develop
a thickened epiplastral lip, recessed behind, exists in all tortoises.
It is Aveakest in Homopus, Malacockersus and some Testudo, such
a thickened lip apparently being one of the many testudinatc
characters displaying i)arallel development. Its significance is
obscure, especially so in the present case in which, as Boulenger
(1889a -.142) has noticed, the length of the thickened portion may
be appreciably more than a fourth of the total length of the plas-
tron. In this respect no other tortoises approach Kinixys except
Chersina, in which the very similar hypertrophy of the epi-
plastral lip ma}' even exceed tlie development shown in some
Kinixys (cf. Figs. 37B and 49).

Among continental African genera the skidl of Kinixys is
distinctive in its pointed snout, as seen from above. This charac-
ter is discernible in hatchlings and led to further examination
and re-identification of the desiccated juvenile from the Rondo
Plateau mistakenly assigned to Malacochersus tornieri by Love-
ridge (1942e:248). However a similarly pointed snout is present
in Pyxis of ^Madagascar.

According to Hewitt (1937a :791) another cliaracteristic of
Kinixys is the lack of fusion of the distal carpals. Such fusion is
usual in Geochelone, Testudo and PsoiiDitohates. It occurs also
in Homopus signaius and houlengeri but is absent in H. fc-
moralis and areolatus {pde Hewitt :1937a). It is also lacking in
two British Museum specimens of Chersina examined by
one of us (E.E.W.). However, according to Siebenrock (1906b)
this is a purely individual character wdthout taxonomic signifi-
cance. The matter rccjulres re-examining with more adequate
material.

Species groups. It has been suggested (for example by
Schmidt: 1919) that bcUiana should be removed from Kinixys,
the type of which is erosa, and placed in the genus Cinothorax
Fitzingi'r. However, homcana so clearly bridges the gap between

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA

37!)

erosa and belliana that we caii see no advantage in the recogni-
tion of Cinuthorax either as genus or subgenus. Tliis natural
grouping is sufficiently indicated in less formal fashion by de-
scribing (as much on the ground of ecology as structure) two
species groups within the genus, one with the two rain-forest
species, K. erosa and K. homeana, the other with only the savanna
form K. belliana.

Kinixys hclliana, the best known species, occurs over a vast area
of African savanna (climatic region 2 of Finch and Trewartha)
and borders in a great arc on three sides the range of the other
two members of the genus. Both in habitus and coloration it dis-

B

Fig. 4-1. Forelimb sealatiou in Kinixys, (all x 1).
(M.C.Z. 41945); B, K. Iwmeana (M.C.Z. 38371); C,
22486).

A, K. h. belliana
K. erosa (M.C.Z.

(P. Washer del.)

plays great variability. At times these differences are individual,
at others they appear to characterize populations, and conse-
({uently have resulted in the proposal of numerous species and
subspecies. All but one of these we reject as not being sufficiently
distinct, either morphologically or geographically. The name
spekii Gray is an example. One of us (A.L.) has used it on sev-
eral occasions, but when the species helliana was considered as a
whole we did not find it possible to retain spekii, except in a very

380

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

OSlnaia, 50 ka. from Drnkmr
O Kaolaot '; q Bakal
OWacCarthy leiand
'/•° Oanbla

S

Fore carl ah 0/

O Labe, Fouta-DJalen.

OHueala ofankao ^

t| labala ^^'^-'X^

Kadlna
0 dafsrabe

northeaat Blerra Leone Tario'
i»r.m • ^-^ P.i'it.
,.,?.""i? • ■-' ♦ Bonnta
~^^^e< • '^"•^

.■^'

«■*

Ji*"

/ — ^_^,iOr fasia. Oounaa Olatr

', ( O Allele .'

) O ClJStaiDpo I
y ; EaoollaO

' . «~ ' "^ KleahShe

, Aihantl** , oTohbun (or Tohouj/
1 Abetlfl* \ o 6 o® iron

fQUOfor

O Kinixys belliono noqueyi (Lotoste^

® Kinixys homeono Bell

• Kinixys croso (Schweigger)

subjective and geog-raphically meaningless way. A subspecies
based on this name would have neither range nor boundaries ; it
would more easily be applicable to individuals than to popula-
tions; it would in fact be a morphological type rather than a geo-
graphically differentiated segment of the species. On the avail-
able evidence we level this same criticism at the majority of the
proposed subspecies of helliana.

We are conscious that we may have undervalued some segment
(perhaps, for example, K. spckii, which was originally described
as a full species) of what appears to us, on the available evidence,
as one highly variable species. There are obviously characters
worthy of study which require better analysis than we have been
able to give them. An example is the condition of the postorbital
bone. In the material studied by us the postorbital bone may be
absent, or a loose thin bar of bone, or a thin bar of bone solidly
sutured to the frontal above and the jugal below. However, we
have not had material to check in adequate fashion the correla-

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA

381

-.../

Oao Hauasarl
Ml Oarua-Benue-jJi

^IZ^

M-

OKanaara (as Mandaaa) Hl^aanda
/ O Mayo Bala
P Oaroua J Oania ) , . -' \
^ Korm/' ^-

^SaJdJ* (Sakdje)

O. Ngaouyanga
O Tlbatl'

U-

"' sou 'ip' ^ ~- ^--,

Tleterla ®;p®»S«kbay»a»» Sou* \

Detia?.^ A^f .^°*°'"^ •Yaounde* Rol.

• lank

iTlbl

• Blplndl
J Hetat
£fonl«a

Oaopo ^

langha »- ■■■■"^-
Hlo Manl'

Wto* J

Oabontl French Congo

Ogoue V •
Fernaa •va»

Kyangav •
\N\

Loango 9
)U(aa>s ,r<

Landana' .#
CMnohozo

.^.

\

Cklbonde •
Banalla O Poke •
Oaaangnl •

Hlapn«He4Je«
Panga • OJalaalnda^
Baf»abaka»AT»*uM»ffl«Baliu''» Tillage
Stanlerrlile,* Ma»aBbl® -/ Mablra '

{

Vallkala • KalKl •

tlndn*

t Lake TangaoTlka

tabambara ar: a
Laka TanganTlka ^^

region

Fig. 45. Locality records of Kinixys (exclusive of K. 6. hellkina whose
lange extends from the Cameroon east and south). Other erosa localities
encircle Cameroon Mountain, (4), viz. Bonge (1) and Buea (5); with
homeana recorded from Bibundi (2); Isongo (3); Mpanga (6); Mujuka
(7) ; Mosake (8) and Likomba (9).

(P. Washer del.)

382 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

tioii of these three conditions with other characters and with
g-eography. "We have therefore been conservative in our taxo-
noniic approach. We agree with Hewitt (1931:466) who, while
thinking- "that a number of allied species and subspecies can be
recognized" among what is here called lielliana, admitted that
"the data for sharply defining them are not yet available."
(Italics ours.)

The only subspecies of belliana that we have retained is nogueyi
Lataste. It is distinguished by having lost a claw on each fore-
foot, a condition correlated with a definable range within which
it is constant so far as our evidence goes. Though the character
of a 4-clawed forefoot is pure in the populations we call nogueyi,
it crops up occasionally within the range of what we call h. bel-
liana. This fact, first noticed in a juvenile from Beira by Boulen-
ger, induced both him and Siebenrock to synonymize nogueyi.
However, the overwhelming predominance of 5-claw^ed forefeet
throughout the range of b. belliana affords ample justification
for the retention of nogueyi as a western race.

The other ecological species group consists of K. hoineana and
crosa which are restricted to the rain forests of West and Central
Africa (climatic region 1 of Finch and Trewortha). Morpho-
logically, all three species are readily identifiable, as may be seen
by reference to Table 7 on p. 383. As indicated there, A', home-
ana and crosa, which occur together in at least a macrogeographic
sense, agree in coloration, serration of carapace margin, and in
having an elongate and coiled trachea. They differ in a combina-
tion of other characters, sufficiently distinct and closely corre-
lated to preclude the possil)ility that we are dealing with a poly-
morphic species. In spite of a certain amount of individual vari-
ation in single characters, the total differences do hold up in
series. K. homcana is the rarer of the two, but neither species is
well understood biologically and no information is available re-
garding differences in dietetic or other ecology.

LOVERTDGE AND WILLIAMS : AFRICAN" CRTPTODIRA

383

Table 7
Characters of the Species of Kintxys

Kinixys bcUiama
Carapace pattern usu
ally radiate, variable

Carapace margin neither
spinoso in young nor
serrate in adults

Bear end of carapace
steeply sloping from
middle of 5th vertebral

Xuchal scute present,
rarely absent

5-9 scutes on forelimb
from elbow to outer
claw

Postorbital bar of skull
very thin, elongate,
loose or sometimes ab-
sent

Habitat in savannas

Kinixys homeana Kinixys erosa

Carapace pattern indistinguishable
as between these two species^

Carapace margin spinose in young
and sharply serrate in adults

Rear end of carapace Rear end of carapace

overhanging or vertical sloping; if vertical then

from anterior end of from middle of 5th ver-

5th vertebral tebral

Xuchal scute present. Nuchal scute absent,
rarely absent rarely present

5-8 scutes on forelimb 3-5 scutes on forelimb
from elbow to outer from elbow to outer
claw claw

Postorbital bar of skull
moderately thin (thin-
ner and longer than in
erosa )

Postorbital bar of skull
rather short and stout

Habitat in virgin forests

Key to the Species and Subspecies of Kinixys

Posterior margin of carapace not or but slightly reverted and serrated,
never spinose in young; nuchal shield normally present 2

Posterior margin of carapace more or less strongly reverted and strongly
serrated, spinose in young ; nuchal shield present or absent 3

Forelimb with 5 claws (though occasional 4-clawed individuals oe( nr
throughout much of the range). Range: Madagascar. Eritrea south-
west to Natal, northwest through Bechuanaland and Angola to Frencli
Cameroon 6. helliana Gray (p. 384)

Forelimb with 4 claws (all known specimens have 4). Range: Nigeria
west to Senegal b. nogueyi Lataste (p. 396)

Nuchal present, rarely absent; posterior portion of carapace from an-

1 For full description, see p. 408 and PI. 14 B, C.

384

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

terior part of the fifth vertebral descending abruptly, either vertically
or overhanging. Eange: Belgian Congo northwest to French Guinea

Jwmeana Bell (p. 400)

Nuchal absent, rarely present; posterior portion of carapace descending
gradually, if vertical, then from middle of the fifth vertebral. Eange:

Uganda and Belgian Congo northwest to Gambia

erosa Schweigger (p. 404)

KiNIXYS BELLIANA BELLIANA Gray

1831c. Einixys Belliana Gray, Synopsis Reptilium, p, 69: No locality ("W.
Africa,?" was added later by Boulenger, but appears doubtful).

1831b. Gray, 6 (as "Test. (Einyxis) Belliana").

1835. Dumeril and Bibron, 168.

1855. Gray, 13, pi. ii.

1863f. Gray, 196.

1864a. Gray, 169 (reprint of 1863f).

1865. Stxauch, 40.

1870e. Gray, 13.

1872c. Gray, 6.

1873b. Gray (part), 16.

1910. Meek, 414.

1919. Schmidt, 406, 600, pi. xi, fig. 2.

1928d. Loveridge, 49.

1929. Flower, 25.

1929h. Loveridge, 14.

1931. Hewitt, 466.

1933. Schmidt, 4.

1934. Pitman, 307.
1936h. Loveridge, 19.
1937a. Flower, 5.
1937f. Loveridge, 495.

1937. Euckes, 101, pi. x, fig. 8.

1939b. FitzSimons, 18.

1940. Parker, Moreau aud Pakenham, 311.
1940a. Scortecci, 126.

1941a. Angel, 151.

1941. Moreau and Pakenham, 109.
1942e. Loveridge, 247.

1943. Scortecci, 270.

1946b. Mertens, 39, 68.

1949. Conant and Hudson, 4.

1950. Eose, 346, figs. 209-210.
1950f. Laurent, E., 133.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA

385

1950. Williams, 551,

1951. Monard, 168.

1835. Cinothorax belliama Fitziager, 108.

1843. CinLrys helliana Fitzinger, 29.

1849. Smith, A., App., 1.

1854a. Peters, 215.

1866a. Bocage, 40.

1866b. Peters, 887 (Cmi/xw).

1869a. Peters, 11.

1870. Blanford, 444.

1871b. Sclater, P. L., 544.

1878a. Peters, 202,

1880c. Vaillant, 34, 88.

1882a. Peters, 5.

1886. Vaillant, 137.

1887a. Bocage, 209. (Cinnixys).

1887b. Bocage, 86.

1887a. Miiller, F., 295 {Cinyxis).

1889a. Boulenger, 143.

1890. Strauch (part), 62.

1893a. Boettger, 8.

1893. Trimen, 79.

1894c. Gunther, 85 (Cinyxis).

1895a. Bocage, 2.

1895. Prato, 19 (Cynixis).
1895c. Werner, 127 (Cininyxis) .
1896a. Bocage, 97,

1896e. Boulenger, 15.

1896. Tornier (part), 2 (Cinicys).
1897g. Boulenger, 277.

1897. Tornier, 63.
1898a. Boulenger, 716.

1898. Jeude, 5.

1898. Johnston, 361 (Cinyxis).

1898. Sclater, W. L., 96.

1898. Tornier, 282.

1898a. Werner, 203.

1899a. Mocquard, 218 (Cinyxis).

1901. Gadow, 365.

1901. Sordelli, 111, figs, 1-2,
1902d. Boulenger, 445 (Cinyxis).

1902. Scherer, 253, photo.
1902b. Tornier, 580.

386

BULLETIX : MUSEUM OF COMPARATIVE ZOOI.OGV

1902r. Tornier, 66.">.

1905h. Boulenger, 252.

1905. iSTeumann, 389.

1 9001). Siebenrock, 2, 39, fig. 1 .

1907n. Boulenger, 6.

1907J. Boulenger, 482, fig. UO.

1907a.. Duerden, 9.

1907. Siebenrock, 6.

1908. diiibb, 220.
iy08i'. Kamnierer, 770.
19081). Mocqiiard, 557.
1908. Odhuer, 2.
1909a. Chul)b, 592.
19091). Chubb, 34.

1909a. Siebenrock (part;, 510.

1910. Miiller, L., 622.

1910. Roux, 100.

1910. Sternfeld, 4.

1911c. Boulenger, 162.

1912c. Sternfeld, 200.

19121). Werner, 424.

1913. Boettger, 315.

1913c. Nieden, 54.

1913d. Werner, 40.

1916. Calabresi, 42.

1916. Siebenrock, 6.

1917. Sternfeld, 412.

1919g. Boulenger, 12 (Cinyxis).

1919. Siebenrock, 276.

1921d. Loveridge, 50.

1922d. Loveridge, 523.

1922b. Procter, 515.

1923g. Loveridge, 924, 931.

1924b. Loveridge, 2.

1924a. Werner, 266.

1927. Calabresi, 37.

1928. Cott, 952.
1928b. Scortecci, 334.

1929. Eose, 189.
1930a. Scortecci, 215.

1931. Monard, 1 09 (Cynixis ) .

1933m. Witte, 67.

1937b. Monard, 146.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA iJST

1954. Noel-Huuie, 7J.

1845a. Kinixys schoensi.i Riippell, Mus. Sonckenhorjj, 3. ]). 22(5, \)\. xvi, figs.

1-3 : Shoa, southern Ethiopia.
1845b. Riippell, 297.

1922a. Mertens, 168 (lists type in Mus. Senckenberg).

1847. Testudo geometrica Bianconi (not of Linnaeus), 52, pi. vi, tigs. 1-2.
1882a. Peters, 3 (corrects Bianconi 's misidentification of belliana).

1863b. Kinixys SpeMi Gray, Ann. Mag. Nat. Hist. (3), 12, p. 381: Central

Africa (i.e. probably northwest Tanganyika Territory).
1870e. Gray, 14.

1931. Hewitt, 469.

1935. Hewitt, 347,

1936.J. Loveridge, 218.

1937f. Loveridge, 492, 495,

1943. Scortecci, 270, 282.

1S90. Testudo areolata Strauch (part; not of Thunberg), 58 (Victoria

Nyanza specimen only).
Ji)02b. Ilomopus darlingi Boulengcr, Proc. Zool. Soc. London, 2, p. 15, pi.

iv: Salisbury District, Mashoualand, Southern Ehodesia.

1906. Duerden, 408, pL xi, fig. 3.

1907. Duerden, 68, fig.

1920c. Testudo Loveridgii Bouleuger (part: the small specimen mthout

locality was erroneously attributed to loveridgii) , p. 264.
1920. Uinged Land Tortoise Carpenter, 186 (Lake Victoria).
1927c. Cinixys lohatsinna Power, Trans. Roy. Soc. S. Africa, 14, p. 410,

pig, xix-xx: Lobatsi, Bechuanaland Protectorate.
1931. Power, 46, 49, figs. 1-2.

1931. Kinixys belliana sombensis Hewitt, Ann. Natal Mus., 6, p. 469, fig.

Ic, pi. xxxviii, fig. 4: Zomba, Nyasaland.
1934a. Mertens and Miiller in Rust, 8.

1935. Hewitt, pi. xxxiv, fig. 1 (not mentioned in text) .

1931. Kinixys belliana suluensis Hewitt, Ann. Natal Mus., 6. p. 471, figs.

la-lb, pi. xxxviii, figs. 1-3, 5-10: Richard's Bay, Zululand.
1934a. Mertens and Miiller in Rust, 8.

1935. Hewitt, 350, pi. xxxv, figs. 1-2.

1931. Kinixys australis Hewitt, Ann. Natal Mus., 6, p. 477, pL xxxvi, figs.

4-6 : White River, eastern Transvaal.
1931. Kinixys darlingi Hewitt, 481.
1934a. Mertens and Miiller in Rust, 8.

1934. Pitman, 307.

1931. Kinixys jordani Hewitt, Ann. Natal Mus., 6. p. 481, pi. xxxvii, figs.

7-9 (not figs. 1-3 as stated) : Isoka, Northern Ehodesia.
1934. Pitman, 307.

388 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1931. Kinixys youngi Hewitt, Ann. Natal Mus., 6, p. 486, fig. Id, pi. xxxvii,

figs. 4-5 : Shore of Lake Nyasa below Livingstonia, Nyasaland.
1934a. Mertens and Miiller in Eust, 8.

1949. Moyle, 582.

1931. Kinixys lohatsiwria Hewitt, 488, figs, le-lf and 2, pi. xxxvii, figs. 6-7.
1934a. Mertens and MiiUer in Rust, 8.

1932. Kinixys australis mababiensi-s FitzSimons, Ann. Transvaal Mus., 13.

p. 37: Tsotsoroga Pan, Mababe Flats, Bechuanaland Protectoi-ate.
1934a. Mertens and Miiller in Rust, 8.

1935b. FitzSimons, 302, pi. x, figs. 1-3.

1934a. Kinixys australis australis Mertens and Miiller in Rust, 8.
1934a. Kinixys helliana helliana Mertens and Miiller in Rust, 8.
1937b. Mertens, 5.

1938e. Mertens, 430.

1952. Witte, 12.
1953e. Loveridge, 158.
1953h. Loveridge, 140,

1953. Witte, 15, pi. iv, fig. 1.
1955b. Mertens, 52.

1934a. Kinixys belliuna si)e'kei Mertens and Miiller in Rust, 8.

1935. Kinixys natalcnsis Hewitt, Rec. Albany Mus., 4, p. 353, pi. xxxv,

figs. 3-4: Jameson's Drift, Tugela River Valley, Natal.
1937a. Kinixys sulucnsis Hewitt, 791, fig. 5 (of foot).
1942e. MalacocliersHS tornieri Loveridge (not of Siebenrock), 248 (misdet.,

juvenile found dead, crushed and dried).
1946. Kinixys somhensis Mitchell, 19.

Further citations of Kinixys, or Cinixys, helliana will be found
under K. h. nogueyi.

Common names. Bell's Eastern Hin^ed-Tortoise ; doha (at
TeteiPeters) ; enfudu (Ganda iCarpenter) ; fudi (Pokomo :Love-
ridge) ; furgohi (Gogo :Loveridge) ; furu (Cewa :Mitchell) ; im-
hati (at Ngangela :Monard) ; kakafida (eastern Angola :Lau-
rent) ; Kamba (Manganja & Sena :Mitchell) ; kohi (Mwera:
Loveridge) ; namhi (Makonde and Mawiha :Loveridge) ; ngongo
(Yao:Loveridge; Mitchell); nguru (Sagella and TaitatLover-
idge) ; ufutu (Matabele :Chubb) ; umheo (in Benguela iBocage).

Description. Beak not or but weakly to moderately hooked,
uni- or tricuspid ; edge of jaws not serrate ; prefrontal large, en-
tire, semidivided or divided longitudinally, rarely broken up ;
frontal large (occasionally divided longitudinalljO , moderate or
broken up ; upper head shields small, irregular ; forelimb anteri-

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 389

orly covered with a few or many, lar<i:e, unequal, scattered or
juxtaposed, strongly imbricate or non-imbricate, more or less
pointed (subcircular or pointed in young, subacute in adults)
scutes which on anterior edge form a longitudinal series of 5-9
from elbow to outer claw ; claws 5, rarely 4 ^ ; hinder side of
thigh without enlarged tubercles; heel with or without well-de-
fined spurlike tubercles ; claws 4, rarely 3 ^ j tail terminating in a
more or less distinct hornj^ tubercle that is sometimes much
larger in c5 $ .

Carapace moderately convex or markedl}^ flattened dorsally,
sides sloping, scarely or not (though sometimes deeply ^) notched

Fig. 46. Skull of Kinixys h. helliana (A.M.N.H. 10029). Condylobasal
length 37 mm.

(S. McDowell del.)

in nuchal region ; anterior margin not or but slightly expanded,
not or but slightly reverted, not spinose in young, posterior mar-
gin not expanded, not or but slightly reverted, not or but slightly
(occasionally moderated"*) serrated, not spinose in young, rear

1 Four in 2 out of 23 tortoises from Uele region of Congo, a third had 4 on
one forefoot and 5 on the other (Schmidt :1919 :407) ; four in 3 (M.C.Z. 40011,
40013, 40017) out of 8 Kenya spfcimens, and four on both forefeet of 1 (M.C.Z.
44451) of S Southern Rhodesia tortoises examined by us. This Khodesian tortoise
is unusual in that it is the fourth claw that is laclving ; four in a juvenile only
of a series of at least 6 from Beira (Boulenger : 1907J : 482 and Hewitt : 1935 :
353). The missing claw on each forefoot is represented by a light-colored, rounded
tubercle (Battersby in letter of 15.xii.54). All 14 Tanganyika and all 24 Xyasa-
land specimens in the M.C.Z. have the normal five claws of h. helliana.

2 Three in a Belgian Congo tortoise mentioned by Schmidt (1919 : 407).

3 Deeply notched in an Mtimbuka juvenile (M.C.Z. 50328) as the uu
bsent.

4 Moderately serrated in type of lohatsiana Power (1927c :pls. xix-xx).

390 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

end in 3'oiing sloping more or less steeply, in adults even more
SO; dorsal shields concentrically striated, the striations more or
less reduced by wear in aged specimens, not or only sometimes
swollen,^ not convex; nuchal usuallj^ elongate, though as broad
as, or much broader than, long in verj^ young, occasionally di-
vided, minute- or absent;'^ vertebrals 5, rarely 4,"* 6 or 7,'' not
or but slightly convex, first broader than long in young, broader
than long, as broad as long, or longer than broad in adults ; sec-
ond to fifth as broad as, or broader than, long; second broader,
subecpial to, or narrower than, fifth in young, narrower in
adults; fifth fiat in young, more or less convex in adults; costal.s
4, rarely 3** or 5,'^ usually not forming an angle with the mai'-
ginals; marginals 9-12,*^ usually 11; supracaudal undivided,
rarely divided,** with or without a tendency to be incurved in
S $ , otherwise not or but slightly reverted in both sexes.

Front lobe of plastron anteriorly truncate, very thick, not or
but slightly or moderately projecting beyond the anterior border
of the carapace, not bifid, not or very openly notched; gulars
paired; pectorals ^^ moderate, their anterior borders straight or
widening more or less gradually towards the axillary notch ;
axillaries 2-4, the innermost small, inconspicuous or absent, the
outermost more or less concealed; inguinal 1,'^ large, in contact

1 Strongl.v swollen, with deep gutters between the shields, in .". Matia Islaiul
tortoises (fide Siebeurock : 1906b : 2).

-Divided in a juvenile (U.S.N.M. 41699). When rudimentary it may rosiilt in
the marginals meeting either before or behind it (Strauch : 1890).

3 Absent in an Mtimbuka Juvenile (M.C.Z. 50328) and in one of a Beira serie:^
(Hewitt : 1931).

■i Pour in a carapace from Moroni figured by Sordelli (1901:111) with con
sequent elongation of the third vertebral, etc.

5 SiY or seven in an Ujiji and some Mtimbuka adults (M.C.Z. 48001 ; 5031S :
50323).

6 Three on right side of a Garamba 9 (A.M.N.H. 10047) where the missing
i-ostal is replaced by an extended vertebral that is in contact with the marginals.

VFive in an Mtimbuka $ (M.C.Z. 50318).

« Nine on left, ten on right of Moroni carapace (Sordelli :1901) ; ten on right,
eleven on left in a Sandown Q (Hewitt) ; twelve on left side of an Ithanga
juvenile (M.C.Z. 8158 ( and twelve in an Mtimbuka $ (M.C.Z. 50323) with the
hinge between marginals 8 and 9 instead of between 7 and 8, the normal arrange-
ment; twelve in an Ujiji $ (M.C.Z. 48001) whose hinge is between marginals 7
and 8 ; in an Angolan tortoi.se the hinge was between 6 and 7 (fide Bocage : 1866a).
as also in a Natal specimen mentioned by Hewitt (1935).

9 Divided in a Mussera tortoise (fide P. L. Sclater :1871b) ; also in several from
Natal, maile cotypes of nataletisin (Hewitt : 1935).

10 Pectoral on right side of a Cheren $ is fused with the corresponding ab-
dominal shield (Scortecci : 192Sb) ; pectorals fail to meet in an Tele tortoise
(A.M.N.H. 10031) mentioned by Schmidt (1919 : 407).

11 Absent in a Natal tortoise (fide Hewitt :1935 :353).

LOVERIDGE AND AVILLIAMS : AFRICAN CRYPTODIRA ."'i'l

with the femoral ; hind lobe very short, truncate, not or but
slightly notched posteriorly.

Plastral formula: Al)h>, h>(g, j), f, an, all extremely vari-
able).

Color. Carapace straw-colored, yellowish brown, olivaceous
brown or reddish brown ; in young uniform, or the areolae deei>
brown surrounded by a zone of yellow which may extend to the
margins of each shield or be interrupted by fine black radiations
from the darker areolae or be entirely replaced by black edging.
The juvenile coloring may persist in adults, though usually with
the modification that the black edging of the shields is irregularly
or symmetrically broken up by yellow pigment, or only the
black areolae of the juvenile pattern persists, in which event it is
frequently interrupted by a median, longitudinal, yellow streak.
In a few individuals, chiefly S S , the coloring may be uniform.

Plastron horn color, uniform or smudged with black, the
latter sometimes in the form of radiations from around the
areolae.

The literature teems with descriptions of deviations of the
above, for the coloring of heUiana is extremely variable — due in
part to the normal zonary pattern of the young being replaced
by the secondary or radiate arrangement found in adults, in
part because S S tend towards uniformity while 9 9 are apt
to be darker.

Size. Carapace length of largest S (M.C.Z. 18151) from
Turiani, 193 mm., breadth 125 mm., height 83 mm.; carapace
length of largest 9 (M.C.Z. 50321) from Mtimbuka, 207 mm.,
breadth 135 mm., height 88 mm. Possibly the smallest, from
tlliya, length 50 mm., breadth 46 mm., height 25 mm. (All
three collected by Loveridge.)

That the heUiana of open savanna have higher shells, while
the more depressed spekii type live in rocky terrain, where they
may seek shelter beneath boulders, was suggested by Loveridge
(1923g:924) who, both then and later (1936j :1942e :1953e).
furnished tabulated measurements. In 1942 he reluctantly re-
tvirned spekii to the synonymy in the absence of evidence that
the low-vaulted form, which predominates in certain localities
like Mtimbuka, has any zoogeographical significance. With this
decision we concur after careful reappraisal of all available
material and data.

392 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Sexual dimorphism. In addition to a certain amount of sexual
dichromatism among adult helliana, as indicated above, the most
promising aids to determining sex are :

Plastron usually somewhat concave along the median axis ;
anal shields forming an almost straight edge posteriorly ;
tail very long, normally terminating in a large horny

tubercle 5

Plastron flat ; anal shields forming an incurved edge pos-
teriorly; tail short, normally terminating in a very

small horny tubercle ?

The caudal tubercle is not always so well-developed in males as
to be the infallible guide to sex supposed by Sternfeld (1917:
412).

Breeding. On October 10, at Lumbo, a 207 mm. 9 held a
mass of ovules ranging from pea-sized to 28 mm. in diameter;
of the latter there were about 15 (Loveridge :1921d).

In May, 2 oval eggs were present in one tortoise, and 3 each
in two other Mafia Island 9 9 (Siebenroek :1906b :fig. 1), their
dimensions correlated with those of the parent, for the eggs
of a 168 mm. tortoise measured 41 x 33 mm., those of a 192
mm. reptile 45 x 36 mm.

Longevity. Nine years and 9 months ( Conant and Hudson :
1949) ; 9 years, 3 months, 28 days (Flower :1937a).

Diet. Frequently seen feeding on fallen bananas and man-
goes ; two were observed eating sugar cane in a plantation during
a downpour (Scherer :1902). Four found feeding on fungi at
Lobatsi (Power :1927e) ; fungi (Mitchell :1946) ; take pawpaw
readily (Loveridge :1928). Though plenty of grass was present
in the enclosure, Loveridge (1923g:925) found one of these tor-
toises eating hard Indian corn; a single kernel would be taken
up and dropped five or six times before being swallowed. An-
other Kinixijs, munching something at the time, w^as found nos-
ing a large yellow carpenter bee {Xylocopa nigrita) of whose
hind legs one was missing. Though the tortoise was seen to return
again and again to the bee, it never actually bit it. The insect,
which had only recently died, had probably fallen from a beam
above. A $ , feeding in bright sunshine beside the Voi road, made
off with surprising agility when approached (Loveridge :1936j :
220).

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 393

Parasites. Ticks (Amhlyomma yiuttalUy have been recorded
from many localities (Loveridge). Nematodes are commonl}-
found in the intestines of Congo tortoises ( Lang, in Schmidt :
1919).

Enemies. In Angola the natives esteem these tortoises as food
(Anchieta, in Bocage:189r)a). In Faradje district of the Belgian
Congo, following the grass fires of February and March, the
Azande and Logo organize tortoise-hunts. The reptiles are
located by the natives' dogs, and the animals unearthed by the
iron-shod butts of spears. The rear portion of the carapace is
readily separated at the hinge and the meat removed for stewing.
The stomach and cloaca are usually rejected but the rest of the
reptile, including nematodes which are thought to add to the
flavor, are dropped in the cooking pot. These tribesmen of the
northeast Congo apparently prefer tortoises to chicken, and fancy
them more than the semi-aquatic terrapin (Pelomedusidae) which
have a strong musky odor (Lang, in Schmidt :1919 :410).

Apart from man, Kinixys appears to have few enemies. How-
ever, Lang records finding the shell of a recently eaten one whose
carapace and plastron were marked by the teeth of a leopard the
tracks of which were plainly visible. In East Africa tortoises
may sometimes be found wandering about with shells that have
been bitten by hyenas or other carnivores, the scars completely
healed. The empty shell of one specimen (M.C.Z. 18153) is stud-
ded with tooth marks and the bone re-ossified where an extensive
piece has been broken right off.

A Kinixys with carapace fractured and bleeding, as if dropped
from a height by some bird of prey, was found by Carpenter
(1920:186).

Defense. When picked up, a Bell's Box-Tortoise may snap its
beak and struggle wildly, so that its captor is liable to be
scratched by the flailing forelimbs or wet by the pungent cloacal
fluid that is discharged. Another individual may emit a hiss as
it withdraws its head within the sheltering carapace, completing
its concealment by bringing forward and folding the heavily
armored forelimbs. Simultaneously, air is expelled from the
lungs as the rear portion of the carapace comes down to enclose
the hind limbs and tail (Lang, in Schmidt :1919).

1 Early records of A. marmoreum were based on ticks that have been re-examined
by Dr. .1. C. Bequaert and found to he nuttalli.

894 BULLETIN : JMUSEUM OF COMPARATIVE ZOOLOGY

Aestivation. At the commencement of the dry season (Decem-
ber to May) in the Uele, by burrowing among the roots of trees,
into termitaria, or a few inches below ground, Kmixys usually
escapes the devastating grassfires, remaining quiescent until
aroused by the onset of the rains (Lang, in Schmidt :1919 :41()).

Habits. In the coastal belt of the Congo, where it inhabits low
granite hills, Kinixys appears only in the hot rainj- season (Octo-
ber to May) remaining underground during the cold weather
according to the natives (Sclater :1871b :54-4). Fairly common
in the savanna of the northeast Uele where they find shelter and
food in the dense undergrowth of the moister areas bordering
swamps and shallow watercourses. During the rainy season they
wander further afield so that the natives have difficulty in locat-
ing them in their dry-season haunts. At the beginning of the
rains (flune) three of the tortoises were found in a shallow j)Ool.
overgrown with vegetation. One, released near a river, walked
to the edge of the steep bank and unhesitatingly dropi)ed into
the water where, though powerless against the strong current, it
ap]5eared at ease (Lang, loc. cit.). Found beneath bushes and in
thickets along the river banks of Eritrea (Blanford :1870). In
Tanganjdka commonly encountered during the rainy season in
damp areas of the savanna {fide Bohm), also in a papyrus swamj)
according to Stuhlmann (Tornier :1896). Plentiful at Broken
Hill during rains (Pitman :1934). J. M. McCallum {in Hewitt:
1935:494) reports on the readiness with which "lohatsiaiui" will
enter water. A 75 muL juvenile was found in a termitarium at
Enteudweni, Zululand (Odhner :190b).

Though members of this genus are said to usually walk un the
claws of their forefeet, lohatsiana places "the palms on the
ground at every step and can remain standing on the palms, the
carapace being raised a little from the ground" fide Hewitt
(1931:500;footnote).

In captivity these tortoises thrive best when kept under humid
conditions at high temperatures fide Kammerer (1908b :770). In-
stead of taking advantage of the shelters provided for them at
Kilosa, seven A'. &. hcUinna were apt to stay out at night, par-
ticularly when raining, si)ending the entire night in etforts to
escape from their enclosure (Loveridge in Procter :1922b :523,
where its habits are contrasted with those of Malacochersus) .

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 'AU^)

Localities. i<udan: Khartoum; i^iraiiga; Moroni; Talodi, Nuba
Mountain Province; Sudan (as '"Sandoin" ^) ; *Torit. Eritrea:
^Vnseba Valley; Asmara; Cheren. Ethiopia: El Direre, Tertale ;
Errer (Erer) Kiver near Harar; Shoa (Selioa). British Soynali-
land: Berbera. Somalia: "Adden Cabola"- ( '/ Caboba) ; "Batl-
ditu to Dime"; Webi Kiver in Ennia-(jiallaland. Uganda:
Busu Hill; Busu (Bussu) near Jinja; Masindi; Mount El-
gon. Kenya Colony: Athi Plains; *Golbanti; *Itlianga Hills;
*Kibwezi; Kitui; Moyale; Mtito Andei ; Nairobi; Taru ; Tercale
(Pozzi-) ; *Voi; Yatta Plains. Tanganyika Territory: Ambon i
near Taiiga ; Bubu near Kondoa; ]5ukome ; "Central Africa"
(for type of spekii) ; Dodoma: Godegode; Ikikuyu; Kakoma ;
Korogwe (Kerog"^ve) ; Kigoma; Kilimanjaro Mtn. ; *Kilosa;
*Kiponda to Mitungu; Kiserawe; *Kitaya ; Kongonda near Do-
doma; Lake Rutamba; Lake Tanganyika; Lake Victoria; Lon-
gido West; Mafia Id.; Maji ya Cliumvi; Mbulu near Arushri :
*Mikindani ; Mkata River; .\lkonmsi River; Morogoro; Mugango.
24 km. S. W. Musoma ; Myombo ; *Ncliingidi, Rondo Plateau ; Ngo-
ga; Nyangesi, Usukuma; Pangani River bank; Pwaga; *Sinili()
near Tabora ; Tamborale (Muana Tomboloo) ; Tanga; *Turiani :
*Ujiji; Uliya. Zanzihar Island: Mafia Id.; *Zanzil)ar (]\Iiiller:
1887a ; Strauch :1890 ; etc.) . Mozambique: Amatongas ; Beira ; Bo-
ror ; Caia ; Cliarre ; Chitau ; Gauca ; Inhambane ; Lumbo ; Mesuril ;
Sena; Tete. Ayusaland: Angoniland ; Chibotela ; Chikwawa ;
Chiromo; Cliitala River; *Cholo Mtn.; Port Johnston; Lake
Shirwa (Chilwa) ; Lakeshore below Livingstonia; Likangala ;
Monkey Bay; *Mpimbi near Zomba; *]\Itimbuka ; Port Herald;
Zomba. Nortliern Bhodesia: Broken Hill; *Jsoka; Petauke.
Southern Rhodesia: *Birclienough Bridge; *Bulawa3'o ; Chan-
gadzi River; Essevale; *Hot Springs; Marandellas to Umtali ;
Mazoe ; Salisbury district ; Sandown ; *Selinda Mtn. and valley
at its foot; Shangani River. Bcchuanaland Protectorate: Lo-
batsi; Tsotsoroga Pan, Mababe Flats. Transvaal: Glentig near
Nylstrom; Naauwpoort; White River. Natal: Dimani's Stream;
Entondweni (Entendweni) ; Lupanza near Greytown; Jame-
son's Drift; Manaba; Matubatuba; Natambana; Richard's Bay;
Tugela River and Valley; Umfolosi Station. Angola; Caconda;

1 Id P.oulenger (lSS!»a), correction confirmed by J. C. Battersby (15.xii.l954).

-In Calabresi (1916), reporting on the Stefani and Paoli material with many
misspellings.

396 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Capangombe; Caquindo (Kakindo) ; Chitaii (CM.) Cubal;
Cubango (Kuvangu) ; Duque de BragaiiQa; Ebanga; Galanga;
Gaiica (CM.) Lui River, Muata-Yamvo region; Lunda; Mbale
River; Muita; Osi; Quissange; Vila da Ponte. Belgian Congo:
Abimva ; Avakubi ; Bowa River ; Cape Tembwe ; Dika ; Faradje ;
Ganza ; Garamba ; Kabambaie ; Kabulnmba ; Kansenia ; ' ' Kanz-
luze ' ' ; *Kapiri ; Kasonga to Uvira ; Kaswabilenga ; Kateke ;
*Lukai'u; Lukonzolwa; Mahagi Port; Manda; Masombwe;
Mpala; Mulungwe River; Musosa ; Musserra; Niangara; North-
west shore of Lake Tanganyika ; Uele region. French Cameroon :
Meiganga; Yoko (5 claws present on forefeet of both).

Madagascar: Ambauja (two alive: Angel :1941a :151) ; Ambas-
suara (Peters :1878c :792) ; Nosy (Nossi) Be (Boettger :1889 :
295).

Range. Eritrea southwest to Natal, northwest through Bechu-
analand and Angola to French Cameroon where it meets with
the western race. Also Madagascar. (The possibility of its hav-
ing been introduced here should not be overlooked. But for
Angel's recent record from northern Madagascar, we might have
questioned its occurrence, half a century having elapsed since
the indefinite records of Miiller, Strauch and others.)

KiNIXYS BELLI ANA NOGUEYI (Lataste)

1844. Kinixys lelliana Gray, part, not of Gray 1831, p. 12.

1855. Gray (part), 13, pi. ii.

1873b. Gray (part), 16.

1884a. Eochebrune, 16.

1857. Cinixys Belliana Jan (not of Gray 1831), 35.

1890. Strauch (part), 62.

1893c. Matschie, 208.

1896a. Socage, 74.

1901c. Tornier, 66.

19061. Boulenger, 197 (as Cinyxis).

1906b. Siebenroek, 3.

1907 j. Bouleuger, 482.

1907. Siebenroek, 6.

1909. Gendre, cvl.

1921a. Qiabanaud, 461.

1921b. Chabanaud, 522.

1937. Andersson, 3.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 397

1886. Homopits Nogueyi Lataste, Le Naturaliste (2), 8, p. 286: Medina

(as Mediue), Upper Senegal.
1889a. Boulenger, 148.

1901c. Tornier, 67.

1906c. Duerden, 408.

1888a. Cinixys Dorri Lataste, Le Naturaliste (2), 10. pp. 164, 226, fig. 1:

Bakel, Upper Senegal.
1903b. Cinixys nogueyi Siebenroek, 185, 442.
1931. Kinixys nogueyi Hewitt, 468.

1934a. Kinixys belliana nogueyi Mertens and Miiller in Rust, 8.
1952a. Loveridge, 231.

1955. Cansdale, 89, 103.

Common 7iame.<i. Bell's Western Hinged-Tortoise. Nkounou
(in Senegal :Roehebrune).

History. The species was named for M. Gustave Noguey of the
Linnaean Society of Bordeaux. Siebenroek (1903b) was the first
to detect Lataste 's error in assigning- yiogueyi to the genus Homo-
pus; he also re-examined the eleven Togo tortoises referred to
heUiana by Tornier (1901c) and found that all had 4 claws on
the forelimbs. Boulenger (19061) questioned Siebenroek 's action
in making nogueyi a full species, pointing out that in some West
African examples radiating black streaks are present, while the
gular/anal proportions do not differentiate nogueyi from heUi-
ana. Boulenger regarded nogueyi as a 4-clawed variety of
belliana. Later (1907j) he reported a young belliana from Beira,
Mozambique, as having only 4 claws (though the adults from the
same locality had 5), so concluded that nogueyi was inseparable.
Loveridge (1952a), after studying five 4-clawed tortoises from
Togo and Dahomey, regarded it as a West African race and de-
fined the range.

Description. Beak weakly to moderately hooked, uni- or bicus-
pid, edge of jaws not serrate ; prefrontal divided longitudinally ;
frontal large or small, occasionally divided longitudinally ; upper
head shields small, irregular ; f orelimb anteriorly covered with a
few, large, unequal, scattered or juxtaposed, strongly imbricate,
subcircular or subacute scutes which on the anterior edge form a
longitudinal series of 6-7 from elbow to outer claw; claws 4;
hinder side of thigh without enlarged tubercles ; heel without, or
with a trace of an enlarged tubercle ; claws 4.

3!)8 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Carapace moderately convex, flattened dorsally, sides sloping
in young, descending abruptly in adult, not or but openly notched
in nuchal region; anterior margin somewhat expanded and not
or but slightly reverted, not serrated, not spinose in young;
posterior margin not expanded, not or but slightly reverted, not
serrated, not spinose in young ; rear end in young sloping gradu-
ally, in adults vertically from the middle of the fifth vertebral ;
dorsal shields concentrically striated (the striations more or less
reduced by wear in aged specimens), neither swollen nor convex.

Nuchal about as long as broad in young, elongate in adults;
vertebrals 5,^ without median keel (except in young which may
show the barest trace), not convex, first broader than long in
young, longer than broad in adults, second to fifth broader than
long, second as broad as fifth in young, narrower in adults, fifth
flat in hatchlings, somewhat convex in adults ; costals 4, not form-
ing an angle with the marginals; marginals 11-12 -; supracaudal
undivided, more or less reverted ; tail terminating in a horny
tubercle.

Front lobe of plastron very thick, not or but slightly project-
ing beyond the anterior border of the carapace, not or but
scarcely notched ; gulars paired ; pectorals moderate, their anterior
border widening gradually towards the axillary notch ; axillaries
3, the innermost small, the outermost more or less concealed ;
inguinal large, in contact wath the femoral; hind lobe very
short, truncate, not notched posteriorly.

Plastral formula : Abd>h>g>or=or<f >p>or<an.

Color. Carapace light olive green or reddish brown, uniform,
or each dorsal shield with yellowish green areolae and bordered
with black, w'hich may be interrupted at the angles. Plastron
yellow, or greenish yellow, more or less marked bj' black or
reddish brown.

Size. Carapace length of a <$ (I. F.A.N. 2052) from Bassila,
148 mm., carapace length of largest Togo tortoise, unsexed,
in Berlin Museum, 220 mm.; largest 9 (I.F.A.N. 2052 also)
from Bassila, 152 mm. ; of the type 2 (B. M.) from Medina, 135
mm., breadth 99 mm., height 60 mm. Carapace length of a .juv.
(M.C.Z. 51755) from Tohoun, 48 mm.

1 A relatively small azygous shield split off from left side of fifth vertebral in a
Kabala $ (M.C.Z. 54118).

2 Twelve on left side only of Kabala ^ (M.C.Z. 54118).

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA ^00

Diet. Lataste, finding much debris of Coleoptera as well as
vegetable matter in the intestines of the 9 cotype, cites this as
evidence of the reptile being insectivorous as well as vegetarian.

Enemies. Ornaments are made from the emptied shell (Cans-
dale :19r)r)). Loveridge (1952a), noting that the claws and soles
of eaeh foot of a Tohoun tortoise appear to have been sliced off
with a sliai'p knife, suggests it may have been done by an African
who was afraid of being scratched.

Aestivation. Burrows into the ground during the dry season
according to native reports (Cansdale :1955).

