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BREVIORA
MUSEUM OF COMPARATIVE ZOOLOGY
AT
HARVARD COLLEGE, IN CAMBRIDGE
NuMBERS 1-66
1952-1956
CAMBRIDGE, MASS., U.S.A.
1957
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Nov 3
No. 4
No: 5
No. 6
No. 7
No. 8
No. 9
No. 10
CONTENTS
BREVIORA
MuSsEUM OF COMPARATIVE ZOOLOGY
NuMBERS 1-66
1952
A new Panamanian tree frog. By Epwarp H. Taytor.
3 pp., 1 pl. February 8.
A staurotypine skull from the Oligocene of South Da-
kota. (Testudinata, Chelydridae). By Ernest WIL-
LIAMS. 14 pp., 2 pls. February 8.
Notes on siphonophores. 38. Nectopyramis spinosa n. sp.
By Mary Sears. 4 pp. May 23.
A unique remopleuridid trilobite. By H. B. Warirtrne-
TON. 10 pp., 1 pl. June 9.
Tricholimnas conditicius is probably a synonym of T'rt-
cholimnas sylvestris. (Aves, Rallidae). By James C.
GREENWAY, JR. 4 pp. August 29.
Contributions toward a reclassification of the Formicidae.
[. Tribe Platythyreini (Hymenoptera). By WILLIAM
L. Brown, JR. 6 pp. August 29.
On the nomenclature of the Pacific gray whale. By W1L-
LIAM E. ScHEVILL. 3 pp. September 29.
A new species of the cyclostome genus Paramyzine from
the Gulf of Mexico. By Henry B. BickELow and W1t-
uIAM C. ScHROEDER. 10 pp. October 21.
New species of earthworms from the Arnold Arboretum,
Boston. By G. E. Gates. 3 pp. October 21.
On the earthworms of New Hampshire. By G. E. GatTEs.
3 pp. December 29.
No.
mB ol
eli
1953
Characters and synonymies among the genera of ants.
Part I. By Wiuutam L. Brown, Jr. 13 pp. March 20.
A eave faunule from western Puerto Rico with a diseus-
sion of the genus Isolobodon. By Tuomas E. Reyn-
oups, Kart F. KoopMan and ERNEST 1B WILLIAMS.
7 pp., 1 pl. April 23.
Fossils and the distribution of chelyid turtles. 1.
‘“Hydraspis’’ leithii (Carter) in the Eocene of India
is a pelomedusid. By Ernest WiuuiamMs. 8 pp., 3 pls.
April 23.
Record of a hermaphroditic horseshoe crab, Limulus
polyphemus L. By Joun P. Baptist. 2 pp., 2 pls.
May 28.
further notes on the earthworms of the Arnold Arbore-
tum, Boston. By G. E. Gates. 9 pp. May 15.
Notes on the races of Micrurus frontalis (Duméril,
Duméril and Bibron). By BENJAMIN SHREVE. 6 pp.
May 28.
A new Ordovician trilobite from Florida. By H. B.
WHITTINGTON. 6 pp., 1 pl. May 28.
Characters and synonymies among the genera of ants.
Part II. By Wiuuram L. Brown, Jr. 8 pp. Septem-
ber 23.
Lower Cretaceous nautiloids from Texas. By BERNHARD
KumMEL. 11 pp., 2 pls. September 23.
Lower Triassic Salt Range nautiloids. By BERNHARD
KummMeEL. 8 pp., 2 pls. September 23.
The ancestry of the family Nautilidae. By BERNHARD
KuMMEL. 7 pp., 1 pl. September 23.
Notes on the ant, Leptothorazr obliquicanthus Cole (Hy-
menoptera: Formicidae). By Rosert E. Greee. 3 pp.
October 13.
A new species of Hypognatha from Panama. By ARTHUR
M. Cuickerine. § pp. October 23.
No.
No.
No. 35.
24.
eA
. 30.
voile
. 32.
. 33.
. 34.
1954
A new family, a new genus, and two new species of
batoid fishes from the Gulf of Mexico. By Henry B.
BIGELow and WILLIAM C. SCHROEDER. 16 pp. Janu-
ary 27.
A new Miocene species of Pelusios and the evolution of
that genus. By Ernest Wiuutams. 7 pp., 4 pls. Janu-
ary 28.
A preliminary list of the earthworms of northern New
Jersey with notes. By H. Davies. 13 pp. February 3.
Anterior regeneration in a sexthecal species of lumbri-
cid earthworm. By G. E. Gates. 5 pp. February 5.
Clemmydopsis Boda a valid lineage of emydine turtles
from the European Tertiary. By Ernest WILLIAMS.
9 pp. February 8.
Absence of mesoplastra in a Pelomedusa (Testudines,
Pelomedusidae). By Ernest WiLuiAms. 4 pp.. 2 pls.
February 8.
Aestivation in a Permian lungfish. By ALFRED S. RoMER
and Everetr C. Ouson. 8 pp., 1 pl. February 8.
New freshwater gastropod mollusks of the African
genus Lanistes. By T. Pan. 4 pp. March 3.
Fossils and the distribution of chelyid turtles. 2. Addi-
tional reputed chelyid turtles on northern continents:
Palaeaspis conybearu (Owen) —a pelomedusid. By
ERNEsT WILLIAMS. 6 pp. March 12.
Systematic and other notes on some of the smaller
species of the ant genus Rhytidoponera Mayr. By
Wiuuiam L. Brown, Jr. 10 pp. May 14.
A review of the coxalis group of the ant genus Sticto-
ponera Mayr. By Witu1am L. Brown, Jr. 10 pp.
July 20.
New or rediscovered pelomedusid skulls from the Tertiary
of Africa and Asia (Testudines, Pelomedusidae). 1.
Dacquemys paleomorpha, new genus, new species from
the Lower Oligocene of the Fayum, Egypt. By Ernes?
WiuuiaMs. 8 pp., 1 pl. July 28.
No.
No.
No.
36.
noo:
. 40.
41.
. 43.
. 44.
. 46.
47.
Present knowledge of the snake Elachistodon wester-
manni Reinhardt. By Cart Gans and ERNEsT E.
WiuuiamMs. 17 pp. August 6.
On the evolution of an oriental earthworm species,
Pheretima anomala Michaelsen 1907. By G. E. GATEs.
S pp. August 18.
Onnia (Trilobita) from Venezuela. By H. B. Wurr-
TINGTON. 5 pp., 1 pl. November 28.
New or redescribed pelomedusid skulis from the Terti-
ary of Africa and Asia (Testudines, Pelomedusidae ).
2. A podoenemide skull from the Miocene of Moghara,
Egypt. By Ernest Witiiams. 8 pp., 2 pls. Novem-
ber 24.
Some mollusks from the continental slope of northeast-
ern North America. By ArTHUR H. CLARKE, JR. 11
pp. November 29.
Comments on the classification of rodents. By ALBERT
E. Woop. 9 pp. December 17.
1955
A new salamander of the genus Parvimolge from Mexico.
By Gerorce B. Rass. 9 pp. February 28.
Speed-induced skin folds in the bottle-nosed porpoise,
Tursiops truncatus. By FRANK 8. EssaPIAN. 4 pp.,
9 plss April i:
A new Murex from Matanzas, Cuba. By WILLIAM J.
CLENCH. 2 pp., 1 pl. April 8.
Palaeotaricha oligocenica, new genus and species, an
Oligocene salamander from Oregon. By RicHarp
vAN FRANK. 12 pp., 3 pls. June 13.
Cave-fossil vertebrates from Camaguey, Cuba. By Karu
F. KoopMan and Ropoutro RurBAu. 8 pp. June 24.
A new species of whiptailed lizard (genus Cnemido-
phorus) from the Colorado plateau of Arizona, New
Mexico, Colorado, and Utah. By Cuaries H. Lowe,
Jn. Ipp. July 12.
No.
No.
48.
149:
. 00.
54.
. 60.
Notes on American earthworms of the family Lum-
bricidae. I-II. By G. E. Gates. 12 pp. October 10.
Three new shark records from the Gulf of Mexico. By
Henry B. Bicgetow, W. C. SCHROEDER, and STEWART
SPRINGER. 12 pp. November 18.
New frogs of the genera Asterophrys and Oreophryne
from New Guinea. By ArtrHuR LovertpaGr. 5 pp.
November 30.
1956
A small mustelid from the Thomas Farm Miocene. By
STANLEY J. OLSEN. 5 pp. January 27.
Remarks on some Miocene anurans from Florida, with
a description of a new species of Hyla. By WALTER
AUFFENBERG. 11 pp. April 6.
Food-finding by a captive porpoise (Tursiops trun-
catus). By WiuuiAM E. ScHrvini and BarBara Law-
RENCE. 15 pp. April 6.
A revision of the genus Brachymeles (Seincidae), with
descriptions of new species and subspecies. By WALTER
C. Brown. 19 pp. June 6.
Rediscovery of. Hyla dorsalis and Lechriodus papuanus
in New Guinea. By ARTHUR LOVERIDGE, 4 pp. June 6.
Notes on the Jamaican and Cayman Island lizards of the
genus Celestus. By PENNY NORSEEN COUSENS. 6 pp.
June 15.
Three new species of Mimetidae (Araneae) from Pan-
ama. By Artuur M. Cuickerina. 14 pp. June 29.
Sequence of Passerine families (Aves). By E. Mayr
and J. C. GREENWAY, JR. 11 pp. June 29.
A new subgenus of Chamaeleo from Rhodesia and new
race of Mabuya from Kenya Colony. By ArTHUR
LOVERIDGE. 4 pp. September 12.
A new species of Agriognatha from Jamaica, B.W.I.
By ArtHur M. CHICKERING. 7 pp. September 12.
No.
No.
No. 6
No
GD)
. 66.
On regeneration by earthworms of a species of the lum-
bricid genus Dendrobaena Eisen 1874. By G. E.
GATES. 6 pp. September 14.
A third leaf-nosed species of the lizard genus Anolis
from South America. By JaAmMEs A. PETERS and Gus-
‘pavo OrcEs-V. 8 pp. October 3.
New bathyal Isopoda from the Caribbean with observa-
tions on their nutrition. By Roperr J. Menzies. 10
pp. October 11.
Rare species of Copepoda, Calanoidea, taken from the
Izu Region. By Oronrro Tanaka. 8 pp. October 11.
A new species of Agriognatha (Araneae, Argiopidae )
from Panama. By ArtHuR M. CHICKERING. 7 pp.
November 20.
The Caninae of the Thomas Farm Miocene. By StTan-
LEY J. OLSEN. 12 pp. November 27.
INDEX OF AUTHORS
BREVIORA
MUSEUM OF COMPARATIVE ZOOLOGY
NuMBERsS 1-66
1952-56
No.
PSERENBERG, WALTER «2: oie Sc cldn skies °. 52
EP ISA ROLUN Me 3) atehes t 1 o oats te She BRINE, «A .. 6 Sng 14
BiegELow, Henry B. and WILLIAM C. SCHROEDER ......... 8, 24
BigELow, Henry B., W. C. ScHROEDER and STEWaRT
RIBRUN GER aes ais a oh. pee lee tes eee OG. say aan 49
RON MATER Co pcisc.)-5 ane See 2 ery = ee Os!
ISROW NE WVIGLTAMG Ii) Ral. 0 2. veut tat se) 6, dl 18188. 84
WHICKERING/ ARTHUR DE. .(: 5.1.5 «tee. sates ee 20, Di 0009
CLARKE, ARTHUR H., JR. .... 40
(INC ET paNV TITAN elie we ots. 2s. 2 2) fp eh aa cl nde cea rie ie 44
WONSENS SO ENN WNORSEEN ... ..:. ha ve ine). slo cane 56
Daviss, H. 26
FISSAPTAN, URANEK (93.05... -.-.- + « ie Chis aera" hs as a 3
Gans, CaRL and Ernest E. WILLIAMS .................. 36
Golmigorgye 161) SRN Ue IU oe ee 9, 10, 15, 27, 37, 48, 61
SREMNAUVAEE AIMS COUN el Rey sc. 2 ase > Padawan ohepciares See 5
GEEMINWAY.t- CdR. and H. MAYR.% 3; hee: 58
PREGGO MIROBERT (Hi eile ag. ose ae 99
Koopman, Karu F., and RopoLro RUIBAL ... 46
Koopman, Karu F., THomas E. REyYNoups and ERNEst BL
WV RUBIA MS 24S boro ae ope 12
PO MMEE ISERNEARD (0% 052725 wjsirge wedtae hf aol. ee 19, 20, 21
LAWRENCE, BARBARA and WILLIAM E. SCHEVILL ..... 53
LGVERIDGEA AG TUR caer ese e Sem eede 50, 55, 59
Own CHAR ES Bil. cman choo: eee gee eee Fe amo oe 47
MAwR Hy sand: Jit CaGREbINpwaAty, c) Rad) <7 geee ene ete he ce eer eae 58
MENZIES) GORI Tee oe See. aes oe eee oe ie), Se heel 63
(ONES TOTS ea SUW UNb) a as MAN cei or CMR eny ah Aco) Ct Aa 51, 66 -
OLsoN, Everett C. and ALFRED S. ROMER ................ 30
Orcrs-V., Gustavo and Jams A. PETERS .:............¢. 62
1 PUN eid NR rca Stat bon ES I ey, 1. a ee, Sete 8 31
PETERS, JAMES Apand:GuSTAVO ORGHS-V ols)... ..22s507e 506 62
RARBG RORGE DO aaer te a2 22 sin tie REI arn Sok 2a eee 42
REYNOLDs, THoMAsS E., Karu F. KoopMAN and ERNEsT EK.
Wigs TAUNIGREPIERE ee ds if We ee ER tod Ake a 12
Romer, Auennp is, and HveRErTr ©. OUSON 02.2.) 2.4 seee 30
RUIBAL; RopoLro and Karu F’. KoOPMAN ..............-- 46
SCHEV wa EbaNAME JH), 22k). ss ces nee Gata oc ee 7
SCHEVILL, WILLIAM E. and BARBARA LAWRENCE .......... 5S
ScHROEDER, WILLIAM C. and Henry B. BIGELow ........ 8, 24
ScHRoEDER, W. C., Henry B. BigELow and STEWART
SPRINGIR 08 [eso ccs ace eee ee 49
SEARS. MAR Wi. ceStet s. 5 ec ER: Geen ee hat ee tae 3
SEREVE, (SENUAMING: . . sl:+ oat. Meera cho, eo eee eee ee 16
SPRINGER, STEWART, Henry B. BiaELow and W. C.
SCHROEDERS —c4. 54 Sone Ce Eee el. ee ee 49
iDanacas) OPOBITO: 2. 3 ...1 J serve eae te coat ae eee 64
WAviOR.. MDWARD: J” .%.- .,: secaateigeat oils serene eee il
WAN GM RANK? (RIGEDARD: |< 5.0. \. 2 dey eke, cig Set 8 ee aes ee 45
VEERING DON Ely Bern 2 pain apne naee teeta ls rata 4,17, 38
NV PANS HIRST o2.05 24).2 3 2). yee 2, 13, 25, 28,29 S2e some
WaburAmMs, HRNESsT EH. and Caru,GANS 2°34 45-25 ee ee 36
WiuiAMs, Ernest E., Kart F. KoopMAN and THomas E.
IRE VNOMDS. “sos Moun ch. A ic a One LO ee eee a
WioODTALBERT Bs avis. sass 2 See eee eh ge ete eel 41
- i
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass.
Frsruary 8, 1952
NUMBER |
A NEW PANAMANIAN TREE FROG
By Epwarp H. TayLor
Department of Zoology, University of Kansas, Lawrence, Kansas
While studying Central American frogs in the Museum of Com-
parative Zoology, Harvard College, a small, presumably undescribed
frog was noted, and Mr. Arthur Loveridge, with his characteristic
kindness, suggested that I study the form and describe it if it proved
new. The most characteristic of the differential features of this di-
minutive frog is the very considerable enlargement of the horny spines
on the nuptial swelling of the first finger. It is presumed that it is a
rivulet species, living in the neighborhood of small streams, rather than
a bromeliad species.
After my study of the specimen and comparison with other known
species of southern Central America, | conclude that the species is new
and I present herewith its description.
Hy La SHREVEI sp. noy.
Type. M.C.Z. No. 26769 o. La Loma, Chiriquicito, Reptblica de
Panama; EK. R. Dunn and C. Duryea, collectors.
Diagnosis. A diminutive frog (25.2 mm.) with a broad, rather flat
head; skin smooth; outer fingers with a trace of web, and well-developed
dises; canthus rostralis wanting; nostrils nearly terminal; tympanum
large directed strongly upward, its diameter at least half of eye length;
2 BREVIORA No. |
eye directed obliquely forward; vomerine teeth in two rounded fasci-
culi, lying almost completely behind posterior level of large choanae;
no web between two inner toes; others one third, to one half webbed;
first finger of male with 19-22 large horny spines.
Description of the type. Head flattened, broader than body, its width
(10 mm.) greater than its length (9.1 mm.); tympanum large, semi-
transparent, pigmented, facing more upward than outward, the upper
edge obscured by a fold arising at eye and curving back to jaw angle,
its diameter (1.5 mm.) equal to half length of eye opening (8 mm.);
distance between tympanum and eye 2.8 mm.; eye prominent, directed
somewhat forward; length of snout less than eye length (2.85 mm.);
canthus absent, loreal region concave, sloping obliquely to lip; nostrils
nearly terminal, the areas about them strongly swollen with a de-
pression between; no notch in upper lip; groove from angle cf jaw
touches tympanum; choanae subcircular, the distance between them
2 mm.; diameter (transverse) of choana .65 mm.; vomerine teeth in
two somewhat circular fasciculi almost completely behind the posterior
level of choanae, the teeth in somewhat curving rows; they are equally
separated from choanae and from each other, a fasciculus as large as
a choana; tongue rather thick, broader than long, not or scarcely
notched behind; vocal slits very large, lateral; openings of the palatal
gland forming a series of separate slitlike openings arranged in a broad
V-shape about the middle of palate.
Arm very short, the wrist not reaching tip of snout; first finger
short, greatly expanded at base by a very large nuptial swelling and
a large metacarpal tubercle; swelling covered with a black, horny plate
bearing 19-22 large horny spines; dises of fingers smaller than tympa-
num, that on first finger only little smaller than those on outer fingers;
a small trace of webbing between outer fingers; subarticular tubercles
single; supernumerary tubercles numerous; palmar tubercle partly
divided; a slight axillary web present; heel reaches forward to edge of
eye; tarsal fold represented by a series of tubercles growing more
distinct on the proximal end of tarsus; a large inner metatarsal
tubercle and a small distinct outer; no web between first and second
toes; one fourth webbed between second and third; approximately
half webbed between the third and fourth and two fifths webbed be-
tween the fourth and fifth, the web forming a slight ridge to dises on
sides of digits; subarticular and supernumerary tubercle distinct. Skin
on dorsum smooth, slightly wrinkled or corrugated on sides; chin
smooth; breast, venter, part of the ventral surface of thighs and region
lateral to vent, strongly granular; a rather elongate anal flap, the vent
opening midway between upper and lower surface of thighs, followed
“
1952 NEW TREE FROG
by a groove bordered by small granules and on ventral surface by a
pair of distinctly larger granules; fold above tympanum curves down to
jaw angle.
Color. In preservative, light brown above with a brownish-white
shade below; the pigment is in minute chromatophores, that can be
clearly seen under a lens; no pigmentation on under surfaces except
on edge of jaw, under surface of thigh, part of under surface of tibia
and some scattered flecks under hand and foot; upper eyelids appearing
dark from the dark covering of the eyeball; tip of snout somewhat
darker than dorsum.
Measurements in mm. Snout to vent 25.2; width of head 10; length
of head 9.1; diameter of tympanum 1.5; length of eye opening 3; snout
length 2.85; arm 13; hand 7.2; leg 38; tibia 12; foot and tarsus 15.
Remarks. The species is probably related to Hyla zeteki. There are
numerous similarities in structure, and the measurements are similar
but the most striking differences appear to be in the remarkable
nuptial asperities on the first finger, the position and direction, and
the larger size of the tympanum (14 instead of 14 of eye); the absence
of the spectacle-like markings about eyes. The characteristic anal
decoration of zeteki is absent, and the strong granulation on under
surface of arm, and on sides is absent.
The species is named for Mr. Benjamin Shreve of the Museum of
Comparative Zoology, Harvard College.
4 BREVIORA No. |
Hyla shrevei sp. nov. M.C.Z. No. 26769, La Loma, Chiriquicito, Reptblica
de Panama. KE. R. Dunn and C. Duryea, collectors. (Type o X 2.) 1. Dorsal
view. 2. Lateral view of head. 3. Enlargement of nuptial spines on inner
digit.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. Fepruary 8, 1952 NuMBER 2
A STAUROTYPINE SKULL FROM THE
OLIGOCENE OF SOUTH DAKOTA
(Testudinata, Chelydridae)
By ERNEST WILLIAMS
INTRODUCTION
The modern representatives of the Staurotypinae (two genera,
Staurotypus and Claudius, and three described species) are restricted
to Southern Mexico and Central America. No fossils belonging to these
living genera are known. A form known only from a single perfect
shell from the Oligocene (Chadron) of South Dakota, Xenochelys
formosa Hay, seems to be the only previously described fossil record
of this subfamily.
Recognition of a staurotypine from shell material as good as that
of the unique type of Xenochelys (A.M.N.H. No. 1097) is quite easy.
Staurotypines differ from chelydrines in having only 23 instead of 25
marginals and 21 rather than 23 peripherals. The same feature dis-
tinguishes them from dermatemydids. They differ from kinosternines
in possessing an entoplastron. In these characters Xenochclys is clearly
staurotypine. In the elongation of the first vertebral scute and in its
precise shape Xenochelys closely resembles Staurotypus. The neural
series is much as in Staurotypus. The plastron of Xenochelys has a
reduced number of shields, the pectorals and abdominals having ap-
parently fused. This again is a staurotypine or chelydrine feature.
The general form and height of the shell are very Stawrotypus-like.
There is also some suggestion of the tricarinate condition found in
Staurotypus in the carapace of Xenochelys.
2 BREVIORA NO. 2
In other respects the shell of Xenochelys is more primitive (more
dermatemydid-like) than any other member of the chelydrid series.
The plastron is relatively large (although the bridge is quite narrow as
in chelydrids, not as in dermatemydids). There is a xiphiplastral notch.
The nuchal scute is small. There is a trace of nuchal indentation.!
In shell characters, therefore, Xenochelys is a good structural inter-
mediate between Dermatemys and Staurotypus, and it is also a temporal
and phylogenetic intermediate between the complex of fossil forms
called dermatemydid and the staurotypine section of the Recent
Chelydridae.
It should be noted that Hay was not under any misapprehensions
as to the affinities of Xenochelys. He quite explicitly cited Staurotypus
and Claudius as ‘‘the nearest living relatives of Xenochelys’’ (1908,
p. 282), and in his assignment of the form to the Dermatemydidae he
merely followed Boulenger’s 1889 catalog of the Recent turtles in
which the Staurotypinae were associated with the Dermatemydidae.
When he published his 1908 monograph he had probably not seen
Siebenrock’s 1907 monograph of the “Cinosternidae” in which the
close affinities of the Staurotypinae and Kinosterninae were for the
first time made clear; he thus missed an opportunity to point up more
clearly the apparent ancestral position of the South Dakota genus.
In my 1950 classification of the testudinate order I united the
chelydrines, staurotypines and kinosternines in the family Chelydridae
as understood in a broad sense. To do so somewhat obscures the ex-
treme closeness of relationship of staurotypines and kinosternines,
which differ in the simultaneous loss of an entoplastron and acquisition
of more or less of a box turtle habitus by the latter. The chelydrid
series is surely a natural group, but within that group the staurotypines
and kinosternines stand very much closer to one another than to the
chelydrines.
In the Oligocene this specialized section of the Chelydridae sensu
lato was evidently fully distinct in shell characters, if still somewhat
primitive in a few features. It has, however, not previously been
known to what extent the skull had evolved concomitantly with the
shell.
In the collection of the Department of Geology, Princeton Uni-
versity, I have now found a skull (No. 13686), likewise from the
Chadronian Oligocene of South Dakota, which surely belongs to a
member of the staurotypine-kinosternine section of the Chelydridae.
It is distinctly more specialized in a number of ways than any previ-
1 Most of these characters might also be counted as kinosternine resemblances.
1952 A STAUROTYPINE SKULL 3
ously known staurotypine or kinosternine skull. Although incomplete
it merits extended description and discussion. In the section which
follows, the skull is described in detail and compared with Claudius
augustatus, Staurotypus salvinii, and Sternotherus carinatus.
DESCRIPTION OF THE PRINCETON SKULL
The skull is complete as far as the postorbital bar. Behind this,
however, only the parietals, pterygoids, basisphenoid and basioccipital
are retained, all of them somewhat fragmented and incomplete.
Sutures are rather difficult to make out because of fractures in critical
areas.
The profile of the face is strikingly like that of Stawrotypus or
Sternotherus. The prefrontals project dorsally above the narial opening
but laterally are somewhat retracted, so that in lateral view the nostril
is seen as a distinct angular indentation. The premaxillae unite in the
formation of a distinct median beak, while posterior to this median
projection the contours of premaxilla and maxilla form a smoothly
sinuous curve, which, however, is doubly incurved in the fossil in
contrast to both Stawrotypus and Sternotherus. The depth of the
premaxilla is markedly greater than in Stauwrotypus. The orbit is rela-
tively smaller than in Stawrotypus and even more distinctly lateral
(in contrast to dorsolateral) than in that form. The maxilla below the
orbit is marked by a distinct groove running down to the second
incurving of the lateral festooning of the jaw. The postorbital bar is
rather wide, half the rostro-caudal length of the orbit rather than
between 14 and 14 as in Staurotypus.
In lateral view the prootic is seen to project far forward about the
pterygoid in a very exceptionally developed “‘crista praetemporalis”’
(Siebenrock 1897). Only a narrow channel separates these two bones.
The dorsal margin of the prootic is continued also in a ridge on the
parietal. A similar forward projection of prootic is seen in Staurotypus
(and in Graptemys) but is far less extreme.
The “‘crista praetemporalis” which is the feature exaggeratedly de-
veloped in the fossil and in Staurotypus and Graptemys serves to
modify and increase the leverage of the jaw muscles (Zdansky 1924,
pp. 101-104). All three genera have widened alveolar surfaces of the
maxilla with strong tendency to formation of a secondary palate. In
the case of Graplemys the wide alveolar surfaces are known to be
associated with a malacophagous diet. Probably in the case of the
fossil a strengthening of the jaw action and a similar diet are to be
inferred.
4 BREVIORA No. 2
The dorsal planum of the parietal continues the remarkably flat
dorsal margin of the face.
In anterior view the nostril is very small, much as in Sternotherus,
not as in Staurotypus. The prefrontals are anteriorly distinctly convex,
posteriorly are very flat. Behind them the frontoparietal area rises
somewhat more sharply than in Staurotypus.
In palatal view the maxillae have united in the formation of an
extensive secondary palate, complete in front with a pronounced
median ridge but incomplete in the midline behind. In this secondary
palate the palatines share to about the same extent as in Staurotypus
and to a significantly greater extent than in Sternotherus. A striking
and unique feature is the very impressive dorsomedial slope of the
palatal roof. To a very slight extent this condition is prefigured in the
other genera, particularly Stawrotypus, but the difference is very con-
siderable: in this respect no other genus is at all close. There are no
ectopterygoid processes and the pterygoids also are bowed dorsally at
the midline, so that their lateral flanges are very strong and high,
though spreading wide apart. Anteriorly the premaxilla has the deep
pit for the tip of the lower jar characteristic of chelydrids. In the
specimen it breaks through into the narial region. As in Stawrotypus
and Claudius (differing in this from all examined kinosternines and
chelydrines), foramina incisiva appear to be lacking in the fossil. At
the postorbital bar the palate, and thus the whole outline of the skull,
is very expanded from the side: this form must have been decidedly
brachycephalic. The waist of the pterygoids, however, is only moder-
ately broad, as in Sternotherus, not very broad as in Staurotypus or
narrow as in Claudius. The basisphenoidal suture is not clearly dis-
tinguished from breaks in this region, but it seems probable that the
exposed portion of this bone was very short and broad, not tending
to be elongate craniocaudally as in Staurotypus. The infracondylar
depression, so marked in Staurotypus, is less distinct in this form, as
also in Sternotherus, but not so weak as in Claudius.
In dorsal view the great breadth of the skull at the postorbital bar
is again evident, along with the considerable incurving of the skull
contours just in front of the orbits. The origin of the supraoccipital
spine is indicated by a triangular plane surface with well defined lateral
margins, as in Staurotypus and Sternotherus, not as in Claudius.
The sutures bounding the frontals are somewhat obscured by breaks,
but it is extremely probable that as in other Chelydridae the frontals
occupy a very small area and do not enter the orbits.
1952 A STAUROTYPINE SKULL 5
SKULL CHARACTERS IN THE CHELYDRIDAE
AND THE ALLOCATION OF THE FOSSIL
For the comparisons made in the course of the description just given
I have had available the skeletal collections of the Museum of Com-
parative Zoology and of the American Museum of Natural History.
Included in the M.C.Z. collection are a young and an adult skull of
Staurotypus (the young specimen, M.C.Z. No. 4989, is figured), while
the A.M.N.H. collection has furnished for study a skull of Claudius
(A.M.N.H. No. 65865).
Because the fossil is a fragment only, comparisons must remain
incomplete and portions of the skull which might be diagnostically
significant are unavailable. In this circumstance and in the absence
of more complete knowledge of variation and difference within the
genera Kinosternon and Sternotherus (valuable information which we
may hope to learn in the forthcoming revision of these genera by Dr.
Norman Hartweg) I do not attempt to discriminate too finely the
affinities of the fossil.
However some rather general discussion is possible. We may first
consider what characters define a skull as chelydrid in the broad sense,
then what features are chelydrine, staurotypine, or kinosternine, and
finally what provisional allocation of the fossil skull is possible and
expedient.
There are six genera of living chelydrids (Chelydra, Macroclemys,
Staurotypus, Claudius, Kinosternon, Sternotherus), and of these Sterno-
therus might quite properly be relegated to the synonymy of Kino-
sternon. The fossil record adds a few more (Acherontemys, Chelydrops,
Chelydropsis, Xenochelys). In contrast the Testudinidae has about 30
living genera while the fossil record brings the count up to about 50.
It is not surprising, therefore, that the Chelydridae seem a more closely
knit group than do the Testudinidae, even if the rather isolated
Platysternon is omitted from the latter assemblage.
If the skulls of chelydrids (all living forms North America, a few
Tertiary fossils European) are compared only with the skulls of North
American or European testudinids very clear distinctions are evident.
If, however, the comparison is extended to the very rich testudinid
fauna of Southeast Asia where, so far as known, chelydrids have never
occurred, some of the forms to be found there bridge over the differences
which were thought to be significant. Even so astute an observer as
Baur, and one so familiar with the testudinate order, found it possible
‘ A discussion of the anatomy of Claudtus is in preparation.
6 BREVIORA NO. 2
to refer Adelochelys (= Orlitia) to the ‘‘Chelydroidea’”’ when he had
the skull only, though the shell would have placed the genus without
question in the Emydinae.
The distinctions between the Chelydridae and Testudinidae are wide
enough, when all parts of their anatomy are taken into account and
the trends within them are considered, that there is no doubt that,
although related, they have long been separated, perhaps since the
Cretaceous, certainly since earliest Tertiary. Thus there are differ-
ences in cervical formula (Williams 1950), in the form of the eighth
cervical vertebra, in degree of development of the costiform processes
of the nuchal, in the presence versus absence of inframarginals, in the
characteristic reduction of the plastron or its elements in the Chely-
dridae, in the proximal end of the femur, and in the absence in chely-
drids of gaudy or bright patterns.
However, in a way which seems very characteristic of turtles, few
of these features hold good with complete fidelity in all cases. In
cervical formula there is a striking difference between the two families
in that the eighth cervical is biconvex in the Testudinidae, procoelous
in the Chelydridae. A few individuals of the Testudinidae (mostly
advanced tortoises) vary in the direction of the condition of the
Chelydridae, but no chelydrids are known to vary in the direction of
the testudinid condition. The eighth cervical vertebra tends to differ
in the two families, the Testudinidae generally showing three ventral
crests on the centrum, though the lateral ones may be barely indicated,
the Chelydridae showing a single median crest which may divide into
two (some Chelydra, kinosternines).
The costiform processes of the nuchal are typically much developed
in the Chelydridae, relatively little developed in adult testudinids, but
the kinosternine section of the chelydrids approaches the condition of
the testudinids, and young emydines have this process rather strongly
developed. Inframarginals are never normally present in most testu-
dinids, but there is an exception in the case of Platysternon, and
inframarginals do occur as individual variations in Chrysemys picta
(A.M.N.H. specimens to be reported on by Samuel McDowell). The
plastron is never reduced in testudinids; it is strikingly reduced in
protective efficiency in chelydrines and staurotypines; in some species
of the kinosternines it regains its complete coverage of the ventral
surface, but prior to this redevelopment it had lost one of the bones
normally present in unreduced plastra. In the testudinid femur a
fusion of the trochanters tends to limit the intertrochanteric fossa to
a shallow dorsal pit, though a number of emydines (and Platysternon)
ry
‘
1952 A STAUROTYPINE SKULL
have the juncture barely suggested. In chelydrids as in most turtles
the intertrochanteric fossa is a widely open groove. The lack of bright
patterns in the Chelydridae, while characteristic enough of the rela-
tively few genera and species involved, is not consistently contrasted
with the presence of such patterns in the testudinids (see, for example,
the uniform pigmentation of Galapagos tortoises and of some of the
Asiatic emydines.)
Nevertheless, in spite of these exceptions, these postcranial charac-
ters, external and internal, permit in combination a clear discrimination
of the two families.
Several characters distinguish the skulls of chelydrids and testu-
dinids, but here even more than in postcranial characters exceptions
reduce the utility of single characters.
Chelydrids usually possess at least traces or indications of a sharp
median beak or ‘“‘tooth” on the upper jaw. But this is absent in some
kinosternines, and while many testudinids have a notch here, a few
(e.g., Terrapene, Cuora) have a beak quite similar to that of chelydrids.
Most testudinids (but not Malayemys) have the temporal bar deeply
emarginate from below; the chelydrids have this bar at most shallowly
emarginate. Chelydrids have the nostril, orbits, and otic opening
somewhat smaller than is frequently the case in testudinids. The
supraoccipital crest is higher or more steeply arched, the premaxillary
pit is usually deeper, and in staurotypines and kinosternines there is
a more marked festooning of the contours of the upper jaw than occurs
in testudinids. The frontals are always small and excluded from the
orbit in chelydrids; this feature is variable in testudinids.
A combination of most of the characters mentioned defines a chely-
drid skull. The absence of all but one or two, most often of all, defines
a testudinid as contrasted with a chelydrid skull.
Within the Chelydridae determination of chelydrine skulls from
staurotypine and kinosternine skulls is at once possible on one key
character which offers no difficulty. All chelydrines have the temporal
region more fully covered than do any of the more advanced genera.
Discrimination of staurotypine as against kinosternine skulls is more
difficult on the basis of any characters which have the smallest proba-
bility of holding good if more genera are discovered. Perhaps the
premaxillary beak is always more strongly developed in staurotypines
and the temporal bar narrower vertically in the same group.
This difficulty in finding differences in the skulls of these two groups
is akin to the difficulty in distinguishing dorsal shells. The shells of
both subfamilies differ from those of chelydrines in the loss of one pair
8 BREVIORA NO. 2
of marginal scutes and one pair of peripheral plates. Shape and height
of the shells and scute shape are essentially the same. Keeling is
variable. Only in the plastron is a key difference at once apparent in
the absence of an entoplastron. The mobility of anterior and posterior
plastral lobes in kinosternines is another differentiating feature.
In which of these groups does the fragmentary Oligocene skull find
its natural place?
It is clearly chelydrid sensu lato. It has the sharp premaxillary beak
and deep premaxillary pit of a chelydrid. The temporal bar is broken
but there is no suggestion of ventral emargination. The nostrils and
the orbits are quite small. The supraoccipital crest is only partially
preserved but its root gives evidence of a high arch as in typical
chelydrids.
The skull is, however, clearly not chelydrine, since the temporal
region is fully exposed by posterior emargination as in the two ad-
vanced subfamilies, not as in chelydrines. But the postorbital bar is
wider than in any staurotypine or kinosternine; this may be a primitive
feature.
Is it staurotypine or kinosternine?
It has resemblances to both groups. The premaxillary beak is very
strong as in staurotypines, but the temporal bar is very stout in
vertical depth as in kinosternines. The pterygoid waist is moderately
broad as in kinosternines, not very broad as in Stawrotypus or very
narrow as in Claudius. The nostril is very small as in kinosternines
and in contrast to the condition in staurotypines. Foramina incisiva
are lacking as in staurotypines.
Some features, however, are extremely specialized. The degree of
development of the secondary palate is greater than in any presently
recognized chelydrid species, significantly greater than in Stawrotypus.
Quite unique (unique in the order) is the extreme obliquity and dorsal
arching of the secondary palate.
All in all, the skull seems more specialized than that of any living
staurotypine, but at the same time more primitive in at least one
respect (the strong premaxillary beak) than any living kinosternine,
and perhaps more primitive in the width of the postorbital bar than
either modern staurotypines or modern kinosternines.
The skull is Oligocene in age. The only known shell to which it
might belong is Xenochelys, of the same age and not distant in locality.
But the shell of Xenochelys is quite primitive for its group. Can so
specialized a skull be assigned with any probability of correctness to
so primitive a shell?
1952 A STAUROTYPINE SKULL g
Such an association is by no means impossible. Indeed, in the
Princeton skull a few features like the very strong premaxillary beak
and the wide postorbital bar may point to a stage of differentiation
not very different from that of the Xenochelys shell: advanced in some
respects, primitive in others. For the present it seems expedient to
refer the Princeton skull — with doubt — to Xenochelys formosa Hay.
THE GEOLOGIC RANGE OF THE CHELYDRIDAE
The Princeton skull and the American Museum shell of Xenochelys
formosa are the oldest known representatives of the Chelydridae
(Chadronian Oligocene). An older fossil from the Eocene of Tunis
was indeed referred to the family by De Stefano (1903), but his de-
scription was based on a few bone fragments associated with the mold
of three pleurals. The generic and species names, Gafsachelys phospha-
tica, erected on this very insufficient basis may be disregarded as a
nomen vanum and need no longer be considered as part of the fossil
history of the Chelydridae.
It is, of course, surprising that the oldest representatives of the
family should be staurotypine rather than chelydrine. It is, however,
possible that future more complete knowledge may connect some of
the other forms called dermatemydine by Hay (1908) with the Chely-
dridae. The type species of Hoplochelys Pay was first called Chelydra
crassa by Cope, and this genus, though possessed of a full complement
of marginals and peripherals was regarded as possibly related to
Staurotypus by Hay himself. Both this genus (Paleocene of North
America) and Baptemys (Lower and Middle Eocene of North America)
have the plastron considerably reduced, the bridge narrowed and the
posterior lobe pointed, and an arrangement of plastral scutes like that
of Recent Chelydra (pectorals meeting femorals). The shape of the
shell in both genera is quite like that of staurotypinesand kinosternines,
and in Hoplochelys the shell is tricarinate, as it is also in one species of
Baptemys. The humerus of Baptemys tricarinata is very like that of
Chelydra. However, the first vertebral is never elongate as in stauro-
typines and kinosternines. The other vertebrals are never as wide as
in chelydrines, though wider in later (Torrejon) than in earlier (Puerco)
Hoplochelys. The costiform process of the nuchal is said to be short
in Baptemys (Hay, 1908). The skull of Baptemys wyomingensis is
known and is quite un-chelydrid in its major characters: the temporal
bar is deeply emarginate from below, and the temporal region widely
open above, there being a much greater caudal emargination than in
10 BREVIORA NO. 2
chelydrines. The postorbital bar is in consequence relatively narrow.
The orbit also is rather large.
None of the conditions just mentioned in which Baptemys and
Hoplochelys differ from chelydrids positively debars them from ancestry.
For the present, however, and until they are better known and tran-
sitional forms are discovered, it seems convenient to retain them in
the Dermatemydidae, merely calling attention to their possible special
relation to the Chelydridae.
Unless these forms, perfectly suitable in age, are ancestral chelydrids,
there is no record of the family until the early Oligocene, and it is then
first represented, as the shell and the referred skull of NXenochelys
formosa show, by an advanced subfamily.
The first occurrence of apparent Chelydrinae is in Europe and later
in the Oligocene. Fragments of doubtful value from the middle
Oligocene of Germany have been assigned to “Chelydra sp.” by
Reinach (1900), and in the later Oligocene of Germany rather good
remains are found of an undoubted chelydrine, “‘Chelydra” decheni v.
Meyer. Reference of the latter form to the Recent genus Chelydra is,
as Zangerl (1945) has already pointed out, extremely doubtful: though
the shell shape is that of a chelydrine, there are curious resemblances
to Staurotypus and to Macroclemys rather than to Chelydra. Indeed,
H. v. Meyer himself in 1852 expressly admitted that the inclusion of
this form in Chelydra depended upon a very wide generic concept, and
his idea of the genus was very definitely much wider than that current
today.
In the Miocene of both Europe and North America there is a
flowering out — real or apparent — of chelydrine types. In Europe
“Chelydra’”’ murchisoni Bell and four other named species of ‘‘Chelydra”’
and Chelydropsis carinata Peters record the rather widespread oc-
currence of the subfamily. The remarks above for ‘‘C.”’ decheni apply
also to “C.” murchisoni and the other European forms referred to
“Chelydra.”’
The carapace of Chelydropsis carinata has been excellently figured
by Peters (1869). Unfortunately it does not seem possible to verify
in his figure the features upon which Peters relied in distinguishing
this form generically. I am unable to interpret the plate as showing
the presence of supramarginals, and while a division of the nuchal
bone into two parts is clearly shown, I (as also Boulenger in 1889)
doubt that this reflects the normal condition of the animal. Never-
theless, I consider it probable, if only on the grounds of zoogeography
and age, that the genus will stand, though requiring redefinition. It
1952 A STAUROTYPINE SKULL 11
may at least be pointed out that if the eventual much needed restudy
of the European chelydrines should reveal that they all belong to one
genus, the name Chelydropsis is available.
From the Miocene of North America three chelydrines have been de-
scribed. Two are known from skulls only: Chelydrops stricta Matthew
and Macroclemys schmidti Zangerl. Both of these are from Nebraska,
M. schmidti from the Middle Miocene, Chelydrops stricta from the
Upper Miocene. Both are certainly related to Recent M. temminckit,
but they are distinct from that form and from one another. Chelydrops
is unique among known chelydrids in having a ridged alveolar surface
of the maxilla.!. M. schmidti differs from Chelydrops and from M. tem-
minckw by the considerably shorter antorbital portion of the face.
The other Miocene North American form (from the Roslyn Miocene
of Washington) is known from the carapace only, no portion of the
plastron nor any skeletal parts having been recovered. Hay described
this form as a new genus and species, Acherontemys heckmani, because
of the close articulation of pleurals and peripherals and because the
vertebrals were even broader than in living chelydrines. This shell
may belong to either or neither of the forms represented by the skulls
before mentioned.
Zangerl (1945) has described a skull fragment from the Pliocene
(Clarendonian) of South Dakota, which is indistinguishable from
Recent M. temminckvi. Gilmore (1923) has described from the San
Pedro Valley of Arizona, either Pliocene or Pleistocene, a Kinosternon
which is said to differ from Recent K. flavescens mostly in size.
Two species of Chelydra and one of Macroclemys have been described
by Hay from the Pleistocene of Florida. The value of these forms,
based on fragmentary material, will be difficult to determine. There
are also scattered Pleistocene records for the Recent species Chelydra
serpentina and Macroclemys temminckit.
Acknowledgments. I am indebted to Dr. Glenn L. Jepsen for the
privilege of studying and describing the Princeton skull. Mr. Arthur
Loveridge and Mr. C. M. Bogert have generously made available the
comparative Recent material under their care in the Museum of
Comparative Zoology and the American Museum of Natural History
respectively. Dr. E. H. Colbert permitted examination of the types
of Xenochelys formosa and Chelydrops stricta. Dr. A. S. Romer and
Mr. L. I. Price have read the manuscript. Mr. Sam McDowell is to
be credited with the drawings and a number of useful suggestions.