Localities. French Cameroon: Garoua (Garua) ; Konn; Man-
dara (as Mandana) Iliglilands (shell only, so race questionable) ;
Mayo Sala: Xdikinimeki ; Ngaouyanga; Sajdje (Sakdje) ;
Tibati.^ Nigeria: Gando (Gandu) ; "Gao Haussari am Garua-
Benue "■•-'; Lagos. Dahomey: *Bassila ; Porto Novo."' Togo:
Adele (as BLsmarekburg) ; Lome; Misahohe (see Hohe) ; Tohoun
(or Tohou). Gold Coast: Kintampo; Somanya, Krobo (M.G.
Lesage). Sierra Leone: *Kabala, Koinadugu district; Musaia."*
French Guinea: Beyla; Forecariah ; Kankan; Labe, Fouta-
Djalon. Portuguese Guinea: Bolama. Gamhia: MacCarthy
Island. French Sudan: Diafarabe; Pama, Gourma district (as
Pama-Gurma in Tornier :1901c). Senegal: Bakel; Kaolack; Me-
dina; Sindia, 50 km. from Dakar (A. Villiers).

Range. French Cameroon west to Senegal.

1 We are indebted to Dr. V. Aellen and Mods. W. Lanz for examining the speci-
mens from six of tliese localities in French Cameroon and ascertaining that the
forefeet bore 4 claws in every one of the eight tortoises involved.

- Apparently this refers to some Gao in the Haussa country of northern Nigeria
west of Garoua on the Benue River in French Cameroon, in an effort to dis-
tinguish it from the well l<no\vn Gao on the Niger River. Passare was the
collector, states Tornier (1001c : 67).

3 Specimens from Porto Novo : Lome ; Forecariah : Kanlian ; Diafarabe and
Kaolack, all have forefeet bearing 4 claws, writes A. Villiers (ll.ii.55, etc.) in
whose care they are at I.F.A.N., Dakar, Senegal.

4 A limbless ."iS mm. shell, originally said to have measured 64 mm., so shrink-
age during drving may account for the suiT>rislngly abrupt, homeana-likp declivity
of the carapace posteriorly. Collected in northern Sierra Leone in 1048 by T. S.
.Tones. Now in the British Museum and seen by us.

400

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

KiNIXYS HO ME ANA BeLL

1827. Einixys Homeana Bell, Trans. Linn. Soc. London, 15. p. 400, pi. xvii,
fig. 2: West Africa (The statement that the types came from
Sierra Leone, was later amended by Boulenger (1889a: 143) to
Cape Coast, Ashanti, i.e. Gold Coast. Both Bell and Boulenger
give Lt. M. C. Friend as the collector.)

1831b. Gray, 6.

1831c. Gray, 15.

1835. Dumeril and Bibron, 161, pi. xiv, figs. 2-2a.

1844. Gray, 11.

1845. Wyman, 38.

1850. Berthold, 423, pis. xliii-xlv.

1855. Gray, 13.

1863f. Gray, 170 (reprint of 1863f).

1864a. Gray, 197.

1865. Strauch, 38.

1870e. Gray, 14.

1873b. Gray, 17.

1884a. Eochebrune, 17.

1919. Schmidt, 403, 598.

1929. Flower, 25.

1930a. Barbour and Loveridge, 770.

1934a. Mertens and Muller in Eust, 8.

1937c. Loveridge, 269.

1937. Euekes, 101.

1938b. Mertens, 33.

1941e. Loveridge, 115.

1950. Williams, 551.

1951. Monard, 168.

1955. Cansdale, 89, 103, fig. H5.

1856a. Cinixys Homeana Dumeril, 372.

1860. Dumeril, 162.

1875a. Peters, 196.

1887a. Miiller, F., 295.

1889a. Boulenger, 143.

1890. Strauch, 61.

1893c. Matschie, 208.

1897. Sjostedt, 6.

1898a. Werner, 203.

1899. Siebenrock, 566, pi. i, figs. 1-2,

1900b. Boulenger, 447.

1901. Gadow, 364.

1901b. Tornier, 61.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA

401

1902c.

Tornier, 665.

1906a.

Mocquard, 479.

1909a.

Siebenrock, 510.

1910.

Miiller, L., 622.

1910.

Steinfeld, 5, fig.

1911.

Lampe, 145.

1911a.

Masi, 35.

1912c.

Sternfeld, 200.

1916.

Siebenrock, 5.

1917.

Sternfeld, 412.

1919.

Siebenrock, 276.

1921a.

Chabanaud, 461.

1921b.

Chabanaud, 522.

1954.

Noel-Hume, 74.

Fig. 47. Skull of Einixys homeana (A.M.N.H.
length 41 mm.

43306). Condylobasal
(S. McDowell del.)

Common name. Home's Hinged-Tortoise (Gray).

Illustrations. Bell's (1827) finely executed drawing consists
of a lateral view of the shell in black and white to show its
abruptly descending rear end in striking contrast to that of e^'osa
(as castanea) on the same plate. Dumeril and Bibron (1835)
furnish a colored figure (also lateral), together with a line draw-
ing of the plastral scutes.

Description. Beak moderately to strongly hooked, unicuspid;

402 BULLETIN: MUSEUM OF (OMPARATIVF, ZOOLOGY

edge of jaws not dentate ; prefrontal divided longitudinally ;
frontal large, small, or broken up ; upper head sliields small,
irregular ; f orelimb anteriorly covered with a few scattered, large,
unequal, strongly imbricate, more or less pointed scutes which on
the anterior edge form a longitudinal series of .l to 8 from elbow
to outer claw ; claws 5 ; hinder side of thigh without enlarged
tubercles ; heel with or without a spurlike tubercle ; claws 4 ; tail
terminating in a small, somewhat clawlike tubercle that is larger
in ,$ 5 .

Carapace flattened dorsally, sides sloping, not or but openly
notched in nuchal region, anterior margin strongly expanded
and not or but slightly reverted, spinose in young, posterior mar-
gin not expanded, more or less strongly reverted and strongly
serrated, spinose in young, rear end in young slopiug gradually,
in adults vertically from the anterior part of the fifth vertebral ;
dorsal shields concentrically striated though sometimes scarcely
distinguishable, through wear, in aged specimens, neither swollen
nor convex (except the fifth vertebral) ; nuchal elongate, very
rarely absent^ ; vertebrals 5, with a slight median keel, first
broader than long in young, as long as, or longer than, broad in
adults, second to fifth broader than long, the tliird broader thau
the third costal ; costals 4, rarely 5- or 7,^ not or but rarely form-
ing an angle with the marginals; marginals 11, rarely 12,^ the
anterior much longer than the posterior ones ; supracaudal undi-
vided, rarely divided,^ more or less strongly reverted in both
sexes.

Front lobe of plastron very thick, not or but slightlj- project-
ing beyond the anterior border of the carapace, openly notched ;
gulars paired ; axillaries 2-4, the innermost, if present, small, the
outermost more or less concealed ; inguinal large, in contact with
the femoral; hind lobe very short, rounded, truncate, notched
posteriorly.

Plastral formula : Abd>h> (g, p, f, an, very variable), f or an
shortest.

1 Absent in one Zorzor tortoisu (U.S.N.M. l(iM698) conflrniing Loveridgc
ni>41e:113), and in an Oda specimen (C.N.H.M. 53630) studied \>y E. K.
Williams.

2 Five on left sidt; only of a Nliolembembe tortoise (Sternfeld :iyi7) .

3 Five on right, 7 on left, of a Zorzor specimen (U.S.X.M. 109098) wliicb lias
12 marginals on left side only (Loveridge : 1941e).

i Divided in a juvenile from Gabon (Dum6ril :1S60).

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 403

Color. Essentially similar to that of K. erosa, which see.

Size. Carapace length of largest S (U.S.N.M. 10962), from
Zorzor, 200 mm., not exceeded by a Mujuka $ (S.M. 156) of
210 mm. mentioned by Mertens (1938b :33) as it was apparently
measured over the curve. Carapace length of largest 2 (U.S.
N.M. 109685), also from Zorzor, 210 mm. (fide Loveridge :1941e :
115). The holotype was said to be 190 mm. long, 132 mm. broad,
and 85 mm. in height.

Sexual dimorphism. As in erosa. The length of the gular
suture in relation to the length of the plastron, was found by
Loveridge (1941e:115) to have some significance in seven tor-
toises from Zorzor. It might not hold good for larger series.

Localities. Belgian Congo: Mawambi (Mavambi) ; northwest
of Lake Tanganyika (Siebenrock:1916) ; near Saidi's Village
(Loveridge :1937c). French Equatorial Africa: French Congo.
Rio Muni: Yenghe (Jengwe, Cameroon). French Cameroon:
Bipindi; Dehane; Dibongo near Edea; Kribi; Nkolembembe
(Nkoubembe) ; Nkoolong (CM.) ; *Sakbayeme. British Camer-
oon: Bibundi; Bonge, Meme River; Isongo; Johann Albrechts-
hohe; Likomba; Mapanja; Mosake; Mujuka; Victoria. Nigeria:
Bonny (Boni) ; *Ifon, Ondo district (M.C.Z.). Gold Coast: Abe-
tifi; Ada (C.N.H.M.) ; Ashanti ; Cape Coast Castle. Ivory
Coast: Adiopodoume (V. Aellen) ; Liberia: *Cape Palmas; *Len-
ga, Farmington River; Zorzor.

Range. Belgian Congo northwest to French Guinea, or at least
Liberia.

Wyman (1845:38) received a shell from the Cape Verde
Islands, presumably a human importation. Senegal and Gambia
(Rochebrune :1884a) are similarly rejected pending confirmation.
The Mawambi record (Sternfeld :1912c), being based on a head
and limbs, was questioned by Schmidt (1919:403). However,
later records from the Belgian Congo (Siebenrock:1916 ; Love-
ridge :1937c) validate the presence of homeana in this general re-
gion, so we have tentatively allowed the Mawambi record to

stand.

We emphatically reject the alleged occurrence at Kiu, Ukamba,
Kenya Colony of such a rain-forest species as homeana. Sieben-
rock (1916 :5) evidently entertained doubts for he made enquiries
of Lampe respecting it and was assured that the specimen in

404 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

question had been presented to the AViesbaden Museum by a
missionary who had personally picked it up at Kiu about 1912.
Dr. F. Neubar, Director of Wiesbaden Museum, informs us
(7.i.55) that no reference to such a specimen can be found in
their catalogue, and also that much of their material was lost in
a fire.

In the absence of the specimen, speculation seems idle for many
alternative explanations are possible. The simplest is to assume
a transposition of specimens in the museum ; such things have
been known to occur ! Again there may have been a mistake as
to the provenance of that particular specimen on the part of the
collector, who may have previouslj'^ been stationed at some mis-
sion in West Africa. It might be pointed out that in 1912 there
were no dining cars on the trains plying between Lake Victoria
and Mombasa, so that it was customary to disembark all passen-
gers at Kiu (alt. 4861 feet) for meals at the station restaurant.
Thus the possibility of the young tortoise having escaped from
the custody of some passenger, homeward bound from the eastern
Congo, should not be overlooked.

K. homeana does occur in the extreme eastern Congo ; erosa,
continues eastward, though s]jarsely, in Uganda. The two agree
in the serrated nature of the posterior edge of their carapaces so
it is barely possible that Siebenrock might have confused them,
though this is unlikely as he {loc. cit.) cites the differing scutel-
lation of their respective forearms — 4 or 5 for erosa, 5 to 8 in
the case of homeana. In this respect homeana approaches helli-
ana, which has 5 to 9 scales on the forearm, and helliana is a
relatively common reptile in Ulvamba. A misidentification with
helliana would be likely only if the carapace of the Kiu juvenile
seemed abnormally serrate to Siebenrock.

K. homeana has been erroneously recorded as occurring in
Guadeloupe and Demara, South America, by Gray (1831c),
followed by Dumeril and Bibron (1835).

KiNixYS EROSA (Scliweigger)

1802. Teatudo Denticulata Shaw (not of Linnaeus), 3, p. 59, pi. .xiii.
1812. Testudo erosa Schweigger, Konigsberger Arch. Naturwiss. Math., 1,
p. 321: "America septentrional! (Shaw)."

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA

405

1814.

Schweigger, 52 (reprint of 1812).

1826.

Testudo Schopfii Fitzinger, Neue Classif, Kept., p. 44: No localitr

{nom. nov.).

1836.

Fitzinger, 121,

1827.

Kinixys castanea Bell, Trans. Linn. Soc. London, 15: p. 398, pi. xvii,

fig. 1 : Africa.

1831c.

Kinixys^ erosa Gray, 16.

1835.

Dumeril and Bibron, 165.

1844.

Gray, 12.

1851.

Dumeril and Dumeril, 6.

1855.

Gray, 13.

1856a.

Dumeril, 372.

1857.

Jan, 35.

1859.

Cope, 294.

1860.

Dumeril, 162.

1862a.

Strauch, 24.

1863f.

Gray, 197.

1864a.

Gray, 170 (reprint of 1863f).

1865.

Strauch, 39.

1866a.

Bocage, 40.

1870e.

Gray, 14.

1873b.

Gray, 17.

1873h.

Gray, 320, pi. xii, fig. 2.

1874.

Reicheuow, 298.

1875a.

Peters, 196.

1877c'.

Peters, 611.

1880c.

Vaillant, 34, 88.

1882a.

Muller, F., 165.

1882.

Pechuel-Loesche, 278.

1884a.

Rochebrune, 17 (ignored).

1884b.

Sauvage, 200.

1886.

Vaillant, 137.

1887a.

Miiller, F., 295.

1888a.

Boettger, 12.

1889a.

Boulenger, 141, figs. 40-41.

1890.

Biittikofer, 436, 478.

1890.

Strauch, 61.

1893a.

Boettger, 8.

1893c.

Matschie, 208.

1895a.

Bocage, 1.

1895e.

Werner, 127 (Cininyxis).

1897.

Sjostedt, 6.

1 Variously misspelled Kinyxis, Cinyxia and Cinixys in a few of the citations
following.

406

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY'

1898. Jeude, 5.

1898a. Werner, 203.

1900. Boulenger, 447.

1901. Gadow, 364, fig. 82.
1902e. Tornier, 664.

1 905. Barbier, 84, pi. iii.
19051". Boulenger, 183.
1906i. Boulenger, 197.

1906. Johnston, 820, 833.
190Ga. Mocquard, 479.

1907. Johnson, 11, 68, photo.
1907. Siebonrock, 2, fig. — , pi. i.
1908c. Kanimerer, 775, fig. 15.
1909a. Siebenrock, 509.

1910. Sternfeld, 4, fig. 6.

1911. Lampe, 145.
1912b. Werner, 424, fig.

1916. Siebenrock, 4, figs. 1-2.

1917. Sternfeld, 408, 411.
1919g. Boulenger, 11.

1919. Schmidt, 403, 598, pi. xi, fig. 1, map 3.

1919. Siebenrock, 275.

1920. Werner, 424, fig. — .
1929. Flower, 25.
1929h. Loveridge, 14.

1930a. Barbonr and Loveridge, 770.

1933. Hamerton, 455.

1933ni. Witte, 67.

1934a. Mertens and Miiller in Bust, 8.

1937c. Loveridge, 269.

19371'. Loveridge, 502.

1937b. Monard, 145.

1937. Buekes, 101, pi. x, fig. 7.

1941. Witte, 106.

1941e. Loveridge, 115.

1942b. Pitman, 214.

1949. Conant and Hudson, 4.

1949. Moyle, 582.

1950. Williams, 551.

1951. Monard, 168.

1952. Aurelio Bassilio, photo facing p. 97 (as Cinisis).

1953. Witte, 15.
1954d. Laurent, 296.

LOVERIDGE AXP WILLIAMS: AFRICAN CRYPTODIRA 407

1954. Noel-Hiune, 73, pi. v.

1955. Cansdale, 89, 103.

1839. Klniris dentwulata Hallowcll (not of Linnaeus), 161-169, pis.
viii-ix.

Common names. Schweigger's Hinged-Tortoise (Flower) ;
mbulutobe (on Loango Coast :Pechiiel-Loesche).

Illustrations. Both Shaw (1802) and Bell (1827) provide good
drawings of the shell of this distinctive species; more recently
Siebenrock (1907) figures the entire tortoise, and excellent pho-
tographs from life appear in Johnson (1907), Schmidt (1919)
and Noel-Hume (1954).

Description. Beak weakly to strongly hooked, unicuspid; edge
of jaws not dentate; prefrontal divided longitudinally; frontal
large or broken up; upper head shields small, irregular; fore-
limb anteriorly covered with a few scattered, extremely large,
unequal, imbricate or non-imbricate, more or less pointed scutes
which on the anterior edge form a longitudinal series of 4 to 5
from elbow to outer claw ; claws 5 ; hinder side of thigh without
enlarged tubercles; heel usually without any spurlike tubercle,
if present situated low on the heel ; claws 4 ; tail terminating in a
horny, clawlike tubercle that is much larger in S $ .

Carapace flattened dorsally, sides sloping, not or but scarcely
to moderately notched in nuchal region, anterior margin strongly
expanded and slightly to strongly reverted, spinose in young,
posterior margin not expanded, more or less strongly reverted
and strongly serrated, spinose in young, rear end in young slop-
ing gradually, in adults more steeply, if vertical only from the
middle of the fifth vertebral ; dorsal shields concentrically stri-
ated, neither swollen nor convex (except as noted below) ; nuchal
absent or rarely present, elongate, ^ occasionally developed on the
underside only ^ ; vertebrals 5, first broader than long in young,
broader than long, as broad as long, or longer than broad in
adults, second to fifth broader than long, second broader or sub-
equal to fifth in young, narrower in adults, fifth flat in hatch-
lings, more or less convex in adults, broader than the costals;
costals 4, rarely 3,^ not or but rarely forming an angle wdth the

1 Present in a Xola tortoise (M.C.Z. 44285).

2 In a Sangmelima specimen (Siebenrock : 1916).

'■ Three in a Landana tortoise whose missing costal was replaced by broadened
fourth and fifth vertebrals (Strauch : 1890).

408

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

marginals; marginals 11, rarely 12/ the anterior much longer
than the posterior ones; supracaudal undivided, more or less
reverted in both sexes.

Front lobe of plastron very thick, projecting beyond the
anterior border of the carapace, openly or deeply notched, bifid ;
gulars paired ; axillaries 3-4, the innermost small, the one or two
outermost more or less concealed ; inguinal large, in contact with
the femoral ; hind lobe very short, truncate, not or very broadly
notched posteriorly.

Plastral formula: Abd>h>(g, p, f, an, all extremely vari-
able).

Fig. 48. Skull of E'mixys ei'osa (A.M.N.H. 69727). Condylobasal length
53 mm.

(S. McDowell del.)

For description of a pathological individual, cf. Tornier
(1902c).

Color. Carapace of hatchlings pale brown, nearly uniform,
darkening with age until almost black while the centers of the
shields remain somewhat lighter ; in older juveniles and all adults
irregularly stellate patches of light yellow appear on the outer
margins of the costals, also in smaller areas on the vertebrals,
upper portions of the costals and the anteriormost and posterior-
most marginals.

1 Twelve on left side of a Sakbaj-eme specimen (M.C.Z. 34014).

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA

409

Fig. 49. Internal view of plastron of Kinixys erosa (Yale Mus. 4586).

(P. Washer del.)

410 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Plastron yellow, more frequently retained on the gulars than
on the remaining shields whose centers are brown or black,
sometimes so extensive as to reduce the yellow to narrow lines
along the sutures or occasionally a midplastral yellow zone.

Size. Carapace length of largest S (A.M.N.H. 10023), from
Banalia, 323 mm., breadth 236 mm., height 122 mm. (Schmidt).
Carapace length of largest 9 (M.C.Z. 52169), from Inferri
Chiefdom, 260 mm., breadth 172 mm., height 106 mm.

Sexual dimorphism. The plastron of older <J <5 is more or
less concave, their tails considerably longer than those of $ 9 .

Breeding. Herbert Lang was told by the Congolese that
erosa deposits its eggs in sandy ground and covers the site with
leaves. One of four eggs laid by a captive tortoise measured
31 X 36 X 40 mm. (Schmidt :1919 :406) .

Longevity. Three years and five months in Philadelphia Zoo
(Conant and Hudson :1949).

Diet. Apparently omnivorous. In captivity one ate apples,
pears, sweet potatoes and meat — both raw and cooked ( Hallo-
well :1839). Possibly Biittikofer (1890:436) is repeating native
stories when he states that erosa eats insects and snails.

Hamerton (1933:455) records a captive tortoise djdng of
faecal concretions.

Enemies. Eaten by Liberians (Biittikofer) and by Gold Coast
natives (Cansdale^) ; also by the inhabitants of the Loango
Coast, who convert the shells into ornamental receptacles
(Pechuel-Loesche :1882). Eaten also by the Congolese who em-
ploy dogs to track the tortoises whose odor betrays them (Lang
in Schmidt).

Habits. In Liberia erosa frequents the shady banks of rivers
(Biittikofer). On the Loango Coast Pechuel-Loesche (1882) was
informed by the natives that these tortoises actually live in the
brooks and rivers, exclusive of tidal estuaries, and bury in the
mud. Despite their unwebbed feet they swim well, and captive
erosa dived to obtain their food in a rather deep basin of water.
These aquatic habits, continues Werner (1920), were confirmed
by Ussher who informed him that his captive specimen remained
in water for as much as a month.

The oft-repeated statement that these tortoises are slow and
lethargic, says Kammerer (1908c), is due to captive specimens

1 In letter of 30.xi.l954.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 41 1

being subjected to uneongeuial conditions such as too little mois-
ture or too low temperatures.

The best account is given by Lang {in Schmidt :1919 :405), who
states that erosa "is fond of marshy sites. During the day it
hides beneath logs, roots, and heaps of dead and living vegeta-
tion, sometimes comijletely covering itself with debris. The pro-
jecting forked gular portion of the plastron and the slightly
upturned edges of the carapace are very useful in this task ; the
head is partly retracted, and the feet then push and wedge until
the turtle is hidden from view.''

Localities. U<j(ni(la: JNIabira Forest; Nabea. Cabinda: Chin-
choxo ; Landana. Jielgian Coiuju: *Avakubi; Bafwabaka, "Baf-
wasikuli" ^ ; Banalia; Bcni ; Bikoro; Djalasinda; Duma, Ubangi;
Ekibondo; Gamangui ; Kabambare; *Kindu (Kindo :U.S.N.M.) ;
Koloka; Kunungu; Lake Tanganyika region; "Lissimu"^;
Masisi in Pare National Albert; Medje (Madie) ; Niapu ; Panga;
Poko; *Saidi's Village; iStanlej'ville ; Walikalc. French Equuto-
rial Africa: Fenian Vaz (Fernand Vas) ; Gabon; Kama (Cam-
ma) ; Loango; ]\iassabe (Massaba; Massabi) ; *Nola; *Ntyanga
(? Nyanga, or Ntchonga Lake), French Congo (U.S.N.M.) ;
Ogoue (Ogobai). Eio Mimi: Rio Muni. French Cameroon: *Bi-
pindi (CM.) ; Campo (or Kampo) ; Douala; *Efoulen (Efulen) ;
*Kribi; *Lolodorf (CM.); *Metet (CM.); Ndikinimeki ; *Sak-
bayeme ; *Saiigmelima (CM.); Yaounde (Jaunde). British
Cameroon: Bonge, Meme River; Buea; Cameroon Mtn. ; Johann
Albrechtshohe ; Nssakpe; Ossidinge. Gold Coast: Aburi; Anko-
bra (Ancober) River; Ashanti. Ivory Coast: Adiopodoume ;
Vapo Nord (both V.Aellen). Liberia: *Bonuta (Bonutah) ;
*Cape Palmas; *Cuttington College, Suacoco ; *Harbel ; *Lenga,
Farmington River; Mesurado River; *Paiata (Peahtah) ; St.
Paul River ; *Zorzor. Sierra Leone : Barri Chief dom ; *Imf erri
Chiefdom; ^northeast Sierra Leone. Gambia. (B.M. 43.2.29.1
and i shell).

Range. Uganda and Belgian Congo northwest to Gambia.

Senegal (Rochebrune) is rejected pending confirmation. Erro-
neously recorded from North America (Shaw), Guadeloupe
(Dumeril and Bibron), and Mozambique (Gray).

1 In Stanleyville district : a Cliiisly Ideality (Roiilen-er : 1919};).

412 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Family TRIONYCHIDAE

1828b. Trionychidae Bell, Zool. Jour., 3, p. 515.

Definition. Cryptodirous testudinates of fully aquatic habit.
Horny shields except for obscure rudiments absent, represented
by naked skin.

Skull without nasal bones; prefrontals always in contact dor-
sally, usually with descending processes that are well separated
when present; temporal region deeply emarginate posteriorly;
parietal never meeting squamosal ; postorbital sometimes re-
duced ; quadrate surrounding stapes ; post-otic antrum well devel-
oped; parietal with well developed descending processes; upper
jaw without ridges on its triturating surfaces; vomer usually
present, if present short, noi separating the palatines ; mandible
with well developed coronoid bone.

Neck vertebrae without a biconvex centrum, all opisthocoelous
except the eighth which is doubly convex in front but united to
the centrum of the first dorsal only by ligament ; entire coracoid
blade broad and saber-shaped ; tuberosities of humerus widely
separated ; trochanteric fossa of femur widely open ; phalanges
with condyles ; claws 3.

Carapace not directly united to plastron ; carapace not hinged,
but plastron with some flexibility betAveen anterior elements;
pleural bones always forming a solid disk, but peripheral bones
absent (possibly represented by a few ossifications in one genus) ;
nuchal overlapping, or overlapped by, the first pleural, without
typical costiform processes or attachment surface for eighth
cervical ; neural bones hexagonal, mostly short-sided behind, vari-
able in number ; pygals absent ; plastron with some development
of fontanelles; entoplastron absent, mimicked by a separated
posterior portion of the epiplastra.

Bange. North America; Africa; Asia.

Fossil record. First known from the Lower Cretaceous of Asia
(with a possibly ancestral type — Sinaspideretes Young and
Chow, 1953, x\cta Scientia Sinica 2:226-227, pi. 6 — in the Up-
per Jurassic or Wealden of the same region). A good record in
North America since the Cretaceous ; in Europe from Paleocene
to Pleistocene. Known in Africa since the Lower Miocene.

Remarks. The trionychids or "soft-shelled turtles"^ have

1 A niisudiner that has I'requcutly caused misconceptions. Actually iu these
turtles the carapacial disk is at least as solidly ossified as in other testudinates ;
only the fleshy margins are soft, except in Lissemys, which retains peripherals.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 413

often been separated as one of the major groups within the
turtles, coordinate in rank with the Pleurodira, with the marine
turtles, and with the land and freshwater forms of more normal
or primitive habitus. There are many characters to support such
a separation. The peculiar plastron with its callosities and the
unique modification of the anterior elements (Williams and Mc-
Dowell :1952) ; the skull with its peculiar intermaxillary fora-
men ; the singular fragmented basal plate of the hyoid ; the amaz-
ing connection of cervical column and dorsal vertebrae (by zyga-
pophyses only) ; the sickle-shaped scapula; the unique hyper-
phalangy of the digits; the remarkable penis; the absence of a
urogenital sinus, etc., are a formida!)le complex of features, by
any one of which a trionychid may be recognized.

A full analysis of the Trionychidae, setting up phyletic lines
and considering carefully both fossil and Recent forms, has not
yet been attempted. Yet there are indications such an analysis
is less difficult than reputed. It is true that the lierpetologist
dealing only with externals, is perplexed by the lack of characters
he can utilize, and is further disturbed by ontogenetic change (as
in the shape and number of the plastral callosities), as well as by
individual variation (as in the breadth of the maxillary triturat-
ing surfaces). In these turtles, however, even more than in
others, it is necessary to go below the surface and, for example,
use osteological characters. Consequently for every species we
offer two descriptions, one of externals, including color, and a sec-
ond dealing with its osteological characteristics.

When the osteology is taken into account the African species
are remarkably distinct. We have not carried through a stud}'
of the osteological characters of the extra-African species, but
preliminary explorations by one of us, and by previous workers,
convince us of its practicality. Siebenrock (1902b), for example,
has demonstrated the usefulness of plastral characters. He pre-
sents a key to the living trionychids based entirely on the plas-
tron, also an atlas of plastral bone conformation. Both atlas and
key are still very useful. In his key Siebenrock did not empha-
size the number or shape of the callosities — very wisely, for it is
the latter that (because of individual and age variation) are
most likely to make use of his atlas difficult and deceptive. On
the contrary, if attention is paid to the shapes and processes of

414

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

the underlying bones, the characters employed by Siebenrock suc-
cessfully separate all but the most closely related species. All
the African species are clearly separable by their plastral char-
acters.

The skulls of trionychids, to mention another area of osteo-
log'ical study that has proved rewarding in the African species,
have not been exploited sufficiently. More attention has been paid
to general shape and to the varying breadth of the maxillary
triturating surfaces than to suture patterns and details of mor-
phology which promise to be much more useful.

Table 8

Characters of the two sections of the Trionychidae
Group I Group II

Cydanoriis ;

Trionyz; Pelochelys ; Chitra.

Plastron without cutaneous femoral
valves.

Sculpture characteristically of
anastomosing ridges.

Nuchal bone without conspicuous
ventral ridges, its posterior margm
overlying the first pleural (costal
bone).

Hyo- and hypoplastra distinct.

Xiphiplastron gripping lateral
prong of posteromedial process of
hypoplastron.

Posterior border of pterygoids
without an ascending process that
makes sutural contact with the
opisthotic.

Lissemys;
derma.

Cyclo-

Plastron with cutaneous femoral
valves under which the hind limb
may be concealed.

Sculpture more often tuberculato.

Nuchal bone with a conspicuous
ventral ridge on each side extend-
ing posterolaterally to underlie the
first pleural (costal bone).

Hyo- and hypoplastra fused.

Xiphiplastron gripping middle
prong of the three prongs of the
posteromedial process of hypo-
plastron.

Posterior border of pterygoids
with an ascending process forming
a suture with the opisthotic and
greatly restricting the fenestra
postotica.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 41 5

Ilelationships and Grouping of the Living Trionyehids

Since no thorough analysis of all living trionyehids has been
carried out. any discussion of relationships must necessarily be
tentative. Nevertheless, it seems advisable to attempt to place
the African species in their proper perspective within the family.
Our approach is a conservative one. African trionyehids, and
the family as a whole, fall into two very distinct major groups
which have often been separated as distinct subfamilies. The
division, very clear-cut, is based on a conspicuous external char-
acter and several internal ones, as set out in Table 8.'

As Group 1 contains only a single African representative —
Trionyx triunguis — it would be out of place here to discuss its
subdivisions in detail. Our purpose is merely to indicate the
probable position of triunguis in our t( ntative arrangement of
the group, which may be subdivided as follows :

Subgroup ]. Relatively unspecialized species primarily Ori-
ental in distribution, primitively with a dorsal pattern of large
ocelli.

A. Those with a preneural.

1. Trionyx hurum

2. T.leithii

3. T. nigricans

4. T. gangeticus

B. Those with no preneural and with a mandibular syai-
physial ridge.

5. T. formosus

6. T. cartilagineus

C. Those with no preneural but without a symphysial
ridge.

7. T. suhplanus

Subgroup 2. Species with highly modified skulls, primarily
Oriental in distribution.

1. Pelochelys hihronii

2. Chitra indica

1 Siebenrock (1907; 25 footnote) invites attention to a partial restriction of
the fenestra postotica in Dogania (= Trionyx) subplana and Trionyx sinensis bv
a descending process of the opisthotlc. This condition, which we have vcfified for
Trionyx suhplanus, differs strikingly from the condition in Li.ssemys, (■tc, in
which the pterygoid forms an ascending process. However, in some Trionyx
triunguis (see p. 429 and Fig. 54) the pterygoids may send up an ascending process
like that of Lissemys, but in this case it fails to niake contact with the opisthotic.

416

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Subgroup 3. Relatively unspecialized species primarily extra-
Oriental in distribution, primitively without a dorsal pattern of

large ocelli.

muticus

spinifer
emoryi

ferox

cartilagineus

euphraticus
swinhoei

sinensis
steindachneri

aubryi
frenatum

formosus

Fig. 50. Dendrogram of trionychid species relationships. The purely

hypothetical horizontal line divides those species which may have evolved

north of tho ancient Tethys sea from those which may have evolved south

of it.

(P. Washer del.)

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 417

A. Those whose 8th pleurals are typically not reduced.

1. Trionyx triicnguis

2. T. sinensis

3. T. steindachneri

B. Those whose 8th pleurals are typically partially reduced.

3. T. eupJwaticus

4. T. swinJioei

C. Those whose 8th pleurals are typically much reduced or
absent.

5. T. ferox

6. T. emoryi

7. T. spinifer

8. T. muticus

This conception of the most probable relationships within the
Trionychidae is presented as a dendrogram (see Fig. 50).

Trionyx tri^inguis, as indicated on the dendrogram, is phyleti-
eally quite remote from the other African trionychids. It and
they represent tAvo invasions of Africa. Curiously, however, so
far as the available fossil record indicates, the two invasions may
have been simultaneous. Both groups are clearly represented in
the Lower Miocene, which offers the first record of trionychids in
Africa.

An overall view of the characters of African trionychids is
given in the following table.

In Group II there is, in addition to the two African genera,
one Indian genus — Lissemys. Lissemys, while clearly a member
of the group, seems not to be very closely related to Cyclanorbis
or Cycloderma. From both, as well as from all other trionychids,
it differs in the possession of peripheral ossifications (usually
regarded as non-homologous with the peripherals of other tur-
tles).

The African members of Group II appear to form a series of
their own. Each species has its individual specialization and
yet appears to form part of a sequence which, beginning with
the most primitive form, is as follows: Cyclanorhis elegans, C.
scnegalensis, Cycloderma fre7iatum, C. aubryi. Alone of the
African series, Cyclanorhis elegans retains prefrontal connec-
tion with the vomer (primitively present in all turtles; lost,
except in this series, only in Chitra and the suborder Pleuro-

418

BULLETIN: MUSf:TT:\I OF COMPARATIVE ZOOLOGY

Table 9
Characters of the African Species of Trionychidae

triunguis
premaxillary
bone small but
distinct

elegans
premaxillary
bone small but
distinct

.lenegalenms
as in elrgans

frrvatum
preinaxillarv' tione
a|>par'>ntly absent,
at most scalelike

aubryi
aa in frrnai'm

distance from
tip of premaxil-
lary to orbit
much greater than
long diameter
of the orbit

distance from
tip of prejnaxil-
lary to orbit
less than the
long diameter
of the orbit

distancf from
tip of premaxil-
lary to orbit
creator than the
long diameter
nf the orliit

distance from
tip of premaxil-
lary to orbit
suhequal to long
diameter of the orbit

as in frenatum

height of orbit
somewhat greater
than the inter-
(irbital width

height of orbit
about twice the
interorbital

.fiilth

as in eletjanx

as in cUgn-n.'i

height of orbit
about one and one
third the inter-
orbital width

prefrontal bones
meet the vomer

prefrontal bones
meet the vomer

prefrontal bones
not meeting vomer

as in }tenegali'n.<;is

as in sene'gahn.iu

intermaxillary
foramen large,
ovoid

intermaxillary
foramen absent

Miterinaxillary
foramen small,
narrowed
anteriorly

as in senegalensi.'; as in senegaiens-^s

prechoanal part
of vomer con-
cealed by union
of maxillary

xipliiplastra
long, triangular

prechoanal part
of vomer con-
cealed by union
of maxillary
medial processes

vomer I'xposed
throughout its
entire length
or concealed

tlegans

as in elfgam

jngal enters

jugal enters

as in tl<-gans

ii.s in elt'jayis

jugal excluded

orbit

orbit

jugal separated

jugal narrowly

as in clfijaus

jngal briiadly

as in frcnaium

from or only

in contact with

ill contact with

narrowly in con-

the parietal

the parietal

tact with parietal

neurals 8 ;

neurals 6-8 -f

neurals l-.j +

neurals 7-8 +

neurals 7-8 +

no preneural

preneural

preneural

p^'eneural

preneural

callosities in

callosities in

callosities in

callosities in

as in fifnatuni

adults 4-5.

adults 2-4

adults 7-0 or mure

adults 7

sometimes an

never any

supernumerary

callosities on

as in frenaliim

azygous callosity

callosities in

callosities in

prejdaslra and

in gular region

gular region

gular region

fused epiplastra

xiphiplastral

xiphijilastral

as in fitgans

xiphiplastral

us in frenatum

callosities

callosities

callosities

always present

weak or absent

strongly ileveloped

xiphiplastra

posteriorly

pointed

xiphiplastra
posteriorly
broad and
notched

as in sciieyaUnsis as in senegtdf nsis

color of
hatchltng:
head profusely
spotted with
lighter; thrott
spotted and
vermiculated ;

Mlor of
hatchling:
head profusely
spotted with
yellow ; throat
vemiiculated ;

color of
hatchling;
head vaguely
mottled ;
throat
immaculate ;

color of
hatchling:
head grayish
with .") lines;
throat nearly
immaculate ;

f oJor of

hatchling:

head light brown

with 5 lines; throat

indistinctly

flecked ;

carapace olive
to dark brown,
uniform or much
speckled :

carapace green
with irregular
yellow blotches ;

carapace gray-
brown vaguely
mottled ;

carapace gray
or green,
nearly uniform ;

carapace light
brownish with a
chocolate brown
vertebral stripe ;

plastron white,
immaculate or
with dusky
vermiculations
anteriorly

plastron spotted
with yellow to
periphery

plastron with a
brownish blotch
in centre, peri-
phery immaculate

plastron whitish
with small black
blotches

plastron with two
broad V-shaped,
chocolate-brown
markings, a small
one on gular
region, a larger one
from axillae to tail

size of disk to
950 mm.

size ;jf disk to
440 mm.

size of disk to
350 mm.

size of disk to
470 mm.

size of disk to
330 mm.

LOVERIDGE AND WILLIAMS: AFRICAN CRTPTODIRA

419

dira).^ C. senegalensis lacks the vomer-prefrontal connection
but, as in elegans, has a narrow postorbital bar. This bar is tre-
mendously broadened in both species of Cycloderma, resulting
in extensive contact between parietal and jugal, though in primi-
tive fashion C. frenatum still retains the jugal entering the orbit.
In C. auhryi the jugal is excluded from the orbit, thus complet-
ing the series of cranial modifications by one that is unique in the
family.

B

Fig. 51. Forefoot scalation in certain trionychida. A, Trionyx trmnguL%
(M.C.Z. 16509) ; B, Cyclanorhis elegans (B.M.) ; C, Cyclanorbis senegalensis
(Z.M.U.).

(P. Washer del.)

Key to the Trionychidae of Africa

1. Femoral flaps absent; nuchal somewhat overlapping second rib; hTo- and
hypoplastra separate; lateral prong of posteromedial process of hypo-
plastron gripped by xiphiplastron; pterygoids not joining opisthotic ;
fenestra postotica not or not much restricted; head spotted. Eange:
Egypt and Eritrea south to Laie Eudolf, southwest to Lake Albert and

Angola, northwest to Senegal Trionyx triunguis Forskal (p. 3)

Femoral flaps present; nuchal not overlapping second rib, more or less
projecting under second pleural; hyo- and hypoplastra fused; middle
prong of posteromedial process of hyo-hypoplastron gripped by xiphi
plastron; pterygoids joining opisthotic, greatly restricting fenestra
postotica 2

1 In trlonychids the connection Is mostly formed by ascending processes of the
vomer meeting the prefrontals. In primitive turtles and in many modern forms
(e.g. marine turtles), the prefrontal and vomer share about equally in this pe-
culiar bony strut. In testudinids the connection is almost entirely formed from
the prefrontals.

420 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

2. Head spotted or partly vermiculated, at least in young; postorbital arch

narrower than diameter of orbit; preplaatra long, angular; no trace

of a xiphiplastral commissure 3 (CycJanorhis Gray)'

Head striped, at least in young ; postorbital arch wider than diameter
of orbit; preplastra short, straight; a xiphiplastral commissure
present 4 {Cycloderma Peters)

3. Carapace of hatchlings with large yellow blotches; plastral area with a

large dark central blotch profusely spotted with yellow, at least an-
teriorly; no trace of gular callosities at any age; xiphiplastra pointed
posteriorly; prefrontal bones meeting vomer. Range: Sudan; Nigeria;

Togo Cyclanorbis elegans Gray (p. 438)

Carapace of hatchlings with small black spots; central plastral area
more or less blotched with dark; gular callosities, 4 or more in adults,
present even in the half grown; xiphiplastra broad and notched pos-
teriorly; prefrontal bones not meeting vomer. Range: Sudan west

through French Cameroon to Senegal

C. senagalenxis (Dumeril and Bibron) (p. 443)

4. Carapace of hatchlings orange or orange brown with a narrow black

vertebral line anteriorly; jugal bone excluded from orbit; in adults
the azygous callosity is subcircular, large; hyo-hypoplastral callosities
in contact with xiphiplastral callosities by long straight sutures.
Range: Portuguese, French, and adjacent rivers of western Belgian

Congo Cycloderma aubryi (Dumeril) (p. 453)

Carapace of hatchlings pale gray or leaden 2 without a vertebral streak;
jugal bone enters orbit; in adults the azygous callosity is crescentic,
subcrescentic or semilunar, and smallest of all the callosities; hyo-
hypoplastral callosities separated from, or in contact with the xiphi-
plastral callosities by only a short suture. Range: Tanganyika
Territory (Rovuma River and Lake Nyasa) southwest through Nyasa-
land (Lake Nyasa and Shire River) to Mozambique (Zambezi River
and affluents) C. frenatum Peters (p. 459)

Genus TeiOXYX Geoffroy
1809a. Trionyz Geoffrey, Ann. Mus. Hist. Nat. (Paris), 14, p. 1. Type by

original designation: Trionyx aegyptiacus = Testudo triunguis

Forskal.
1830. Aspidonectes Wagler, Natiir. Syst. Amphib., p. 134. Type by Stej-

neger's designation: Trionyx aegyptiacus Geoffroy = Testudo

triunguis Forskal.
1835. Gymnopus Dumeril and Bibron, Erpet. Gen., 2. p. 472. New name

for Aspidonectes Wagler.

1 A single juvenile specimen from the northwest shore of Lake Tanganyika will
not key out even generically on the characters here used and may be a new form.
See page 450. ,.,„„„ , ,>

2 Green in Zambezi turtles according to Peters (1822a, pi. i).

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 421

1835. PlatypeUis Fitzinger, Ann. Wiener Mus., 1. pp. 109, 120, 127. Type
by Fitzinger 's 1843 designation: Testiido ferox Schneider.

1835. Pelodiseus Fitzinger, Ann. Wiener Mus., 1. pp. 110, 120, 127. Type
by Fitzinger 's 1843 designation: Trionyx sineiusis Wiegniann.

1835. Amyda Fitzinger, Ann. Wiener Mus., 1. pp. 110, 120, 127. Type by
original designation : Trionyx suhplamis Geoff roy.

1843. Potamochelys Fitzinger, Syst. Rept., p. 30. Type by original desig-

nation: Trionyx javanious Wiegmann = Testudo cartilaginea Bod-
daert.

1844. Tyrse Gray, Cat. Tortoises Brit. Mus., p. 47. Type by tautonomy:

Trionyx nilotimi^ Gray =; Testudo triunguis Forskal.
1844. Dogania Gray, Cat. Tortoises Brit. Mus., p. 49. Type by monotypy:

Trionyx suiplanus Geoffroy.
1864e. Eafetus Gray, Proc. Zool. Soc. London, p. 81. Type by monotypy:

Testudo euphratica Daudin.
1864e. Aspilus Gray, Proc. Zool. Soc. London, p. 83. Type by original

designation: Trionyx cariniferiis Gray = Testudo cartilaginea

Boddaert.
1869c. Landemania Gray, Proc. Zool. Soc. London, p. 215. Type by mono-
typy: L. irrorata Gray = Trionyx sinensis Wiegmann.
1869c. Fordia Gray, Proc. Zool. Soc. London, p. 219. Type by monotypy:

F. afri<;ana Gray = Testudo triunguis Forskal.
1869c. Callinia Gray, Proc. Zool. Soc. London, p. 221. Type by Stejneger's

designation: Trionyx spinifer Lesueur.

1872. Nilssonia Gray, Ann. Mag. Nat. Hist. (4), 10. p. 332. Type by

monotypy: Trionyx formosus Gray.

1873. Isola Gray, Proc. Zool. Soc. London, p. 51, fig. 2. Type by monotypy:

Trionyx peguensis Gray = Trionyx formosus Gray.

1873. Ida Gray, Proc. Zool. Soc. London, p. 55, fig. 7. Type by monotypy:
Trionyx ornuta Gray = Testudo cartilaginea Boddaert.

1873. Oscaria Gray, Ann. Mag. Nat. Hist, (4), 12, p. 157. Type by mono-
typy: 0. swinhoei Gray.

1880. Yuen Heude, M6m. Hist. Nat. Emp. Chinois, I. p. 18. Type fide
Stejueger: Y. leprosus Heude = Oscaria sivinhoei Gray.

1880. Psilognathus Heude, Mem. Hist. Nat. Emp. Chinois, 1. p. 24, pi. ii.
Type by monotypy: P. laevis Heude = Trionyx sinensis Weig-
mann.

1880. Temnognathus Heude, Mem. Hist. Nat. Emp. Chinois, 1. p. 25, pi. iii.
Type by monotypy: T. mordax Heude = Trionyx sinensis Wieg-
mann.

1880. Gomphopelta Heude, Mem. Hist. Nat. Emp. Chinois, 1. p. 27, pi. iv.
Type by monotypy: G. officinalis Heude = Trionyx sinensis Wieg-
mann.

422 BULLETIN : MUSEUM OP COMPABATIVE ZOOLOGY

1880. Coelognathus Heude (not of Hessling: 1852), M6m. Hist. Nat. Emp.

Chinois, 1, p. 29, pi. v. Type by monotypy: C. novemcosiatus

Heude = Trionyx sinensis Wiegmann.
1880. Tortisternum Heude, Mem. Hist. Nat. Emp. Chinois, 1, p. 31, pi. vi.

Type by monotypy : T. novemcostatum Heude = Trionyx sinen.-<i.s

Wiegmann,
1880. Ceramopelta Heude, Mem. Hist. Nat. Emp. Chinois, 1, p. 33, pi. vii.

Type by monotypy : C. latirostris Heude = Trionyx sinensis Wieg

mann.
1880. Coptopelta Heude, Mem. Hist. Nat. Emp. Chinois, I, p. 34, pi. viil.

Type by monotypy: C. septemcostata Heude = Trionyx sinensis

Wiegmann.
1880. Cinctisicrnum Ileude, Mem. Hist. Nat. Emp. Chinois, 1, p. 36, pi. \x.

Type by monotypy: C. hicinctum Heude^= Trionyx sinensis Wieg-
mann.
1903. Aspidereies Hay, Proc. Amer. Philos. Soc, 42. p. 274. Type by

original designation : Trionyx gangeticus Cuvier.
1944. Euamyda Stejneger, Bull. Mus. Comp. Zool., 94, p. 9. New name for

Amyda mutica Agassiz.
Synonymy. The listing of Amyda Schweigger MS in the
synonymy of his Trionyx by Geoff roy-Saint-Hilaire (1809a :15),
neither constitutes the proposal of a new name nor validates it.
Nor is Amyda validated by its brief mention by Oken (1816:
Lehrbuch der Naturgeschichte, Teil 3, Abt. 2, 348). The first
acceptable usage would appear to be that of Fitzinger (1835:
120) as listed above, and the first explicit designation of a type
species that by Fitzinger (1843:30) — Amyda subplana. Since
we regard subplanus as a species of Trionyx, the necessity of
substituting Amyda Fitzinger, 1836, for the currently used
Dogania Gray, 1844, does not arise.

In regard to the type of Trionyx Geoffroy, we concur with
Schmidt (1953, Checklist of North American Amphibians and
Reptiles, p. 108) in believing that Geoffroy, when citing T.
aegyptiacus as giving an exact idea of the habitus and generic
characters of Trionyx, was consciously and with full intent des-
ignating the type of that genus. Admittedly the word 'Hype"
was not used by Geoffroy, but we must point out that no species
except the genotype can give an exact idea of the habitus and
generic characters of a genus, and that this is the sole function
and meaning of a genotype. Furthermore, we invite attention
to evidence that Geoffrey's words were accepted as a type

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 423

designation by at least one of liis contemporaries. Bory de
Saint Vincent (1828, Resume d'Erpetologie ou d'Histoire nat-
urelle des Reptiles [Paris], p. 77) has this to say: "C'est au
eelebre professeur Geoffroy de Saint-llilaire que Ton doit
I 'etablissement de ee genre [Trionyx] dont le type fut une tres
singuliere tortue du Nil que nous avons fait representer dans
notre planche 6^. ' '

Definition. Cutaneous femoral flaps absent.

Skull without maxillary ridging; intermaxillary foramen
moderate to large ; prefrontal always with connection to vomer ;
jugal not or but scarcely in contact with parietal ; postorbital
arch narrower than diameter of orbit ; pterygoids never joining
opisthotic; fenestra postotica unrestricted.

Carapace without prenuchal bone, without peripheral ossifica-
tions ; nuchal bone not notched laterally, no raised ventrolateral
processes, its lateral margin tending to overlie the second rib;
between the first pleurals are 1 or 2 neurals; neurals 7 to 9 ;
pleurals 7 or 8 pairs, the last one or two in contact medially.

Plastron wnth preplastra long and angular; hyo- and hypo-
plastra separate, the latter 's posteromedial process with a prong
that is inserted between the two anterior prongs of the adjacent
xiphiplastron ; callosities variable.

Range. North America; Africa; Asia.

Fossil record. A good record since the Cretaceous. First re-
ported in Africa from the Lower Miocene of Kenya; first re-
corded in North Africa from the Pliocene ; present in the western
portion of North Africa in the Pleistocene.

Trionyx triunguis (Forskal)

1775. Testudo triunguis Forskal, Descr. Anim. Avium. AmpMb., p. ix:

Xile River.
1789. Gmelia, 1039.

1792. Schneider, 280.

1809a. Trionyx aegyptiacics Geoffroy, Ann. Mus. Hist. Nat. (Paris), 14,

p. 12, pis. i-ii: Nile River, Egypt.
1809b. Geoffroy, 368 (proof of priority over 1809a unobtainable).

1812. Schweigger (part), 286, 328, 364.

1814. Schweigger (part) (reprint of 1812), 16.

1820. Goldfuss. 179.

424

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1820. Merrem, 20.

1824. Mohring (on osteology: not seen).

1827a. Geoffrey, 115, pi. i, figs. 1-1'.

1828. Bory de Saint Vincent, 77.

1829. Bory de Saint Vincent, 2, pi. vi.
1829. Guerin, 6, pi. i, fig. 7 (as Tryonix).
1835. Eiippell, 3, footnote.

1856a. Dumeril, 377.

1856. Lichtenstein and Martens, 1.

1857. Jan, 36.

1860. Dumeril, 168, footnote.

1862b. Peters, 271.

1862a. Strauch, 175.

1865. Strauch, 126.

1870a. Steindachner, 326.

1874. Eeichenow, 298.

1877. Bruhl, pi. xxxiii, figs. 9, 13 ; pi. xxxiv, figs. 7, 10.

1880. Boettger, 208.

1880. Bruhl, pi. Ixix, figs. 5-6 (skull).

1880b. Vaillant, 797.

1880c. Vaillant, 54, pi. xxx, fig. 14.

1887. Lortet, 24.

1897. Sjostedt, 33.

1905. Barbier, 96.

1831c. Trionyx Niloticus "Shaw", Gray, Syn. Eept., p. 46: Nile Eiver.

1831b. Gray, 18.

1835. Temminck and Sehlegel, 31.

1845b. Eiippell, 298.

1855. Gray, 68.

1867. Baker, 44, 203, 374 (as Trionis).

1882a. Miiller, F., 164.

1882. Pechuel-Loesche, 277.

1929. Worthington, 123.

1835. Gymnopus aegyptiacus Dumeril and Bibron, 484.

1851. Dumeril and Dumeril, 22.

1851. Guichenot, 190.

1867a. Bocage, 218.

1884a. Eochebrune, 31, pi. iii, figs. 1-2.

1884b. Sauvage, 200.

1837. Trionyx labiatus Bell, Monog. Testudinata, text to col. pis.

(=: xviii-xx) : Sierra Leone.

1872b. Gray, p. 11, in Sowerby and Lear pis. lii-liv.
1843. Aspidonectes Aegyptiacus Fitzinger, 30.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA

425

1855.
1844.

1844.

1864c.

1870e.

1872c.

1873b.

1873g.

1859.

1869c.

1869b.

1870.

1873.

1873b.

1875a.

1876a.

1877c.

1882.

1889a.

1889.

1890.

1890.

1893a.

1893c.

1895a.

1896.

1896c.

1897g.

1897b.

1897.

1898.

1898.

1898a.

1900b.

1901.

1901c.

1902b.

1902c.

Fitzinger, 252.
Trionyx Mortoni Hallowell, Proc. Acad. Nat. Sci. Philadelphia, p.

120: Africa.
Tyrse nilotica Gray, 48.

Gray, 88.

Gray, 108.

Gray, 27.

Gray, 84.

Gray, 470.
Aspidoncctes aspilus Cope, Proc. Acad. Nat. Sci. Philadelphia, p.
295: Ovenga and Eembo Rivers, Fernan. Vaz (as Fernando Vas),
Gabon.
Fordia africana Gray, Proc. Zool. Soc. London, p. 219: Nile Riyer
at Khartouni, Anglo-Egyptian Sudan.

Gray, 191.

Gray, 100 (reprint of 1869c).

Gray, 43, figs. la-Id.

Gray, 77.
Trionyx triunguis Peters, 196.

Peters, 117.

Peters, 611.

Pechuel-Loesche, 277.

Boulenger, 254.

Hesse, 261.

Buttikofer, 437, 478.

Strauch, 113.

Boettger, 16.

Matschie, 208.

Bocage, 7.

Anderson, 95.

Boulenger, 17.

Boulenger, 277.

Moequard, 5.

Sjostedt, 7.

Anderson, 32, col. pi. iii.

Jeude, 10.

Werner, 204.

Boulenger, 447.

Gadow, 410.

Tornier, 68.
Siebenrock, 826, fig. 8.

Tornier, 665.

+26

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1902a. Werner, 348.

1904. Andersson, 9.

1906. Johnston, 820, 83:5.

1906a. Mocquard, 481.

1906a. Siebenrock, 827.

1908. Johnston, 929 (generic name only).

1908. Werner, 1826.

1909b. Pellegrin, 413.

1909a. Siebenrock, 600.

1910. Miiller, L., 623.

1910. Sternfeld, 7, fig. 16.

1911a. Masi, 37.

1917. Sternfeld, 417.

1923-?. Loveridge, 933.

1924b. Loveridge, 3.

1925b. Flower, 935.

1926a. Mertens, 152.

1927. Calabresi, 38.

1928b. Scortecci, 336.

1928b. Witte, 49.

1929. Flower, 51.

1929. Hummel, 376.

1930a. Scortecci, 216.

1932a. Parker, 229.

1983. Flower, 753.

1933ni. Witte, 68.

1933. Worthington, 38, 189 (as Trionix).

1937:1. Flower, 16, 37.

1937f. Loveridge, 489, 495, 503.

1943. Scortecci, 270, 283.

1948e. Cansdale, 71 (as Trionyx only).

1948. Deraniyagala, 29.

1950. Williams, 552.

1953. Durrell, 253, fig.

1884a. Gymnopus aspiltis Eochebrune, 32.

1893. Pelodiscus triunguis Baur, 220.

1899. Lcailiery tortoises Ansorge, 289.

1919. Amyda triunguis Schmidt, 598, 601.

1930a. Barbour and Loveridge, 770.

1934a. Mertens and Miiller in Eust, 12.

1940c. Mertens, 236, figs.1-5.

19.^5. Cansdale, 93, 104, fig. H6.

1948. Trionyx triunguis rudolfiamu<> Deraniyagala, Spolia Zeylanii-a, 25,

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 427

part 2, p. 30, fig. 5, pi. xii, fig. c: Ferguson's Gulf, Lake Eudolf,
Kenya Colony.
1955. Amyda triunguis triunguis Hellmich, 11, 12, 15 (of reprint).

Synonymy. The young soft-shelled turtle, allegedly from the
Gabon River, described by Strauch (1890:113, pi. iii, figs. 3-4)
as Trionyx vertehralis, later proved to be a juvenile Trionyx
suhplanus Geoff roy, according to Siebenrock (1909:606).

Common names. Nile Soft-shelled Turtle (Flower :1929) ; Soft-
shelled River-Turtle (Cansdale) ; ahihi (Alua and Gang of West
Nile:Pitman MS.) ; ahu geda (Arabic on White Nile:Werner) ;
bekoom (Liberia :R. P. Currie) ; ger (Alva and Gang for any
aquatic turtle: Pitman MS) ; gondo (in Catumbela: Anchieta) ;
kiinda (Banziri: Johnston); navingo (Bagungu of Bunyoro:
Pitman MS.); neko (Sango: Johnston); terseh, tirse or tyrsc
(Arabic: fide Anderson, Forskal and Geoffroy respectively; also
cirse or thirse: fide Burton) ; um diraga (riverine Arabs:
Flower).

Illustraiions. Bell's fine colored plates (1837) of this species
(as labiatus) are reproduced in Sowerby and Lear (1872:pls. lii-
liv) ; equally good are the colored figures in Anderson (1898 :pl.
iii), reproduced in black and white as our Plate 13.

Descripiion. Proboscis projecting, the distance from its tip t(»
the orbit longer than orbital diameter in both young and adults ;
nostril with a papilla-like process projecting laterally from the
median wall; forelimb with 3 sharp-edged, crescentic skin-folds
on its anterior surface and two weal-like sldn-thickenings (ab-
sent in young) on outer aspect; hind foot with another sharp-
edged crescentic fold under the heel posterior to the base of the
fifth toe; tail very short, pointed, not projecting beyond the
posterior leathery margin of the carapace.

Carapace of young with an indistinct keel and covered with
tubercles arranged in more or less wavy longitudinal lines ; cara-
pace 1 of adults smoother in life but showing, when dried, the
underlying bon^' sculpture ; leathery margin relatively extensive,
posteriorly extending well beyond tail.

Plastron of young smooth, without callosities, which are late in
developing (not visible in a specimen [M.C.Z. 53573] with a
carapacial length of 217 mm.) ; in adults absent on preplastra

1 A kyphotic shell Ircni the mouth of thp Catiiiiibela Uiver is meutionpd by
Mertens" (19'i6a), more fully with illustrations {1940c).

428

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Fig. 52. Skull of Trionyx triunguis (B.M. 1947.3.6.12). x %. Bo = basi-
occipital; bs =z basisphenoid ; eo = exoccipital; / = frontal; j = jugal;
m = maxillary; op = opisthotic; pa = parietal; pal =i palatine; pm
= premaxillary; po = postorbital; prf = prefrontal; pro =: prootic; pt
= pterygoid; q = quadrate; so = supraoccipital ; sq = squamosal.

(E. R. Turlington del.)

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 429

and the fused epiplastra, present on both hyo- and hypoplastra
and on xiphiplastra ; anterior edges of the hypoplastral callosi-
ties relatively straig-ht; hypoplastral and xiphiplastral callosi-
ties not in contact ; xiphiplasti'al callosities not in contact
medially; no caudal or femoral flaps but a pronounced crural-
caudal skinfold.

Osteological description. Skull. Distance between tip of pre-
maxilla and orbit much greater than the long diameter of orbit;
lieight of orbit greater than the interorbital wddth, which is sub-
equal to the width of naris; premaxilla single, not separating
maxillae luiderneath the naris, anteriorly bordering a large,
ovoid, intermaxillary foramen that is laterally and posteriorly
bordered by the maxillae, which are in contact medially for a
distance greater than tlie length of the foramen ; vomer stout, in
contact with descending processes of the prefrontals, its inter-
choanal portion relatively short; choanae narrowed, narrower
than the least distance between the maxillary triturating surfaces
and not bordered by them ; postorbital arch not half as wide
as the long diameter of orbit; jugal not or but rarely meeting
])arietal, and then but narrowly; orbit entered by jugal which
may, or may not, meet squamosal; squamosal crests thin, sharp,
and curving medially ; pterygoids sometimes approaching ^ but
never meeting opisthotic, not restricting postotic fenestra ; man-
dible without symphysial ridge, its symphysial width equalling
or exceeding the long diameter of the orbit.

Carapace. Carapacial bones coarsely pitted and vermiculated ;
no prenuchal bone ; nuchal bone not notched laterally, overlying
the second rib ; between the first pleurals a single neural ; neurals
8; pleurals 8 pairs, the eighth (and sometimes the seventh)
wholly or partially in contact medially.

Plastron. Preplastra widely separated, the anterior branch
short, about half as long as the posterior ; epiplastra fused, form-
ing an acute or right angle; hyo- and hypoplastra distinct, the
anteromedial process of the latter with 3 to 5 prongs, medial
process broad with several prongs, both medial and posteromedial
processes usually united to form an extensive dentate margin ;

1 The pterygoid sends up a triangular, tongue-like process which almost meets
the opisthotic in a Lobito Bay turtle (A.M.N.H. 50724) and a second specimen
(A.M.N.H. 36399) from "Africa." In another Lobito Bay turtle (A.M.N.H.
50723), as well as in B.M. and M.C.Z. material, this process does not exist.

430

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

xiphiplastra very long, triangular, medioanterially united by 2
or 3 prongs inserted into 3 or 4 notches, a long oval fontanelle
intervenes in front of the posterior portions which are also in
contact.

Callosities 4 (hyo-hypoplastral and xiphiplastral pairs) or ,5
(an additional azygous element on the fused epiplastra), sculp-

Fig. 53. Skull of Trionyx triunguis (B.M. 1947.3.6.12) x %. Letter sym-
bols as in Figure 52 except i' ^ vomer. A, Lateral view; B, anterior view;
C, posterior view.

(E. E. Turlington del.^

tured like the carapace, well developed but never completely cov-
ering the bones.

Color. Carapace olive, dark olive, olive brown or dark brown,
uniform or profusely speckled with dark-margined, white (yel-
low) spots, often well-marked in young, obscure or obsolete in
adults ; fleshy disk very narrowly edged with yellowish.

Plastral area Avhite or creamy white, immaculate or with
coarse, dusl^' vermiculations anteriorly ; fleshy underside of disk
dark, laterally with large, dark-bordered, light spots tending to
become much smaller posteriorly, also edged with yellowish,
either narroAvly or presenting a scalloped appearance.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 431

Head and limbs above dark, profusely spotted with lighter;
throat of young dark with larger spots and vermieulations, of
adults immaculate white or yellow ; limbs below lighter, uniform,
spotted, or vaguely vermiculate.

Size. Over-all measurements are discarded on account of the
extensile neck. It is not clear whether Anderson (1898), when
stating that this turtle "attains to a metre in length," was re-
ferring to over-all or carapace length. Carapace length of larg-
est ^ 950 mm., from Gaboon River (A.H.A. Dumeril :1860 :169) ;
another witli a length of 800 mm. was taken in Lake Albert
(Witte :1928b) ; a third was 780 mm., breadth 570 mm., from
Lake Fisherman (Biittikofer :1890) ; a Nile specimen of 680 mm.,

Fig. 54. Skull of Trionyx triunguis (A.M.N.H. 36599). Occipital view
to show the occasional presence of pterygoid ascending processes.

(S. McDowell del.)

breadth 540 mm., was caught near Duem (Siebenrock :1906a).

A $ from Khartoum measured 630 mm., a 9 from Giza 700
mm. (Flower :1933).

Weight. Largest of sixteen turtles taken in Lake Albert by
Worthington (1929) weighed 80 lbs. (as 37 kilos.). The 950 mm.

1 If this is Dum^ril's meaning wlieti he says : "sa longenr totale est de 1 m, 33 :
le bouclier seul a 0 m, 95."

432 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

triunguis from the Gaboon weighed 65 lbs. (as 29.7 kilos.), yet
the 780 mm. specimen from Lake Fisherman was 75 lbs. Pos-
sibly Baker (1867:44), who estimated the weight of one that he
hooked (but which escaped before being landed) as "at least 100
lbs." is the source of Worthington 's mentioning this weight
(1933), but we do not know the basis for that of Mocquard
(1906a), who puts the weight of this species as about "100 kilo-
grams," i.e. 220 lbs.!!

Breeding. On March 13, 1928, a 9 from Butiaba held 27
eggs with shells, measuring rather more than an inch in diame-
ter ; there were possibly 100 more about half that size developing
in the ovary (Worthington :1929). During April, in Egypt,
from 50 to 60 eggs are laid in a sandbank where they are hatched
by the solar heat (Anderson :1898). On July 10, 1825, Riippell
observed a triunguis on an island in the Nile. The reptile was
gently scraping the sand on the upper part of a steep bank.
There, three inches below the surface, she deposited 29 spherical
eggs, each having a diameter of about I14 inches (as 16 lines).
The eggs were removed to the ship where, after only two hours,
a single young one broke through the shell and began immedi-
ately to swim in a basin of water (Riippell :1835 :3).

Longevity. Twenty-five and a half years; another, still alive
on 16.ix.36, had been in the London Zoo for 22 years, 1 month,
17 days (Flower :1937a).

Diet. Stomachs of the soft-shelled turtles captured by Riip-
pell (1835) held nothing but vegetable matter, viz. fragments of
dates, gherkins and pumpkins. The reptiles came ashore at night
in search of such food, according to local fishermen, who averred
they would not take meat bait. This was confirmed by Riippell 's
experience, for none took his meat bait but two were hooked
when bread-dough was used. The stomachs of two turtles taken
in Liberia contained many palm nuts; these Biittikofer (1890)
thought they must procure on shore. Anderson (1898:33) thinks
it highly improbable that "so thoroughly an aquatic animal as
Trionyx should leave the water in search of food," and with
Flower (1933), apparently regards Arab statements as to triun-
guis digging up and devouring crocodile eggs or hatchlings, as
unproven. The allegation that they eat hatchling crocodiles dates
back at least to Goldfuss (1820).

LOVERIDGE AXD WILLIAMS: AFRICAN CRYPTODIRA 433

Anderson (1898) states that shellfish and fish appear to form
tlieir staple iood, and these are eaten under water. Worthington
(1929) agrees that moUusks and fish, both living and rotting,
constitute their principal food, but from one he recovered scraps
of goatskin and feet, from another ten large dragonfly nymphs.

P^lower (1933), when feeding captive triunguis on live bolti
fish (Tilapia), observed that if a turtle seized a fish in its jaws
that was too large to be swallowed whole, it impaled the fish on
its foreclaws, tore off the head, and employed its forefeet in
finishing the meal. One captive ate ham (Hesse :1889) ; more
unfortunate was the type of lal)iatus which refused all food and
died after three weeks (Bell:1837).

Enemies. Being regarded as a delicacy, the Nile Soft-shelled
Turtle has been hunted for hundreds, possibly thousands of
years, yet a few manage to survive in Egypt. In the Sudan cer-
tain natives specialize in the capture of these reptiles when they
come ashore to lay their eggs (Flower :1933). Keferences in the
literature reveal that the species is preyed upon by Africans in
many parts of its range. In the Gold Coast they are sometimes
taken in cane fish traps, or the larger turtles are hooked by bot-
tom fishers (Cansdale :1948c).

Durrell (1953) reports that the flesh, when stewed, was "most
palatable," tasting "like a rich and slightly oily veal." Baker
(1867:374), who removed upwards of a hundred eggs from one
of the turtles he hooked, regarded the flavor of resulting ome-
lettes as ' ' rather strong. ' '

Near Kasenyi, Lake Albert, Africans do not eat triunguis —
possibly for superstitious reasons — and allegedly release any
turtles tliat become entangled in their nets, as P. Janseens in-
formed Dr. J. P. Chapin (fide label of vii.1937 on A.M.N.H.
60263).

By stretching the untanned skin of a fish over a turtle cara-
pace, Africans convert it into a guitar (Baker :1867 :203). An-
derson (1898 :33 :footnote) quotes Prospero Alpini as saying, in
1735, that the larger carapaces were utilized for making shields,
and adds that the practice still persisted in Nubia.

Defense. Sir Samuel Baker (1867:44) relates how, using a
strip of glistening perch on a stout hook fitted to treble-twisted
wire, he caught an enormous turtle "at least 100 lbs. weight."

434 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

However, when his Arab assistant drew the reptile, strugglingr
and snapping, close to the steep bank, its jaws cut through the
triple brass wire as with shears.

Durrell (1953) states that the jaws of a decapitated turtle
snapped on, and splintered, a piece of wood. Hesse (1889) writes
of one snapping at his fingers. According to Bell (1837) the
movable and extended coriaceous margin of the carapace en-
abled the t3'pe of lahiatus to conceal itself in ver}^ shallow mud
in much the same manner as would a flounder or plaice.

Temperament. Vicious and always ready to bite (Cansdale:
1948c).

Habits. Soft-shelled Turtles run extremely fast when taken
out of the water, according to Baker (1867:374) who landed
three. A 9 in the Giza Zoological Gardens frequently travelled
from one i:)ond to another. To do so she had to climb a metre-
liigh, vertical fence of wire netting and, after gaining the top,
flop down on the farther side (Flower :1933) . A very small turtle
surprised Worthington (1933:38) by climbing from a bucket.
He describes them as wallowing in the crater lakes of an island
in Lake Rudolf, where he landed a fifty-pounder accidentally
hooked by one of its feet. When occasion demands, however, these
turtles can swim fast in their chosen element ( Hesse :1889).

Habitat. Ponds, lakes and rivers, but Auguste Dumeril (1860 :
168 :footnote) records a triungids with a carapace length of 950
mm., and weighing 29.7 kilograms, as having been taken in the
ocean about 3 or 4 km. from the mouth of the Gaboon River. The
collector did not believe such an able swimmer as Trionyx could
have been carried so far by the river in flood.

However, Flower (1933 :754) believes that when the Nile is in
spate these turtles are carried into the Mediterranean along with
the carcasses of cattle and clonkej's, and so up the Palestine coast
by the powerful inshore current.

Localities. Egypt ^ Embaba district, Giza Province; Ezbet
Semeda Saleh, Fayum Province; Kom Ombo, Aswan Province;
Nile near Cairo; Philae ; Wadi Haifa. Sudan: *Atbara village
(U.S.N.M.); Berber Province; Blue Nile; Delladilla, deep in
Base; Dongola; Duem; Haifa Province; Khartoum; Kulgeila,

1 For discussion of questionable records, see Flower (1933).

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 435

Dongola Province; lioseires (Rosairos in B.M.), Blue Nile; Sen-
nar; Shandi ; *8ul River; Upper Nile (1873). Eritrea: Asmara;
Setit River (Fiume). Ethiopia: (as Abyssinie :1851) Lake
Tana (Zana). Somalia: Ganana; Lugh. Uganda: Butiaba,
l.ake Albert. Kenya Colony: Ferguson's Gulf, Lake Rudolf;
K'aliokwell River .Mouth, Lake Rudolf. Angoln: Benguela;
Cuanza; *Cunga ; Catunibela; Loanda; *Lobito Bay (A.M.N. II.).
Cabinda: Chinehoxo. Belgian Congo: *Congo River at Lukolela
(A.M.N.H.); *Lake Albert at Kasenyi (A.M.N.H.) ; Mahagi
Port; Mayombe (Mayumbe). French Equatorial Africa: Bil-
cliem: Fernan Vaz (Fernand Vas) ; *Gabon River; Kuilu River
below Bumina; Lambarene; Loaugo Coast; Ogoue (Ogowe) ;
Ovenga River; Rembo River. French Cameroon: Uibongo, near
Edea ; Lake Ossa ; *Sakbayeme. British Cameroon: Bonge, Meme
River; Lake Soden: Ossidinge. Togo: Kete Krachi (Kratje).
Gold Coast: Accra; Birrim, tributary of Pra River. Liberia:
Fisherman Lake; Grand Bassam (G. Massam) ; *Saint John
River near Gl)anga; Saint Paul River near Muhlenberg Mission,
Mt. Coffee. Sierra Leone: *AVanje River near Gbap ; *near Kale;
*near Subn. Portuguese Guinea: Bissao (Bissan). Gambia. Sen-
egal: Dagana (1870). French West Africa: Bilchem on Niger-
chad frontier.

Range. Egypt and Eritrea south to Lake Rudolf, southwest to
Lake Albert and Angola, northwest to Senegal. Accidental along
Palestine coast of Asia Minor (cf. Flower :1933).

Erroneously included in the fauna of Lake Victoria by Lover-
idge (1924b) on the basis of alleged remains in the Miocene de-
posits of Karungu Bay, Lake Victoria, for, as pointed out by
Worthington (1929) no living triungnls has been taken above
the Murchison Falls, White Nile.

Werner (1924a :269), unaware of any record for triunguis
above Duem on the White Nile, suggests that it occurs only in
the lower and middle reaches of that river, besides the entire
length of the Blue Nile. Werner makes no mention of its pres-
ence in Lakes Albert and Rudolf, and thinks that in the Upper
Nile. Sobat River, and Bahr el Ghazal, Trionyx is replaced by
Cyclanorbis.

436 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Genus CyCLANOEBTS Gray

1854. Cyclanorbis Gray, Proc. Zool. Soc. London, 1852, p. 135. Type:

petersii Gray (by monotyi)y) = scnegalensis (Dumeril and Bib-

ron).

1856. Cyclanosteus Gray, Proc. Zool. Soc. Loudon, 1855, p. 201. Type:

petersii Gray (designation by GUnther 1865, in Zool, Kec. for 1864).

1865e. Tetrathyra Gray, Proc. Zool. Soc. London, p. 323. Type: petersii

Gray (by monotypy).
1870e. Baikiea Gray (not Bailcia Gray: 1865), Suppl. Cat. Shield Kept.,
Part 1, p. 114. Type: elegans Gray (by monotypy).

Definition. Cutaneous femoral flaps present, permitting con-
cealment of the hind limbs.

Skull without maxillary ridging ; intermaxillary foramen small
or absent; prefrontals with or without connection with the
vomer ; jugal narrowly in contact with parietal ; postorbital arch
narrower than diameter of orbit ; pterygoids joining opisthotic,
greatly restricting fenestra postotica.

Carapace with prenuehal bone present or absent, without peri-
pheral ossifications; nuchal bone not notched laterally, ventrally
with raised posterolateral processes which tend to underlie the
first pleurals, between which are two neurals; neural number
variable ; pleurals 8 pairs, one to all meeting medially.

Plastron with preplastra long and angular; hyo- and hypo-
plastra fused, their posteromedial process with 3 prongs between
which are inserted the 2 anterior prongs of the adjacent xiphi-
plastron; callosities variable in number, present or absent on
gulars.

Range. Southern Sudan west to Senegal.

Fossil record. No fossils certainly identifiable ; however, see
under Cycloderma. The fragment reported by Dacque (1912:
Geol. Pal. Abhandl. :14:322) is at present indeterminable, as
Siebenroek suggested when consulted by Dacque.

Remarks. Both in skull and shell, as a glance at the table on
p. 418 will show, the two species of Cyclanorhis are strikingly
different. These differences, however, reflect the relative primi-
tiveness of elegans and relative specialization of senegalensis.
but the two are still closer to one another than to any
other African trionychids. Nevertheless, should it ever seem
desirable to separate them generically, the name Baikiea Gray is

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 437

available for elegmis while the type of Cyclanorhis is sencgalen-
sis.

C. seneyalensis offers no problems at the species level, though
ontogenetic changes in the callosities induced Gray to propose
names which he very soon synonymized. On the other hand, our
identification of oligotylus with elegmis clarifies a muddle of
names that began in 1864 when Gray described and figured a
skull from Gambia as being possibly that of an adult senegalen-
sis. Later Gray associated this skull and similar ones with the
mottled juveniles which he described as BaiJdea elegans. This ac-
tion was reversed by Boulenger ( 1889a :272) who returned the
skulls to sencgalensis, restricting elegans to the two young cotypes
described by Gray. In consequence, Siebenroek (1902b :810),
accepting Boulenger 's verdict without seeing the British Museum
skulls, described a new species oligotylus, based on an adult
shell.

The persistence of this confusion till the present, is largely
attributable to the absence of an adequate ontogenetic series.
While there is still no such series of this species, more material
is now available and the situation in the freshwaters of north
central Africa appears to be as follows :

In this region, so far as is known, there are 07ily two kinds of
juvenile trionychids with femoral flaps. The two are readily
distinguished by color and pattern. One of these is clearly ele-
gans, the other is definitely senegalensis.

So far as known, there are only two kinds of adult cyclanor-
bide shells from this region. They differ in maximum size, num-
ber of neurals ^ (and correspondingly in the number of pleurals
in contact), the extreme development of callosities in one versus
their great reduction in the other. One corresponds to Sieben-
roek's description of oligotylus, the other is definitely senegalen-
sis.

Similarly there are only two kinds of adult cyclanorbide skulls
known from this region. The larger are clearly referable to
oligotylus but are also identical with the large skulls which

1 We consider that size in conjunction with the neural number is a good guide
for the reparation of the two species. Thus Werner's Mongalla collections re-
ported on bv Siebenroclj (1906a) and Werner (1924a) separate readily into two
series — aU'the large shells having 6 or more neurals, all the smaller ones 5 or
less. Neural number does not increase during ontogeny.

■i'SS BULLETIN : MUSEUM OP COMPAKATIVE ZOOLOGY

formed part of Gray's concept of elegans; the smaller are defi-
nitely senegalensis.

At each age level there is therefore good evidence for the ex-
istence of only two cyclanorbide turtles in north central Africa.
This has become increasingly clear over the years and the newer
collections have been identified on this basis. Thus for some time
the British Museum has been labeling its strongly mottled cycla-
norbide juveniles oligotyhis, correctly assuming that they were
juveniles of the large adults they had also received. It is only
because no comparison with the juvenile cotypes of elegans was
made that the identity of oligotylus and elegans has remained so
long undiscovered.

Cyclanorbis elegans (Gray)

1864c. Cyclanosteus senegalensis (adult ?), Gray (part: omit figs. 16-18

only), 97, figs. 19-21.
l<S65d. Tetrathyra baikii Gray (part: omit shell), 205.
1865e. Gray (part: omit shell), 324 (reprint of 186od).

1869c. Bailcica elegans Gray, Proc. Zool. Soc. London, p. 222, pi. xv, fitr. 2:

Africa (West ?).
1870e. Gray (part: omit C. sonegalensis var. callosa), 114. fig. 39.

1873. Gray, 69.

1873b. Gray, 86.

1 889a. Cyclanorhis elegans Boulenger, 272.
1909a. Siebenrock, 594.

1919. Schmidt, 601.

1934u. Mertens and Miiller iii Bust, 12.

1889a. Cyclanarbis senegalensis Boulenger (part: not of D. and B.), 272

(cotypes r, s, t of Bailiea elegans).
1897. Siebenroek, 248, pi. ii, fig. 5 ; pi. iv, fig. 20.

]901i;. Tornier, 68 (material lost, but agrees with elegans with possible

exception of Jeudi specimen).
1902b. Cyclanorbis oligotylus Siebenroek, Sitzb. Akad. Wiss. Wien, 111,

pp. 810, 842, fig. 17: Nubia (Upper Nile?).
1905a. Siebenroek, 465, fig. 2.

1906a. Siebenroek, 838, fig. 8.

1908. Werner, 169.

1909a. Siebenroek, 594.

1912b. Werner, 493.

1919. Schmidt, 601.

1924a. Werner, 269.

1934a. Mertens and Miiller in Bust, 12.

LOVERIDQE AND WILLIAMS: AFRICAN CBYPTODIRA 439

Synonymy. Further citations of "baikii (part) " will be found
under Cyclanorhis senegalensis.

Common 'name. Nubian Soft-shelled Turtle.

Description of hatchlinij .^ Proboscis projecting, the distance
from its tip to the orbit (not eye) greater than the orbital diame-
ter ; upper lips meeting in front to form a flat arch ; nostril with
a papilla-like process projecting from the median wall; forefoot
with 4, sharp-edged crescentie skinfolds on its upper surface,
another indistinct weal-like thickening on outer aspect ; hind foot
with a sharp-edged crescentie fold under the heel posterior to the
base of the fifth toe; tail rudimentary, not projecting beyond the
posterior rim of the carapace.

Carapace of hatchling with a moderate vertebral keel and
numerous, raised, rather wavj^ longitudinal lines comprised of
small individual tubercles anteriorly converging towards a
strongly tubercular nuchal area; leathery margin posteriorly
smooth, not extensive, scarcely extending beyond the tail.

Plastron of hatchling smooth, without callosities, femoral (but
no caudal) flaps permit concealment of hind limbs.

Osteological description. Skull. Distance between tip of pre-
maxilla and orbit greater than the long diameter of orbit ; height
of orbit about one and one-third to one and one-half times the
interorbital width, which is less than the width of naris; pre-
maxilla single, not separating maxillae underneath the naris ;
behind the premaxilla the maxillary triturating surfaces meet,
their inward extensions closing or partly closing over the inter-
maxillary foramen ; behind the triturating surfaces more dorsal
flanges of the maxillae meet mesially ventral to the vomer for a
distance greater or les.s than the palatal length of the premax-
illa; vomer slender, in contact with the prefrontals, its inter-
choanal portion moderately long; ehoanae only slightly narrowed
by inward extensions of the triturating surfaces of the maxillae ;
postorbital arch somewhat more than half the long diameter of
the orbit; orbit entered by jugal, which is narrowly in contact
with the parietal; squamosal crests long, sharp, straight or in-
curved; pterygoids meeting opisthotic, greatly restricting post-
otic fenestra; mandible without symphysial ridge, its ventral
symphysial width less than the diameter of the orbit.

1 Based oil a Khartoum specimen (B.>L 1909.3.25.3).

440

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Fig. 55. Skull of Cyclanorbis elegans (Cotype. B.M. 65.5.9.22 = 1947.
3.6.28), X %. Dorsal and ventral views.

(E. E. Turlington del.)

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA

441

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442 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Carapace. Carapacial bones distinctly granulated ; no prenu-
chal bone ; nuchal bone not notched laterally bnt with multiple
digitations underlying to a small extent the first pleurals, be-
tween which are two neurals; neurals 8 or 9, forming a continu-
ous series, or the last two separated ; pleurals 8 pairs, the eighth
in contact medially.

Plastron. Preplastra widely separated, the anterior branch
long, longer than the posterior; epiplastra fused, forming an
acute angle ; hyo- and hypoplastra fused, their anterior border
straight or somewhat concave, 7iot convex, posterior border of
this compound bone very deeply excavated, anteromedial process
with indistinct prongs; medial process short, onlj' slightly pro-
jecting, posteromedial process with only 8 prongs between whicli
are inserted the 2 anterior prongs of the xiphiplastron ; xiphiplas-
tra rodlike, anteriorly broader, nowhere in contact though con-
verging posteriorly, neither notched nor broad but pointed
posteriorly.

Callosities 2 or 4, sculptured like the carapace ; usually 2 on
the hyo-hyoplastra onh', covering most of these bones but widely
separated medially; occasionally small callosities on the xiphi-
plastra.

Color. Carapace of hatchling ^ dark olive brown with large
irregular blotches of yellow, the periphery sometimes with a few,
scattered, somewhat smaller, roundish, yellow spots. Carapace
of adults light brown to olive green, the i)eriphery spotted and
vermiculated with light green.

Plastron of hatchling yellowish, its central area dark, pro-
fusely spotted with lighter, not, or only sometimes, extending to
the periphery ; fleshy underside of disk edged or spotted with
yellow. Plastron of adults dirty yellow, obscurely blotched with
darker.

Head and neck of hatchling gray brown above, anteriorly spot-
ted with lighter, posteriorly darker and unspotted; chin and
throat anteriorly' vermiculated ; neck below, yellow, immaculate ;
limbs uniform or mottled with yellow. Head of adults above dark
brow^n with light green vermiculations ; temporal region olive
green ; neck lighter, speckled and spotted with yellow ; throat and
neck below, uniformly yellow.

^ Checked l).v a Khartoum spocimon (B.M. 1009.3.25.3) with carapace length
of 67 mm., and breadth of 50 mm.

LOVERIDGE AM) WILLIAMS: AFRICAN CRYPTODIRA 443

Size. Carapace length, including fleshy margin, of oligotylus
type, G05 mm., its breadtli 465 mm., its height 150 mm.; disk of
same specimen 440 mm., breadtli 400 mm.; exceeded by a dorsal
di.sk from Mongalla with a length of 490 mm. (Siebenrock :1906) .

Enemies. At Mongalla, on the Bahr el Jebel, thirteen Cycla-
norbis carapaces, representing both species, were purchased by
Werner, but he could not get the Bari tribesmen to part with an
entire turtle. Even the plastron and fleshy margins were sought
after for culinary purposes ( Siebenrock :1906).

Localities. Sudan: Bahr el Jebel at Mongalla; Nubia; Sobat
River at Khor Attar; Upper Sobat at Nasser; White Nile at
Khartoum and at *Tonga. French Equatorial Africa: Schari
River near Fort Archambault (V.M.). Nigeria: Niger River at
Lohata, Kabba Province. Togo: Kete Krachi (Kratje) ; Mango
(Mangu) ; Yendi (Jendi).

Range. Sudan ; Nigeria ; Togo.

Cyclanorbis senegalensis (Dumeril and Bibron)

183."). CryptoptLS Scnegaleruiis Dumeril and Bibron, Erpet, Gen., 2, p. 504:

Senegal.
1851. Dumeril and Dumeril, 23.

1844. Emyda senegalensis Gray, 47.
1855. Gray, 64.

1860e. Gray, 316.

1854. Cyclanorbis Petersii Gray, Proc. Zool. Soc. London, 1852, p. 135:

Gambia Eiver, Gambia.

1855. Gray, 69.

1855. Cyclanostcus Petersii Gray, 64, pi. xxix.

1860e. Gray, 315.

1860. Cycloderma senegalense Dumeril, 168.

1865. Strauch, 131.

1862a. Cycloderma Petersii Strauch, 56.

1864e. Cyclanostcus senegalensis Gray (part: omit figs. 19-21), 95, figs.

16-18.
1870d. Gray, 717, pi. xliii.

1872c. Gray, 27.

1873. Gray, 70.

1873b. Gray, 87.

1884a. Eochebrune (ignored).

1865d. Tctrathyra haiMi Gray (part: omit young and skulls), Ann. Mag.

Xat. Hist. (3), 16. p. 205, fig. — : ? Niger River, West Africa.

444 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1865e. Gray, 324, fig. — (reprint of 1865cl).

1873. Gray, 70.

1873b. Gray, 86.

1884a. Rochebrune, 35 (ignored).

1865f. Cyclanosteus senegalensis var. equilifera Gray, Proc. Zool. Soe. Lon-
don, p. 423, fig. 2 : Niger River, West Africa.

1870e. Gray, 113, fig. a.

1865f. Cyclanosteus senegalensis var. normalis Gray, Proc. Zool. Soc. Lon-
don, p. 423 : Niger River, West Africa.

1865f. Cyclanosteus senegalensis var. callosa Gray, Proc. Zool. Soc. London,
p. 423, fig. 1 : Niger River, West Africa.

1870e. Gray, 113, fig. 38.

1884a. Tetrathyra vaillantii Rochebrune, Faune Senegambie, Reptiles, p.
36, pi. iv, figs. 1-2: Senegal (localities unreliable).

1889a. Cyclanorbis senegalenisis Boulenger (part: omit Baikiea elegans
cotypes), 271.

1898. Siebenrock, 425.

1899. Siebenrock, 566.

1900. Flower, 967.

1902b. Siebenrock, 839, fig. 16.

1905a. Siebenrock, 467.

1906a. Siebenrock, 835, fig. 7.

1908. Werner (1907), 1826, 1924.

1909a. Siebenrock, 594.

1910. Miiller, L., 624.

1910. Sternfeld, 8.

1912b. Werner, 493.

1913. Siebenrock, 43, fig. 22.

1917. Sternfeld, 416.

1919. Schmidt, 601.

1924a. Werner, 269.

1925. Flower, 935.

1929. Flower, 54.

1934a. Mertens and Miiller in Rust, 12.

1937. Andersson, 3.

1937a. Flower, 16.

Further citations of "senegalensis" and "haikii (part) " will
be found under CycJanorbis elegans.

Common names. Senegal Soft-shelled Turtle (Flower :1929) ;
African Bungoma (Gray:1844).

IllustraUo'ns. Dorsal, ventral and lateral views of the shell are
furnished by Gray (1855 rpl.xxix), also the skull and jaws

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 445

Fig. 57. Skull of Cyclanorhis senegalensis (B.M. 65.5.9.20), x Yz. Dorsal

and ventral views.

(E. E. Turlington del.)

446 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

(Gray :1864c :ti^. 16-18), while Siebenrock supplies dorsal
(1913b:fig.22) and ventral (1902b :fig. 16) views of the plastron.

Description of hatchlings and young.^ Proboscis projecting,
the distance from its tip to the orbit (not eye) shorter than, sub-
equal to, or greater than the orbital diameter ; nostril with
papilla-like process projecting upwards from its infra-median
corner ; upper lips meeting in front to form a flat arch ; forefoot
with 5 or 6, sharp-edged, crescentic skinfolds on its upper surface
(see Fig. 51C), another indistinct weal-like thickening on outer
aspect; hind foot with a sharp-edged crescentic fold under the
heel posterior to the base of the fifth toe ; tail rudimentary, not
projecting beyond the posterior rim of the carapace.

Carapace of young with a moderate vertebral keel and numer-
(jus, raised, rather wavy, longitudinal lines comprised of small
individual tubercles anteriorly converging toward a strongly
tubercular nuchal area, leathery margin posteriorly smooth, not
extensive, scarcely extending beyond the tail.

Plastron of young, smooth, without callosities; femoral and
caudal flaps permit concealment of hind limbs and (probably)
tail.

Osteological description. Skull. Distance between tip of pre-
maxilla and orbit greater than the long diameter of orbit ; height
of orbit about twice the interorbital width, which is less than the
width of naris ; premaxilla single, not separating maxillae under-
neath the naris ; inward extensions of the maxillary triturating
surfaces anteriorly narrowing the small intermaxillary foramen,
almost excluding the premaxillae from its medio-anterior margin,
elsewhere bordering the foramen (except sometimes mediopos-
teriorly where the maxillae may be separated by the vomer, which
then also separates them behind the foramen) ; length of the
foramen subequal to its distance from the anterior choanal mar-
gins ; vomer slender, without ascending processes to prefrontals ;
its interchoanal portion moderately long; choanae bordered by
the triturating surfaces, postorbital arch «about equal to half the
long diameter of the orbit ; orbit broadly entered by jugal, which
is in contact with the parietal ; squamosal crests short and blunt ;
pterygoids meeting opisthotic, greatl^^ restricting postotic fenes-

1 Based on two Maccarthy Id. hatchlings (B.M. 1901.7.16.26-27), a hatchling
and two young from Togo (iu the Berlin Museum), and, seven specimens trom
the Schari River near Port Archambault, French West Africa.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA

44'

tra; mandible without syinphysial ridge, its symphysial width
less than the long diameter of the orbit.