1 The type and figured adult skull fragment has this ridge. The young specimen referred
by Matthew to this form lacks the ridge.
12 BREVIORA NO. 2
TABLE 1, FOSSIL SPECIES OF THE CHELYDRIDAE
OLIGOCENE:
Xenochelys formosa Hay
Chadronian Oligocene (S. Dakota) North America
“Chelydra” decheni v. Meyer
Upper Oligocene (Siebengebirge) Europe
MIOCENE:
“Chelydra”’ murchisoni Bell
Miocene (Oeningen) Europe
Chelydropsis carinata Peters
Miocene (Eibiswald) Europe
“Chelydra”’ meilheuratiae Pomel
Miocene (Allier) Europe
“Chelydra” lorettana (v. Meyer) Glaessner
Miocene (Leithagebirge) Europe
“Chelydra” argillarum Laube
Miocene (Preschen) Europe
“Chelydra” allinghensis E. Fuchs
Miocene (Viehhausen) Europe
Macroclemys schmidti Zanger]
Middle Miocene (Nebraska) North America
Chelydrops stricta Matthew
Upper Miocene (Nebraska) North America
Acherontemys heckmani Hay
Miocene (Washington) North America
and additional European records for ‘‘Chelydra sp.”” and
“Macroclemys sp.”
PLIOCENE:
Macroclemys temminckii (Holbrook) Zanger!
Early Pliocene (S. Dakota) North America
PLEISTOCENE:
Macroclemys floridana Hay
Pleistocene (Florida) North America
Chelydra laticarinata Hay
Pleistocene (Florida) North America
Chelydra sculpta Hay
Pleistocene (Florida) North America
Kinosternon arizonense Gilmore
Pleistocene (Arizona) North America
and additional North American records for M. temminckii and
C. serpentina
1952 A STAUROTYPINE SKULL 13
LITERATURE CITED
BouLENGER, G. A.
1889. Catalogue of the chelonians, rhynchocephalians and crocodiles in
the British Museum (Natural History). London. 311 pp.
Fucus, ErtKa
1938. Die Schildkrétenreste aus dem oberpfalzer Braunkohlentertiar.
Palaeontographica, Abt. A, Vol. 89, pp. 57-104.
GILMoRE, C. W.
1923. A new fossil turtle, Kinosternon arizonense. Proc. U.S. Nat. Mus.,
Vol. 62, pp. 1-8.
GLAESSNER, M. F.
1933. Die Tertiirschildkréten Niederdsterreichs. Neues Jahrb. Min.
Geol. Pal., Abt. B, Vol. 69, pp. 353-387.
Hay, iO; PF:
1908. Fossil turtles of North America. Carnegie Institution of Washing-
ton Publication No. 75. 568 pp.
1916. Description of some Floridian fossil vertebrates belonging mostly
to the Pleistocene. Rept. Florida Geol. Survey, Vol. 8, pp. 41-76.
Horrman, C. K.
1890. Schildkréten in Bronn’s Klassen und Ordnungen des Tierreichs.
Leipzig. 442 pp.
LauBg, G. C.
1900. Neue Schildkréten und Fische aus der bohm Braunkohlenformation:
Abhandl. deutsch. naturwiss.-med. Ver. B6hmen ‘‘Lotos’’, Vol. 2,
no. 2, pp. 37-56.
Martruew, W. D.
1924. Third contribution to the Snake Creek fauna. Bull. Amer. Mus.
Nat. Hist., Vol. 50, pp. 59-210.
Meyer, H. von
1852. Ueber Chelydra Murchisoni und Chelydra Decheni. Palaeonto-
graphica, Vol. 2, pp. 237-247.
Peters, K. F.
1855. Schildkrétenreste aus den 6sterreichischen Tertiér-ablagerungen.
Denkschr. math-naturwiss. K]. Akad. Wiss. Wien, Abt. 2, Vol. 9,
pp. 1-22.
1869. Zur Kenntniss der Wirbelthiere aus den Miociinschichten von
Eibiswald in Steiermark. J. Die Schildkrétenreste: Denkschr.
math-naturwiss. K]. Akad. Wiss. Wien, Vol. 29, pp. 111-124.
14 BREVIORA NO. 2
PomEL, A.
1854. Catalogue méthodique et descriptif des vertébrés fossiles décou-
verts dans le bassin hydrographique supérieur de la Loire et surtout
dans la vallée de son affluent principal l’Allier. Paris. 193 pp.
REINACH, A. VON
1900. Schildkrétenreste in Mainzer Tertiirbecken und in benachbarten
ungefahr gleichalterigen Ablagerungen. Abhandl. Senckenberg.
naturf. Ges., Vol. 24, pp. 3-135.
SIEBENROCK, F.
1897. Das Kopfskelet der Schildkréten. Sitz.-Ber. Akad. Wiss. Wien,
Abt. 1, Vol. 106, pp. 245-328.
1907. Die Schildkréten Familie Cinosternidae m. monographisch
bearbeitet. Sitz.-Ber. Akad. Wiss. Wien, Abt. 1, Vol. 116, pp.
527-599.
1909. Synopsis der rezenten Schildkréten mit Berucksichtigung der in
historischer Zeit ausgestorbenen Arten. Zool. Jahrb., Suppl.,
Vol. 10, pp. 427-618.
STEFANO, G. DE
1903. Nuovi Rettili degli strati a fosfato della Tunisia. Bol. Soc. Ital.
Ecol., Vol. 22, pp. 51-80.
TEppPNER, W.
1915. Ein Chelydra-Rest von Goriach. Mitt. naturwiss. Ver. Steiermark,
Vol. 51, pp. 474-475.
WIuuiaMs, E. E.
1950. Variation and selection in the cervical central articulations of
living turtles. Bull. Amer. Mus. Nat. Hist., Vol. 94, pp. 511-561.
ZANGERL, R.
1945. Fossil specimens of Macrochelys from the Tertiary of the plains.
Fieldiana, Geol. Ser., Vol. 10, pp. 5-12.
ZDANSKY, O.
1924. Ueber die Temporalregion des Schildkrétenschadels. Bull. Geol.
Inst. Univ. Upsala, Vol. 19, pp. 89-114.
1952 A STAUROTYPINE SKULL 15
PLATE 1
Staurotypus salvinit M.C.Z. 4989: Dorsal, ventral, anterior and lateral views
of skull. X about 114.
16 BREVIORA NO. 2
TES +
Lor*5
PLATE 2
Xenochelys formosa Princeton 13686: Dorsal, ventral, anterior and lateral
views of referred skull fragment. About natural size.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. May 23, 1952 NUMBER 3
NOTES ON SIPHONOPHORES
3. Nectopyramis spinosa n. sp.
By MARY SEARS!
In a number of samples collected by the ‘‘Dana”’ in tows with 1000
to 3000 meters of wire out, there were several nectophores and bracts
with rows of peculiar jagged spines (Fig. C). These spines are quite
unlike those on other deep-dwelling species, such as V ogtia pentacantha
Kolliker, V’. spinosa Keferstein and Ehlers, and Athorybia rosacea
Forskal. In fact, the specimens have little in common with either the
Hippopodidae or the Anthophysidae. However, one nectophore was
described by Moser (1925, p. 425, Pl. 25, figs. 5-7) as Hippopodius (?)
cuspitata, although it actually closely resembled the specimens found
in the “Dana” collection. The chief difference is that hers had the
peculiar jagged spines scattered irregularly over its surface, whereas
on the ‘‘Dana” specimens these are arranged in discrete rows. Never-
theless, the spines themselves seem to be identical in structure. Like
the ‘‘Dana’’ nectophores, most of the characteristics described and
figured by Moser (1925) make it appear almost certain that her speci-
men was also a Nectopyramis, not a Hippopodius. In all examples, the
shape of the nectosac, its relation to the hydroecium, and the fact that
the nectophores have all been taken singly and at considerable depths
are all peculiarities of Nectopyramis. On the other hand, the soma-
tocyst is tubular and not “represented by a series of divergent canals”’
(Bigelow, 1911a, p. 338) as defined in the brief description of the genus.
The original account was based on one species, Nectopyramis thetis.
Soon thereafter a second species, Nectopyramis diomedeae, with many
of the same characteristics was recorded (Bigelow, 1911b). Since then
no additional species have been reported, and little more has been
1 Papers from the ‘‘Dana”’ Collection No. 38, and Contribution No. 607 of the Woods Hole
Oceanographic Institution.
2 BREVIORA NO. 3
learned about the genus. Hence, there has been little reason for
modifying the original description. Nevertheless, our ideas concerning
the relationships of this genus have changed considerably. Thus, the
family of the Monophyidae has been broken up and the individual
genera transferred to other well-established families (Totton, 1932,
p. 328) with which they had obvious affinities. Nectopyramis, there-
fore, is now referred to the Prayidae. In this family, the somatocyst
and radial canals are structurally quite varied; in some, they are
simple, in others branched. At first, these were used as a criterion for
separating genera (Bigelow, 1911b), but more recently, a transitional
form with branched radial canals and a simple somatocyst has been
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discovered (Leloup, 1934, p. 11). This suggests that when more is
known about this family, differences in the structure of somatocyst
and canals may prove to be of specific rather than generic value.
If, then, we consider that the shape of the somatocyst is in all
probability a specific character in Nectopyramis, as may be the case
among other prayids, we have two species originally described by
Bigelow (1911la; 1911b), N. thetis and N. diomedeae, both of which had
branched somatocysts, but simple radial canals and two species,
N. cuspitata Moser and N. spinosa n. sp. (described below) with simple
tubular somatocysts. The two latter are both spiny, but the two are
readily distinguished for Moser’s species is irregularly covered by the
spines, but in spinosa these are arranged in regular rows along more
or less distinct ridges.
1952 NOTES ON SIPHONOPHORES 3
NECTOPYRAMIS SPINOSA N. sp.
The type specimen is a nectophore about 5 mm. in length taken at
“Dana” St. 3933" at 11°18’S., 50°13’E. on 20 December, 1929, in a
stramin net 150 cm. in diameter towing with 2000 meters of wire out.
The type specimen will be deposited in the University Museum,
Copenhagen, Denmark.
Other specimens were taken as follows:
St. 3556". 2°52’N., 87°38’W. 14. IX. 1928.
S150. 2000 m. wire. 1 nectophore.
Stabi(.4 15 28's5 160739 B.. 4230 TIT... 1929)
S150. 2000 m. wire. 2 bracts.
St. 3920"". 1°12’N., 62°19’E. 9. XII. 1929.
S150. 1000 m. wire. 1 bract.
St, 3964")., 25:19'S., 36:13 BE.) 15. 11. 1930.
E300. 3000 m. wire. 1 bract.
St. 3998. 7°34’S., 8°48’W. 1. III., 1930.
S150. 1000 m. wire. 1 bract.
Nectophore (Fig. A). The unique rows of jagged spines make it
obvious that the nectophore is new. Although probably flattened on
preservation, in dorsal or ventral view, it is roughly triangular much
as are the nectophores of the other known species of Nectopyramis.
The nectosac opens into the base of the triangle and like N. thetis “is
comparatively small, shallow, and saucer-shaped” (Bigelow, 1911a,
p. 338) with four primary radial canals along its sides. It is uncertain
whether or not these eventually branch because of preservation. A
large deep hydroecium lies just above it with a slit-like opening on the
ventral surface. This extends the entire length of the hydroecium.
The somatocyst insofar as can be determined appears to be a tubular
rod extending from the base of the hydroecium to slightly above its
apex. The stem and appendages are entirely missing.
Bract (Fig. B). Although bracts have not been found attached to
the nectophore, there seems to be little doubt that the bracts with
rows of the same jagged spines belong to this species. These are
globular, about 2.5 mm. high and slightly more than 3 mm. wide. As
in the nectophore, the hydroecium is so deep that it extends somewhat
into the upper half of the bract. Closely associated with the hydroe-
cium is a long tubular somatocyst which follows the outline of the
bydroecium. On two of the specimens, two threadlike branches are
given off dorsally and appear distally to have a globular connection.
4 BREVIORA No. 3
BIBLIOGRAPHY
BicELow, H. B.
191la. Biscayan plankton collected during a cruise of H.M.S. ‘“‘Research”’
1900. Pt. XIII. The Siphonophora. Trans. Linn. Soc., London,
Zool., 2nd ser., 10(10): 337-357, pl. 28.
1911b. XXIII. The Siphonophorae. Reports on the Scientific Results of
the Expedition to the Eastern Tropical Pacific, 1904-1905... .
Mem. Mus. Comp. Zool. 38: 171-402, 32 pls.
1931. Siphonophorae from the Arcturus Oceanographic Expedition.
Zoologica. N. Y., 8(1): 525-592, figs. 185-220.
BicELow, H. B., and M. Spars
1937. H. 2. Siphonophorae. Rept. Danish Ocean. Exped., 1908-10, to
the Mediterranean and Adj. Seas, Biol. 2: 1-144, 83 text figs.
LELoup, E.
1932. L’eudoxie d’un siphonophore calycophoride rare, le Nectopyramis
thetis Bigelow. Bull. Mus. Hist. Nat. Belg. 8(3): 1-8, 5 text figs.
1933. Siphonophores calycophorides provenant des campagnes du Prince
Albert I de Monaco. Rés. Camp. Sci., Monaco, 87: 1-64, 1
double plate.
1934. Siphonophores calycophorides de l’Océan Atlantique tropical et
austral. Bull. Mus. Hist. Nat. Belg. 10(6): 1-87, 15 figs.
1936. Siphonophores calycophorides (suite) et physophorides provenant
des campagnes du Prince Albert ]* de Monaco. Rés. Camp. Sci.,
Monaco, 93: 1-35, 2 double plates.
Le.oup, E., and E. HENTscHEL
1935. Die Verbreitung der calycophoren Siphonophoren im Siidatlan-
tischen Ozean. Wiss. Ergeb. Deutschen Atlantischen Exped... .
“Meteor” 1925-1927, 12(2): 1-31, 17 text figs.
Moser, F.
1925. Die Siphonophoren der Deutschen Siidpolar-Expedition. Deut-
schen Siidpolar Exped. 17, Zool. 9: 1-541, 33 pls., 61 text figs.
Totton, A. K.
1932. Siphonophora. Great Barrier Reef Exped., 1928-29, Brit. Mus.
(N. H.), Sci. Repts. 4(10): 317-374, 36 text figs.
1936. Plankton of the Bermuda Oceanographic Expeditions. VII.
Siphonophora taken during the year 1931. Zoologica, N. Y., 21(4):
231-240.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. JUNE 9, 1952 NuMBER 4
A UNIQUE REMOPLEURIDID TRILOBITE
By H. B. Wuirrincton
INTRODUCTION
The species described below, and made the type of a new genus,
has been known for more than 75 years. Previous illustrations are
both few and inadequate, and no attempt had been made to reconstruct
the exoskeleton. When such an attempt is made (text-fig. 1) this
unique remopleuridid is shown to have been a most unusual trilobite,
in which the long, forked hypostome reached back to the anterior
margin of the pygidium. It could not enroll, as it would seem that
many (or most) trilobites could. If the hypostome was rigidly attached
to the cephalic doublure, then only very restricted movements of the
thorax and pygidium, relative to the cephalon, were possible.
I am indebted to Dr. G. Arthur Cooper, U. S. National Museum,
Washington, D. C., for allowing me to study and photograph material
in his charge.
SYSTEMATIC DESCRIPTION
Family REMOPLEURIDIDAE Hawle and Corda, 1847
Genus HYPODICRANOTUS gen. nov.
Type species. Remopleurides striatulus Walcott, 1875, pp. 347-349,
fig. 27, from C. D. Walcott’s locality in the Trenton limestone of
Trenton Falls, Oneida Co., New York.
Discussion. The cephalon of Hypodicranotus striatulus differs from
that of the type species of Remopleurides (Whittington, 1950, pp. 540-
2 BREVIORA NO. 4
543, Pl. 70, figs. 1, 2, 4, 5) in that: (a) the anterior part of the glabella,
the tongue, projects for a greater distance in front of the eyes; (b) three
pairs of glabellar furrows are present, and the second is relatively
farther back than that of Remopleurides; (ec) the broad spine that
curves back beside the thorax is not the genal spine, but a lateral
cephalic spine; (d) the hypostome is long and deeply forked, not a
Text-figure 1. Hypodicranotus striatulus (Walcott, 1875). Reconstructions
of ventral (left) and right lateral (right) appearance of outline of exoskeleton,
approx. X 3, based upon M. C. Z. Nos. 1616, 1617. The positions of the raised
lines on the cephalic and thoracic doublure and the hypostome are indicated
in the ventral view. m = median suture; p = pit in doublure. In both draw-
ings a general outline of the pygidium is indicated by dotted lines.
sub-rectangular plate wider than long. The thorax of Hypodicranotus
probably consists of eleven segments, and is like that of Remopleurides,
with the wide axis, prominent articulating processes and sockets,
diagonal pleural furrows, and backwardly curved and pointed pleurae.
1952 UNIQUE REMOPLEURIDID TRILOBITE 33
The axial and pleural furrows in //ypodicranotus are shallow, there are
no axial spines, and no segment has the pleural spines exceptionally
elongated. The pygidium of Hypodicranotus is poorly known, but
appears to be rectangular in outline, longer than wide, with the axis
much shorter than that of Remopleurides. The dorsal exoskeletons of
other remopleuridid genera may readily be distinguished from that
of Hypodicranotus or Remopleurides.
Hypodicranotus is known from the Trenton Group of New York
and Ontario, from rocks of the same age in Wisconsin, the Prosser
limestone of Minnesota, the Kimmswick limestone of Missouri and
Illinois, the Viola limestone of Oklahoma, and from the Ordovician
strata at Silliman’s Fossil Mount, Baffin Island.
HypopIcRANOTUS STRIATULUS (Walcott, 1875)
Plate 1, figs. 1-10; text-figure 1.
Lectotype (selected Raymond, 1925, explanation of Plate 3). Mus.
Comp. Zool. No. 1616, original of Walcott, 1875, p. 347, fig. 27A,
from the dark-gray colored Trenton limestone of Trenton Falls,
Oneida County, New York. The large collection from this locality in
the Museum of Comparative Zoology includes the trilobites Ceraurus
pleurexanthemus Green, 1832; Calliops callicephalus (Hall, 1847);
Leonaspis ? trentonensis (Hall, 1847); Diacanthaspis ? parvula (Walcott,
1879); Ilaenus cf. americanus Billings, 1859; [sotelus gigas DeKay,
1824; Flevicalymene senaria (Conrad, 1841), as well as bryozoans,
brachiopods, crinoids and asteroids. This locality, from which both
W. P. Rust and Walcott collected, is probably in the Denmark member
of the Sherman Fall formation of Kay (1937, pp. 267-268).
Paratypes. Mus. Comp. Zool. Nos. 1617, 1618, 1537, same locality
and horizon. Additional material, cranidia and hypostomes, is included
under Mus. Comp. Zool. Nos. 3267-3269. No. 1617 is original of
Walcott, 1875, p. 347, fig. 27a.
Description. Dimensions of lectotype, Mus. Comp. Zool. No. 1616,
in millimetres:
Cephalon:
WWemeule Ga Clbtall) ie cape an. Sock. ye cust areas a ee tA fos igicy: 10.4
emma (CXS OTOL ree cena mac. Nec ett teen eye ds eect she ay ee 18.2
Wircthecttsceneall ingles en eee tye teres eo gh 16.4
height atsecond elabellar furrow... 2. ~.2. 6. sete oe oe 6.5
Maximuniswidth across palpebral lobes .......-:...-...5.- Ieee
WEN HME OMEN CU GWC neh. nhs Cos cto noae mae aac ree se P ez
4 BREVIORA NO. 4
Thorax:
Widthvaxis first Seamentees ecu eee ae ory eo oe eee 10.2
: ‘Smumith)> “> approxi ee ee eee ee 4.3
“‘ first segment (across tips of pleurae)................13.6
‘* seventh segment (across tips of pleurae)............ Tet
Length (sagittal) complete thorax, approx................. 13.0
Dimensions of hypostome, Mus. Comp. Zool. No. 1617, in milli-
inetres:
bength: (Saritbtall ic meh ets doko ose heed het a oe eee OZ
Re (ExSAriblalmn utp hue a1. oo eae eee ae ee 20.2
Maximum width (at about half the length)............... 7.4
Widthsofantadledbodyss... ee. pei ioe ee eee 3.7
The cranidium includes the occipital ring, glabella and tongue,
and palpebral lobes. The occipital ring is longitudinally flat, trans-
versely moderately convex, bounded laterally by the posterior branches
of the facial sutures, which are straight but run diagonally back from
the posterior corner of the eye lobe, then curve to run into the posterior
margin at right angles. Inside and parallel to the suture is a faint
depression, the axial furrow, and just inside the point where the suture
cuts the posterior margin is a rounded notch, the articulating socket.
The occipital furrow is shallow. Between the eye lobes the glabella
and palpebral lobes are gently and evenly convex both longitudinally
and transversely. In the mid-line the tongue of the glabella of the
lectotype projects forwards 1.8 mm. in front of the eye lobe, and is
convex and bent down so that the anterior part overhangs the sutural
margin. There are three pairs of glabellar furrows, visible because they
are both unornamented and slightly depressed (PI. 1, figs. 2, 4).
Each furrow runs in a curve convex forwards, and they are equally
spaced from each other, the distance (exs.) between them slightly
less than the equal distance (exs.) of the outer end of the first from a
point opposite the posterior end of the eye lobe, and the outer end of
the third from a point opposite the anterior end of the eye lobe.
The furrows commence a short distance inside the palpebral furrow,
and do not extend to the mid-line. The first is the most strongly
convex, the second commences farthest out, and the third farthest in.
The palpebral rim is flat, depressed slightly below the lobe, outwardly
sloping, widest posteriorly, narrowing forwards and disappearing at
the anterior end of the eye lobe. Posteriorly the palpebral furrow runs
into the junction of the extremity of the occipital furrow and the
axial furrow. The eye lobe is long, in dorsal aspect curved, most
strongly in the posterior part. The eye surface is gently convex
1952 UNIQUE REMOPLEURIDID TRILOBITE 5
transversely, and slopes steeply. The arrangement of the minute,
convex facets is shown in Plate 1, figure 10. The outer margin of the
eye lobe is defined by a narrow, convex border which commences at
the posterior corner and runs forwards to merge anteriorly with the
border of the cheek. The anterior branches of the facial suture curve
to run at first forwards and inwards from the outer, anterior, corner
of the eye lobe, and then run inwards and upwards to the mid-line.
Thus in anterior aspect (PI. 1, fig. 5) the margin of the tongue is a curve
convex upwards. The free cheek is triangular in outline, outwardly
sloping, widest behind the eye lobe, the genal spine short and pointed.
The cheek narrows forwards to disappear opposite the anterior part of
the eye lobe. From the lateral border opposite the median part of
the eye lobe a broad spine curves back, narrowing and reaching to a
point opposite the axis of the fifth thoracic segment. The inner margin
of the proximal part of this spine is parallel to, and just outside of,
the outer margin of the cheek and genal spine. The doublure of the
cephalon is broad anteriorly, gently convex, crossed by a median
suture. Antero-laterally and laterally the doublure is narrower and
bent upwards. A small pit is situated on the exsagittal line passing
through the anterior end of the eye lobe, and about mid-way across
the doublure (PI. 1, fig. 8; text-fig. 1). The hypostome is long and
forked, and its position relative to the rest of the exoskeleton is shown
in text-figure 1. It is gently convex, in both longitudinal and
transverse directions, the transverse convexity greatest at the curved
anterior margin. The convexity (in ventral view) is interrupted only
by the shallow depression in the median region just in front of the
crotch of the fork. The middle body is circular in outline, faintly
elevated, defined postero-laterally by a distinct furrow. It is divided
into three sub-equal sections by shallow radial depressions, the strong-
est leading back to the crotch of the fork, the other two directed
antero-laterally (Pl. 1, fig. 7). On the inside of the exoskeleton of a
smaller specimen (PI. 1, fig. 9) these sections appear more pronounced,
and on both sides the postero-lateral pair are the more prominent.
Anteriorly the middle body merges with the border, and the anterior
wings appear to be short, triangular, and upwardly directed. Each
postero-lateral border is extended as a prong of the fork, the inner
margin of the prong straight, the outer margin curved, so that the
maximum width of the hypostome is in front of half the length, and
the prong tapers to a sharp point. The doublure of the hypostome
(Pl. 1, figs. 4, 9) is narrow along the lateral border, but extends in
under the fork to a line almost under the margin of the middle body.
6 BREVIORA NO. 4
The inner edge of the doublure is flexed up sharply. Mus. Comp. Zool.
No. 1618 (PI. 1, fig. 4) shows that the doublure of the prong is convex
dorsally, with a low, median, longitudinal ridge.
The thorax is moderately convex transversely, the axis broad,
narrowing rapidly backwards. It seems to consist of eleven segments.
On the left side of the lectotype 10 segments can clearly be seen
(Pl. 1, fig. 2). Behind this only what may be part of the left pleura
of the eleventh segment, and part of the pygidium, are preserved.
Another specimen (PI. 1, fig. 6) shows ten partly disarticulated seg-
ments and an incomplete pygidium The last segment has the pleura
pointing almost directly posteriorly, whereas that next in front has
the pleura directed back and slightly outwards, and the lateral margins
are curved. This penultimate segment is very like the tenth of the
lectotype. The axial ring is moderately convex transversely, the
articulating furrow shallow, the articulating half-ring of length
(sag. = sagittal) about half that of the ring (PI. 1, fig. 6). There is no
axial furrow as such, but inside the articulating sockets and processes
a triangular area (broadest anteriorly) of the outermost part of each
ring is slightly depressed. The inner margin of this area is a diagonal
curved line sub-parallel to the pleural furrow. The articulating
process, on the anterior margin of the segment, is slightly raised as
well as forwardly projecting. Postero-laterally the inflation is con-
tinued across the pleura as a low, curved ridge, dying out at about
two-thirds (exs. = exsagittal) the length. This ridge, and a depression
on the inner side, define the pleural furrow. The pleurae are gently
convex (tr. and exs.), outwardly sloping, the narrow (tr. = transverse)
inner part directed transversely, the outer part curved back and
pointed. The shape and inclination of the pleurae are shown in
Plate 1, figures 1, 2, 6.
Only a small part of the pleural lobes of the pygidium is preserved
in the lectotype. The only other specimen known (PI. 1, fig. 6) is
also incomplete. The outline was evidently rectangular, longer than
wide, the ill-defined axis convex, short, and wide. Behind and beside
the axis the pleural lobes slope down in a curve concave in longitudinal
profile. The lateral and posterior margins, and the doublure, are
unknown.
Scattered over the cranidium, but largest and most closely spaced
on, and adjacent to, the palpebral lobes (though absent from the
palpebral rim), are small crescentic depressions, the points of the
crescent facing forwards, and the concave, anterior margin raised
(Pl. 1, figs. 1, 2, 4). Similar but larger structures are present on the
1952 UNIQUE REMOPLEURIDID TRILOBITE
free cheeks, the genal spine, and the base and outer part of the long
lateral spine (Pl. 1, fig. 1). There is a tiny median tubercle on the
occipital ring, closer to the furrow than the posterior margin. Faint,
small crescentic depressions are scattered on the median part of the
axis of the thorax. Strong, well-spaced, raised lines run approximately
longitudinally on the inner part of the lateral cephalic spine, the
thoracic pleurae, and on the doublure of these areas and the cheeks
(Pl. 1, figs. 1-4, 6, 8). The lines tend to run in curves convex inwerds
on the dorsal surface of the pleurae, and transversely in curves convex
forwards on the dorsal surface of the pygidium. On the inner part of
the doublure of the thoracic pleurae, however, they run in curves
concave inwards. These lines die out on the median part of the
cephalic doublure, and are replaced by faint, anastomosing, transverse
grooves. On the lateral borders and fork of the hypostome (PI. 1,
fig. 7) the longitudinal lines are strongest, and that which runs just
inside the inner margin of the prongs is a prominent ridge. The middle
body shows a faint pattern of narrow, anastomosing ridges running
longitudinally on the postero-lateral sectors. On the anterior sector of
the middle body and the anterior border, fine, well-spaced, anasto-
mosing grooves, like those on the median part of the cephalic doublure,
run transversely.
Discussion. This species was first described by Walcott (1875,
pp. 347-349, fig. 27), and the original material later redescribed
by Raymond (1925, pp. 57-58, PI. 3, figs. 4, 5). An incomplete cran-
idium from the same general locality as the type (U.S. Nat. Mus.
92528) was also figured by Foerste (1920, p. 222, Pl. 22, figs. 18 A-C).
Raymond (1921, p. 31, Pl. 9, fig. 7) described a hypostome from the
Middle Trenton of Trenton, Ontario, and referred it to H. striatulus,
and stated that a cranidium had also been found at Governor Bay,
near Ottawa. Specimens kindly loaned to me by Professor G. Winston
Sinclair, from the Middle Trenton of Lakefield, Ontario, contain
cranidia and hypostomes like those of the type material. A second
species of Hypodicranotus is H. missouriensis (Foerste, 1920, pp.
220-222, Plz 21, fig. 17; Pl. 22, figs. 17A, 17B; Bradley, 1930, pp. 246-
247, Pl. 30, figs. 4-9) from the Kimmswick limestone of Missouri and
Illinois. The type material (U. S. Nat. Mus. No. 78438) includes
cranidia and a hypostome. The outline of the latter, and the prom-
inence of the postero-lateral areas of the middle body, distinguish it
from H. striatulus. The characteristic hypostome of Hypodicranotus
also occurs in the Trenton of Duck Creek Quarry, near Green Bay,
Wisconsin (U.S. Nat. Mus. Nos. 72181, 87687), the Prosser limestone
Ss BREVIORA NO. 4
of St. Paul, Minnesota (U.S. Nat. Mus.), and I have collected one
from about 100 ft. above the base of the Viola limestone, in the road
cut on U.S. highway 77, in Carter County, 21 miles north of Springer,
Oklahoma. The matrix at this locality was a finely-granular, light
grey-brown limestone, which yielded abundant graptolites and the
trilobites Cryptolithoides ulrichi Whittington, 1941, Trinodus_ sp.,
Robergia sp., and an asaphid. The specimen figured by Roy (1941,
p. 155, fig. 114), as Remopleurides sp., from Silliman’s Fossil Mount,
Baffin Island, seems also to be the hypostome of this genus The
genus thus occurs in central and eastern North America in rocks of
Trenton age, and the Baffin Island beds may be of a similar age.
The exoskeleton of Hypodicranotus striatulus, apart from the
hypostome, is like that of other remopleuridids of Middle Ordovician
age, e.g. the Irish specimens (Whittington, 1950, pp. 540-548, PI. 69,
figs. 5-10, Pl. 70, figs. 1-6) and undescribed species from the Edinburg
limestone of Virginia. The latter have a sub-rectangular hypostome,
not forked, like that of the type species (Whittington, 1950, Pl. 70,
fig. 2). The posterior margin reaches back to a point lying no farther
back than the occipital furrow. The long, forked hypostome of Hypo-
dicranotus is not only unique, so far as is known, among remopleuridids,
but its relatively great length is in excess of that of any other trilobite
known to me. It Is evident that here is one genus of trilobites that
could not enroll.
The pit in the antero-lateral cephalic doublure (PI. 1, fig. 8, text-
fig. 1) is a feature which I have observed in several different species
of Middle Ordovician remopleuridids. Silicified specimens show the
pit to be the opening of an upwardly-directed tube, which narrows
inwards and terminates near the lower, anterior corner of the eye
surface. The position and nature of this opening do not suggest that
it is homologous with the Panderian opening, which, if present in the
cephalon, is situated in the postero-lateral cephalic doublure. Not all
remopleuridids appear to show this opening (e. g. those described
by Ross, 1951, pp. 84-91, Pl. 20, do not), and I have not observed a
similar opening in any other group of trilobites.
It is tempting to speculate on the mode of life of the holaspid
Hypodicranotus. It occurs in company with shallow-water marine
forms, many of which are indisputably benthonic. The long hypostome
prevented enrollment, but did provide some protection for the ventral
surface. If no movement was possible at the hypostomal suture, then
the amount of possible movement of the thorax and pygidium in the
vertical plane, relative to the cephalon, must have been severely
1952 UNIQUE REMOPLEURIDID TRILOBITE Q
limited. The mode of articulation of the thorax precludes any con-
siderable movement relative to the cephalon in the horizontal plane.
There is ample room for the appendages to project downwards and
outwards between the hypostome and the thoracic pleurae. But we
know nothing of the type of appendage possessed by remopleuridid
trilobites, and without this information have little basis for speculation
on the mode of locomotion, manner of feeding, ete. Was Hypodicran-
otus a burrowing, crawling, floating, or swimming form? No definite
answer is possible but Iam inclined to think of it as either floating or
swimming.
REFERENCES
Brab.ey, J. H.
1930. Fauna of the Kimmswick Limestone of Missouri and Illinois.
Contrib. Walker Mus., vol. 2, no. 6, pp. 219-290, pls. 23-30.
Forrste, A. F.
1920. The Kimmswick and Plattin Limestones of Northeastern Missouri.
Denison Univ. Bull., J. Sci. Lab., vol. 19, pp. 175-224, pls. 21-23.
Kay, G. M.
1937. Stratigraphy of the Trenton Group. Bull. Geol. Soc. Am., vol.
48, pp. 233-302, pls. 1-10.
RAYMOND, P. E.
1921. A Contribution to the Description of the Fauna of the Trenton
Group. Geol. Surv. Canada, Mus. Bull. 31, Geol. Ser. 38, pp. 1-64,
pls. 1-11.
1925. Some Trilobites of the Lower Middle Ordovician of Eastern North
America. Bull. Mus. Comp. Zool., vol. 67, no. 1, pp. 1-180, pls.
1-10.
Ross, R. J.
1951. Stratigraphy of the Garden City Formation in Northeastern
Utah, and Its Trilobite Faunas. Peabody Mus. Nat. Hist., Yale
Univ., Bull. 6, pp. 1-161, pls. 1-36.
Ox 1S. KK
1941. The Upper Ordovician Fauna of Frobisher Bay, Baffin Land.
Mem. Field Mus. Nat. Hist., Geol., vol. 2, pp. 1-212, 146 figs.
Wa.tcort, C. D.
1875. New Species of Trilobite from the Trenton Limestone at Trenton
Falls, N. Y. Cincinnati Quart. J. Sci., vol. 2, pp. 347-349, fig. 27.
WHITTINGTON, H. B.
1950. Sixteen Ordovician Genotype Trilobites. J. Paleont., vol. 24,
no. 5, pp. 531-565, pls. 68-75.
EXPLANATION OF PLATE I
Hypodicranotus striatulus (Walcott, 1875). Trenton limestone, probably
Denmark member of the Sherman Fall formation, Trenton Falls, Oneida
County, New York, al] X 3, except figure 10, X 17144, Mus. Comp. Zool.
collections.
Figs. 1-3, 5, 8. Lectotype, No. 1616, left lateral, dorsal, right lateral,
anterior, and antero-ventral views, presumed original of Walcott, 1875, p. 347,
fig. 27A, and original of Raymond, 1925, PI. 3, fig. 4.
Figs. 4, 10. Paratype, No. 1618. 4, dorsal view. The right half of the
dorsal exoskeleton of the cephalon and three thoracic segments are preserved;
on the left is seen the lateral cephalic spine, and external moulds of the doublure
of several pleurae. The left, and tip of the right, prongs of the fork of the
hypostome are exposed from the inner side. The hypostome has been slightly
displaced. 10, the anterior portion of the right eye surface, showing the
arrangement of the facets.
Fig. 6. Paratype, No. 1537, dorsal view.
Fig. 7. Paratype, No. 1617, ventral view, presumed original of Walcott,
1875, p. 347, fig. 27a, and Raymond, 1925, Pl. 3, fig. 5.
Fig. 9. No. 3269, dorsal view of interior of hypostome.
BREVIORA Unique Remopceuripip TRicopite. PLATE 1
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. AveustT 29, 1952 NuMBER 5
TRICHOLIMNAS CONDITICIUS
IS PROBABLY A SYNONYM OF
TRICHOLIMNAS SYLVESTRIS
(Aves, Rallidae)
By JAmeEs C. GREENWAY, JR.
Comparison of the type of Tricholimnas conditicius Peters and
Griscom (Proce. New England Zool. Club, vol. 10, 1928, pp. 99-103)
with a series of T'richolimnas sylvestris (Sclater) makes it appear most
probable that the type specimen is an immature female of sylvestris.
Furthermore, some recently discovered notes in the handwriting of
collector Andrew Garrett cast grave doubt upon the theory that
Garrett shot the bird on Apaiang Atoll, between Makin and Tarawa,
in the Gilbert Islands, as Thomas Barbour postulated in the original
description of the supposed species. Both Mr. Peters and Mr. Griscom
agree to this.
Naturalists have wondered since then whether a habitat, such as
this low sandy island affords, could have supported a population of
these rails. The very distance from congeners on New Caledonia and
Lord Howe Island, off the coast of Australia, made this seem unlikely,
quite apart from the disparity of habitats. However, the arguments
brought forward in the original description were difficult to circumvent.
This type and only known specimen of conditicitus was found pre-
served in alcohol in the Museum of Comparative Zoology after a period
of long oblivion. With it was a label: “Kingsmill Islands, 1861,
Andrew Garrett, Collector’. Since it could be proved without question
that Garrett was actually on Apaiang Atoll of the Kingsmill, or what
is now called the Gilbert group, in the autumn of 1859, it seemed likely
that he did collect the bird there and that the date ‘1861’’ represented
the year of acquisition by the museum. However, it now seems quite
2 BREVIORA NO. 5
as probable that the specimen has been confused with a consignment
of specimens that came from the Kingsmills and that the label was
written in the museum under a misapprehension.
Andrew Garrett was a dealer who had in his stock specimens from
all over the world. In a letter to Charles E. Hamlin at the Museum of
Comparative Zoology, written on April 10, 1878, from Huahine,
Society Ids., he says: “In return I shall be glad to receive any land or
fresh water shells from the East Indies and marine shells from any
part of the world except Eastern States, California, and Polynesia.’”
And again, ‘“You can send me in return for this box everything you
can spare from your duplicates no matter from what part of the world
... don’t forget the museum publications for which I will send the
ferns in compensation.”
From Hilo, Hawaii, he writes on Oct. 10, 1856, that he plans a
voyage on the whaler ‘Lydia’, the captain of which was an acquaintance
of his.
In a letter to Jas. M. Barnard dated April 29, 1857, and written at
Hilo, he gives a short history of his life. He had been a sailor and had
visited in past years the ‘“‘Ladrone, Bonin, Loochoo, some of the
Australian and several of the East India Islands ...my taste for
Natural History, which I dearly love, was acquired while visiting
foreign places.”’
The most important bit of evidence has recently been found. A
short note in Andrew Garrett’s handwriting and on the same paper
used by him in his correspondence is entitled “Remarks upon the
Birds of Apaiang one of the Kingsmills Is.’”’ and is reproduced here in
its entirety. It bears no date but was probably written in 1859.
“During my short residence at the above location [Apaiang Atoll,
Sept.-Oct. 1859] I collected one or two specimens of every bird I could
get. As I had not time to prepare their skins, I packed them in alcohol.
There is only one species of land bird, and this I never saw but obtained
one of its tail feathers which I send you. I think it is a species of Hawk.”
(Italics are mine.) Unfortunately this feather cannot be found. Two
species of migrant hawks have been reported from the Palau Island;
possibly a stray bird reached Apaiang.
Andrew Garrett was an excellent naturalist; Barbour has called him
“an extraordinary genius.”’ Although his interests were centered upon
fishes and shells, he knew birds. In a contemporary list of birds,
mammals of Hawaii, sent to the Museum of Comparative Zoology, he
says, (no.) “7 — Phaethon — This fine bird builds its nests and roosts
among the precipices bordering the sea in Hamakua and Hilo.” He
1952 TRICHOLIMNAS CONDITICIUS 3
then describes the bird accurately. And again, (no.) “8 — Fulica —
common about our fresh water ponds.” It is quite unlikely that he
would consider Tricholimnas to be a sea bird. Nor is it likely that he
would mistake the short, decomposed tail feather of T'richolimnas for
that of a hawk.
Unfortunately there is no record of the actual provenance of the
specimen but the weight of evidence points to its having come from
Lord Howe Id. It is probable that it found its way into Andrew
Garrett’s collection by an exchange, or perhaps through the good offices
of a whaling friend.
Further questions occur. Assuming that the type was a member of
a population which had for some time inhabited Apaiang Atoll, why
was it not found by the contemporaries of Garrett? The answer is
that none of them visited the island. However, Kubary, Finsch and
others did land upon islands of neighboring groups and no such speci-
men was recorded. No bird even faintly resembling this has ever been
recorded except from Lord Howe Id. and New Caledonia. If the
assumption is correct, we must presuppose that wind born or ship born
individuals reached Apaiang and no other neighboring islands and that
the population was extirpated a very short time after Garrett visited
there. It would be strange if such a population had established itself
on this flat, almost desert, island in view of the fact that its un-
mistakable relatives prefer forests and hills and that the intervening
islands afford more suitable habitat. That it became extinct in a
number of island groups during the twenty years 1860-1880 is not at
all probable.
Granted that no proof can be offered and that at best we are dealing
with probabilities, the best calculated guess would appear to be that
the type of Tricholimnas conditicius came from Lord Howe Id.
No specimens of sylvestris were to be found in the United States in
1928 when conditicitus was described. Comparison of the type with a
series of twelve in the Rothschild collection, now in the American
Museum of Natural History, New York, cause most of the characters
ascribed to geographical variation to disappear. The paler crown,
throat and underparts, the browner head and throat are doubtless the
result of long immersion in alcohol and it is surprising that more color
change has not taken place. The only differences are then the length
of wing and bill, which are two millimeters shorter than those of the
smallest female of sylvestris available. Since G. M. Allen, in the
original description indicated that the skeleton was obviously that of
4 BREVIORA NO. 5
an immature bird, perhaps it may reasonably be assumed that this
character is due to age.
Measurements, in millimeters, are as follows:
“conditicius” sylvestris lafresnayanus
CE so
WViini oe. | eS ihre ee 132 135-139 133-142 185
Dail. avs. S30 58 58-63 61-68 109
Bill pha: Sarin 45 54-56 47-50 52
Marsus 1K: ahs ae 47 47-50 43-48 54
In the original description the length of the tail of the type of
conditicius is said to be 68 mm. Mr. Peters agrees with me that it
should read 58 mm.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. August 29, 1952 NUMBER 6
CONTRIBUTIONS TOWARD A
RECLASSIFICATION OF THE FORMICIDAE
I. Tribe Platythyreint (Hymenoptera)
By Wiuu1AMmM L. Brown, JR.
Museum of Comparative Zoology
Harvard University
The tribe Platythyreini has included the sole genus Platythyrea
Roger as treated by former authors. My own investigations show that
three additional genera (Probolomyrmex, Escherichia and Eubothro-
ponera) must be included. Probolomyrmex Mayr and Escherichia
Forel have heretofore been placed among the Proceratiini because of
their depigmented condition, atrophied eyes (workers), and especially
their vertical, fused and approximated frontal carinae and the closeness
of the antennal insertions to the median line and to one another,
accompanied by fusion of frontal carinae with the greatly crowded
clypeus. Also, these two genera have, according to the describers of
the included species, only one tibial spur to each of the two posterior
pairs of tibiae. I believe that all of the characters just mentioned are
correlated with adaptation of the insects to hypogaeic or other crypto-
biotic conditions of life; they appear in widely separated genera of ants
and other hymenopters, such as Proceratium, Discothyrea and others
in the Formicidae, Psilobethylus, etc. in the Bethylidae, and so on, as
rather consistent combinations. The similar modifications of doryline
and some other ants may be partly due to hypogaeic or subhypogaeic
adaptation, but it would seem that the legionary habit may somehow
be more important in accounting for this particular structural modi-
fication.
For our present purposes, it will be sufficient merely to recognize
two facts: (1) the characters combining to produce the “‘proceratiine
y BREVIORA NO. 6
habitus” can and do act to form similar-appearing groups of genera,
even though these genera may have the most diverse ancestry; (2) the
number and condition of the spurs of the two posterior pairs of tibiae
have been found to possess much less taxonomic value than Emery
and Wheeler granted in their widely-used keys; in fact, the spurs are
now known to be present as pairs, single units, or even to be absent
in different species of the same genera in tribes where once the spurs
were considered universally constant and tribally diagnostic.
With these facts in mind, one can proceed at once to a more natural
classification of the genera and tribes of Ponerinae. Probolomyrmex
and Escherichia have always seemed anomalous as members of the
Proceratiini, and they are not, in fact, at all closely related to any
other proceratiine genus. Following a slight clue dropped by Mann in
1923, when he mentioned the similarity of the pilosity and sculpture
between his Probolomyrmex boliviensis and the species of Platythyrea,
I have carefully compared the two last-named genera and find that
they agree in an astoundingly complete way. In fact, the point-by-
point agreement is so close that I must consider Probolomyrmex to
represent a direct derivative of Platythyrea modified for a highly
cryptobiotic existence in the same way, as I also believe, that the
proceratiines are only modified ectatommines. If Platythyrea and
Probolomyrmez are to be associated, then Escherichia should probably
go along with them. While I have never seen an example of the latter
genus, it appears from all accounts to be very close to Probolomyrmez,
and may even be synonymous and representative of an ergatoid or
other intermediate female caste of one of the Ethiopian Probolomyrmex
species.
A fourth genus that must be included in the Platythyreini is Eubo-
throponera, in spite of its unfortunate name. In his original diagnosis
of this Australian group, Clark stated, “ . . . certainly close to Bothro-
ponera Mayr;” he called the tarsal claws “simple’’ and gave the palpal
formula as maxillary 4, labial 2 segments. I have examined types
(Museum of Comparative Zoology) of all the Eubothroponera species
described to date except E. brunnipes Clark, and find that all species,
at least in the worker caste, possess single, small but distinct median
teeth on all tarsal claws. The palpi in this genus are unusually long
for a ponerine group, and the maxillary pair may reach the foraminal
border behind if fully outstretched. Most of the few specimens
available were inconveniently situated or else had the maxillae re-
tracted, but in spite of this it is clear that all six species seen have
more of both maxillary and labial palpal segments than Clark claims
1952 PLATYTHYREINI (HYMENOPTERA) 3
for them. In E. reticulata Clark and E. tasmaniensis (Forel), a full
count was made under good conditions, and in these species the formula
was maxillary palpi 6, labial palpi 4 segments. The basal segment,
especially in the maxillary palpus, is short, fairly broad, and the
segment most likely to be overlooked. Comparing the proportions of
the visible segments of the palpi of the less favorably situated speci-
mens with those upon which a direct count could be made, I feel quite
safe in assuming that the six species I have seen all possess a 6, 4
formula in the worker.
Now this formula is the primitive one in Platythyrea, as I can con-
firm (vide infra), and since all other characters save one combine to
link the two genera, they must at least be placed in the same tribe.
In fact, the characters separating them are relatively minor ones,
though constant and consistent in combination in the series before me.
The body in Eubothroponera is smaller and generally more compact
(less slender and elongate), and the characteristic sculpture of Platy-
thyrea is less well developed in Eubothroponera, but still basically the
same on head and alitrunk. The pilosity must be used to separate the
two genera until a study can be made including all castes of a large
number of species belonging to both (see key, below).
In examining the palpi of some representative species of Platythyrea
for this work, it was confirmed, as has been long known, that certain
_ of them have a palpal formula of 6, 4. What is not so generally known,
or perhaps even unknown until now, is that certain species of the Old
World tropics possess lesser numbers of segments. The New World
species P. angusta Forel and P. strenua Wheeler and Mann have
elongate palpi, with the 6, 4 segmentation; the basal segment, par-
ticularly of the maxillary palpi, is very short, and the total similarity
of the palpi to those of Eubothroponera is very striking. Among the
Old World forms, P. cribrinodis (Gerstaecker) very definitely has short
palpi, formula 3, 2. In P. schultzei Forel, the maxillary palpi have 2
clearly visible segments each, and there may possibly be a third small
basal one, though no third segment could be seen in the single specimen
examined; the labial palpi are definitely 2-segmented. Other species,
such as P. sagei Forel, were not advantageously placed for a definite
palpal count, but had short palpi with formulae almost certainly under
those of angusta for both pairs. Also seen in one doubtfully determined
species from the Old World were palpi with definite counts of 6 and 4,
but very short in overall proportions. It seems that Platythyrea
species vary considerably from one to the next in palpal characters,
and anyone possessing the material needed for a thorough survey of
4 BREVIORA NO. 6
the group by means of dissection may be able to correlate this vari-
ation with other characters, and thereby be able to recognize more
than one group or genus among the array of Platythyrea species.
Furthermore, the males of Platythyrea seem, from external exami-
nation only, to have very distinctive terminalia worthy of further
study.
Key to the genera of the tribe Platythyreini, based on the workers
and probably applicable to the females
1. Frontal carinae and clypeus fused and projecting anteriorly over the
mandibles; antennae somewhat incrassate apically, inserted close together
on the anterior part of the clypeo-carinal process. (Small, usually pale-
colored forms, under 4 mm. in length. Ethiopian, Neotropical, Indo-
TATISOr TAT) eb te ee ea, hes Re RE ac nell ie cia ee ha oh Probolomyrmex Mayr'
Escherichia Forel
Frontal carinae and clypeus not projecting anteriorly over mandibles;
antennae inserted distinctly posterior to the clypeus and apart from one
another, the insertions covered by the broad lobes of the frontal carinae,
funiculus not or scarcely incrassate in most forms. (Larger, usually
pigmented forms, mostly over 4 mm. in length).................... 2
2. Distinct erect pilosity present and widely distributed (widespread in
Acasbrallis; siecertet Sei Soak biatch Sen aa oe Eubothroponera Clark
Distinct erect pilosity limited to mouthparts and gastric apex. (Tropico-
politan; occurring naturally or as tramp species in some warm temperate
L:CPat Otc) | eee dy OMEN: Aa ace Rae mR NE ROSE Berend Platythyrea Roger
Below are listed the platythyreine genera as known to me at present,
and also a citation of the genotypes and the principal references to the
literature. The references are not intended to be complete, but are
designed to give the interested worker a start toward the gathering of
pertinent titles.
PLATYTHYREA Roger
1863, Berlin. Ent. Zeitschr., 7: 172. Genotype: Pachycondyla punctata Fred.
Smith, 1858, soldier (recte worker), male; designated by Bingham, 1903.
Besides the characters cited in the key (above), it should be men-
tioned that this genus, and also Eubothroponera, are further marked by
1 Probolomyrmex and Escherichia are supposed to differ in that the latter has small compound
eyes and a discernible postmesonotal groove or line, but it is important to note that Forel
himself, in the original description of Escherichia, compared the novelty with several proceratiine
genera, but made no mention of Probolomyrmex! Certain Probolomyrmezx (P. boliviensis Mann,
P. parvus Weber) are known only from the female caste.
1952 PLATYTHYREINI (HYMENOPTERA) 5
having a complete and apparently functionally flexible suture sepa-
rating the pro- and mesonotum, and also by having a large and a
smaller spur on each one of the two posterior pairs of tibiae.
Emery’s survey of the world species (1911, Gen. Ins., Fase. 118,
pp. 28-30) is comprehensive for its time, but is now far out of date
due to the addition of many forms since it was written. Wheeler’s
“Ants of the Belgian Congo” (1922, Bull. Amer. Mus. Nat. Hist.,
45: 57-60, 758-761, 1007) gives references to African and Malagasy
species.
EUBOTHROPONERA Clark
1930, Proc. R. Soc. Victoria, Melbourne, (n.s.) 43: 8-9. Genotype: Eubothro-
ponera dentinodis Clark, 1930, worker; original designation, nec E. tasmani-
ensis (Forel) designated by Clark in 1934.
With the original generic description, Clark characterized and
figured three species (pp. 9-11, fig. 1): FE. dentinodis Clark (p. 9,, fig. 1,
nos. 6, 6a); E. micans Clark (p. 10, fig. 1, nos. 7, 7a); FE. bicolor Clark
(p. 11, fig. 1, nos. 8, 8a); he also included in the genus Forel’s Pachy-
condyla (Bothroponera) tasmaniensis (p. 11), and gave a “‘key”’ (p. 9)
to the three Western Australian species newly described.
In 1934 (Mem. Nat. Mus., Melbourne, No. 8, pp. 32-34, pl. 2,
figs. 15-17) Clark redescribed the worker of E. tasmaniensis (Forel)
(p. 32, fig. 15) from Tasmania and described the new species E. reticu-
lata (p. 33, fig. 16) from New South Wales and F. septentrionalis (p. 34,
fig. 17) from Queensland.
A seventh and last species was added by Clark when he described
E. brunnipes (1938, Proce. R. Soc. Victoria, Melbourne, 50: 361-362,
fig. 3) from Reevesby Island, Sir Joseph Banks Group, South Australia
(biology, loc. cit., p. 356).
The above, I think, includes all the recognized species of Eubothro-
ponera, each reported only from the worker caste. The species are
quite uniform in structure, and it is felt that Clark has failed in some
instances to properly differentiate them; consequently, a review of the
genus is necessary.
6 BREVIORA NO. 6
PROBOLOMYRMEX Mayr
1901, Ann. Naturhist. Hofmus. Wien, 16: 2-3. Genotype: Probolomyrmex
filiformis Mayr, 1901, worker; monobasic.
M. R. Smith (1949, Proc. Ent. Soc. Washington, 51: 38-40) has
reviewed briefly the five species known to that date. A sixth species
was described at about the same time by Weber (P. parvus Weber,
1949, Amer. Mus. Novit., No. 1398, pp. 3-4, fig. 2, female) from
Africa. A description with a good habitus drawing of the female type
of P. boliviensis is given in the original reference by Mann (1923,
Psyche, 30: 16-18, fig. 2). A seventh species, apparently remaining
undescribed, has been taken in the Canberra region of eastern Australia
by Mr. Tom Greaves.
EscHERICHIA Forel
1910, Zool. Jahrb. Syst., 29: 245-246. Genotype: Escherichia brevirostris
Forel, 1910, op. cit., pp. 246-247, worker; monobasic.
Although there are entries in the catalogs of Emery and Wheeler,
this genus and species seems not to have been reported a second time.
It should be noted, however, that Weber’s description of Probolomyrmex
parvus, cited above, may just possibly apply to the female of Esche-
richia brevirostris when the latter becomes known in association with
workers, as the characters and type localities given for the two species
do not, to my mind, exclude the possibility of conspecificity.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. SEPTEMBER 29, 1952 NUMBER 7
ON THE NOMENCLATURE OF THE
PACIFIC GRAY WHALE
By WILLIAM E.. SCHEVILL!
The gray whale of the Pacific has until recently been called
Rhachianectes glaucus (Cope). In 1937 van Deinse and Junge published
an important paper of which the main part is a thorough comparative
study of subfossil Eschrichtius robustus bones from the Netherlands,
comprising one adult cranium and two juvenile crania, one of which
was associated with the mandibles, scapulae, and other parts of the
skeleton. These they compared with an adult Rhachianectes glaucus
skeleton in the British Museum, as well as with the published material
on both forms. Although the subfossil bones are naturally worn and
fragmentary, as well as being for the most part juvenile, they present
ample characters to indicate that they and the gray whale are conge-
neric. At this point I would like to emphasize that the following
criticisms of van Deinse and Junge’s conclusions are directed solely at
their taxonomy; this involves no disparagement of their important and
valuable osteological work, which has demonstrated the generic
identity of Eschrichtius Gray 1864 and Rhachianectes Cope 1869
(= Agaphelus Cope 1868). But they also argue for specific identity,
and thus unite Rhachianectes glaucus (Cope) 1868 with Eschrichtius
robustus (Lilljeborg) 1861. Moreover, they argue that Dudley’s scrag
whale (1725), named Balaena gibbosa by Erxleben (1777, p. 610), is
the same form, and therefore say (1937, p. 181), ‘After this it is clear
that the name of this whale must be changed into Eschrichtius gibbosus
(Erxleben). We must emphasize that with the bones we have now at
hand we cannot give any argument that the Pacific and Atlantic
representatives of Eschrichtius should be specifically different.” But
by the same token these bones cannot prove that these whales are
specifically identical.
1 Contribution No. 626 of the Woods Hole Oceanographic Institution.
2 BREVIORA NO. 7
Additional and better material of VY’. robustus may present more
specific characters, but until such material turns up, it seems unfortu-
nate to take a well-grounded name of a well-characterized living form,
known from complete specimens, and replace it with that of a form
so imperfectly known from such incomplete material that specific
characters cannot be entirely established; instead it seems better not
to change the name of the well-known form.
It is even less desirable to use the name (gibbosa Erxleben) of a form
which from the beginning has been an unrecognizable literary curiosity.
Considering the available information, which is scanty though oft
repeated, the most that can be said for Balaena gibbosa Erxleben is
that it may be an Eschrichtius, but since it is not specifically well
characterized and since nothing sufficiently like it has ever again been
reported from anywhere near the type locality (‘the Coast of New
England’’) or elsewhere in the Atlantic, it seems to be stretching
probability to submerge in it known forms from other regions. De-
tailed consideration of the uncertainties in attempting to identify
Dudley’s secrag whale (Balaena gibbosa Erxleben) are omitted here.
From time to time cetologists have tried to identify this beast, but
have always been frustrated by various irreconcilable data, the chief
of which is perhaps the utter lack of specimens. Cope’s attempt to
embody gibbosa Erxleben was undermined by his confusion over a
dilapidated specimen of Balaenoptera acuto-rostrata, so that he formally
withdrew in 1884 (p. 1124). Although he did not publish details, it
is probable that Cope used much the same reasoning as van Deinse
and Junge in associating Dudley’s scrag and the Pacific gray whale
in the same genus. Thus it appears best to continue to use robusta
Lilljeborg for the subfossil European form.
As to the nomenclature of the Pacific gray whale, it seems reasonable
to retain the trivial name g/aucus until better material of E'schrichtius
robustus may demonstrate specific identity (although Cederlund (1939,
p. 282) believes the holotype sufficient for this). It is therefore sug-
gested that the living Pacific gray whale be called Eschrichtius glaucus
(Cope).
This status may be summarized thus:
Genus
Eschrichtius Gray 1864 (p. 350). Type species (by subsequent
designation by Lilljeborg 1865, p. 494): Balaenoptera robusta
Lilljeborg 1861.
Species
Eschrichtius robustus (Lilljeborg)
Balaenoptera robusta Lilljeborg 1861, p. 602.
The Grias6 whale.
Subfossil in northwestern Europe (English Channel to Baltic
Sea).
1952 PACIFIC GRAY WHALE 3
Uschrichtius glaucus (Cope)
Agaphelus glaucus Cope 1868a, p. 160, 1868b, p. 225.
Rhachianectes glaucus Cope 1869, p. 15.
The California or Pacific gray whale; devilfish.
Recent in North Pacific Ocean (Baja California to Korea).
REFERENCES
Cederlund, B. A.
1939. A subfossil gray whale discovered in Sweden in 1859. Zoologiska
Bidrag fraan Uppsala, 18, pp. 269 - 286, 5 pls.
Cops, E. D.
1868a. [Remarks on Cetacea at meeting of 23 June.] Proc. Acad. Nat.
Sci. Phila., 1868, pp. 159-160.
1868b. On Agaphelus, a genus of toothless Cetacea. Proc. Acad. Nat.
Sci. Phila., 1868, pp. 221-227.
1869. Systematic synopsis of the species of the cetaceans of the west
coast of North America. Proc. Acad. Nat. Sci. Phila., 1869,
pp. 14-32.
1884. Catalogue of aquatic mammals of the United States, by F. W.
True. Review. American Naturalist 18, no. 11 (Nov.), pp.
1123-1124.
vAN DeinsE, A. B., and G. C. A. JUNGE
1937. Recent and older finds of the California gray whale in the Atlantic.
Temminckia, 2, pp. 161-188, pls. 4 - 11.
Dup.tey, Pau
1725. An essay upon the natural history of whales with a particular
account of the ambergris found in the sperma ceti whale. Philos.
Trans. Roy. Soc. London, 33, no. 387, pp. 256-269.
ERXLEBEN, J. C. P.
1777. Systema regni animalis ... Classis I. Lipsiae, pp. xlvili +
636 + [66].
Gray, J. E.
1864. Notes on the whalebone-whales; with a synopsis of the species.
Ann. Mag. Nat. Hist. (8) 14, no. 83 (Nov.), pp. 345-353.
LILLJEBORG, W.
1861. Hvalben, funnaijorden paa Grason i Roslagen i Sverige. Forhandl.
Skandinav. Naturf., 8de Mgde, 1860, Kigbenhavn, pp. 599-616.
1865. [Note quoted in] Gray, J. E., Notice of a new whalebone whale
from the coast of Devonshire, proposed to be called Hschrich-
tius robustus. Ann. Mag. Nat. Hist. (8) 15, no. 90 (June)
pp. 492 - 495.
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BREVIORA
Museum of Comparative Zoology
(AMBRIDGE, Mass. OcToBER 21, 1952 NUMBER 8
A NEW SPECIES OF THE
CYCLOSTOME GENUS PARAMYXINE
FROM THE GULF OF MEXICO
By Henry B. BiagELow and WILLIAM ©. SCHROEDER!
Up to the present time the myxinids, represented in the North
Atlantic by the unibranchiate genus Myzine alone, have been unknown
in the Gulf of Mexico. During 1951, however, the experimental
trawlings of the U. S. Fish and Wildlife Service vessel “Oregon’’, in
the northern part of the Gulf, yielded three specimens of a myxinid
with six pairs of external gill openings that agree closely with the
genus Paramyxine Dean 1904, reported previously from Japan only.’
Paramyzxine, originally described from a single specimen, as the new
species P. atami Dean 1904, is characterized, by its author, by having
the outer “branchial ducts of distinctly unequal length, the most
anterior several times the length of the posterior” (Dean, p. 22), so
that the most anterior pair of external gill openings is opposite the
4th or 5th pair of internal gill sacs; and by having the successive pairs
of gill openings close together. In other known myxinids with more
than one pair of gill openings the successive pairs of external branchial
ducts are of approximately equal lengths, or the Ist pair is only a little
longer than the rearmost pair, and the successive pairs of gill openings
are more widely spaced. On the type specimen the gill openings them-
selves were oval, with their long axes transverse (except for the 6th
opening on the left-hand side, which — receiving the oesophageal
duct — is larger than the others), not round as they are in other
polybranchiate species. But the gill openings were of various shapes
in other specimens described subsequently by Matsubara (1937),
showing that their shape is not a generic character as Dean supposed,
‘ Contribution No. 610, from the Woods Hole Oceanographic Institution.
2 We owe to Mr. Stewart Springer the opportunity of studying these interesting specimens.
2 BREVIORA NO. 8
or even a specific character, but is, likely, dependent to a considerable
extent on the degree of muscular contraction. This leaves only the
relative lengths of the external branchial ducts and perhaps the close
spacing of the gill openings, as compared with other myxinids in which
they are few in number, as bases for separating the genus Paramyxine
from other polybranchiate myxinids. We accept the genus provision-
ally, awaiting the opportunity of examining other myxinids with
corresponding numbers of gill openings.
Characters that Dean (1904, p. 22) included as specific for P. atami
were the precise number (6) of pairs of gill openings; Ist pair of ex-
ternal branchial ducts about 3 times as long as the 6th pair; 12 and
13 lingual teeth in the outer rows with 13 in each of the inner rows;
anterior 3 lingual teeth of each row fused with one another at their
base; white gill openings and conspicuously dark mucous pores. His
illustration also shows a rather pointed snout, resembling that of
Myzxine, but a somewhat broader and more paddle-shaped caudal fin
fold.
The gill openings of the type specimen of atamz occupied a space
about 9 per cent as long as the distance from the tip of the snout to
the most posterior pair of gill openings,! with those on each side
arranged close together in a regular row, and they are arranged simi-
larly in an excellent specimen of atami, from Suruga Gulf, Japan,
received recently by the U. S. National Museum (U. S. Nat. Mus.
No. 161442). But Matsubara’s (1937) observations have shown that
the serial arrangement of the gill openings, like their shapes, may vary
widely from individual to individual, and even between the two sides
of the body of a given specimen, for they were irregularly arranged in
13 of his 14 specimens. Muscular contraction may play a role here.
Matsubara also found that the dental formula is not a precise one, for
his specimens had only 10 or 11 lingual teeth in each row, outer and
inner, with only the first two fused at the base instead of three as was
true of the type specimen.
The Gulf of Mexico specimens agree in general with the accounts
and illustrations of the Japanese atam7 and with the National Museum
specimen, in body form, with rather pointed snout and a caudal fin
fold somewhat wider than that of Myaine. Their barbels are similar
to those of atami in length and in arrangement; they have the same
number (6 pairs) of external gill openings; the external branchial ducts
(as measured by probing) are about as long relative to the 6th pair;
1 As scaled from Dean’s (1904, pl. 1, fig. 3) illustration.
1952 NEW SPECIES OF CYCLOSTOME 33
the foremost pair is situated about as far rearward while the gill
openings cover about as long a space relatively; the distribution of
mucous pores is about the same; also the linear dimensions in general,
as appears from the following tabulations:
Table 1
Distances, in Per Cent of Total Length
Gulf of Mexico atami, atami, U.S.
specimens, 338, Japan type Nat. Mus.
055 and 590 mm. 550 mm. specimen
500 mm.
Snout to foremost mucous pore 7.5- 8.7 6.8 9.0
Snout to foremost gill openings 22.5-23.5 26.0 27.0
Ist gill opening to 6th gill
OVEDIN Pesag 7 Sins susie Shae a 2.4— 5.7 2.5 2.4
6th gill opening to tip of caudal
(TA S10) fo ae ae eee ee 72.0-74.3 (alee: 73.0
Snout to origin of anal fin fold 37.1-50.0 41.4 43.4
Cloacal pocket (center) to tip
of caudal fin fold:.....:..- 13.4-17.5 12.8 1322,
Table 2
Numbers of Mucous Pores on Each Side
Gulf of Mexico atami, type! atami, U.S.
specimens Nat. Mus.
specimen
Snout to foremost pair of gill
openings................. 15-15 to 19-19 19-20 16-16
6th pair of gill openings to
cloacalkpocketinus. + 4244¢-- 44-45 to 57-57 58-59 47-47
Center of cloacal pocket to tip
of caudal fin fold.......... 11-11 to 13-14 ? 10-10
The number of lingual teeth, too, is about the same in a Gulf of
Mexico specimen of 505 mm. (14 and 13 in the outer rows, 11 and 12
in the inner rows) as it was in the type specimen of atami, with the
same number (3) of anterior teeth fused basally in each of the outer
two rows. And while two only, of the anterior teeth in each of the
inner rows are fused in the Gulf of Mexico specimen, three were fused
in the type, and Matsubara (1937, p. 15, text fig. 3) has already
reported and figured Japanese specimens that agreed with the Gulf of
1From Dean 1904, p. 11, text fig. 2, D.
4 BREVIORA NO. 8
Mexico one in this respect. The series of mucous pores on each side
is interrupted near the anterior end of the cloacal pocket on the U. 8.
National Museum specimen of atami (Fig. 6), just as it is in the Gulf
of Mexico specimens (Fig. 5) — in Myzine too, for that matter — to
continue rearward at a higher level. Representation of it in Dean’s
(1904,pl. 1, fig. 3) illustration as continuing unbroken past the cloacal
pocket seems to have been an error by the artist. But the rearward
pores are situated higher above the ventral edge of the caudal fin fold
in the Japanese atami (Fig. 6) than in the Gulf of Mexico specimens
(Fig. 5).
The depth, too, of the body relative to the distance from tip of
snout to Ist pair of external gill openings is greater on all three of the
Gulf of Mexico specimens (34 per cent to 37 per cent) than is pictured
for atami either by Dean (30 per cent) or by Matsubara (22 per cent),
or than it is on the U. S. National Museum specimen of atami (29
per cent). And while the hagfishes are so soft of body that measure-
ments of different specimens cannot be accepted as precisely com-
parative unless taken under equal tension, the difference in this respect
between the two geographic populations may be large enough to be
significant. The shape of the caudal fin fold also differs rather notice-
ably, its lower margin being nearly straight in all three of the Gulf of
Mexico specimens while it is pictured as moderately convex in the two
Japanese specimens of atami which have been illustrated (Dean 1904,
pl. 1, fig. 3; Matsubara 1937, pl. 1, fig. A), likewise in the National
Museum specimen of atami (compare Fig. 5 with Fig. 6).
Furthermore, neither of the two larger Gulf of Mexico specimens
shows any trace of a dorsal fin fold anterior to the level of the cloacal
pocket, whereas Matsubara’s (1937, pl. 1, fig. A) illustration of atami
clearly shows such a fold, extending forward for one third to one half
the distance from the level of the cloacal pocket toward the level of
the 6th pair of gill openings while the dorsal fin fold on the National
Museum specimen of atami is continued forward as a definite though
low dermal ridge to abreast of the 6th pair of gill openings.!
These differences between the Gulf of Mexican and the Japanese
representatives of the genus seem sufficient, in combination, for recog-
nition in nomenclature. We therefore propose for the former the new
specific name springeri, as defined below. What is most interesting,
however, in this case, is not that the Gulf of Mexican population
1The anal fin fold also, of the National Museum specimen of atam7 is continued forward
nearly to the 6th gill opening as a prominent dermal ridge. But the appearance of the latter,
in alcohol, suggests that it was the result of aulmuscr contraction, not a normal] characteristic,
1952 NEW SPECIES OF CYCLOSTOME 5
differs in minor ways from the Japanese — it would be astonishing if
such were not the case — but that two populations so widely separated
should resemble each other so very closely, and that myxinids with
five to seven pairs of external gill openings should have been found
at localities as widely separated as the Gulf of Mexico, Japan and
China, New Zealand and South Africa, but nowhere else in the oceans.!
This instance does not stand alone as it concerns the Gulf, for the
chimaeroid Hydrolagus alberti Bigelow and Schroeder 1951a, recently
discovered in deep water there, seems more closely allied to the Japa-
nese H. mitsukurti (Dean) 1904, than it is to either of the two species
of its genus that are known from the North Atlantic, i.e., affinis (Brito
Capello) 1868, and mirabilis (Collett) 1904, or than it is to H. collet
(Lay and Bennett) 1839, of the Pacific coast of North America.
Equally unexpected was the recent discovery, in the Gulf, in moder-
ately deep water, of a representative (Springeria folirostris Bigelow
and Schroeder 1951), of the rajoid family Anacanthobatidae, for the
latter had been known before only off the Natal coast of southeast
Africa; also of two representatives of the skate genus Cruriraja in
Cuban-Floridan waters (C. atlantis and C. poeyi Bigelow and Schroeder
1948), for this genus, too, had been known only off Natal, southeast
Africa, previously.
Altogether, the bottom zone in the Gulf and around Cuba at 75 to
600 fathoms has yielded three new species of sharks, eleven new
batoids, and one new chimaeroid since 1938, a foretaste of the rich
harvest of unknown fishes to be expected from a more thorough ex-
ploration of the deep slopes and of the sea floors of the Gulf and of
the Caribbean.
PARAMYXINE SPRINGERI, new species”
Type. A specimen, 590 mm. long, Oregon Station No. 489, Lat.
27°44’N., Long. 85°09’W., 254 fathoms, Sept. 29, 1951 (U. S. Nat.
Mus. No. 161512).
Additional material. Another specimen 505 mm. long from the
same trawl haul, and a third, 338 mm. long, from Oregon Station
No. 321, Lat. 27°27’N., Long. 87°19’W., 220 fathoms, April 28, 1951.
The sex has not been determined for any of these specimens.
Description. Dimensions, in per cent of total length, of type
(590 mm.), 505 mm. specimen, and 338 mm. specimen.
Depth of body, at Ist gill openings, 9, 7.8, 8.9.
1 For a synopsis of the species of myxinids, see Holly, 1933.
2 Named after Stewart Springer who discovered this interesting cyclostome.
6 BREVIORA NO. 8
Distance, snout to Ist gill openings, 23.5, 23.2, 22.5.
Distance, Ist gill openings to 6th gill openings, 2.4, 2.6, 5.7.
Distance, snout to origin of anal fin fold, 50.0, 37.1, 39.8.
Distance, center of cloacal pocket to tip of tail, 14.5, 13.4, 17.5.
Depth, in per cent of distance, snout to Ist gill openings, 38, 34, 37.
Number of mucous pores on each side, type (590 mm.), 505 mm.
specimen and 338 mm. specimen:
Snout to Ist gill openings, 19-19, 18-18, 15-15.
1st gill openings to 6th gill openings, 4-3, 3-3, 6-6.
6th gill openings to anterior end of cloacal pocket, 55-56, 57-57,
44-45,
Anterior end of cloacal pocket to tip of caudal fin fold, 13-14, 11-11,
12-12.
Snout obtuse-conical, much as in Myzine; body cylindrical an-
teriorly, about 80 to 100 per cent as thick (transversely) at region of
gill openings as deep there, but increasingly compressed posteriorly,
until only about 38 to 54 per cent as thick as deep at level of cloacal
pocket, and flat sided thence rearward to paper-thin margin of caudal
fin fold. Depth of body about 5.4 to 6 per cent as great as total
length at level of most anterior mucous pores, about 6.3 per cent at
anterior end of cloacal pocket on type, and 6.6 to 8.6 per cent on
smaller specimens.
Fleshy rostrum, anterior to nasal opening, broadly rounded an-
teriorly and wider than long,! the nasal opening conspicuous below it.
Barbels in the typical myxinid arrangement, the Ist pair about two-
thirds as long as the 2nd pair, the 3rd _ pair a little the longest.
Mouth, when closed, with the usual puckered or irregularly
stellate appearance, the conical projection that flanks it on either side
about as long as the 2nd pair of barbels. Thirteen and fourteen lingual
teeth in the outer rows, eleven and twelve in the inner rows of 505 mm.
specimen’, individual teeth conical with slightly blunted tips, curving
a little rearward, the most posterior of each inner row strongly so;
the 5th and 6th tooth longest in each row; the 7th and the following
teeth successively shorter along each row; the rearmost only about
one-third as long as the 5th and 6th. The first three teeth in each
outer row, and the first two in each inner row fused at their bases;
the outer row in each side about 1.2 times as long as the inner row;
the longest teeth in the outer rows about 1.3 times as long as the
longest in the inner rows.
1 Longer than wide in Myzine. :
2 The teeth of the type specimen cannot be counted without dissection.
=~
1952 NEW SPECIES OF CYCLOSTOME
The six pairs of gill openings are low down on the sides, their fore-aft
position as given in Table | (p.3), occupying a shorter space on larger
specimens (2.4 per cent of total length on the type) than on smaller,
the series on the two sides converging rearward, with the 6th pair only
about three-fourths as far apart as the Ist pair on the type, about
four-fifths as far apart on the smallest specimen; the 6th opening on
the left-hand side (receiving the oesophageal duct) larger than the
others as is the rule among myxinids, and irregular in shape, the other
gill openings ranging in shape from round to oval with the long axis
transverse or slightly oblique, the precise shape seemingly dependent
on muscular contraction; the largest (apart from the 6th left-hand) a
little less than one-half as long as the 3rd (longest) pair of barbels.
The gill openings are either in regular serial arrangement on each side
as on the type (Fig. 2) and on the smallest specimen, or more or less
irregular at least on one side, as on the 505 mm. specimen (Fig. 3),
a difference perhaps associated with muscular contraction. The Ist
pair of external branchial ducts (as sounded by probing, on one side
of the body) are about three times as long as the 6th pair, both on the
type and on the 505 mm. specimen.
The mucous pores range in size from easily visible to so minute as
to be discernible only on close examination. Their location on the
body and in serial arrangement is as in the other myxinids; 19 pores
on each side from snout to Ist gill openings on the type, 4 on one side
and 3 on the other abreast the gill openings, 55 and 56 between gill
openings and cloacal pocket and 13 and 14 rearward from the anterior
end of the cloacal pocket on the type specimen; their numbers on the
other specimens as given in Table 2 (p.3). Totals of 77 pores on the
338 mm. specimen, of 88 on the 505 mm. specimen, and 92 on the
590 mm. specimen (type) show that the pores increase in number with
growth. The pores along the section occupied by the gill openings are
ventral from the latter, which it seems is characteristic of the poly-
branchiate myxinids in general.
The pores rearward from the gill openings are nearly in line with
the latter, are low down on the sides nearly to the anterior end of the
slit-like cloacal pocket, where the series is interrupted, with the nine
posterior pores situated at a definitely higher level, a distinctive
character of this species as contrasted with the Japanese atami (p. 4).
On the type, those rearward from the cloacal pocket are also noticeably
larger (hence more conspicuous) than those farther forward, and they
are somewhat more conspicuous than the more anterior pores on the
8 BREVIORA No.8
505 mm. specimen as well. But this regional difference in size of
mucous pores is not a specific character; rather it is likely a phase of
muscular contraction, for the pores of the caudal series on the smallest
specimen are no larger than those anterior to the cloacal pocket.
The point of origin of the anal fin fold, about midway of the body
on the type specimen, is considerably farther forward on the other
two specimens (see Table 1, p. 3) showing that its precise situation
is variable within rather wide limits. On the type, the fold itself is so
narrow that its width is hardly measurable, and it is traceable rearward
only about half the way from its point of origin toward the cloacal
pocket. But it is wider on the other specimens (8 to 13 per cent as
wide as the body is deep), and traceable rearward nearly or quite to
the cloaca. To what degree these differences are associated with
different stages in growth, or in the sexual cycle, is a question for the
future.
Neither the type, nor the 505 mm. specimen shows any trace of a
dorsal fin fold anterior to the level of the cloacal pocket, their backs
being evenly rounded, in which respect they contrast with P. atamz
as pictured by Matsubara (1937, pl. 1, fig. A). And while the smallest
(338 mm.) specimen has a low fleshy ridge all along the back nearly
to the level of the mouth, the fact that it is flanked on either side by
a shallow and indefinite furrow suggests that it is the result of muscular
contraction, rather than a definite and persisting fin fold. The caudal
fin fold fringes the rear section of the trunk, from about opposite the
anterior end of the cloacal pocket on the dorsal side, around to the
posterior end of the cloaca on the ventral side. Its lower outline is
nearly straight as remarked above (p.4). The irregular fluting of its
broadly rounded, posterior-dorsal margin on the smallest specimen
(Fig. 5) seems to represent its normal outline better than its margin
on the type, (Fig. 1) where it seems to have been injured.
One of the most interesting features of P. springeri, reported to us
by its discoverer, is that no one of the specimens discharged any mucus
at all when brought on board, or so little that none was noticed.
Color. All three specimens are greyish brown, faintly tinged with
reddish, below as well as above, the edges of the caudal fin fold and
of the anal fin paler; the gill openings encircled with white, rendering
them conspicuous against the dark background; the mucous pores
appearing as dark brown dots, each of them at the bottom of a shallow
depression of the skin.
Size. The maximum size is not known.
1952 NEW SPECIES OF CYCLOSTOME 9
Range and habits. So far known only from the northern side of the
Gulf of Mexico, at the localities listed on page 5. The depths of
capture, added to the fact that fishermen have never reported it, show
that it is a deep-water species. Nothing else is known of its habits.
REFERENCES
BriceLtow, Henry B., and WILLIAM C. SCHROEDER
1948. New genera and species of batoid fishes. Journ. Marine Research,
vol. 7, no. 3, pp. 548-566.
1951. A new genus and species of anacanthobatid skate from the Gulf
of Mexico. Journ. Washington Acad. Sci., vol. 41, no. 3, pp.
110-113.
195la. Three new skates and a new chimaerid fish from the Gulf of
Mexico. Journ. Washington Acad. Sci., vol. 41, no. 12, pp. 8383-392.
Brito CapELLO, FELIX DE
1868. Catalogo dos peixes de Portugal. Jorn. Sci. Math. Phys. Nat.
Acad. Lisboa, vol. 1, no. 3, pl. 3, fig. 1A.
1868a. Descripcaao de dois peixes novos de Portugal. Jorn. Sci. Math.
Phys. Nat. Acad. Lisboa, vol. 1, no. 4, pp. 314-317.
Co..etTt, R.
1904. Diagnoses of four hitherto undescribed fishes. . . . Selsk. For.,
1904, no. 9, 7 pp.
DrAN, BASHFORD
1904. Notes on Japanese myxinoids. Journ. Coll. Sci. Imp. Univ. Tokyo,
vol. 19, art. 2, 23 pp., 1 pl.
Ho.uiy, MAxIMILiAN
1933. Cyclostomata, in Schultze and Kukenthal, Das Tierreich, lief 59,
62 pp.
Lay, G. T., and E. T. BENNETT
1839. Fishes, in Richardson and others, Zoology of Captain Beechey’s
voyage, pp. 43-75.
MatsuBaRa, K1YoMATsu
1937. Studies on deep sea fishes of Japan. III. On some remarkable
variations found in Paramyzxine atami Dean. Journ. Imper.
Fisheries Inst. Tokyo, vol. 22, no. 1, pp. 18-15, 1 pl.
10 BREVIORA NO. 8
Fig. 1. Type specimen, x 14; the mucous pores somewhat exaggerated.
Fig. 2. Ventral view of anterior part of type specimen, x 14; the mucous
pores somewhat exaggerated.
Fig. 3. Ventral view of gill region of specimen 505 mm. long; natural size;
the mucous pores somewhat exaggerated.
Fig. 4. Lingual teeth of specimen 505 mm. long, from the right-hand side;
outer row above, inner row below, anterior ends to the left, x 2.
Fig. 5. Side view of caudal region of specimen 338 mm. long, x 14; the
mucous pores slightly emphasized.
Fig. 6. Side view of caudal region of P. atami, 500 mm. long, from Suruga
Gulf, Japan, U.S. Nat. Mus.,No. 161442;the mucous pores slightly emphasized.
7 ai? [ 1% ; : i iu \ Rie ania iA bd Dia cae
is OH eh | Td
TA eR Na
ma - iy yk
J f er
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. OcToBER 21, 1952 NuMBER 9
NEW SPECIES OF EARTHWORMS FROM THE
ARNOLD ARBORETUM, BOSTON
By G. E. GAtTEs
if
From Allolobophora caliginosa (Savigny) 1826 there have recently
been split off: A. nocturna Evans 1946, and A. zowana Evans 1948.
The new species were thought to be distinguished by differences in
location of first dorsal pore, extent of male porophores, etc., and es-
pecially by number of segments. These characteristics had not previ-
ously been given such weight in lumbricid taxonomy, may be subject
to variation and, as in the case of segment number, may be tedious
to use when dealing with large numbers of worms.
While working on collections from the Arnold Arboretum and other
localities near Boston, it was found that material with characteristic
caliginosa tubercula pubertates could be rapidly sorted into two groups
according to the presence or absence of genital tumescences on segment
xxxlii. Further study showed that those specimens with tumescences
on xxxili were referable to caliginosa as apparently restricted by Evans
after breeding the worms in the laboratory. The worms with no
tumescences on xxxill were found to be consistently distinguishable
from caliginosa by the same sort of characteristics as in the case of
nocturna and iowana and, as well as those species, worthy of specific
status.
ALLOLOBOPHORA ARNOLDI N. sp.
Type. Museum of Comparative Zoology, cat. no. 4441.
Length, 55-100 mm., diameter, 4-6 mm. Segments, 152-194
(normal specimens). Unpigmented and almost white, or of greyish
appearance or, especially in older worms of second season, with a
brownish coloration. First dorsal pore, on 10/11, 11/12, or 12/13.
Clitellum, on xxvii, 46xxvii, or xxviii to xxxiv or Yoxxxv. Tubercula
3, BREVIORA No. 9
pubertates of double origin, on xxxi-xxxill. Genital tumescences, in-
cluding a and b setae, on 1x—Xi, XXX, XXxll, xxxiv, and occasionally also
On XXVI.
Spermathecal pores, on 9/10-10/11 on ¢ lines, setae, male poro-
phores, calciferous sacs, calciferous glands, typhlosole and last hearts,
etc., as in caliginosa.
Type locality. Arnold Arboretum, Boston, Mass.
Distribution. Known at present only from Boston and vicinity.
Remarks. A. arnold? is distinguished from caliginosa by the constant
absence of genital tumescences on xxxili, occasional presence of tumes-
cences on xxvi (instead of xxvii), slightly greater anterior extent of the
clitellum (xxvii or xxviii instead of xxviii or xxix), greater number
of segments, and perhaps by a slightly more anterior first dorsal pore.
From iowana, arnold: is distinguished by the constant presence of
genital tumescences on xxxiv, constant absence of those tumescences
on xxxill, by the two-part tuberculum about as in calzginosa, possibly
also by a slightly more posterior first dorsal pore, greater number of
segments and absence or slighter development of pigmentation (not
dark reddish brown).
From nocturna, arnoldi is distinguished by the constant absence of
tumescences on xxxiii and xii, smaller size, fewer segments, less re-
stricted male porophores, and possibly by a more posterior first dorsal
pore.
Data as to variation in both caliginosa and arnold: in the Boston
area will be presented, if possible, in another contribution.
He
In lumbricid taxonomy considerable weight has been given in the
past to rather small differences in the tubercula pubertates. Thus, for
instance, the species in the following pairs have been distinguished
from each other by a one-segment homoeosis of the tubercula: Octo-
lasium cyaneum (Savigny) 1826 and O. lactewm (Orley) 1881, xxx-xxxili
and xxxi-xxxiv; Lumbricus rubellus Hoffmeister 1848 and L. castaneus
(Savigny) 1826, xxvili-xxxi and xxix—xxxiil. Further, Dendrobaena
rubida (Savigny) 1826 has been distinguished from D. subrubicunda
(Eisen) 1874, by a one segment difference in length of the tubercula,
XXIX-xxx and xxvili-xxx. Genital tumescences, as indicated above,
may also be of considerable value in lumbricid taxonomy.
Accordingly, it seems advisable to treat a worm distinguishable at
present from the caliginosa-complex only by differences in tubercula
1952 NEW SPECIES OF EARTHWORMS 3
and tumescences as specifically distinct.
ALLOLOBOPHORA MOLITA Nn. sp?
Type. Museum of Comparative Zoology, cat. no. 4442.
Length, 81 (+ ?) mm., diameter, 5 mm. Segments, 140 (+ ? pos-
terior amputee?). Pigmentation lacking. First dorsal pore in 712/13.
Clitellum on xxviii-xxxiv, possibly reaching slightly onto xxvii.
Tubercula pubertates, longitudinally placed bands in bc, extending
across entire lengths of xxx—xxxiil; with lateral margins nearly smooth,
but median margins (quite definitely lateral to b lines) slightly indented
by 30/31, 31/32, 32/33. Genital tumescences, including a and b setae,
present on ix—xi, xxxli and xxxili (feebly developed).
Spermathecal pores, male porophores, setae, calciferous sacs and
glands, typhlosole, last hearts, as in caliginosa.
In the clitellum intersegmental furrows are obliterated and dorsal
pores are occluded but the epidermal thickening is slight. Sperma-
thecae (in x and xi) are iridescent and presumably contain spermatozoa
obtained from a copulatory partner. Male funnels are iridescent, also
indicating sexual maturity in spite of the feeble development of the
clitellum.
Remarks. A. molita is distinguished from all of the caliginosa-
complex: caliginosa, nocturna, iowana and arnoldi, by the four-segment
tubercula pubertates. Further distinction may be provided by an
apparent restriction of genital tumescences in clitellar region to xxxii-
xxxill. From all of the complex, except possibly iowana, molita is also
distinguished by the quadripartite (instead of double) origin of the
tubercula. In zowana, tubercula are band-like but nothing was said
as to origin.
Type locality. Arnold Arboretum, Boston, Mass.
Distribution. At present known only from the type locality.
REFERENCES
Evans, A. C.
1946. A new species of earthworm of the genus Allolobophora. Ann.
Mag. Nat. Hist. (11) 138: 98-101.
1948. On some earthworms from Iowa, including a description of a new
species. Ann. Mag. Nat. Hist. (11) 14: 514-516.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. DECEMBER 29, 1952 NuMBER 1(0)
ON THE EARTHWORMS
OF NEW HAMPSHIRE
By G. E. GATES
So far only one record of earthworms in New Hampshire has been
encountered in the literature. The worms were found “under the
bark of trees” at elevations of 2,000-3,000 feet in the White Moun-
tains by a German delegate to the International Zoological Congress
at Boston in 1907. The specimens went to an Irish Museum where
they were identified as Dendrobaena rubida (Savigny) 1826, by South-
ern (1910). The species had not previously been reported from North
America and since 1910 there has been no further record from this
continent. The record was given in a short article on another kind of
worm and, though the article was published in Philadelphia, the
record was not included in Smith’s review (1917) of the Lumbricidae
of North America.