448 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Carapace. Carapacial bones rather finely granulated and ver-
mieulated ; a prenuehal bone in adults ; nuchal bone not notched
laterally, extending slightly below the first pleurals, between
which are two neurals ; neurals variable in number, not forming
a continuous series ; pleurals 8 pairs, a variable number in con-
tact medially.

Plastron. Preplastra widely separated, the anterior branch
longer than the posterior ; epiplastra fused, forming an acute
angle, their pointed posterior prong inserted in a deep notch
which separates the rounded inner margin from the strongly
convex anterior border of the fused hyo- and hypoplastra;
posterior border of this compound bone very deeply excavated,
the anteromedial process with several almost juxtaposed prongs,
the medial process very short, scarcely projecting beyond the
rim, the posteromedial process with only 3 prongs between which
are inserted the 2 anterior prongs of the xiphiplastron ; xiphi-
plastra oblong, slightly wider at their anterior and posterior ends,
with 1 or 2 notches posteriorly, converging posteriorly but no-
where in contact.

Callosities 7 to 9, sculptured like the carapace, extremely de-
veloped in adults except on the xiphiplastra where they may be
absent, even in adults; regularly present anteriorly, but not
overlying any bones, are two small, subquadrilateral callosities in
contact medially; posteriorly these are in contact with the pre-
plastral callosities, Avhich are large oblique ovals in contact
medially ; fitting between the latter posteriorly is the forwardly
directed apex of the fused epiplastra bearing a large triangular
callosity ; in this general anterior region additional callosities are
sometimes present {fide Gray:1865d); hyo-hypoplastral callosi-
ties, covering almost the entire surfaces of these bones, nearly
meet medially ; xiphiplastral callosities poorly developed or ab-
sent.

Pointing out that the callosities increase in number and size
with age, but that their development appears to proceed more
slowly in the Nile than in West African rivers, Siebenrock (1906)
states that the hypoplastral callosity changes in shape as it devel-
ops. This is borne out by the B.M. and Vienna material exam-
ined by E.E.W.

Thus while fully adult senegalensis have the hyo-hypoplastral

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 449

callosity strong^ly convex anteriorly, as described above, in
younger specimens the anterior edge of this element is straight —
as in adult elegans. From the Schari River series in Vienna
Museum it would appear that the anterior convexity occurs in
specimens with a carapace length between 210 and 220 mm. But
while the anterior convexity of the hyo-hypoplastral callosity is
not developed in senegalensis with carapace lengths of 200 mm.
or under, there is a greater development of all the callosities in
all senegalensis (except hatchlings and near hatchlings) than
there is in elegans. Thus, while callosities are already well indi-
cated on a Schari River senegalensis with a carapace length of
128 mm., there is still no trace of them on a Schari River elegans
of 170 mm.

In Schari River senegalensis the first traces of preplastral cal-
losities appear in specimens with a carapace length of 128 mm.,
and these have become quite large in those with a carapace
length approaching or exceeding 200 mm. In elegans, so far as
is known, preplastral callosities are never developed. This fact,
together with the more retarded condition of the hyo-hypoplas-
tral callosity in elegans as compared with senegalensis of the
same size, and differences in coloration, should always permit
separation of young elegans from halfgrown senegalensis.

Color. Carapace of hatchling grayish or brownish, sparsely
punctate with black and indistinctly mottled with lighter, most
clearly around the posterior periphery which may be edged with
lighter. Plastron yellowish or cream colored, centrally quite
heavily blotched with darker. Head, neck and limbs gray or
brownish, finely spotted with white or indistinctly vermiculate
with lighter; throat, and neck below, whitish (based on five
specimens seen by us, viz. three hatchlings from Maccarthy Id.
and Togo, also two young from Togo).

Carapace of adult blackish olive, the periphery narrowly
edged with white ; plastron pure white or yellowish, clouded
with brown. Head above, olive or blackish, sparsely punctate
with lighter, sides of head indistinctly vermiculated with dusky
(Gray:1870d:pl. xliii, not text).

Carapace of adult light brown, its fleshy margin dark brown,
immaculate. Plastron yellow. Head and neck above, olive
green; below, yellow {fide Siebenrock :1905a).

450 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Size. Length of dorsal disk of a hatchling (Berlin Mus. 26687)
from Kundja Konkomba, Togo, ca. 46 mm. ; of a gravid 9 from
Duem, 225 mm. (Siebenroek :1906a) ; length of largest disk, 350
mm. (Boulenger :1889a).

Breeding. On April 12th, at Duem, 6 hard-shelled, almost
spherical eggs measuring 36 mm. in diameter, were removed from
a 9 whose cloaca was so flaccid that Werner thought she must
iiave been captured while laying (Siebenroek :1906a).

Longevity. Ten years, 2 days, and still living in London Zoo
at the time of the report (Flower :1929).

Enemies. These turtles are hunted by the Bari tribesmen of
the Bahr el Jebel who utilize the carapacial disks as basins, and
esteem the fleshy margins for their food value (Werner; cited
by Siebenroek :1906a).

Defense. When these turtles are molested they withdraw their
heads and forelimbs, enclosing them by raising the anterior
plastral lobe to meet the carapacial margin ; the hind limbs are
similarly withdrawn beneath the protecting femoral flaps (Stern-
feld:1917).

Habits. During the (iambian rainy season young senegalensis
move into the marshes where four were captured. Africans
place these turtles in their wells, possibly to keep them free of
putrifying matter as Anderson (1937) suggests.

Localities. Sudan: Bahr el Zeraf ; White Nile at Duem, and
north to Renk, and at Tonga. French Equatorial Africa: *Gabon
(skull in M.C.Z.) ; *Schari River near Fort Archambault (V.M.).
French Cameroon: Lake Tchad (also as Chad and Tsade). Togo:
Kundja Konkomba; Mango (Mangu). Nigeria: Lagos; Niger
River at Lohata, Kabba Province. Gambia: Gambia River; Mac-
earthy Island. Senegal: (Type locality).

Range. Sudan west through French Cameroon and Gabon
to Senegal.

-■is'-

"Cyclanorbis" sp.

By the courtesy of Dr. Eiselt of the Naturhistorisches Museum,
Vienna, we have been able to examine the soft-shell-turtle
hatchling said by Werner (1924a :270) to have come from the
northwest shore of Lake Tanganyika. This specimen, as noted
on p. 420, will not key out even generically on the characters

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 451

used by us. In having the distance from tip of the proboscis to
the orbit greater than the orbital diameter (in so young a
specimen), it resembles Cyclanorhis rather than Cyclodcrma. In
possessing a caudal flap as well as femoral flaps it resembles
Cyclanorhis senegalensis or the species of Cycloderma. In colora-
tion, being without spots or vermiculations on head, neck, cara-
pace or plastron, it differs from all other African soft-shells
examined by us. This hatchling may be a local variant of
senegalensis, but at present no soft-shell turtle is known from
Lake Tanganyika. It may well represent a new form, but we
consider it undesirable to erect a new name on the basis of a
single hatchling of somewhat indefinite origin.

Descriptiori of hatchling (V.M. 14826) from northwest shore
of Lake Tanganyika. Proboscis projecting, the distance from
its tip to the orbit greater than orbital diameter; nostril with
papilla-like process projecting upwards from the inframedian
corner; forefoot with 4, sharp-edged, crescentic skinfolds on its
upper surface, another indistinct weal-like thickening on outer
aspect; hind foot with a sharp-edged crescentic fold under the
heel posterior to the base of the fifth toe ; tail rudimentary, not
projecting beyond the posterior rim of the carapace.

Carapace with a moderate vertebral keel and numerous raised,
wavy, longitudinal lines comprised of small tubercles anteriorly
converging toward a tubercular nuchal area; leathery margin
posteriorly smooth, not extensive, scarcely extending beyond the
tail.

Plastron smooth, without callosities; femoral and caudal flaps
permit concealment of hind limbs and (probably) of tail.

Color. Carapace uniform olive (olive green fide Werner).
Plastron uniform yellowish wdiite. Head and neck above, gray-
brown (gray-green : Werner) without spots, lines or vermicula-
tions; throat and neck below, yellowish white.

Size. Length of dorsal disk (V.M. 14826) 70 mm. ; breadth 60
mm. (as given by Werner).

Genus CycLODEEMA Peters

18.54a. Cycloderma Peters, Monatsb. Akad. Wiss. Berlin, p. 216. Type:
Cycloderma frenatum Peters.

452 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1859. Heptathyra Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 294. Type:

Cryptoptis anbryi A. Dumeril.
1856. Cryptopodus Dumeril, Eevue Mag. Zool. (2), 8. p. 374. Lapsus for

Cryptopus Dumeril and Bibron, 1835.
1860a. Aspidochelys Gray, Proc. Zool. Soc. London, p. 6. Type: A. Jivivg-
stonii Gray = frenatuvi Peters.

Defi7iition. Cutaneous femoral flaps present, permitting con-
cealment of the hind limbs.

Skull without maxillary ridg'ing ; intermaxillary foramen
small; postfrontals not meeting vomer; jugal broadly in contact
with parietal; postorbital arch wider than diameter or orbit;
pterygoids joining opisthotic, greatly restricting fenestra post-
otica.

Carapace without prenuchal bone, without peripheral ossifica-
tions; nuchal bone notched laterally, ventrally with raised pos-
terolateral processes which tend to underlie the first pleurals,
between which are two neurals, rarely one; neurals 8 or 9,
rarely 7, pleurals 8 pairs, the seventh and eighth, or eighth only,
in contact medially.

Plastron with preplastra short and straight; hyo- and hypo-
plastra fused, their posteromedial process with three prongs
between which are inserted the two anterior prongs of the ad-
jacent xiphiplastron ; callosities in adults 7, always present on
gulars.

Range. East (Tanganyika Territory south to Mozambique)
and West (Portuguese, Belgian and French Congo) Africa.

Fossil record. First reported in the Lower Miocene of Kenya ;
also known from the Pleistocene of Lake Rudolf. Although these
remains are certainly cyclanorbide, the generic identification is
uncertain; they may be referable to Cyclanorbis.

Remarks. The unity of the genus Cycloderma is clearer than
that of Cyclanorbis. Discrimination of the two species is also
very easy. There is, however, the possibility of a taxonomic
problem within the species frenatum. The original Zambezi
series were described by Peters as being dark green in general
color, as also shown on his plate. Our material^ from Lake
Nyasa and the Ruvuma River, however, consists of pale gray to
leaden hatchlings and pale to dark olive adults. Other minor
differences are noted in the color description below (p. 464).

1 Twenty-one juveniles and 11 adults (Loveridge coll.).

LOVERffiGE AND WILLIAMS: AFRICAN CRYPTODIRA 453

There may be also a parallel osteological difference. The
skulls^ of all our frcnatwm from Lake Nyasa and the Ruvuma
lack a vomer, for which, in fact, there is no evident place of
attachment. On the other hand, a vomer, though very narrow
and much reduced, is shown in Peters' figure of a Zambezi skull.
The vomer is present in a Zambezi specimen (B.M.) as well as
another skull (Senck. Mus.), unfortunately without locality,
seen by one of us (E. E. W.).

Thus the possibility of constant color and skull differences
between the nominate population of the Zambezi, and those
from further north needs to be investigated. Should these dif-
ferences hold good, the more northerly populations will merit
subspecific distinction.

No suspicion of geographical differentiation attaches to aubryi,
but in this case there is an interesting biological problem. The
pattern of au-hryi is very distinctive, indeed unique among
trionychids. On the head and nape is a chocolate brown vertebral
streak which is continued on to at least the anterior third of
the rather light brownish disk. Curiously enough, in the same
West African rain-forest region inhabited by C. auhryi, a very
similar pattern is present in two species of the pleurodiran genus
Pelusios. The juveniles of both P. gahonensis and niger have
similarly light brown carapaces with a black vetebral stripe
which, in gahonensis, is continued on to the head. While in C.
auhryi the pattern persists throughout life, in P. niger it occurs
only in the hatchling stage, and has an intermediate duration in
P. gaho7iensis. The significance of this apparent mimicry is quite
unknown.

Cycloderma AUBRYI (Dumeril)

1856a. Cryptopodus Aubryi Dumeril, Eevue Mag. Zool., (2), 8, p. 374, pi.
XX : Gabon, French Congo.

1859. Eeptathyra aubryi Cope, 294.
1864c. Gray (part), 93, figs.
1870e. Gray, 93.

1884a. Eochebrune (in error), 28, pi. ii, figs. 1-2.

1860. Cycloderma aubryi Dumeril, 166.
1875a. Peters, 196.

1876a. Peters, 117, pi. — , figs. 1-2.

1 Seven skulls (Loveridge coll.).

45-1

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1877c. Peters, 611.

1880b. Vaillant, 797.

1880e. Vaillant, 53, pi. xxx, fig. 13.

1889a. Boulenger, 267.

1895a. Bocage, 8.

1896. Giinther, 263.

1897. Sjostedt, 33.
1898a. Werner, 204.
1900b. Boulenger, 447.
1902b. Siebenroek, 836, fig. 14.
1902e. Tornier, 665.

19061. Boulenger, 197.

1906a. Mocquard, 480.

1909a. Siebenroek, 593.

1910. Sternfeld, 8, fig. 15.

1917. Sternfeld, 415.

1919. Schmidt, 598.

1929. Flower, 53.

1933m. Witte, 68.

1934a. Mertens and Miiller in Rust, 12.

1950. Williams, 552.

1953a. Laurent, E., 21, 26.

1953. Witte, 21.

1864c. Heptathyra frenuta Gray (part), 94.

1873b. Gray (part), 76.

Common name. Aubry's Soft-shelled Turtle (Flower :1929).
Illustrations. The black and white drawings on plate xx of
Dumeril's original description (1856a), copied here as our Plat€
18 (top), clearly show the principal characteristics.

Description. Proboscis projecting, the distance from its tip to
the orbit (not eye) shorter than or subequal to (in young), or
longer than (in adults), the orbital diameter; nostril without
papilla-like process projecting from its infra-median corner;
upper lips meeting in front to form a more or less pointed arch ;
forefoot with 6 or 7 sharp-edged, crescentic skin-folds on its
upper surface, another w^eal-like thickening on outer aspect;
hind foot with a sharp-edged crescentic fold under the heel pos-
terior to the base of the fifth toe ; tail rudimentary, ^ not project-
ing beyond the posterior leathery rim of the carapace.

Carapace of young with a moderate vertebral keel and numer-
ous tubercles, those in center not arranged in longitudinal lines,

1 Possibly slightly longer in ^ |J than in 9 9 .

l.OVEBLDQE AND WILLIAMS : AFRICAN CRYPTODIHA 45.1

the lateral ones showing a tendency to such arrangement ; cara-
pace of adults smooth in life, at least centrally, but showing,
when dried, the underlying bony sculpture; the disk with a
distinct nuchal excavation, less convex, shorter and rounder than
in frcnatiim; leathery margin not extensive, posteriorly scarcely
extending beyond tail.

Plastron of hatchlings smootli, without callosities; in young
(90 mm., M.C.Z. 4300) callosities present on preplastra, on the
fused epiplastra, hyo-hypoplastra and xiphiplastra ; at this age
the anterior edges of the hyo-hypoplastral callosities are almost
straight, later becoming strongly convex anterolaterally ; in
adults the azj-gous callosity is subcircular, large ; hyo-hypo-
plastral callosities in contact with xiphiplastral callosities by long
straight sutures; femoral and caudal flaps permit concealment
of hind limbs and tail.

Osteological description} Skull. Distance between tip of pre-
maxilla and orbit greater than long diameter of orbit ; height of
orbit about one-third greater than the interorbital width, M'hich
is somcM'hat less than the width of naris; premaxilla apparentl}^
absent ; intermaxillary foramen small, pear-shaped with apex
directed forwards; both in front and behind this foramen the
maxillae in contact medially, the posterior medial suture much
longer than the foramen ; vomer slender without ascending proc-
esses to prefrontals, its interchoanal portion relatively elongate ;
choanae restricted anteriorly by inward expansion of the tritur-
ating surfaces, expanding abruptly posteriorly ; postorbital arch
much wider than the long diameter of orbit ; orbit not entered
by jugal, which is excluded by broad contact of postorbital and
maxilla but is broadly in contact with parietal ; squamosal crests
long, sharp, incurved ; pterygoids meeting opisthotic, greatly re-
stricting postotic fenestra ; mandible without symphysial ridge,
its ventral symphysial width much less than diameter of orbit.

Carapace. Carapacial bones finely granulated and vermicu-
lated ; no prenuchal bone ; nuchal bone notched laterally, under-
lying the first pleurals, between which are two neurals ; neurals
8 or 9, forming a continuous series ; pleurals 8 pairs, the eighth
in contact medially.

Plastron. Preplastra (no information) ; epiplastra fused, form-
ing an obtuse angle, their pointed posterior prong inserted

1 Based on B.M. 63.6.13.5 only.

456

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

in a deep notch which separates the rounded inner margin from
the strongly convex outer margin of the fused hyo- and hvpo-
plastra ; the posterior border of this compound bone very deeply
excavated, its anteromedial process with 2-3 juxtaposed prongs,

Fig. 59. Skull of Cycloderma aubryi (B.M. 63.6.13.5), x %. Dorsal and
ventral views.

(E, E. Turlington del.)

LOVERIDGE AND WILLIAMS : AFRICAN CBYPTODIRA 457

the medial process very short, not projecting beyond the rim, the
posteromedial process with 3-4 prongs, between the lateral three
of which are inserted the 2 anterior prongs of the xiphiplastron ;
xiphiplastra rather wide with 3 prongs posteriorly, in contact
medially by a prong of one fitting into a notch on the other.

Callosities 7, sculptured like the carapace, extremely developed
in adults, covering most of the plastral area ; preplastral callosi-
ties large, nearly half-moon-shaped, in contact medially by
a long straight suture ; fused epiplastra bearing a large, almost
round, azygous callosity; hyo-hypoplastral callosities large, cov-
ering most of the surfaces of these bones and almost meeting
medially, meeting in an oblique straight line the large trape-
zoidal xiphiplastral callosity ; xiphiplastral callosities in contact
medially over their entire length.

Color. Carapace of young dull orange to yellow brown with
a few scattered brown or black specks or blotches and a chocolate
brown line on the anterior third of the vertebral keel.

Plastron mostly yellow, anteriorly a broad chocolate brown,
more or less V-shaped marking whose truncate arms are directed
posteriorly; from the axillae the arms of another, but much
larger, black, V-shaped figure converge posteriorly to cover the
femoral and caudal flaps ; under aspect of the fleshy margin yel-
low.

Head above, yellow brown with five almost hairlike (some-
times converging anteriorly) longitudinal lines, a median one
beginning on the crown extends down the neck (in line with the
one on the carapace), two arise in the interocular region and
terminate on the occiput, each of the remaining pair commence
at the nostril and, passing through the eye, continue along the
side of the neck; chin and throat whitish indistinctly flecked
with darker; limbs largely dark chocolate brown, each bearing
a large yellow patch.

Size. Carapace length over curve of the Gabon type, 455 mm.,
the disk alone over curve 330 mm., its breadth 310 mm. (fide
Dumeril :1856a) ; head and neck together 280 mm. A. Fernan
Vaz turtle with head and neck of exactly the same length (11")
is said by Cope (1859) to have a total length of 760 mm. (2' 6").
Carapace length of an Eala specimen (M.C.Z. 43000) is 90 mm.,
breadth 80 mm. ; both measurements being taken in a straight

458

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

line. Carapace lenajth of a Lake "Ogemwe" ( ? Azinguo) hatch-
ling (B.M. 1908.5.25.3), 55 mm., breadth 44 mm.

Ilahitat. Inchided in the rain forest fauna by Schmidt (1919).

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LOVERIDOE AND WILLIAMS: AFKICAX CRYPTODIRA 4.")! I

Localities. Cahinda: Chinchoxo. Belgian Congo: *Eala, Flau-
ilria, (i.e. Bokele), Equateur Province; *Tumba, Lake LooikIu
(B.M.). We concur Avith de Witte (1953) that the "Katanga:
lcS98" specimen is a very questionable record. French Equa-
torial Africa: Fernan Vaz (Fernando Vas) ; Gabon; *Lake
''Ogemwe" (B.M.) ( ?Azinguo) ; Lanibarene (Limbareni) ; Ogoue
(OgOAvai) ; Oubangui (Ubangi) River near Libengue (Libenge).

We know of no specimen actually taken in the Cameroons.
Its alleged presence there dates from a list by Sjostedt (1897)
who failed to collect it. Subsequently cited in the lists of Werner
(1898a) and Sternfeld (1910), but without material.

Range. Portuguese, Belgian (western) and French Congo.

Cycloderma frenatum Peters

lS.')4!i. Ci/clodrrma frenatum Peters, Monatsb. .'Xkad. Wiss. licrlin, p. 'JIH:

Zambezi Eivcr, Mozambique.
1882a. Peters, 14, pis. i-iiia.

1889a. Boulengcr, 26.5.

1894a. Giinthcr (1893), 618.

189(ia. Bocage, 97.

1897. Johnston (also 1898 ecL), 356, 361.

1900b. Tornier, 583.

iy02b. Sicbenroek, 834, fig. 13.

1902b. Tornier, 580.

1909a. Siebenrock, 592.

1913c-. Nieden, 64.

1929. Lindholm, 291.

1934a. Mertens and Miiller in Bust, 12.

1934. Pitman, 307,

193 7f. Loveridge, 489, 495.

1942e.' Loveridge, 251.

1946. Mitchell, 21.

1950. Rose, 330, figs. 198-199.

1950. Williams, 552.

1953c. Loveridge, 160.

1855. Cijclanostem frenalus Gray, 64 (ex Peters MS).
1860a. A.'spidochelys livingstonii Gray, Proe. Zool. Soc. London, p. 6, iil.

xxii, figs. 1-2: Tributaries of the Zambezi River, Mozambique.
1860f. Gray, p. 430.
1864c. Jleptathyra livingstonii Gray, 94.
1864c. Heptathyra frenata Gray (part), 94.

460 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1870e. Gray, 93.

1873b. Gray (part), 76.

1884a. Rochebrune (in error), 30.

Syno7iymy. Further citations of "frenata" will be found
under Cydoderma aubryi, with which this species was confused
by Gray (1864c). Aspidochelys livingstonii was supposed to be
generically distinct because of certain characters of the callosi-
ties; when Gray realized these developed with age he himself
(1870e) synonymized livingstonii with frenatum.

Cofnmon names. Zambezi Soft-shelled Turtle (English) ; Zam-
bezi Mud-Turtle (Loveridge) ; " cassi or ncassi" (at Sena and
Tete: Peters, but see below) ; kalibungu (Sena: Mitchell) ; lite-
tamera (Yao: Loveridge); ynhidundwe (Yao: Mitchell); nalii
(Makonde: Loveridge) ; nkhasi (Manganja and Cewa : Mitchell).

Illustrations. Peters (1882a) furnishes tine colored drawings
(pi. i, copied as our black and white Plate 18 [bottom]), of
dorsal and ventral aspects of what is presumably a $ ; also a
figure (pi. iiia) showing plastral marbling in the 9 ; skulls, skele-
ton, etc. (pis. ii-iii).

Description. Proboscis projecting, the distance from its tip
to the orbit equal to (in young), or longer than (in adults)
the orbital diameter ; nostril with papilla-like process projecting
upwards from its inframedian corner; upper lips meeting in
front to form a flat arch; forefoot with 4 or 5 sharp-edged,
crescentic skin-folds on its upper surface, another weal-like
thickening on outer aspect ; hind foot with a sharp-edged crescen-
tic fold under the heel posterior to the base of the fifth toe ; tail
rudimentary,^ not projecting beyond the posterior leathery rim
of the carapace.

Carapace of young with a moderate vertebral keel and numer-
ous, raised, rather wavy, longitudinal lines; carapace of adults
smooth in life, at least centrally, but showing, when dried, the
underlying bony sculpture; the disk is less excavated anteriorly'
than in aiibryi, besides being more convex, longer and narrower;
leather}' margin not extensive posteriorly, scarcely extending
beyond tail.

Plastron of hatchling smooth, without callosities (in the young
these first appear as small pairs on the preplastra, hyo-hypoplas-
tra and xiphiplastra, while the last to develop is the azygous

1 Cf. Sexual dimorphism below.

LOVERIDQE AND WILLIAMS : AFRICAN" CRYPTODIRA

461

element on the fused epiplastra) ; femoral and caudal flaps per-
mit concealment of hind lim])s and tail.

Fig. 61. Skvdl of Cycloderma frenatuvi (B.M. 84.2.4.1), (Zambezi speci-
men), X %. Dorsal and ventral views.

(E. E. Turlington del.)

462 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Osteological description. Skull of adult. ^ Distance between
tip of premaxilla and orbit subequal to long diameter of orl)it ;
height of orbit about twice the interorbital width, which is less
than the width of naris; premaxilla apparently absent (rudi-
mentary, fide Peters in Zambezi specimens) ; intermaxillary fora-
men small, heart- or pear-shaped with apex directed forwards;
both in front and behind this foramen the maxillae in contact
medially, the posterior medial suture longer than the foramen ;
vomer very slender, without ascending processes to the pre-
frontals, its interchoanal portion relatively elongate, or vomer
absent ; internal choanae narrowed anteriorly by inward expan-
sion of the triturating surfaces, expanding gradually posteriorly ;
postorbital arch much wider than the long diameter of orbit ;
orbit entered by jugal, which is broadly in contact with parietal ;
squamosal crests long, sharp, incurved ; pterygoids meeting
opisthotic, greatly restricting postotic fenestra; mandible with-
out symphysial ridge, its symphysial width less than the long
diameter of the orbit.

Carapace. Carapaeial bones finely granulated and vermicu-
lated ; no prenuchal bone ; nuchal bone notched laterally, under-
lying the first pleurals, between which are two neurals, rarely
one ;^ neurals 8 or 9, rarely 7,- usually forming a continuous
series; pleurals 8 pairs, the seventh and eighth, or eighth only
in contact medially.

Plastron. Preplastra widely separated, only the anterior
branch present ; epiplastra fused, forming an acute angle, their
pointed posterior prong inserted in a notch which separates the
rounded inner margin from the strongly convex anterior border
of the fused hjo- and hypoplastra ; posterior border of this com-
pound bone very deeply excavated ; the anteromedial process
with 3 prongs, the medial process typically very short and, as a
rule, scarcely projecting beyond the rim; the posteromedial
process with only 3 prongs between which are inserted the 2
anterior prongs of the xiphiplastron ; xiphiplastra short and

1 The skull, with condylobasal length of .31 mm., of a juvenile skeleton
(A.M.N.H. 56479) from Lake Nyasa, Nyasaland, shows striking differences in the
pi-oportions to those of the adults described above. For examine, the distance
between tip of premaxilla and orbit is very much shorter than the long diameter
of the orbit. Presumably similar discrepancies due to ontogenetic change would
have been noticed in other African trionychids had skeletal preparations of their
young been available.

2 One only between the first pair of pleurals, and only 7 neurals In one (M.C.Z.
48032) of a series of seven from the Rovuma River.

LOVERIDGE AND WILLIAMS: AFRICAN" CRYPTODmA

463

broad with 3 or 4 notches posteriorly, on the posterior half of
each notch a short process joining a similar process on the other
element.

Callosities 7, sculptured like the carapace, very well developed

.a

0

o

00

Pi

Ci

i ^

S o

V^ CO

e pa

'Si

02 ^
eg ^

si .-C

464 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

in adults; preplastral callosities large oblique ovals, in contact
medially; fused epiplastra bearing a small, crescentic to semi-
lunar callosity, smallest of any and last to develop; hyo-hypo-
plastral callosities covering most of the surfaces of these bones
but widely separated medially; hyo-hypoplastral callosities sep-
arated from, or in contact with the xiphiplastral callosities by
only a short suture ; xiphiplastral callosities large oblique ovals,
in contact medially.

Color. Carapace in Lake Nyasa hatchlings ranges from pale
green to leaden, the periphery usually edged with white ; cara-
pace of adults pale to dark olive, uniform or with a trace of
blotching. Zambezi turtles would appear to differ somewhat,
for Peters (1883a) describes young specimens as green and
adults as dark green, uniform, or with interrupted black bands
and scattered white spots.

Plastron of Lake Nyasa hatchlings was often almost white,
more usually an irregular black blotch was present in the um-
bilical region, an elongate, oblique blotch near each forelimb, a
rounder one anterior to each hind limb, and a smaller sixth spot
in the anal region. Plastron of adult $ 9 is china white to
flesh-pink almost obscured by pearly gray reticulations. These
reticulations are sometimes absent in what may he $ $ . For
example, the subject of Peters' colored plate (1882a, pi. i),
possibly a Zambezi $ , is shown as having a cream-colored plas-
tron extensively infuseated with dusky blotches.

The head and neck in Lake Nyasa hatchlings were gray with
a dark, light-edged, interorbital crossbar, five similar, but wavy
and sometimes broken, longitudinal lines from near occiput to
base of the long neck ; throat and underside of neck pure white,
uniform or showing some dusky streaks. Head and neck of adults
very dark olive, the dark longitudinal lines of the young turtle
obsolete or, more usually, absent. Peters (1882a), writing of
Zambezi turtles, states that the head and neck in both age
groups display numerous white dots. Feet gray above, lighter
below.

Size. Carapace length over curve of a Zambezi $ , 560 mm.,
the disk alone over curve 470 mm., its breadth 420 mm. (fide
Peters :1882a, whose figures do not quite tally, for he gives the
total length as 970 mm., though head and neck together are only

LOVERIDGE AND WILLIAMS: AFRICAN CRTPTODIRA 465

340 mm., the tail, which would not project beyond the carapacial
margin, 65 mm.).

Overall carapace length {in a straight line) of three Nyasa
$ 9 averaged about 560 mm., breadth 418 mm. The head
and neck of one (M.C.Z. 50357) measures 420 mm.; breadth of
head 100 mm. Disk length of largest gravid Rovuma 9 (M.C.Z.
48030) 390 mm., its width (also without leathery margin) 310
mm. (Loveridge :1942e and 1953e).

Overall carapace lengths of 31 Nyasa hatchlings were from
40 to 48 mm., their breadths 30 to 36 mm. Peters mentions a
Zambezi hatchling of 45 mm.

Weight. AVeight of largest Zambezi $ between 28 to 30 lbs.
(as 13-14 kilos., Peters :1882a) ; that of largest Rovuma 9 , 25
lbs. (Loveridge :1942e) .

Sexual dimorphism. Possibly the sexes of adults, though not
of young, may be told by the fact that the tail of the $ is visible
in ventral view (cf. Peters :1882a :pl. i), while that of the 5
is concealed beneath the caudal flap (ibid., pi. iiia, fig. 1). We
are unable to confirm Peters' statement that the carapace is nar-
rower in S S than in 9 9 (the relative proportions said to
be 1:1^/^ and 1:1^) or that the union betAveen carapace and
plastron is longer in (J o than in 9 9.

Breeding. On January 11th, at Tete, a hatchling with umbili-
cal scar was found by Peters (1882a). Between December 24th
and earl}^ February, at Fort Johnston, nine hatchlings were
taken by B. L. Mitchell (letter of 10.iv.47). On February 7th,
at Mtimbuka, natives brought in three clutches of eggs number-
ing 15, 16 and 19, respectively. On February 10th, 19th and
28th, three 9 9 were captured when coming ashore to lay. All
three held spherical, hard-shelled eggs ranging from 33 to 35 mm.
in diameter. One 9 laid 3 eggs before being killed, 19 more
were present in her oviducts, and many shell-less ova of various
sizes were present in the ovaries. On February 7th, the fourth
day after a heavy downpour that terminated about eight months
drought (but did not inaugurate the rains as was hoped), a
young turtle, its carapace caked in mud, was found. Twenty
other hatchlings were collected during the three weeks following
(Loveridge :1953e). On March 27th and 28th, at Kitaya, two
9 9 held 17 and 19 hard-shelled eggs ranging from 31 to 32

466 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

mm. in diameter; in addition one turtle held 55 small ova in
various stages of development. (Loveridge :1942e).

Diet. The powerful, all-pervading, fishy odour of Kitaya
turtles led Loveridge (1942e) to suggest that these reptiles sub-
sist largely on fish in the Rovuma River. Mitchell found that
captive hatchlings from Lake Nyasa eagerly took fine strips of
fish or meat from his fingers. However, he concluded (1946:21)
that feral adults subsist principally on aquatic snails and mus-
sels. At Mtimbuka the former were Lanistes eUipticus and L.
sordidus; the clams proved to be Mutela alata and simpsoni,
with a robust cockle-like shell {Caelatura nyassensis) particu-
larly abundant (Loveridge :1953e).

These large mussels embed themselves vertically in the mud
or sandy bottom of lakes and ponds, and considerable effort is
required to dig them out. For the purpose, Cycloderma must use
the powerful claws on its forefeet. Usually the shells are
crushed and swallowed, though occasionally undamaged valves
have been defecated by freshly captured turtles (Mitchell:
1946).

Parasites. Leaches were present on turtles at Kitaya and
Mtimbuka; two removed from the neck of a Lake Nyasa 9
(M.C.Z. 50357) were of the family Glossiphonidae, possibly
Placohdella jaegerskioeldi.

Enemies. In 1926 Dr. J. 0. Shircore observed two small
rufous otters running around a dessicating puddle about 100
yards from the right bank of the Kilombero River, Mahenge
District. The object of their interest appeared to be a soft-
shelled turtle that had withdrawn within its defenses. At least
this was its position when he reached the spot after shooting one
of the otters. Subsequently, Dr. Shircore sent a photo of turtle
and otter to one of us (Loveridge :1942e).

On February 11th, 1949, Loveridge saw scores of eggshells on
the village middens around Mtimbuka. A Yao, whose home was
only about ten miles north of Mtimbuka, accepted turtle eggs
gladly, but the more sophisticated members of the staff scorned
the idea of eating them. Loveridge, himself, ate 18, finding them
quite edible though not as palatable as fowl's eggs. It is when
coming ashore to lay that these turtles fall an easy prey to
Africans; consequently S S are rare in collections (Loveridge:
1953e).

LOVEBIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 467

Defense. Though presumably, wheu molested, Cycloderma is
capable of inflicting a severe bite with its strong crushing jaws,
instead it withdraws both head and forelimbs within the fleshy
margins of its shell, the hind limbs beneath dermal flaps, thus
completely closing the plastron before and behind. When picked
up, however, a Cycloderma is occasionally difficult to handle, for
vigorous kicking of the clawed feet, which can reach any part of
the shell's edge, may displace the captor's hands (Mitchell:
1946).

Temperament. In stating that Cycloderma "are very fierce,"
Johnston (1897) w^as apparently attributing to them the well-
merited reputation of Trionyx. Actually these turtles are timid
and inoffensive, withdrawing their heads and limbs within the
protection of the shell at the slightest disturbance. When all is
quiet, the head is protruded -with the utmost caution by very
gradual stages and withdrawn precipitately to the accompani-
ment of a kind of snort at the least sound or movement in their
vicinity (Loveridge :1942e :252, which consult for a concrete ex-
ample of such timidity). This inoffensiveness is confirmed by
Mitchell (1946).

Hahits. Surprisingly active according to Peters (1882a), both
in the water and on land where, using their forefeet and snout,
they quickly burrow into soft mud in an effort to hide. One of
these turtles, wnth head held well above the surface, paddled
swiftly past Loveridge (1942e) as he was wading thigh-deep in
a lake. When twenty feet away the reptile dived and was seen
no more.

Mitchell (1946) states that during the daytime two captive
turtles, that he kept in a pond at Salima, remained for long
periods with their nostrils protruding from the w^ater. They did
not rise to the surface between 4.00 P.M. and 8.30 A.M. and re-
mained on the bottom during cold weather. He suggests that at
such times they obtain oxygen by means of the mass of blood
vessels on the unpigmented ventral surface in the vicinity of the
flaps. Their only attempts at escape w^ere made at night during
rainy weather, or after the water in their pond had been changed.
One turtle that got away was recaptured a fortnight later;
though wdthout water during this time the reptile was apparently
none the worse.

468 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

In Januar}^ when Loveridge (1953e) endeavoured to secure
topotypic frenatum near Tete, local native fishermen asserted
that these turtles would not be seen until the rains, already two
months overdue, broke.

Habitat. Rivers, lakes and stagnant ponds.

Localities. Tanganyika Territory: Kilombero River, Mahenge
District (sight record by Shircore ; photo seen by A.L.) ; Lake
Nyasa at Manda (as Wiedhafen) ; *Rovuma River at Kitaya.
Mozambique: Lake Inhalutanda near Tete; Licuare River, trib-
utary of the Zambezi ; *Zambezi River near Tete. Nyasaland.
*Chowe; *Lake Nyasa at Mtimbuka and at Fort Johnston^;
Lake Shirwa (Chilwa) ("said to occur": Mitchell); Upper
Shire; Lower Shire ("said to occur, but rare": Mitchell).
'] Northern Rhodesia: chelonian remains near Munyamadzi River,
Luangwa Valley, Mpika District {fide Pitman).

The erroneous recording by Tornier (1900b) of Fiilleborn's
specimen as from "Wiedhafen, Victoria Nyassa, " resulted in
later authors adding Victoria Nyanza or Lake A^ictoria to the
range.

Range. Southern Tanganyika west and south through Nyasa-
land and possibly Northern Rhodesia to Mozambique (Zambezi
River).

Gray's records (18G4c: 1873b) for Gabon were due to con-
fusion with aubryi.

MARINE TURTLES

General Remarks

The treatment being given to the marine turtles of the families
CHELONIIDAE and DERMOCHELYIDAE, differs substan-
tially from that accorded to the other Cryptodira of Africa.
While African to the extent that, without exception, the}^ land on
the continent or its offshore islands to deposit their eggs, the
group is so cosmopolitan that any revisionary study of them
should be undertaken on a worldwide rather than a regional
basis. This, lacking adequate material, we have not attempted.

The literature dealing with African records, to which — with

1 We presume that this is also the correct locality for a British ^Museum speci-
men labeled "Zomba." Zomba, where Sir Harry Johnston lived, was presumably
only the dispatching point of the donor.

LOVEBIDQE AND WILLIAMS: AFRICAN" CBYPTODIBA 469

few exceptions — we have restricted our citations, is scanty in
the extreme. Though there are good reasons to suppose that
turtles are by no means rare in the waters surrounding the Afri-
can continent, yet for extensive areas there are curious gaps in
the record. Such hiatuses are presumably due to the failure of
travellers to collect and preserve such admittedly cumbersome
reptiles.

In the following account, therefore, we present no figures, no
analyses of the species, no fossil history, and relatively little gen-
eral discussion or comment on habits, as we were averse to draw-
ing on extra-territorial sources.

In conformity with this treatment, and in the absence of the
necessary material, we have refrained from discussing the alleged
Atlantic and Pacific races of the green turtle, hawksbill, logger-
head and leatherback. In all these instances we have listed only
the species name and, following it as synonyms, the putative
subspecies. Only in the case of the much more distinct Lepi-
dochclys races have we registered a decision. We have omitted, as
not germane to our area, the problem of the status of Chelonia
depressa Garman.

For those who desire more complete discussions of the Recent
marine turtles, we recommend Deraniyagala (1939) and Carr
(1952). The fossil marine turtles are currently being studied by
Dr. Rainer Zangerl, to whose publications those interested must
be referred.

Incidence of occurrence of the various species on the Senegal
coast has been furnished by Cadenat (1949:17) but the figures
also reflect edibility or marketable value, suggesting that selec-
tion practiced by local fishermen may have influenced the results.

Table 10
Marine turtles caught on the Senegal Coast.

Species Total Individuals Captured Incidence

Chelon,ia mydas 256 86.48%

Eretmochelys imbricata 23 7.77%

Caretta and Lepidodhelys 10 3.39%

Dermochelys coriacea 7 2.36%

470 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Nomenclature. On several occasions Deraniyagala (e.g. 1943)
has asserted, though not formally proposed, that on grounds of
priority the names of the loggerhead turtles require to be totally
altered from present usage, Caretta being the correct name of
the forms currently called Lepidochelys, and Thalassochclys the
correct name of those nov7 called Caretta. The use of the name
Thalassochelys would involve little hardship since this name is
classic, having been applied to both loggerheads by Boulenger
(1889a) in the catalogue that all discreet students of this order
still faithfully consult. How^ever, an exchange of names such as
would be required by the use of Caretta for the olive-green log-
gerheads, rather than for the red-brown forms, is a perversion
of nomenclature to be avoided at almost any cost. We therefore
venture to comment upon Deraniyagala 's reason for proposing
the change. He calls attention to the fact that Linnaeus' species
Testudo caretta was a composite. He believes that the first un-
ambiguous use of the Linnaean name is that of Schoepff (1792),
who presents several figures under this name. From the scala-
tion shown in these figures, Deraniyagala deduces that Schoepff
was in effect restricting Testudo caretta to the form now known
as Kemp's Loggerhead.

However, Schoepff' 's figures appear in editions both with and
without colored plates. There are at the Museum of Comparative
Zoology four copies of Schoepff, two without and two with col-
ored plates. In the latter editions, apparently not examined by
Deranij^agala, the figures of Testudo caretta are colored a very
fine red-brown, in shade and texture a splendid representation
of the red-brown loggerhead. Thus, accepting Deraniyagala 's
interpretation of the scalation at face value, the color of these
plates seems to demonstrate that Schoepff 's concept of Testudo
caretta was, like that of Linnaeus, a composite one.

We have not concerned ourselves with Deraniyagala 's reason-
ing in other regards, nor have we pursued the matter further.
We merely wish to protest against the spirit of antiquarian and
wholly bibliographic research which leads to these proposals for
name changes. We point out that such researches (e.g. in the
case of Trionyx) almost invariably lead to confusion and insta-
bility, insoluble except by the plenary powers of the International
Commission. This is the more certain since antiquarian re-

LOVERIDGE AND WLLLIAMS : AFRICAN CRYPTODIRA 471

searches will rarely be complete and thus will leave us for all
future time at the mercy of the chance discovery of some rare
and (nomenclature apart) worthless book.

We deprecate also the forcing of text or figures of the oldest
writers into modern standards of precision. As everyone is
aware, the figures of the older naturalists were as often as not
composite and (with praiseworthy exceptions) rarely accurate,
while not infrequently descriptions are unrecognizable beyond
the order. The most exhaustive discussion and analysis of such
figures and texts can never carry conviction or produce agree-
ment.

In the present case the Copenhagen rules may prevent in fact,
as they certainly prohibit in spirit, the changing of these long-
accepted names.

Key to ihe Marine Turtles Breeding in Africa

1. Upper jaw conspicuously bicuspid at symphysis; upper shell covered with

smooth skin (or i^inaU scales in juveniles) overlying a mosaic of sninll
bones and showing 7 prominent longitudinal ridges ; limbs clawless ....

Bermochelys coriacea (Linnaeus) (p. 499).

Upper jaw not bicuspid; upper shell covered with large horny shields

overlying large bony plates, and ridges 0 to 3 ; limbs with 1 or 2

claws 2

2. Upper shell with 4 pairs of costal shields of which the foremost pair is

never the smallest and is separated from the nuchal shield 3

Upper shell with 5 or more pairs of costal shields of which the foremost
pair is the snmllest and normally in contact with the nuchal shield . . 4

3. Snout not compressed; 2 prefrontal shields on head; shields of upper

shell not overlapping (except in very young turtles) ; usually a single

claw on each limb .Chelonia mydas (Linnaeus) (p. 474)

>Snout compressed ; 4 prefrontal shields on head ; shields of upper shell
strongly overlapping (except in very young or very old turtles) ;

usually 2 claws on each limb

EretTuochelys imbricata (Linnaeus) (p. 485)

4. Upper shell normally with only 5 pairs of coastal shields; bridge on either

side of lower shell with 3 enlarged inframarginal shields without pores ;

color of adults and young predominantly reddish brown

Caretia carctta (Linnaeus) (p. 490)

Upper shell normally with 6 to 9 (rarely 4 or 5) pairs of costal shields;

bridge on either side of lower shell with 4 enlarged inframarginal

shields, each with or without a pore; color of adults predominantly

olive, of young olivaceous black

Lepidochelys olivacea oUvacea (Eschscholtz) (p. 495)

472 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Family CHELONIIDAE

1882. Cheloniidae Cope, Proc. American Philos. Soc, 22. p. 143.

Defiyiition. Cryptodirous testudinates adapted to marine life.
Horny shields normally present ; costal scutes 4 or 5 pairs ; mar-
ginals, exclusive of the nuchal and supracaudals, 11 or 12 pairs ;
inf ramarginal series complete ; additional axillary shields also
present ; plastron with 6 pairs of scutes and one unpaired scute ;
an intergular commonly present.

Skull without nasal bones; prefrontals always in contact dor-
sally, always with descending processes that are moderately sep-
arated inferiorly; temporal region posteriorly very little
emarginate ; parietal meeting squamosal ; no bones tending to be
reduced; quadrate never enclosing stapes; postotic antrum ali-
sent ; parietals with reduced descending processes ; upper jaw
with or without ridges on its triturating surfaces ; vomer alwaj^s
present, separating palatines ; mandible with coronoid bone.

Neck vertebrae with only one biconvex centrum, usually the
fourth, typically a plane joint between the sixth and seventh
centra, the eighth centrum concave in front, doubly or not ; cora-
coids with their median borders moderately expanded; humerus
of a specialized marine type with deltopectoral crest far down
the shaft; trochanteric fossa of femur reduced by union of the
trochanters; phalanges without condyles; claws 1 or 2.

Carapace heart-shaped, united to plastron only by ligament ;
neither carapace nor plastron hinged; pleural bones somewhat
reduced and often peripheral fenestrae persisting till late in
life ; nuchal without costiform processes but with a ventral at-
tachment area for the eighth cervical ; neural bones hexagonal,
short-sided in front, variable in number ; pygals 3 ; plastron never
cruciform but \^dth some development of fontanelles; entoplas-
tron always present, more or less lance-shaped.