Through the kindness of Dr. P. J. Darlington, Jr. who donated the
bait remaining from a fishing trip, it is now possible to list a second
species for New Hampshire.
Southern (1910, p. 18) quoted his collector who stated that “earth-
worms were very rarely met with in the forests at elevations of 2,000-
3,000 feet.’ Similar statements have been made in the past with ref-
erence to forests of northern New England as well as of northern New
York. Accordingly, Dr. Darlington’s observations appear to be
worthy of record:
‘““My experience has been that it is always possible to find earth-
worms in fair numbers even in the heavy woods in the more remote
parts of New England, or at least in the White Mts. and the Connecti-
cut lakes districts of New Hampshire. I have looked for them many
times when I wanted them for bait for trout and I have always been
able to find them under cover along the banks of brooks. I am fairly
2 BREVIORA No. 10
sure that there are at least two species. One,... (see below) .. . al-
though it occurs on the stream banks, is usually well above the water
level under stones or logs or in the roots of vegetation in sandy places
that are damp but not wet. The other species is darker and much
smaller, in fact hardly large enough to use for bait and it seems to oc-
cur in much wetter places, under stones on gravel bars or sandy banks
very close to the water level... . Practically every brook in New
England is fished these days, and many of the fishermen bring worms
with them and throw out what may be left at the end of their fishing
trips, so that different species must have been introduced many
times, even in the deep woods.” (Darlington, 7n Jit.)
ALLOLOBOPHORA ARNOLDI Gates 1952
In steep bank of East Inlet Brook, about 2,000 ft. above sea level,
near spruce, in heavy but not virgin forest, above Second Connecticut
Lake, May 30, 1952. Four clitellate specimens. Dr. P. J. Darling-
ton, Jr. collector.
Length, 95-105 mm. Diameter, 5.5 mm. Segments, 91 (amputee),
106 (amputee), 156, 158. First dorsal pore, 712/13 (1), ??13/14 (2).
(@ litellum, $xxvil-xxxiv. Tubercula pubertates bipartite and.as in
A. caliginosa (Savigny) 1826. Genital tumescences, on ix-xi (4), XXX,
Xxxil and xxxiv (4), xxix (1);
Remarks. Alive, these worms appeared to be quité without pigment
and they looked in that respect much as Octolasiwm cyanenm it Savigny)
1826 usually does.
The epidermis of segments xxvi, anterior half of xxvii, and of XXXV,
is somewhat thickened and of a translucent appearance quite distinct
from that of normal epidermis as well as from the yellow or white
opacity of the clitellar epidermis, on three specimens. The demarca-
tion of opacity and translucence on xxvii is indistinct, one gradually
passing into the other, on the fourth worm.
On the last segment of the amputees, rudiments of setal follicles and
of nephropores are still recognizable. In one of those specimens the
new anal region is small and scarcely distinguishable from the ninety-
first segment but probably is demarcated by a quite fine, greyish trans-
lucent furrow (presumably a rudimentary interségmental furrow).
In the other amputee no special anal region is recognizably demarcated.
A. arnoldi was split off from A. caliginosa when it was found in
Boston collections that all specimens having genital tumescences on
1952 EARTHWORMS OF NEW HAMPSHIRE 3
XXX, XXxll and xxxiv (with none on xxxill) could also be distinguished
from caliginosa by greater clitellar length and by larger segment
number.
The presence of worms with arno/di characteristics in northern New
Hampshire shows that those characteristics by which the species was
recognized and defined are not merely those of a local variant in the
Boston area. The occurrence of arnold? in many other states, as well
as in Canada, may be anticipated.
REFERENCES
Gatss, G. E.
1952. New species of earthworms from the Arnold Arboretum, Boston.
Breviora, No. 9: 1-3.
Smitu, F.
1917. North American earthworms of the family Lumbricidae in the col-
lections of the United States National Museum. Proc. U. 8. Nat.
Mus. 52: 157-182.
SouTrHERN, R.
1910. A new species of enchytraeid worm from the White Mountains.
Proc. Acad. Nat. Sei. Philadelphia, 62: 18-20.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. MarcH 20, 1953 NuMBER 11
CHARACTERS AND SYNONYMIES AMONG
THE GENERA OF ANTS
Part I
By Wiuiiam L. Brown, JR.
Museum of Comparative Zoology
Harvard University
Work now under way is aimed at a revision of the basic classification
of the Formicidae down to generic and subgeneric levels. The present
classifications are considered both unnatural and impractical, con-
cealing as they do large numbers of generic and specific synonyms. I
propose to differentiate genera only on the basis of discontinuities of
a largely morphological kind, and it is hoped that by this method a
useful and accurate key to the genera will become possible. Syn-
onymies given here require discussion that is best set forth separately,
so as not to clutter more nearly final, synoptic classifications and keys
now being prepared.
Myrmecta Fabricius
Myrmecia Fabricius, 1804, Syst. Piez.: 423. Genotype: Myrmecia gulosa
Fabricius, by designation of Emery, 1911.
Myrmecia (Promyrmecia) Emery, 1911, Gen. Ins., Fasc. 118: 18-19. Sub-
genotype: Myrmecia aberrans Forel, by original designation. NEW
SYNONYMY.
Myrmecia Clark, 1952, Formic. Australia, Melbourne, 1: 21.
Promyrmecia Clark, 1952, [bid., p. 119.
The references of Clark should be consulted for further synonymy.
Clark has stoutly defended the separation of Myrmecia and Pro-
myrmecia as distinct genera, first on the basis of ability or inability to
jump of the species concerned, and later, when saltation proved to
have numerous exceptions in bothdirections, on morphological grounds.
2 BREVIORA no. 11
Years of study of the myrmeciines by this author have produced in
the 1952 reference (p. 20) a couplet summarizing the characters
supposed to separate Myrmecia and Promyrmecia. The three charac-
ters contrasted are: (1) fraction of length by which the antennal scape
surpasses the occipital border, (2) distinctness of metanotum, and
(3) size, as mirrored in total length. In the first character, it may be
observed from the descriptions of species of Myrmecia given by Clark
himself in the pages following the couplet, that no less than one-third
of the 59 forms recognized disagree with the couplet, and disagree in
the direction of Promyrmecia. In at least some cases, the scape pro-
portions are exactly as stated in the couplet for the upper limit of
Promyrmecia. The distinctness of the metanotum is subject to varying
interpretation, but from any reasonable point of view, the generality
of the indistinctness of the metanotum in Promyrmecia cannot be
defended. In fact, Clark’s figures (op. cit.) in rapid survey are the
best contradiction of his couplet that I can offer anyone without a
large representation of material at hand for direct study. The “‘size”
figures, even as cited in the couplet, are broadly overlapping and
useless for the purpose to which they are put. Study of a good repre-
sentative series of the species is convincing proof for me that size, as
based on any measurable dimension or proportion of the worker or
female so far utilized, is graded from the largest Myrmecia to the
smallest Promyrmecia without a break.
It is entirely possible that study of the male genitalia, when a
sufficient number of males becomes available, will demonstrate a set
of cleavages that along with other characters will serve as disconti-
nuities for the proper splitting of Myrmecia. Clark’s and other di-
visions are here rejected for lack of evidence. It should be pointed
out, in order to save possible future confusion, that Clark has syno-
nymized Halmamyrmecia Wheeler with Promyrmecia, although he
wrongly maintains Myrmecia nigrocincta Fr. Smith, its type, in the
genus Myrmecia as he has defined it.
RHYTIDOPONERA Mayr
Ectatomma (Rhytidoponera) Mayr, 1862, Verh. zool.-bot. Ges. Wien 12: 731.
Genotype: Ponera araneoides Le Guillou, by designation of Emery, 1911.
Rhytidoponera (Chalcoponera) Emery, 1897, Ann. Mus. Stor. Nat. Genova
38: 548. [Sub] genotype: Ponera metallica Fr. Smith, by designation of
Emery, 1911. NEW SYNONYMY.
Rhytidoponera and Chalcoponera treated as distinct genera, Wheeler, 1922,
1953 ANT CHARACTERS AND SYNONYMIES 3
Bull. Amer. Mus. Nat. Hist. 45: 643-644. Clark, 1936, Mem. Nat. Mus.,
Melbourne, 9: 14-15.
The large genus Rhytidoponera in the present broad sense includes
species of typical ectatommine characteristics, distinguished in having
the inferior pronotal margins just in front of the fore coxae armed on
each side with an acute tooth; the hind coxae are completely unarmed
above. Distribution is primarily Australian and Papuasian, with
outliers in the ‘“‘Wallacia”’ region and in the southern Philippines to
the west, and in New Caledonia in the east.
The separation of Rhytidoponera from Chalcoponera, either generic
or subgeneric, has been accepted from the time of Emery’s first di-
vision of the few species then known. Rapid accretion of species in
both groups has frequently led to puzzlement of authors trying to
place new species in one group or the other, and this puzzlement has
resulted in several published expressions of doubt accompanying
specific descriptions.
The separation has been based on (a) proportions of certain antennal
segments in worker and male, (b) development and pectination of the
spurs of the middle and hind tibiae in the worker, (c) development of
notaulices in the male, and (d) presence or absence of a normal winged
female caste (Wheeler, loc. cit. 1922). If one applies these Emery-
Wheeler key characters rigorously to a wide variety of species in
Rhytidoponera and Chalcoponera, discrepancies are not long in ap-
pearing.
The tibial spurs of the middle pair of legs are reduced and narrow
(rarely absent) in all species of both genera examined for this work,
and may be safely disregarded. The extremes of development of
breadth and pectination of the posterior spurs are largely in accord
with the conventional separation of the two groups, but exceptions are
glaring, and gradual transition from one type to the other is evident
among a restricted selection of eastern Australian species. In the
species tenuis Forel, which has Chalcoponera antennal funiculi and
which has been placed in Chalcoponera by all authors, the posterior
tibial spurs are minute and show the strongly reduced pectination
supposed to be characteristic of Rhytidoponera s. str. Conversely,
certain Rhytidoponera s. str. species, such as R. malandensis Forel, have
the posterior spurs large, broad, and strongly pectinate; in fact,
R. malandensis has the spurs considerably more broadly pectinate
than in any Chalcoponera species I have seen, including the genotype.
In the worker funiculi, the proportions of the ‘‘critical’’ segments
4 BREVIORA no. Il
are intergradient and fully ambiguous in a number of species, among
them R. reticulata Forel, C. lamellinodis Santschi (paratypes), C. dubia
Crawley, and all the species of the C. impressa Mayr complex. More
ambiguous examples or outright contradictory species could be cited,
but the above cases should serve to prove the point so far as the worker
is concerned.
Concerning the male characters, a case similar to that for the
workers can be drawn, but it will serve merely to focus on the New
Caledonian fauna, consisting of a handful of forms in which the
workers have been placed in Chalcoponera on the usual characters.
The males of these species are, unfortunately for the generic division,
endowed with the characters of Rhytidoponera s. str. Emery (1914, in
Sarasin and Roux: Nova Caledonia, Zool. 1: 397) covers this situation
adequately when he states: “The males of the New Caledonian
species of Chalcoponera (Rhytid. fulgens, numeensis, atropurpurea and
acupuncta) make an exception to the character that I have attributed
to the subgenus concerning the structure of the antennae (Genera
Insectorum, fase. 118, p. 39). For the present, I am incapable of
distinguishing the males of the two subgenera of Rhytidoponera.”
Apparently Wheeler overlooked this paragraph when he raised Chalco-
ponera to generic rank in 1922, citing in his key the same discredited
male characters. I have dissected the male genitalia of several species
representing Chalcoponera and Rhytidoponera s. sir., and the prepa-
rations fail to show differences except minor ones among the various
species, without regard to the old division. The volsellae are uniformly
much like those of Myrmica and the Dacetini in the subfamily Myrmi-
cinae, and the other parts are on the usual formicid pattern.
It seems fairly obvious that most describers have pretty consistently
ignored the formal characters in assigning new species to one group or
the other; rather, assignment seems to have been made chiefly on the
basis of size and habitus. On such a basis, I can make out not two,
but several, species-groups of fairly distinctive relationships, but
broadly intergradient one to the next. Of these, the impressa complex
seems to have a combination of characters approaching nearest the
hypothetical generalized Rhytidoponera. Disregarding certain aberrant
species and complexes, such as the turneri group and the New Cale-
donian stock, the development of the majority of species seems to
have followed two lines: one toward large species of the type of mayri
Emery, araneoides Le Guillou, and punctata Fr. Smith, and one toward
the type of metallica Fr. Smith and victoriae André. In the farflung
1953 ANT CHARACTERS AND SYNONYMIES 5
regions penetrated by Rhytidoponera s. lat. on the Australian continent,
these two broad adaptive types have met with relative success, while
the more generalized connecting types have been restricted to favorable
forested areas, largely in the mountains of eastern Australia. Thus,
while the Rhytidoponera species inhabiting the deserts, scrubs, heaths
and eucalypt woodlands of the great, arid, open regions of Australia
may seem to be amenable to assortment into two distinct groups, it
must be borne in mind that the more generalized forest-loving species,
while less familiar to the observer, form an effective bridge joining
these two groups.
Before leaving Rhytidoponera, a word is required on the females.
It has been assumed that Chalcoponera species always produce normal
winged females, and that Rhytidoponera s. str. species do not. In 1950,
Clark (in litt.) indicated to me that he had found females of some sort,
presumably differentiable from the workers, in the latter group. To
date, he has published nothing concerning this find, so that it is fitting
that the old assumption be continued until definite information appears
in print. My own field investigations in Australia (1950-51) indicate
that the R. impressa group (¢mpressa, chalybaea, splendida) normally
produces a regular yearly crop of males and females in most mature
nests during the first part of the dry season in each of the regions
inhabited. Both sexes are winged before the nuptial flight, and ferti-
lization and nest-founding are presumed to follow conditions more or
less normal among the Ponerinae, as isolated females have been found
dealated and inhabiting small cells in various situations in normal
habitats. Outside the season for production of winged forms, only
dealate females have been found in established nests in addition to
the workers and such brood as occurs. When winged forms are found
in a nest, these are always of both sexes so far as my observations go.
In species of the metallica group, a situation of a quite different sort
is apparent. Among the common and widespread members of this
group, such as metallica, tasmanicnsis, and their very close allies,
winged or obviously dealate females are very rare or unknown. The
victoriae group is similar in this respect, although I have found a
dealate female in a mature nest of the species common at Kuranda, in
North Queensland, and have also found a lone winged female of
victoriae floundering in the damp sand at the surf edge at Seaford, on
Port Phillip Bay, in Victoria. Of several thousand nests of victoriae,
metallica, tasmaniensis and very close allies of these species that I have
opened, not one (with the single noted exception above) has yielded
6 BREVIORA no. ll
a female of the normal type, either alate or dealate. Yet there are a
few isolated specimens of normal females in these groups in the Mu-
seum of Comparative Zoology collection, most of which seem to have
been taken singly on the wing or otherwise.
While it is conceivable that the rarity of females in the nests of such
species may reflect the methods of investigation, and not true absence
of this caste, another circumstance makes this seem unlikely. I refer
to the very widespread occurrence of winged males in nests of the same
species at all seasons of the year, a phenomenon that stands out among
my Australian observations even though I failed to take quantitative
notes that would have made it much more convincing. These males
are usually present in small numbers, and may be found in the upper
as well as the lower chambers of a given nest. They are very active,
and either run to hide quickly or else take to ready flight. A more
limited number of observations on the large deserticolous members of
Rhytidoponera s. str. indicates that males are to be found in their nests
on a somewhat similar basis, though observations through several
seasons will be needed to confirm this.
Among some other, more rare and distributionally restricted species,
R. croesus Emery appears to have normal females most or all of the
time, while R. aspera (Roger) is recorded as having this caste at least
some of the time. For most of the species of the “inter-subgeneric”’
and aberrant groups, workers only are known at present, but these
forms are so rare and so little-investigated that presence or absence of
winged females cannot be assumed on any reasonable grounds. Under
the circumstances as outlined, utilization of the presence or absence
of a true female caste as a taxonomic character would seem entirely
premature.
The situation in the female-less or female-rare species that have
been fairly extensively investigated is, however, of considerable interest
apart from taxonomic considerations. In two such species, I have seen
a male seeking to enter an alien nest, apparently of the same form.
In the spring, in southeastern Australia and the dry Northern Flinders
Ranges of South Australia, at least, males of these forms are often seen
hawking in rapid flight low over the ground in the manner of certain
mutillid and thynnine males during the sexual search. Observations
in 1951:
On the 12th of September, in a strip of open eucalypt woodland at
Burwood, near Melbourne, Victoria, I noticed such a male in flight.
on a cool, sunny afternoon. While following him, I suddenly saw him.
a
‘
1953 ANT CHARACTERS AND SYNONYMIES
alight and disappear without hesitation into a small hole in the ground.
This hole, when attacked with a trowel, proved to be the entrance to
a nest of R. tasmaniensis, a nest containing, so far as I could dig on
this occasion, workers and two males, the latter indistinguishable from
males taken in nests of this species on other occasions. One of these
males, found just below the entrance, must have been the male I saw
entering just before I started to dig.
At Wilpena Pound, a sort of natural oasis in the arid Flinders Ranges
of South Australia, the evening of November 25 brought an abundant
flight of a large brown Rhytidoponera species, entirely males, to our
camp pressure lanterns. Later that night, some rain fell and there
were electrical storms on the adjacent peaks. On the next day, in an
area of T'riodia grass within the Pound, I found a low, gravel-studded
mound with wide, slit-like entrance, of the type made by certain arid-
land species of Rhytidoponera and Camponotus. The nest was seen in
the path taken on the way out, and was not excavated until returning
toward camp, late in the afternoon when the sun had fallen very low.
While I was bending over to deal the hard clay mound a first blow, a
large male of Rhytidoponera came, flying in low over the ground from
a distance, and landed directly on the lip of the entrance slit, which it
immediately entered. This was surprising, as no sign of life had been
detected around the mound at either time of inspection (the species
concerned, probably R. mayri Emery, is, with its close relatives, a
primarily crepuscular and nocturnal forager). The nest was immedi-
ately attacked, and the male recovered quickly several inches down
along the main entrance passage. Further digging secured only a few
workers of R. mayri Emery.
While these two incidents, observed by chance, are scarcely to be
considered definite proof of an established behavior pattern, the obser-
vations made to date on various Rhytidoponera species suggest the
following hypothesis, now being tested by Haskins (in litt.).
Rhytidoponera (s. lat.) varies by species and species-groups in
presence, absence or rarity of a normal female caste, and varies corre-
spondingly in behavior of and toward the males. In the species in
which normal females are rare or absent, it may be assumed that
deposition of fertile eggs is wholly or largely taken over by workers
1 The species described by Clark as R. stridulator and R. dixoni, along with some other forms‘
are supposedly distinguished from R. mayri by details of petiolar structure and body sculpture.
Single nests of what I take to be mayri, however, show a wide range of variation in these same
characters, and Clark has not made the differences sufficiently clear to convince me that his
species are really distinct.
8 BREVIORA No. ll
or highly modified ergatoids. It is possible that apparatus for the
retention of sperm isstrongly impaired in such hypothetical individuals;
if so, then the need for constant refertilization would seem to call for
the constant presence of consort males in the nest. Meager obser-
vations seem to indicate that males sometimes, perhaps normally,
leave the parent nest and enter another nest of the same species,
where they may remain as guests for long periods. If observation and
controlled experiment corroborates the above hypothesis, a new and
extremely interesting kind of behavior will be added to the multiplicity
of patterns known among the ants, and the origin of certain similar
male habits among the Dorylinae may be easier to understand.
CENTROMYRMEX Mayr
Centromyrmex Mayr, 1866, Verh. Zool.-bot. Ges. Wien 16: 894. Genotype:
Centromyrmex bohemant Mayr, monobasic.
Typhioteras Karawajew, 1925, Konowia 4: 128. Genotype: Typhloteras hamu-
latum Karawajew, monobasic. NEW SYNONYMY.
Karawajew based his genus on a specimen with only a single, large
pectinate spur on the posterior tibiae, and thereby carried it out to
Emery’s Group III of the tribe Ponerini. This placement is only
another example of the great faith formerly held by many authors in
the constancy and taxonomic importance of the number and condition
of the spurs of the middle and hind tibiae. In fact, the character in
question is a very poor one upon which to base a classification, es-
pecially in the Ponerini. If one reviews the situation in various
Centromyrmex species, including Centromyrmexr hamulatus NEW
COMBINATION, it is at once apparent that the number and con-
dition of the spurs in question differ by species, and possibly even
within species. Thus, either the middle, or the posterior, or even both
pairs of tibiae may possess the extra lateral spur in a more or less
rudimentary condition, while the degree of development of the medial
spurs and their pectination is also variable from one species to the next.
In other characters, including the striking general habitus and the
probably general termite-eating proclivities, the species here included
in Centromyrmex form a very homogeneous and natural-seeming group.
To break up this combination on the basis of spur characters, one
must, as did Karawajew, go to extraordinary lengths in invoking
‘““convergence”’ as a possible explanation of the common resemblances;
to be thorough in applying the same logic, one would have to distribute
the few species involved among three or four genera. In any case, the
1953 ANT CHARACTERS AND SYNONYMIES 9
tibial apices of several of the species bear such a dense growth of heavy,
spine-like setae that the identification of a spur rudiment of similar
size becomes an academic exercise. In addition to the synonymy of
Typhloteras with Centromyrmex, it becomes necessary to point out
that the subtribe Centromyrmicini of Emery is a NEW SYNONYM
of tribe Ponerini if the spur characters will not hold.
PRISTOMYRMEX Mayr
Pristomyrmex Mayr, 1866, Verh. Zool.-bot. Ges. Wien 16: 903. Genotype:
Pristomyrmex pungens Mayr, monobasic.
Odontomyrmex André, 1905, Rev. Ent. Caen 24: 207.
Hylidris Weber, 1941, Ann. Ent. Soc. Amer. 34: 184, 190. Genotype: Hylidris
myerst Weber, monobasic. NEW SYNONYMY.
Hylidris defended, Weber, 1952, Amer. Mus. Novit. 1584: 15-22.
When, in 1941, Weber first described Hylidris, he did so without
realizing that it might be closely related to another genus like Pristo-
myrmex. By 1952, he has realized this relationship and is at some
pains to mark out its boundaries. His new approach to the situation
lies in segregating the African species of Pristomyrmex from the Indo-
Australian ones; the former group, rallied about H. myers? as genotype,
is given the name Hylidris. He is vague about what genus the Indo-
Australian species are to belong to, but these are presumably to remain
in Pristomyrmex. The characters of Hylidris are cited briefly, but are
not contrasted with those of Pristomyrmea s. str. in any direct state-
ment. Instead, Weber gives briefly his views on generic limits, which
he feels should be arbitrarily drawn in some cases. (For my contrasting
opinion, see the introduction to this paper.)
Summing up, it may be said that Weber’s division of Pristomyrmex
is based upon characterization of only one of the resultant groups, and
no assurance is given that the characters are exclusive to that group.
In point of fact, these characters are not exclusive to the African group.
Since Weber claims to have examined the Indo-Australian species, at
least in part, it may be wondered that he did not note this fact for
himself; perhaps he did note it, and offered the generic-limit discussion
as a defense against protests he felt would be lodged against Hylidris.
I cannot follow Weber’s generic split, which I regard as wholly
arbitrary. Significantly, Weber fails to mention the old synonym
Odontomyrmex, a name put forward by André for a species that appears
to have the chief definitive characters of Hylidris. Mann (1919, Bull.
Mus. Comp. Zool. 63: 341) delivered the coup de grace to Odonto-
10 BREVIORA no. ll
myrmex when he noted two specimens of his Pristomyrmex obesus
melanoticus having a prominent tooth on one side of the pronotum,
but none on the other side. We do not need to appeal to abnormal
specimens in this case, however, as the known, normal specimens of
the Indo-Australian and African regions form a tightly intergradient
series with respect to the development of the posterior propodeal and
anterior alitruncal pairs of teeth and the smooth to foveate-reticulate
sculpture. This series, in my opinion, does not even split into re-
spectable species-groups on the basis of the known characters, let alone
genera or subgenera. Series of several undescribed species in the
Museum of Comparative Zoology and J. W. Chapman collections
serve only to fortify this opinion.
GAUROMYRMEX Menozzi
Gauromyrmex Menozzi, 1933, Natuurhist. Maandblad 22: 146. Genotype:
Gauromyrmex bengkalist Menozzi, monobasic.
Solenomyrma Karawajew, 1935, Treubia 15: 103. Genotype: Solenomyrma
acanthina Karawajew, monobasic. NEW SYNONYMY.
Acalama M. R. Smith, 1948, Jour. N. Y. Ent. Soc. 56: 205-207. Genotype:
Acalama donisthorpet M. R. Smith, monobasic. NEW SYNONYMY.
This genus is very doubtfully distinct from Vollenhovia Mayr, from
which it may at present be distinguished by means of the 11-segmented
antennae and the bidentate propodeum, vs. 12-segmented antennae
and unarmed propodeum in Vollenhovia. The characters are very weak
ones, and may be compromised in species in this complex that are
presently not available to me. Vollenhovia emeryi Wheeler, with
12-segmented antennae, has minute propodeal teeth, and the median
funicular segments are so reduced in length as to be virtually obsolete;
the step to Gauromyrmex from this species is a very short one. The
amber species Vollenhovia beyrichi (Mayr), if properly placed by
Wheeler, would be intermediate in the critical characters.
Types of Gauromyrmex bengkalisi and Acalama donisthorpet were
examined, and are considered specifically distinct, but not generically
so. Specific synonymy is formalized as follows:
GAUROMYRMEX ACANTHINUS (Karawajew) new combination
Solenomyrma acanthina Karawajew, 1935, Treubia 15: 103-104, fig. 23,
worker.
Acalama donisthorpei M. R. Smith, 1948, Journ. N. Y. Ent. Soc. 56: 207-208,
figs. 1, 2, worker. NEW SYNONYMY.
Types of Smith’s species and Gauromyrmex bengkalist have been
examined through the kindness of Dr. Smith. The former species has
1953 ANT CHARACTERS AND SYNONYMIES 1
been compared with Karawajew’s description and figure, and also
with the West Chinese series taken by myself, and mentioned by
Smith (loc. cit., p. 206). The comparison gives the distinct impression
that one is dealing with a single variable species. Variation is chiefly
in size (slight), distinctness and acuteness of propodeal teeth, and
depth of pigmentation, but the present evidence does not warrant
subspecific distinction in my opinion. This ant appears to be rather
common in India and China, and I believe that I have seen it on
several occasions in Bengal Province nesting under loose bark, though
the actual specimens were lost in a wartime shipment. In life, the
distinctive point of recognition lies in the extremely depressed ap-
pearance of the ant, even when moving about. It appears to the
naked eye somewhat like a flattened Leptothorax, and its attachment
to tree trunks and plant cavities may be correlated with the habitus.
At the time of Dr. Smith’s investigation of this insect, he sent
specimens to me, and I failed to recognize it as any described species.
The published descriptions of Menozzi and Karawajew came to light
later. The entire case of these synonymous genera should prove my
frequently-held point that the classification of the Myrmicinae is long
overdue for complete revision.
Tribe AMBLYOPONINI
Onychomyrmicini Ashmead, 1905, Canad. Ent. 37: 382.
Examblyoponini Donisthorpe, 1949, Ann. Mag. Nat. Hist. (11) 15: 401.
Reneini Donisthorpe, 1947, Ann. Mag. Nat. Hist. (11) 14: 183. NEW
SYNONYMY.
PRIONOPELTA Mayr
Prionopelta Mayr, 1866, Sitzb. Akad. Wiss. Wien 53: 503. Genotype: Pri-
onopelia punctulata Mayr, monobasic.
Ponera Fr. Smith (partim), 1860, Journ. Proc. Linn. Soc. London, Zool. 4
(suppl.): 105, nec Latreille.
Examblyopone Donisthorpe, 1949, Ann. Mag. Nat. Hist. (11) 15: 401. Geno-
type: Examblyopone churchilli Donisthorpe, monobasic: vide infra.
Renea Donisthorpe, 1947, Ann. Mag. Nat. Hist. (11) 14: 183. Genotype:
Renea testacea Donisthorpe, monobasic. (Nec Renea Nevill, 1880, in
Mollusca.) NEW SYNONYMY. Vide infra.
Incredible as it may-seem, Donisthorpe based his two new genera
and two new tribes on two new species, both of which are synonymous
with Prionopelta majuscula Emery. The same author had already
proposed a new name for a synonym of the same species, so he has
2 BREVIORA No. ll
achieved the unequalled feat of proposing for the same species three
new specific, two new generic and two new tribal names. The syn-
onymy of Prionopelta majuscula follows:
PRIONOPELTA MAJUSCULA Emery
Ponera simillima Fr. Smith, 1860, Journ. Proc. Linn. Soc. London, Zool. 4
(suppl.): 105, “worker,” 2, nec Fr. Smith, 1860, op. cit., p. 104. NEW
SYNONYMY.
?Rhopalopone simillima, Emery, 1900, Term. Fiizetek 23: 311. 1911, Gen.
Ins. 118: 35.
Prionopelta majuscula Emery, 1897, Term. Fiizetek 20: 595-596, worker, 9.
Brown, 1951, Bull. Brooklyn Ent. Soc. 46: 102, Examblyopone churchilla
synonymized.
Prionopelta poultoni Donisthorpe, 1952, Ann. Mag. Nat. Hist. (10) 10: 462,
nom. pro Ponera simillima Fr. Smith, I. NEW SYNONYMY.
Examblyopone churchilli Donisthorpe, 1949, Ann. Mag. Nat. Hist. (11) 15:
401-402, 2.
Renea testacea Donisthorpe, 1947, Ann. Mag. Nat. Hist. (11) 14: 183-186,
fig., worker, o’; pp. 590-591, 9. NEW SYNONYMY.
In his original description of Ponera simillima I, Frederick Smith
describes a ‘“‘worker”’ and also gives characters purporting to be those
of a winged female. In his treatment of 1932, Donisthorpe mentions
as Smith’s type “1 dealated @” from “Dor.” [Dory, A. R. Wallace].
It is by now fairly well known that some of the Dory insect material,
at least among the coleopterous collections in the British Museum, is
suspected to be from other East Indian localities. Smith’s description
fits Prionopelta mayuscula better than it does either of the other two
Prionopelta species known from the Indo-Papuasian area (P. kraepelina
Forel and P. opaca Emery), so there is no reason to question this
particular record for a widespread New Guinea species. The as-
sumption must be made, of course, that Donisthorpe’s 1932 assignment
to Prionopelta was correct; there is every reason to believe that it
was so.
In 1951, I determined that a paratype female of Examblyopone
churchilli was synonymous with P. majuscula and returned the type
to Dr. E. S. Ross at the California Academy of Sciences. Dr. Ross
was the original collector (at Maffin Bay, Dutch New Guinea) of both
E. churchilli and Renea testacea, and he has recently sent me workers
and males from the type series of the latter species. The workers
fit Emery’s diagnosis of P. majuscula very neatly, except for the
usual small size difference resulting from Emery’s habitual under-
1953 ANT CHARACTERS AND SYNONYMIES 13
measurement. This being the case, I asked Dr. Ross to compare the
female specimens assigned by Donisthorpe to R. testacea with the
female holotype (defective) of FE. churchill. He has replied (in litt.)
that the correspondence of form, etc. is as good as can be expected,
allowing for the missing parts of the EL. churchilli type. The R. testacea
workers differ as expected from authentic workers of Prionopelta opaca
and P. kraepelini in the Museum of Comparative Zoology and match
the differences from P. opaca listed by Emery in the original de-
scription of P. majuscula.
P. majuscula is larger than the other two Indo-Papuasian species,
and its worker has the dorsum of the head and alitrunk very definitely
shining, with minute, spaced punctulation. The female is considerably
larger and darker than the worker, and has stronger punctulation; the
head is darker than the rest of the body. Both P. kraepelini and
P. opaca are not only smaller, but also there is only a slight difference
between the worker and female stature in these species; the dorsum
of the head is densely and more coarsely punctulate and opaque,
especially in opaca. The worker and female of kraepelini and the
worker of majuscula are pale to bright yellow in color, while the females
of majuscula and opaca and the worker of opaca are darker, ranging
from ferrugineous brown to blackish-brown. P.opaca and P. majuscula
appear to be widespread on New Guinea and neighboring islands,
while P. kraepelini is a more westerly, Indomalayan species that has
spread into the Pacific as a tramp and has reached the Philippines in
the north at Dumaguete, Negros Oriental: several series (J. W.
Chapman and D. Empeso).
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BREVIORA
Museum of Comparative Zoology
CaMBRIDGE, Mass. APRIL 23, 1953 NuMBER 12
A CAVE FAUNULE FROM WESTERN PUERTO
RICO WITH A DISCUSSION OF THE
GENUS ISOLOBODON
By Tuomas E. ReyNoLps
Zoology Department, Massachusetts College of Optometry
Karu F. KoopMAN
Biology Department, Queens College
ERNEST E. WILLIAMS
Harvard University
In spite of the diligent work of H. E. Anthony nearly 40 years ago,
the vertebrate paleontology of Puerto Rico must still be described as
incompletely known. Anthony’s labors have not been followed up in
any systematic fashion by other investigators, and much therefore
remains to be done.
The present paper places on record a collection from an area of the
island relatively neglected by Anthony — the extreme western portion
of Puerto Rico. The collection, though small and apparently repre-
senting a fauna of no high antiquity, permits new locality records for
certain forms and includes a very interesting specimen of the genus
Isolobodon which has compelled us to inquire into the variability of
that genus and into the propriety of taxonomic subdivisions of it.
The materials here recorded were obtained by the senior author
during the fall and winter of 1948-49 at the mouth of the Cueva Monte
Grande between Mayagiiez and Cabo Rojo (Distrito de Mayagiiez,
Pueblo de San German, Barrio de Monte Grande). The specimens
have been deposited in the Museum of Comparative Zoology.
There were several levels in the deposit, and part of the deposit was
kitchen midden, but no record was kept of the level at which specific
material was found.
2 BREVIORA No. 12
We cite below only the mammalian component of the faunule.
There are, however, also fish, bird, turtle, snake and lizard bones.
The lizard jaws have been identified by Rodolfo Ruibal and Max K.
Hecht as all belonging to Anolis cuvieri, the living giant anole of
Puerto Rico. No attempt was made to identify limb bones in any
group. In the case of the mammals we take this opportunity to cite
also localities not previously published but represented in the col-
lections of the Museum of Comparative Zoology (MCZ) or the
American Museum of Natural History (AMNH).
The Monte Grande Faunule
INSECTIVORA
NESOPHONTES EDITHAE
Two mandibular halves.
This large extinct insectivore of Puerto Rico has previously been
reported from Hacienda Jobo, Morovis and Utuado. The MCZ col-
lections show it also from Ciales, Manati and Rio Piedras.
CHIROPTERA
NOCcTILIO LEPORINUS MASTIVUS
One complete skull. Two mandibles.
Though recorded by Anthony (1926) only from Old Loiza, this form
was collected by Anthony and Goodwin at Vega Baja (AMNH speci-
men) during a 1926 West Indian expedition which has never been fully
reported in the literature. The present is therefore the third record for
this large fish-eating bat in Puerto Rico.
CHILONYCTERIS PARNELLII PORTORICENSIS
One skull.
The specimen so referred seems somewhat small for the species, but
it is much too large to be the small Puerto Rican species of the genus
(C. fuliginosa inflata). It is probably a young specimen.
This subspecies seems to be relatively rare in collections, only 11
specimens having been previously obtained, representing four locali-
ties: Cayey, Morovis, Pueblo Viejo and Trujillo Alto.
MoNOPHYLLUS PORTORICENSIS
One skull.
1953 A PUERTO RICAN CAVE FAUNULE 3
This species was recorded by Anthony (1926) from five localities:
Bayamon, Cayey, Morovis, Pueblo Viejo and Trujillo Alto.
BRACHYPHYLLA CAVERNARUM
Four more or less complete rostra. Six complete mandibles. Seven
half-mandibles.
Recorded by Anthony from six localities: Cayey, Comerio, Corozal,
Morovis, Pueblo Viejo and Trujillo Alto.
ARTRBEUS JAMAICENSIS JAMAICENSIS
One skull. Six mandibles.
This very common form was obtained or reported by Anthony from
14 localities.
EPTESICUS FUSCUS WETMOREI
One mandible.
Anthony (1926) reported this species from four localities (Maricao,
Morovis, Pueblo Viejo and San German). He had collected it also at
Trujillo Alto (AMNH specimen) during the 1916 expedition but
somehow overlooked this record in his publication.
RODENTIA
RaTTUS sp.
Two mandibles. One half-skull.
Rattus, as always in the Western Hemisphere, indicates that at least
part of the Monte Grande faunule was post-Columbian in age.
ELASMODONTOMYS OBLI QUUS
A fragment of an upper incisor and a third right upper molar appear
to represent this species.
This large extinct rodent has previously been reported only from
Ciales, Morovis and Utuado. The MCZ collections add it also from
Rio Piedras.
ISOLOBODON PORTORICENSIS
Nineteen skull fragments. Fifty-five half- or partial mandibles.
The bulk of the material from Cueva Monte Grande is of a medium
sized hypsodont rodent. The enamel folds of the molars are nowhere
completely separated to form laminae. The lateral surfaces of these
4 BREVIORA No. 12
teeth have the pattern of striations characteristic of the genera or
subgenera [solobodon and Aphaetreus. Even a cursory inspection of
the molar pattern, however, rules out A phaetreus (Miller 1929a, pl. 2).
The specimens then seem referable to [solobodon. The only species of
that genus which has previously been recognized on Puerto Rico is
Isolobodon portoricensis, and almost all of the specimens agree well
with this form in both size and pattern and may immediately be
placed as representatives of this species
Specimens of [solobodon portoricensis from Monte Grande afford a
new record of locality but less clearly than the other forms recorded
above, since Anthony has already reported this species from nearby
Cabo Rojo as well as from Ciales, Manati, Salina, San German and
Utuado. (The MCZ adds Saliche and Aguirri.)
Four of the Monte Grande mandibles, however, are not immediately
classifiable as I. portoricensis and require special attention. All are
much smaller in size than fully adult J. portoricensis. Two are obvi-
ously immature. One of these is edentulous; the other has completely
unworn crowns to the teeth, which are, therefore, not readily compared
with the worn adult molar pattern. However, by breaking the ventral
surface of the mandible, it has been possible to discover from under-
neath the pattern of the enamel ridges which will be eventually re-
vealed by growth. The enamel pattern so revealed is exactly that of
typical I. portoricensis.
The third small mandible is larger than the other two and although
considerably smaller than adult J. portoricensis shows no morphological
evidences of immaturity (last molar not fully erupted, unworn molar
pattern, absence of bony shelf behind the last molar). The fourth
mandible is similar to the third in size but is edentulous.
Subadult mandibles equivalent in size to these last two Monte
Grande specimens are known from many localities elsewhere in Puerto
Rico and in Mona and the Virgin Islands and are always indis-
tinguishable from the adults except in size. It is, therefore, clear that
small size is in itself no bar to considering the four questionable Monte
Grande mandibles as I. portoricensis, and we refer the two obviously
immature specimens to that species without further discussion. The
fourth mandible, lacking teeth, manifests no distinctive characters and
may tentatively be referred to the same species.
The third questionable specimen differs, however, in molar pattern
from all the numerous specimens of Jsolobodon with which it has been
1953 A PUERTO RICAN CAVE FAUNULE 5
compared. In any [solobodon three important enamel folds are evident,
a single main fold from the labial side and two counterfolds from the
lingual side. In the Monte Grande specimen in question the anterior
counterfold of the first molar shows a distinct constriction about
midway along its length and, correlated with this, there is a marked
broadening of the lingual lobe anterior to the counterfold. No similar
constriction is present on the second and third molars; the anteriormost
lingual fold may have been broadened on both these teeth, but fractures
in the critical regions make it impossible to confirm this. No con-
striction comparable to that on the first molar, nor any broadening of
the anterior lingual lobe at all similar to that seen on the first molar
and probable on the second and third molars of the Monte Grande
specimen, have been found in the more than 250 Isolobodon portori-
censis mandibles examined. This third, small, Monte Grande mandible
may, therefore, represent an exceedingly rare species, possibly reaching
a smaller adult size, but closely related to Isolobodon portoricensis.
On the other hand, it may represent a rare mutant condition in a
subadult individual of J. portoricensis. Of these two alternatives we
consider the second the more economical hypothesis, since we have
been unable to find any other consistent differences between the third,
small, Monte Grande mandible and the mass of compared J. porto-
ricensis. We feel that the counterfold constriction might have been
caused by a single mutant gene. It is interesting that this apparently
aberrant individual probably never reached full adulthood.
In the course of this investigation, and while small size as a possible
species character was still in question, the Monte Grande mandible
and other Jsolobodon material from Puerto Rico was carefully com-
pared with material identified as [solobodon levir from Hispaniola. It
is worthwhile to consider here the status of the latter species, but
before doing so a brief resume of its taxonomic history seems in order.
I. levir was first described by Miller (1922) as a distinct genus and
species, [thyodontia levir, from two molars found fossil at St. Michel.
He did this believing that these isolated teeth were lower molars.
Twelve mandibles from the same locality were identified as [solobodon
portoricensis without comment. In 1929, after studying much more
material from the same locality, he realized that the two molars of
“Tthyodontia’’ were actually upper molars of Jsolobodon. In an attempt
to save I. levir from complete synonymy, he noted that all St. Michel
specimens were smaller than J/solobodon from other Hispaniolan
6 BREVIORA No. 12
(kitchen midden) localities. The latter, in turn, were indistinguishable
from Puerto Rican J. portoricensis. Although he could find no morpho-
logical characters to distinguish them, he nevertheless separated the
St. Michel specimens as a distinct species on size alone. He pointed
out, however, that Hispaniolan I. portoricensis came from kitchen
middens, whereas J. /evir came from ow] pellet deposits. As he indi-
cated, smaller species and individuals are usually found more fre-
quently in owl deposits than in kitchen middens, but he believed that
the presence in St. Michel caves of relatively large mandibles of the
related genus or subgenus A phaetreus ruled out the suggestion that in
the two types of Hispaniolan Jsolobodon he was dealing with large- and
small-sized samples from the same population. In later papers (Miller
1929b, 1930) he records [. levir from several other Hispaniolan locali-
ties (San Gabriel, Monte Cristi, Constanza, Trujin), in Monte Cristi
actually in association with I. portoricensis.
After extensive comparisons of I. portoricensis from Puerto Rico
and surrounding islands (AMNH material) with J. levir from Hispa-
niola (MCZ material from Fort Liberté as well as material from the
United States National Museum, especially mandibles from St.
Michel, Monte Cristi and Anadel, but also skulls from these and other
localities), we agree with Miller that no morphological characters are
to be found to separate the two species. We fail, however, to see any
clearcut size difference, since the smaller species ‘“‘levir’’ appears to us
to grade into the larger species portoricensis. “‘I. levir’? mandibles
from Hispaniola can be matched by equally small mandibles from
Puerto Rico. We believe also that in some cases Miller introduced a
false dichotomy into his measurements by comparing the largest speci-
mens identified as J. levir with the largest I. portoricensis: it is unlikely
that a bimodal distribution would have resulted in the one case of
actual association of the two forms if measurements of all measurable
specimens had been used. For these reasons we consider [thyodontia
levir Miller a complete synonym of [solobodon portoricensis J. A. Allen.
Isolobodon thus emerges as a monotypic genus or subgenus, undiffer-
entiated from Hispaniola through Mona and Puerto Rico to the Virgin
Islands. As has already been pointed out (Miller, 1918) this wide
uniform distribution may well have been brought about by human
transport. If that be true just what was the original range of the
genus before the coming of man to the Antilles. This would seem
impossible to determine now.
1953 A PUERTO RICAN CAVE FAUNULE of
Acknowledgments. We are indebted to the Departments of Mammals
of the Museum of Comparative Zoology, the American Museum of
Natural History, and the United States National Museum for the
privilege of examining and comparing specimens.
REFERENCES
ALLEN, J. A.
1916. An extinct octodont from the island of Porto Rico, West Indies.
Ann. N. Y. Acad. Sci., vol. 27, pp. 17-22.
AnTuony, H. E.
1926. Mammals of Porto Rico, living and extinct. N. Y. Acad. Sci.,
Scientific Survey of Porto Rico and the Virgin Islands, vol. 9,
pp. 1-238.
MILER, G. 8.
1918. Mammals and reptiles collected by Theodoor de Booy in the
Virgin Islands. Proc. U. 8. Nat. Mus., vol. 54, pp. 507-508.
1922. Remains of mammals from caves in the Republic of Haiti.
Smithsonian Misc. Coll., vol. 74, no. 3, pp. 3-5.
1929a. A second collection of mammals from caves near St. Michel, Haiti.
Smithsonian Misc. Coll., vol. 81, no. 9, pp. 14-18.
1929b. Mammals eaten by Indians, Owls, and Spaniards in the coast
region of the Dominican Republic. Smithsonian Misc. Coll., vol.
82, no. 5, pp. 6-8.
1930. Three small collections of mammals from Hispaniola. Smithsonian
Misc. Coll., vol. 82, no. 15, pp. 4-8.
8 BREVIORA NO. 12
PLATE
Crown views of mandibles of Isolobodon. A. “‘Isolobodon levir’’. U.S.N.M.
No. 255874 from Monte Cristi, Dominican Republic. B. Isolobodon portori-
censis. A.M.N.H. ‘0.1’? Utuado, Puerto Rico. C. Aberrant Monte Grande
mandible. M.C.Z. Cueva Monte Grande, Puerto Rico. (These specimens, all
of about the same size, have been selected to show the extremes of variability
of molar pattern.) 5 x natural size.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. APRIL 23, 1953 NuMBER 13
FOSSILS AND THE DISTRIBUTION
OF CEE EMID Wiens
1. ““Hydraspis’’ leithii (Carter) in the
Eocene of India is a Pelomedusid
By Ernest WILLIAMS
For SO years a fossil turtle from the Eocene of India has been
referred to the Recent South American chelyid genus [/ydraspis (not
of Bell 1828, correctly called Phrynops according to Stejneger 1909,
and Lindholm, 1929). In spite of the zoogeographical interest of this
record the evidence for this assignment has not previously been re-
examined critically.
In view of the prevalent idea that the family reached Australia from
Asia, the occurrence of a chelvid in the Eocene of India would be
neither surprising nor unwelcome. Reference, however, of an Eocene
Indian fossil to a modern South American genus is more suspect and
obviously calls for re-investigation. Examination of the original de-
scription leads to a verdict unfavorable to both the generic and the
family assignment.
The fossil in question was brought to scientific attention just over
100 years ago. In 1852 H. J. Carter in a study of the geology of the is-
land of Bombay described as 7T’estudo leithii* the remains of a small
turtle from the Intertrappean beds. Carter was not deceived as to the
affinities of the form; he was using the generic name T'estudo in a
Linnaean sense and explicitly stated that he regarded his fossil as close
to Sternothaerus” (= Pelusios). He published two good plates giving
a reconstruction of the fossil made from nine partial specimens. These
Testudo leithii Carter 1852 preoccupies Testudo leithii Giinther 1869 for the tortoise of
Egypt. The available and correct name of the latter then appears to be Testudo kleinmanmni
Lortet 1883
2 BREVIORA NOwlS
plates and his text description are the basis of all subsequent discussion.
The type material cannot now be located in India and has never been
restudied.
On the basis of the 1852 description, Gray in 1871 reassigned the
Indian fossil. He remarked: ‘‘The description and figure of the cara-
pace induce me to believe that the fossil is nearly allied to some of
our existing South American species of the restricted genus Hydraspis;
and the remains of the head, which are unfortunately imperfect, lead
to the same conclusion....’’ This determination has been very
generally copied, in spite of the zoogeographical anomaly upon which
Gray himself commented.
I find it necessary to disagree with Gray on the basis of the figured
morphology of both shell and skull. Plates 1 and 2 are reproductions
of Carter’s plates X and XI with a few inessential modifications for
clarity.
According to Carter’s plate X (though the area is given in dotted
lines only) and according to his express statement in the text there is
no nuchal seute in the Indian fossil. Lydekker (1S89b, p. 170) stated:
“The omission of a nuchal shield in the restoration of the anterior
border of the carapace is probably incorrect.”” Perhaps, however,
Lydekker made this statement only on the ground that if the form
were []ydraspis it should possess a nuchal scute. Absence of a nuchal
scute would rule out all Recent genera of Chelvidae except Chelodina
and Emydura (in both of which the scute may be present or absent)
and Elseya (in which it is regularly absent). The latter genera are all
natives of the Australian region.
In Carter’s fossil the first vertebral is much smaller than the second
vertebral. This precise condition is not met with in the living C helyi-
dae. In most Recent South American forms including most cf the
species of Hydraspis (= Phrynops) the first vertebral] is on the contrary
much larger than the second. There is, it is true, an appreach to the
condition of the fossil in the Australian genera Ek mydura, Pscudemiydura
and Elseya and in the South American Hydraspis tubcrosa (specimers
in the British Museum) and perhaps in some specimers of the Scuth
American genus Hydromdusa, but in the latter only if the anterior
median scute is interpreted as a nuchal withdrawn from the margin
rather than as a transversely divided first vertebral. In none of these,
however, is the first vertebral as much smaller than the second as it
is in the fossil.
In the Indian fossil the gulars are small, and the very bread inter-
1953 FOSSILS AND DISTRIBUTION OF CHELYID TURTLES i)
gular extends from the anterior margin to the humero-pectora! sulcus,
separating the humerals in the midline. This pattern of scutes on the
anterior plastral lobe is different from any which is known in the
modern Chelyidae. In all the genera except Chelodina, Pscudemydura,
and rarely in Emydura (Siebenrock, 1907) the humerals meet in the
midline for a significant distance behind the intergular. The intergular
also is rarely as broad as in the fossil. In Chelodina the intergular is
very large and separates the humerals, indeed extending deep into the
area of the pectorals, and, except in C. intergularis Fry, it does not
reach the anterior plastral margin, the gulars meeting in front of it.
In Pseudemydura (Siebenrock, 1907) the intergular is like that of the
Indian fossil in its breadth but as in Chelodina dips deeply between
the pectorals. The gular-intergular pattern in the exceptional Emydura
subglobosa in which Siebenrock found the intergular separating the
humerals is also quite unlike that of the Indian fossil, the intergular
being narrow and of quite different shape.
The feeble xiphiplastral notch is another feature in which the fossil
differs from Hydraspis and other chelyids except the forms of the
Australian region and Batrachemys.
The shell, therefore, is not a good match for that of any known
genus of chelyid. It is perhaps most like those of the Australian genera
but differs from all of these in significant details, for example, in the
presence of neurals, which are lacking in all the Australian genera.
The skull and mandible in their turn provide conclusive evidence
against chelyid affinities. The mandible, although incomplete, is
stouter and broader than in any known chelyid. The symphysis must
have been long, in strong contrast to the condition in chelyids. The
skull, shown by Carter only in dorsal view, is radically different from
that of any chelyid. The skull roof has undergone emargination from
behind as in the Pelomedusidae or most Cryptodira. As a result, the
parieto-squamosal arch is absent, but a jugal-quadratojugal bar is
present. In the Chelyidae and in no other turtles the skull is emargi-
nated from the ventral margin only, and a parieto-squamosal con-
nection is (except in Chelodina) always preserved. In chelyids the
quadratojugal and the bar of which it was a part are always absent.
These are as crucial and clearcut differences as it is possible to obtain
between skulls of turtles. Carter’s fossil cannot be a chelyid.
It is most probable that it is a pelomedusid. Reference to this
family wou'd, it will be recalled, be a return to the opinion of the
original dese-iber, wh» thought the fossil was closest to the African
4 BREVIORA NOn Ie
pelomedusid genus Pelusios. The characters of the shell fit such a
reference extremely well. The nuchal scute is almost always absent in
pelomedusids. The first vertebral is almost always smaller than the
second (exception in Palaeaspis Gray, and sometimes in Pelusios).
An intergular separating the humerals is found in Stereogenys podocne-
motdes and in Elochelys perfecta as well as in occasional individuals of
Podocnemis expansa. The xiphiplastral notch is very variable in
pelomedusids. Vertebral shields 2 to 4 of the Indian fossil have
strikingly convex anterolateral borders, conspicuously concave postero-
lateral borders, as in some Recent Podocnemis and a number of fossil
pelomedusids.
A final feature which, as described and figured by Carter, is anoma-
lous, may be clinching proof of the pelomedusid affinity of this form.
Gray mentioned that Carter’s form was “peculiar also for the under-
side of the marginal opposite the (pectoral-humeral) suture being
rather broader than the rest and angular on the inner edge, which I
have not seen in any of the Recent species.” If we have to do here
with sulci between scutes, the situation is indeed peculiar and unique,
but it is noteworthy that the lateral marginal scute boundaries are
represented by dotted lines in Carter’s original reconstruction (solid
lines in plate 2 here) except for the anomalous ‘‘marginal”’ in question.
It is noteworthy also that the ventral view of the lateral marginals in
Carter’s plates does not match the dorsal view of the same marginals.
It is evident that Carter was not certain of the exact scute boundaries
here, and it is possible that he has figured as the anomalous “‘mar-
ginal” the sutures between bones rather than the sulci between
scutes. The lines drawn solidly by Carter in this region are in nearly
the right position and have the right aspect to represent in their
lateral portions the sutures bounding small mesoplastra and medially
the hyo-hypoplastral suture. This interpretation is the more probable
because Carter’s figure is a reconstruction from nine specimens, one
of which may have shown the sutures in this critical region and not
the sulci. If small laterally placed mesoplastra were present, this fact
would definitely place the Indian form in the Pelomedusinae of Zan-
gerl (1948) with which, on the basis of other resemblances in its shell,
it is most plausibly linked.
Carter’s turtle is, therefore, most probably a new genus of pelome-
dusine. I cannot distinguish it from all previously proposed genera
of pelomedusines because not all of these are themselves well-delimited.
Thus I cannot distinguish it from Rosasia (Carrington da Costa 1940)
1953 FOSSILS AND DISTRIBUTION OF CHELYID TURTLES 5
because at present that genus does not seem to be definable. (On its
known characters Rosasia might be a synonym of any one of several
genera. The carapacial shield, which alone is known, is not sufficiently
diagnostic.) I cannot distinguish the Indian fossil from Dacochelys
(Lydekker, 1889a) because there are no comparable parts, that genus
having been founded on a mandibular symphysis, a part that is missing
in Carter’s fossil. There is also no evidence that Dacochelys is a pelome-
dusid.* I distinguish Carter’s form, with some hesitation, from
Elochelys Nopcsa 1931 because the gular-intergular pattern is not
quite that of E. perfecta, the type of the genus, and I am not persuaded
that the other species referred by Nopcsa to that genus (/. major)
belongs there. The critical feature of Elochelys also, the absence of a
suprapygal, is not determinable in Carter’s form. A possibly trivial
feature, the feeble xiphiplastral notch, distinguishes the Indian fossil
from Stereogeni's podocnemoides (Reinach, 1903), but with Schmidt
(1940) I do not believe that podoenemoides belongs to the genus
Stereogenys. Podocnemoides and leithii may indeed belong to the same
genus, but that genus is then unnamed.
From the better defined genera of the Pelomedusidae the Indian
form is distinguished by the following combination of characters:
(CARTEREMYS, new genus
Type. Testudo leithii Carter 1852.
Diagnosis. Skull roof much emarginate from behind; opisthotics
prolonged backwards in sharp crests; mandible with a moderately long
symphysis; nuchal absent; first vertebral not divided transversely ;
intergular large and very broad, separating humerals; xiphiplastral
notch narrow and shallow; pubic and ischial sears distinctive in shape
and position.
* An ingenious device by which Lydekker avoided the possibility of a change in the trivial
name of Dacochelys has been the source of confusion in regard to this point. The type of the
genus is Dacochelys delabechei Lydekker 1889, but Lydekker considered his form probably
synonymous (largely on size alone) with Emys conybeariti Owen and therefore (according to
Lydekker and Boulenger 1887) with Emys delabechei Bell. No name change is necessary 7f this
synonymy is correct and, since the shell of Emys conybearii shows small lateral mesoplastra
(Lydekker and Boulenger 1887), Dacochelys delabechei is then a pelomedusine. But the only
valid physical type of Dacochelys (the type of the species upon which the genus is based) is
the very peculiar mandibular symphysis, which I regard as quite impossible to assign to family.
Lydekker’s device has most unfortunately brought the name Dacochelys into the literature (for
example in Zangerl, 1948) as a pelomedusine — which it may be but which it certainly cannot
at present be proved to be.
It should be mentioned that if Dacochelys delabechei were in fact a synonym of EF. delabechei
Bell as Lydekker assumed, and if E. conybearii Owen were a synonym of E. delabechei Bell as
Lydekker also assumed, Dacochelys Lydekker 1889 would be a straight synonym of Palaeaspis
Gray 1870, type Emys conybearzi Owen
6 BREVIORA NO. 13
Horizon. Intertrappean Eocene of Bombay.
Comment may now be made on other records and alleged records of
this form.
In 1890 Lydekker reported an entoplastron showing half of an
intergular scute from the Intertrappean beds in the Nagpur district
as a second occurrence of ‘“Hydraspis leithii.”’ The new fragment was
much larger than typical Carteremys leithii and differed also in the
much narrower intergular. Lydekker passed over the differences
casually with a mention of variability in Hydraspis (= Phrynops)
hilar. It is unlikely that he had in hand any member of the genus
Hydraspis or Carteremys leith. It is probable that he had some other
pelomedusid, and he had himself previously (1887) described a Podoc-
nemis indica in the Eocene of India.
Sukheswala (1947, an abstract only) has reported a find of a shell
of Testudo (= Carteremys) leithii in the Intertrappean of Worli Fill,
Bombay. Here, as in the case of the specimens described by Carter,
remains of frogs (Indobatrachus pusillus) were associated. In this
instance there is no doubt of the identification.
Dr. Sukheswala has kindly sent me a photograph of this specimen
(plate 3). The outline of the shell (somewhat different from that
figured by Carter) is clearly shown, as is also the characteristic
sculpture of the surface (mentioned by Carter), which while somewhat
like that of some chelvids also resembles that of, for example, the
American pelomedusine genus Taphrosphys. More important is the
evident presence of several neurals, the first vertebral clearly much
smaller than the second, and (less certainly) the absence of a nuchal
shield. The plastron and the skull are unfortunately missing. As with
Carter’s specimens the new shell is small, eight inches long by six wide.
The specimen is now in the possession of the Geological Survey of
India.
Two other specimens have since been found by Dr. Sukheswala and
have been sent by him to the Geological Survey of India. These are
recorded in the general report of the Survey for 1948 (West, 1950).
These specimens, while recognizable, afford no additional information.
Also recorded by the Survey in the same report was a possible young
shell of Carteremys leithii from the carbonaceous shales of an Inter-
trappean band near Raibasa in the Chhindwara district, Central
Provinces, India. The Survey has generously sent me a photograph
of this specimen. The identification must be pronounced doubtful as,
indeed, the Survey has regarded it. While this new locality may be
“J
1953 FOSSILS AND DISTRIBUTION OF CHELYID TURTLES
thought of hopefully, it would appear that Carteremys leith is at
present known with certainty only from the Intertrappean of the
island of Bombay.
Acknowledgments. Dr. A. S. Romer, Dr. P. J. Darlington, Dr. P. E.
Vanzolini, Dr. Karl F. Koopman, Dr. W. E. Swinton and Mr. L. I.
Price have read the manuscript and given critical advice. The allo-
cation of the Carter fossil has been discussed with Dr. Rainer Zangerl.
Mr. Arthur Loveridge, Mr. C. M. Bogert and Dr. H. W. Parker have
permitted the examination of Recent skeletal material in their de-
partments. I wish to thank also the authorities of the Bombay
Natural History Society for endeavoring to locate Carter’s types, the
Geological Survey of India for furnishing me all the information in
their possession in regard to the new specimens of Carteremys and for
photographs of certain of them, and especially Dr. R. N. Sukheswala
of St. Xavier’s College, Bombay, for his prompt and courteous replies
to iny insistent questions and for the gift of the photograph reproduced
as plate 3.
REFERENCES
CARRINGTON DA Costa, J.
1940. Um novo quelonio fossil. Portugal Com. Serv. Geol., vol. 21,
pp. 107-123.
Carter, H. J.
1852. Geology of the island of Bombay. Jour. Bombay Branch Roy.
Asiatic Soc., vol. 21, pp. 161-215.
Gray, J. E.
1870. Supplement to the Catalogue of Shield Reptiles in the Collection
of the British Museum. Part I. Testudinata. London. 120 pp.
1871. Notice of a fossil hydraspide (Testudo leithii Carter) from Bombay.
Ann. Mag. Nat. Hist., ser. 4, vol. 8, pp. 339-340.
LINDHOLM, W. A.
1929. Revidiertes Verzeichnis der Gattungen der rezenten Schildkréten
nebst Notizen zur Nomenklatur der Arten. Zool. Anz., vol. 81,
pp. 275-295.
Lortet, L.
1883. Poissons et reptiles du lac de Tiberiade et de quelques autres
parties de la Syrie. Arch. Mus. Hist. Nat. Lyon, vol. 3, pp.
99-194.
LYDEKKER, R.
1887. Eocene chelonians from the Salt Range. Paleontographica Indica,
ser. 10, vol. 4, pp. 59-65.
8 BREVIORA No. 13
1889a. On the remains of Eocene and Mesozoic Chelonia and a tooth of
2(Ornithopsis). Quart. Jour. Geol. Soe. London, vol. 45, pp.
227-246.
1889b. Catalogue of the fossil Reptilia and Amphibia in the British
Museum (Natural History). Part III. London. 239 pp.
1890. Notes on certain vertebrate remains from the Nagpur district.
II. Part of a chelonian plastron from Phisdura. Rec. Geol. Surv.
India, vol. 23, pp. 22-23.
LypEKKEmR, R. and G. A. BOULENGER
1887. Notes on Chelonia from the Purbeck, Wealden and London Clay.
Geol. Mag., ser. 3, vol. 4, pp. 270-275.
Nopcsa, F.
1931. Sur des nouveaux restes de tortues du Danien du Midi de la
France. Bull. Soc. Geol. France, ser. 5, vol. 1, pp. 223-234.
1934. The influence of geological and climatological factors on the distri-
bution of non-marine fossil reptiles and Stegocephalia. Quart.
Jour. Geol. Soc. London, vol. 90, pp. 76-140.
REINACH, A. VON
1903. Schildkrétenreste aus dem aegyptischen Tertiir. Abhandl.
Senckenberg. naturf. Ges., vol. 29, pp. 1-64.
Scumipt, K. P.
1940. A new turtle of the genus Podocnemis from the Cretaceous of
Arkansas. Geol. Ser. Field Mus. Nat. Hist., vol. 8, pp. 1-12.
SIEBENROCK, F.
1907. Beschreibung und Abbildung von Pseudemydura wmbrina Siebenr.
und iiber ihre systematische Stellung in der Familie Chelydidae.
Sitz-Ber. Akad. Wiss. Wien. Math-naturw. KI. vol. 116, pp.
1205-1211.
STEJNEGER, L.
1909. Generic names of some chelyid turtles. Proc. Biol. Soc. Wash-
ington, vol. 22, pp. 125-127.
SuKHESWALA, R. N.
1947. A fossil tortoise (Testudo leithii) from the Intertrappean of the
Worli Hill, Bombay. (Abstract.) Proc. Indian Sci. Congress,
33rd Session, vol. 3, p. 97.
West; W. D:
1950. General report of the Geological Survey of India for the year 1948.
Rec. Geol. Surv. India, vol. 82, pp. 1-253.
ZANGERL, R.
1948. The vertebrate fauna of the Selma Formation of Alabama. Part
II. The pleurodiran turtles. Fieldiana, Geology Memoirs, vol, 3,
pp. 23-56.
Plate 1. Carteremys leithii, dorsal view of shell and skull, slightly modified
for clarity from the original reconstruction (Plate X of Carter, 1852). Skull
restored from the more perfect half.
Plate 2. Carteremys leithii, ventral view of shell and mandible, slightly
modified for clarity from the original reconstruction (Plate XI of Carter, 1852).
Plate 3. Carteremys leithii, photograph of dorsal surface of shell. (Courtesy
of Dr. R. N. Sukheswala.)
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. May 28, 1953 NUMBER | 4
RECORD OR A
HERMAPHRODITIC HORSESHOE CRAB,
ON CTEL UGS VOY EY Gels NUE ES Ge
By JoHn P. Baptist
lishery Research Biologist
U.S. Fish & Wildlife Service
Newburyport, Mass.
A hermaphroditic horseshoe crab, Limulus polyphemus L., was
picked up by a small scallop dredge from the channel of Plum Island
Sound, Massachusetts on May 16, 1952. As far as can be determined,
this is the first such Limu/us to be recorded.
The general appearance was that of a mature male. Its width
(125 mm.) was about average for males in this area. (The widths of
606 mature males measured this summer ranged from SS mm. to
140 mm., with an average of 117 mm., while 489 mature females
ranged from 130 mm. to 199 mm. in width, with an average of 155 mm. )
It had mating claspers, and the anterior edge of the prosoma was
curved upward in the middle, which also is a male characteristic. The
left genital aperture was typically male, round and located on a definite
papilla. The right genital aperture was typically female, a horizontal
slit and not on a papilla (Lochhead, 1950). The genital operculum,
showing both male and female genital apertures, may be seen in the
upper illustration of Plate 1. On the dorsal surface of the opisthosoma
near the telson were darkened areas normally present on mated
females. These may be barely discernible in Plate 2. They are pro-
duced by the abrasive action of the anterior edge of the male as he
clings tenaciously to the female during the mating period. Females
which have mated for prolonged periods exhibit deep scars on the last
pair of immobile spines, caused by the powerful claspers of the male.
BREVIORA no. 14
bo
Such scars were not evident on the hermaphrodite, suggesting that it
had been mated as a female for a comparatively short time.
Gross dissection of the prosoma and histological sections revealed
both male and female gonads. The right and anterior portions were
full of eggs in various stages of development (Plate 2), comparable to
to those of mature females. Histological sections of the tissue on the
left side demonstrated sperm sacs full of tail-less sperm, diverticula of
the hepatopancreas and connective tissue (Plate 1, lower illustration).
The same structures were visible in sections of gonads from norma!
males. According to Benham (1885) the tails are apparently produced
as the sperm approach the aperture.
The presence of both eggs and sperm confirms the external evidence
that this specimen is a true hermaphrodite.
REFERENCES
BENHAM, W. B.S.
1885. On the testis of Limulus. Trans. Linn. Soc. Lond. (Zool.), (2)
2: 363-366.
LocHHEAD, J. H.
1950. NXiphosura polyphemus. In Selected Invertebrate Types, edited by
F. A. Brown, Jr. pp. 360-381. John Wiley & Sons, Inc., N. Y.
1953 A HERMAPHRODITIC HORSESHOE CRAB 3
Upper — Ventral view of posterior surface of genital operculum (enlarged),
showing right genital aperture (Q ) and left genital aperture (<7).
Lower — Photomicrograph of section through testis (X 120). S, sperm sacs
containing tail-less sperm. H, hepatopancreas. C, connective tissue.
(Photographs by Alden P. Stickney)
BREVIORA No. 14
PAG 2
Dorsal view of Limulus polyphemus with carapace of the prosoma removed,
showing mass of eggs on the right side, but only a few scattered eggs on the
left side. (Telson has been cut off). (Photograph by Alden P. Stickney.)
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. Niay 1, 1953 NuMBER 15
FURTHER NOTES ON THE EARTHWORMS
OF THE ARNOLD ARBORETUM, BOSTON
By G. E. Gatss
An opportunity to search once more for mature specimens of the
Pheretima sp., represented in collections of the previous spring by a
single, specifically unidentifiable juvenile, was unexpectedly provided
on September 25, 1952. After securing the desired material, as many
of the previous sites as time permitted were re-examined, to permit
comparison of fall with spring populations.
In the natural woods, during the summer, the leaf pile of previous
collections had been grown over and had become rather dry. Some
fifty feet away, and at about the same distance from the site where
the juvenile of Pheretima sp. had been found, another large leaf pile,
possibly older and with more gravel, had remained damp and without
any plant overgrowth. In the peat bog, where vegetation had been
mowed and removed, water was no longer running in the brook though
still standing in small, scattered puddles, but previously disturbed soil
still seemed moist. In the ponds, water level had fallen several inches
exposing a marginal strip of bottom muck that was still saturated.
The Newton garden had been planted and could not conveniently be
re-examined.
The species obtained in the September collections are listed in the
~ following table which also shows sexual stages and number of specimens
secured.
2 BREVIORA no. 15
EARTHWORMS OBTAINED IN THE ARBORETUM IN SEPTEMBER, 1952
Natural Woods
Species Original Second Leaf Pile Peat Bog
Leaf Pile Near Side Far Side
Allolobophora T =
arnoldi 0-2-2 0-1-0 0-0-1 0-0-1
caliginosa 0-0-2 0-0-1
limicola 0-2-0
sp. (1) 4-0-0-0
Dendrobaena
subrubicunda 3-5-32 1-0-1
Eisenia
foetida 12-5-6 7-1-7 2-0-1
Lumbricus
castaneus 0-0-2
rubellus 3-1-7 2-2-3-16 6-11-17
terrestris 4-1-0-4 1-0-1
Octolasium
lacteum 0-0-1
Pheretima
agrestis 0-0-35 0-0-21
Papas indicate juvenile-aclitellate-clitellate or small juvenile-late juvenile-aclitellate-
clitellate.
+ Postsexual aclitellate.
* Specimen in poor condition.
(1) Probably of arnoldi and /or caliginosa.
A. arnoldi. Two specimens from original leaf pile listed as clitellate had
spermatophores externally but the clitellum had almost completely re-
gressed, the site being indicated only by a brownish discoloration. In an
amputee from the bog, after enteroparietal healing at 106/107, cvi had
been much elongated and at the same time narrowed so as to taper
terminally and its setae had been lost though follicle apertures were still
visible. One worm from the leaf pile showed the same changes in its last
segment.
L. terrestris. A small juvenile from the leaf pile, 27 x 2.5 mm. has 149 segments.
O. lacteum. Segments in front of the anus were packed with coelomic corpuscles
and parasitic bodies many of which were discharged through dorsal pores
during preservation.
1953 EARTHWORMS OF THE ARNOLD ARBORETUM 3
DENDROBAENA SUBRUBICUNDA (Eisen) 1874
Clitelluin, on xxvi-xxxi but occasionally with slight extension onto
xxxli (32 + 26), xxvi-xxxii (1). Epidermis of xxv (several + 8) and
xxxil (several) slightly modified. Tubercula pubertates, on xxvili-xxx
(85 + 27), occasionally reaching onto xxxi. Each tuberculum nearly
always is bounded laterally by a deep groove. In two clitellate speci-
mens in which tubercula were quite indistinguishable the groove was
obvious, on both sides of the body. The tuberculum is longitudinally
elliptical in outline. A central depressed portion of similar outline
may be more or less clearly demarcated from a band-like marginal
area by a slight groove, or, instead, there may be recognizable only a
single longitudinal groove at the center. The lateral groove, in some
specimens, apparently passes, though less clearly, around ends of
tuberculum to become completely circumferential. Genital tume-
scences, including ab, on viii (5), ix (14+ 11), x (5), xvi (83 + 27),
xvii (2+ 1), xviii (1), xx (8), xxi (1), xxii (+1), xxiii (+2), xxiv
(12 + 12), xxv (1+ 3), xxvi (+1), xxvii (26 + 25), xxviii (2), xxxi
(19 + 23), xxxii (1), xxxiii (+1), xxxiv (+1), xxxv (+1). Tumescences
on xvi may extend well into be and aa or even be united mesially.
Those on xxxil to xxxv are especially well developed.
Homoeosis. Male pores, on xvi. Female pores, on xiv. Clitellum,
on XXvi-xxxil and on xxxii-xxxiv of left side only. Tubercula pu-
bertates, on xxvili-xxx (right side), xxix-xxxili (left side). Genital
tumescences, on left sides of xxiv and xxxiv. No metameric abnor-
mality recognized.
Remarks. In live worms a short terminal region had a brilliant
yellow appearance that disappeared during preservation and which,
presumably, was due to accumulations of coelomocytes in the coelomic
cavities of a region with rather transparent parietes. The anal region
was not marked off by an intersegmental furrow from the last seti-
gerous segment in 19 specimens. Spermatophores were present ex-
ternally on eight specimens.
Data from a series of 27 specimens (collected at Bangor, Maine)
that has been available for comparison with arboretum worms, are
included in parentheses above, preceded by the + sign. Very little of
such information has been reported.
4 BREVIORA No. 15
LUMBRICUS RUBELLUS Hoffmeister 1845
Number of segments: 67, 73, 75, 79, 85, 88, 91, 96, 100, 101. 109 (2),
110, 111, 112 (4), 113 (3), 114 (8), 115 (2), 116 (8), 117 (4), 118 (3),
119 (4). Clitellum, on xxvii-xxxii (40). Tubercula pubertates, on
XXVili-xxxi (5), but extending onto xxvii (40). Genital tumescences,
including ab, on vi? (1), vil? (1), vili (2), x (6), xi (42), xii (50), xxvi
(56), xxvu (3).
Abnormality. No. 1. One segment near the hind end split, on
opposite side, into three segments each with setae. No. 2. Two
metameric abnormalities in region shortly in front of anus.
Homoeosis. No. 1. Male pores, as well as female, on xiv. No. 2.
Male pores on xvi, female pores on xiv. Clitellum on xxvii-xxxil,
tubercula on xxvili-xxxi but extending onto xxvil. Genital tume-
scences, on xxvi, and possibly also (?) on vii-xiil. No. 3. Female pores
on right side of xiii and left side of xiv. Male pores on right side of
xiv and left side of xv. Aclitellate, tubercula on right side of xxvi-xxx
and left side of xxvii-xxxi. Genital tumescences, on right sides ef ix
and xxv, left sides of x and xxvi.
Remarks. Worms with 79, 88, and 100 segments have large nephro-
pores, pit-like depressions at sites of apertures of setal follicles but no
setae on the last segment, and probably were amputees. Two speci-
mens, of 109 and 119 segments, had setae in the last segment and
presumably were also amputees. If any of the other worms having 67
to 101 segments are amputees, no indications thereof were recognized
and the terminal region must have been subsequently reorganized so
as to look like that of a normal worm.
In the clitellar region, intersegmental furrows are always more or
less clearly indicated. The worms can be arranged in a finely graded
series of stages showing gradual deepening of the furrows, decrease
in thickness of the epidermis, reappearance of dorsal pores, and
gradual reddening of the parietes. In the final stage, boundaries of
the clitellum were not certainly determinable but in much of the
clitellar region the red color has a sort of slight whitening that is
lacking on other segments. No spermatophores were found. No
brownish discoloration such as is present in the clitellar region of
postsexual specimens of some species, was recognized in any of the
worms.
Smith (1917) and Olson (repeatedly) have given the number of
1953 EARTHWORMS OF THE ARNOLD ARBORETUM 5
segments in this species as 95 to 150 or 90 to 145, but with no data
as to individual counts in American specimens. Segment numbers in
all of the arboretum collections are in better agreement with those of
English worms, 95 to 120, as reported by Cernosvitov and Evans (1947).
PHERETIMA AGRESTIS Goto and Hatai 1899
External characteristics. Length, 70-150 mm. Diameter, 5-8 mm.
Number of segments: 63 (3***), 64 (*), 66 (2**), 67, 69 (*), 71, 76,
77, 78 (2**), 82, 84, 85 (*), 86, 90, 92 (2**), 95, 97 (2), 98 (2), 99 (3),
100 (7), 101 (6), 102 (7), 103 (5), 104 (1). First dorsal pore, on 12/13
(56); a dark and somewhat pore-like marking shortly behind 11/12
on several specimens, definitely on 11/12 though not a functional pore,
in one worm. Setae: 52, 55, 56, 62, 62, 64, 64, 65, 69/xii; vii/2, 4, 4,
See emi lyme 2h, 4, De AA ee lay 2s 25: Villy Oycd,.0,, le 2,2, a0.
2, 2, 2, 5, 3, 4, 3, 0, 0, 0, 1, 1. (Clitellum, on xiv—xvi; annular.)
Sexthecal, spermathecal pores minute and superficial, on 5/6-7/8,
about 3C apart. Each pore may be at center of a definitely demarcated
circular porophore or the latter may have the appearance of two semi-
circles, one in front of and the other behind the pore. (Female pore,
single, median, on xiv.) Male porophores, male pores, as well as any
rudiments thereof, completely lacking.
Definite genital markings also are lacking. The setal circles of
vii-vili are uninterrupted (several specimens), or have a small gap on
one side (several) or one large median gap which may extend nearly to
spermathecal pore lines (25), or a pair of gaps leaving a few setae
midventrally (as indicated above). In the latter case, in ten specimens,
in region of each setal gap there is recognizable, in good optical con-
ditions, an area where the epidermis is thin or especially cross-hatched
with fine furrows. These areas, which have no definite boundary
though appearing to be of transversely elliptical outline, are sym-
metrically placed across the setal equator. No special pigment, brown
or otherwise, was recognized in any of those areas.
Internal anatomy. A low collar of iridescent tissue is present on the
oesophagus just behind the gizzard. The intestine begins in xv (6)
and has a double row of sacculations on each side from about xvi to
about xxv, the ventral row on each side usually more definite than the
dorsal and with one or more of the sacculations so constricted as to
have an appearance of dorsally directed pockets. The caeca are mani-
6 BREVIORA No. 15
cate, with seven or eight secondary caeca, the dorsalmost the longest.
The typhlosole, which begins in the region of the caecal segment, is
low, simply lamelliform and from about xxxix gradually decreases in
height, becoming more irregular in shape and finally unrecognizable
behind lxvi (worm of 92 segments), Ixxiv (100, 104 segments), Ixxvii
(102 segments). A very small but definite and double ridge runs
posteriorly on the floor of the gut from the caecal metamere for fifteen
or twenty segments.
Last hearts in xiii (5), heart of ix on left side (3) or right side (2),
hearts of x lacking (5). Subneural trunk much smaller anterior to
xiii but continued into 11. The dorsal trunk of one specimen passes
superficially through tissues of the pharyngeal bulb and on emerging
anteriorly bifurcates under the brain, the branches passing around the
gut median to the nervous commissures and reuniting midventrally to
become the ventral trunk.
Testis sacs unpaired, apparently above the nerve cord and with
ventral blood vessel in, on or just above the roof. Testes present,
male funnels large and nearly filling the sacs which have very little
coagulum. Seminal vesicles laterally flattened, filling coelomic cavities
of xi and xu, reaching into contact with dorsal vessel, in xii with a
primary ampulla marked off distinctly from the lamina. Vasa de-
ferentia, in region of xvi to xxiv, slightly swollen for a short distance
and ending blindly without passing into parietes. No trace of male
terminalia. Ovaries appear to be unusually large.
Spermathecal ducts shorter than contracted ampullae, with thick
wall (having a muscular sheen), abruptly narrowed at parietes, lumen
in coelomic portion large, irregularly slit-like in cross section as a
result of presence of high longitudinal ridges. Diverticulum longer
than combined lengths of duct and ampulla, into median face of duct
close to parietes, with slender stalk (having muscular sheen) longer
than the duct and a longer, more or less sausage-shaped seminal
chamber.
Distribution. Japan: Takahashi, Tokorosawa, Oarai, Sapporo,
Hokkaido, Sendai, Oshima Island, Matsuyama, Tomitaka, Kago-
shima. Said to be widely distributed throughout the Aomori and
Iawati prefectures. P. agrestis has been reported once before outside
of Japan and then from Baltimore but several attempts to secure
specimens for confirmation of the identification have been fruitless.
Abnormality. Specimen of 82 segments with one metameric abnor-
1953 EARTHWORMS OF THE ARNOLD ARBORETUM 7
mality in region of lii. First dorsal pore on 12/13 but a definitely
pore-like though obviously non-functional marking exactly on 11/12.
Left spermathecal pore of 7/8 and clitellum on right side of xiv lacking.
Female pores: on left side of xiii, in setal circle, and on right side of xv.
Heart present in xiv on right side. A testis on posterior face of left
rudiment of 8/9, a male funnel on anterior face of left rudiment of
9/10 (no testis sac but sacs and vesicles of x—xii as usual. Left ovary
lacking but a rudimentary female funnel present in left side of xii;
right ovary and female funnel in xiv. Vasa deferentiaend blindly in xxiv.
Autotomy and Regeneration. On being touched three worms in-
stantly autotomized a posterior portion (of 31, 34, 35 segments). In
one case autotomy was either through Ixiv or at 64/65. In a specimen
of 85 segments, Ixii-lxiii were deeply constricted all around the body
but apparently without recognizable break in the musculature. Pos-
terior amputees are indicated above by asterisks after setal numbers
(one for each amputee). Specimens with 63, 69, 77, 78, 86, 90 and 92
segments each have an anal region that is not marked off by an inter-
segmental furrow from a terminal segment with a complete circle of
setae. The last metamere in a 78-segment worm (a) is unusually long,
without setae, but with minute pits still representing sites of apertures
of original follicles. A similar long terminal metamere but without
setal pits characterizes a worm (b) of 76 segments but in this case
setae are recognizable, in the penultimate metamere, only within the
parietes of the ventrum. Tail regenerates, in spite of the high incidence
of posterior amputation, were not found.
In the (a) worm, loss of setae with resorption of setal follicles, and
elongation of the last segment, are regarded as external indications of
a process of reorganization, as one result of which a new growth
region, in addition to an anus, is reestablished in place of that which
had been lost by amputation. In the (b) worm the. penultimate
segment is regarded as one that had already been marked off (by an
intersegmental furrow from a reestablished growth region), but in
which equatorial setae, one of the stigmata of a metamere, had not
yet been sufficiently developed to be externally visible except in the
ventrum. Differentiation of the new metamere had not yet been fully
completed. Further, the length of the terminal (growth) region is
such as to indicate possibility of production of yet another segment.
If these interpretations are correct, P. agrestis presumably does not,
at least ordinarily replace lost posterior portions at levels behind 62/63
8 BREVIORA No. 15
by the supposedly usual process of tail regeneration. Instead, a
growth zone is reconstituted that may produce new segments, but only
slowly and one at a time. In new segments so produced, the usual
differential stigmata of regeneration, so easily recognizable in long
tail regenerates that have been produced by a much more rapid
process, may well be lacking.
The penultimate segment (counting anal region as a segment), in
several specimens that presumably had not undergone posterior ampu-
tation, had no externally visible setae and no vestiges of former setal
pits were recognizable. Both ante- and pen-ultimate segments in
another specimen (of 100 segments) were similarly incompletely differ-
entiated. In the rather small anal region of two specimens (of 100
segments each), a short rudiment of an intersegmental furrow is
present, in the ventrum (1) or dorsum and then with a rudiment of a
dorsal pore. ‘These conditions are regarded as indicating that new
segments may be produced, even in unamputated adults, by a slow,
one-at-a-time method.
Remarks. The clitellum seems to be fully developed in every speci-
men. Nevertheless no spermatozoal iridescence was noticed on any
male funnel of the six dissected specimens. Nor was spermatozoal
iridescence recognizable in any of the thirty-five spermathecae. If
sperm are produced they cannot, in absence of male terminalia, be
transferred to another individual in copulation. Every specimen in a
fair-sized sample being anarsenosomphic, reproduction must take
place by self fertilization or parthenogenesis, with the latter more
likely because of apparent absence of mature sperm in the probable
breeding season. In either case, introduction of a single worm, even
a single cocoon, theoretically could have enabled establishment of
P. agrestis in the arboretum.
Presence of a full set of seminal vesicles, and of a complete battery
of spermathecae, all now of no apparent use in reproduction, on a
principle of economy, indicates that the anarsenosomphic development
in this species has been recent.
DISCUSSION
During the summer the population of the pond periphery quite
obviously had increased considerably. Exposed but still saturated
margins were almost covered with castings of Sparganophilus eisena,
1953 EARTHWORMS OF THE ARNOLD ARBORETUM 9
the only species found (the rubellus sites, just at the high water level
not re-examined).
In the original leaf pile, E. rosea apparently had disappeared, and
two species, E. foetida and D. subrubicunda, absent in the spring, had
appeared. One of them, D. subrubicunda, had become dominant nu-
merically. JL. terrestris, however, still remained dominant by sheer
volume even though not as common as in the spring.
From the earth of the original peat bog site all worms had been
removed in the spring. Several weeks later more than fifty specimens
had moved into the disturbed soil from below or round about. Although
that same soil still appeared moist in September only seven worms
were found in it. Three species, FE. rosea, FE. tetraedra and L. rubellus
were no longer represented, but two, O. lactewm and L. terrestris, that
had been lacking at time of second collecting were again present.
In the newly examined leaf pile, Pheretima agrestis, an exotic from
Japan (and possibly brought directly from there) was clearly dominant
by weight though equalled in numbers by L. rubellus.
No cocoons or copulating individuals of any species were found in
fall collecting. A clitellum sufficiently mature for cocoon deposition
apparently characterized specimens of L. terrestris, O. lactewm, D.
subrubicunda, P. agrestis and E. foetida but appearance of the clitellar
region indicated that the reproductive period had ended in A. arnoldi
and L. rubellus. Breeding period in P. agrestis is probably in the fall.
ACKNOWLEDGMENTS
The collecting and identifying of the material considered in this
contribution constituted spare time recreation during part of a fellow-
ship period. The author’s thanks are extended to the John Simon
Guggenheim Foundation for making this contribution possible and to
the Director of the Arnold Arboretum for permission to collect therein.
REFERENCES
Crrnosvitov, L. and A. C. Evans
1947. Lumbricidae. No. 6 in Synopses of the British fauna. London,
Linnean Society.
Gates, G. E.
1953. On the earthworms of the Arnold Arboretum, Boston. Bull. Mus.
Comp. Zool. Harvard, 107, No. 10: 497-534.
Smitu, 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.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. May 28, 1953 NuMBER 16
NOTES ON THE RACES OF Micrurus frontalis
(Duméril, Duméril and Bibron)
By BENJAMIN SHREVE
As Micrurus frontalis and its associated forms appeared in need of
revision, 46 specimens were assembled for the purpose. This total
consisted of all the material in the Museum of Comparative Zoology
(M.C.Z.), two specimens from the American Museum of Natural
History (A.M.N.H.), and all those in the Chicago Natural History
Museum (C.N.H.M.).
It was found that the material fell into three groups: M. f. frontalis
(21 ex.), M. f. altirostris (21 ex.), and M. f. pyrrhocryptus (4 ex.)..
I have retained altirostris as a race of frontalis as evidence is lacking
that the two do not have separate ranges joined by an area where both
occur together and interbreed, although the supposed area of hybrid-
ization is decidedly large and there is doubt how much, if any, inter-
breeding occurs within it. Further discussion of this matter is
presented below.
I regard pyrrhocryptus as a race owing to its close relationship to
frontalis and the fact that it apparently has a range separate from
either altirostris or frontalis. :
This opportunity is taken to thank the curators, Messrs. C. M.
Bogert, C. H. Pope and K. P. Schmidt, for these loans, also Mr. K. P.
Schmidt for furnishing data on a pyrrhocryptus belonging to the
American Museum of Natural History. I also wish to thank Mr.
Arthur Loveridge for furnishing data on the head coloration of the
Uruguay series of altirostris in the British Museum (N.H.).
2 BREVIORA No. 16
Key to the Subspecies
1. Parietals without pale area except sometimes along the anterior border;
triads|on’ body 7—l'53\ventrals 26-2427 5 oso eee 2
Parietals exhibit a pale area at least anteriorly; triads on body 13-17
(l3¥only-rarely)!: vemtrals*@9) 203-220"... 222. ee eee altirostris
2. Light nuchal area between posterior end of parietal suture and first black
ring from 14 to 3% scales in width; body triads 9-15 (9-10 only rarely),
black median ring of triad usually not broader or but little broader than
the blackjouter Tings’. a4; .. chao | os ki. lakes eee frontalis
Light nuchal area between posterior end of parietal suture and first black
ring from 51% to 8 scales in width; body triads 7-9, black median ring
of triad usually twice as broad as the black outer rings. . pyrrhocryptus
MICRURUS FRONTALIS ALTIROSTRIS (Cope)
Elaps altirostris Cope, 1859, Proc. Acad. Nat. Sci. Philadelphia, p. 345:
Type locality unknown.