Range. Tropical, subtropical and — as an occasional visitor —
temperate oceans.

Genus ChELONIA Bronguiart
1800. Chelonia Brongniarti (part), Bull. Soc. Philom. Paris, 2, p. 89.

1 In 1801 this uame was used by Sonnini and Latreille (Hist. Nat. Kept., 1,
p. 22) but they correctly attributed it to Brongniart, whose article we have
examined.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 473

"Ce sont les tortues de nier. " Type: Testudo mydas Linnaeus
(designation by Fitzinger: 1843).

1837. Chelona Buinieister, Handbuch Naturg., 2, Abt. Zool., p. 731. Type:

T. mydas Linnaeus (by nionotypy).

1838. Mydas Cocteau (not of Fabricius: 1799), Kept, in De la Sagra, Hist.

Fis. Pol. Nat. Cuba 4, p. 22. Type: T. mydas Linnaeus (by tau-
tonomy).
1843. Mydasea Gervais, Diet. Hist. Nat., 3. p. 457. Type'- T. mydas Lin

naeus (by nionotypy).
1845. Euchelonia Tschudi, Fauna Peruana, p. 22. Tj-pe: I. mydas Lin-
naeus (by monotypy).
1848. Megemys Gistel, Naturg. Thier., p. viii. Nomen novum for Chelonia.
1858. Euchelys Girard, U. S. Explor. Exped. 1838-1842, Herp., p. 447.

Type: E. macropus Girard =^ T. mydas Linnaeus (by monotypy).
1862a. Chelone Strauch, Mem. Acad. Imp. Sci. St. Petersbourg, (7), 5.
No. 7, p. 59. Type: Testudo viridis Schneider = T. mydas Lin-
naeus (by original designation).
Definition. Head with 1 pair of prefrontal shields ; postoculars
3-4; claws 1.

Skull with triturating surface of maxilla ridged, the ridge
rising to a dentate projection at suture with the preinaxillae ; pre-
maxillae in contact with vomer which separates the maxillae;
maxilla with vertical ribbing on inner surface of its cutting edge ;
descending processes of prefrontals in contact with vomer and
palatines ; a blunt ridge on vomer and palatines at the anterior
margin of the internal choanae; choanae in ventral view con-
cealed by the extensive secondary palate; pterygoids flat pos-
teriorly ; frontal usually entering orbit ; crista pretemporalis -re-
duced; mandibular symphysis short; the somewhat serrate labial
margin (the horny sheath is sharply serrate) rising to a sharp
point at the symphysis; the lingual margin surmounted by a
wider blunt ridge rising to a higher point at the symphysis, the
two points united by a sharp symphysial ridge.

Carapace with persistent lateral fontanelles; neurals 9-11,
usually hexagonal, short-sided in front; peripherals 11 pairs;
the tenth pair not in contact with the ribs ; nuchal shield not in
contact with first costals ; costals typically 4 pairs.

Plastron with a large intergular shield; inframarginals 4
pairs, without pores.

Range. Tropical, subtropical and — as an occasional visitor —
temperate oceans.

474 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOQY

Chelonia mydas (Linnaeus)^

1758. Tcstudo Mydas Linnaeus, Syst. Nat., ed. 10, 1, p. 197: Ascension

Island.
1802h. Daudin, 10, pi. xvi, fig. 1.

1860. Tristram, 405.

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

locality.

1783. Tcstudo Vifidis Schneider, Natur. Sehildkroten. p. 299: No locality.

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

vii, fig. 1 : Japan.

1788. La Tortue Franche Lacepede, Hist. Nat. Quad. ovip. Serpens, L p.

5-i; and Testudo marina (sen vulgaris) in Synopsis Methodica, a
table (in which binomials are used) at end of same volume:
Torrid Zone.

1788. La Tortue ecaille-vcrte Lacepede, Hist. Nat. Quad. ovip. Serpens, 1.
p. 92; and Testudo viridi-s-squamosa in Synopsis Methodica, a
table (in which binomials are used) at end of same volume:
Amazon Eiver, Brazil.

1802b. Testudo cepediana Daudin, Hist. Nat. Rept., 2, p. 50, p!. xvii, fig. 1 :
No locality.

1812. Chelonia virf/atn (Dumeril; Schweigger, Konigsberger Arch. Natur-
wiss. Math., pp. 291, 411 : Seas of the Torrid Zone.

1814. Schweigger, 21.

1835. Dumeril and Bibron, 541.

1S49. Smith, A., App., 2.

1855. Gray, 74.

1873b. Gray, 93.

1820. Caretta esculenta Merrem, Vers. Syst. Amphib., p. IS: Atlantic Ocean.

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

1820. Caretta Thunbergii Merrem, Vers. Syst. Amphib., p. 19: Japan.

1829. Chelonia maculosa Cuvier, Eegne Animal, ed. 2, 2, p. 13: No locality.

1829. Chelonia ladirymata Cuvier, Regue Animal, ed. 2, 2, p. 13: No lo-
cality.

1834. Clielonia hicarinata Lesson, in Belanger, Voy. Tndes-Orient., Zool..

p. 301 : Atlantic Ocean.

1835. Chelonia Marmorata Dumeril and Bibmri, Erpet. Gen., 2, p. 546,

pi. xxiii, fig. 1 : Ascension Island.
1844. Clielonia viridis Gray, 54.
1855. Gray, 75.

1 For tho niosf pnrt this bihlingraiihy is rrstrictecl to synonyms, togcthpr with
citations dealing with African material. Nonienclatorial changes such as Chelone,
Chelonia, midas and mydas call lor carel'ul checking.

LOVERIDGE AND WILLIAMS: AFRICAN CBYPTODIRA

475

1875.
1858.

1858.

1865.

1885a.

1885.

1888a.

1890.

18G6a.

1890.

1868.

1872.

1878.

1882.

1882a.

1906a.

1909a.

19121).

1929.

1933.

1936J.

1937a.

1937e.

1937f.

1949.

19-49.

1950.

1950.

1873b,

1884a,

1880.

1887.

1889a

1890.

1893a

1895a

Melliss, 99.
Chelonia formosa Girard, U. S. E-xplor. Expcd. 1838-1842, lU'ip.,

p. 456, pi. x.\xi, figs. 1-4: Fiji Islands.
Chelonia tennis Girard, U. S. Explor. Exped. 1838-1842, Ilorp., p.
459, pi. xxxi, fig. 8: Ilondcn Island, Pauniotu Group, Tahiti and
Eimeo ; Rosa Island.
ChcJonc viridis Strauch, 141.

Boettger, 172 (as marine turtle, later identified as viridis).
Greeff (1884), 49. footnote.
Boettger, 174.
Biittikofer, 1. 302.
Chelonia midas Bocage, 41.
Biittikofer, 1, 266, 269.
Chelonia Agassizii Bocourt, Ann. Sei. Nat. Zool. (Paris), (5), 10,

p. 122: Nagulate River mouth, Guatemala.
Chelonia mydas Sowerby and Lear, pis. lix-lx.
Reichenow, p. 92.
Pechuel-Loesche, 277.
Peters, 18.
Mocquard, 481.
Siebenrock, 545.
Werner, 454.
Flower, 38.
Flower, 750.
Loveridge, 221.
Flower, 13.
Hewitt, 15.
Loveridge, 488.

Cadenat, 16, figs. 1-3, 10, 14, 15a.
Villiers, 165.
Rose, 327.
Williams, 552.
Mydas viridis Gray, 95.

Rochebrune, 40.
Chelonia deprcssa Garman (part at least), Bull. Mus. Comp. Zool.,
6, p. 124: East Indies juv. (N. Australian adult may be distinct.
Also following Fry (1913), we omit Xatator tessellatus McCuUoch.
1908, from Port Darwin, X. Australia, from the synonymy).
Chelonia lata Philippi, Zool. Garten, 28. p. 84: near Valparaiso, Chile.
, Chelone mydas Boulenger, 180.
Biittikofer, 2. 438. 478.
Boettger, 12.
Bocage, 6.

476 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1896a. Bocage, 74, 98.

1896. Lonnberg, 11, 12 (on Linnaeus' type).

1901. Gadow, 381.

1901. Steindachner, 326.

1903a. Bocage, 52.

1903a. Boulenger, 92 (footnote), 96.

1906. Johnston, 819, 833, fig. 311.

1906b. Siebenrock, 39.

1908. Soidelli, 17,

1911d. Sternfeld, 51.

1913. Boettger, 318, 330, 332, 335.

1915. Eawitz, 657, pi. xlviii, figs. 59-61.

1923g. Loveridge, 930, 933.

1924b. Loveridge, 3.

1925b. Flower, 932.

1927. Calabresi, 37.

1938. Cozzolino, 241, graphs.

1947. Irvine, 309.

1955. Cansdale, 95, 104.

1925b. Chelone virgata Flower, 933.

1931. Chelone midas Ingrams, 429.

1934a. Clielonia my das my das Mertens and Miiller in Eust, 10.

1934a. Clielonia mydas japonica Mertens and Miiller in Rust, 10.

Common names. Green Turtle; Edible Turtle (English);
anjwa or apuhulu (Nzima of Gold Coast :Irvine) ; apuhuru
(Fante of Gold Coast :Irvine) ; assa (Cape Delgado :Peters)
hala (Ga of Gold Coast :Irvine) ; itataruca (Mozambique Id.
Peters); kassa (Swaliili Jngrams) ; klo (Ewe at Keta:Irvine)
nruvi (Mozambique Id. :Peters) ; taza (Bajun of havnu -.fide
"Ngamba," 1932: The Field :159 :421).

Illustrations. The finest colored plates of this turtle, dorsal
and ventral views, are those in Sowerby and Lear (1872 :pls.
lix-Lx).

Description.^ Snout short; beak not hooked, without cusps;
edge of jaws apparently smooth ; prefrontals elongate, a single
pair ; frontal azj'gous, small ; frontoparietal large ; parietals 2 ;
supraocular rather large ; postoculars 4 ; supratemporals 2 ; fore-
limb with moderately enlarged scales along anterior edge, behind
them several rows of scales, posterior edge with a series of en-

1 Based exclusively on a single specimen from Lamu Island (M.C.Z. 40019) and
one from Madagascar (M.C.Z. 16866).

LOVERroGE AND WILLIAMS : AFRICAN" CRYPTODIRA 477

larg'ed scales; each fore- and hindlimb with 1 claw ;^ tail short.

Carapace ovate, sulitectiform (juv.) or smooth (adult), nuchal
region truncate, mar<;in serrate posteriorly; dorsal shields juxta-
posed ; nuchal broader than long, not in contact with first costals,
not in contact with second marginals; vertebrals 5, first much
the broadest, much broader than long, the rest as broad as, or
broader than long, or II to IV longer than broad; costals 4,
fourth smallest; marginals 11 pairs; supracaudal divided.

Plastron anteriorly subtruncate, no obvious lateral keels ; plas-
tral shields juxtaposed ; intergular moderate ; brachials 2 or
3 with some smaller scales extending anteriorly ; inf ramarginals
4, without pores; inguinal small; interanal minute or absent.

Color. Carapace of young olive to dark brown ; plastron white
or yellow. Crown and sides of head dark, throat yellowish ; limbs
above olive to brown margined with white or yellow, below pure
white or yellow with a large dark blotch, or blotches, or almost
entirely dark.

Carapace of adult essentially similar to that of young, but
with radiating lines or marbling; plastron yellow, immaculate.
Head dark above, but on the sides the shields are more conspicu-
ously edged with yellow.

The foregoing color description of the species is general as
our African material is so scanty. For more detailed descrip-
tions consult Deraniyagala (1939:228-230) or Carr (1952:348-
349).

Size and Weight. Three juveniles, each about 5 inches (.127
meter) in length and weighing a lb. (.45 kilo), when received at
the London Zoological Gardens in July-August, 1924, developed
in the course of about 9 years and 4 months to 50 lbs. (22. .68
kilos) each on November 8, 1933 (Flower :1937a).

Biittikofer (1890:1:260) mentions a Kobertsport specimen as
being 1 meter long by 75 cm. broad. The largest individuals
from Europa Island were from 314 to 4 feet in length with
weights up to 999 lbs. (450 kilos), fide Siebenrock (1906b).
This is considerably in excess of the average adult weight of 336
lbs. mentioned by Gadow (1901 :381) .

Sexual dimorphism. Males have much the longer tails accord-
ing to Voeltzkow {in Boettger :1913).

1 Two claws are present on both fore- and hindlimbs of very young specimens,
iit which stage they correspond to Euchelys macropus Girard, as noted by Peters
(1882a) and also observed by Deraniyagala (1939 : 22-1).

478 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Breedi7ig. On the West African coast the laying season is
from September to January (Gadow :1901 :382). More specifi-
cally on Rolas and San Thome Islands it is from December to
January (Greef :1885). Where the warm waters of the Gulf of
Guinea break on the beaches of the northern Loango Coast, Green
Turtles emerge to lay, at the beginning of the November rains.
They are scarcer on the southern shores of Loango which are
washed by the colder Atlantic currents.

That breeding takes place off the South African coast is ad-
duced by Hewitt (1937e) from the occurrence of an occasional
hatchling with a carapace length of 50 mm. and umlnlical scar
still unhealed.

In Aldabra Island lagoon the S $ fight desperately with
each other during the breeding season, the natives told Voeltzkow
{in Boettger :1913), but at Europa Island they usually remain
outside the reef, rarely visit the bay, and never come ashore. At
low tide during the period — December 4th to 20th — of Voeltz-
kow's visit, one might observe the 9 9 as black patches against
the white sand of the lagoon. As night fell they began coming
ashore in considerable numbers. From his vantage point on the
ship Voeltzkow observed seven emerge at the same time. He
notes that the turtles made numerous trial excavations without
laying, and describes at some length the technique they em-
ployed. He frequently found eggs, sometimes an entire laying,
decomposed, apparently as a result of heavy rain and too much
moisture. This led him to conclude that Europa Id. was by no
means an ideal breeding site for these reptiles. The mortality
among laying 9 9 was high for their corpses were encountered
along the beach and among the dunes, as many as five being met
with in the course of one walk. On Aldabra, said the natives,
the 9 9 disappear after reaching a certain age and are covered
with barnacles when they return to the island for laying. This
is chiefly between May and December when, at intervals of six
weeks, a Green Turtle will come ashore three times, on each
occasion laying as many as 125 eggs. Voeltzkow 's full account
should be compared with that of Cozzolino, synopsized below,
as they differ in details. Probably the most detailed account
of a Green Turtle laying in Africa is that of Cozzolino (1938)

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 470

whose observations were made on a small island 10' S off
the Somali coast. Gravid 9 2 appeared offshore cruising to
and fro with their heads above the surface as if selecting a
suitable spot to land. Landing takes place chiefly during the
waxing, rarely during the waning, of the moon. On emerg-
ing from the water, leaving a characteristic imi)rint on the
sand, each turtle makes for the more elevated portions of the
beach beyond the reach of the highest tides.

As soon as the turtle reaches a suitable spot she promptly
starts excavating, the sickle-shaped forelimbs alternating Avith
the hind limbs. The former fling the sand aside, the latter
throw it backwards. In a very short time the extent of the oval-
shaped excavation begins to be apparent, and within twenty
minutes is carried to a depth of about two feet ("60 cm."), the
turtle disappearing from sight. She betrays her presence, how-
ever, by emitting a loud puffing sound that is audible to anyone
with normal hearing for a distance of sixty yards (50 meters).

At this juncture a second operation commences. The sand
being relatively loose at this depth, the tail tip is used to move
it over a small area corresponding to the cloacal orifice.^ Then
the hind flipper, being turned inwards to form a scoop, is em-
ployed to lift out the displaced sand until a second cavity is
excavated to a depth of about fifteen inches (35-40 cm.) with
a diameter of from ten to twelve inches (25-30 cm.).

On completion of this second operation the turtle brings her
hind limbs together to form an arc-shaped covering extending
from the posterior margin of the carapace to the rim of the
hole. Into the hole one or two eggs at a time are then dropped at
irregular intervals of a second or two's duration. With the aid
of a flashlight, which in no way disturbed the turtle, Cozzolino
was able to watch the process and counted a total of 163 eggs.
As soon as the full complement was laid the turtle quickly filled
in the smaller, then the larger, hole with sand and briskly re-
turned to the sea.

1 It seems io us the turtle's action may liuve been misinterpreted and that the
rluacal oritice was beint' moved about as urine was being discharged in order to
hind the sand in preparation for its removal.

480 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Cozzolino thinks that as many as 600 eggs may be laid by a
single turtle during one month, basing his belief on the fact that
more than 1000 eggs were present in the ovaries of the specimens
he dissected. He also thinks that a 9 , though remaining in the
vicinity where she first laid, never deposited subsequent batches
of eggs in the immediate vicinity of the first site. He measured
some of the spheroid eggs whose diameters ranged from 42-46
mm.,^ their weights from 45-65 grams. For the first 48 hours
after deposition the chalky white shells remained soft so that an
egg might be dropped from a height of three feet or so without
breaking. Under normal temperature conditions dehydration
sets in after 48 hours, the shell shrivels and becomes brittle
and the shape of the egg changes.

Incubation takes from 30 to 34 days, at which time the
hatchlings struggle to the surface, sometimes taking an oblique
course from the nest. This Cozzolino discovered after surround-
ing a nest-site with fishing net at a distance of 20 inches (50 cm.)
and the turtlets appeared outside the net, usually on the seaward
side. Thereafter he set his nets at a distance of 271/2 inches (70
cm.) and captured about 153 hatchlings from each of three nests.

Hatching takes place at night and the young turtlets instinc-
tively orient themselves and make for the sea. Many never
reach it, however, for in crossing the beaches they are attacked
by famished crustaceans. Survivors of the massacre which do
reach the ocean are then preyed upon by other predators.

Longevity. Fifteen years for a Pacific Green Turtle in New
York Aquarium, but only six years for an Atlantic specimen
(Flower :1925b).

Enemies. On Aldabra Island hatchling turtles are preyed
upon by Gray Herons {Ardea cinerea cinerea) and Frigate Birds
{Fregata minor aldabrensis) ; sharks swallow young turtles
entire and attack larger ones by biting off their flippers. Natives
harpooned the adults when asleep or sunning in lagoons or along
the shallower stretches of the coastline. About 3000 were
annually taken at the time of Voeltzkow's visit — 1903-1905 (cf .
Boettger:1913).

As only about 500 of these could be sent to Mahe, owing to
the scarcity of shipping, the remainder were processed locally.
The flesh, cut into strips and salted, was sun-dried for export to

1 85 mm. for a Liberian egg measured by Biittikofer (1890).

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 481

the Seychelles where workers favored it as an article of diet.
The fat was cooked for shipment to France where it enjoyed
a reputation as a remedy for chest diseases.

On Juan de Nova Island the natives have a curious way of
dealing with the discarded portions of the turtles they kill. On
the sloping dunes one sees a great many scaffoldings a yard or
more in height, strengthened by cross sticks and the whole
securely bound together by fiber. Behind the scaffolding a small
bank is constructed. In front of the scaffolding an area of from
one to two meters square is marked off and covered with the
twigs of Pemphis ocidula. Upon these twigs are laid the turtle
plastra; the carapaces, each overlapping the one in front, are
arranged in long rows to the right and left while from the scaf-
folding hang the heads of the turtles. No such scaffolding was
seen on Madagascar where the Sakalava Fody merely impale the
discarded heads on stakes.

On Ascension Id., according to D. Krlimmel (Werner :1912b),
Green Turtles are captured and held in lagoons connected with
the ocean. By May, when the season ends, there may be several
hundred. The meat appears twice weekly in the Commandant's
mess, and one or more turtles are presented to each visiting
warship.

The use of sucker fish {Bemora remora) to capture turtles
has long been practiced on the coast of East Africa from
Mombasa to Natal. The earliest references appear to be those
of Andrew Sparrman who, on his return from Natal, briefly de-
scribed (1787)^ how the natives, having attached one cord to
the fish's head and another around the tail, released the remora
in the sea in the vicinity of a turtle. As soon as the fish attached
itself to a turtle, both fish and reptile were drawn to the boat
where the reptile could be seized or harpooned. Sparrman wrote
that turtles were hunted in similar fashion on the coasts of
iladagascar.

During a voyage from the Cape to Ethiopia, Henry Salt
stopped at Mesuril (as Masuril), a village in Mozambique har-
bour. There, on September 9, 1809, he was presented with a
large example of Echeneis naucrates which the Bishop of Mesuril

1 For all Remora references see the extensive bibliography by E. W. Gudger,
1919, American Nat., 53, pp. 520-525.

482 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

informed him would attach itself to the plastron of a turtle with
such tenacity that the reptile rarelj- got away.

In 1829 Philibert Commerson's more detailed description of
the method employed at Mozambique was published by Lacepede.
Commerson states that a ring, small enough not to slip over the
caudal fin, is placed about the tail of a remora; to the ring is
attached a long cord. After this preparation the fish is placed
in a receptacle, containing salt water, in the bottom of the
fisherman's boat. Sail is set and the craft headed towards an
area where turtles are likely to be basking on the surface.
Aroused by the approaching boat, however, the lightly sleeping
reptiles seek safety in diving.

It is at this juncture that the remora is released, and sufficient
cord paid out to enable the fish to reach the turtle. Commerson
states that it is only after the fish has made futile attempts to
escape that it seeks shelter beneath the turtle to whose plastron
it adheres by means of a sucker on its head and nape, thus
enabling its employers to pull the turtle to the boat.

When returning from Pemba Island to Zanzibar, Frederic
Ilolmwood^ (1884) observed several fish dart out from beneath
the steam launch, on which he was travelling, whenever garbage
was thrown overboard. These chazo, as the boatmen called the
fish, after feeding, returned and attached themselves to the
bottom and sides of the launch. All Holmwood's efforts to pull
one off by force failed, but a Zanzibari readily detached it by
drawing the fish sideways. Later, Holmwood discovered that
the local fishermen were accustomed to employ these remora,
ranging in length from 2 to 4l^ feet and in weight from 2 to 8
lbs., in catching turtles. He was informed that sharks and even
large crocodiles were captured by this means in Malagasy waters.
Between trips, the remora are kept in canoe-like dug-out logs
stored in the huts of the fishermen who periodically change the
water.

The rings worn by the fish were attached in various ways,
though generally welded to a simple iron band. In some in-
stances the band had been worn for so many .years that it had
become imbedded in a thick fleshy formation. One chazo had an
extensive wound where the ring had been torn off. This fish.

1 Holmwood, 1SS4, '•Qn the Eniployment of the Remora by Native Fishermen
on the East Coast of Africa." Proc. Zool. Soc. London, pp. 411-413, figs. 1-2.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 483

said its owner, had caught the two turtles which were lying in
his canoe and then affixed itself to a shark. It continued to hold
on until all the spare line had been paid out and the ring
torn off. The injureci fish had then left the shark and returned
to the boat, a by no means unusual occurrence so Holmwood
was told.

Photographs of turtle hunting by means of remora are shown
in the film "West of Zanzibar" (1954). However, the technique
is not confined to East Africa, being practiced by fishermen in
Chinese and Australian waters as well as in the West Indies.

In the belief that citrus leaves mitigate the pungent odor of
boiling turtle flesh, it is customary in the Gold Coast to place
leaves and meat together in the cooking pot. The eggs too are
considered a great delicacy by the Gold Coast people. After first
boiling them in sea-water the shells are perforated to drain off
their watery contents, leaving the yolks. These are then baked
and eaten, the taste being compared to rich cake (Irvine :1947).

Man is unquestionably the w^orst enemy of the Green Turtle,
famous for soup though its carapace is of no commercial value.
With natives everywhere hunting these turtles for their flesh
and fat, besides searching for their eggs which are also highly
esteemed as food, the extermination of this reptile would appear
to be only a matter of time. Further information regarding
their exploitation in other parts of the world wall be found in
Gadow (1901:382), Deraniyagala (1939:320) and Carr (1952:
353-357).

Temperament. The placidity with which a Green Turtle
ignores inspection, making no attempt to bite even when touched,
is commented on by Voeltzkow (in Boettger :1913).

Hahits. Two captive Mozambique Green Turtles emitted
croaking ("qakenden") sounds, says Peters (1882a).

Habitat. Green Turtles frequent the submarine "prairies a
Posidonies ou Cymodocees" off the coast of Senegal, and where
these occur near the shore, as at Joal, the proportion of mydas
taken is considerable. In such areas they are captured at all
seasons, though no systematic hunting takes place, being either
harpooned or taken when they venture ashore during the egg-
laving season (Cadenat:1949 :22).

484 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Localities. Egypt: Port Said. Somalia: Mogadiscio (Mo-
gadish). Socotra Islayid: Abd el Kuri Id.; Shadwan Id.
Kenya Colony: *Lamu Id; *Mombasa Id. (B.M.). Tanganyika
Territory: *Dar es Salaam; Tauga. Zanzibar Island. Mozam-
bique: Europa Id. ; Juan de Nova Id. ; Mozambique Id.; Querim-
ba Id. Cape Province: Bird Id.; Cape of Good Hope; East
London ; Kei River mouth ; Kleinemonde. Ascension Island.
Angola: off Loanda. Cabinda: Cliinchoxo. Belgian Congo:
Banana at mouth of Congo River. French Equatorial Africa:
Loango Coast. 8do Thome Island. Principe Island. Bolas Island.
Gold Coast. Liberia: Monrovia; Robertsport. Portuguese
Guinea. Senegal: Banc d'Arguin (Argain) ; Hann ; Joal;
Mbour ; other localities of questionable authenticity are listed by
Rochebrune (1884a). Cape Verde Islands: Sao Vicente (Saint
Vincent) Id. Canary Islands: Teneriffa Id.

Range. All African coasts, Indian and Atlantic oceans and
— as an accidental A'isitor — the temperate seas.

Genus ErETMOCHELYS Pitzinger

1828. Caretta Ritgen (not of Kafinesque: 1814), Nova Acta Acad. Leop.-
Carol., 14, p. 270. Type: Testudo imhricata Linnaeus (by mono-

typy).

1843. Eretmochelys Fitzinger, Syst. Eept., p. 30. Type: T. imiricata

Linnaeus (by original designation).
1868. Herpysmostes Gistel, Die Lurche Europa, p. 145. Type: T. im'bricata

Linnaeus (fide Mertens, 1936, Senckenbergiana, 18, p. 75).

1873J. Onycliocliehjs Gray, Proc. Zool. Soc. London, p. 397, figs. 1-2. Type:

0. Tcraussi Gray ^ T. im'bricata Linnaeus (by monotypy).

Definition. Head with 2 pairs of prefrontal shields ; postocu-
lars 3 ; claws 2.

Skull wdth triturating surface of maxilla ridged, the ridge
strongest anteriorly, extending somewhat on to the premaxillae ;
premaxillae in contact with vomer which separates the maxillae ;
maxillae without vertical ribbing on inner surface of its cutting
edge ; descending processes of prefrontals in contact with vomer
only ; a blunt and rather low ridge on vomer and palatines at
the anterior margin of the internal ehoanae ; ehoanae in ventral
view not concealed by the moderate secondary palate ; pterygoids
deeply concave posteriorly; frontal entering orbit; crista prae-

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 485

temporalis strong ; mandibular symphysis pointed, concave, as
long as broad ; labial margin sharp, straight, not rising to a
point at the symphysis ; lingual margin lower than the labial,
without ridge; no symphysial ridge.

Carapace without lateral fontanelles in fully adult indi-
viduals; neurals 9-11, usually hexagonal, short-sided in front;
peripherals 11 pairs, the ninth pair not in contact with the ribs;
nuchal shield not in contact with first costals; costals tj^pically
4 pairs.

Plastron with a large intergular shield ; inf ramarginals 4
pairs, without pores.

Range. Tropical, subtropical and — as an occasional visitor —
temperate oceans.

Eretmochelys imbricata (Linnaeus)

1766. Tcstudo imbricata Linnaeus, Syst. Nat., ed. 12, 1. p. 350: American

and Asiatic Seas.
1S.34. Chelonia pseudo-mydas Lesson, in Belanger, Voy. Indies-Orient.,

Zool., p. 299: Atlantic Ocean.

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

Zool., p. 302 : Atlantic Ocean.

1835. Caretta Bissa Riippell, Neue Wirbelth. Fauna Abyss., Amphib., p. 4,

pi. ii: Red Sea.
1922a. Mertens, 168 (lists type, as syn. of imbricata).

1835. Chelonia imbricata Temminek and Schlegel, p. 13, pi. v.
1872. Sowerby and Lear, pis. Ivii-lviii.

1875. Melliss, 99.

1882a. Peters, 17.

1890. Buttikofer, 2. 438, 478.

1896a. Boeage, 98.

1906a. Mocquard, 481.

1909a. Siebenrock, 547.

1913c. Nieden, 55.

1921d. Loveridge, 51.

1849. Caretta imbricata A. Smith, App., 2.
1873b. Gray, 92.

1884a. Rochebrune, 39.

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

Indian and Pacific Oceans.

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

442, pi. XXX, figs. 1-7 : Sulu Seas and Indian Ocean.

486

BULLETIN : MUSEUM OF COMPAKATIVE ZOOLOGY

1858. Caretta rostrata Girard, IT. S. Explor. Exped. 1838 1842, Herp., p.

446, pi. XXX, figs. 8-13 : Fiji Islands.

]87aj. Onychochclys Jcraussi Gray, Proe. Zool. Soe. London, p. 398, .figs.

1-2 : Atlantic Ocean off French Guiana.

1885. Chelone imhricata Greef (1884), 49, footnote.

1889a. Boulenger, 183.

1893a. Boettger, 12.

1896. Tornier, 3.

1897. Tornier, 63.

1898. Sclater, W. L., 97.
1898. Tornier, 283.
1900b. Tornier, 582.
1901. Gadow, 385.
1901. Steindachner, 32(i.
1903a. Boulenger, 92, 96.
19061. Boulenger, 197.
1906. Johnston, 819, 833.
1910. Sternfeld, 5, fig. 10.
1911(1. Sternfeld, 51.

1914. Fuchs, 1914, pp. 1-325, figs. 1-182, pis. i-vi.

]923g. Loveridge, 933.

1924b. Loveridge, 3 (as imprieata)

1925b. Flower, 932.

1929. Eose, 184,

1930a. Scortecci, 215.

1931. Ingrams, 429.

1937 b. Angel, 1696,

1955. Cansdale, 96, 104.

1929. Eretmochelys imhricata Flower, 39.

1929. Lindholm, 287.

1933. Flower, 750.

1933h. Loveridge, 207.

1937a. Flower, 13.

1937e. Hewitt, 16, pi, ivB.

1937f. Loveridge, 488.

1947. Irvine, 311,

1949. Cadenat, 16.

1949. Villiers, 165.

1950. Rose, 327.
1950. Williams, 552.

1952. Williams and McDowell, pi. i, fig. 4.

1934a. Eretmochelys imhricata imhricata Mortens and Miiller in Rust, 10.

1934a. Eretmochelys imhricxita squamosa Mertens and Miiller in Rust, 10.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 487

Common names. Ilawksbill Turtle (English); anyite (Ewe-.
Irvine); apuhuru (Fante :lrvine) ; ayikplonto (Ga:Irvine) ;
baga for 9, hissa for $ (Arabic in Red Sea :Riippell) ; hala-
apatadzi (Ga:Irvine, q. v. for other Gold Coast names) ; ingappa
(Mozambique :Peters) ; ngamba (Cape Delgado Island :Peters;
Zanzibar :1 ngrams ) .

Illustrations. A reasonably good colored plate will be found
in Riippell (1835), and very fine ones of a subadult in Sowerby
and Lear (1872b :pls. Ivii-lviii).

Description. (Based exclusivelj^ on six African and Malagasy
specimens in the Museum of Comparative Zoology).

Snout elongate, compressed ; beak drawn out but not hooked,
without cusps ; edge of jaws apparently smooth ; prefrontals not
elongate, in 2 pairs; frontal azygous, small, rarely^ fused with
frontoparietal ; frontoparietal large, entire or semidivided anter-
iorly; parietals 2, transverse in juveniles, somewhat elongate in
adults; a small interparietal sometimes present; supraocular
rather large; postoculars 8, rareh^- 4; supratemporals 2 or 3,
one sometimes fused with a tempoi'al ; f orelimb with moderately
enlarged scales along anterior edge, behind them several rows
of scales, posterior edge with a series of enlarged scales; both
fore- and hind limbs with 2 claws; tail short.

Carapace ovate, subtectiform in adults, with 3 interrupted
keels in young but only a trace of the lateral ones persisting in
adults, nuchal region truncate, margin weakly serrate in young,
strongly in adults ; dorsal shields juxtaposed in hatchling, im-
bricate in adults, juxtaposed in aged individuals; nuchal broader
than long, not in contact with first costals, not in contact with
second marginals; vertebrals 5, exposed portion of each broader
than long at all ages, the first no broader than the others in
young, broader than the others in adults ; costals 4, fourth small-
est ; marginals 11 pairs ; supracaudal divided.

Plastron anteriorly rounded, 2 prominent lateral keels; plas-
tral shields of adult more or less imbricate ; intergular moderate ;
brachials 2 or 3 with some smaller scales extending anteriorly ;
inframarginals 4, without pores; inguinal small; interanal
minute or absent.

1 Fused In a Zanzibar spccinipu (M.<'.Z. 1141) and a Malagas.v tiirtU- (M.C.Z.
1G867).

2 Four in a Zanzibar spcciinon (M.C.Z. 1141).

488 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Color. Carapace of young light brown flecked with darker, or
black blotched with lighter ; plastron substantially similar. Head
and limbs light brown to black, more or less uniform.

Carapace of adult horn to reddish brown heavily marbled or
streaked with black; plastron yellow, the posterior portion of
each shield blotched with black. Head and limbs dark above, but
on the sides and below yellow heavily blotched with black.

The foregoing color description is based solely on our African
and Malagasy material.

Size. Carapace length of largest in the Paris Museum 740
mm.; surpassed by one of Riippell's "hissa" which was 765 mm.,
by 690 mm. broad ; the lectotype was only 750 mm., by 635 mm.
broad; the record is apparently 850 mm. (B.M.).

Breeding. On Rolas and Sao Thome Island the 9 2 come
ashore to lay from December to February (Greef :1885). In the
Red Sea region they lay from February to March ; after laying,
the 5 5 regularly return to the site, according to Riippell
(1835).

Longevity. Fifteen years, 7 months, 28 days, and still alive
about 1937 (Flower :1937a). The alleged record of 32 years,
quoted by Gadow (1901:386), has been traced to Tennant
(1861:294), who quoted Bennett (1843:275) who followed it
back to 1826 and 1794 (Flower :1925b, where full references to
preceding citations will be found).

Enemies. In Liberia it is eaten by the Africans but not by
Europeans (Johnston :1906). The plates of the $ $ are thinner
than those of the $ 2 , and consequently are of less value com-
mercially (Riippell :1835). Between 700 and 800 kilograms of
high grade tortoise-shell was annually taken from the Hawks-
bill Turtles in the vicinity of Cape Delgrado (Peters :1882a). As
much as 8 lbs. of shell may be removed from a single large speci-
men (Gadow :1901). Great numbers of stripped carapaces were
observed around the natives' huts on Abd el Kuri Island by
H. 0. Forbes, who remarks that though it still occurs off the
coast of Socotra Island it is not nearly so common as it was
eighteen centuries ago when it was an article of trade much
sought after by merchants from Arabia (Boulenger :1903a).

Temperament. In marked contrast to the Green Turtle, the
Hawlvsbill bites so freelv that ]^lalagasv fishermen carrv in their

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 489

boats a piece of wood Avhich they hold out to a harpooned turtle.
Once the reptile has seized the proffered wood it does not readily
let go. If this procedure is not followed, so the men told Voeltz-
kow {in Boettger :1913), the turtle is apt to bite on their frail
craft when being hauled aboard, and may inflict considerable
damage.

Localities. Egypt: Saint John Id. ; Red Sea coasts from Zebejir
to Zukur. Eritrea: Massaua. Socotra Island: Abd el Kuri Id.;
Socotra Id. Kenya Colony: Frere Town; *Mombasa Id. Tangan-
yika Territory : Dar es Salaam ; Lindi ; Tanga. Zanzibar Island :
*Zanzibar. Mozamhique: Cape Delgado; Mozambique Id.; Que-
rimba Id. near Ibo. Cape Province: False Bay; Simonstown
Beach; Table Bay. Ascension Island: *(M.C.Z. 4095). French
Equatorial Africa: Gabon Coast (P.M.). Sao Thome Island. Prin-
cipe Island: Holas Id. (only). French Cameroon: Longuy (Long-
ji). Gold Coast: *Tenia (B.M.). Liberia: xingel River. Sierra
Leone: *Bonthe, Bonthe district. Senegal: Banc d'Arguin (Ar-
gain) ; Joal (Joalles) ; other localities of questionable authen-
ticity are listed by Rochebrune (1884a). Cape Verde Islands:
Fogo Id.

Range. East, South and West coasts of xVfrica, Indian and
Atlantic oceans and — as an accidental visitor — the temperate
seas.

Genus Caeetta Rafinesque

1814. Caretta Eafinesque, Specehio Sei. (Palermo), 2, No. 9, p. 6G. Type:

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

Tj'pe: T. caouana Fitzinger = T. caretta Linnaeus (designation

by Fitzinger, 1843).
1838. Caouana Cocteau, Kept, in De la Sagra, Hist. Fis. Pol. Nat. Cuba, 4,

p. 31. Type: T. cephalo Sclmeider = T. caretta Linnaeus (by

monotypy).
1843. Halichelys Fitzinger, Syst. Rept., p. 30. Type: E. atra Fitzinger =

T. caretta Linnaeus (by original designation).
1873J. Cephalochelys Gray, Proc. Zool. Soc. London, p. 408. Type: C.

oceanica Gray = T. caretta Linnaeus (by monotypy).
1873J. Eremonia Gray, Proc. Zool. Soc. London, p. 408. Type: E. elongata

Gray = T. caretta Linnaeus (by monotypy).

490 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Definition. Head with 2 pairs of prefrontal shields; postocu-
lars 3 ; claws 2.

Skull with triturating surface of maxilla not ridged; premax-
illae not reached by vomer which does not separate the maxillae ;
maxilla without vertical ribbing or a sharp cutting edge, its
whole surface sloping dorsomedially as part of a large crushing
surface; descending processes of prefrontals in contact with
vomer and palatines; a blunt ridge on vomer and palatines at
the anterior margin of the internal choanae ; choanae in ventral
view concealed by the extensive secondary palate; pterygoids
deeply concave posteriorly; frontal excluded from orbit; crista
praetemporalis strong ; mandibular symphysis rounded, concave,
longer than broad; neither labial nor margins raised as cutting
edges; no symphysial ridge.

Carapace without lateral fontanelles in adults; neurals 7-11.
occasionally some separated, typically hexagonal, short-sided in
front; peripherals 12-13 pairs, the ninth or tenth pair not in
contact with the ribs ; nuchal usually in contact witli first costals :
costals typically 5 pairs.

Plastron without, or with only a very small, intergular shield;
inframarginals 3 pairs, without pores.

Range. Tropical, subtropical and — as an accidental visitor —
temperate oceans.

Caretta caretta (Linnaeus)

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

America.
1783. Testudo Cephalo Schneider, Naturg. Sohildkroten, p. 303: No locality.
1788. La Caouane Lacep^de, Hist. Nat. Quad. ovip. Serpens, I- p. 95; and

Testudo caouana in Synopsis Methodiea, a table (in which bi

nomials are used) at end of same volume: Jamaica, British West

Indies (restricted).
1788. La Tortue nasicorne Lacepede, Hist. Nat. Quad. ovip. Serpens, 1,

p. 103; and Testudo nasicornis in Synopsis Methodiea, a table (in

which binomials are used) at end of same volume: America.
ISU. Caretta imsuta Rafinesque, Specchio Sci. (Palermo), 2, No. 9, p. (ifi:

Sicily (/ideLindholm: 1929).
1820. Caretta atra Merrem, Vers. Syst. Amphib., p. 17: Ascension Island.
1833. Chelonia pdasgorum Valenciennes, pi. vi, in Bory de Saint Vincent,

Exped. sci. de Moree, Zool.: Modon, etc., Mediterranean Sea.

LOVERIDGE AND WILLIAMS: AFRICAN CBYPTODIKA 49.1

1844. Caouana elongata Gray, Cat. Tortoises Brit. Mus., p. 5.3: No locality.
18o8. Thalassochelys corticata Girard, U. S. Explor. Exped. 1838 184"J.

TTerp., p. 431, pi. xxix : Madeira Islands.
1894. Oliver, 100.

18961). Oliver, 118.

1862a. Chelonia corticata Strauch, 19.
1903. Mayet, 9.

1873J. Cephalochelys occanica Gray, Proc. Zool. Soc. London, p. 408:

? Mexico.
1884a. Caouana caretta Rochebrunc, 38.
1887. Thalassioclielys tarapacona {sic) Philippi, Zool. Garten, 28, p. 85:

Iquique, Chile (identified by subsequent description. See Philippi,

1899).
1889a. Thalassochelys rarrita Boulenger (part), 184.
1891c. Steindaehner, 305, 306.

1893a. Boettger (part), 13.

1895a. Boeage, 6.

1896a. Boeage, 66.

1898. Jeude, 11.

1S9S. Sclater, W. L., 97.

1901. Gadow (part), 320, 387, figs. 69, 84 (diagrams based on a

misconception involving young of one species, adults of an-
other).

1902. Boeage, 208.
1903a. Boulenger, 96.
1906a. Mocquard, 481.
1912a. Pellcgrin, 256.
1923g. Loveridge, 933.
1924b. Loveridge, 3.
1925. Flower, 932.
1926c. Pellegrin, 49.
1929. Rose, 184, fig. 122.
1932d. Witte, 57.

1899. Thalassochelys tarapacana Philippi, Anales Univ. Chile (Santiago),

No. 104, p. 731 (validating redescxiption).

1899. Chelonia Caouana Doumergue, 253.

19n9a. Caretta caretta Siebenroclc, 549.
191 Id. Sternfeld, 52, fig. 64.

1912b. Werner, 465.

1929. Flower, 39.

1929. Lindholm, 287.

1933. Flower, 751.

1934. Mosauer, 52.

492 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1937b. Angel, 1696.

1937a. Flower, 14, 37.

1937e. Hewitt, 15, pi. ivB.

1949. Cadenat, 17, figs. 4, 5, 7-12, 15b, 16a.

1949. Villiers, 165.

1950. Rose, 327, fig. 127.
1950. Williams, 552.

1930. Caretta [caretta inferred] Babcock (part), 95.

1933. Caretta gigas Deraniyagala, Ceylon Jour. Sci., Sect. B, 18, p. 66,

figs. 4-6, pi. v: Gulf of Mannar, Ceylon.
1934a. Caretta caretta caretta Mertens and Miiller in Rust, 10.
1935. Tortue caret Witte, 94.
1939. Caretta caretta gigas Deraniyagala, 162.

Common names. Red-brown Loggerhead (English) ; fahroiiyi
el harr (in Tunisia .-Mosauer) ; nduvi (at Lindi :Loveridge MS).

Description} Snout relatively short, not compressed; beak
distinctly hooked, unicuspid ; edge of jaws apparently smooth ;
prefrontals not elongate, in 2 pairs with an additional central
pair ; frontal azygous, small, longer than broad ; frontoparietal
large, as broad as long, semidivided posteriorly; parietals not
elongate, broken up ; supraocular moderately large ; postoculars
3 ; supratemporals in 3 pairs ; f orelimb with moderately enlarged
scales along anterior edge, behind them numerous small scales,
posterior edge with a series of distinctly enlarged scales; both
fore- and hind limbs with 2 claws; tail moderately short.

Carapace ovate, tectiform, with a strong interrupted vertebral
keel and nearly obsolescent lateral keels (in the young Madeira
specimen) ; nuchal region truncate, margin somewhat serrate ;
dorsal shields juxtaposed; nuchal broader than long, in contact
with first costals on right side, excluded on left; vertebrals 6,
first smallest ;- costals 5, first smallest ; marginals 13 pairs ; su-
pracaudal divided.

Plastron anteriorly rounded, 4 strong, obtuse, interrupted, lat-
eral keels ; plastral shields juxtaposed ; intergular present, small ;
brachials 3 with some smaller scales extending anteriorly ; infra-
marginals 3, without pores ; inguinal absent ; interanal absent.

Color. Carapace reddish brown,^ points of keels and marginals

'• Based on the almost foot-long, Madeiran type of T. rorticatn Girard
(U.S.N.xM. 7778) examined by E. E. W.

2 The normal first vertebral in the type of corticata (U.S.N.M. 7778) is asym-
metrically divided.

3 In the now dried type of corticata more uniformly reddish than described by
Girard.

LOVERIDGE AND WILLIAMS : AFRICAN" CRYPTODIRA 493

lighter; plastron whitish yellow to yellowish brown. Head scales
reddish centrally, narrowl}' margined with yellow; eyelids con-
spicuously black {fide Doumergue) ; neck yellowish; throat yel-
lowish brown ; limbs a})Ove, reddish brown, below, yellowish
brown.

Size and Weight. In the almost complete absence of African
material we offer the following remarks.

The carapace of a Florida loggerhead at hatching had a length
of about 48 mm., a breadth of 35 mm., and a Aveight of about 20
grams. When four-and-a-half years old this turtle weighed 811^
lbs. (37 kilos.) and its carapace measured about 2 feet (630 mm.)
in length, and 1 foot, 11 inches (590 mm.) in breadth. Its rate
of growth was much more rapid than that of three other logger-
heads reared under almost similar conditions. Yet all three re-
vealed that these turtles attain to maturity much more rapidly
than was generally supposed (Parker :1939).