Schmidt (1936, p. 199) gives the type locality as South America,
but there is nothing to suggest this in the description or title of
Cope’s article.
Range. Uruguay (where frontalis is apparently unknown). Outside
of Uruguay, except where otherwise noted, it occurs alongside frontalis
with which it may possibly hybridize. Jt also occurs in the Brasilian
states of Rio Grande do Sul, Parana, Sao Paulo (probably also in
Santa Catarina and southern Mato Grosso). In Argentina it occurs
in the Gobernacién of Misiones (probably also in Corrientes Province
and possibly those of Entrerios and Buenos Aires, as well as Paraguay).
Material examined. ARGENTINA: Misiones: Iguazu Falls (C.N.H.M.
9260-2, 9383-4), Monte Carlo (C.N.H.M. 9357). Brasiz: Parand:
eastern part (C.N.H.M. 37736-7, 37743-4), Pinheiro Machado
(M.C.Z. 51495); Rio Grande do Sul: Itaqui (M.C.Z. 17853), Nova
Teutonia (C.N.H.M. 16121-2, 18475), Pelotas (M.C.Z. 27651),
Santa Maria (M.C.Z. 43337-9); State of Sao Paulo (M.C.Z. 20848),
eastern part (C.N.H.M. 37742).
Ventrals (197) 203-219 in oo", (206) 208-220 in 2 9 ; subcaudals
(16) 19-25 in oo", (16) 18-23 in 2 9; also ventrals 215-220, sub-
caudals 19-22 in two unsexed juveniles; body triads 13-17; total
length: largest co (M.C.Z. 20848) 918 (859 + 59) mm.; largest Q
(C.N.H.M. 37737) 881 (834 + 47) mm.
1 12-15 body triads according to Boulenger (1896, p. 428) on Uruguay snakes he assigned to
Elaps frontalis, but which are probably altirostris.
1953 RACES OF MICRURUS FRONTALIS 3
As regards the possibility of hybridization between altirostris and
frontalis, it has not reached a point where doubts are raised as to the
assignment of any individual. For example, an altirostris Q (C.N.
H.M. 37736) from eastern Parana has the pale area on the anterior
part of the parietals reduced, though otherwise the dark head markings,
except for being rather brownish, are those of altirostris. The body
triads are 16, which is higher than in any frontalis; ventrals 217.
Only one frontalis, a Nueva Italia o (M.C.Z. 47028) with 216
ventrals has so low a count, possibly resulting from an admixture of
altirostris “blood”. Admixture with frontalis is further suggested by a
2 (C.N.H.M. 37737) and juvenile (C.N.H.M. 9261) altirostris with
220 ventrals, as well as by other material. Further support for the
idea of hybridization is afforded by the ventral and subcaudal counts
of Uruguayan altirostris which, being presumably free from association
with frontalis, average lower than those from elsewhere. It seemed
likely that all Uruguayan altirostris might have almost entirely white
parietals such as is found in an apparently pure bred o& (C.N.H.M.
16121) from Brasil, whose ventrals number 204, subcaudals 21, and
body triads 15, but an examination of the Uruguay series in the
British Museum by Arthur Loveridge reveals that they are as variable
in this character as many from elsewhere.
The tendency for one form to show one or more characters of the
other may be due to individual variation in the nature of a “throw
back” to more primitive characters. In this event possibly hybrid-
ization does not occur and the two forms should be treated as specifi-
eally distinct, a course which probably also should be followed if
limited hybridization occurs.
Available material is insufficient to allow of a definite opinion, but
I incline to the theory of hybridization which is possibly limited by
some factor such as infertility of the resulting offspring. The evidence
is against general bybridization, though the two forms occur in many
Brasilian states and elsewhere as the following figures show. Where
the number of specimens is small those from several areas are grouped
together, and all arranged from North to South so that a comparison
may be made of the varying incidence of the two forms.
altirostris —_ frontalis
Bahia?; Minas Gerais; Rio de Janeiro 0 5
Sao Paulo 2 11
4 BREVIORA No. 16
Parana 5 2
Misiones, Argentina; Villeta, Paraguay 6 1
Rio Grande do Sul 8 2
M. lemniscatus multicinctus Amaral (1944, p. 91), with Teixeira,
Soares, Parana, Brasil, as a type locality, appears to be a composite of
altirostris and frontalis. It is not known which of the two the type
might be.
MIcRURUS FRONTALIS FRONTALIS (Duméril, Duméril and Bibron)
Elaps Frontalis Duméril, Duméril and Bibron, 1854, Erp. Gen., 7, p. 1223:
Brasil (part).
The type locality would appear to be Brasil, rather than Corrientes
and Misiones, Argentina, as given by Schmidt (1936, p. 199), for
Claussen’s two Brasilian specimens appear to be the only ones whose
description conforms to frontalis (sensw stricto) as currently regarded.
The rest of the material listed under this form in the Paris Museum
is apparently altirostris. It is interesting to note that Boulenger (1896,
p. 428) lists three Brasilian frontalis in the British Museum from
Clausen (sic). (Actually it is not known which way the name should
be spelled.) Dr. P. E. Vanzolini tells me that Claussen collected in
Lagoa Santa, Minas Gerais; so that is probably the type locality of
frontalis.
Range. Southern Brasil, where it is known from: southern’ Bahia?,
Minas Gerais, Rio de Janeiro (possibly also Espirito Santo and southern
Goias, altirostris being apparently unknown and not expected to occur
in these five states), Sado Paulo, Paranda, Rio Grande do Sul (probably
also southern Mato Grosso and Santa Catarina); Paraguay; while in
Argentina it probably occurs in the Province of Buenos Aires; and
possibly those of Corrientes and Entrerios and the Gobernacién of
Misiones.
Material examined. Brastu: Bahia?: Santa Cruz (M.C.Z. 3298);
Minas Gerais: Belo Horizonte (M.C.Z. 13954, A.M.N.H. 22573),
Uberaba (M.C.Z. 12698); State of Parané (C.N.H.M. 37206-7); Rio
Grande do Sul: Itaqui (M.C.Z. 17854), Pelotas (M.C.Z. 27652): State
of Rio de Janeiro: Rio de Janeiro (M.C.Z. 1375); State of Sao Paulo
(M.C.Z. 12699, 16684, 20844~7, 20849-50), eastern part (C.N.H.M.
37738), Butantan (M.C.Z. 17759-60). Paraauay: Villeta: Colonia
Nueva Italia (M.C.Z. 47028).
1 As M.C.%. 3298 is only labelled Santa Cruz, Brasil, it is uncertain whether it came from
Bahia or one of the other states.
1953 RACES OF MICRURUS FRONTALIS 5
Ventrals 216-236 in oo", 221-242 in 9 9 ; subcaudals 20-26 in oc",
17-23 in 2 9; body triads 9-15; total length: largest &@ (M.C.Z.
27652) 1121 (1058+ 63) mm., largest @ (M.C.Z. 13954) 860
(819 + 41) mm.
The possibility of hybridization between frontalis and altirostris has
already been dealt with under the latter. Comparison of frontalis from
different parts of its range furnished no grounds for present recognition
of further subspecies to those enumerated here. However, some
variation was noted. The light bands of the triads, normally 2 or 3
seales in width, are about 4 or 5 scales in width in a Rio de Janeiro
(M.C.Z. 1375). Sao Paulo snakes tend to have more body triads than
specimens from elsewhere, the counts not overlapping with those of
more northern snakes but overlapping with more southerly ones; these
counts in turn overlap with those of northern snakes. In the event of
a further race being recognizable, the availability of Amaral’s
multicinctus should not be overlooked.
I follow Schmidt (1936, p. 199) in regarding Elaps baliocoryphus
Cope (1859, p. 346) as a synonym of frontalis frontalis.
MicRURUS FRONTALIS PYRRHOCRYPTUS (Cope)
Elaps pyrrhocryptus Cope, 1862, Proc. Acad. Nat. Sci. Philadelphia, p. 347:
Vermejo River, Chaco, Argentina (ex Schmidt, 1936).
Range. Northern Argentina, where it is known from Mendoza and
Cordoba provinces, and likely to occur in the intermediate areas
between them and the Department of Santa Cruz, Bolivia, from
which area it is known. Probably occurs east to the Paraguay River
and east to the Parana River below its confluence with the Paraguay.
Note also the type locality.
Material examined. ArcEentiINA: (A.M.N.H. 17592); Province of
Mendoza (M.C.Z. 15911-2). Bottvra: Santa Cruz: Buenavista, 450
meters (M.C.Z. 20622).
Ventrals 224-232 in oc", 228-(231) in 2 9; subcaudals 24-25 in
oo", 24-? (tail truncated) in @ @ ; ventrals about 225 and subcaudals
25 in an unsexed juvenile. Subcaudals appear to average slightly
higher than in the other races. The count in parenthesis is that of a
9 (A.M.N.H. 65273) from La Cumbre, Cordoba, Argentina, kindly
supplied by Mr. K. P. Schmidt. Body triads 7-9; total length:
largest o’ (A.M.N.H. 17592) 948 (896 + 52) mm., largest 9 (M.C.Z.
20622) 955 (900 + 55) mm.
6 BREVIORA No. 16
The median black ring in the Bolivian example is not as wide in
relation to the other two black rings of a triad as in the two Mendoza
snakes, and it has one triad more than in any others counted. Whether
this is an indication of a Bolivian race remains to be seen.
It is probable that the ranges of all three races are more extensive
than here indicated, at least in Argentina, according to Serié (1936,
pp. 52, 53 and 55) whose nomenclature differs, but from whose
descriptions it is possible to deduce with some degree of certainty
which of the forms he had in hand.
BIBLIOGRAPHY
AMARAL, AFRANIO DO
1944. Notas sobre a Ofiologia Neotropica e Brasilica. Pap. Avul. Dept.
Zool., Sio Paulo, 5: 83-94.
BouLEnGER, G. A.
1896. Catalogue of the Snakes in the British Museum. 3: xiv + 1-727,
figs. 1-37, pls. 1-25.
Corps, E. D.
1859. Catalogue of the Venomous Serpents in the Museum of the
Academy of Natural Sciences of Philadelphia, with Notes on the
Families, Genera and Species. Proc. Acad. Nat. Sci. Philadelphia,
1859: 332-347.
Scumipt, K. P.
1936. Preliminary Account of Coral Snakes of South America. Field
Mus. Nat. Hist. Zool. Ser., 20: 189-203.
Serié, PEDRO
1936. Nueva Enumeracién Sistematica y Distribucién Geografica de
los Ofidios Argentinos. Inst. Mus. Univ. Nac. La Plata, 1936:
33-61, maps.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. May 28, 1953 NuMBER 17
A NEW ORDOVICIAN TRILOBITE
FROM FLORIDA
By H. B. Wurtrrincton
Introduction. The trilobite described below came from a core
recovered from a deep well in Madison County, Florida. It is the only
trilobite found in the numerous wells that have penetrated the
Palaeozoic of Florida, and is relatively complete. Of special interest is
its relationship to trilobites of central and southern Europe and north-
west Africa (not with any so far known in North America), and its
being by far the earliest calymenid to appear in North America. I am
indebted to Dr. J. Bridge, U.S. Geological Survey, for inviting me to
study the specimen, and to Drs. Bridge and Jean M. Berdan for
allowing me access to the results of their preliminary study of the
Palaeozoic rocks from test wells in Florida and adjacent parts of
Georgia and Alabama. The advice on the identification of the trilobite
offered by Dr. C. J. Stubblefield, Geological Survey of Great Britain,
is gratefully acknowledged.
Age. The Florida trilobite is here regarded as related to early
representatives of the calymenid-homalonotid groups, in particular to
species of Colpocoryphe, but less closely to Plaestacomia. The former
genus occurs in strata of Llanvirn (and probably early Llandeilo) age
in Bohemia and Thiiringia (R. and E. Richter, 1927), northwestern
France (Rouault, 1849; Barrois, 1900), Portugal (Delgado, 1908),
central Spain (Verneuil and Barrande, 1855), Morocco and Algeria
(Gigout, 1949; G. and H. Termier, 1950). Plaesiacomia occurs in
younger strata, of Llandeilo age, in Bohemia and possibly Normandy
(Prantl and P¥ibyl, 1948). The Florida strata may therefore be of
Llanvirn-Llandeilo age, 1. e. in North American terms approximately
late Canadian or early Champlainian.
bo
BREVIORA NO. 17
Paleogeographical Implications. Evidence brought forward recently
by Harrington and Kay (1951) and Wilson (1952) has served to
strengthen the impression that the early Ordovician faunas of South
America, the Ouachita and Appalachian troughs, and northwestern
Europe belong to the same general province. This likeness, at least
between the Appalachians, Ireland, Scotland, and the Baltic, con-
tinues into Middle Ordovician times (cf. Stubblefield, 1939, pp. 57-60).
Ordovician faunas of central and southern Europe (including West
Shropshire and South Wales), and especially those of Llanvirn-
Llandeilo time, include a characteristic group of genera, elements of
which are rare or unknown farther north and west, 1.e. in Scotland,
Ireland, and the Appalachians (cf. Stubblefield, 1939, pp. 52-55).
The Florida trilobite seems to be a lone representative of this more
southerly fauna, and its presence may indicate that the Florida
Palaeozoic rocks were laid down in a province faunally separated from
that of the Appalachian-Ouachita trough (cf. King, 1950, pp. 657-658).
In addition, Professor H. J. Harrington, University of Buenos Aires,
informs me (personal communication) that the Florida trilobite is
unlike any known in South America.
The oldest calymenid-homalonotid trilobites so far known in the
Appalachian trough (and elsewhere in North America) are Flex-
calymene and “Brongniartella”’, from Trentonian rocks of late Cham-
plainian time. This is much later than the first appearance of caly-
menids in Lower Ordovician deposits of England and Wales, and
considerably younger than the presumed age of the Florida strata in
question. Thus the Florida trilobite is a representative not only of a
different faunal province but of the earliest calymenids in North
America. On the other hand, the Ordovician of Florida is a clastic
sandstone and shale sequence, dark in color, not unlike the Llanvirn-
Llandeilo sediments of central European areas. The occurrence of a
“Bohemian’”’ type trilobite in Florida may be because of the suitable
environment there, rather than because of the existence of some other
type of barrier to faunal migration between Florida and the Appa-
lachian province.
SYSTEMATIC DESCRIPTION
Family CALYMENIDAE Milne Edwards, 1840.
Genus CoLPOCORYPHE Novak in Novak and Perner, 1918.
19538 A NEW ORDOVICIAN TRILOBITE 3
Type species. Calymene arago Rouault, 1849, from the Ordovician
(Llandeilo) of north-western France.
Discussion. The outline of the glabella, position of eyes, thorax of
13 bluntly-terminated segments, and triangular pygidium combine
to suggest that the Florida specimen belongs with the calymenid-
homalonotid group of trilobites, which appears in Lower Ordovician
times. It was first compared with illustrations of the homalonotid
Plaesiacomia rara Hawle and Corda, 1847 (p. 55, PL. 3, fig. 30; see also
Barrande, 1852, pp. 581-582, Pl. 29, figs. 21, 22; 1872, p. 40, Pl. 5,
fig. 27; Prantl and Piibyl, 1948, pp. 19-21, PI. 2, fig. 6). The outline of
the glabella, presence of two faintly defined pairs of lateral lobes, posi-
tion of the eyes (far outwards and forwards), form of the thoracic
segments and pygidium, are strikingly similar. P. rara is distinguished
by the angulate outline of the pseudo-frontal glabellar lobe, the lack
of ring furrows in the pygidial axis, lack of broad border and border
furrow on the pleural lobes of the pygidium, and presence of the first
pair of pygidial pleural furrows. The ‘small circular elevation” on the
inner posterior part of the fixed cheek of P. rara, mentioned by Prantl
and Pribyl (1948, p. 19), is apparently not present in the Florida
specimen.
Dr. C. J. Stubblefield (personal communication) suggested that the
Florida trilobite should also be compared with species of the calymenid
Colpocoryphe Novak, 1918 (in Novak and Perner, 1918), and especially
with C. inopinata Novak (in Pompeckj, 1898, p. 249; see also Novak
and Perner, 1918, p. 37, Pl. 4, figs. 1-16; R. and E. Richter, 1927, pp.
73-75, PI. 5, figs. 10, 11). The pygidium of Colpocoryphe is characteris-
tic, the axis showing several ring furrows, the pleural lobes with few,
faint pleural furrows, but with a broad, gently convex border. The
border furrow runs from about the mid-point of the anterior margin
of the pleural lobe, sub-parallel to the margin, to abut against the tip
of the axis (e.g. Barrande, 1872, Pl. 2, figs. 35, 39). The Florida
trilobite has this type of pygidium. The cephalon, however, differs
from that of C. arago (Barrande, 1872, Pl. 2, figs. 34, 35; PI. 8, figs. 11,
12) in that the eye is farther forwards and outwards, and the glabella
lacks the deep glabellar furrows and well-defined lateral lobes. The
cephalon of C. inopinata has the eyes as far out and forwards as the
Florida specimen, and the glabellar furrows are faint, the lobes lacking
independent convexity. Thus the Florida trilobite, while it bears a
considerable resemblance to Plaesiacomia rara, has the distinctive
4 BREVIORA No. 17
pygidium of Colpocoryphe, and the cephalon is like that of one species
of the latter. It is therefore placed in Colpocoryphe, and regarded as a
distinct species.
COLPOCORYPHE EXSUL sp. nov.
1 IS aaa niece ee
Holotype. External mould in dark-grey, micaceous shale from core
taken between 5154 and 5162 feet depth in Hunt Oil Company’s
J. W. Gibson No. 2 well, sec. 6, T 15, R 10E., Madison County,
Florida. This well was drilled in May, 1944, surface elevation 107 ft.,
and reached the top of the Palaeozoic rocks, beneath probable Lower
Cretaceous rocks, at 4628 ft. depth. The total depth drilled to was
5381 feet, so that 753 feet of Palaeozoic rocks were passed through,
mainly dark shale, some sandstones.
Geological Horizon. Late Lower or early Middle Ordovician (see
above).
Description. Glabella of length (sagittal) 3.7 mm., maximum width
immediately in front of occipital ring 3.8 mm., narrowing to 1.6 mm.
across pseudo-frontal lobe, gently convex, antero-lateral margins
straight, anterior margin bluntly rounded. Occipital ring .17 mm.
long (sagittal), defined by a shallow furrow curving slightly forwards
to the mid-line, deeper in the outer part. In front of the outer one-
third of the occipital furrow is a low, narrow, transversely-directed
inflation, strongest at the inner end, dying out dista!ly. In front of this
inflation the basal glabellar lobe is exceedingly faintly outlined, the
first furrows commencing at the mid-length and running diagonally
inwards to about one-third the width. The posterior edge of the lobe
is defined by the transverse inflation. Second glabellar lobes are
indicated by the second furrows, commencing at one-third the
glabellar length, running parallel to the first and extending in to one-
third the width at that point. Axial furrows deep, continuous with
deep, preglabellar furrow. Fixed cheeks gently convex, broad, outer
parts and borders not preserved. Anterior to the glabella a sagittally
short preglabellar field is continuous with the cheeks, and the anterior
margin of this field (probably the rostral suture) is a curve concave
forwards, so that the length (longitudinal) of the preglabellar field
increases outwards from the mid-line. Right palpebral lobe repre-
sented by a small, low swelling at the margin of the fixed cheek, situated
1953 A NEW ORDOVICIAN TRILOBITE 5
in line with the preglabellar furrow and 1.6 mm. out from the antero-
lateral corner of the glabella.
Thorax of 18 segments. Axial ring of second segment 2.7 mm. wide,
of last segment 1.6 mm. wide. Axis gently convex, articulating
furrows deep. Pleurae of left side only preserved, width (transverse)
of pleura of second segment 2.7 mm., of last segment 2.1 mm. Pleural
furrow deep, straight, slightly diagonal, ending against broad facet.
Tips of pleurae bluntly rounded.
Pygidium of length (sagittal) 2.7 mm., width measured over surface
approximately 4.8 mm. Axis more than one-third total width, gently
convex, the blunt tip not reaching the posterior margin. First three
ring furrows distinct, fourth (situated just beyond half length) faint.
Pleural lobe gently convex, no pleural furrows, divided at about half
width by prominent longitudinal furrow which runs from the anterior
margin sub-parallel to the lateral margin to abut against the tip of the
axis. Narrow border behind tip of axis.
External surface of exoskeleton with reticulate pattern of fine,
raised lines.
REFERENCES
BARRANDE, J.
1852. Systéme Silurien du Centre de la Bohéme. vol. I. Prague and
Paris.
1872. Systéme Silurien du Centre de la Bohéme. Supplément au vol. I.
Prague and Paris.
Barrors, C.
1900. Bretagne, Livret-Guide, VIIIS Congrés géolog. internat. 36 pp.
Paris.
Deteapo, J. F. N.
1908. Systéme Silurien du Portugal. Etude de Stratigraphie Paléonto-
logique. Comm. Serv. Géol. Portugal, pp. 1-245, pls.
Gicout, M.
1949. Le synclinal silurien et dévonien de Foucauld (Maroc occidental).
C. R. Acad. Sci., Paris, vol. 228, no. 15, pp. 1303-1304.
Harrineton, H. J., and M. Kay
1951. Cambrian and Ordovician Faunas of Eastern Colombia. J.
Paleont., vol. 25, no. 5, pp. 655-668, pls. 96, 97.
HAWwLgE, I., and A. J. C. Corpa
1847. Prodrom einer Monographie der b6hmischen Trilobiten. Prague.
pp. 1-176) pls: 1-7.
fon)
BREVIORA No. 17
Kine, P. B.
1950. Tectonic Framework of Southeastern United States. Bull. Am.
Assoc. Petrol. Geol., vol. 34, no. 4, pp. 635-671, figs.
NovAk, O., and J. PERNER
1918. Die Trilobiten der Zone. Ddyy, von Prague und Umgebung.
Paleontogr. Bohemiae, Ceska Ak. C. Frant. Jos. pro védy ete.,
no. 9, pp. 1-51 (Czech and German text), pls. 1-4.
e
PRANTL, F., and A. Prisyi
1948. Classification of the Bohemian Homalonotidae (Trilobitae).
Bull. Int. de Acad. tchéque des Sci., 46th yr., no. 9, 24 pp.,
pls: 1=2:
PompecKJ, J. F.
1898. Uber Calymmene Brongniart. Neues Jahrb., Min. Geol. Pal.,
1898 (vol. I), pp. 187-250.
RicuTer, R. and E.
1927. Uber zwei fiir das deutsche Ordovicitum bedeutsame Trilobiten.
Senckenbergiana, vol. 9, pt. 2, pp. 64-82, pl. 5.
Rovav tt, M.
1849. Memoire 1) sur la composition du test des Trilobites; 2) sur les
changements de formes. ... Bull. Soc. Géol. France, ser. 2, vol.
6, pp. 67-89, pls. 1, 2.
STUBBLEFIELD, C. J.
1939. Some Aspects of the Distribution and Migration of Trilobites in
the British Lower Palaeozoic Faunas. Geol. Mag., vol. 76, pp.
49-72.
Trermier, G. and H.
1950. Paléontologie Marocaine, vol. 2. Actual. Sci. Industr. no. 1095,
Paris.
VERNEUIL, I. bE, and J. BARRANDE
1855. Description des fossiles trouvés dans les terrains silurien ct
dévonien d’Almaden, d’une partie de la Sierra Morena et des
montagnes de Toléde. Bull. Soc. Géol. France, ser. 2, vol. 12,
pp. 964-1025, pls. 23-29.
Witson, J. L.
1952. Cambrian and Ordovician Faunas from the Marathon Region.
Guidebook, 1952 Spring Field Trip. W. Texas Geol. Soc., pp.
62-64.
EXPLANATION OF PLATE 1
Colpocoryphe exsu' sp. nov. x 6. U. 8S. Nat. Mus. Collections.
Fig. 1. Rubber mould of holotype. The black spots on the left side of the
cephalon are caused by minute holes in the mould. Fig. 2. Holotype, ex-
ternal mould.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. SEPTEMBER 23, 1953 NuMBER 18
CHARACTERS AND SYNONYMIES AMONG
THE GENERA OF ANTS
‘Part If
By Wiuuiam L. Brown, JR.
Museum of Comparative Zoology
Harvard University
What follows is a continuation! of the inquiry into the characters
and interrelationships among the genera of ants, having as its goal
the eventual reclassification of the Formicidae along more natural
and useful lines than those presently holding. Perhaps some readers
will be surprised to find that several of the genera mentioned below
were originally described in the wrong tribe or even in the wrong
subfamily, and that many monotypic genera are actually synonymous
with more familiar genera with which they have never been associated.
Nevertheless, such cases are commonplace among the largely very
artificial tribes of the larger subfamilies, and more of them will duly be
exposed, and must be exposed, before the really basic work on ant
classification can begin. A careful perusal of these cases, as well as
those treated in Part I and in some of my other papers, will make clear
the impossibility of constructing a workable key to the ant genera, and
should also open the eyes of those who now regard the existing generic
keys as basically sound. I think that I need not labor the fact that a
key bringing what now prove to be synonymous genera out to different
tribes is a key of very limited usefulness and is scarcely reliable. Only
after the synonymy is fully uncovered, justified, and formally recorded
can we begin to think of large-scale generic keys. It is suggested,
therefore, that useful generic keys will be greatly hastened if more
myrmecologists will spend a little of their time investigating and
formally publishing new synonymy.
1 Part I: Breviora, Mus. Comp, Zool. 11: 1-13 (1953).
yy BREVIORA No. 18
ANocHETUS Mayr
Anochetus Mayr, 1861, Europ. Formicid., p. 53. Genotype: Odontomachus
ghilani Spinola, 1851, by designation of Bingham, 1903.
Myrmapatetes Wheeler, 1929, Amer. Mus. Novit., 349: 6. Genotype: Myrma-
patetes filicornis Wheeler, 1929, loc. cit., fig. 8, male, monobasic. New
synonymy.
Wheeler described Myrmapatetes filicornis, “‘obviously one of the
Dolichoderinae,” from a unique male taken on Larat Island, near New
Guinea. His original figure, and even more clearly the type specimen
in the Museum of Comparative Zoology, show that he had before him
not a dolichoderine, but a ponerine male apparently representing one
of the smaller Indo-Australian species of Anochetus, and typical of
males of the latter genus in every respect save for its larger-than-usual
compound eyes. When males and workers are associated in one nest
series, further synonymy may result at the species level. Meanwhile,
Anochetus filicornis (Wheeler) becomes the necessary New com-
bination.
ASPHINCTOPONE Santschi
Asphinctopone Santschi, 1914, Boll. Lab. Zool. Portici, 8: 318. Genotype:
Asphinctopone silvestrii Santschi, 1914, loc. cit., fig. 6, worker, monobasic.
Asphinctopone (lucida) Weber, 1949, Amer. Mus. Novit., 1898: 7, figs. 5-7,
worker.
Lepidopone Bernard, 1953 (1952), Mém. Inst. Frangaise Afr. Noire, 19 (J):
207. Genotype: Lepidopone lamottei Bernard, 1953, op. cit., p. 208,
fig. 4, worker, monobasic. New synonymy.
Professor Bernard has raised the genus Lepidopone for a species
(lamottec) having the general characters of Asphinctopone, but sup-
posedly differing in details of the coxae of the posterior legs, post-
peduncle of the petiole and basiventral process of the gaster. How-
ever, it is difficult to understand how these features may be cited as
differences marking a separate genus when none of them has as yet
been reported upon for either species of Asphinctopane. From the
descriptions and figures of A. silvestrit and A. lucida, it seems probable,
not that the characters are absent, but rather that they are obscured
by glue in the types or that they otherwise escaped observation.
Bernard offers no further evidence concerning these structures of
either older species, so it is premature to insist that their presence in
the new species constitutes a valuable difference. The basiventral
1953 CHARACTERS AND SYNONYMIES AMONG ANT GENERA o
process particularly should not be singled out as a diagnostic feature
unless it is found to be developed in some unique direction; actually,
this process in some form is found widely throughout the Ponerinae
in many genera and species. However, even should the two Asphincto-
pone species prove, in the long run, to lack the unusual development
of the characters in question as seen in lamottez, it would seem that the
strong relationship of all three species named above overrides con-
siderations based on minor characters. It seems preferable by far to
avoid setting up what is at best a very weak monotypic genus when
the generic unity of the three species is so clear. Asphinctopone
lamottei (Bernard) becomes the necessary new combination.
HytomyrMa Forel
Pogonomyrmex (Hylomyrma) Forel, 1912, Mém. Soc. Ent. Belg., 20: 16.
Genotype: Pogonomyrmex (Hylomyrma) columbicus Forel, 1912, loc. cit.,
by original designation.
Lundella Emery, 1915, Bull. Soc. Ent. France, p. 191. Genotype: Tetramorium
reitterL Mayr, 1887, by original designation. New synonymy.
When Forel described Hylomyrma, he neglected to mention the
spurs of the middle and posterior tibiae. By his association of the new
group with Pogonomyrmex, the original author at least created the
impression that the spurs were present and pectinate as in other
members of the tribe Myrmicini. We still do not know whether
pectinate spurs are present in Forel’s two Hylomyrma species, but this
question hardly matters now in the face of several exceptions to the
same character in members of the Myrmicini. Forel also mentions as
a character that “the labrum, with two lateral teeth, surpasses the
anterior clypeal border.” This is certainly incorrect; what Forel
refers to here is the depressed anterior clypeal border which, in an
indeterminate Hylomyrma specimen from Central America, has a low,
blunt clypeal tooth on each side in such a position that a superficial
examination might lead to the belief that they were on the labrum.
This same indeterminate specimen has the spurs present and very
minutely and sparingly barbulate as seen at great magnification. It
seems evident from Forel’s description of the mandibles and petiolar
node, as well as other features, that Hylomyrma agrees well with
Emery’s Lundella, even though Emery assigned his genus to tribe
Tetramoriini. Lundella speciosa Borgmeier, 1937, [described in Arch.
Inst. Biol. Veg. 2: 241, figs. 25-29 (worker)] appears to me indis-
4 BREVIORA no. 18
tinguishable from Hylomyrma gocldi: Forel, if the descriptions con-
cerned are accurate. Surely, L. speciosa is a typical Hylomyrma, and
should be compared with authentic specimens of H. goeldii. Another
specimen from Nova Teutonia, Santa Catharina (F. Plaumann), con-
sidered to be a Lundella near reitteri (Mayr) by Father Borgmeier, is
also a Hylomyrma, and may be the same as goeldii, with the description
of which it agrees. The recognition of the generic synonymy thus
throws five species together and creates problems of specific synonymy
that will only be certainly settled when more is known about the types
concerned than is given in their mostly inadequate descriptions.
I believe that Jrogera, considered by Emery to be a subgenus of
Rogeria, is either very close to or synonymous with Hylomyrma. The
two Irogera species from the New World, I. procera Emery and
I. tonduzi (Forel), are known to me only from the original characteriza-
tions, which are very inadequate, and I therefore suspend judgment
until the types can be examined. Jrogera should, however, be con-
sidered as a genus distinct from Rogeria for the time being. Further
it is clear that the Oriental-Pacific species placed by Mann in Irogera,
and by Santschi in Rogeria, do not belong there, but instead should be
shifted to Lordomyrma Emery. The further study of species belonging
to the genera mentioned in this paragraph requires the review of cer-
tain crucial types before a solid rearrangement can be made. It is
apparent now, however, that Rogeria has never been clearly defined,
and that it has served as a dumping ground for ambiguous myrmicine
species throughout its existence. While the species concerned are
mostly rare and economically unimportant ones, little-known taxo-
nomically or biologically, their systematic placement will affect
myrmicine classification most profoundly.
BLEPHARIDATTA Wheeler (tribal transfer)
Blepharidatta Wheeler, 1915, Bull. Mus. Comp. Zool., 59: 484. Genotype:
Blepharidatta brasiliensis Wheeler, 1915, monobasic.
Wheeler and other authors have considered Blepharidatta to belong
to tribe Attini or tribe Dacetini, but a recent study of the types of
B. brasiliensis convinces me that the genus is really a member of the
Ochetomyrmicini. The single species is very closely related to the
species of Wasmannia Forel, differing chiefly in its more elongate head
with produced posterior angles and in having a long, low petiolar node.
Wasmannia seems scarcely to be separable from its sister-genus
1953 CHARACTERS AND SYNONYMIES AMONG ANT GENERA 5
Ochetomyrmex Mayr by means of the presently-employed character
based on the form of the elypeus. Wasmannia williamsoni Kusnezov
is one ambiguous species, and there may be still others in which the
clypeal form is intermediate. I have already shown that Hercynia
J. Enzmann is a junior synonym of Wasmannia (Brown, 1948, Ent.
News, 59: 102).
TECHNOMYRMEX Mayr
Technomyrmex Mayr, 1872, Ann. Mus. Civ. Stor. Nat. Genova, 2: 147.
Genotype: Technomyrmex strenuus Mayr, 1872, by designation of Bingham,
1903.
Aphantolepis Wheeler, 1930, Psyche, 37: 44. Genotype: Aphantolepis quadri-
color Wheeler, 1930, loc. cit., fig. 2, worker, monobasic. New synonymy.
In his figure of A. quadricolor, Wheeler shows an ant agreeing in
every respect with a Technomyrmex of group sophiae Forel, but with
a small, clearly-drawn poison-ejecting conule at the tip of the gaster.
This conule is characteristic of the subfamily Formicinae, and Wheeler
assigned his new genus to this subfamily. The type worker of A.
quadricolor has long since been dislodged from its point and lost, but
two whole legs and other fragments remain stuck to the glue on the
point. Placed beside the type pin in Wheeler’s collection are other
specimens, determined as this species by Clark and evidently con-
firmed by Wheeler, collected by T. Greaves at Cairns, northern
Queensland (the type locality is ‘Cairns dist., rotting leaves; A. M.
Lea”). These specimens agree in detail with Wheeler’s characteriza-
tion and figures, and their legs correspond with the parts on the type
point, but they do not have an ejaculatory conule. I myself took a
specimen of this ant among dead leaves on the rain-forest floor at
Kuranda, near Cairns (probably the precise locality of the original
type collection), and this agrees perfectly with the other material. It
is obvious that the ant in question is a dolichoderine, not a formicine,
and that it is most closely related to Technomyrmex sophiae Forel of
southern Queensland; Wheeler’s depiction of the gastric apex is due
to an illusion or an artifact that led him to think the conule was
present in the type; he was probably further misled by certain large
paired setae on the alitruncal dorsum that resemble those of Para-
trechina, etc. The necessary new combination is Technomyrmex
quadricolor (Wheeler); the ant is distinguished from other T'ech-
nomyrmex species by its rounded propodeum and its rather smooth,
6 BREVIORA no. 18
shining integument; it is ferrugineous yellow in color, with blackish
gaster, whereas 7’. sophiae is black throughout.
EupRENOLEPIS Emery
Prenolepis (Euprenolepis) Emery, 1906, Ann. Soc. Ent. Belg., 50: 134. Geno-
type: Prenolepis procera Emery, 1901, by original designation.
Paratrechina (Euprenolepis) Emery, 1925, Gen. Ins., 183: 223-224; synonymy
and characterization.
Chapmanella Wheeler, 1930, Psyche, 37: 41. Genotype: Chapmanella negrosensis
Wheeler, 1930, op. cit., pp. 42-44, fig. 1; worker (minor), monobasic.
New synonymy.
Wheeler described his new genus and species from a single badly
mauled worker taken from the column of a raiding Aenictus army. He
differentiated his genus from Prenolepis and Paratrechina (sensu
Emery in Gen. Ins.) by means of its small eyes, short palpi, form of
petiole and gastric base, very long appendages, and the absence of
spurs on the middle and hind tibiae. He was wrong about the last
character; the type possesses minute spurs, one to each of the two
posterior pairs of tibiae. The “apparently 6-jointed” maxillary palpi,
shown with five segments in the original figure, are four-segmented in
_the type, with the basal and apical segments short and the second and
third long. In these and other characters of habitus, eye size, etc.,
C. negrosensis shows itself to be a minor worker of one of the light-
colored species of Ewprenolepis, and it agrees well with a Euprenolepis
species (undetermined) seen from New Guinea and Cape York, except
that it has a somewhat thicker, lower petiolar node. Ewprenolepis
negrosensis (Wheeler) is the new combination.
Euprenolepis may as well be considered as an independent genus for
the time being, though further investigatién may show that it is
connected to Paratrechina by intergrades. The workers are larger
than most Paratrechina-Nylanderia species, and are, so far as is known,
markedly polymorphic; the appendages are very long and slender,
and like the rest of the body are set with numerous long, fine, erect
hairs. The species vary in pigmentation, eye size, and shape of the
petiole, but are otherwise rather homogeneous. In cabinet specimens,
the mandibles are frequently closed so tightly as to cross over one
another and lie largely hidden beneath the clypeus; this is seen in
certain other formicine and dolichoderine genera on occasion.
The species geet Wheeler apparently does not belong to EKwpreno-
“J
1953 CHARACTERS AND SYNONYMIES AMONG ANT GENERA
lepis, as Wheeler believed; by its habitus, it is more like Prenolepis,
and should be shifted to that genus pending revision of the whole
complex. I have taken geez in moist deciduous forest at 4000 feet in
the mountains behind Kuanhsien, Szechuan Province, West China; its
type locality is Mokanshan, China.
ANISOPHEIDOLE FROGGATTI (Forel)
Pheidole froggatti Forel, 1902, Rev. Suisse Zool., 10: 414, female, male.
Monomorium lippulum Wheeler, 1927, Boll. Lab. Zool. Portici, 20: 89-90,
fig. 3, worker (minor). New synonymy.
A. froggatti has a very broad range of polymorphic workers, although
the largest soldier forms have so far only been reported in flourishing
nests from extreme southwestern Australia. The distribution of this
species is broader than commonly believed, and it occurs widely in
central Australia and as far to the southeast as the Victorian Mallee.
Cotypes of M. lippulum from the MacDonnell Ranges in central
Australia (J. W. Finlayson) agree perfectly with an A. froggatti
ergatotype and other specimens, all minor workers, from South and
Western Australia. Localities for collections in the Museum of Com-
parative Zoology are as follows: Western Australia: Lion Mill ergato-
type minim (Hamburg Expedition). Manjimup; Augusta (W. S.
Brooks). Beverly (F. H. du Boulay). Woorooloo, on Darling R.;
Latham (L. Glauert). Rottnest I. (P. J. Darlington). Ludlow (J.
Clark). Kukerin (A. Douglas). Margaret River (W. M. Wheeler).
South Australia: Mt. Remarkable, Southern Flinders Ranges, ca.
1300 feet, under rock in dry open eucalypt woodland (W. L. Brown).
Cape Borda Lighthouse, Kangaroo Island, under stone in low head-
lands sand heath (Brown). Victoria: Sea Lake, mallee district (J. C.
Goudie). Northern Territory, in addition to M. lippulum type
collection: 3-5 miles west and southwest of Alice Springs, depauperate
colonies under stones in dry gullies, mulga (Acacia aneura) dominating
vegetation (Brown). Other M. lippulum cotypes, in the South
Australian Museum, are from Port Lincoln, South Australia, collected
by A. M. Lea.
Anisopheidole Forel forms with its curious monotypic sister genera
Adlerzia Forel and Machomyrma Forel a closely interrelated
group showing certain features in common with Pheidole and Stenamma.
The ranges of the single species of each of the three genera are largely
separate so far as known; Adlerzia froggatti occupies southeastern
8 BREVIORA No. 18
Australia, while Machomyrma dispar occurs in tropical and sub-
tropical Queensland. When the sexual forms of these three species
have been thoroughly compared, it may prove advisable to combine
them into one genus in spite of notable differences in the form of the
clypeus and petiole now serving to separate them generically.
The elimination of lippulwm and Adlerzia froggatti from Mono-
morium helps considerably to relieve the heterogeneity of this large
and taxonomically confused genus. Kusnezov has eliminated the
South American group Martia Forel from consideration under
Monomorium in a recent paper. It is by such small but necessary
revisionary steps as these that confusion in the Myrmicinae will
finally be eliminated.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. SEPTEMBER 23, 1953 NuMBER 19
LOWER CRETACEOUS NAUTILOIDS
FROM TEXAS
By BERNHARD KUMMEL
INTRODUCTION
Nautiloids from the Lower Cretaceous of north central Texas are
very limited in numbers of species and individuals. A recent paper by
Miller and Harris (1945) on North American Cymatoceratidae noted
only three species of nautiloids from the Washita group of north
central Texas. These are Cymatoceras hilli (Shattuck), Cymatoceras
loeblicht Miller and Harris, and Paracymatoceras texanum (Shumard).
The present study is based on 13 specimens of immature nautiloids of
the “pyrite faunae” of the Duck Creek, Pawpaw, and Grayson for-
mations, plus three specimens of ‘‘calcareous”’ preservation from the
Pawpaw formation. To the best of my knowledge this is the first
description of nautiloids of the “‘pyrite faunae’’ from the Washita.
group. Adkins (1923, p. 57) records Nautilus sp. in his faunal list of
the pyrite fauna from the upper Grayson formation, 4.5 miles south
of McGregor, McLennan County, Texas. The pyrite fauna of the
Pawpaw formation has been described by Adkins (1918) but he listed
no nautiloids, and Bose (1927) has described numerous species of
pyritized ammonites from the Grayson formation of north central
Texas. Pyrite faunae are known from the Duck Creek, Denton,
Pawpaw, and Grayson formations of the Washita group in north
central Texas.
The pyritized nautiloids are all juvenile specimens of approximately
one volution. They are tentatively placed in Cymatoceras and Para-
cymatoceras largely because they are associated with large mature
species of these two genera. One of the specimens of calcareous preser-
vation from the Pawpaw formation belongs in the genus Angulithes.
This is the first reported occurrence of this genus in North America.
There are recognized at the present 15 genera of nautiloids in the
2 BREVIORA No. 19
Cretaceous. In all of the Cretaceous formations of North America
only five genera of nautiloids are known, Cymatoceras, Paracymatoceras,
Eutrephoceras, Heminautilus, and Angulithes.
The pyrite faunae of the Washita group have been considered as
dwarfed by various authors (Scott, 1924, 1940, Winton, 1925). The
question of dwarfism of these faunae has also been discussed by
Kummel (1948). The nautiloids described here are juvenile individuals
and are not dwarfed forms.
The author expresses his thanks to Dr. J. T. Lonsdale, Director of
the Bureau of Economic Geology, Austin, Texas, for the loan of the
specimens and permission to publish this note. The Department of
Geology, University of Illinois, furnished the photographic equipment
and facilities to complete the study for which the author is grateful.
Throughout the report the Bureau of Economic Geology is abbreviated
as B.E.G.
SYSTEMATIC DESCRIPTIONS
(CYMATOCERAS sp. indet.
Plate 1, figures 1-12; plate 2, figures 1-3
Eleven small immature specimens are available for study. The
smallest specimen measures 12 mm. in diameter and the largest 21 mm.
All of the specimens include approximately one volution of the conch,
they are septate throughout, and no living chambers are preserved.
In all of them the conch is nautiliconic, the whorl section is higher
than wide, and the venter is rounded. In some of the specimens,
portions of the venter are more sharply rounded. The number of
camerae present ranges from 6 to 8. The most adoral sutures are
strongly projected adorally forming a rounded ventral saddle, a broad
concave lateral lobe and a small rounded saddle on the umbilical
shoulder. The first suture is essentially straight; however, the second
suture already shows distinctly the ventral saddle and the lateral lobe.
A small annular lobe is present in the middle of the dorsal lobe (pl. 2,
fig. 3). The siphuncle is small, rounded, and located very near the
dorsum. All the specimens are internal molds and have no surface
markings of any kind.
Small immature nautiloids are usually impossible to identify spe-
cifically and very difficult to determine generically. All the specimens
studied came from the Duck Creek, Pawpaw, and Grayson formations
and are part of the characteristic pyrite faunae of these formations.
1953 TEXAS LOWER CRETACEOUS NAUTILOIDS 3
There are also mature and relatively large specimens of calcareous
preservation of Cymatoceras hilli, Paracymatoceras texranum, and Para-
cymatoceras sp. indet. Hyatt (1894) described and figured juvenile
specimens of Cymatoceras elegans? (Paracymatoceras texanum), C. de-
slongchampsianus, C. simplex?, C. radiatum. The simple outline figures
by Hyatt of the early whorls of the above species compare very
favorably with the specimens figured here. The degree of involution,
ontogeny of the suture, and shape of the cross section are quite similar.
The writer has also compared his material with specimens of approxi-
mately the same size of various species of Cymatoceras from the Gault
of England in the British Museum of Natural History.
Branco (1880) figured the first and second camerae of Nautilus
clementinus d’Orbigny (=Cymatoceras) For Nautilus cf.
champsianus d’Orbigny (=Cymatoceras) he figured the first four septa.
The first two septa are essentially straight. The third septum shows
a well developed ventral saddle with only a very shallow lateral lobe.
The fourth septum has an even more pronounced ventral saddle and
a wide lateral lobe. This fourth septum is very similar to the septa
of the specimens of Cymatoceras described here.
Durham (1946, p. 482, pl. 68, figs. 3, 5, 7) described and figured a
juvenile specimen of Heminautilus etheringtoni from the upper Aptian
of Colombia. This specimen includes approximately one volution and
measures 6 mm. in diameter. The whorl section is subtrigonal. The
venter is slightly flattened and with a faint groove. This specimen
with its subtrigonal outline compares favorably with those of Cyma-
toceras sp. figured on plate 1, figures 10-12.
deslong-
Nautiloids in the Cretaceous of Texas are not abundant and repre-
sentatives of only four genera, namely Cymatoceras, Paracymatoceras,
Eutrephoceras, and Angulithes are known. In the Washita group
Cymatoceras and Paracymatoceras have been known for a long time.
The small immature specimens described here probably belong in
Cymatoceras. Juvenile specimens of Paracymatoceras of comparable
size already have very sinuous sutures. The adorally projecting suture
could possibly develop into the sinuated suture of Paracymatoceras in
more advanced ontogenetic stages. In respect to these two genera
Miller and Harris (1945, p. 10) state that, ‘“‘Presumably these two
genera will be found to grade more or less into each other.”” When a
more complete ontogenetic series of specimens can be assembled there
should be no difficulty in determining the proper affinities of these
specimens.
4 BREVIORA NO. 19
Occurrence. (1) Duck Creek formation, cut along Frisco R.R. about
one mile north of Denison, Texas; (2) Pawpaw formation, halfway
between Glen Garden Country Club and Sycamore Creek, 3 miles
southeast of Fort Worth, Tarrant County (locality 723 of Adkins,
1918, p. 51); (8) Grayson formation, 4.5 miles south of McGregor and
100 yards east of the highway, McLennan County (locality 966 of
Adkins, 1923, p. 52); 14 miles south of Bosqueville on the Belmont
place, McLennan County; and at Bosqueville, Mcl ennan County.
Repository. B.E.G. 297 (pl. 1, figs. 7-9), 3057 (pl. 1, figs. 4-6), 3002
(pl. 1, figs. 10-12; pl. 2, figs. 1, 2), 18750 (pl. 1, fgs. 1-3), 3273 (pl. 2,
hig. 3)):
CYMATOCERAS HILLI (Shattuck)
This species has recently been described and figured by Miller and
Harris (1945) and the description need not be repeated here. Two
specimens from the Pawpaw formation are available. They are well
preserved internal molds 70 mm. and 65 mm. in diameter, and have
no surface markings of any kind.
Occurrence. Pawpaw formation, at Blue Mound, 2 miles southwest
of Haslet, Tarrant County, B.E.G. loc. 219-T-1; and along east side
of Mansfield Road, halfway between Glen Garden Country Club and
Sycamore Creek, 3 miles southeast of Fort Worth, Tarrant County,
B.E.G. loc. 219-T-17 (locality 723 of Adkins, 1918, p. 51).
Repository. B.E.G. 17414.
PARACYMATOCERAS sp. indet. 1
Plate 2, figures 6-7
A unique specimen forms the basis of this discussion. It measures
46 mm. in diameter, is partially crushed, septate throughout, and has
the shell preserved. The conch is involute, compressed, and essentially
smooth. The venter is broadly rounded, ventral shoulders distinct
and rounded. The flanks are broadly convex and converge towards
the venter. The maximum width of the whorl is just ventral of the
umbilical shoulder. The umbilical shoulder is broadly rounded and
the umbilical wall slightly convex, sloping at an angle toward the
umbilicus which has the shape of a broad open funnel. The suture
forms a broad rounded ventral saddle, a deep lateral lobe of approxi-
mately the same width as the ventral saddle, and a second smaller
saddle on the umbilical wall. The siphuncle is not observable. The
1953 TEXAS LOWER CRETACEOUS NAUTILOIDS 5
surface of the shell has very faint ribs that are slightly sigmoidal on
the flanks and form a deep sinus on the venter. The specimen is
slightly crushed and several of the cracks in the shell follow along the
line of the ribbing. The ribbing is at most very faint but is most
distinct at the adoral part of the specimen.
Paracymatoceras differs from Cymatoceras only in the greater sinu-
osity of the suture. Miller and Harris (1945) suggest that these two
genera may be gradational. Only one species of Paracymatoceras,
namely P. texanwm (Shumard), has been recorded from North America.
That species is not uncommon in the Washita group of Texas. Para-
cymatoceras sp. indet. 1, differs from P. texanum primarily in its very
subdued ribbing. It is undoubtedly an immature specimen which
makes direct comparison difficult. Most of the specimens of P. texanwm
that have been collected in the Washita group of north Texas are
internal molds of argillaceous limestone and few of the specimens have
the shell preserved.
Spath (1927) and Miller and Harris (1945) have pointed out that
the ribbing in the Cymatoceratidae can be very subdued. The suture
of Paracymatoceras is very similar to that of Hercoglossa of the late
Cretaceous and early Cenozoic. Pseudaganides is a late Jurassic genus
also with a hercoglossid suture. In the Upper Triassic the Clydo-
nautilidae are characterized by very sinuous sutures. Each of these
nautiloid groups with sinuous sutures are probably independent de-
velopments and are not genetically related.
There are only five species of late Jurassic and Cretaceous nautiloids
that can be placed in Paracymatoceras at this time; these are: P. asper
(Oppel), the type species from Tithonian formations of Europe, P.
trichinopolitensis (Blanford) from the Ariyalur group (Senonian) and
P. rota (Blanford) from the Uttattur group (Albian) of south India,
P. tecanum (Shumard) from the Washita group of Texas, and P. sp.
indet. 1, from the Pawpaw formation of Texas. The south Indian
Cretaceous nautiloid fauna described by Blanford and Stoliczka (1861—
66) and Spengler (1910) contains numerous cymatoceratids and also
some species of Hercoglossa. As more Cretaceous nautiloids are dis-
covered it will become increasingly difficult to differentiate those
essentially smooth or weakly ribbed species of Paracymatoceras
from Hercoglossa.
Occurrence. Pawpaw formation, from a pit of the Cobb brick yard,
14 mile east of Sycamore Creek and 3 miles southeast of Fort Worth,
Tarrant County (locality 716 of Adkins, 1918, p. 47).
Repository. B.E.G. 196 (pl. 2, figs. 6, 7).
6 BREVIORA No. 19
PARACYMATOCERAS sp. indet. 2
Plate 2, figures 8-10; text figure 1
One small, immature specimen measuring 28 mm. in diameter
warrants separate discussion. The specimen is a smooth internal mold
preserved in the typical manner of the pyrite fauna of the Pawpaw
formation. It is very involute, compressed, and with a well rounded
venter. The suture forms a broadly rounded ventral saddle, and a
deep lateral lobe followed by a smaller saddle. The siphuncle was not
observed.
B
Figure 1. Diagrammatic representation of sutures. A, Paracymatoceras
sp. indet. 2, at a diameter of about 24 mm., X 4; B, Paracymatoceras sp. indet.
3, at a whorl height of 6 mm., X 7.
I was at first inclined to place this specimen in Hercoglossa because
it is perfectly smooth and has a typical hercoglossid suture, but
possible pre-Danian Cretaceous species of Hercoglossa are known only
from south India, and since Paracymatoceras sp. indet. 2 is associated
with P. sp. indet. | and P. texanum, it is highly probable that P. sp.
indet. 2 is a juvenile smooth specimen of Paracymatoceras. Very little
is known of the ontogenetic development of ribbing in Paracymatoceras.
The possibility exists, however, of this specimen being a juvenile
Hercoglossa.
Loesch (1914) has described numerous species of Upper Jurassic
nautiloids with very sinuous sutures. He includes figures of juvenile
~I
1953 TEXAS LOWER CRETACEOUS NAUTILOIDS
specimens for Nautilus schneidi (pl. 1, fig. 5a, b, ¢), N. ammoni (pl. 3,
figs. 5a, b, c), N. strambergensis (pl. 5, figs. 6a, b), and N. sp. (pl. 6,
figs. 6a, b, c). In these four species of Upper Jurassic hercoglossids
the sinuosity of the suture is very pronounced by the end of the first
quarter whorl. On the mature whorls of these species the ventral
saddle has a shallow lobe, likewise the ventral shoulders generally
become angular. The specimen from the Pawpaw formation figured
here is very similar to specimens of comparable size figured by Loesch
(1914) of Nautilus schneidi (pl. 1, fig. 5c) and N. ammoni (pl. 3,
figs. 4a, b).
Occurrence. Pawpaw formation, one-fourth mile south of the Inter-
national and Great Northern railway bridge across Sycamore Creek,
414 miles southeast of Fort Worth, Tarrant County, B.E.G. loc.
219-T-23.
Repository. B.E.G. 17425 (pl. 2, figs. 8-10).
PARACYMATOCERAS sp. indet. 3
Plate 2, figures 4, 5; text figure 1
One small internal mold consisting of only two and one-half camerae
is sufficiently distinct to be kept separate from the other specimens of
Paracymatoceras described here. The specimen is from the pyrite
fauna of the Pawpaw formation. The whorls measure 6 mm. in height
and 8 mm. in wdth. The venter is broadly rounded, flanks convex,
and the umbilical shoulders are sharply rounded. The suture has a
broad ventral saddle and a deep rounded lateral lobe followed by asmall-
er saddle. There is a small annular lobe in the middle of the dorsal
lobe. The small siphuncle is slightly dorsad of the center of the whorl.
The internal mold is perfectly smooth.
This specimen differs from the other specimens of Paracymatoceras
in having a more subquadrate outline. As with P. sp. indet. 2, this
fragmentary specimen is thought to be a portion of a juvenile indi-
vidual. Whether or not it is conspecific with the other two specimens
of Paracymatoceras described above cannot be told. The general
proportions of P. sp. indet. 3, are so different that they do not suggest
specific identity.
Occurrence. Pawpaw formation, at headwaters of Buffalo Creek,
0.5 miles southeast of Blue Mound and 2.2 miles south of Haslet,
Tarrant County, B.E.G. 219-T-2.
Repository. B.E.G. 17415 (pi. 2, figs. 4, 5).
S BREVIORA NO. 19
ANGULITHES sp. indet.
Text figure 2
This is the first representative of this genus recorded from North
America. The specimen is a somewhat weathered internal mold of
five camerae and a portion of the living chamber. The flanks are
moderately convex and strongly converge to a narrowly rounded
venter. The greatest width of the whorl is just ventrad of the broadly
rounded umbilical shoulder. The suture forms a narrowly rounded
ventral saddle, a broad shallow lateral lobe which occupies the whole
flank, and a small low saddle on the umbilical wall. The siphuncle is
3.5 mm. in diameter and located rather close to the dorsum.
Figure 2. Diagrammatic cross section of Angulithes sp. indet., X 1.
The genus Angulithes Montfort (type species Nautilites triangularis
Montfort) is characterized by a tightly coiled conch, strongly con-
vergent flanks and a venter that is angular or narrowly rounded.
Foord and Crick (1890, p. 404) point out that the venter on A. tr-
angularis is alternately rounded and sharply angular. There are only
five species of Cretaceous nautiloids that appear to belong in the genus
Angulithes; these are A. triangularis, the type species, from Ceno-
manian formations of France and England; A. tamulicus (Kossmat)
from the Danian of south India; A. westphalicus (Schluter) from the
lower Senonian of Germany; A. fleuriausianus (d’Orbigny) from the
Cenomanian of England and Europe; and A. sp. indet. from the
Pawpaw formation of Texas. Due to the fragmentary nature of the
specimen at hand direct comparison with the other species of this
genus is difficult. It is most similar in general outline to the type
species and to A. fleuriausianus. The specimen is too poorly preserved
1953 TEXAS LOWER CRETACEOUS NAUTILOIDS Q
to warrant a specific name but the record of a species of the genus
Angulithes in North America is of interest.
Deltoidcnautilus — type species Nautilus sowerbyi Wetherell — was
proposed by Spath (1927) for Eocene nautiloids with nautiliconic
conchs and more or less sagitate cross sections with narrowly rounded
to angular ventral zones. The sutures form narrowly rounded ventral
saddles, rounded lateral lobes, and a small saddle near the umbilical
wall. The siphuncle is located near the dorsum. Spath pointed out
the similarity of whorl shape of Deltocdonautilus to Angulithes; how-
ever. he thought that Angulithes “ . . . with its less sinuous suture-line
was more closely allied, via Ang. fleuriausianus (d’Orbigny) with the
regular Nautiloid stock that produced Pseudocenoceras in the Creta-
ceous, as it had given rise before to the less specialized Paracenoceras
of the calloviensis type.”
Examination of the various species of Angulithes from the European
Cretaceous and the specimen from the Pawpaw formation described
here shows no appreciable differences between Deltocdonautilus and
Angulithes. At the time Spath proposed the genus Deltocdonautilus it
was known only from the Eocene. The range of Deltoidonautilus is
now known to be Paleocene to Oligocene (Miller, 1951). The few
species of Angulithes range well throughout the Cretaceous. Because
of the similarity of Deltotdonautilus and Angulithes the genus Deltoido-
nautilus should be suppressed and its species placed in the genus
Angulithes which has priority.
Occurrence. Pawpaw formation, at hillside on north side of east-west
road, 3 miles southeast of Haslet, Tarrant County, B.E.G. loc. 219-T-8
(locality 724 of Adkins, 1918, p. 51).
Repository. B.E.G. 17421.
10 BREVIORA » No. 19
REFERENCES
ADKINS, W. 3.
1918. The Weno and Pawpaw formations of the Texas Comanchean.
Texas Univ. Bull. 1856, pp. 1-172, pls. 1-11 (1920).
1923. Geology and mineral resources of McLennan County. Texas Univ.
Bull. 2340, pp. 1-202, pls. 1-4 (1924).
BLANFoRD, H. F., and Ferp. SrouiczkKa
1861-1866. The fossil Cephalopoda of the Cretaceous rocks of southern
India; Belemnitidae-Nautilidae (by H. F. Blanford), Ammoni-
tidae, with a revision of the Nautilidae, ete. (Ferd. Stoliczka).
India Geol. Survey Mem. (Palaeontologia Indica), ser. 1 and 2,
vol. 1, pp. 1-216, pls. 1-94.
Bose, Emini
1927. Cretaceous ammonites from Texas and northern Mexico. Texas
Univ. Bull. 274s, pp. 1438-312, pls. 1-18 (1928).
Branco, W.
1880. Beitrage zur Entwickelungsgeschichte der fossilen Cephalopoden,
Theil II, Die Goniatiten, Clymenien, Nautiliden, Belemnitiden
und Spiruliden. Palaeontographica, Bd. 27, pp. 17-81, pls. 4-11.
DuruaM, J. Wyatr
1946. Upper Aptian Nautiloids from Colombia. Jour. Paleontology,
vol. 20, pp. 428-434, pls. 62-64.
Foorp, A. H., and G. C. Crick
1890. On some new and imperfectly-defined species of Jurassic, Creta-
ceous, and Tertiary nautili contained in the British Museum
(Natural History). Ann. Mag. Nat. Hist., 6th ser., vol. 5, pp.
388-409.
Hyarr, A.
1894. The phylogeny of an acquired characteristic. Am. Philos. Soc.
Proc., vol. 32, no. 143, pp. 349-647, pls. 1-14.
KUMMEL, B.
1948. Environmental significance of dwarfed cephalopods. Jour. Sedi-
mentary Petrology, vol. 18, pp. 61-64.
Lorscu, K. C.
1914. Die Nautilen des weissen Jura, erster Teil. Palaeontographica,
Bd. 61, pp. 57-146, pls. 10-15 (1-6).
MILLER, A. K.
1951. Tertiary nautiloids of West-Coastal Africa. Annales du Musée du
Congo Belge Terruren (Belgique), ser. 8, Sci. Geol., vol. 8, pp-
1-88, pls. 1-31.
1953 TEXAS LOWER CRETACEOUS NAUTILOIDS hat
— , and R. A. Harris
1945. North American Cymatoceratidae (Mesozoic Nautiloidea). Jour.
Paleontology, vol. 19, pp. 1-13, pls. 1-6.
Scorr, GAYLE
1924. Some gerontic ammonites of the Duck Creek formation. Texas
Christian Univ. Quart., vol. 1, no. 1, pp. 4-31, pls. 1-9.
1940. Paleoecological factors of Cretaceous ammonoids. Jour. Pale-
ontology, vol. 14, pp. 299-323, figs. 1-9.
Spatu, L. F.
1927. Revision of the Jurassic cephalopod fauna of Kachh (Cutch).
India Geol. Survey Mem. (Palaeontologia Indica), n. ser., vol. 9.
no. 2, pp. 1-84, pls. 1-7.
SPENGLER, ERICH
1910. Untersuchungen iiber die sudindische Kreideformation. Die
Nautiliden und Belemniten des Trichinopolydistrikts. Beitr.
Paliontologie Osterreich-Ungarns u. des Orients, Bd. 23, pp.
125-157, pls. 11-14 (26-29).
Winton, W. M.
1925. The geology of Denton County. Texas Univ. Bull. 2544, pp. 1-86,
pls. 1-27.
EXPLANATION OF PLATE 1
Figs. 1-12. Cymatoceras sp. indet. all figures X2. 1-3, B.E.G. 18750, from
Grayson formation, 4.5 miles south of McGregor, McLennan County, Texas.
4-6, B.E.G. 3057, Grayson formation, 1.5 miles south of Bosqueville, McLennan
County, Texas. 7-9, B.E.G. 297, Duck Creek formation, one mile north of
Denison, Texas. 10-12, B.E.G. 3002, Grayson formation, Bosqueville,
McLennan County, Texas.
PLATE 1
EXPLANATION OF PLATE 2
Figs. 1-3. Cymatoceras sp. indet. 1, 2, B.E.G. 3002, Grayson formation,
Bosqueville, McLennan County, Texas, X2. 3, B.E.G. 3273, front view of
camerae showing annular lobe and siphuncle, Grayson formation midway
between McGregor and Moody, Texas, X2.
Figs. 4-5. Paracymatoceras sp. indet. 38, B.E.G. 17415, 2 views of a specimen
from the Pawpaw formation, 2.2 miles south of Haslet, Tarrant County,
Texas, X2.
Figs. 6-7. Paracymatoceras sp. indet. 1, B.E.G. 196, 2 views of a specimen
from the Pawpaw formation, 3 miles southeast of Fort Worth, Texas, X1.
Figs. 8-10. Paracymatoceras sp. indet. 2, B.E.G. 17425, 3 views of a speci-
men from the Pawpaw formation, 4.5 miles southeast of Fort Worth, Texas, X1,
PLATE 2
BREVIORA
Museum of Comparative Zoology
NUMBER 20
CAMBRIDGE, Mass.
SEPTEMBER 23, 1953
LOWER TRIASSIC SALT RANGE
NAUTILOIDS
By BERNHARD KUMMEL
INTRODUCTION
Triassic nautiloids in contrast to the ammonoids are not abundant
in the fossil record nor are they very diverse in kinds. A monograph in
press by the writer on late Paleozoic and Triassic nautiloids demon-
strates that the evolutionary pattern of Triassic nautiloids is merely a
culmination of trends that started in the Carboniferous. There is no
marked change in nautiloid evolution at the Permo-Triassic boundary.
In contrast to this pattern the ammonoids suffered severe extinction in
the Permian, and in the early Triassic went through an “eruptive”
development of numerous new stocks. For both the ammonoids and
the nautiloids the Triassic-Jurassic boundary was a period of greater
crisis. Only one stock of ammonoids survived to give rise to the great
host of forms known in the Jurassic and Cretaceous. The nautiloids
likewise became nearly extinct at the end of the Triassic but a stock
probably out of the Syringonautilidae led to a new but more modest
radiation of the group.
Of the approximately 250 species of Triassic nautiloids in 30 genera,
17 are from the Seythian, 74 from the Anisian, 28 from the Ladinian,
90 from the Carnian, 39 from the Norian, and none is certain from the
Rhaetian. The Lower Triassic or Scythian species are the only ones
that will be mentioned here. These Lower Triassic nautiloids include:
Metacoceras (Mojsvaroceras) subaratum (kXeyserling) 1860, from
northern Siberia; Pleuronautilus dienert Wrafft and Diener. 1909,
Grypoceras (Grypoceras) brahmanicum (Griesbach) 1880, Grypoceras
(Grypoceras) hexagonalis Diener, 1897, Grypoceras (Grypoceras)
lilangense Krafft and Diener, 1909, from the Himalayas; Plewronautilus
subquadrangulus Tien, 1933, from south China; Pleuwronautilus kokent
Frech, 1905, Tainionautilus trachyceras Frech, 1905, Menuthionautilus
kieslingert Collignon, 1933, Grypoceras (Grypoceras) bidorsatoides n. sp.,
~
2 BREVIORA NO. 20
Grypoceras (Grypoceras) aemulans n. sp. from the Salt Range, Pakistan.
Menuthionautilus kieslingert was originally described from Seythian
strata of northern Madagascar (Collignon, 1933). The Lower Triassic
Dinwoody and Thaynes formations of Idaho have yielded new species
of Mojsvaroceras, Pleuronautilus, Enoploceras, Germanonaut'lus, and
urypoceras. The descriptions of these new species are in press (Kum-
mel, 1953). Muller and Ferguson (1939) list Grypoceras cf. brahmani-
cum (Griesbach) from the Candelaria formation of Nevada.
In the Lethaea Geognostica (Frech, 1903-1908), in the chapter on
the Asiastic Trias (written by Fritz Noetling), there are described two
species of nautiloids from the Lower Triassic of the Salt Range.
These are Tainionautilus trachyceras from the Ceratite Sandstone
(zone of Flemingites flemingianus) and Pleuronautilus kokeni from the
Upper Ceratite Marls (zone of Koninckites volutus). The chapter on
the faunal characteristics of the Salt Range Lower Triassic formations
states that there are 5 to 6 species of nautiloids of which 4 to 5 species
are undescribed (Frech, 1903-1908, pp. 167-168). These undiagnosed
specimens of nautiloids from the Lower Triassic formations of the
Salt Range are in the British Museum of Natural History as part of the
Koken Collection and are described here. Besides the three species
described below, there are two specimens of Plewronautilus (B.M.N.H.
C10429 and C10426) from the Ceratite Marls, Chideru, Salt Range,
to which Koken had applied manuscript names but they are too poorly
preserved to warrant description.
The writer is pleased to acknowledge the kind hospitality of Dr.
W. N. Edwards, Keeper of Geology of the British Museum, and
Dr. L. F. Spath, and their permission to study these specimens. The
photographs were taken at the British Museum by its staff photog-
rapher.
SYSTEMATIC DESCRIPTIONS
Family GRYPOCERATIDAE Hyatt, 1900
Genus GRYPOCERAS Hyatt, 1883
Type species. Nautilus mesodicum Hauer, 1846.
There are approximately 22 species of Upper Paleozoic nautiloids
assigned to Domatoceras (type species, Domatoceras umbilicatum
Hyatt, 1891). They are characterized by being flattened ventrally and
laterally and slightly impressed dorsally. The sutures form rounded
“~
53 LOWER TRIASSIC SALT RANGE NAUTILOIDS
ventral, lateral, and dorsal lobes. The siphuncle is subcentral in posi-
tion and orthochoanitic. Species of Domatoceras show much variation
in the degree of involution and in the position of the siphuncle.
The Triassic genus Grypoceras has long been recognized as very
similar to late Paleozoic domatoceratids (Mojsisovies 1902, p. 227).
Grypoceras differs from Domatoceras in generally being more involute,
the ventral area more rounded, the lobes more concave, and in having
an annular lobe. The Lower Triassic grypoceratids are especially
similar to typical domatoceratids. The degree of involution of 12
species of Domatoceras shows the percentage of the diameter of the
umbilicus to the diameter of the conch to range from 16 to 50. The
two most involute species, D. klethegei (17°7) and D. moorei (16%), are
transitional forms to Stenopoceras. The same data on 9 species of
Grypoceras show a percentage range from 8.5 to 30. Species of both
genera, likewise, show variability in the position of their siphuncle.
The Lower Triassic species of Grypoceras have angular ventral
shoulders, the Middle and Upper Triassic species generally have more
rounded shoulders. These data suggest that Grypoceras is a direct
evolutionary continuation of the late Paleozoic Domatoceras, and the
two genera are gradational in most characters. The principal differ-
ences are in the presence of an annular lobe and the greater involution
of Grypoceras. Under these circumstances full generic status for
Domatoceras does not seem warranted and it is here placed as a sub-
genus of Grypoceras since the latter genus has priority.
Grypoceras is widely distributed geographically, being more or less
common in the Alpine and Himalayan Triassic regions. It is also
reported in Triassic rocks from North America, the Ussuri region,
Timor, and New Zealand. Stratigraphically it is equally widespread.
There are five species in the Scythian, eight in the Anisian, three in
the Carnian, three in the Norian, and there is a form allied to G.
mesodicum in the Rhaetic of Kossen (Mojsisovics, 1873, p. 22).
GRYPOCERAS (GRYPOCERAS) BIDORSATOIDES (Koken MS) n. sp.
Plate 1, figures 1, 2
This species is based on one well preserved specimen of phragmocone
and living chamber. It measures 55 mm. in diameter, 27 mm. in height
of the last whorl, and 30 mm. in the width of the last whorl. The
umbilicus is 8 mm. in diameter. The whorl section is subquadratic in
outline. The flanks are flattened and converge toward the venter. The
4 BREVIORA No. 20
venter is flattened on the phragmocone and becomes concave on the
living chamber. The ventral shoulders are rounded and prominent.
The umbilical shoulders are more broadly rounded and the umbilical
wall is vertical. The shell is preserved on parts of the conch and is
smooth except for faint lines of growth. The septa form a broad,
shallow, ventral and lateral lobe. The siphuncle is in a subventral
position, being only 4 mm. from the venter at a point where the whor!
height is 22 mm.
Fig. 1. Diagrammatic cross-section of Grypoceras (Grypoceras) bidorsatoides
IMs Soy cll.
Remarks. G. (Grypoceras) bidorsatoides is closely related to G.
(Grypoceras) brahmanicum of the Otoceras beds of the Himalayas. The
Salt Range species is more involute and more depressed in cross section.
The siphuncle of G. (Grypoceras) brahmanicum is in a near ventral
position in the early whorls but on the last whorl has a position closer
to the center of the whorl. G. (Grypoccras) lilangense from the “ Meeko-
ceras’”’ beds of the Himalayas has a very rapidly expanding and large
conch, and no comparison can be made with G. (Grypoceras) bidorsa-
toides. The Dinwoody formation in Idaho has yielded a new species of
Grypoceras that is somewhat similar to this Salt Range species but
more compressed and with a high whorl section. Except for greater
involution, G. (Grypoceras) bidorsatoides is similar to several species of
upper Paleozoic domatoceratids.
Occurrence. Lower Ceratite Marl, zone of Prionolobus rotundatus,
Chideru, Salt Range, Pakistan.
Repository. B.M.N.H. C10428, holotype.
1953 LOWER TRIASSIC SALT RANGE NAUTILOIDS o
GRYPOCERAS (GRYPOCERAS) AEMULANS (Koken MS) n. sp.
Plate 2, figure 4
Only one moderately well preserved specimen is available for study.
It consists of a complete phragmocone and only a quarter volution of
body chamber. The conch measures 60 mm. in diameter; the most
adoral part of the last whorl measures 30 mm. in height and 27 mm. in
width. The umbilicus is 10 mm. in diameter. The whorl sides are
convex and converge toward the venter. The ventral shoulders are
angular. The venter is broad and slightly arched. The umbilical
shoulders are sharply rounded and the umbilical walls vertical. The
suture forms a very shallow ventral lobe, and a broad sweeping lateral
lobe. The position of the siphuncle is not known; it is not, however, in
a near ventral position. No part of the shell is preserved.
Fig. 2. Diagrammatic cross-section of Grypoceras (Grypoceras) aemulans
iy Op, BOE
Remarks. Grypoceras (Grypoceras) aemulans differs from G. (G.)
bidorsatoides in being more compressed and in having an arched venter,
not concave on any part of the conch. G. (G.) brahmanicum is more
evolute and with rounded ventral shoulders. G. (G.) hexagonalis
differs also in its greater involution and in the shape of the whorl
section. G. (G.) aemulans like G. (G.) bidorsatoides is similar to late
Paleozoic domatoceratids except for the greater involution of the
conch.
6 BREVIORA No. 20
Occurrence. Lower Ceratite Marl, zone of Prionolobus rotundatus,
Chideru, Salt Range, Pakistan.
Repository. B.M.N.H. C47068, holotype.
Genus MENUTHIONAUTILUS Collignon, 1933
Type species. Nautilus (Menuthionautilus) kieslingert Collignon, 1933
MENUTHIONAUTILUS KIESLINGERI Collignon
Plate 1; figures 3, 4; plate 2, figures 1-3
Nautilus (Menuthionautilus) kieslingert Collignon, 1933, Annales de Paleonto-
logie, tome 22, pp. 164-166, pl. 1, figs. 1, la.
Four incomplete but well preserved specimens of this interesting
species are available. The conch is involute, compressed, and smooth.
Fig. 3. Diagrammatic cross-section and representation of an external
suture of Menuthionautilus kieslingert, X1.
(
1953 LOWER TRIASSIC SALT RANGE NAUTILOIDS
The whorls are much higher than wide. The largest specimen (pl. 2,
fig. 1) has a whorl height of 67 mm. and a width of approximately
41 mm. The whorl sides are slightly convex and converge toward the
venter. The ventral shoulders are well rounded and the venter broadly
arched (text fig. 3).
The umbilicus is small in diameter. On the figured paratype (pl. 2,
fig. 3), 40 mm. in diameter, the umbilicus measures 6 mm. The umbili-
cal shoulders are abruptly rounded and the umbilical wall convex and
vertical. The suture forms a shallow, acutely rounded ventral lobe, a
broad, rather prominent lateral lobe which occupies the whole whorl
side ending at an indistinct saddle on the umbilical wall. The siphuncle
is In an extreme ventral position and in the largest specimen measures
2 mm. in diameter. The ventral position of the siphuncle is well
illustrated in the immature specimen figured on Plate 1, figure 3.
Remarks. Collignon’s specimen came from the Lower Triassic
(Scythian) formation at Barabanja, Madagascar. The Salt Range
specimens described and figured here are very similar in nearly all
features to the type specimen from Madagascar. The Salt Range
forms are the only other recorded specimens of this genus and species.
Collignon (1933) established Menuthionautilus as a subgenus of
Nautilus and he recognized the uniqueness of this Triassic nautiloid
with its ventral siphuncle. He compared his species with Paranautilus
and Grypoceras. There appears to be little actual affinity with Para-
nautilus. The Paranautilidae are a stock that tended toward involute,
globular conchs and simplified septa. There are very few compressed
species of Paranautilus, Indonautilus, or Sibyllonautilus. The Para-
nautilidae stem from the upper Paleozoic Liroceratidae and themselves
gave rise to the Clydonautilidac of the Upper Triassic with their very
sinuous sutures.
The inner whorls of Menuthionautilus are more quadratic in cross
section, having low arched venters and sharply rounded ventral
shoulders. The characters of the conch of Menuthionautilus show
clearly affinities to Grypoceras of the Triassic and Grypoceras (Domato-
ceras) of the Pennsylvanian and Permian. The Grypoceratidae include
the following genera: Grypoceras, Grypoceras (Domatoceras), Steno po-
ceras, Stearoceras, Titanoceras, Menuthionautilus, and Gryponautilus.
The evolutionary patterns in this family show three main trends:
(a) towards involution, (b) towards rounding of the ventral shoulders,
and (c) towards greater sinuosity of the suture. At the same time
there is great variability in the position of the siphuncle among the
S BREVIORA NO. 20
various species. In an analysis of 10 species of G. (Domatoceras) the
percentage of the distance from the venter to the siphuncle, to the
height of the whorl varied from 12.5 to 50. The same measurements
for species of Grypoceras range from 30 to 50 per cent. As mentioned
above under the discussion of Grypoceras and G. (Domatoceras) these
two forms are believed to belong to a continuous evolutionary series.
Menuthionautilus is most likely a development out of this Domatoceras-
Grypoceras line. The similarity of the whorl section, general plan of
the suture, and known variability of the siphuncle in the parent stock
strongly support this relationship.
Occurrence. Lower Ceratite Marl, zone of Prionolobus rotundatus,
Chideru, Salt Range, Pakistan.
Repository. B.M.N.H. C10483 (pl. 2, fig. 1), C10488a (pl. 1, figs.
3, 4), C10438b (pl. 2, fig. 3), C10438e (pl. 2, fig. 2).
REFERENCES
CoLLicNon, MAURICE
1933. Les Cephalopodes du Trias Inferieur, Paleontologie de Madagas-
ear, XX. Annales de Paleontologie, tome 22, pp. 151-180, pls.
14-20 (1-7).
Frecu, Fritz
1903-1908. Lethaea Geognostica, Teil 2, Das Mesozoicum, Band 1,
Trias. pp. 1-623, pls. 1-68. Stuttgart.
MojJstsovics, EDMUND VON
1873, 1875, 1902. Das Gebirge um Hallstatt. — 1. Abt. Die Cephalo-
poden der hallstatter Kalke. Abh. geol. Reichsanst. Wien, Band 6,
Heft 1, 2; Band 1, Suppl. - Heft, pp. 1-356, pls. 1-70 and 1-23.
(Heft 1, pp. 1-82, pls. 1-32, 1873; Heft 2, pp. 83-174, pls. 33-70,
1875; Suppl. — Heft, pp. 175-356, pls. 1-23, 1902.)
Mut.er, 8. W. and H. G. Ferauson
1939. Mesozoic stratigraphy of the Hawthorne and Tonopah Quadrangles,
Nevada. Geol. Soc. Am. Bull., vol. 50, pp. 1573-1624.
see
EXPLANATION OF PLATE I
The specimens illustrated on this plate are from the Lower Ceratite Marl, zone
of Prionolubus rotundatus, Chideru, Salt Range, Pakistan. They are deposited
in the British Museum of Natural History, London. All figures X 1.
Figs. 1, 2. Grypoceras (Grypoceras) bidorsatoides n. sp., B.M.N.H. —
C10428, holotype.
=);
Figs. 3, 4. Menuthionautilus kieslingert Collignon. Immature specimen;
figure 3 shows clearly the ventral position of the siphuncle. B.M.N.H. —
C10488a, paratype.
1
=
4
PLATE
EXPLANATION OF PLATE 2
The specimens illustrated on this plate are from the Lower Ceratite Marl,
zone of Prionolobus rotundatus, Chideru, Salt Range, Pakistan. They are
deposited in the British Museum of Natural History, London. All figures X 1.
Figs. 1-38. Menuthionautilus kieslingeri Collignon. 1. Mature phragmo-
cone (the cross section is illustrated in text figure 3), B.M.N.H. C10433, holo-
type. 2. Ventral view of small paratype, B.M.N.H. C10438c. 3. Side view of
small paratype, B.M.N.H. C10438b.
Fig. 4. Grypoceras (Grypoceras) aemulans n. sp. B.M.N.H. C47068, holo-
type.
BREVIORA
Museum of Comparative Zoology
9
SEPTEMBER 23, 1953 NuMBER 21
One wien
THE ANCESTRY OF THE FAMILY
NAUTILIDAE
By BERNHARD KUMMEL
The evolutionary history of the ammonoids and nautiloids is quite
parallel. The nautiloids had their greatest radiation in the early stages
of their history —in the Ordovician and Silurian — and declined
rapidly in numbers of genera per period until the Recent. The
ammonoids, however, displayed a slow start in the middle Paleozoic
but reached a peak of development in the Upper Triassic and again
in the Jurassic and Cretaceous. Both groups experienced periods of
mass extinctions with few surviving lines which formed the starting
point for new and generally even greater radiations. The transition
from the Permian to the Triassic witnessed wholesale extinctions
among the ammonoids with only the Nenodiscidae and Pronoritidae
giving rise to the early Triassic stocks (Spath, 1934). The nautiloids
at this critical boundary did not undergo such radical changes. A
number of late Paleozoic families did become extinct but three major
stocks continued uninterrupted into the Triassic.
The Triassic-Jurassic boundary was, however, a time of crisis for
both the ammonoids and the nautiloids. Only a single stock of
ammonoids survived the Rhaetic to give rise to the vast numbers of
Jurassic and Cretaceous forms (Spath, 1934). The Triassic families of
nautiloids became extinct by the Norian. No Rhaetic nautiloids have
as yet been described. The general character of Triassic nautiloids is
quite distinct from those of the Jurassic.
Spath (1927) was the first to present a comprehensive summary of
post-Triassic nautiloids. He placed all the post-Triassic nautiloids in
5 families and 28 genera, many of which were established by him.
On the problem of origin of these post-Triassic nautiloids Spath (1927.
p. 23) discussed ‘“Grypoceras cf. mesodicum” (Hauer) recorded by
Trechmann (1918, p. 181) from the Upper Triassic (Carnian, bed C)
of the Hokonui Hills, New Zealand. Spath considered this species a
, BREVIORA NO. 21]
Bisiphytes and stated that “. .. there is more probability of Bisiphytes
being an involute, globose, development of the family Syringo-
nautilidae Mojsisovies, with annular lobe.”’
Neither Trechmann (1918) nor Spath (1927) illustrated this species
and Trechmann’s description is inadequate. The purpose of this
paper is to illustrate this species and discuss its affinities, and review
the problem of ancestry of post-Triassic nautiloids. The author is
indebted to Dr. L. F. Spath and Dr. W. N. Edwards for many courte-
sies during his visit to the British Museum (Natural History) and for
permission to study Trechmann’s specimen. The photographs were
taken in the British Museum by its staff photographer.
The uniqueness of this species (here named Bisiphytes trechmanni
n. sp.) lies in the fact that it is the only tangible clue we have to
deciphering the evolutionary patterns of the nautiloids across the
Triassic-Jurassic boundary. Spath (1927, p. 23) recognized that
Trechmann’s specimen was not a Grypoceras and goes on to mention it
as “...indistinguishable from typical Bisiphytes, with their strong
strigations, especially on larger whorls, annular lobe and only slightly
sinuous septa.” With this statement I am in complete agreement.
The evolutionary patterns of Triassic nautiloids can be summarized
as those of culminating trends which began in the Carboniferous.
Three distinct lines of development are represented. One includes the
Paranautilidae, Clydonautilidae, and Gonionautilidae characterized
by globular, occluded conchs and tendency towards sinuosity of the
suture. The second major group consists of the Tainoceratidae in-
cluding most of the “ornamented” Triassic nautiloids. This stock is
generally evolute and with nodes and ribs. The Tainoceratidae in-
cludes Pennsylvanian, Permian, and Triassic genera. The third
Major group comprises the Grypoceratidae and Syringonautilidae.
The Grypoceratidae include evolute to invelute, generally smooth,
compressed nautiloids with tendency for modification of the venter.
This family also includes Pennsylvanian, Permian, and Triassic
genera. The Upper-Triassic developments of this family, Grypoceras
and Gryponautilus, are completely unlike Bisiphytes trechmanni.
Within the Syringonautilidae there are forms which are very similar
in conch patterns to Bisiphytes trechmanni. Syringoceras and Syringo-
nautilus are the most similar to the species described here. These two
genera include evolute nautiloids with rounded whorl sections in the
early volutions that become more quadrate in outline adorally. The
outer whorls have distinct umbilical and ventral shoulders and steep
1953 ANCESTRY OF THE FAMILY NAUTILIDAE 3!
JURASSIC
Nautilidae
Norian |Rhaetian
Gonionautilidae
Clydonautilidae
Bisiphytes
trechmanni
Wavasae
Scythian] Anisian |Ladinian Carnian
Syringonautilidae
Tainoceratidae
Paranautilidae
Grypoceratidae
Fig. 1. Phylogenetic diagram illustrating the relationships of the Liassic
Nautilidae with the Triassic nautiloid families. The columns representing the
Triassic families have been terminated at an arbitrary even line in the Norian.
The exact relative extinction dates of these families in the Norian are not
known.
umbilical walls. The shell bears fine radial and longitudinal lines.
These two genera differ only in the position of the siphuncle; Syringo-
ceras has the siphuncle very near but not at the venter, Syringonautilus
has its siphuncle in a more central position. There are about 14
species of Syringonautilus, with 9 species in the Anisian, 5 in the
Ladinian, 3 in the Carnian, and | indeterminate species in the Norian
(Some of these species occur in more than one epoch.) Of Syringoceras
4 BREVIORA No. 21
there are 17 species recorded, | from the Anisian, 3 from the Ladinian
1? from the Carnian, and | from the Norian. The pattern of develop-
ment of these two genera is markedly different in that Syringonautilu :
had a large radiation soon after its appearance and then declined
rapidly; Syringoceras had its maximum radiation in the Carnian, after
a slow development in the Anisian and Ladinian. Both stocks are
represented by a single species in the Norian.
The Syringonautilidae also include three aberrant genera of Norian
age, namely Clymenonautilus, Juvavionautilus, and Oxynautilus.
Clymenonautilus has a conch pattern like that of Syringonautilus, but
has a very sinuous suture convergent to the type of suture present in
the Clydonautilidae. Juvavionautilus, in its conch shape, is a hetero-
chronus homeomorph of Domatoceras of the Pennsylvanian and Per-
mian. Oxynautilus is an involute oxycone similar in its conch pattern
to Stenopoceras of the Pennsylvanian and Permian. Clymenonautilus
and Oxynautilus are monotypic and only 6 species are known of
Juvavionautilus.
On the basis of the prominent peripheral strigations this species is
placed in bisitphytes and not in Syringoceras or Syringonautilus. The
latter two genera are more evolute than Bisiphytes, the umbilicus
approximating 25 to 30 per cent of the diameter of the conch. The
umbilicus of B. trechmanni is about 19 per cent of the diameter of the
conch. Among typical Liassie Bisiphytes the width of the umbilicus
ranges from around 20 per cent to completely occluded conches as in
B. simillimus Foord and Crick. A general evolutionary trend among
the coiled cephalopods, except the heteromorph ammonoids, is
towards greater involution.. Among Triassic nautiloids this trend is
very apparent in the Domatoceras-Crypoceras line in the Grypo-
ceratidae and the Metacoceras-Mojsvaroceras \ine in the Taino-
ceratidae. The shape of the whorl section and suture of B. trechmanni
is quite similar to such species of Syringoceras as S. credneri Mojsiso-
vies, S. evolutus Mojsisovies or species of Syringonautilus as S. lilianus
(Mojsisovies), and S. longobardicus (Mojsisovies).
Early Jurassic Nautilidae are accomodated in 5 genera including
Bisiphytes Montfort, 1808, Cenoceras Hyatt, 1883, Sphaeronautilus
Spath, 1927, Digonioceras Hyatt, 1894, Ophionautilus Spath, 1927.
Sphaeronautilus is monotypic, Digonioceras has 4 species, and Ophio-
nautilus only 5 species. The remaining species of early Jurassic
Nautilidae belong to Bistphytes and Cenoceras. The Paraceno-
ceratidae, Hercoglossidae, and Cymatoceratidae are derived from the
1953 ANCESTRY OF THE FAMILY NAUTILIDAE 5
Nautilidae.
Liassic nautiloids have received rather thorough treatment by
Prinz (1906) and Pia (1914). Pia considered the attempt at a phylo-
genetic arrangement of Liassic nautiloids by Prinz as un-uccessful
and premature. Spath (1927) came to the same conclusion. Although
Pia (1914, p. 45) was not prepared to offer a detailed phylogenetic
arrangement of Liassic nautiloids, he made the following tentative
thesis on the evolution of the great majority of Liassic nautiloids:
1.) The conch was probably originally rounded without angular
shoulders (perhaps broader than high).