Carapace length of a hatchling (M.C.Z. 4017) from Key West.
Florida, 49 mm., breadth 41 mm., length of forelimb from axilla
to tip, 35 mm. ; the carapace length being contained in that of
the forelimb .71 per cent.

Carapace length of a young turtle (M.C.Z. 28202) from Ber-
muda, 83 mm., breadth 72 mm., length of forelimb from axilla
to tip 65 mm. ; the carapace length being contained in tliat of the
forelimb .79 per cent.

Carapaces rarely exceed 700 mm. in length at Oran (fide
Doumergue :1899). Carapac.e lengths of adults from Hann and
Joal, Senegal, were 1040 and 1080 mm., respectively (Cadenat:
1949:19).

Carapace length over the curve of the largest known " gigas"
$, from China {fide Fang:1934 in Deraniyagala : 1939), 940
mm.; largest ''gigas" ? from Moratuva, Ceylon 960 mm. {fide
Deraniyagala :1939:183).

Longevity. Possibly 33 years in the Vasco da Gama Aquarium
in Lisbon, where several lived from 1898 until 1931, when they
succumbed during a spell of exceptionally hot weather {fide Dr.
Rodrigo Boto, who so informed Flower :1937a). Another logger-
head, whose age when received at the Berlin Zoological Gardens
in September, 1913, Avas estimated at 2 years, was still alive on
September 19, 1936. Consequently the animal was at least 23,
possibly 25, years old.

494 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Diet. Ill spring, loggerheads appear within the narrow sand-
bar on the west coast of Lanzarote Island in order to feed on the
abundant mussels (Gari depressa, recorded as Psammol)ia ves-
pertina, and Venus verrucosa) which occur there ( Steindachner :
1891c).

Enemies. The flesh of the caouanne is good to eat, according to
Doiimergue (1899).

Localities. French Morocco: Mogador. Algeria: Oran. Tun-
isia: Bizerte; Gabes; Sfax market; Tunis. Egypt: Alexandria;
BruUos; Damietta; Port Said. Socotra Island: Abd el Kuri Id.;
Bandar Saleh. Kenya Colony (Loveridge :1923g). Mozambique:
*Inhambane (U.S.N.M.). Cape Province: Kalk Bay; Muizen-
berg; Peddie Coast. Southwest Africa. Ascension Island. Bel-
gian Congo: Banana. French Equatorial Africa: Gabon. Ivory
Coast: San Pedro. Senegal: Fadiouth; Hann; Joal. Bio de Oro
{fide Carr :1952 ;383, but Cameroous may be included on basis
of Lepidochelys, which see). Cape Verde Islands: Sal Id. near
Sao Vicente Id. Canary Islands: Lanzarote Id.; Teneriffa Id.
Madeira Islands: Madeira Id.

Range. All African coasts, Indian and Atlantic oceans, the
Mediterranean and — as an accidental visitor — the temperate
seas.

Genus LePIDOCHELYS Fitzinger

1843. Lepidodhelys Fitzinger, Syst. Eept,, p. 30. Type: Chelonia olivacea

Eschscholtz (by original designation).
1880. Colpochelys Garman, Bull. Mus. Comp. Zool., 6. p. 124. Type:
kempi Ganiian (by monotypy).

Definition. Head with 2 pairs of prefrontal shields; postocu-
lars 3-4 ; claws 2-3 (though adults sometimes have only 1).

Skull with triturating surface of maxilla ridged, the ridge sub-
equal in height throughout, not extending on to the premaxillae ;
premaxillae in contact with vomer which separates the maxillae ;
maxilla without vertical ribbing on inner surface of its cutting
edge ; descending processes of prefrontals in contact with vomer
only; no obvious ridge on vomer and palatines at the anterior
margin of the internal choanae; ehoanae in ventral view not con-
cealed by the moderate secondary palate ; pterygoids deeply con-
cave posteriorly; frontal entering orbit; crista praeteraporalis

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIBA 495

strong; mandibular sj-mphysis pointed, about as long as broad;
labial and lingual margins slightly raised into cutting edges; a
more or less distinct symphysial ridge.

Carapace without lateral fontanelles iu adults; neurals 12-15,
some quadrate, others hexagonal, short-sided in front; peri-
pherals usually 12-13 pairs, the tenth pair not in contact with the
ribs; nuchal shield usually in contact with first costals; costals
,) or more pairs.

Plastron without, or with only a very small, intergular shield ;
inframarginals 4 pairs, each with a pore near its posterior edge.

Range. Tropical, subtropical and — as an accidental visitor —
temperate oceans.

LePIDOCHELYS OLIVACEA ULIVACEA ( EscllScholtz )

1820. IChelonia multiscutata Kuhl, Boitr. Zool. Anat., p. 78: No locality.
1829. Cheloniu oUvacea Eschst-holtz, Zool. Atlas, p. 3, pi. iii: Manila Bav.

Philippine Islands, China Sea.
IStiO. Dumeril, 170.

1835. Chclonia Dussumierii Dumeril and Bibron, Erpet. (i('-n., 2, ]>. 5~^7. pi.

xxiv, figs. 1-la: China Sea and Malabar Coa.st.
1835. Caretta olivacea Riippell, 7, pi. iii.
1839. Parker, 129.

1949. Cadenat, 17, 23, figs. 6, 16b.

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

? India (nomen nudum).
1849. Caouana dessumierii (sie) A. Smith, App., 2.
1857. Cheloniu polyaspis Bleeker, Nat. Ti.idshr. Ned. Indies, 14, j). 239:

Batavia, Java.
1884a. Caouana olivacea Kochebrune, 38.
1888a. Thalassoclielys olivacea Boettger, 18.
1889a. Chelonia dubia Bleeker (nomen nudum) in Boulcuger, Cat. Chclon.

Pihyncho. Croeod. Brit. Mus., p. 186: Borneo.
1899. TJuilassochelys controversa Philippi, Anales Univ. Chile (Santiago),

No. 104, p. 732: No locality.
1900b. Thalassoclielys caretta Tomier (not of Linnaeus), 582.
1902c. Tornier, 665 (head lost, but hatchlings studied by us).

1910. Sternfeld, 5 (based on the 1902c material just mentioned).

1913c. Nieden, 55,

1930a. Scortecci, 215.

1908. Caretta remivaga Hay, Proc. U. S. Nat. Mus., 34. p. 194, pi. x, figs.

1-3, pi. xi, fig. 5: Ventosa Bay, Gulf of Tehuantepec, Mexico.

496 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1934a. Caretta caretta oUvacea Mertens and Miiller in Eust, 10.
1937f. Loveridge, 488.

1943. Lepidochelys olivaoea olivacea Deraniyagala, 81, 92, figs. 1, 3a,
5a, 6a.

Common names. Olive Loggerhead (English) ; hage (in Erit-
rea :Riippell).

Illustrations. A clear and detailed drawdng of the dorsal view,
together with a lateral view of the head and plastron from be-
low, is furnished by Riippell (1835).

Description.^ Snout relatively short, not compressed; beak
scarcely or slightly hooked, not or but slightly bicuspid (tricus-
pid in Riippell 's figure) ; edge of jaws apparently smooth; pre-
frontals not elongate, in 2 pairs f frontal azygous, small, as broad
as, or broader than, long (in hatchlings), longer than broad (in
adult) ; frontoparietal large, broader than long, semi- or wholly
divided ; parietals transverse (in hatchlings) , elongate (in adult) ,
or somewhat broken up (in either) ; supraocular moderately
large ; postoculars 3-4 ; supratemporals broken up ; f orelimb with
moderately enlarged scales along its anterior edge, behind them
several rows of scales, posterior edge with a series of distinctly
enlarged scales; both fore- and hind limbs with 2 claws, though
on the former scarcely distinguishable; tail short.

Carapace ovate, hardly tectiform. with a weak, interrupted,
vertebral keel, in hatchlings similar lateral keels are also present,
nuchal region truncate, margin scarcely serrate; dorsal shields
juxtaposed ; nuchal broader than long, normally ^ in contact with
the first costals, not in contact with second marginals ; vertebrals
5-7, in hatchlings all broader than long except the last M'hich is
usually longer than broad, in adults I is somewhat broader than
long, II-VI as long as, or longer than, broad, VII much the
broadest; costals^ 6 or 7,^ first smallest; marginals 12 pairs;

1 Based on five hatchlings from the Cameroon (Berlin Mus. 15513). three
hatchlings from the Gabon (Paris Mus. 41.58) and Riippell's (1835) figures and
description of a Massaua specimen. Checked on nine hatchlings from Ceylon ;
India ; New Britain and Mexico, but their deviations are not included in this
description.

2 The second pair of frontals are fused with the supraocular on one or both
sides in three of the five Cameroon hatchlings.

3 Nuchal separated from first costal by a marginal in one Cameroon hatchling.

4 Of the five Cameroon hatchlings two have 5 costals on both sides ; one has 5
on the right, 6 on the left : two have 6 on both sides. The series, obviously from
one nest, must be regarded as intergrades between L. o. olivacea (Eschscholtz)
and L. o. kempi (Garman). Incomplete supernumerary costals, split off from the
larger complete ones, occur in the Gabon hatchlings ; we have not included these
incomplete scutes in the costal count.

5 Seven In a Senegal specimen (Cadenat :1949).

LOVERIDQE AND WILLIAMS : AFRICAN CRYPTODIRA 497

siipracaudal divided.

Plastron anteriorly rounded, 2 obtuse, interrupted, lateral
keels; plastral shields juxtaposed; intergular small, rarel}^ di-
vided ^ or absent ; brachials 2-3 with some subequal or smaller
shields extending anteriorly; inframarginals 4, each with or
without a small pore posteriorly ; inguinals minute or absent ; in-
teranal minute or absent.

Color. Carapace of hatchlings (from Gabon) uniformly oliva-
ceous black, or (in Cameroon intermediates) wholly black; plas-
tron, head and limbs suffused with brown.

Carapace of adult (from Massaua, Riippell:1835) olive green;
plastron sulphur yellow. Head, neck and foot above, olive green ;
below sulphur yellow. Iris dark brown, ej^eball blue-gray.

Size. Carapace lengths and breadths of five hatchlings from
Cameroon (Berlin Mus.) range from 42 x 38 to 44 x 40 mm.,
those of three hatchlings from Gabon (Paris Mus.), 49 x 45,
50 X 44 and 50 x 45 mm., respectively. This will give some idea
of the variation in length to breadth ratio in hatchlings that
almost certainly came from only two broods. Riippell's Massaua
turtle was said to be 630 mm. in length as well as breadth. Cara-
pace lengths of specimens from Ngaparo and Joal, Senegal, were
660 and 690 mm. respectively (Cadenat :1949), while the possible
maximum length for an adult may be 790 mm. ; breadth 680 mm,
(fide Deraniyagala :1939) .

Breeding. In May on the Eritrean coast (Riippell:1835).

Localities. Egyptf Eritrea: Massaua. Socotra Island. Tan-
ganyika Territory: Lindi. Cape Province: Table Bay. Belgian
Congo: Banana; Moanda (Royal Museum of the Belgian Congo,
fide Carr in litt.). French Equatorial Africa: Gabon Coast,
British Cameroon: *Victoria (intermediates olivacea X kempi).
Gold Coast: *Tenia (B.M.). Ivory Coast: (Paris Mus.). Sene-
gal: Goree Id.; Guet N'Dar; Harm, Joal; Ngaparo.

Range. East, South and West coasts of Africa, Indian and
Pacific Oceans, also — as an accidental visitor — the temperate
seas.

From the Azores we have seen a young L. o. kempi (Monaco
Mus. 2660), taken by the Prince of Monaco in 1913, while hatch-

1 Divided on a Cameroon (Berlin Mus.) and on a Gold Coast hatchling (Brit.
Mus.) seen by us.

498 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

lings from the Cameroons (Z.M.U. 15513) must be considered
intermediates between the two races (see footnote to p. 497).

Family DERMOCHELYIDAB

1902. Dermochclyidae Wieland, Amer. Jour. Sci., (4), 14, p. 107.

Definition. Cryptodirous testudinates adapted to a marine
life. Horny shields present in young, replaced in adults by
smooth skin.

Skull without nasal bones ; prefrontals in contact dorsally,
with descending processes that are moderately separated ; tem-
poral region hardly emarginate posteriorly ; parietal meeting
squamosal; postorbital very large; no bones tending to be re-
d;..ced; quadrate not enclosing stapes; postotic antrum absent;
parietals without descending processes ; upper jaw without ridges
on its triturating surface; vomer present, separating palatines;
mandible without a coronoid bone.

Neck vertebrae with only one biconvex centrum — ■ the fourth,
typically an articular connection between the sixth and seventh
centra, the eighth centrum concave in front, doubly or not ; eora-
coids with their median borders not expanded ; humerus of an
exaggerated marine type with deltopeetoral crest very far down
the shaft ; trochanteric fossa of femur reduced by union of the
trochanters; phalanges without condyles; claws absent.

Carapace with epithecal component very strongly developed,
represented by a mosaic of small bones; thecal carapace and
plastron greatly reduced, not directly united; neither carapace
nor plastron hinged; pleural bones reduced to the endochondral
ribs, separated by wide f enestrae ; peripherals absent ; nuchal a
butterfly-shaped bone with a ventral attachment surface for the
eighth vertebra; neural bones absent, represented only by the
neural arches of the vertebrae ; pygals absent ; plastron reduced
to a narrow ring of bones surrounding a great f ontanelle ; ento-
plastron absent.

Range. Tropical, subtropical and — as an accidental visitor —
temperate oceans.

LOVERTDGE AND WILLIAMS : AFRICAN CRYPTODIRA 490

Genus DekMOCHELYS Blaiiiville

1816. Dermochelys Blainville.i Bull. Soc. Philom. Paris, p. "Ill" (mis
print = 119). Type: Testudo coriacea Linnaeus (by monotypy in
Cuvier: 1829).

1820. Sphargis Merrem, Vers. Syst. Amphib., p. 19. Type: T. coriacea
Linnaeus (by monotypy).

1822. Coriudo Fleming, Phil. Zool., 2. p. 271. Type: T. coriacea Linnaeus
(by monotypy).

1828. Seytina Wagler, Isis von Oken, p. 861. (Substitute name for Sphar-
gis Merrem.)

1830. Dermatochelys Wagler, Natur. Syst. Amphib., i^. 13.3. (Emendation
for Dermochelys.)

1832. Chelyra Rafinesque, Atlantic Jour., 1, p. 64. Tyjie: T. coriacea Lin
naeus (by monotypy).
Definition. Similar to that given under DERMOCHELY-

IDAE, the family being monotypic.
Range. Tropical, subtropical and — as an accidental visitor —

temperate oceans.

Dermochelys coriacea (Linnaeus)

1766. Testudo coriacea Linnaeus, Syst. Nat., ed. 12, 1, i). 350: Mediter-
ranean Sea.

1789. Poiet, 283.

1802b. Daudin, 62, pi. xviii, lig. 1.

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

1788. Le Luth Lacepedc, Hist. Nat. Quad. ovip. Serpens, 1. p. Ill; and
Testudo lyra in Synopsis Methodica, a table (in which binomials
are used) at end of same volume: Barbary Coast.

1792. Testudo tub.erculata Schoepff, Hist. Testud., p. 144: No locality .^

1814. Chelonia lutaria Eafinesque, Specchio Sci. (Palermo), 2. No. 9, p. 66:
Sicily (^(ie Lindholm: 1929).

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

1 In the same year Dermochelys was also proposed by Blaiuville in Jour. Phys.
(.'him. Hist. nat. (Paris), 83, p. 2.59. We cannot s:iy which publication had
priority, but it is the Bulletin that is currently cited by herpetolo^ists. In neither
paper Vas any species mentioned. However, coriatia Linnaeus was the only
included species assigned to Dermochelis (sic) by Cuvier (1S29, Regn6 Animal,
ed. 2. 2, p. 14) and this has been accepted as the type by the "Official List of
(generic Names in Zoology."

2 Attributed by Schoepff to Pennant, 1771, Philos. Trans. Royal Soc. London,
61, p. 271 ()iot 2"75), pi. X, figs. 4-5, but Pennant merely provides an English name
— Tuberculated Tortoise — to a specimen which he thinks may be Testudo
coriacea Linnaeus. In Schoepff's 1801 ed. tuberculata appears on p. 123.

500

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1835. Temniinck and Schlegel, 6, pis. i-iii.

1849. Smith, A., App., 2.

1829. Dcrmochelis Atlantioa Lesueur, in Cuvier, Regne Animal, 2. p. 14,

footnote: No locality (nomen nudum).

1830. Dermatochelys porcata Wagler, ISTatlir. Syst. Amphib., pi. i, figs.

1-28: No locality (uses corincea in text, p. 133).
1862a. Spliargh coriacea Straueh, 19.
1872. Gervais, p. 199, pis. v-ix.

1873b. Gray, 96.

1884a. Rochebrune, 40.

1894. Oliver, 101.

1896. Doumergue, 477.

1896b. Oliver, 118.

1899. Doumergue, 255.

1933. Heldt, 1-40, figs. 1-17.

1949. Cadenat, 16, fig. 13.

1844. Sphargis coriacea var. Schlegelii Garman, U.S. Nat. Mus. Bull., 25,

pp. 295, 303 : Tropical Pacific and Indian Oceans.
1889a. BermocTielys coriacea Boulenger, 10.
1890. Buttikofer, 1. 266 ; 2. 437, 478, fig.

1893c. Matschie, 208.

1899. Sclater, 96.

1901c. Tornier, 66,

1906a. Mocquard, 483.

1909a. Siebenrock, 553.

1911b. Stemfeld, 52,

1925b. Flower, 918.

1929. Lindholm, 287.

1929. Rose, 184.

1933. Flower, 752.

1937e. Hewitt, 16, pi, ivB.

1937f. Loveridge, 488.

1947. Irvine, 312.

1949. Villiers, 165, fig.

1950. Rose, 327.
1950. Williams, 552.
1955. Cansdale, 97, 104.

1899. Sphargis angusta Philippi, Anales Univ. Chile (Santiago), No. 104,

p. 730, pi. i: Tocopilla, Chile.
1901. Dermatochelys coriacea Gadow, 325, 333, fig. 73.
1906. Dermoehelis coriacea Johnston, 819, 833.

1934a. Dermochelys coriacea coriacea Mertens and Miiller in Rust, 10.
1934a, Dermochelys coriacea schlegeli Mertens and Miiller in Rust 10.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 501

Common names. Leatherback; Leathery Turtle; Luth or
Trunk Turtle (English); hosange (Ga:Irvine, whom see for
other Gold Coast names); 7ioa (Swahili at Lindi :Loveridge

MS).

Illusirations. Those of Temminek and Schlegel (1835 :pls.
i-iii) are quite good.

Bescription. (Based exclusively on a juvenile (M.C.Z. 21055)
from the Guinea Coast, West Africa.)

Snout moderate; beak deeply notched in middle, strikingly
bicuspid ; edge of jaws denticulate ; prefrontals in 2 pairs, of
which the posterior pair may be broken up ; frontal broken up ;
frontoparietal large; parietals, supraocular, postoculars, and
supratemporals broken up ; f orelimb with slightly enlarged scales
along anterior edge, behind them many rows of smaller scales,
posterior edge without noticeably enlarged scales ; both fore- and
hind limbs clawless; ta.il short.

Carapace a mosaic of small polygonal plates, 7 longitudinal
rows of which are enlarged to form conspicuous ridges, ovate,
slightly arched, posteriorly prolonged into a caudal point.

Plastron also a mosaic of polygonal plates, 5 longitudinal rows,
of which the median is double, forming conspicuous keels, an-
teriorly truncate, posteriorly pointed.

Color. Of juvenile from Guinea coast. Carapace and plastron
black, each keel tipped with yellow; crown and sides of head
dark, throat yellowish ; limbs black narrowly edged with yellow.

Of adult from Gulf of Gabes. Above, blackish brown, the
black ground color of the neck and fleshy parts sometimes washed
with rose and finely spotted with whitish yellow ; sides yellowish
gray ; the brownish black limbs show patches of white at their ar-
ticulations. Below, the limbs are spotted with whitish yellow.
Plastron grayish black flecked with white and yellowish spots
(Heldt:1933:13).

Size and Weight. Overall length of juvenile (ex Guinea
coast), from tip of snout to terminal tip of carapace, 95 mm.;
carapace length approximately 65 mm. ; breadth 43 mm. ; length
of forelimb from axilla to tip 58 mm.

A 9 killed at Mahfa River mouth was 6 feet long, and 4 feet
broad, with an estimated weight of 400 kg. (Biittikofer, 1890:
1:268).

502 BULLETIN : MUSEUM OF COMPAKATIVE ZOOLOQ?

A Luth taken in 1885 on the beach of the Bay of Arzew,
measured over 8 feet (21/^ metres) long, and 61/2 feet (2 metres)
broad, according to M. Bouty, Comptroller of Mines, who ex-
amined it (Doumergue:1896).

A slightly smaller one taken in the Gulf of Gabes on Septem-
ber 11, 1930, had an estimated weight of 650 kg., but this may
be exaggerated according to Heldt (1933:9), who should be con-
sulted for measurements of this specimen and more detailed
ones of another taken in the Gulf of Gabes on May 27, 1933.
Maximum weight has been given as 681.82 kg. (1,500 lbs.) (cf.
Deraniyagala, 1939:100).

Breeding. Besides many hundreds of smaller ova, one Luth
held 750 eggs with diameters ranging from 10-50 mm., but even
the largest were still immature (Heldt :1933). Lays on the Bar-
bary coast (Daudin :1802b). On November 24 and 27 respec-
tively, two Luths came ashore at the mouth of the Mahfa River,
Liberia. Hundreds of partially and fully developed eggs were
present in the turtle examined by Biittikofer (1890:1:260).

Longevity. Three weeks in New York Aquarium (Flower:
1925).

Parasites. Trematodes {Astrorchis renicapite Leidy) of the
family Pronocephalidae were present in the intestines of a Gulf
of Gabes Luth. Parasites determined by R. P. DoUfus (Heldt:
1933:33).

Companions. A Luth from the Gulf of Gabes was accompa-
nied by about 40 Sucking-fish {Bemora remora) and 100 Pilot-
fish (Echeneis ductor) says Heldt (1933 : 9 and 31).

Enemies. The flesh, though tough, oily, and an unappetizing
red, tasted better than it looked. The abundant green fat was so
nearly fluid that it partially melted in Biittikofer 's hand. The
eggs, though not without an oily taste, were considerably more
palatable than the flesh (Buttikofer, 1890:1:268). See also
Heldt (1933 :35) . On the Gold Coast the flesh and eggs are eaten
only by the Kru people (Irvine :1947).

Localities. Algeria: Golfe d'Arzeu (Baia d 'Arzew). Tunisia:
Gabes Gulf; Hafacha near Tarf il Ma; Monastir; Sidi Daoud,
Cap Bon. Egypt: Alexandria market. East Africa: Seychelles
(no coastal record known to us). Cape Province: Algoa Bay;
Cape of Good Hope; Table Bay. Southwest Africa. French

LOVERIDQE AND WILLIAMS : AFRICAN CRYPTODIRA 503

Equatorial Africa: Gabon. Togo: Sebbe (Sebc; Zebe). Gold
Coast: Tenia (B.M.). Liberia: Malila Kiver mouth near Rob-
ertsport. French Guinea f: *" Guinea Coast." Senegal: llann ;
Ilufisque.

Range. All African coasts, Indian and Atlantic Oceans, the
Mediterranean and — as an accidental visitor — the temperate
seas.

BIBLIOGRAPHYi

Works concerning African Cryptodira referred to in the text. For refer-
ences to generic and specific descriptions see synonymies.
Aellen, ViUjY

1951. Contribution a I'llerpetologie du Maroc. Bull. Soc. Sci. Nat.
Maroc, 31, pp. 153-199, figs. 1-6. {Not 1952 as in Zool. Rec).
Agassiz, Louis

1857. Contributions to the Natural History of the United States. Bos-
ton, 1 and 2. lii + 643 pp., pis. i-xxxiv.
AxDEBSON, John

1892. On a Small Collection of Mammals, Reptiles, and Batrachians

from Barbary. Proc. Zool. Soc. London, pp. 3-24.
1896. A Contribution to the Herpetology of Arabia, with a Pre-
liminary List of the Reptiles and Batrachians of Egypt. Lon-
don, pp. 1-122.
1898. Zoology of Egypt, 1. Reptilia and Batrachia. London, Ixvi -f
371 pp., figs. 1-14, pis. i-1.
Andersson, L. G.

1903b. List of Reptiles and Batrachians collected by the Swedish
(1904) Zoological Expedition to Egypt, the Soudan and the Sinaitic
Peninsula. In JLigerskiold, L. A., Results of the Swedish Zoologi-
cal Expedition to Egypt and the White Nile 1901, L No. 4, pp.
1-12 (This separate appeared in 1903, fide wrapper).
1937. Reptiles and Batrachians collected in the Gambia by Gustav
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Arkiv Zool., 29A. No. 16, pp. 1-28, figs. 1-8.
Angel, Fernand

1937b. Sur la Faune herpetologique de I'Archipel du Cap-Vert. Comptes
Rendus XII. Congrcs Internat. Zool. Lisborme 1935, 3. pp. 1693-
1700.

1 Where a date is foUowed by a letter of the alphabet, it indicates that during;
the year cited the author in' question published more than one paper on African
herpetology. The letter has chronoloj;ical sifrniticance in a more comprehensive
bibliography of African herpetology (1880-1955) which it is hoped may tie pub
lished in the not-too-distant future.

504 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1941a. Sur la Presence a Madagascar de Tortues du Genre Kinixys
Bull. Mus. Hist. Nat. (Paris), (2), 13. pp. 151152.
Angel, F., and H. Lhote

1938. Eeptiles et Amphibiens du Sahara central et du Soudan. Bull.
Com. Etudes Hist. Sci. Afr. Occid. Franc, 21. pp. 345-384.
Ansorge, J. W.

1899. Under the African Sun. New York, pp. 1-355, 134 text illus.;
2 col. pis.
Archer, W. H.

1948a. The Mountain Tortoise (Geochelone pardalis). African Wild

Life (Johannesburg), 2. No. 2, pp. 74-77, photos.
1948b. The Mountain Tortoise (GeodJielone pardalis). African Wild
Life (Johannesburg), 2. No. 3, pp. 75-78, photos.
AsTBE, Gaston

1948. Eeproduction de la Tortue mauresque a Toulouse. Bull. Soc.
Hist. Nat. Toulouse, 83. pp. 60-64.
AuBussoN, Louis Magau D'

1894. Esquisse de la Faune egyptienne. Bull. Inst. Egypt, 4. pp. 217-
(1893) 238, 293-314 (5.V.1893).
AuRELio Basilio, R. p.

1952. La Vida Animal en la Guinea Espanola. (Inst. Estudios Afri-
canos: Madrid), xii + 146 pp., pis.
Babcock, H. L.

1930. Variations in the Number of Costal Shields in Carelta. Amcr.
Nat., 64. pp. 95-96.
Bailly-Maitre, (M.)

1928. Une jeune Tortue d'Algerie vivante ... a Narbonne. Bull. Soc.
Nat. d'Acclimat. France, 75. pp. 111-112.
Baker, S. W.

1867. The Nile Tributaries of Abyssinia. Philadelphia, xxiv + 575
pp., photo, illus., maps.
Bally, P. R. O.

1946. Tortoises Eating Bones. Jour. East Afr. Uganda Nat. Hist. Soc,
(1945) 18, p. 163.
1952. Einige Beobachtungen an der ostafrikanischen Pantherschild-
krote, Testudo pardalis. Zool. Garten (Leipzig), 19, pp. 236-238.
Barbier, H.

1905. Les Cheloniens du Musee d'llistoire Naturelle d'Elbeuf. Bull.

Soc. Etude Sci. Nat. Elbeuf, 23. pp. 68-102, pis. iii-iv.
1907. Les Crocodiliens et les Batracieus du Musee d'Histoire Naturelle
d'Elbeuf. Bull. Soc. Etude Sci. Nat. Elbeuf, 25, pp. 35-74.

LOVERIDQE AND WILLIAMS : AFRICAN CRYPTODIRA 505

Barbour, T., and A. Loveridge

1930a. Reptiles and Amphibians from Liberia. In Strong, Richard,
Report of the Harvard-African Expedition upon the African
Republic of Liberia and the Belgian Congo. Cambridge, 2.
pp. 786-796.
Bateman, G. C.

1897. The Vivarium. London, pp. 1-424, figs. 1-98; pi.
Battersby, J. C.

1954. Reptiles and Amphibians collected in the Northern Frontier
Division of Kenya in 1951. Ann. Mag. Nat. Hist., (12), 7,
pp. 241-248.
Baur, Georg

1892. Bemerkungen iiber verschiedene Arten von Scliildkrotcii. Zool.
Anz., 15, pp. 155-159.

1893. Notes on the Classification and Taxonomy of the Testudinata.
Proc. Amer. Philos. Soc, 31. pp. 210-225.

Bechstein, J. M.

1800. In Lacepede, B. G. E. de " Naturgeschichte der Amphibien . . .
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Bedriaga, J. V.

1882. Die Amphibien und Reptilien Grieehenlands. Bull. Soc. Imp.
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Bell, Thomas

1827. On Two New Genera of Land Tortoises. Trans. Linn. Soc. Lon-
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1828a. Descriptions of Three New Species of Land Tortoises. Zool.

Jour., 3, pp. 419-421, (pis. xxiii-xxv in Supplement).
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1842. bered pages accompanying 40 unnumbered col. pis.
Bernstein, R. E.

1938. Blood Cytology of the Tortoise, Testudo geom.etrica. S. Afr.
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Berthold, a. a.

1850. ijber Cinixys homeana Bell. Nova Act. Acad. Leopold-Carol.
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BlANCONI, J. J.

1847- Specimina zoologica mosambicana. Bononiae, pp. 1-282, pis.
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1st. Bologna).
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1870. Observations on the Geology and Zoology of Abyssinia, made

during the Progress of the British Expedition to that Country.

London, xii -f- 487 pp., figs., pis. i-viii + i-iv.

506 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Blumenbach, J. r.

1804. Abbildungen naturhistorischer Gegenstande. Text unpaged, \W
illus. (the pertinent ones — noa. 66-70 — were published in
1804).

BOCAGE, J. V. BaRBOZA DU

1866a. Lista dos Reptis das Possessoes portuguezas d 'Africa Occidental

que existem no Museu de Lisboa. Jorn. Sci. Lisboa, 1. pp.

37-56.
1867a. Segunda Lista dos Reptis das Possessoes portuguezas d 'Africa

Occidental que existem no Museu de Lisboa. Jorn. Sci. Lisboa,

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1887a. Melanges erpetologiques. Jorn. Sci. Lisboa, 11, pp. 177-21 1.
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1895a. Herpetologie d 'Angola et du Congo. Lisbon, xx + 203 pp., pis.

i-xix.
1896a. Reptis de alguinas Possessoes portuguezas d 'Africa que existem

no Museu de Lisboa. Jorn. Sci. Lisboa, (2), 4. pp. 65-104.
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206-210.
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1874. Reptilien von Marocco und von den canarischen Inseln. Abhandl.
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Ber. Senckenberg. Naturf. Ges., pp. 132-220, pis. iii-iv.
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1885b. Liste der von Hrn. Dr. med. W. Kobelt in Algerien und Tunisien

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Naturf. Ges., pp. 3-108, pis. i-ii.
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267 316.
1893a. Katalog der Reptilien-Sammlung im Museum der Senckenberg

ischen Naturforschenden Gesellschaft iu Frankfurt am Main.

Frankfurt a. M., ix -f 140 pp.

LOVERIDGE AND WILLIAMS : AtltlCAN CBYPTODIRA 507

1893b. Ubersieht dcr von Prof. C. Keller anliisslic-h der Ruspoli'schen
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lS94a. Aiifziihlung der Arten. In Dr. Eduard Fleck's Reiseausbeute aus
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1913. Reptilicn und Aniphibien von Madagascar, den Inscln und deni
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1900. Eine ornithologische Forschungsreise durcli Tunesien. Zweiter
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1819- Anatome Testudinis Europaeae . . . Vilnae, vi + 178 pp., figs.
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1831. Saggio di una Distribuzione metodica degli Animali Vertehrati.
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BONNATERRE, AbBE

1789. Erpetologie, In Tableau encyclopedique et methodique dcs troip
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1920b. Exhibition of, and Remarks on, Living Specimens of a Remark
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BoULENGER, G. A.

1886b. On the South-African Tortoises Allied to Testudo geometrica.

Proc. Zool. Soc. London, pp. 540-542, pis. Ivii-lviii.
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Nat. Hist., (6), 2. pp. 136-141.
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508 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1891c. Catalogue of the Eeptiles and Batrachians of Barbary (Morocco,
Algeria, Tunisia), Based Chiefly upon the Notes and Collections
made in 1880-1884 by M. Fernand Lataste. Trans. Zool. Soc.
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1896a. A List of the Eeptiles and Batrachians Collected by Dr. Ragazzi
in Shoa and Eritrea. Ann. Mus. Civ. Stor. Nat. Genova, (2), 16,
pp. 545-554.

1896c. Report on Capt. Bottego's Second Collection of Reptiles and
Batrachians from Somaliland. Ann. Mus. Civ. Stor. Nat. Genova
(2), 17. pp. 15-23.

1897g. A List of the Reptiles and Batrachians of Somaliland and Galla-
land. Ann. Mus. Civ. Stor. Nat. Genova, (2), 17. pp. 275-280.

1898a. Concluding Report on the Late Capt. Bottego's Collection of
Reptiles and Batrachians from Somaliland and British East
Africa. Ann. Mus. Civ. Stor. Nat. Genova, (2), 18. pp. 715-723,
pis. Lx-x.

1900b. A List of the Batrachians and Reptiles of the Gaboon (French
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1902b. A List of the Fishes, Batrachians, and Reptiles Collected by Mr.
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Species. Proc. Zool. Soc. London, 2. pp. 13-18, pis. ii-iv.

1902d. List of the Cold-blooded Vertebrates Hitherto Recorded from
the Uganda Protectorate. An appendix to Johnston, H. H., The
Uganda Protectorate. London, 1. pp. 445-447.

1903a. The Reptiles of Sokotra. In Forbes, H. 0., The Natural History
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1905f. Reptiles de la Guinee espagnole, Mem. Real Soc. Esp. Hist. Nat.
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19061. Report on the Reptiles Collected by the late L. Fea in "West
Africa. Ann. Mus. Civ. Stor. Nat. Genova, (3), 2, pp. 196-216,
figs. 1-9.

1907a. On a Collection of Fishes, Batrachians, and Reptiles, made by
Mr. S. A. Neave in Rhodesia, North of the Zambesi, with Field
Notes by the Collector. Mem. Proc. Lit. Philos. Soc. Man-
chester, 51, pp. 1-12.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 509

1907J. Second Report on the Reptiles and Batrachians Collected in

South Africa by Mr. C. H. B. Grant, and Presented to the British

Museum by Mr. C. D. Rudd. Proc. Zool. Soc. London, pp. 478-

487, figs. 140-141, pis. xxi-xxii.
1911c. On a Third Collection of Reptiles and Batrachians Made by

Dr. E. Bayon in Uganda. Ann. Mus. Civ. Stor. Nat. Genova, (3),

5. pp. 161-169.
1912b. Missione per la Frontiere Italo-Etiopica sotto il Comando del

Capitano Carlo Citerni. Resultati zoologici. List of the Reptiles

and Batrachians, Ann. Mus. Civ. Stor. Nat. Genova, (3), 5,

pp. 329-332.
1919g. Batraciens et Reptiles recueillis par le Dr. C. Christy au Congo

beige dans les Districts de Stanleyville Haute-Uelo et Tturi en

1912-1914. Revue Zool. Af r., 7. pp. 1-29.
1920c. Una Tortue extraordinaire: Testudo Loveridgii, sp. n. Coniptes

Rendus Acad. Sci. (Paris), 170. pp. 263-266.
1920d. Presidenee de Mm. Boulenger, President d'honneur et Rabaud

Vice-President. Bull. Soc. Zool. France, 45. pp. 58-69.
BOWKEE, F.

1926. Tortoise Eggs and Nests. S. Afr. Jour. Nat. Hist., 6. p. 37.
Braestbup, F. W.

1947. Remarks on Faunal Exchange through the Sahara. Vidensk.

Medd. Dansk. Naturh. Foren, 110. pp. 1-15, fig. 1.
Bruhl, C. B.

1874- Zootomie aller Thierklassen fiir Lernende nach Autopsien skiz-
1882. zirt. Wien, pis. i-eiv.
Burton, Maurice

1953. Tortoise Tango and Dog Dance. Illus. London News, 4.vii.53,

p. 34, fig., photo.

BUTTIKOFER, JoHANNES

1890. Reisebilder aus Liberia. Leiden, 1 and 2. pp. 1-510, figs., pis.
i-xxxii.
Buxton, R. D.

1937. A Natural History of the Turkana Fauna. Jour. E. Afr. Uganda
(1936) Nat. Hist. Soc, 13. pp. 85-104, pis. A-H -f i-vi.
Cadenat, Jean

1949. Notes sur les Tortues marines des Cotes du Senegal. Bull. Inst.
Franc. Afrique Noire, 11. pp. 16-35, figs. 1-18.
Cairncross, B. L.

1946. Notes on South African Tortoises. Ann. Transvaal Mus., 20,
pp. 395-397, pi. i.
Calabresi, Enrica

1916. Batraci e Rettili raccolti nella Somalia meridionale dai Dott.

510 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Stefani e Paoli. Monitore Zool. Ital. (Firenze), 27. pp. 33-45,

pi. ii.
1923a. Missione Zoologica del Dr. E. Festa in Cirenaica. Boll. Mus.

Zool. Anat. Comp. Univ. Torino, 38. No. 7, pp. 1-28.
1927. Anfibi e Rettili raccolti nella Somalia dai Proff. G. Stefanini e

N. Puccioni. Atti. Soc. Ital. Sci. Nat. (Milano), 66. pp. 14-60,

pi. i.
Camerano, Lorenzo

1877. Dei Caratteri Sessuali Secondari della Testuda ibera Pallas. Atti

Eeal Accad. Sci. Torino, 13. pp. 97-102, pi. iv.
Cansdale, G. S.

1948c. Reptile Collecting in West Africa. II. Crocodiles, Turtles and

Snakes. Scottish Zoo and Wild Life (Edinburgh), 1. pp. 70-78,

pi.
1955. Reptiles of West Africa. Penguin Books: London, pp. 1-104,

figs. A-H.
Carpenter, G. D. H.

1920. A Naturalist on Lake Victoria. London, xxiv -f- 333 pp., figs.,

pis. i-ii, map (chapter IX. Reptiles, etc.).
Carb, Archie

1952. Handbook of Turtles. The Turtles of the United States, Can-
ada and Baja California. Ithaca, N. Y., xv + 542 pp., figs. 1-37,

pis. i-lxxxii.
Cert, F. Le

1907. Reptiles et Batraciens observes k Maison-Car6e (Algerie). Ann.

Assoc. Natur. Levallois-Perret, 13, pp. 22-26.
Chabanaud, Paul

1916e. Sur divers Reptiles et Batraciens du Maroc recueillis par M.

Pallary. Bull. Mus. Hist. Nat. (Paris), 22. pp. 228-233, figs. 1-2.
1917c. Enumeration des Reptiles non encore etudies de I'Afrique occi-

dentale, appartenant aux Collections du Museum, avec la Descrip-
tion des Especes nouvelles. Bull. Mus. Hist. Nat. (Paris), 23,

pp. 83-105, figs. 1-13.
1921a. Contribution a 1 'Etude de la Faune herpetologique de I'Afrique

occidentale. Deuxieme Note. Bull. Com. Etudes Hist. Sci. Occ.

FrauQ.. (Paris), pp. 445-472, pis. i-iv, map.
1921b. Mission Paul Chabanaud en Afrique occidentale (1919-1920).

Liste des Batraciens et des Reptiles. Bull. Mus. Hist. Nat.

(Paris), 27. pp. 519-525.
Chaignon, H. de

1904. Contributions k I'Histoire naturelle de la Tunisie. Bull. Soc.

Hiat. Nat. Autun, 17. pp. 1-166.

LOVEFilDGE AND WILLIAMS: AFRICAN CRYPTODIRA 511

Chubb, E. C.

1908. List of iiutracliia and Keptilia collected in Nortliern Matabele-

land. Ann. Mag. Nat. Hist., (8), 2. pp. 218-221.
1909a. The Batrachians and Reptiles of Matabeleland. Proc. Zool. Soc.

London, pp. 590-597.
1909b. List of Ehodesian Batrachians and Reptiles in the Rhodesia
Museum Collection. Rhodesia Mus., Bulawayo, Ann. Rep., 8, pp.
34-36.
CoNANT, Roger, and Alexander Downs

1940. Miscellaneous Notes on the Eggs and Young of Reptiles. Zoolog
ica (New York), 25. pp. 33 48.
CoNANT, Roger, and R. G. Hudson

1949. Longevity Records for Reptiles and Amphibians in the Philadel
phia Zoological Garden. Herpetologica (San Diego), 5. pp. 1-8.
Cooper, B.

1948. Note on a Land Tortoise. Ann. Rep. Game Pres. Dept. Tangan
yika Territory, 1947, p. 11.
Cope, E. D.

1859. Notes and Descriptions of Foreign Reptiles. Proc. Acad. Nat.
(1860) Sci. Philadelphia, pp. 294-297.
CoTT, H. B.

1928. Report on the Zoological Society's Expedition on the Zambesi,
1927. Proc. Zool. Soc. London, pp. 923-961, pis. i-iv.
COZZOLINO, A.

1938. Osservazioni etologiche sulla Chelone mydas delle Coste Somalia
italiana. Riv. Biol. Colon., 1, pp. 241-248, graph.

Cuvier, G. L. C. F. D.

1817. La Rdgne Animal distribu6 d'apr^s son Organisation, pour servir

de base a I'Histoire naturelle des Animaux et d 'Introduction a

['Anatomic comparee. ed. 1, Paris: Deterville, 2, Lcs Reptile.s

. . ., xviii + 532 pp.

1829. La Regne Animal . . . ed. 2, Paris, Deterville, 2. Les Reptiles

. . ., xvi 4- 406 pp.
1838. La RSgne Animal ... ed. 3, Paris, Fortin, Masson et Cie, 3,
Les Reptiles avec un Atlas par M. Duvernoy, pp. 1-169, pla.
i-xlii.
Daubenton, E. L.

1784. Les Animaux Quadruples ovipares et Les Serpents. In Ency-
clopedie m^thodique, 2, pp. 545-712.
Daudin, F. M.

I8O2II Histoire naturelle, g^nfirale et particuliSre, des Reptiles. Paris,
2. pp. 1-432, pis. xvixxviii.
Debaniyaqala, p. E. P.

1939. The Tetrapod Reptiles of Ceylop. I. Testudinates and Croco-
dilians. xxxii + 412 pp., figs. 1-137, pis. i-xxiv.

512 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1943. Subspecies Formation in Loggerhead Turtles (Carettidae).
Spolia Zeylanica (Colombo), 23. pp. 79-92, figs. 1-7.

1948. Some Scientific Eesults of two Visits to Africa. Spolia Zeylan-
ica, 25, Pt. 2, pp. 1-42, pis. i-xiv.
DoiiLFUS, E. P.

1929 Notules herpetologiques marocaines. Bull. Soc. Sci. Nat. Maroc
(Rabat), 9. p. 112.

DOUMERGUE, F.

1896. Contributions k la Faune erpetologique de la Province d'Oran.

Compte Rendu Assoc. Franc. Avanc. Sci., Carthage, 25. pt. 2,

pp. 477-478.
1899. Essai sur la Faune erpetologique de I'Oranie. Soc. Geogr. Arch.

Oran, 19. pp. 197-260, 501-532, pis. i-v.

DUERDEN, J. E.

1906a. How the Berg-Schilpad Lays her Eggs. (Published in local
paper of Feb. 2 as Albany Mus. Notes, XIII. Not seen.)

1906c. South African Tortoises of the Genus Eomopus, with Descrip-
tion of a New Species. Ree. Albany Mus., 1, pp. 405-411, figs.
1-6, pi. xi. (Actually the description of boulengeri was first pub-
lished in local paper of May 5 as Albany Mus. Notes, XVIII.)

1907a. Report of the Albany Museum for 1906. Pp. 9-10. (A smiple
list of species together with localities which are apparently the
donors' addresses and not necessarily the place of origin.)

1907b. Genetics of the Colour Pattern in Tortoises of the Genus
Eomopus and its Allies. Rec. Albany Mus., 2. pp. 65-92, figs. 1-9,
pis. vi-viii.

1907c. Variations in the Geometrica Group of South African Tortoises.
(1906) Rep. S. Afr. Assoc. Advane. Sci. for 1905-1906, pp. 178-208.

DUMERIL, A. H. A.

1856a. Note sur les ReptUes du Gabon, Revue Mag. Zool., (2), 8, pp.

369-377, pi. XX.
1860. Reptiles et Poissons de I'Afrique occidentale. Arch. Mus. Hist.
Nat. (Paris: 1858-1861), 10. pp. 137-240, pis. xiii-xix. (The
date for this part is 1860, fide Roy. Soc. Cat., though frequently
cited as 1859 or 1861.)
DuMERiL, A. M. C, and G. Bibron

1835. Erpetologie generale ou Histoire naturelle complete des Reptiles.
Paris, 2, ii -j- 680 pp., pis. xi-xxiv.
DuMERlii, A. M. C, and A. H. A. DUM^EIL

1851. Catalogue methodique de la Collection des Reptiles du Museum
d 'Histoire naturelle de Paris. Paris, iv -|- 224 pp. *
DURRELL, G. M.

1953. The Overloaded Ark. New York, xv + 272 pp., figs.

LOVERIDGE AND WILLIAMS: AFRICAN CRTPTODIRA 513

Eade, Petee

1886. Tortoises. Trans. Norfolk Norwich Nat. Soc, 4. pp. 316-322.
1893. A further Note upon Tortoises. Trans. Norfolk Norwich Nat.
Soc, 5. pp. 368-372.

ElCHWALD, C. E. VON

1851. Naturhistorische Bemerkungen iiber Algier und den Atlas. Nouv.
Mem. Soc. Imp. Nat. Moscou, 9, pp. 331-464.
EscHERiCH, Karl

1896. Beitrag zur Fauna tunisischen Insel Djerba. Verh. Zool. Bot.
Ges. Wien, 46. pp. 268-279.

EVERSMANN, EdWARD

1854. Einiges aus meinem Tagebuche wjihrend einer Eeise in's Ausland
im Winter 1852-53. Bull. Soc. Imp. Nat. Moscou, 27. pp. 398-445.

Falk, Kurt

1922. Siidwestafrikanische Eeptilien und ihre Heimat. Wochensch.
Aquar.-Terrarienkunde (Braunschweig), 19. pp. 160 164, figs.
Fejerva'ry, G. J. F. DE

1927. On a Small Collection of Reptiles from Morocco. Ann. Mag. Nat.
Hist., (9), 20. pp. 512-518.
FiSK, G. H. B.

1878. Two Leopard Tortoises {Testudo pardalis). Proc. Zool. Soc.
London, p. 790.

FiTZIXGER, L. J. F. J. DE

1826. Neue Classification der Eeptilien nach ihren naturlichen Ver-
wandtschaften . . . Wien, viii + 66 pp., pi. i.

183.3. Entwurf einer svstematischen Anordnung der Schildkroten nach
(1836) den Grundsatzen der natiirlichen Methode. Ann. Wiener Mus., 1,
pp. 103-128.

1843. Systema Eeptilium . . . Fasciculus primus: Amblyglossae (Con-
spectus geographicus). Vindobonae, pp. 1-106.

1855. Bericht an die kaiserl. Akademie der Wissenschaften iiber die
von dem Herrn. Consulatsverweser Dr. Theodor v. Heuglin fiir
die Menagerie zu Schonbrunn mitgebrachten lebenden Thiere.
Sitzungsb. K. Akad. Wiss. Wien, 17. pp. 242-253.

1861. Die Ausbeute der osterreichischen Naturforscher an Siiugethieren
(1800) und Eeptilien wahrend der Weltumsegelung Sr. Majestiit Fre-

gatte Novara. Sitzungsb. K. Akad. Wiss. Wien, 42. pp. 383 416.
FiTzSlMONS, V. F.

1932. Preliminary Description of New. Forms of South African Eep-

tilia and Amphibia, from the Vernay-Lang Kalahari Expedition,

1930. Ann. Transvaal Mus., 15. pp. 35-40.
1935a. Notes on a Collection of Eeptiles and Amphibians made in the

Southern Kalahari, Bushmanland and Great and Little Nama-

qualand. Ann. Transvaal Mus., 15. pp. 519-550.

514 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1935b. Scientific Results of the Vernay-Lang Kalahari Expedition,
March to September, 1930. Eeptilia and Amphibia. Ann. Trans-
vaal Mus., 16, pp. 295-397, figs. 1-30, pis. x-xi.

1937a. Notes on the Reptiles and Amphibians Collected and Described
from South Africa by Andrew Smith. Ann. Transvaal Mus., 17,
pp. 259-274, pi. X.

1938. Transvaal Museum Expedition to South-West Africa and Little
Namaqualand, May to August 1937. Reptiles and Batrachians.
Ann. Transvaal Mus., 19. pp. 153-209, pis. ii-iv, map.

1939b. An Account of the Reptiles and Amphibians Collected on an
Expedition to South-eastern Rhodesia during December 1937 and
January 1938. Ann. Transvaal Mus., 20, pp. 17 46.

1946a. An Account of the Reptiles and Amphibians Collected on an
Expedition to the Cape Province, October to December, 1940.
Ann. Transvaal Mus., 20, pp. 351-377, pis. xiv-xvii.

1950. Notes on a Collection of EeptUes and Amphibians from the West
Coast of Southern Africa. Ann. Transvaal Mus., 21, pp. 253-259.
pi. iv.
Fleck, Edou.\rd

1894. Vorkommen und Lebensweise der Reptilien und Batrachier. Ber.
Senckenberg. Naturf. Ges., pp. 83-87.
Flower, S. S.

1900. On some Reptiles and Batrachians from the White Nile. Proc.
Zool. Soc. London, pp. 967-968.

1924a. A remarkable Tortoise of the Genus Testudo. Proc. Zool. Soc.
London, pp. 920-921.

1925b. Contributions to our Knowledge of the Duration of Life in Ver-
tebrate Animals. III. Reptiles. Proc. Zool. Soc. London, pp.
911-981.

1926. Species of Land Tortoises of the Genus Testudo Found in the
Countries Bordering the Mediterranean Sea. Copeia, p. 133.

1928. Exhibition of a great African Tortoise (Testudo sulcata). Proc.
Zool. Soc. London, p. 654, figs. 1-2.

1929. Reptiles. Ln List of the Vertebrated Animals exhibited in the
Gardens of the Zoological Society of London, 1828-1927. Lon-
don, 3. pp. 1-272.

1933. Notes on the Recent Reptiles and Amphibians of Egypt, with a

List of the Species Recorded from that Kingdom. Proc. Zool.

Soc. London, pp. 745-851, map.
1937a. Further Notes on the Duration of Life in Animals. III. Rep

tiles. Proc. Zool. Soc. London, pp. 1-39.
1945. Persistent Growth in the Tortoise, Testudo graeca, for Thirty

nine Years, with other Notes Concerning that Species. Proc.

Zool. Soc. London, 114, pp. 451-455, fig. 1.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 515

P^jCHS, Hugo

1914. Uber den Bau und die Entwicklung dea Schadels der Chelone
imbricata. I. Das Priraordialskelett dea Neurocraniums und
Kieferbogens. In Voeltzkow, W., Reise in Ostafrika in den
Jahren 1903-1905, 5, pp. 1-325, text figs. 1-182, pis. i-iv.
Gadow, Hans

1901. Amphibia and Reptiles. In The Cambridge Natural History.
London, 8, xiii -f- 668 pp., figs. 1-181, map (reprinted in 1920).
Gendre, E.

1909. Liste de quelqucs EspSces de Reptiles du Fouta Djalon. Comptes
Rendus Soc. Linn. Bordeaux, 63. pp. cv-cvi.
Geoffroy St.-Hilaire, E. F.

1809a. Sur les Tortues molles, nouveau genre sous le Nom de Trionyx
et sur la Formation des Carapaces. Ann. Mus. Hist. Nat.
(Paris), 14. pp. 1-20, pis. i-v.
1809b. Synopsis of 1809a. Nouv. Bull. Soc. Pliilom. Paris, 22. pp. 363-

367.
1827a. Description des Reptiles qui se trouvent en Egj-pte. In Descrip-
tion de I'Egypte, ou recueil des observations et des recherches
qui ont ete faites en Egypte pendant 1 'expedition de I'armee
FranQaise (1798-1801), 1. Histoire naturelle. Paris, pp. 115-184,
pis. i-vi.
(iERVAIS, F. L. P.

1836. Enumeration de quelques Especes de Reptiles provenant du Bar

barie. Ann. Sci. Nat. Zool. (Paris), (2), 6, pp. 308-313.
1848. Sur les Animaux vertebres de I'Algerie. Ann. Sci. Nat. Zool.

(Paris), (3), 10. pp. 202-208.
1872. Osteologie du Sphargis Luth {Sphargis coriacea). Arch. Mus.
Hist. Nat. (Paris), (N.S.), 8. pp. 199-228, pis. v-ix.
(tEstro, Raffaele and Decio Vinciguerra

1931. Rettili, Batraei. In Resultati scientifici della Missione alia Oasi
di Giarubub. 1926-1927. Roma, 4. pp. 538-539. (This is a re
print, but under two names, of Vinciguerra 's 1927 paper in Ann.
Mus. Civ. Stor. Nat. Genova).

Ghigi, Alessandro

1913. Materiali per lo Studio della Fauna Libica. Mem. Accad. Sci.
Inst. Bologna, (6), 10. pp. 253-296.

1932. Escursione zoologica all Oasi di Marrakesch nell' Aprile 1930.
Boll. Zool. (Napoli), 3. pp. 197-208.

GiRONS, H. Saint

1953c. Influence de la Mise en Eau du 'Barrage de Bin el Ouidane sur
les Vertebres terrestres. Bull. Soc. Sci. Nat. Maroc, 33. pp. 75-80,
figs. 1-2.

516 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Gmelint, J. F.

1789. Caroli a Linne . . . Sj'stenia Naturae . . . (ed. 13, Lipsiae), 1,
(1788) part 3, pp. 1033-1516 (Date of this part is 1789, fidfi John Hop-
kinson, 1907, Proc. Zool. Soc. London, pp. 1035-1037).
GoLDFtrss, G, A.

1820. Handbuch der Zoologie. Niirnberg, 2, pp. 1-510, pis. iii-iv.
GoRHAM, F. W. and A. C. Ivy

1938. General Function of the Gall Bladder from the Evolutionary
Standpoint. Field Mus. Nat. Hist., Zool. Ser., 22, pp. 159-213.
Gray, J. E.i

1825. A Synopsis of the Genera of Eeptiles and Amphibia, with a
Description of Some New Species. Ann. Philos., (2), 10. pp.
193-217.
1828 Speeilegia Zoologica ; or Original Figures and Short Systematic
and Descriptions of New and TJnfigured Animals. Part I (1828), pt.
1830. II (1830), pp. 1-12 (pis. i-ix were never issued but an unpub-
lished set of them is in the Brit. Mus.).
1831b. Synopsis of the Species of the Class Reptilia. hi Griffith, E.,
The Animal Kingdom Arranged in Conformity with its Organ-
ization by the Baron Cuvier. London, 9, pp. 1-481 -|- 1-110, pis.
i-xlii (Gray is on pp. 1-110 of addenda. In his Bibliography the
date of this work appears as 1830, possibly an earlier printing.
It seems to have come out before 1831c which is usually cited
for the fuller descriptions of new names).
1831c. SjTiopsis Eeptilium or Short Descriptions of the Species of Rep-
tiles. Part I. Cataphracta, Tortoises, Crocodiles, and Enalio-
saurians. London, viii + 85 pp., pis. i-xi.
1844. Catalogue of the Tortoises, Crocodiles, and Amphisbaeniaus, in
the Collection of the British Museum. London, pp. 1-80.

1854. Descriptions of a New Genus and Some New Species of Tortoises.
(1852) Proc. Zool. Soc. London, for 1852, pp. 133-135.

1855. Catalogue of Shield Reptiles in the Collection of the British
Museum. I. Testudinata (Tortoises). London, pp. 1-82, pis.
i-xlii.

1856. On Some New Species of Freshwater Tortoises from North
(1855) America, Ceylon and Australia, in the Collection of the British

Museum. Proc. Zool. Soc. London, for 1855, pp. 197-202.
1860a. Description of a Soft Tortoise (Aspidochelys Livingstonii) from
the Zambesi, Sent to the British Museum by Dr. Livingstone.
Proc. Zool. Soc. London, pp. 5-6, pi. xxii.

I Despite its numerous inaccuracies and errors, we have been greatly helped in
assembling this list of Gray's publications on African testudinates by a "List of
the Books, Memoirs and Miscellaneous Papers by John Edward Gray." Compiled
in 1874-1875 by J. Saunders, London. Privately printed pamphlet listing about
1,162 titles.

LOVERTDGE AND WILLIAMS: AFRICAN CRYPTODIRA 517

lS60b. Description of a New Specdes of Emys Lately Living in the

Gardens of the Zoological Society. Ann. Mag. Nat. Hist., (3),

6, pp. 148-149.
1860c. Eeprint of 1860b. Proc. Zool. Soc. London, pp. 232-233, pi. xxx.
1860d. On the African Trionices with Hidden Feet (Emyda). Ann.

Mag. Nat. Hist., (3), 6. pp. 440-442.
1860e. Reprint of 1860d. Proc. Zool. Soe. London, pp. 314-316.
1860f. Note on Aspkloohelys Living stonii. Proc. Zool. Soc. London, pp.

430-431.
1861. Reprint of 1860f. Ann. Mag. Nat. Hist., (3), 7. pp. 226-227.
1863b. Notice of a New Species of Kinixys and Other Tortoises from

Central Africa. Ann. Mag. Nat. Hist., (3), 12. pp. 381-382.
1863f. On the Species of the Genus Stcrnothaerus, with some Observa-
tions on Kinixys. Proc. Zool. Soc. London, pp. 192197, figs.
1864a. Reprint of 1863f. Ann. Mag. Nat. Hist., (3), 13. pp. 165-170.
1864c. Revision of the Species of Trionychidae found in Asia and

Africa, with the Descriptions of Some New Species. Proc. Zool.

Soc. London, pp. 76-98, figs. 1-21.
1864d. On the Genera of Chelydidae and the Characters Furnished by

the Study of their Skulls. Proc. Zool. Soc. London, pp. 128-135,

figs. 1-3.
1865d. Notes of a New Genus and Species of the Family Trionychidae

from West Africa. Ann. Mag. Nat. Hist., (3), 16. pp. 204-206,

fig.

1865e. Reprint of 1865d. Proc. Zool. Soc. London, pp. 323-325, fig.

1865f. On the Development of the Sternal Callosities in Cyclanosteus
senegalensis, and on the Synonyms of Cyclanosteus and its Allied
Genera. Proc. Zool. Soc. London, pp. 422-428, figs. 1-2.

1866. Notes on some Young Specimens of Tortoises {Testudo). Proc.
Zool. Soc. Loudon, pp. 305 306.

1869b. Descriptions of a New Species of Freshwater Turtle and Chame-
leon. In Petherick, John, Travels in Central Africa and Explora-
tions of the western Nile Tributaiies. London, 2, Appendbc B,
pp. 189-193.

1869c. Notes on the Families and Genera of Tortoises (Testudinata),
and on the Characters Afforded by the Study of their Skulls.
Proc. Zool. Soc. London, pp. 165-225, figs. 1-20, pi. xv. (incor-
porated in 1870e with very few changes).

1869d. Description of Mauremys laniaria, a new Freshwater Tortoise.
Proc. Zool. Soc. London, pp. 499-500, pi. xxxvii.

1869e. Descriptions of Emys flavipes. Proc. Zool. Soe. London, pp. 643-
644, pi. 1 (i.e. 50).

518 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1870c. Notes on Tortoises in the British Museum, with Descriptions of
Some New Species. Proc. Zool. Soc. London, pp. 653-659, pis.
xxxiii-xxxiv.

1870d. Notes on a Specimen of Cyclanosteus senegalensis Living in the
Societj^'s Gardens. Proc. Zool. Soc. London, pp. 717-718, pi. xliii.

1870e. Supplement to the Catalogue of Shield Reptiles in the Collection
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1934. The Reptiles and Amphibians of Tunisia. Publ. Univ. Calif.
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1920. Reptiles recueillis de Mai a Juin au cours d'une Excursion zoolo-
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1949. Nitrogenous Excretion in Chelonian Reptiles. Biochem. Jour.,
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1882a. Erster Nachtrag zum Katalog der herpetologischen Sammlung
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528 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1887a. Fiinfter Nachtrag zum Katalog der herpetologischen Sammlung

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MULLER, LORENZ

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MULLER, p. L. S.

1774. Vollstandiges Natursystem. Niirnberg, 3, 350 + [161 PP^
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1905. Uber nordost-afrikanisehe und arabische Krieehtiere. Zool.
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1894. Herpetologie algerienne ou Catalogue raisonne des Eeptiles et
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1896b. Materiaux pour la Faune de la Tunisie. I. Catalogue de Rep-
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1895. Einige Bemerkungen iiber zwei siidafrikanische Schildkroten.
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Pabenti, Paolo and LuiGi Picaglia

1886. Rettili ed Anfibi raccolti de P. Parent! nel Viaggio di Circuni-
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LOVERIDGE AXD WILLIAMS: AFRICAN CRYPTODIRA 529

Mar Rosso e dell' America meridionale negli Anni 1879-84.

Atti Soc. Nat. Modena (3), 5. pp. 26-96.
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1932a. Scientific Results of the Cambridge Expedition to the East

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Parker, H. W., R. E. Moreau and R. H. W. Pakenham

1940. Notes on the Herpetology of the East African Islands. Ann.
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Pechuel-Loesche, Eduard

1882. Die Loango-Expedition. Leipzig, Part III, pp. 1-503, figs., pis.,
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Peel, C. V. A.

1900. Somaliland. London, xv + 345 pp., figs., pi., map.
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1909b. Reptiles du Soudan recoltes par la Mission Tilho-Gaillard. De-
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1913. Les Vertebres des Eaux douces du Maroc. Comptes Rendus

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1914. Les Vertebres des Eaux douces du Sahara. Comptes Rendus

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1926a. Liste des Reptiles, Batraciens et Poissons d'Eau douce des Col-
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1926c. La Longevite chez les Reptiles en Captivite. Revue Gen. Sci.
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1927a. Les Reptiles et les Batraciens de I'Afrique du Nord fran^aise.
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530 BULLETIN : MUSEUM OF COSrPARATIVE ZOOLOGY

Peters, W. K. H.

1854a. Ubersicht der auf seiner Reise nach Mossambique beobacliteten

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Barnim und Dr. Hartmann auf ilirer Reise durch Aegypten,

Nubien und dem Sennar gesammelten Amphibien. Monatsber.

Akad. Wiss. Berlin, pp. 271-279, pi.
1866b. Ubersicht der aus dem Nachlass des Baron Carl von der Decken

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JS69a. Amphibien. In Decken, C. C, Reisen in Ost-Afrika. Leipzig, 3,

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1873c. Uber eine von Hrn. F. Pollen und van Dam auf Madagascar und

anderen ostafrikanischen Inseln gemachte Sanimlung von Am

phibien. Monatsber. Akad. Wiss. Berlin, pp. 792-795.
1875a. Uber die von Hrn. Professor Dr. R. Buchholz in Westafrika

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196-212, pis, i-iii.
]S76a. Eine zweite Mittheilung iiber die von Hrn. Professor Dr. R.

Buchholz in Westafrika gesammelten Amphibien. Monatsber.

Akad. Wiss. Berlin, pp. 117-123, pi. i.
1877c. Ubersicht der Amphibien aus Chinchoxo (Westafrika) welche

von der afrikanischen Gescllschaft dem berliner zoologischen

Museum iibergeben sind. Monatsber. Akad. Wiss. Berlin, pp.

611-621, pi.
1878a. ijber die von Hrn. .J. M. Ilildebrandt wiihrend seiner letzten

ost-afrikanischen Reise gesammelten Sjiugethiere und Amphibien.

Monatsber. Akad. Wiss. Berlin, pp. 194-209, pis. i-ii.
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Reise nach der Oase Kufra gesammelten Amphibien. Monatsber.

Akad. Wiss. Berlin, pp. 305-309.
1881a. Amphibien der Expedition nach Kufra. In Rohlfs, G., Kufra.

Leipzig, pp. 1-559, charts 1-21, maps. (Reprint of 1880d).
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seiner Majestiit des Konigs Friedrich Wilhelm IV in den Jahren

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Pitman, C. R. S.

1934. A Check List of Reptilia and Amphibia Occurring and Believed

to Occur in Northern Rhodesia. In Pitman, C. R. S., Report on a

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+ Index of i-xxxii, maps A-K.
1942b. A Game Warden takes Stock. London, xvii -f 287 pp. pis.

LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 531

PoiRET, .7. L. M.

1789. Voyage en Barbarie ou Lettres ecrites de I'Ancienne Numidic.
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Power, J. IT.

1927c. On the Herpetological Fauna of the Lobatsi-Linokana Area.
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1931. Part II, Ibid, 20. pp. 39-49, figs. 1-2.
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Sci., 29. pp. 466-472.
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Prato, Alberto del

1895. Vertebrati Eritri aggiunta al Catalogo della Collezione Eritrea
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1922b. A Study of the remarkable Tortoise, Testudo loveridgii Blgr.,
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191. J. Das zentrale Nervensystem einiger madagassischer ReptUien.
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Reichenow, Anton

1874. Eine Samnilung Lurche und Kriechthiere von Westafrika. Arch.

Naturg., 40, Abt. 1, pp. 287-298, pi. ix.
1878. ijber einige nachtraglich von der deutschen Expedition nach
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Rochebrune, a. T. de

1884a. Faune de la Senegambie. Reptiles. Paris, pp. 1-221, pis. i-xx.
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Rose, Walter

1929. Veld and Vlei. Cape Town, xxiii + 240 pp., figs., photos 1-125.
1950. The Reptiles and Amphibians of Southern Africa. Cape Town,
xxviii -f 378 pp., frontis. and photos 1-213.
Roux, Jean

1910. Reise von Dr. .1. Caii ini nordliehen zentral-afrikanischen
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Rozet, C. a.

1833. Voyage dans la Regence d 'Alger . . . I'Histoire naturelle, etc.
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532 IJTJLLETIN : MUSEi.M OK COMPARATIVE ZOOLOGY

RuCKES, Herbert

1937. The Lateral Arcades of Certain Emydids and Testudinids.
Herpetologica (San Diego), 1, pp. 97-103, pi. x.
RiJppELL, Edward (W. P. E. S.)

1835. Neue Wirbelthiere zu der Fauna von Abyssinien gehorig. III.

Amphibien. Frankfurt a. M., pp. 1-18, pis. i-vi.
1845a. Beschreibung und Abbildung einer neuen Art von Landschild-
kroten, zur Gattung Kinixys gehorig. Mus. Senckenberg., 3, pp.
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1845b. Verzeichniss der in dem Museum der Senckenbergischen Gesell-
schaft aufgestellten Sammlung. Amphibien. Mus. Senckenberg.,
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KCJST, H. T.

1937. Interessante Sehildkroten. IV. Die Gattung Homopii.<i Dumeril
et Bibron, 1834. Wochenschr. Aquar.-Terrar.-Kunde (Braun-
schweig), 34, pp. 699-700, photos.

Sauvage, H. E.

1884b. Notice sur une Collection de Reptiles et de Poissons recueillie
a Mayumba, Congo. Bull. Soc. Zool. France, 9. pp. 199-208, pi. vi.
SCHEPERS, G. W. H.

1938. The External Morphology of the Brain of Testudo geometrica.
Jour. Anat. (Cambridge, Eng.), 72. pp. 535-555, figs. 1-5.

1939. The Blood Vascular System of the Brain of Testudo geometrica.
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1948a. A Dorso-lateral Invagination of the Olfactory Bulb va. Testudo
geometrica. S. Afr. Jour. Sci. (Johannesburg), 2. pp. 9-11, figs.
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1948b. Evolution of the Forebrain. The Fundamental Anatomy of the
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ScHERER, Joseph

1902. Herpetologische Reiseskizzen aus Zentral-Ost-Afrika. Blatt.
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ScHLEGEL, Hermann

1841. Bemerkungen iiber die in der Regentschaft Algier gesammelten
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Schmidt, K. P.

1919. Contributions to the Herpetology of the Belgian Congo Based
on the Collection of the American Museum Congo Expedition
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vii-xxxii.

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1933. The Eeptiles of the Pulitzer Angola Expedition. Ann. Carnegie

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Schneider, J. G.

1792. Beschreibung und Abbildung einer neucn Art von Wasseischild-

krote' nebst Bestinimungen einiger bisher wonig bekannten

fremden Arten. Ges. Naturf. Freunde Berlin, 10, pp. 259-284,

pi. vii.
SCHOEPFF, J. 1).

1792. Naturgoschic'hte der Schildkroten mit Abbiklungen erliiuteit.

Erlangae, 12 + 160 pp., pis. i-xxxii (? x.xxiii-xxxiv).
1792- Historia Testudinum. Erlangae, xii + 136 pp., pis. i-xxxii
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SCHWEIGGER, A. F.

1812. Monographiae Chelonioium. Konigsberger Arch. Naturwiss.
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1814. Prodromi Monographiae Cheloniorum. Regiomonti, vi 4- 58 pp.
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RCLATER, P. L.

1871b. On Additions to the Menagerie. Proc. Zool. Soc. London, pp.
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1872. On Additions to the Society's Menagerie. Proc. Zool. Soc. Lon-
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SCLATER, W. L.

1898. List of the Reptiles and Batrachians of South Africa, with
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ScoRTECCi, Giuseppe

1928b. Rettili dell 'Eritrea esistenti nelle Collezioui del Museo Civico
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figs. 1-8, pis. vii-ix.

1930a. Rettili ed Anfibi raccolti dal Prof. E. Zavattari in Eritrea.
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1940a. Reptilia. In Missione biologica nel paese dei Borana raccolte
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1943. Reptilia. In Missione biologica Sagan-Omo. Zoologia. Roma,
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Seba, Albertus

1735. Locupletissimi Rerum Naturalium Thesauri Accurate Descriptio,
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1918. Nematodes de la Clemmyde lepreuse. Bull. Soc. Hist. Nat. Afr.
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534 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1925a. Cistudo europaea aux Environs de la Calle. Bull. Soc. Hist. Nat.

Afr. Nord (Alger), 16. pp. 150-151.
1927. L'Estuaire de I'Akarit (Golfe de Gabes). Bull. Soc. Hist. Nat.

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Shaw, George

1802. General Zoology or Systematic Natural History. 3. Amphibia.
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1797. Naturalists Miscellany. London, 9. letterpress to i)ls. ceci-cccxlviii.
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1897. Das Kopfskelet der Schildkroten. Sitzungsb. Akad. Wiss. Wien.
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1898. iiber den Bau und die Entwicklung des Zungenbein-Apparates
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1899. ijber den Kehlkopf und die Luftrohre der Schildkroten. Sit-
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1902b. Zur Systematik der Schildkrotenfamilie Trionychidae Bell, nebst

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1903b. iJber zwei seltene und eine neue Sehildkrote des berliner Mu

seums. Sitzungsb. Akad. Wiss. Wien, 112. Abt. 1. pp. 439-445, pi.
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Eiickenschale von Testudo tornieri Siebenr. Sitzungsb. Akad,

Wiss. Wien, 113, Abt. 1, pp. 29-34, fig.
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Anz. Akad. Wiss. Wien, 41. pp. 194-195.
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Sitzungsb. Akad. Wiss. Wien, 113. Abt. 1. pp. 307-323, pis. i-v.
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817-839, figs. 1-3.
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1907. tlber zwei Schililkroten aiis Kamerun. Ann. Xaturhist.
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1919. Emydnra macqvarrii Gray und der systematischo Wert dcs

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1897. Eeptilien aus Kamerun, West-Afrika. Bihang till K. Svenska

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1885. Notes sur trois Testudinides dc I'Afrique Australe. Ann. Soc.

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Smith, Andrew

1838- Illustrations of the Zoology of South Africa. Eeptiles. London.
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SORDELLI, FeRDINANDO

1901. Anomalia in una Testuggine {Cinixys belliana) del Sudan ori-
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1908. Vertebrati dell 'Argentina e del Benadir donati al Civico Museo
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SowERBY, J. DE C. and Edward Lear

1872. Tortoises, Terrapins, and Turtles. London, pis. i-lx (letterpress
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1953. Africa: A Study in Tropical Development. (New York, vii +
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1867a. Eeptilien. In Eeise der osterreichischen Fregatte Xovara urn
die Erdc in don Jahrcn 1857, 1858, 1859, unter den Befehlen des

536 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Commodore B. von Wullerstorf-IJrbair. Zoology, I, pp. 1-98,

pis. i-iii.
1870a. Herpetologische Notizen II. 1. Eeptilien gesammelt wahrend

einer Eeise in Senegambien (October bis December 1868). Sit-

zungsb. Akad. Wiss. Wien, 62, Abt. 1. pp. 326-335.
1891c. Ueber die Eeptilien und Batrachier der westliehen und ostlichen

Gruppe der canarischen Inseln. Anji. Naturhist. Hofmus.

(Wien), 6. pp. 287-306.
1901. Expedition S, M. Schiff "Pola" in das Eothe Meer. XVII.

Bericht iiber die herpetologischen Aufsammlungen. Denksehr.

Akad. Wiss. Wien, 69. Abt. 1. pp. 325-335, pis. i-ii.
Stemmler-Morath, Carl

1952. Biologische Beobachtungen an Clemmys leprosa in Siidmarokko.

Aquar.-Terrar. Zeitsehr. (Stuttgart), 5. pp. 217-218, photos 1-2.
Sternfeld, Egbert

1910. Die Eeptilien (ausser den Schlangen) und Amphibien Kameruns.

In Die Fauna der deutschen Kolonien. Berlin, Ser. 1, part 2,

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1911b. Zur Herpetologie Siidwestafrikas. Mitt. Zool. Mus. Berlin, 5,

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1911d. Die Eeptilien (ausser den Schlangen) und Amphibien von

Deutsch-Siidwestafrika. In Die Fauna der deutschen Kolonien.

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1912c. Eeptilia. In Wiss. Ergeb. der deutschen Zentral-Afrika-Expedi-
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1917. Eeptilia und Amphibia. In Wiss. Ergeb. der zweiten deutselien

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Stevenson-Hamilton, J.

1947. Wild Life in South Africa. London, vii + 364 pp.
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1862a. Chelonologische Studien, niit besonderer Beziehung auf die

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LOVERIDGE AND WILLIAMS : AFRICAN CRYPTODIRA 537

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Temminck, C. J. and Hermann Sciileqel

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TORNIEB, GUST.W

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1900b. Neue Liste der Crocodile, Schildkroten und Eidechsen Deutsch-
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1902b. Herpetologisch Neues aus Deutsch-Ost-Afrika. Zool. Jahrb., Syst.
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1905c. Schildkroten und Eidechsen aus Nordost-Afrika und Arabien.
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Trimen, Roland

1893 Reptiles and Amphibians. In Noble, John, Illustrated Hand-
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1860. The Great Sahara: Wanderings South of the Atlas Mountains.
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herpetological list in the Appendix is a mere compilation of

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Vaillant, Leon

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1949. La Faune. In Encyelopedie coloniale et maritime Afriquo Occi-

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1927. Risultati zoologici della Missione inviata dalla E. Societa geo-

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Werner, Franz

1892a. Ausbeute einer herpetologischen Excursion nach Ost-Algerien.

Verh. Zool.-Bot. Ges. Wien, 42, pp. 350-355.
1892b. Tierleben in Ost-Algerien. Zool. Garten, 33, pp. 264-273.
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1898a. Ueber Eeptilien und Batrachier aus Togoland, Kamerun uud

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Zool.-Bot. Ges. Wien, 48. pp. 191-213, figs., pi. ii.
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52, pp. 332-348.
1908. Ergebnisse der mit Subvention aus der Erbschaft Trcitl unter
(1907) nommen zoologischen Forschungsreise Dr. Franz Werner's nach

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIRA 58!)

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1908a. On Some Interesting Reptiles Collected by Dr. C. M. Wenyoji

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1914b. Ergebnisse einer von Prof. F. Werner im Sommer 1910 mit

Unterstiitzung aus dem Legate Wedl ausgefiihrten zoologischen

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331-358, pi.
1915. Reptilia und Amphibia. In Miehaelsen, W., Beitrtige zur Keunt

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Hamburg, pp. 325-376, pi. vii.
1920. Edition 5 of 1912b.

1924a. Wissensehaftliehe Ergebnisse der mit Unterstiitzung der Akade
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F. Werner unternommenen zoologischen Expedition nach dem

anglo-agyptisehen Sudan (Kordofan) 1914. XVIII. Schild

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1929b. Wissensehaftliehe Ergebnisse einer zoologischen Forschungsreise

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138. Abt. 1. pp. 1-34, fig. 1, pis. i-iv.
liCMc. Ergebnisse einer zoologischen Forschungsreise nach Marokko.

111. Amphibien und Reptilien. Sitzungsb. Akad. Wiss. Wien.

140, Abt. 1, pp. 271-318, pis. i-iv, maps 1-3.
Wr.TTSTEiN, Otto

1924. Uber Tcstudo tornieri Siebr. und Tesfudn loveridgii Blgr. Zool.

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Wn.MAMS, E. E.

1950. Variation and Selection in the Cervical Central Articulation of

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1952. A New Fossil Tortoise from Mona Island, West Indies, and a

540 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Tentative Arrangement of the Tortoises of the World. Bull

Amer. Mus. Nat. Hist., 99. pp. 543-560.
Williams, E. E. and S. B. McDowell

1952. The Plastron of Soft-shelled Turtles (Testudiuata: Trionychi-

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WiTTE, G. F. DE

1924d. Deux Tortues a Squelette reduit. Revue Zool. Afr., 12. Supjjl.,

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1928b. Les Tortues Trionyx du Lac Albert. Revue Zool. Afr., 15,

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1932d. Un Tortue marine. Revue Zool. Bot. Afr., 22. Suppl., p. 57.
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1935. Un Tortue marine de Banana. Revue Zool. Bot. Afr., 27. Suppl.,

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1941. Exploration du Pare National Albert. Mission G. F. de Witte

(1933-1935). Batraciens et Reptiles. Inst. Pares Nat. Congo

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1952. Amphibiens et Reptiles. In Exploration Hydrobiologique du Lac
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1953. Exploration du Pare National de I'Upemba. Mission G. F. de
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WORTHINGTON, E. B.

1929. A Report on the Fishing Survey of Lakes Albert and Kioga.
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WoRTHiNGTON, S. and E. B.

1933. Inland Waters of Africa. London, ix + 259 pp., figs. 1-13, pis.
i-xi, map.
Wyman, Jeffries

1845. Exhibition of "Shell of Kinixys homeana, Bell, from Cape de
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Zangerl, Rainer

1939. The Homology of the Shell Elements in Turtles. Jour. Morph.,
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Zavattari, Edoardo

1922. Vertebrati di Cirenaica raccolti dal Generale Medico Prof.
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1929. Elenco degli Anfibi e Rettili della Cirenaica. Notiziario E<-o-
noniieo della Cirenaica (Bengasi), pp. 86-88.

LOVERIDGE AND WILLIAMS: AFRICAN CRYPTODIKA 541

1930a. Erpetologia della Cirenaica. Arch. Zool. Ital. (Torino), 14, pp.

253-287.
1937. I Vertebrati della Libia. In Festschrift zum 60. Geburtstage

von Professor Dr. Embrik Strand. Riga, 2, pp. 526-560.
ZuLUETA, Antonio de

1908. Nota sobre Batracios y Reptiles de Mogador, con Descripcion

de la Forma joven de Saurodaotylus mauritanicus (Dum. et

Bibr.). Bol. Real See. Esp. Hist. Nat. (Madrid), 8. pp. 451-457.

542

INDEX

INDEX TO ALL SCIENTIFIC NAMES ^

Achilemys, 219.

Achhiixys, 218, 259, 296, 297, 343,

377.
Acinixys planicauda, 317.
,Aeg.yptiacuni, Hyalomma, 273.
aegj'ptiacus, Aspidonectes, 424.
aegyptiacus, Gymnopus, 424.
aegyptiacus, Tiionyx, 420, 422, 423.
africana, Foidia, 421, 425.
afiicana, Testudo, 368, 369.
africana, Testudo t., 345, 346.
agaasizii, Chelonia, 475.
agassizii, Gopherus, 186.
alata, Mutela, 466.
nlbanica, Cheisinella t., 301, 337,

339.
albanica, Psammobates t., 337.
Aldabiachelys, 166, 223, 225.
Aldabrachelys gigantea, 225.
Aldabiachelys grandidieri, 225.
Aldabrachelys sumeiri, 225.
aldabrensis, Fregatta m., 480.
amasensis, Chersinella v., 301, 327.
Amblyomnia exornatum, 248.
Amblyomma marmoreum, 248, 293,

393.
Amblyomma nuttalli, 248, 293, 393.
amboineiisis, Testudo m., 319.
Amyda, 421, 422.
Amyda mutica, 422.
Amyda subplana, 422.
Amyda triunguis, 426.
A myda t. triunguis, 427.
angulata, Chersina, 167, 212, 344,

345-352, S13, 214, S17, 344, S47,

349.
angulata, Chersine, 346.
angulata, Neotestudo, 346.

angulata, Testudo, 342, 345, 346.
angulata, Testudo (C.) 346.
angulatus, Goniochersus, 346.
angusta, Sphargis, 500.
Aunamemys, 187, 188.
a,ntiguorum, Emys, 202.
arachnoides. Pyxis, 259, 364, S17.
aralensis, Emys o., 204.
arcuata, Testudo, 499.
Ardea c. cinerea, 480.
arcolata, Chersine, 368.
areolata, Testudo, 352, 367, 369,

387.
areolatus, Ilomopus, 173, 353, 355-

3G0, 367-374. 378, 215, 222, 357,

35S, 370, 372.
armata, Testudo, 251.
Aspideretes, 422.
Aspidochelys, 452.
Aspidochelys livingstonii, 452, 459,

460.
Aspidonectes, 420.
Aspidonectes, 424.
Aspidonectes aspilus, 425.
Aspilus, 421.

aspilus, Aspidonectes, 425.
aspilus, Gymnopus, 426.
Asterochelys, 223.
Asterochelys yniphora, 224.
Astrorchis renicapite, 502.
atlantica, Dermochelis, 500.
astra, Caretta, 490.
atra, Halichelys, 489,
aubryi, Cryptopodus, 453.
aubryi, Cryptopus, 452.
aubryi, Cycloderma, 168, 417-420,

453-459, 460, 468, 456, 458.
aubryi, Heptathyra, 453.

1- Misprints such as Cinicys, Cininixya, Homoeopus, Tisfudo. etc., are not
lucluded. The pagination of a valid name's main discussion is indicated by
boldface type. All synonyms are indexed. Pages on which iUustrations ami
maps occur are cited in italics after the other page references.

rXDEX

543

austrnlis, Kinixys, 387.
nustralia, Kinixys a., 388.
hulH'Ocki, (lOocholono p., 167-ir)9,

227-231, 235-251, 252, 299, 360,

213, 214, 217, 226, 234, 239, 257,

267.
Uaikia, 436.
Baikiea, 436.

Baikica eleg-ans, 436-438, 444.
l)aikii, Tctratliyni, 438, 439, 443,

444.
healei, Cistuda, 191.
Bellcmy.s, 218.
l)ellian:t, Cinixys, 286, 287, 38."),

390.
belliana, Cinothorax, 385.
holliana. Kinixys, 166, 374, 377-

3S0, 383. 384, 388, 396, 404.
I.oliinna. Kinixys b., 167, 287, 289,

292, 321, 369, 376, 383, 384-396.

397, 215, 219, 379, 389.
bellii, Testudo, 346.
bergeii, Chersinella v., 327.
bergeri, Homopus, 289, 296, 299,

301, 309, 310, 326, 330, 331, 358,

362.
bergeri, Testudo, 326.
bergeri, Testudo v., 327..
Iiergiana, Eragrostris, 341.
bettai, Testudo g., 265.
bibronii, Pelochelys, 415.
bicarinata, Chelonia, 474.
liicin<'tum, Cinetisternum, 422.
bigultata, Testudo, 229, 250, 251.
bipum-tata, Testudo, 229, 251.
bissa, Caretta, 485, 488.
blandingii, Cistuda, 189.
blandingii, Eniydoidea, 167, 188,

189.
blandingii, Emys, 184.
boettgeri, Chersinella, 327.
lioettgeri, Chersinella v., 327.
boettgeri, Testudo, 299, 301 326,

327, 329, 330, 332.
boulengeri Clicrsobius, 362.
boulengeri, Homopus, 168, 326, 354-

359, 362-365, 378, 215, 219, 3.57.

358, 363.
Iioulengeri. I^soudoiiiopns, 3()2.
Bulbine caulescens, 367.
Caelatura nyasaensis, 466.
raf ra, Testudo, 360.
cafra, Testudo c, 3(50.
calcarata, Cheraine, 231.
calcarata, Testudo, 228. 231, 2'',S,

241.
Calemys, 191.
Callagur, 187.
Callinia, 421.

rallosa, Cyclanosteus n., 438, 444.
Camallanus microeeplialus, 199.
campanulata, Testudo, 264, 277.
Caouana, 489.
Caouana caretta, 491.
caouana, Chelonia, 491.
Caouana dessumierii, 495.
Caouana elongata, 491.
Caouana olivacea, 495.
Caouana ruppellii, 495.
caouana, Testudo, 490.
caouana, Thalassochelys, 489.
Caretta, 469, 470, 484, 489-490.
Caretta atra, 490.
Caretta bissa, 485, 488.
Caretta caretta, 471, 490-494.
Caretta c. caretta, 492.
Caretta e. gigas, 492.
Caretta c. olivacea, 496.
Caretta esculenta, 474.
Caretta gigas, 492, 493.
Caretta imbricata, 485.
Caretta nasicornis, 474.
Caretta nasuta, 489, 490.
Caretta olivacea, 495.
Caretta reniivaga, 495.
Caretta rostrata, 486.

544

INDEX

Caretta squamosa, 485.
Caretta thunbergii, 474.
caretta, Caouana, 491.
caretta, Caretta, 471, 490-494.
caretta, Caretta c, 492.
caretta, Testudo, 470, 489, 490.
caretta, Thalassoehelys, 491, 495.
cariniferus, Trionyx, 421.
Carteremys, 277.
cartilaginea, Testudo, 421.
cartilagineus, Trionyx, 415.
caspica, Clemmys, 192, 195.
caspica, Emys, 193.
caspica, Testudo, 191.
castanea, Kinixys, 374, 401, 405.
caulescens, Bulbine, 367.
Centrochelys, 221.
Centrochelys (P.) sulcatus, 231.
cepediana, Testudo, 474.
cephalo, Testudo, 489, 490.
Cephalochelys, 489.
Cephaloehelys oceanica, 489, 491.
Ceramopelta, 422.
Ceramopelta latirostris, 422.
Chamaerops liumilis, 275.
Cheirogaster, 219.
Chelona, 473.
Chelone, 473.
Chelone imbricata, 486.
Chelone midas, 476.
Chelone mydas, 475.
Chelone virgata, 476.
Chelone viridis, 475.
Chelonia, 472-473.
Chelonia agassizii, 475.
Chelonia bicarinata, 474.
Chelonia caouana, 491.
Chelonia corticata, 491.
Chelonia depressa, 469, 475.
Chelonia dubia, 495,
Chelonia dussumieri, 495.
Chelonia formosa, 475.
Chelonia imbricata, 485.

Chelonia lachrymata, 474.
Chelonia lata, 475.
Chelonia lutaria, 499.
Chelonia maculosa, 474.
Chelonia marmorata, 474.
Chelonia midas, 475.
Chelonia multiscutata, 495.
Chelonia mydas, 469, 471, 474-485.
Chelonia m. japonica, 476.
Chelonia m. mydas, 476.
Chelonia olivaeea, 494, 495.
Chelonia pelasgorum, 490.
Chelonia polyaspis, 495.
Chelonia pseudocaretta, 485.
Chelonia pseudomydas, 485.
Chelonia tenuis, 475.
Chelonia virgata, 474.
Chelonia viridis, 474.
Cheloniidae, 180, 468, 472.
Chelonoidis, 223.
Chelopus, 191.
Chelydra, 189.
Chelyra, 499.
Chersina, 218, 220, 255, 259, 296,

297, 342-345, 346, 377, 378.
Chersina angulata, 167, 344, 345-

352, 212, 214, 217, 344, 347, 349.
Chersine, 254, 346.
Chersine angulata, 346.
Chersine areolata, 368.
Chersine calarata, 231.
Chersine geometrica, 321.
Chersine pusilla, 345.
Chersine signata, 360.
Chersine tetradactyla, 368.
Chersinella, 254, 294, 299.
Chersinella boettgeri, 327.
Chersinella fiski, 326, 327.
Chersinella f. colesbergensis, 301,

326.
Chersinella f. eronwrighti, 301, 326.
Chersinella f. grica, 301, 327.
Chersinella f . gricoides, 301, 327.

INDEX

545

Chersinella f. orangeusis, 301, 32G.
Chersinella f. seimundi, 326.
Chersinella geometrica, 321.
Chersinella graeca, 264.
Chersinella oculifera, 316.
Chersinella schonlandi, 301, 307,

309, 310, 311, 326, 329.
Chersinella strauchi, 321.
Chersinella tentoria, 337.
Chersinella t. albanica, 301, 337,

339.
Chersinella t. duerdeni, 301, 337.
Chersinella t. hexensis, 301, 309,

333, 338.
Chersinella t. karuella, 301, 337,

339.
Chersinella t. karuica, 301, 337.
Chersinella t. lativittata, 301, 337.

Chersinella t. piscatella, 301, 337.

Chersinella t. subsulcata, 301, 337.

Chersinella t. tentorioides, 301, 337.

Chersinella trimeni, 333.

Chersinella verroxii, 327.

Chersinella v. amasensis, 301, 327,

Chersinella v. bergeri, 327.

Chersinella v. boettgeri, 327.

Chersinella v. smithi, 327.

•:;hersobius, 352, 355, 360.

Chersobius boulengeri, 362.

Chersobius signatus, 360.

Chersus, 254.

Chersus mauritanicus, 264, 276.

Chilensis, Geochelone, 235.

Chitra, 414, 417.

Chitra indica, 415.

Chrysemys, 187.

Cinctisternum, 422.

Cinctisternum bicinctum, 422.

cinerea, Ardea c., 480.

Cinixys, 374.

Cinixys belliaua, 286, 287, 385, 396.

Cinixys dorri, 397.

Cinixys homeana, 400.

Cinixys lobatsiana, 387, 389, 394.

Cinixys nogueyi, 397.

Cinothorax, 374, 378, 379.

Cinothorax belliana, 385.

Cissus, 248.

Cistuda bealei, 191.

Cistuda blandingii, 189.

Cistuda hellenica, 203.

Cistudo europaea, 203.

Cistudo lutaria, 203.

Clemmys, 187, 189, 191, 205, 209.

Clemmj's caspica, 192, 195.

Clemmys c. caspica, 192.

Clemmys c. cretica, 192.

Clemmys c. leprosa, 190, 192-201,

204, 170, 190, 196, 212, 214, 217.
Clemmys c. rivulata, 192.
Clemmys c. sigriz, 195.
Clemmys fuliginosa, 194.
Clemmys laticeps, 193.
Clemmys leprosa, 193.
Clemmys marmorea, 193.
Clemmys oculifera, 316.
Clemmys sigriz, 193.
Coelognathus, 422.
Coelognathus novemcostatus, 422.
colesbergensis, Chersinella f., 301,

326.
colesbergensis, Psammo bates f.,

327.
Colpochelys, 494.
Colpochelys kempi, 494.
comptoni, Eniys, 353, 354.
eomptoni, Testudo, 218, 354.
contortus, Spiroxys, 199.
controversa, Thalassochelys, 495.
Coptopelta, 422.
Coptopelta septemcostata, 422.
coriacea, Dermatochelys, 500.
coriacea, Dermochelis, 500.
coriacea, Dermochelys, 469, 471,

499-503.
coriacea, Dermochelys c, 500.

546

IN'DEX

coriacea, Sphargis, 500.
coriacea, Testurlo, 499.
Coriuao, 499.
cortieatn, Qielonia, 491.
corticata, Thalassochelys, 491, 492.
costaricensis, Geocheloue, 224.
Crassulaceae, 331.
cretica, Clenimys c, 192.
cronwrighti, Chersinella f., oOl,

326.
(Tonwrighti, Psaiuniohates f., 327.
Crrptodira, 180.
Cryptopodus, 452.
Cryptopodus aubryi, 453.
Cryptopus, 452.
Cryptopiis aubryi, 452.
Cryptopus senegalensis, 443.
Cuora, 188.
Cyclanorbis, 414, 417, 420, 435, 436-

438, 451, 452.
Cyclanorbis elegans, 167. 417-420,

436, 437, 438-443, 444, 449, 419.

440, 441.
Cyclanorbis oligotylus, 167, 437,

438, 443.
Cyclanorbis petersii, 436, 443.
Cyclanorbis senegalensis, 417-420,

436-439, 443-450, 451, 419, 445,

447.
Cyclanorbis sp., 450-451.
Cyclanosteus, 436.
Cyclanosteus frenatus, 459.
Cyclanosteus petersii, 436, 443.
Cyclanosteus senegalensis, 438, 443.
Cyclanosteus s. callosa, 438, 444.
Cyclanosteus s. equilifera, 444.
Cyclanosteus s. normalis, 444.
Cyclemys, 188.
Cyclemys mouhoti, 183.
Cycloderma, 166, 414, 417, 420, 436,

451-453, 467.
Cycloderma aubryi, 168, 417-420,

453-459, 460, 468, 456, 458.

Cycloderma frenatum, 173, 417 420.

451-4.53, 455, 459-468, 461, 46.1.
Cycloderma petersii, 44.T.
Cycloderma senegalense, 443.
Cylindraspis, 223, 224.
Cypraea moneta, 323.
Danionia, 187.

darlingi, Honiopus, 296, 387.
darlingi, Kinixys, 387.
Deirochelys, 187, 188.
Deirochelys reticularia. 184, 188,

189.
denticulata, Geochelone, 255.
denticulata, Kinixys, 407.
denticulata, Testudo, 404.
depressa, Chelonia, 469, 475.
depressa, Gari, 494.
depressa, Psammobates, 301, 327,

329, 331.
Dermatochelys, 499.
Dermatochelys coriacea, 500.
Dermatochelys porcata, 500.
Dermochelis, 499.
Dermochelis atlantica, 500.
Dermochelis coriacea, 500.
Dermochelyidae, 180, 468, 498.
Dermochelys, 499.
Dermochelys coriacea, 469, 471,

499-503.
Dermochelys c. coriacea, 500.
Dermochelys c. schlegelii, 500.
dessumierii, Caouana, 495.
Diosnia, 323.
Diosma pulchella, 323.
Dogania, 421, 422.
Dogania subplana, 415.
dorri, Cinixys, 397.
dubia, Chelonia, 495.
ductor, Echeneis, 502.
duerdeni, Chersinella t., 301, 337.
duerdeni, Psammobates t., 338.
dussumieri, Chelonia, 495.
Echeneis, ductor, 502.

TXDEX

547

Echeneis iiaiierates, 48 1 .
Echweria, 273.

elegans, Baikiea, 436-438, 444.
elegans, Oyclaiioibis, 167, 417-420,

436, 437, 438-443, 444, 449, 419,

440, 441.
elegans, Geochelone, 226, 255, 299.
elegans, Testudo, 221, 225.
ellipticus, Lanistes, 466.
elongata, Cauoana, 491.
elongata Eremonia, 489.
Emmenia, 191.
emoryi, Trionyx, 417.
Emyda senegalensis, 443.
Emydinae, 182, 183-189, 190, 37."),

185.
Emydoidea, 167, 187, 188, 189, 299.
Emydoidea blandingii, 167, 188,

189.
Emys, 167, 184, 188, 201-202, 299.
Emys antiguorum, 202.
Emys blandingii, 184.
Emys caspica, 193.
Emys caspia var. leprosa, 195.
Emys comptoni, 353, 354.
Emys europaea, 203.
Emys flavipes, 195.
Emys fraseri, 195.
Emys fuliginosus, 191, 193.
Emys grayi, 191.
Emys hellenica, 203.
Emys hofmani, 203.
Emys iberica, 203.
Emys lamaria, 195.
Emys laniaria, 195.
Emys laticeps, 191, 193.
Emys leprosa, 191, 192.
Emys lutaria, 193, 203, 204.
Emys marmorea, 192.
Emys oculifera, 316.
Emys orbicularis, 167, 170, 184.

188, 191, 201, 202-209, 190, 205.

206, 212, 214, 217,

Emys o. aralensis, 204.

Emys 0. hellenica, 204.

Emys 0. orbicularis, 204.

Emys scutella, 353.

Emys sigriz, 193.

Emys vulgaris, 193.

emys, Geochelone, 224, 255.

equlifera, Cyclanosteus s., 444.

Eragrostris bergiana, 341.

Eremonia, 489.

Eremonia elongata, 489.

Eretmochelys, 484-485.

Eretmochelys imbricata, 469, 471,

485-489.
Eretmochelys i. imbricata, 486.
Eretmochelys i. squamosa, 486.
Eretmochelys squamata, 485.
erosa, Kinixys, 230, 377-381, 383,

384, 403, 404-411, 215, 217, 257,

377, 379, 3S0, SSI, 408, 409.
erosa, Testudo, 374, 404.
Eryma, 191.
Eryma laticeps, 195.
esculenta, Caretta, 474.
Euamyda, 422.
Euchelonia, 473.
Euchelys, 473.

Euchelys macropus, 473, 477.
euphratica., Testudo, 421.
euphraticus, Trionyx, 417.
europaea, Cistudo, 203.
europaea, Emys, 203.
europaea, Lutremys, 203.
europaea, Testudo, 201, 202.
exornatum, Amblyomma, 248.
Falcaustra lambdiensis, 199.
fasciata, Testudo, 369.
femoralis, Homopus, 168, 355-359,

365-387, 378, 357, 358.
ferox, Testudo, 421.
ferox, Trionyx, 417.
fiski, Chersinella, 326.
nski. Testudo, 295, 301, 325, 326-

548

INDEX

329, 331.
fiskii Psammobates, 327.
fiskii, Psammobates f., 327.
fiskii, Testudo t., 326, 333.
flava, Testudo, 202.
flavipes Emys, 195.
floweri, Testudo g., 259, 260.
Fordia, 421.

Fordia africana, 421, 425.
formosa, Chelonia, 475.
formosus, Trionyx, 415, 421.
fraseri, Emys, 195.
Fregatta m. aldabrensis, 480.
frenata, Heptathyra, 454, 459.
frenatum, Cycloderma, 173, 417-

420, 451-453, 455, 459-468, 461,

463.
frenatus, Cyclanosteus, 459.
fuliginosa, Clemmys, 194.
fuliginosa, Mauremys, 194.
fuliginosus, Emys, 191, 193.
gabonensis, Pelusios, 453.
gangeticus, Trionyx, 415, 422.
Gari depressa, 494.
Geochelone, 165, 218, 220, 221-225,

225-230, 238, 255, 256, 284, 293-

295, 299, 342, 343, 344, 354, 355,

378.
Geochelone costaricensis, 224.
Geochelone denticulata, 255.
Geochelone elegans, 226, 255, 299.
Geochelone emys, 224, 255.
Geochelone, gigantea, 225, 255, 295.
Geochelone grandidieri, 224, 225.
Geochelone kaiseni, 224.
Geochelone nana, 224.
Geochelone pardalis, 224, 227, 238,

251, 254, 255.
Geochelone p. babcocki, 167-169,

225, 227-231, 234, 235-251, 252,

254, 299, 360, 212 214, 217, 226,

234, 239, 257, 267.
Geochelone p. pardalis, 227-230,

250, 251-254.
Geochelone platynota, 226.
Geochelone radiata, 224.
Geochelone Senegal ensis, 231.
Geochelone sulcata, 168, 225-229,

230-235, 241, 247, 369, 226, 232.
Geochelone sumeiri, 225, 259.
Geochelone uintensis, 224.
Geochelone yniphora, 342.
Geoclemmys, 191.
Geoemyda, 187.
Geoemyda punctularia, 350.
geographicus, Peltastes, 321.
geometrica, Chersine, 321.
geometrica, Chersinella, 321.
geometrica, Testudo, 173, 201, 294,

299, 301, 316, 319, 323, 337, 387.
geometricus group, 295, 296, 298,

299, 301, 303, 326.
geometricus Peltastes, 321.
geometricus, Psammobates, 168,

174, 253, 291, 300-303, 314, 316,

319-324, 313.
gigantea, Aldabrachelys, 225.
gigantea, Geochelone, 225, 255, 295.
gigas, Caretta, 492, 493.
gigas, Caretta c, 492.
Glossiphonidae, 466.
Glyptemys, 191.
Gomphopelta, 421.
Gomphopelta officinalis, 421.
Goniochersus, 342.
Goniochersus angulatus, 346.
Gopherus, 218, 255, 256, 295, 343,

355.
Gopherus agassizii, 186.
Gopherus polyphemus, 283.
graeca, Chersinella, 264.
graeea, Testudo, 170, 218, 254-256,

258, 259, 261, 276, 282, 298, 299.
graeca, Testudo g., 259, 260, 261-

276, 212, 214, 217, 257, 266, 267.
grandidieri, Aldabrachelys, 225.

INDEX

549

grandidieri, Geochelone, 224, 225.
Graptemys, 187.
<,'rayi, Emys, 191.
grica, Chersinella f ., 301, 327.
gricoides, Chersinella f., 301, 327.
guttata, Testudo, 191.
Gymnopus, 420.
Gymnopus aegyptiacus, 424.
Gymnopus aspilus, 426.
Hadrianus, 218, 222-224, 355,
Halic'helys, 489.
Halichelys atra, 489.
hellenica, Cistuda, 203.
hellenica, Emys, 203.
hellenica, Emys o., 204.
Heptathyra, 452.
Heptathyra aubryi, 453.
Heptathyra frenata, 454, 459.
Heptathyra livingstonii, 459.
hermanni, Testudo, 170, 258, 260,

265, 281, 212, 257, 281.
Herpysmostes, 484.
Hesperotestudo, 223, 224.
hexensis, Chersinella t., 301, 309,

333, 338.
Hievemys, 298.
hofmani, Emys, 203.
homeana, Cinixys, 400, 404.
homeana, Kinixys, 377-381, 383,

384, 399, 400-404, 215, 379, 380,

381, 401.
Homopus, 218, 221, 255, 259, 284,
295-297, 328, 343, 344, 352-359, 360,

377, 378.
Homopus areolatus 173, 353, 355-

360, 367-374, 378, 215, 222, 357,

358, 370, 372.

Homopus bergeri, 289, 296, 299,
301, 309, 310, 326, 330, 331, 358,
362.

Homopus boulengeri, 168, 326, 354-

359, 362-365, 378, 215, 219, 357,
358, 363.

Homopus darliugi, 296, 387.
Homopus femoralis, 168, 355-359,

365-367, :578, 357, 358.
Homopus nogueyi, 296, 397.
Homopus signatus, 168, 173, 237,

316, 353-357, 359-362, 369, 378,

217, 357, 358.
horsfieldi, Testudo, 254, 258, 283.
humilis, Chamaerops, 275.
hurum, Trionyx, 415.
Hyalomma aegyptiacum, 273.
Hyalomma syriacum, 273.
Hydrone, 201.

ibera, Testudo, 170, 262, 265.
ibera, Testudo g., 259, 265.
iberica Emys, 203.
Ida, 421.

imbricata, Caretta, 485.
imbricata, Chelone, 486.
imbricata, Chelonia, 485.
imbricata, Eretmochelys, 469, 471,

485-489.
imbricata, Eretmochelys i., 486.
imbricata, Testudo, 484, 485.
indiea, Chitra, 415.
Indotestudo, 223, 224.
insculpta Clemmys, 186.
insculpta, Testudo, 191.
irrorata, Landemania, 421.
Isola, 421.

jaegerskioeldi, Placobdella, 466.
japonica, Chelonia m., 476.
japonica, Testudo, 474.
javanicus, Trionyx, 421.
jordani, Kinixys, 387.
jujuba, Zizyphus, 275.
kaiseni, Geochelone, 224.
karuella, Chersinella t., 301, 337,

339.
karruica, Psammobates t., 338.
karuica, Chersinella t., 301, 337.
kempi, Colpochelys, 494.
kempi, Lepidochelys o., 496, 497.

550

INiDEX

Kinixys, 166, 218, 220, 259, 288,

296, 342-344, 374-384.
Kinixj's australis, 387.
Kinixys a. australis, 388.
Kinixys a. mababiensis, 388.
Kinixys belliana, 166, 374, 377-380,

383, 384, 388, 396, 404.
Kinixys b. belliana, 167, 287, 289,

292, 321, 368, 369, 376, 383, 384-
396, 397, 215, 219, 379, 3S9.

Kinixys V). nogueyi, 166, 383, 388,
396-399, 380, 3S1.

Kinixys b. spekei, 388.

Kinixys b. zombensis, 387.

Kinixys b. zuluensis, 387.

Kinixys eastanea, 374, 401, 405.

Kinixys dailingi, 387.

Kinixys dontieulata, 407.

Kinixys erosa, 230, 377-381, 383,

384, 403, 404-411, 215, 217, 257,
377, 379, 380, 381, 408, 409.

Kinixys honieana, 377-381, 383, 384,

399, 400-404, 215, 379, 380, 381,

401.
Kinixys jordani, 387.
Kinixys lobatsiana, 388, 389.
Kinixys natalensis, 388, 390.
Kinixys nogueyi, 397.
Kinixys sehoensis, 387.
Kinixys spekii, 379, 380, 387, 391.
Kinixys youngi, 388.
Kinixys zombensis, 388.
Kinixys zuluensis, 388.
Kinotliorax, 374.
kleinmanni, Pseudotestudo, 276.
kleinmanni, Testudo, 218, 220, 258-

261, 264, 276-283, 212, 214, 278,

279, 280.
kraussi, Onyehochelys, 484, 486.
Inbiatus, Trionyx, 424, 427, 433,

434.
Lacerta, 275.
lachrymata, Chelonia, 474.

laevis, Psilognathus, 421.
lamaria, Emys, 195.
lambdiensis, Faleaustra, 199.
Landemania, 421.
Landeniania irrorata, 421.
laniaria, Emys, 195.
laniaiia, Maurcmys, 194.
Lanistes ellipticus, 466.
Lanistes sordidus, 466.
lata, Chelonia, 475.
laticeps, Clenimys, 193.
laticeps, Emys, 191, 193.
laticeps Eryma, 195.
latirostris, Ceramopelta, 422.
lativittata, Chersinella t., 301, 337.
leithii, Peltastes, 277.
leithii, Testudo, 218, 276, 277, 280.
leithii, Trionyx, 415.
Lepidochelys, 469, 470, 494-495.
Lepidochelys olivaeea, 494.
Lepidochelys o. kempi, 496, 497.
Lepidochelys o. olivaeea, 471, 495-

498.
leprosa, Clemmys, 193.
leprosa, Clemmys c, 190, 192-201,

204, 170, 190, 196, 212, 214, 217.
leprosa, Emys, 191, 192.
leprosa, Emys c, 195.
leprosus, Yuen, 421.
Limonium, 282.
Lissemys, 412, 414, 415, 417.
livingstonii, Aspidochelys, 452, 459,

460.
livingstonii, Heptathyra, 459.
lobatsiana, Cinixys, 387, 389, 394.
lobatsiana, Kinixys, 388.
Lorica t. signatae, 359.
loveridgii, Testudo, 287-290, 292,

387.
lutaria, Chelonia, 499.
utaria, Cistudo, 203.
lutaria, Emys, 193, 203, 204.
lutaria, Testudo, 201, 202.

INDEX

551

12:20,
843,

172,
258,

lutcola, Testudo. 253, 299, 301, 321. 1

323.
Lutremys, 201.
Lutreniys europaea, 203.
lyra, Testudo, 499.
iiiahabiensis, Kinixys a., 388.
macropus, Eiu-helys, 473, 477.
maeropus, Testudo, 474.
maeulosa, Chelonia, 474.
Malac'leinys, 187.
Malacoehorsus, 165, 1(56, 219,

255, 257, 283-286, 289, 296,

378, 394.
Malacochersus procterae, 287, .
Malacoehorsus tornieri, 167,

284, 286-294, 378, 388, SIS,

217, 257, 285, 288.
Manouria, 223, 224.
marginata, Testudo, 173, 254,

261, 262, 264, 276, 282.
marina, Testudo, 474.
marmorata, Chelonia, 474.
marniorea, Clemmys, 193.
marmorea, Emys, 192.
marmoreum, Amblyomma, 248,

393.
Mauieniys, 191.
Mauremys fuliginosa, 194.
Maureniys laniaria, 194.
niauritanica, Testudo, 262.
niauritanica, Testudo g., 264.
mauritanieus, Chersus, 276.
Medaesiia, 2.'4.
Megaeheisine, 221.
Megaehersine pardalis, 238.
Megalochelys, 223, 224.
Megeniys, 473.
Mflanemys, 191.
meleagris, Testudo, 202.
mercuralis, Sphargis, 499.
Mesoniln-yanthemum, 332.
micvocephalus, Camellanus, 199
inidas, Cholone, 476.

293,

niidas, Chelonia, 475.
ininiata, Testudo, 368, 369.
ininuta, Testudo, 323.
Monachelys, 223.
nioneta, Cypraea, 323.
mordax, Temnognathus, 421.
Morcnia, 187.
inoitoni, Trionyx, 425.
moubata, Ornithodorus, 332.
niouhoti, Cycleinys, 183.
mouhoti, Pyxidea, 183.
niuhlenbergii, Testudo, 191.
iimltisc'utata, Chelonia, 495.
.Mutela alata,466.
mutica, Aniyda, 422.
niuticus, Trionyx, 417.
Mydas, 473.
Mydas viridis, 475.
mydas, Chelone, 475.
mydas, Chelonia, 469, 471, 474-485.
mydas, Chelonia m., 476.
mydas, Testudo, 473, 474.
Mydasea, 473.
nana, Geoclielone, 224.
Xanemys, 191.
iiasieornis, Caretta, 474.
nasicornis, Testudo, 490.
nasuta, Caretta, 489, 490.
uatalensis, Kinixys, 388, 390.
Natator tessellatus, 475.
uaucrates, Eeheneis, 481.
nebrascensis, Stylemys, 256.
Xeoemys, 189.
Xeotestudo, 342.
Xeotestudo angulata, 346.
uiger, Pelusios, 453.
nigricans, Trionyx, 415.
nigrita, Xylocopa, 392.
nigriventris, Testudo g., 301, 337.
nilotica, Tyrse, 425.
niloticus, Trionyx, 421, 424.
Xilssonia, 421.
nogueyi, Cinixys, 397.

552

INDEX

nogueyi, Homopus, 296, 397.

nogueyi, Kinixys, 397.

nogueyi, Kinixys b., 166, 383, 388,

396-399, 3S0, 381.
normalis, Cyelanosteus s., 444.
Notoehelys, 188.

novemcostatum, Tortisternum, 422.
novemcostatus, Coelognathus, 422.
nuttalli, Amblyomma, 248, 293,

393.
nyassensis, Caelatura, 466.
Oeadia, 186, 187, 188.
oceanica, Cephalochelys, 489, 491.
oculifer, Psammobates, 168, 300-

303, 314, 315-319, 321, 329, 332,

360, 362, 813, 317.
oculifera, Cliersinella, 316.
oculifeia, Clemmys, 316.
oculifera, Emys, 316.
oculifera, Psammobates, 316.
oculifera, Testudo, 299, 301, 31.5,

326.
officinalis Gomphopelta, 421.
oligotylus, Cyclanorbis, 167, 437,

438, 443.
olivacea, Caouana, 495.
olivacea, Caretta, 495.
olivacea, Caretta c, 496.
olivacea, Chelonia, 494, 495.
olivacea, Lepidochelys, 494.
olivacea, Lepidochelys o., 471, 495-

498.
olivacea, Thalassoehelys, 495.
Onychochelys, 484.
Onychochelys kraussi, 484, 486.
Ophisops (as Ophiops), 275.
orangensis, Chersinella f., 301, 326.
orbicularis, Emys, 167, 170, 184,

188, 189, 191, 201, 202-209, 190,

205, 206, 212, 214, 217.
orbicularis, Emys o., 204.
orbicularis, Testudo, 201, 202.
Orlitia, 187.

ornata, Trionyx, 421.
Ornithodorus moubata, 332.
oscarboettgeri, Testudo, 301, 326,

329.
Oscaria, 421.
OscT-ia swinhoei, 421.
Oxvuridae, 332.
pallida, Testudo a., 368.
pardalis, Geochelone, 224, 227, 238,

251, 254, 255.
pardalis, Geochelone p., 228-230,

250, 251-254.
pardalis, Megachersine, 238.
pardalis, Testudo, 221, 235, 251.
pardalis, Testudo p., 238, 251.
peersi, Pseudomopus s., 360.
peguensis, Trionyx, 421.
pelasgorum, Chelonia, 490.
Pelochelys, 414.
Pelochelys bibronii, 415.
Pelodiscus, 421.
Pelodiscus triunguis, 426.
Peltastes, 254.
Peltastes geographicus, 321.
Peltastes geometricus, 321.
Peltastes leithii, 277.
Peltastes m. whitei, 264.
Peltastes semiserratus, 316.
Peltastes sulcatus, 231.
Peltastes tentorius, 337.
Peltastes verreauxii, 324.
Peltastes verroxii, 324.
Peltonia, 254.
Pelusios gabouensis, 453.
Pelusios niger, 453.
Pemphis ocidula, 481.
petersii, Cyclanorbis, 436, 443.
petersii, Cyelanosteus, 436, 443.
petersii, Cyelodernia, 443.
piscatella, Chersinella t., 301, 337.
piseatella, Psammobates t., 337.
Placobdella jaegerskioeldi, 466.
planicauda, Acinixys, 217

INDEX

553

platynota, Geochelone, 226.
Platypeltis, 421.
Platysterninae, 181, 182.
Platysternon, 181, 298.
Pleurodira, 163, 417.
polyaspis, Chelonia, 495.
polyphemus, Gopherus, 283.
porcata, Dermatochelys, 500.
Potamoehelys, 421.
procterae, Malaeochersus, 287, 285.
procterae, Testudo, 287, 288, 289,

292.
Psammobates, 166, 218, 220, 221,

255. 2.19, 294-315, 343, 344, 378.
Psammobates depressa, 301, 327,

329, 331.
Psammobates fiskii, 327.
Psammobates f. colesbergensis, 327.
Psammobates f. cronwrighti, 327.
Psammobates f. fiskii, 327.
Psammobates geometricus, 168,

174, 253, 291, 300-303, 314, 316,

319-324, 313.
Psammobates oculifer, 168, 300-

303, 314, 315-319, 321, 329, 332.

360, 362, 313, 317.
Psammobates oculifera, 316.
Psammobates tentoria, 337.
Psammobates tentorius, 300, 306,

308.
Psauunobates t. albaniea, 337.
Psammobates t. duerdeni, 338.
Psammobates t. karruica, 338.
Psammobates t. piscatella, 337.
Psammobates t. tentorioides, 337.
Psanmiobates t. tentorius, 301, 302,

309, 312, 313, 315, 316, 321, 327,

336-342, 212, 215, 217, 339, 340.
Psammobates t. trimeni, 301, 302,

309-311, 313, 315, 327, 330, 333-

336, 338, 342, 212.
Psammobates trimeni, 333.
Psammobates t. verroxii, 289, 295,

301-303, 313, 314, 316, 324-333,

336, 337, 338, 341, 342, 358,

362, 215, 222, 313, 329.
Psammobia vespertina, 494.
Psammodromus, 275,
Pseudemys, 186, 187, 189.
pseudocaretta, Chelonia, 485.
Pseudomopus, 358, 355.
Pseudomopus boulengeri, 362.
Pseudomopus signatus, 360.
Pseudomopus s. peersi, 360.
pseudomydas, Chelonia, 485.
Pseudotestudo, 166, 218, 258, 276
Pseudotestudo kleinmanni, 276.
Psilognathus, 421.
Psilognathus laevis, 421.
pulchella, Diosma, 323.
pulchella, Testudo, 202.
punctata, Testudo, 191.
punctularia, Geoemyda, 350.
pusilla, Chersine, 345.
pusilla, Testudo, 262, 265, 345, 346,

368.
Pyxis, 218, 256, 259, 296, 297, 343,

376, 377.
Pyxis arachnoides, 259, 364, 217.
radiata, Geochelone, 224, 255.
Rafetus, 421.
remivaga, Caretta, 495.
Remora remora, 480, 502.
remora, Remora, 480, 502.
renicapite, Astrorchis, 501.
retieularia, Deirochelys, 184, 188,

189.
rivulata, Clemmys c, 192.
rostrata, Caretta, 486.
rotunda, Testudo, 202.
rodolfianus, Trionyx t., 426.
ruppellii, Caouana, 495.
Saealia, 191.

Sagittarius serpentarius, 351, 373.
Salsola, 282.
schlegelii, Dermochelys c, 500.

554

INDEX

schlegelii, Sphargis c, 500.
schoensis, Kinixys, 387.
sehoepfi, Testudo, 230, 405.
schonlandi, Chersinella, 301, 307,

309, 310, 311, 312, 326, 329.
seutella, Emys, 353.
scutella Testudo, 354.
Sedum, 273.

seimundi, Chersinella f., 326.
seimundi, Testudo, 301, 326, 329,

330.
semiserrata, Testudo, 299, 301, 316,

319.
semiserratus, Peltastes, 316.
senegalensis, Cryptopus, 443.
scnegalensis, Cyclauorbis, 417-420,

436-439, 443-450, 451, 419, 445,

447.
senegalensis, Cyc lanes teus, 438,

443.
senegalense, Cycloderma, 443.
senegalensis, Emyda, 443.
senegalensis, Geochelone, 231.
senegalensis, Testudo r., 231.
septemcostata, Coptopelta, 422.
serpentarius, Sagittarius, 351, 373.
Seytina, 499.
Siebenrockiella, 187, 298.
signata, Chersine, 360.
signata, Testudo, 352, 353, 359.
signata Testudo C, 360.
signatae, Lorica t., 359.
signatus, Chersobius, 360.
signatus, Homopus, 168, 173, 237,

316, 353-357, 359-362, 369, 378,

217, 357, 358.
signatus, Pseudomopus, 360.
signatus, Testudo, 369.
sigriz, Clemmys, 193.
sigriz, Clemmys c, 195.
sigriz, Emys, 193.
sigriz, Terrapene, 193.
simpsoni, Mutela, 466.

Sinjispideretes, 412.

sinensis, Trionyx, 415. 417, 41.M ,

422, 427.
Sinohadrianus, 219.
smithi, Chersinella v., 327.
smithi, Testudo, 299, 301, 325, 327,

329, 330, 333.
sordidus, Lanistes, 466.
spekei, Kinixys b., 388.
spekii, Kinixys, 379, 380, 387, 391.
Sphargis, 499.
Sphargis angusta, 500.
Sphargis eoriacea, 500.
Sphargis c. schlegelii, 500.
Sphargis mereuralis, 499.
spinifer, Trionyx, 417, 421.
Spiroxys contortus, 199.
squamata, p]retmochelys, 485.
squamosa, Caretta, 485.
squamosa, Eretmoehelys i., 486.
steindachneri, Trionyx, 417.
stellata, Testudo, 221, 225.
Stigmochelys, 221.
strauehi, Chersinella, 321.
strauchi, Testudo, 299, 301, 302,

321-323.
Stylemys 218, 255, 256, 343, 355.
Stylemys nebrascensis, 256.
subplana, Amyda, 422.
subplana, Dogania, 415.
subplanus, Trionyx, 415, 421, 427.
subsulcata, Chersinella t. 301 337.
sulcata, Geochelone, 168, 225-229,

230-235, 241, 247, 369, S26, S32.
sulcata, Testudo, 221, 230, 232, 237.
sulcatus, Centrochelys (P.), 231.
sulcatus, Peltastes, 231.
sumeiri, Aldabrachelys, 225.
sumeiri, Geochelone, 225, 259.
swiuhoei, Oscaria, 421.
swinhoei, Trionyx, 417.
syriacum, Hyalomma, 273.
tarapacana, Thalassochelys, 491.

INDEX

555

tarapacona, Thalassiochelj's, 491.
Temnognathus, 421.
ToDinognathiis niordax, 421.
tentoria, Chersinella, 337.
tentoria, Psaniiiiobates, 337.
tentoria, Testudo, 301, 308, 309,

32d, 33(5, 338, 339.
tentoria, Testudo g., 301, 321.
tentorioides, Cliersinella t., 301,

337.
tentorioides, Psammobates t., 337.
tentorius complex, 309.
tentorius, Peltastes, 337.
tentorius, Psammobates, 306, 308.
tentorius, Psammobates t., 301,

302, 309, 312, 313, 315, 316, 321,

327, 336-342. 212, 215, 217, 339,

340.
tenuis, Chelonia, 475.
Terrapene, 183, 188.
Terrapene sigriz, 193.
tessellatus, Natator, 475.
Testudinella, 254.
Testudinidae, 180, 181, 182.
Testudininae, 182, 209-221. 256,

375, 210.
Testudo, 172, 218, 220, 221, 254-

261, 284, 295, 298, 343, 344, 354,

364, 375, 376, 378.
Testudo afrieana, 368, 369.
Testudo angulata, 342, 345, 346.
Testudo arcuata, 499.
Testudo areolata, 352, 367, 369,

387.
Testudo a. pallida, 368.
Testudo armata, 251.
Testudo bellii, 346.
Testudo bergeri, 326.
Testudo biguttata, 229, 250, 251.
Testudo bipunctata, 229, 251.
Testudo boettgeri, 299. 301, 326,

327, 329, 330, 332.
Testudo cafra, 360.

Testudo calcarata, 228, 231, 238,

241.
Testudo campanulata, 264, 277.
Testudo (C.) angulata, 346.
Testudo caouana, 490.
Testudo caretta, 470, 489, 490.
Testudo cartilagiiiea, 421.
Testudo caspica, 191.
Testudo C. cafra, 360.
Testudo C. signata, 360.
Testudo capediana, 474
Testudo cephalo, 489, 490.
Testudo comptoni, 218, 354.
Testudo coriacea, 499.
Testudo denticulata, 404.
Testudo elegans, 221, 225.
Testudo erosa, 374, 404.
Testudo euphratica, 421.
Testudo europaea, 201, 202.
Testudo fasciata, 369.
Testudo ferox, 421.
Testudo fiski, 301, 325, 327, 329.

331.
Testudo flava, 202.
Testudo geometrica, 173, 294, 299,

301, 316, 319, 337, 387.
Testudo g. nigriventris, 301, 337.
Testudo g. tentoria, 301, 321.
Testudo gigantea, 225, 255.
Testudo graeca, 170, 218, 254-259,

261, 276, 282, 298, 299.
Testudo g. bettai, 265.
Testudo g. tioweri, 259, 260.
Testudo g. graeca, 259, 260, 261-

276, 212, 214, 217, 257 266, 267.
Testudo g. ibera, 259, 265.
Testudo g. mauritaniea, 264.
Testudo guttata, 191.
Testudo hermanni, 170, 258, 260,

281, 212, 257, 281.
Testudo horsfieldi, 254, 258, 283.
Testudo ibera, 170, 262, 265.
Testudo imbricata, 484, 485.

556

INDEX

Testudo insculpta, 191.

Testudo japonica, 474.

Testudo kleinmanni, 218, 220, 258-

261, 264, 276-283, 212, 214, 278,

279, 280.
Testudo leithii, 218, 276, 277, 280.
Testudo loveridgii, 287-290, 292,

387.
Testudo lutaria, 202.
Testudo luteola, 253, 299, 301, 321,

323.
Testudo lyra, 499.
Testudo maeropus, 474.
Testudo m. amboinensis, 319.
Testudo marginata, 173, 254, 258,

261, 262, 264, 276, 282.
Testudo marina, 474.
Testudo mauritaniea, 262.
Testudo meleagris, 202.
Testudo miuiata, 368, 369.
Testudo minuta, 323.
Testudo muhlenbergii, 191.
Testudo mydas, 473, 474.
Testudo nasicornis, 490.
Testudo oculifera, 299, 301, 315,

326.
Testudo orbicularis, 202.
Testudo oscarboettgeri, 301, 326,

329.
Testudo pardalis, 221, 235, 251.
Testudo p. pardalis, 251.
Testudo procterae, 287, 288, 289,

292.
Testudo pulchella, 202.
Testudo punctata, 191.
Testudo pusilla, 262, 265, 345, 346,

368.
Testudo radiata, 255.
Testudo r. senegalensis, 231.
Testudo schoepfii, 230, 405.
Testudo scutella, 354.
Testudo seimundi, 301, 326, 329,

330.

Testudo semiserrata, 299, 301, 316,

319.
Testudo signata, 352, 353, 359.
Testudo signatus, 369.
Testudo smithi, 299, 301, 325, 327,

329, 330, 333.
Testudo stellata, 221, 225.
Testudo strauchi, 299, 301, 302,

321, 322, 323.
Testudo sulcata, 221, 230, 232, 237.
Testudo t. africana, 345, 346.
Testudo tentoria, 301, 308, 325,

336, 338, 339.
Testudo t. fiskii, 326, 333.
Testudo tornieri, 283, 286, 288.
Testudo (M.) tornieri, 287.
Testudo trimeni, 301, 307-311, 333.
Testudo triunguis, 420, 421, 423.
Testudo tuberculata, 499.
Testudo verreauxii, 324, 327, 337.
Testudo verroxii, 301, 324, 327.
Testudo V. bergeri, 327.
Testudo viridis, 473, 474.
Testudo viridis-squamosa, 474.
Testudo vulgaris, 474.
Testudo whitei, 264.
tetradactyla, Chersine, 368.
Tetrathyra, 436.
Tetrathyra baikii, 438, 439, 443,

444.
Tetrathyra vaillantii, 444.
Thalassiochelys tarapacona, 491.
Thalassochelys, 470, 489.
Thalassochelys caretta, 491, 495.
Thalassochelys caouana, 489.
Thalassochelys controversa, 495.
Thalassochelys corticata, 491, 492.
Thalassochelys olivacea, 495.
Thalassochelys tarapacana, 491.
thunbergii, Caretta, 474.
Tilapia, 433.
tornieri, Malacochersus, 167, 172,

284, 286-294, 378, 388, 212, 214,

INT>EX

557

317 257, 285, 288.
tornieri, Testudo, 283, 286, 288.
tornieri, Testudo (M.), 287.
Tortisternum, 422.
Tortisternum novemeostatum, 422.
trimeni, Cliersiiiella, 333.
trinieni, Psammobates, 333.
trimeni, Psammobates t., 301, 302,

309-311, 313, 315, 327, 330, 333-

336, 338, 342, 212.
trimeni, Testudo, 301, 307-311, 333.
Trionychidae, 180, 412-420, 416.
Trionyx, 220, 414, 420-423. 467,

470.
Trionyx aegyptiacus, 420, 422, 423.
Trionyx cariniferus, 421.
Trionyx cartilagineus, 415.
Trionyx emoryi, 417.
Trionyx euphraticus, 417.
Trionyx ferox, 417.
Trionyx formosus, 415, 421.
Trionyx gangeticus, 415, 422.
Trionyx hurum, 415.
Trionyx javanicus, 421.
Trionyx labiatus, 424, 427, 433,

434.
Trionyx leithii, 415.
Trionyx mortoni, 425.
Trionyx muticus, 417.
Trionyx nigricans, 415.
Trionyx niloticus, 421, 424.
Trionyx ornata, 421.
Trionyx peguensis, 421.
Trionyx sinensis, 415, 417, 421, 422.
Trionyx spinifer, 417, 421.
Trionyx steindachneri, 417.
Trionyx subplanus, 415, 421, 422,

427.
Trionyx swinhoei, 417.
Trionyx triunguis, 415, 417-419,

423-435, 419, 428, 430, 431.
Trionyx t. rudolfianus, 426.
Trionyx vertebralis, 427.
triunguis, Amyda, 426.
triunguis, Amyda t., 427.

triunguis, Pelodiscus, 426.

triunguis Testudo, 420, 421, 423.

triunguis, Trionyx, 415, 417-419,
423-435, 419, 428, 430, 431.

tuberculata, Testudo, 499.

Tyrse, 421.

Tyrse nilotica, 425.

uintensis, Geochelone, 224.

vaillantii, Tetrathyra, 444.

Venus verrucosa, 494.

verreauxii, Peltastes, 324.

verreauxii, Testudo, 324, 337.

verroxii Chersinella, 327.

verroxii, Peltastes, 324.

verroxii, Psammobates t., 289, 295,
301-303, 313, 314, 316, 324-
333, 336, 337, 338, 341, 342, 358,
3G2, 215, 222, 313, 329.

verroxii, Testudo, 301, 324, 327.

verrucosa, Venus, 494.

vertebralis, Trionyx, 427.

vesperina, Psammobia, 494.

virgata, Chelone, 476.

virgata, Chelonia, 474.

viridis, Chelone, 475.

viridis, Chelonia, 474.

viridis, Mydas, 475.

viridis-squamosa, Testudo, 474.

viridis, Testudo, 473, 474.

vulgaris, Emys, 193.

vulgaris, Testudo, 474.

whitei, Peltastes m., 264.

whitei, Testudo, 264.

Xylocopa nigrita, 392,

yniphora, Asterochelys, 224.

j-niphora, Geochelone, 342.

youngi, Kinixys, 388.

Yuen, 421.

Yuen leprosus, 421.

Zizyphus jujuba, 275.

zombensis, Kinixys, 388.

zombensis, Kinixys b., 387.

zuluensis, Kinixys, 388.

zuluensis, Kinixys b., 387.

PLATES

PLATE 1
Geochelone sulcata. (From Sowerby and Lear,)

r

<

PLATE 2

Gcocliclonc pdrdali.s subspecies

A. G. p. pardalis juv. (M.C.Z. 22473) ex Aroab, Southwest Africa.

B. G. p. l>ahc<irl-i juv. (M.C.Z. 40005) ex Mt. Mbololo, Kenya Colony.

C. ft. p. panhili.s adult (M.C.Z. I(i713) ex Koliuaiiskop, Southwest
Africa.

D. G. p. hahcocli adult (M.C.Z. 18156) ex Aiusha, Tanganyika Terri-
tory.

(Frank Wliite, photographer.)

2247

^^^li?-

I) Ny*^*^

PLATE 2

PLATE 3

Tv.'itudv (/raced (/lacca

Adult. (M.C.Z. 29909) ex Manioia Foient, Eahat, Morocco. Dorsal and
ventral views.

(Frank Wliite. 3iliotographer.)

•■ -^.-

••L'

PLATE

J

PLATE 4

Testudo Jclcinmanni

Adult (M.C.Z. 54044) ex Salum, Western Desert, Egypt. Dorsal ami
ventral views.

(Frank White, photographer.)

PLATE 4

PLATE 5

Malacochersus tornieri

Adult (M.C.Z. 30004) ex Mangasini, Usandawi, Tanganyika. Dorsal
and ventral views.

(Frank White, photographer.)

PLATE 5

PLATE 6

Valacocliersus iornieri

Juvenile (Holotype of procteruc, B.M. 1923.10.9.102 now 1946.1.22.59)

ex Ikikuyu, south of Guhve, Tanganyika.
B, C. Dorsal and ventral views; A. head and limbs enlarged.

(Courtesy of British Museum. Peter Green, photographer.)

'v.-i

PLATE 6

PLATE 7

Psammobates oculifer

Adult (Type of semistrrata) ex "between Latakoo and the Tropic of
Capricorn. "

a, a. Nuchal shield and forelimb of oculifer, compared with

b, b. nuchal shield and forelimb of geometricus.

(From Andrew Smith.)

PLATE 7

PLATE 8

Psammohates geometricus

A, B. Juv. (luteola phase) (P.M.) ex "Cap".

(Frank White, photographer.)
C, D. Adult (B.M.) ex South Africa.

(Courtesy of British Museum. Peter Green, photographer.)

A

-^j»-

4. -^^ • A

c

I)

PLATE 8

PLATE 9

Psammobates tentorius verroxii

Adult (Type of verroxii) ex "near the sources of tlio Garriep or Orange
River. ' '

(From Andrew Smith.)

PLATE 9

■"^■f"?sl^.

PLATE in

Psaniinobatt s ten fori as subspecies: dorsal patterns.

A. P. t. i-crro.rii Adult (Paratype of schonlaHili. M.C.Z. 42222) ex
Steiiik(.)pl', L. XanuHiualnnd.

B. P. 1. VI rroxii Adult (color form hergevi in B.M.) ex Barby, Tiraz
Mtns., Southwest Africa.

C. P. t. tciilorius Adult (typical pattern. M.C.Z.) ex "South Africa."
I). P. f. te7ttoriut: Adult (aberrant. IM.C.Z. 4(3604) ex "South Africa."

(Frank White, photographer.)

PLATE lU

PLATE 11

Pfiumviobates tentorius tentorius

Plastral patterns of nine Transvaal Museum specimens from Matjes-
fontein, Cape Province.

T.M. 19650. C. T.M. 19497.

T.M. 19649. F. T.M. 19536.

T.M. 19479. I. T.M. 19495.

(Prank Wliite, photographer.)

A.

T.M.

19537.

B.

D.

T.M.

19496.

E.

Cr.

T.M.

19480.

II.

PLATE 11

PLATE 12

Chersina angulata S and $

A. Dorsal view of strongly projecting gular in S .
A'. Plastral aspect of same.

B. Dorsal view of moderately pro.iettuig gulrii- in 9.
D'. I'lastral aspect of same.

(From Vail Ian t. i

.^'

W^=^

PLATE 12

PLATE 13

}[<niuiput: specii-'s: <l(irs;il vifw« of c-<ir;i paces.

A. H. signatus (M.C.Z. 42218) ex Steinkop.

B. 11. bouJnu/eri (M.C.Z. 42L'31 ) ex "Cape Province."

C. E. areola t us (M.C.Z. 17525j ex Tootabi.

D. E.femoralis (M.C.Z. 17523) ex Hanovr.

(Frank White, photographer.)

PLATE 1.

PLATE 14

Kinixys species : dorsal views of carapaces.

A.K. h. helliana (M.C.Z. 42852) ex Lukafa, Belgian Congo.

B. Ji. homeana (M.C.Z. 38371) ex Ifan. Ondo, Nigeria.

C. A', erosa inim. (M.C.Z. 22486) ex Lenga, Liberia.

I). 7v. h. helliana (M.C.Z. 40015) ex Voi, Kenya Colony.

(Frank White, Photographer.)

PLATE 14

PLATE 15

Trionyx triunguis

Juvenile ex Wadi Haifa, Nile, Egypt.

(From J. Anderson.)

>*^„

■^%

)

PLATE 15

PLATE l(j

CiK'hinnihis species: juveniles for comparison.

A. ('. (Jefians (B.M. 1908.4.7.3) ox Anglo-Egypti:ni .Sudan.

IS. ('. ihaans same specinu^n, plastral view.

('. C. .sowgalensis (B.M. 1907.7.16.26) ex Gambia.

1). C. s< itri/dh'iisi.s same specimen, jilastral view.

(Courtesy of British Museum. I'eter Green, pliotograplier.)

^

c

i

»p

/\

B

PLATE 16

PLATE 17

Cycloderma species : juveniles for comparison.

A. C. frenatum (M.C.Z. 50360) ex Chowe, Nyasaland.

B. Same specimen, plastral view.

(Frank White, photographer.)

C. C. aubryi (B.M. 1908.5.25.3) ex Zombo, Congo Eiver, Belgian Congo.

D. Same specimen, plastral view.

(Courtesy of British Museum. Peter Green, photographer.)

PLATE 17

PLATE 18

Cydodcrnia species: aduJts for eoiuparison.

A. C. aubryi (Type: Paris Museum) ex Gabon, French Congo.

(Reduced. Fiom A. Dunieril.)

B. C. frcnafani (Cotype: Z.M.U.) ex Zamliezi River, Mozaniljique.

(From W. K. H. Peters.)

i>

PLATE 18

Harvard MCZ LIbrai

3 2044 066 303 876