2.) The primitive sculpture consisted of radial and longitudinal
striae of equal strength. A smooth shell is a secondary develop-
ment.
3.) The reduction of the longitudinal striae took place first on the
whorl sides, afterwards on the venter.
4.) The siphuncle was originally round.
5.) The annular lobe is a primitive character, its absence a specializa-
tion.
6.) The umbilicus was originally open.
Pia considered Nautilus striatus as being morphologically very
similar to the primitive original stock of Liassic nautiloids.
Of the known Upper Triassic nautiloid stocks, only the Syringo-
nautilidae appear to be possible ancestors to the post-Triassic forms;
the remaining Triassic stocks are specialized developments morpho-
logically unsuited to be potential ancestors to the known. Liassic
nautiloids. Bisiphytes is thus considered a late Triassic off-shoot
of the Syringonautilidae, as first suggested by Spath (1927, p. 23),
and in the direct line of ancestry of all later nautiloids.
SYSTEMATIC DESCRIPTION
Family NAUTILIDAE d’Orbigny, 1840
Genus BISIPHYTES Montfort, ISOS
BISIPHYTES TRECHMANNI N. sp.
Plate 1, figures 1, 2, 3; text figure 2
Grypoceras cf. mesodicum (Hauer). Trechmann, Quart. Jour. Geol. Soc.
London, vol. 73, pp. 181-182, 1918.
The single specimen upon which this species is based is a moderately
large phragmocone. It measures 71 mm. in diameter, 38 mm. in
height of the last whorl, and approximately 50 mm. in width of the
>
6 BREVIORA No. 21
most adoral part of the last whorl. The conch is evolute, the umbilicus
measuring 13.5 mm. in diameter. The venter is broadly rounded, as
are the ventral shoulders. The whorl sides are somewhat flattened
and convergent. The widest part of the whorl is just above the
umbilical shoulders which are more sharply rounded than the ventral
shoulders. The umbilical wall is steep and convex.
Only fragmentary portions of the shell are preserved. The conch
bears fine strigations which are present both on the whorl sides and
the venter of the inner whorls. On the most adoral volution the
strigations are present only on the venter. On this region there are
about 7 lines in a width of 5 mm.
Fig. 2. Diagrammatic cross section of Bistphytes trechmanni n. sp.
The suture forms a broad, very shallow ventral lobe and a slightly
deeper lateral lobe. There is an annular lobe. The siphuncle is sub-
ventral in position. It is 2 mm. in diameter and lies 7 mm. from the
venter.
Occurrence. Upper Triassic, Carnian, Bed C, Otamita, Hokonui
Hills, New Zealand. |
Repository. British Museum (Natural History) C 21947.
1953 ANCESTRY OF THE FAMILY NAUTILIDAE
~
REFERENCES
Pra, JULIUS
1914. Untersuchungen itiber die liassischen Nautiloidea. Beitr. Pala-
ontologie Osterreich-Ungarns u. des Orients, Bd. 27, pp. 19-86,
pls. 4-10.
Prinz, JULIUS
1906. Die Nautiliden in der unteren Jura-Periode. Ann. Hist.-Nat.
Mus. Nat. Hung., vol. 4, pp. 201-248, pls. 3, 4.
Spats, L. F.
1927. Revision of the Jurassic cephalopod fauna of Kachh {(Cutch).
India Geol. Survey Mem. (Palaeontologia Indica), n. ser., vol. 9,
mem. 2, pp. 1-84, pls. 1-7.
1934. Catalogue of the fossil cephalopods in the British Museum
(Natural History), Part 4, The Ammonoidea of the Trias (1).
pp: 1-621), pls. 1-18.
TRECHMANN, C. T.
1918. The Trias of New Zealand. Quart. Jour. Geol. Soc. London, vel.
73, pp. 165-246, pls. 17-25.
‘IN “LPG1IZO (Atoqsuy [BIN}BN) WUNesnyy YsHig “puryeez MoN ‘STL Muoyory
‘RYNUBIC, ‘) peg ‘ueturey ‘orsseny, teddy “ds ‘u tuunuyoos) sajfiydisig
I ALWId fO NOILYNVIdXH
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. OcTOBER 13, 1953 NUMBER 22
NOTES ON THE ANT,
LEPTOTHORAX OBLIQUICANTHUS COLE
(Hymenoptera: Formicidae)
By Rosgrt E. GrRreae
Department of Biology, University of Colorado
A new species of Leptothorax was described recently from New
Mexico (Cole, 1953), together with some memoranda concerning the
distribution of the genus in that state. For some years the writer has
been collecting the ants of the neighboring state of Colorado, and in
the course of these investigations discovered what appeared to be an
entirely distinct form of Leptothorax. This ant was confirmed as new
by Dr. Creighton, and I was in the process of drawing up a description
and the figure of a specimen when Dr. Cole’s paper was published.
Upon comparing his description with my ants, it is certain that we are
both dealing with the same form. Three paratypes were sent to me,
and when compared with my specimens prove to be almost identical.
The only differences worthy of note are that the Colorado specimens
have slight rugulations on the ventral borders of the epinotum (absent
in the types), the dorsal petiolar notch is absent, and the ventral
petiolar spinule is only weakly developed. Dr. Cole’s name for the
species obviously has priority over the one about to have been pro-
posed, but since the figure was already finished, and since the ant is so
distinctive for the genus Leptothoraz, it is thought advisable to publish
it herewith.
As Dr. Cole points out, the ant differs from the closely related
members of the tricarinatus-tecanus complex, and particularly from
L. t. neomexicanus in its shorter scapes, larger epinotal spines, and
differently shaped petiole, ete. By far the most unusual feature, how-
ever, is the pair of huge, subreniform compound eyes, composed of
2 BREVIORA NO. 22
160 to 180 facets. Dr. Cole makes careful note of the existence of this
anatomical character, but unfortunately failed to use it in deriving
the specific name.
In his subgeneric allocation of obliquicanthus, Cole was constrained
to follow the recent changes in the taxonomy of Leptothorax proposed
by Smith (1950), in which the subgenus Leptothorax was replaced by
the name Myrafant. According to this treatment, Cole designated his
new species, Leptothorax (Myrafant) obliquicanthus. But the advis-
ability of Dr. Smith’s revision is open to question, and the problem
has been discussed at length in an article by Creighton and Gregg
submitted to the International Commission on Zoological Nomencla-
ture. In this paper we have taken the position that the changes
advocated by Smith will result in endless confusion, not only because
of the long established concepts concerning the genus as a whole, but
because of the necessity of redefining the characters of the subgenus
Leptothorax. Before 1950 one group of species with a certain set of
traits would belong in the subgenus Leptothorax, and after 1950 an
entirely different set of species with a different set of characters would
be known in the same subgenus. A more incongruous situation could
hardly be imagined. We have attempted to rectify the condition by
asking the Commission to set aside the Rules and adopt certain
names as ‘‘nomina conservanda’’, namely, to retain the taxonomy of
Leptothorax essentially as it was left by Emery.
Pending the action of the Commission, and until a definite decision
is reached by that body, the writer believes unnecessary confusion can
be avoided through adherence to the older nomenclature. Such a
procedure is not without precedent and would appear to be the much
sounder course to follow. Therefore, the treatment which should be
given to the recently described species calls for a restitution of the
subgeneric name Leptothoraz, thus:
LEproTHoRAX (LEPTOTHORAX) OBLIQUICANTHUS Cole
The specimens of obliquicanthus which I collected came from
Higbee, Colorado, in the southeast part of the state, twenty miles
south of La Junta. They were obtained from two general habitat
types, semi-moist meadow near the Purgatoire River, and high dry,
short grass plains above the river valley. The approximate elevations
of these sites are 4400 ft. and 4500 ft. respectively. Dr. Cole’s speci-
mens were taken twelve miles south of Santa Fe, New Mexico, in a
1953 LEPTOTHORAX OBLIQUICANTHUS COLE 3
grassy area. Though the species is now known to exist over a fairly
wide territory, much collecting will be required to learn the precise
nature of its range.
The unusual size, position, and form of the eyes in obliquicanthus
raises the question of their possible function and adaptive value.
Dr. Creighton (1930, p. 121), in discussing a similar phenomenon in
Solenopsis (E.) macrops Santschi, points to Santschi’s observation that
convergent adaptation may be involved. In Oxyopomyrmex, according
Figure 1. Leptothorax obliquicanthus Cole, worker. Drawn from a specimen
collected by the writer at Higbee, Colorado.
to Santschi, the workers stop at the entrance of the nest before going
out, in such a position that their large, elongate eyes are able to scan
the surroundings. It may not be too much to suggest that the over-
sized eyes of these three forms represent analagous developments, and
that their functions are connected with diurnal habits in open areas.
LITERATURE CITED
Coxz, A. C. 1953. Notes on the genus Leptothorar in New Mexico and a
description of a new species. Proc. Ent. Soc. Washington, 55: 27-30.
Cretcuton, W. 8. 1930. The New World species of the genus Solenopsis.
Proc. Amer. Acad. Arts Sci., 66: 39-151.
CrEIGHTON, W. S. and R. E. Greaa, 1953. An appeal to the International
Commission on Zoological Nomenclature for the conservation of certain
generic and subgeneric names of ants. MS.
SmirH, M. R. 1950. On the status of Leptothorax Mayr and some of its
subgenera. Psyche, 57: 29-30.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. OctroBER 23, 1953 NuMBER 23
A NEW SPECIES OF HYPOGNATHA
FROM PANAMA
By Artruur M. CHICKERING
Albion College, Albion, Michigan
Among the less common argiopid spiders found in Panama is a
species of the curious genus Hypognatha. Up to the present time only
one species of this genus has been known from Central America,
H. nasuta O. P. Cambridge, originally described from Mexico. During
the preparation of this paper I have been able to study specimens of
H. nasuta from the collection of the British Museum (Natural History)
and collected at the type locality. I have also been able to examine
at least two species of the genus from South America in the collection
of the Museum of Comparative Zoology. On the basis of this study
I have been compelled to consider the species from Panama as new
to science.
The study of the new species, together with others available to me,
leads to the conclusion that the description of the genus as given by
Simon (94) is wholly inadequate for our present usages and that the
whole genus should be carefully restudied and redefined. Such a study
is not contemplated at this time. Hence, the present paper will be
concerned merely with a description of the new species in accord with
my usual procedure.
Acknowledgements are gratefully extended to the following persons
for their interest and cooperation in aiding me in the preparation of
this paper: Dr. A. S. Romer, Director of the Museum of Comparative
Zoology, Harvard College, and Dr. P. J. Darlington, Jr., Curator of
Entomology in the Museum of Comparative Zoology, where this paper
was completed; Dr. G. Owen Evans and other members of the staff
of the British Museum (Natural History) for the loan of valuable
material; the donors of the Penrose Fund of the American Philosophical
Society, and the Society of the Sigma Xi from both of which I received
grants to enable me to pursue field studies in Panama during the
summer of 1950.
BREVIORA NO. 23
Genus HYPOGNATHA Guerin, 1839
HyYPOGNATHA ELABORATA Sp. Nov.
(Figures 1-10)
Male holotype. Total length 3.12 mm. Other features essentially
as described for the female allotype.
Eyes. Eight in two very strongly procurved rows: PLE pearly white;
central ocular quadrangle equally wide in front and behind; almost as
wide as long. Ratio of eyes AME : ALE : PME: PLE = 11 :6:10:6.
PLE irregularly oval, others circular. AME separated from one
another by 8/11 of their diameter, from ALE by four times their
diameter (straight line distance). PME separated from one another
Fig.
Fig.
Fig.
Fig.
Fig.
OX fea Ct
External Anatomy of Hypognatha elaborata sp. nov.
Figures 1-5
Eyes, clypeus, and cephalic horns of male, from in front.
Median cephalic horn of male in profile.
Lateral cephalic horn and LE of male.
Left male palp, ventro-lateral view.
Left male palp, distal end of tarsus.
1953 A NEW SPECIES OF HYPOGNATHA 3
by 9/10 of their diameter. Laterals separated from one another by
one half the diameter of PLE. Height of clypeus equal to slightly
more than three diameters of AME. Clypeus with a robust curved
median horn arising from just above the middle of this area; with a
lateral horn on each side bearing at its distal end the two lateral eyes
(Figs. 1-3).
Chelicerae. Basal segment .65 mm. long; other features essentially
as recorded for the female allotype.
Mazillae, Lip, and Sternum. Essentially as described for the female
allotype.
Legs. 1243. Width of first patella at “knee” .184 mm., tibial index
of first leg 14. Width of fourth patella at “knee” .13 mm., tibial index
of fourth leg 15.
Femora Patellae Tibiae Metatarsi Tarsi Totals
(All measurements in millimeters)
1 975 487 769 975 .430 3.636
2. .968 462 682 858 A12 3.382
3. .682 202 396 418 308 2.056
4. .902 330 090 770 063 2.915
Palp 352 176 240 SS 012 1.340
Spines. Even more reduced than recorded in the female. Probably
here it would be well to regard them as being replaced by bristles.
Palp. Patella and tibia short, without apophyses. Tarsus compli-
cated; with embolus coiled broadly at distal end of bulb and near the
tip it passes into a deeply grooved conductor and completes a second
circle at a right angle to the first. Near the middle ofthe ventral surface
there is a long, slender, undulating, lash-like apophysis; near the base
there is a stout, two-pronged apophysis (Figs. 4, 5). Altogether
different from the palp of H. nasuta O. P. Cambridge. If what I have
called the embolus at the distal end of the bulb were lacking, then
what I have termed the median apophysis probably would be termed
the embolus. This raises certain questions about the identification of
parts in H. nasuta O. P. Cambridge as well as in this species.
Abdomen. 2.34 mm. long; 2.470 mm. wide about two fifths from
base where it is widest; the chitinous ring nearly surrounding spinnerets
and anal tubercle less well developed than in female; covers somewhat
less of the carapace than in the female; dorsum much less regularly
subdivided into angular areas than in female; four dorsal sigilla very
clear.
4 BREVIORA NO. 23
Color in alcohol. Essentially as recorded for the female except as
recorded below: carapace and chelicerae somewhat lighter in color;
testudinate dorsum without such distinct angular areas; with many
whitish sub-chitinous granules so that nearly the whole dorsum is
whitish; venter much less clearly marked with brown oblique and
transverse bands.
Female allotype. Total length 3.185 mm. Carapace so nearly
covered by abdomen that ordinary description of this part is nearly
impossible. Observations on a paratype show that median longi-
tudinal thoracic groove is lacking; considerably constricted opposite
interval between first coxae and palpi; posterior two thirds smooth;
anterior third granular on surface; appears to be devoid of spinules
and with a scant supply of hair.
Eyes. Eight in two rows; all dark except PLE which appear light
perhaps because of degeneration; both rows would ordinarily be
considered strongly procurved. Viewed from above, posterior row
nearly straight; viewed from in front, both rows strongly procurved;
central ocular quadrangle equally wide in front and behind, slightly
longer than wide. Ratio of eyes AME : ALE : PME : PLE =
11 :8:11:7. AME separated from one another by a little less than
their diameter, from ALE by four times their diameter. PME
separated from one another by a little less than their diameter, from
PLE by five times their diameter. Laterals separated from one
another by one eighth of the diameter of ALE. PLE irregularly oval;
all others circular (Fig. 6). Height of clypeus equal to 37/11 of the
diameter of AME.
Chelicerae. Basal segment .75 mm. long; somewhat receding; quite
robust; basal boss well developed; fang well developed and finely
dentate along inner margin particularly in basal half; fang groove with
five teeth along promargin and four along retromargin (observed
clearly on dissected paratype but confirmed on allotype).
Mazillae. Slightly convergent; broadly rounded along outer margin
where serrula is very extensive.
Lip. Wider than long in ratio of about 2 : 1; gradually narrowed
to a rounded point distally; deeply grooved transversely at base.
Sternal suture distinctly procurved.
Sternum. Of a modified cordiform shape; wider than long in ratio
of about 4 : 3; deeply notched at posterior end; the notch is filled by
a sclerite in the form of a peg; with posterior coxae separated by about
1.3 the width of one of them; with a scant covering of short stiff
bristles (Fig. 7).
A NEW SPECIES OF HYPOGNATHA 5
Fig.
Fig.
Fig.
Fig.
Fig.
External Anatomy of Hypognatha elaborata sp. nov.
Figures 6-10
Eyes and clypeus of female allotype, from in front.
Sternum of female allotype.
Female allotype, dorsal view.
Epigynum of allotype, posterior view.
Epigynum of dissected paratype to show complete posterior exposure.
Legs. 1243. Width of first patella at ‘knee’ .195 mm., tibial index
of first leg 14. Width of fourth patella at “knee” .1624 mm., tibial
index of fourth leg 15.
Onr
—
Femora Patellae Tibiae Metatarsi Tarsi Totals
(All measurements in millimeters)
975 920 15 921 440 3.571
975 _ 020 .650 814 418 3.377
812 380 418 418 352 2.380
1.100 440 616 .780 .400 3.336
6 BREVIORA NO. 23
Spines. True spines almost completely lacking. A few weak spines
may be recorded as follows: First leg: femur dorsal one near middle,
one prolateral near distal end; tibia prolateral one near proximal end.
Third leg: patella apparently with a dorsal distal spinule. Three tarsal
claws throughout. Palpal claw: pectinate in a single row.
Abdomen. 2.73 mm. long; 3.25 mm. wide near middle; very much
flattened; with a soft membranous ring nearly surrounding the six
spinnerets and anal tubercle; tracheal spiracle with a well chitinized
lip just anterior to base of anterior spinnerets.. Covers nearly two
thirds of the carapace. Dorsal angulate areas clearly separated by
narrow lines (Fig. 8).
Epigynum. Epigynal plate a broad, rounded ridge with openings
and other characteristic parts directed posteriorly and partly hidden
by genital groove (Fig. 9). Posterior margin more clearly shown in
dissected paratype (Fig. 10).
Color in alcohol. Carapace a light rich reddish brown with nearly
black indefinite flecks and streaks. Sternum much the same but with
more dark flecks. Chelicerae a rich amber color. Legs yellowish with
many dark flecks varying much on the different segments. Abdomen:
the strongly chitinized dorsum is generally light brown but is finely
pitted and dotted with many brownish specks; the angular areas are
separated by yellowish lines; the venter is progided with several brown
oblique and transverse bands alternating with narrow light lines.
Type locality. Male holotype from the Barro Colorado Island,
C. Z., July, 1934; no mature male paratypes; female allotype from the
same locality July, 1936. Paratype females from the same locality
July-August, 1936; July, 1939; July-August, 1950. Several immature
males and females July, 1936, 1939.
HypoGNnaTHa NAsutTA O. P. Cambridge, 1896
(Figures 11-15)
H. nasuta F. P. Cambridge, 1904
H. nasuta Petrunkevitch, 1911
The following notes are based upon a study of specimens of this
species loaned by the British Museum (Natural History); collected
April, 1905 at Teapa, Tabasco, Mexico; from the Godman and Salvin
collection.
Male. At the base of each lateral cephalic horn there is a groove
ending in a shallow pit beneath each AME. The bases of the lateral
=I
1953 A NEW SPECIES OF HYPOGNATHA
horns are relatively much broader than in H. elaborata sp. nov. and
the clypeus is extended into a broad shelf from near the ventral border
of which the medial horn extends (Figs. 11, 12). The cephalic part of
the cephalothorax is roughened by shallow pits. The palp appears
to follow the same general pattern as in H. elaborata sp. nov. with the
long slender median apophysis extending much as in the new species,
but what is here regarded as the long coiled terminal embolus in the
new species is a relatively short tube following closely the conductor
and very slightly coiled (Fig. 13). The dorsum is essentially like that
in the new species.
External Anatomy of Hypognatha nasuta O. P. Cambridge
Figures 11-15
Fig. 11. Eyes, elypeus, and cephalic horns of male.
Fig. 12. Median cephalic horn of male in profile.
Fig. 18. Left male palp, ventro-lateral view.
Fig. 14. Dorsal view of female abdomen.
Fig. 15. Epigynum of female; a somewhat posterior view.
8 BREVIORA NO. 23
Female. Angular areas on the chitinized dorsum are bordered by
much broader margins than in the new species (Fig. 14) and the whole
abdomen is much deeper and more robust. Total length 3.51 mm.;
abdomen 3.055 mm. long; greatest width 3.445 mm. Ratio of eyes
AME : ALE : PME: PLE = 9:6:9:6. AME separated from one
another by slightly less than their diameter, from ALE by slightly less
than four times their diameter. PME separated from one another by
slightly less than their diameter, from PLE by slightly less than five
times their diameter. The epigynum has a central depressed region
posterior to a strongly chitinized ridge (Fig. 15).
BIBLIOGRAPHY
CAMBRIDGE, O. P. and F. P. CamBripce. 1889-1905. Arachnida-Araneida.
Vols. I-II. Jn: Biologia Centrali-Americana. Dulau & Co,, London.
Smmon, Euckne. 1892-1903. Histoire Naturelle des Araignées. Deuxiéme
édition. 2 Vols. Libraire Encyclopédique de Roret, Paris.
PETRUNKEVITCH, ALEXANDER. 1911. A Synonymic Index-Catalogue of
Spiders of North, Central, and South America, etc. Bull. Amer. Mus.
Nat. Hist., 29: 1-809.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. JANUARY 27, 1954 NuMBER 24
A NEW FAMILY, A NEW GENUS, AND
TWO NEW SPECIES OF BATOID FISHES
BROME DHE, GULE OF MEXICO
By Henry B. BIGELOW
AND
WILLIAM C. SCHROEDER!
The West Indian-Gulf of Mexican region is proverbially poor in
skates as compared with the more northerly coastal waters of the
Atlantic; so much so, indeed, that two specimens only, of Raja akleyi
Garman 1881, from the Yucatan Bank and nearby, were the only
skates that had been reported in scientific literature from any part of
the Gulf prior to 1921. This is fewer than one is likely to find, stranded,
along the beaches of Cape Cod during an hour’s stroll on any summer
day. And while a second species, R. texana Chandler 1921, is now
known to occur in some numbers around the northern shores of the
Gulf from Florida to Texas, it was not until the winter and spring of
1938-1939 that the trawling campaigns of ATLANTIS brought to
light the presence of a varied skate fauna at depths greater than 200
fathoms around the coasts of Cuba.
Examination of these, of collections of skates in the U. S. National
Museum that had been taken by the ALBATROSS many years ago,
and of others taken recently by OREGON of the U.S. Fish and Wild-
‘life Service, had brought to light ten new skates of the genera Raja,
Breviraja and Cruriraja,? up to 1950, from Cuban waters and from the
northern part of the Gulf; also of a new genus, Springeria Bigelow and
1 Contribution No. 657 from the Woods Hole Oceanographic Institution.
2 These are Raja lentiginosa Bigelow and Schroeder 1951; R. olseni Bigelow and Schroeder
1951; R. teevani Bigelow and Schroeder 1951; Breviraja atripinna Bigelow and Schroeder 1950;
B. colesi Bigelow and Schroeder 1948; B. cubensis Bigelow and Schroeder 1950; B. sinus-
mexicanus Bigelow and Schroeder 1950; B. yucatanensis Bigelow and Schroeder 1950; Cruriraja
atlantis Bigelow and Schroeder 1948 and C. poeyi Bigelow and Schroeder 1948,
2 BREVIORA NO. 24
Schroeder 1951, representing the little known family Anacantho-
batidae.
We can now report the capture by OREGON of another new Raja
from the northern part of the Gulf, and of a skate-like fish, from the
southern part, for which a new family seems needed.
Family PSPEUDORAJIDAE, Fam. Nov.
Family characters. Rajoidea without dorsal fins; with well developed
caudal fin extending around tip of tail and forward along lower side
of tail about as far as along upper side, supported by a great number of
very slender ray-like strands, apparently cartilaginous;! with outer-
posterior margins of pelvic fins nearly straight, or even slightly convex
if spread widely; anterior wall of spiracle with a transverse row of low
vertical ridges, representing the vestiges of the embryonic gill fila-
ments; pelvis with transverse element nearly straight, each of its outer
corners with a short projection directed forward.
The batoid fish, described below as Pseudoraja fischeri, falls clearly
among the Rajoidea because of the shape of its pelvis and of the
persistence of vestiges of the embryonic gill filaments on the anterior
wall of its spiracles (mentioned above). It is rajoid, too, in general
appearance; in the nature of its dermal armature; and in the fact that
its snout terminates in a fleshy process, for a corresponding structure
tips the snout in the rajoid family Anacanthobatidae, represented in
the Gulf of Mexico by the genus Springeria Bigelow and Schroeder
1951, and perhaps in the skate Psammobatis mira(Garman)1877? but is
not known to occur in any of the Myliobatoidea. The presence of
a rostral projection (albeit a short one) from the front of its cranium,
and its lack of a tail spine, point toward a rajoid rather than a mylio-
batoid relationship, though neither of these two features is strictly
alternative, between the two suborders, for a few rajoids lack the
rostral cartilage, while it is a matter of common knowledge that some
myliobatoids lack the tail spine.
On the other hand, a myliobatoid relationship, rather than a rajoid,
is suggested for Pseudoraja, superficially, by the shape of its pelvic
fins with straight or slightly convex outer contour, and by the large
size and the shape of its caudal fin. But, again, we are dealing with
1 The caudal fin-folds of various species of Raja, that we have examined under the microscope,
are supported similarly.
2 Garman (Proc. Boston Soc. Nat. Hist., Vol. 19, 1877, p. 207) described the snout of mira
ag terminating in an ‘‘acute point’’, which is pictured as prickly in his classic monograph
(Mem. Mus. Comp. Zool., Vol. 36, 1913, Pl. 27, Figs. 3-5).
1954 NEW FAMILY, GENUS AND SPECIES OF BATOID FISHES 3
characters that are not as strictly alternative as seemed once to be the
ease. Thus the pelvics of Pseudoraja are approached closely, in shape,
by those of Sympterygia (undoubtedly a rajid) in which they are so
weakly concave in outline, when spread, that they can hardly be
characterized as “‘bilobed”’.!. And while the caudal fin is a close counter-
part, in size and shape, of the caudals of the myliobatoid family
Urolophidae, it is similar in shape in the rajid genus Springeria
Bigelow and Schroeder 1951, though smaller.
With Sympterygia and Pseudoraja wholly bridging the gap, in shape
of pelvic fins, between the typical Rajoidea in which these are definitely
bilobed, and the Myliobatoidea, in all of which their outer contour is
continuously convex, it is evident that the precise shape of these fins
must be abandoned, as alternative between these two suborders. But
the shape of the pelvis itself is sharply diagnostic in this regard, for
while it is nearly straight transversely, or bowed forward only very
slightly, in all the Rajoidea for which its shape is known? and has a
projection (longer or shorter) directed forward from each of its outer
corners, its transverse element is bowed strongly forward, and it has
no prepelvic projections at the outer corners in all the myliobatoid
families where it has been studied, but has a prepelvic process in the
mid-line in some of them.’
The presence or absence of vestiges of the embryonic gill filaments
on the anterior wall of the spiracle, after birth, seems equally diag-
nostic, as between the suborders Rajoidea and Myliobatoidea, for
these are present in various species of Raja, in Breviraja, in Cruriraja,
in Springeria and in Pseudoraja, whereas no trace of them is to be seen
in any of the myliobatoids where we have sought them, which include
representatives of Dasyatis, Taeniura, Gymnura, Urolophus, Mylio-
batis, Aetobatus, Rhinoptera and Mobula.
The taxonomic significance of this character was forecast, in fact,
more than a century ago, by Johannes Mueller’s (Arch. Anat. Physiol.
Jahrg. 1841, p. 274) discovery that these vestigial gill folds are present
after birth in skates and in torpedos, as they are in many sharks,
but not in Myliobatis, in Dasyatis (referred to by him as Trygon), or
in Taeniura. But while the contrast in this respect, between skates
and sting rays, has been mentioned repeatedly since Mueller’s day,
1 For illustration of the pelvics in Sympterygia, see Garman, 1913, Pl. 27, Fig. 1.
2 This includes representatives of the genera Raja, Breviraja and Cruriraja (Family Rajidae),
and of Springeria representing the Anacanthobatidae. The shape of the pelvis is not known
for the somewhat problematical family Arhynchobatidae.
3 See Garman (1913, Pls. 53, 54) for illustrations of the pelvis in Dasyatidae, Potamotry-
gonidae, Gymnuridae, Urolophidae, Myliobatidae and Mobulidae.
4 BREVIORA NO. 24
we can not find that any of our predecessors have taken account of it
in defining the two subfamilies in question.
Pseudoraja, in short, falls clearly among the Rajoidea. But its lack
of dorsal fins with its well developed caudal fin forbid its reference
either to the Rajidae or to the somewhat problematical family
Arhynchobatidae, while the nature of its pelvic fins, the large size of
its caudal, and its well developed dermal armature set it apart, equally
sharply, from all known members of the Anacanthobatidae. Hence the
necessity for a new family, lest it be left a taxonomic orphan.
Genus PSEUDORAJA Gen. Nov.
Generic characters. Pseudorajidae with shape of disc, and of tail
relative to disc, as in skates of the family Rajidae; pelvic fins very
large, wing-like, with anterior outline directed outward, about trans-
verse to main axis of disc; front of cranium with a rostral projection,
longer or shorter; a deep pit on ventral surface of head on either side,
close posterior to nostril but entirely separate from latter. Other
characters those of family Pseudorajidae. Type species Pseudoraja
fischeri Bigelow and Schroeder.
PSEUDORAJA FISCHERI,! Sp. Nov.
Figures 1, 2
Type. Female, 479 mm. long, southern part of Gulf of Mexico, near
Campeche Bank, Lat. 22°42’N, Long. 86°41’W, 225 fathoms,
OREGON Sta. 726; bottom temp. 47.7°F; U. S. Nat. Mus. No.
163368.
Study material. Also female 428 mm.; female 262 mm., and juven-
ile male 262 mm. from same station.
Description. Proportional dimensions, in per cent of total length, of
female 479 mm. long (type) and female, 428 mm.
Disc: Extreme breadth, 48.6, 50.3; length 39.5, 40.0.
Snout length in front of:* orbits 9.2, 8.4; in front of mouth 12.3, 12.1.
Orbits: Horizontal diameter 4.4, 4.7; distance between 3.0, 3.0.
Spiracles: length 2.3, 2.2; distance between 6.0, 6.1.
1 Named in recognition of E. N. Fischer’s skillful portrayals of elasmobranchs.
2 From base of rostral filament.
1954 NEW FAMILY, GENUS AND SPECIES OF BATOID FISHES 5
P pPER DR FIR:
DO vp gai?
Fig. 1. Pseudoraja fischeri, type specimen 479 mm. long. Rostral filament
and side view of caudal fin to larger scale. U. S. Nat. Mus. No. 163368.
6 BREVIORA NO. 24
Mouth: breadth 4.7, 4.6.
Nostrils: distance between inner edges 6.4, 6.3.
Gill openings: lengths, Ist 1.0, 1.1; 3rd 1.2, 1.2; 5th 0.7, 0.7; distance
between inner ends, Ist 11.2, 11.8; 5th 6.9, 7.0.
Caudal fin: length of base, upper 7.7, 8.2; lower 6.9, 7.3.
Distance: from tip of snout to center of cloaca 39.5, 38.5; from center
of cloaca to tip of tail 60.5, 61.5.
Pelvics: anterior margin 17.7, 17.7.
28 30 28
Teeth: 26 in type; 6 in female of 428 mm.; 58 in male of 262 mm.
Disc, from base of rostral filament, about 1.2 times as wide as long,
so broadly rounded in front that its anterior contour is not susceptible
to angular measurement; the rostral filament about 14 as long as orbit,
soft, narrow triangular, tapering to very slender but blunted tip.
SaaS SS —ESE—E———
ee
SF
Fig. 2. Pseudoraja fischeri. Left, ventral view of oronasal region with right
hand nasal flap rolled back to show nostril and oronasal pit (0). x about 1.
Right, skin from central part of pectoral, to show arrangement of prickles.
x about 3 1/2.
Anterior margins of pectorals weakly concave abreast of eyes and
spiracles, convex thence rearward; outer corners broadly rounded;
posterior margins moderately convex; posterior corners rather ab-
ruptly rounded; inner margins weakly convex, to axils. Tail slender,
its dorsal surface nearly flat, but its sides and lower surface rather
broadly arcuate in cross section; its breadth at axils of pelvies about
80 per cent as great as length of orbit, its length, from center of cloaca
to tip, about 1.5 times as great as distance from center of cloaca to
base of rostral filament; the lateral caudal folds confined to about
posterior 14 of tail and extending a little beyond origin of caudal fin,
very narrow along their anterior part, but widening rearward until
1954 NEW FAMILY, GENUS AND SPECIES OF BATOID FISHES 7
nearing the level of the caudal fin. The breadth of the folds is about
Vs as great as the height of the caudal above its axis, or about 24 as
great as the length of the first gill openings.
Entire upper surface of dise including skin above eyes, and of tail
close set with minute, sharp pointed prickles curving rearward, except
close along extreme posterior edge of pectorals; also two large and one
small thorn around anterior contour of orbit, the small one the outer-
most; a group of two large and one smaller thorn close behind each
orbit, with the largest member of the group the outermost; two thorns
on each shoulder, with a median line of three from nuchal region to
pectoral girdle; these are followed, after a short gap, by a row of
about 32 along mid-line of dise and of anterior 24 of length of tail, the
first five to pelvic girdle small, the next 17 or 18 large and conspicuous,
with strongly striate bases, those farther rearward, along tail, pro-
gressively smaller; and the posterior 14 of tail without large thorns,
recalling conditions in Raja senta. Either side of mid-dorsal belt of
disc and upper side of tail with 3-4 irregular rows of closely crowded
thornlets; no thorns large or small (apart from the prickles) anywhere
on pectorals. Caudal fin generally prickly above caudal axis, but with
only a few scattered prickles below axis. Upper surface of pelvics
wholly naked. Lower surface of dise and of pelvics naked, also of
anterior part of tail rearward to abreast of rear corners of pelvics, but
densely prickly thence rearward to caudal fin. The dermal armature of
small specimens essentially similar, except the mid-dorsal thorns
fewer (about 26 in 2 specimens of 262 mm.).
Snout (from base of terminal filament) to front of orbits about
2.9 times as long as distance between orbits, its length in front of
mouth about 1.9 times as great as distance between inner edges of
exposed nostrils. Orbit about 1.5 times as long as distance between
orbits and about 2.1 times as long as spiracle. Upper eyelid bowed
downward, and eye with a rounded black velum with crenate margin
above pupil, as in skates of the genera Raja and Breviraja.
Outer lip of spiracle smooth; vestigial gill-ridges on anterior surface
of spiracle about 12 in number, rounded, their edges completely fringed
with minute lobelets. Nasal curtains smooth edged; those of the two
sides of head actually separated by a space about 14 as wide as
distance between exposed nostrils, but seemingly interconnected there
by a weakly outlined fold of skin. Outer (posterior) margin of nostril
smooth, slightly expanded in scoop-like form; the exposed nasal
aperture noticeably small.
8 BREVIORA NO. 24
An interesting feature of this new skate is that if the nasal curtain,
on either side, be rolled inward, and the skin between nostril and
corner of mouth be spread outward, a deep pit is exposed, close behind
the nostril, but its entrance separated from the latter by a bar of stiff
tissue. The pit is directed forward-upward, and it extends so far that
a probe inserted into it can be felt clearly from the dorsal side of the
head, through the overlying skin. And it is so voluminous on the two
larger specimens as to allow the entrance of an ordinary match stick,
or of a slender lead pencil. The presence of this pit was wholly
unexpected, for nothing comparable to it is to be seen, either in Raja,
in Dasyatis, or for that matter in any of the other batoids that we
have examined in this respect, and these include representatives of
all known families, both of Rajoidea and of Myliobatoidea, excepting
only the Arhynchobatidae.
Mouth nearly straight transversely in females and in juvenile males,
28 (type)—-30
26 (type)—28’
females and of juvenile males low, rounded, in quincunx mosaic; those
of mature males not seen; the tooth bands attached rather loosely
to the jaws.
First gill openings about 1 as long as distance between exposed
nostrils; third gills a little longer than first; fifth gills about 7/10.
Distance between inner ends of first gills about 1.8 times as long as
between inner edges of exposed nostrils and about 1.7 times between
inner ends of fifth gills.
No dorsal fins. Caudal fin a little longer than distance between
exposed nostrils, its height above caudal axis about 14 as great as its
length, its depth below axis about 4% as great as its height above
latter; upper and lower caudal origins about even. Upper margin
continuously rounded; lower margin less strongly so; tip broadly
rounded, notched abreast of tip of axis in type specimen, but merely
ragged there on slightly smaller specimen (428 mm.) and continuously
rounded terminally in small specimens (262 mm.), evidence that its
terminal contour on the type has resulted from mutilation.
Pelvic fins more widely spreading than in most other rajoids, and
of very diagnosticshape, the anterior outline directed nearly transversely
to the main axis of disc and tail, and so long that the tips of the pelvies
reach outward considerably beyond the margins of the overlying
portion of the pectorals. Anterior margin nearly straight; outer
corners broadly rounded; outer posterior (distal) margin straight, or
its contour not known for adult males. Teeth those of
1954 NEW FAMILY, GENUS AND SPECIES OF BATOID FISHES 9
very weakly convex if spread widely; only very weakly scalloped to
conform to positions of tips of radial cartilages; the rear corner
abruptly rounded. The inner margin is so short that the pelvics of
the two sides appear to be separated only by a shallow notch. But the
condition of the clasper described below shows that this is not evidence
of a partial fusion of the inner edges of the pelvics with the sides of the
tail, but only that these edges are relatively shorter than they are in
the skates of the genera Raja and Brewiraja. The anterior margin of
the fin soft and fleshy; the first radial cartilage noticeably stout,
especially toward its base; the other radialia slender and flexible.
Point of separation of clasper from inner margin of pectoral, in
juvenile male, about even with the axil of the fin; tip of clasper already
reaching a little beyond rear corner of pelvic. Claspers of mature male
not seen.
It is interesting, in this connection, that the claspers of juveniles
of Springeria originate considerably in advance of the apparent axils
of the fins, evidence of partial fusion of the inner margins of the latter
with the sides of the tail (Bigelow and Schroeder, 1951, Fig. 1).
Front of cranium with a low, blunt tipped rostral projection,
reaching forward a little beyond level of fronts of nasal capsules; tips
of anterior rays of pectorals reaching nearly to tip of snout. Pelvis
with transverse element nearly straight, each outer corner with a short
prepelvic projection.
Color. Upper surface ashy gray, deepening rearward to sooty gray
along posterior 4% of tail; disc rearward from orbits with many small
vaguely outlined pale spots, most conspicuous on head, and along mid-
belt thence rearward, less so outward over pectorals; outer parts of
pectorals unspotted. The large thorns white basally, their tips gray.
Anterior part of disc also with a rather conspicuous pattern of black
dots marking the mucous pores as follows: a) an irregular row along
outer margin of each pectoral from a little behind tip of snout to about
abreast of axis of greatest breadth; b) a row of 7-8 on each side,
diverging from tip of snout to a little in front of level of front of orbits;
c) a row of four, extending in anterior-posterior direction close in front
of each orbit; d) a larger black spot, marking a cluster of three or four
pores a little outward and rearward from the posterior edge of each
orbit; e) a similar black spot (3 or 4 pores) either side of the mid-line
in nuchal region, close in front of the first large mid-dorsal thorn;
f) one black dot (1 pore) on inner part of each pectoral, in shoulder
region; a second in line of orbit a little rearward from shoulder
10 BREVIORA NO. 24
region; and a third about midway thence toward axil of pectoral.
Tail irregularly pale spotted along anterior 14, with a few scattered
pale spots thence rearward to caudal fin; also with a vaguely outlined
dark cross-bar a little posterior to level of tips of pelvics, followed,
after a short gap, by a second such bar. Upper surface of pelvics ashy
gray, without pale spots.
Lower surface of disc ash gray with irregular sooty cloudings, these
most conspicuous on abdominal region; the pectorals of a more
brownish, the mid-belt of a more bluish cast. Lower surface of pelvics
ashy gray, with bluish cast; lower surface of tail pale grayish white
with irregular ashy-gray cloudings, and with the dark cross-bars of
the upper surface encroaching downward across the sides rather
conspicuously.
The pale spots on the disc are much less conspicuous, and less
definitely outlined on small specimens than on large, but the dark
cross-bars on the tail are more conspicuous.
Size. The larger specimens being female, and our only male a
juvenile, their sizes give no clue to the dimensions to which this skate
may grow.
Habits. Apparently confined to rather deep water.
Range. So far known only from the southern side of the Gulf of
Mexico, at the locality listed above, page 4.
Family RAJIDAE
RaJA FULIGINEA, Sp. Nov.
Figures 3, 4
Study material. Type specimen. A juvenile male, 306 mm. long,
OREGON Sta. 534, northwestern part of Gulf of Mexico, Lat. 27°32’/N,
Long. 93°02’W; trawl haul at 400-450 fathoms, April 11, 1952; U. S.
Nat. Mus., No. 163367. g
Distinctive characters. This skate resembles R. bathyphila Holt and
Byrne 1908 so closely in the general arrangement of thorns and
prickles, in proportional dimensions in general, and especially in the
very dark coloration of the lower surface of dise and tail that we took
it for a specimen of bathyphila on our first cursory glance. But a closer
examination showed that it differs from bathyphila in a considerably
more obtuse anterior contour of its dise (Fig. 8A) and especially in
the fact that the entire lower surface of the tail, apart from a very
narrow median stripe, is densely prickly from base to tip (naked in
1954 NEW FAMILY, GENUS AND SPECIES OF BATOID FISHES 11
bathyphila). The prickles, also, on the upper surface of dise and tail
are coarser than on bathyphila, and there are no naked areas, while
its pelvics are largely prickly on the upper surface (naked in bathyphila)
The only skate yet known from the northwestern Atlantic, or from
the Gulf of Mexico, other than R. bathyphila, with which fuliginea
shares a uniformly dark colored lower surface is R. olseni. But it
differs from olseni in blunter snout and tail prickly below. It
shares a tail prickly below with R. mollis. However, it is
marked off from mollis, not only by its dark lower surface (mollis is
pale yellowish or whitish below) but also by a much thornier tail and
dise, by its considerably more convex anterior contour, also by its
harder rostral cartilage. Since these divergences concern characters
that are not generally subject to much individual variation among the
members of its genus, a new specific name seems requisite for it. We
suggest fuliginea because of the sooty chocolate hue of its lower
surface.
Fig. 3. A. Outlines of front of disc; outer, Raja fuliginea, type specimen,
and inner, Raja bathyphila from southern slope of Georges Bank to show
difference in shape. B. Upper teeth of Raja fuliginea from near center of
mouth. x about 15.
R. fuliginea, like bathyphila and olseni, parallels R. badia Garman
1899, R. trachura Gilbert 1892, and R. abyssicola Gilbert 1895 of the
Pacific coast of Central and North America in its uniformly dark
lower surface. But it differs sharply from all three of these in various
respects.
Description. Proportional dimensions, in per cent of total length.
Juvenile male, 306 mm. long; OREGON Sta. 534, northwestern part
of Gulf of Mexico, Lat. 27°32’N; Long. 93°02’W; 400-450 fathoms.
Disc: Extreme breadth 46.3; length 42.8.
Snout length in front of: orbits 9.8; mouth 12.1.
Orbits: horizontal diameter 3.9; distance between 3.6.
1 BREVIORA NO. 24
a
AA
Fy
; ao: oe
Fig. 4. Raja fuliginea, type specimen 306 mm. long, U. 8S. Nat. Mus.
No. 163367, with right hand nostril and nasal curtain x about 3.4, and under
side of tail in advance of first dorsal fin, x about 1.1.
1954 NEW FAMILY, GENUS AND SPECIES OF BATOID FISHES 13
Spiracles: length 2.3; distance between 6.7.
Mouth: breadth 5.9.
Nostrils: distance between inner ends 6.2.
Gill openings: lengths Ist 1.3; 3rd 1.3; 5th 1.0; distance between
inner ends, Ist 12.9; 5th 7.5.
Distance: from tip of snout to center of cloaca 39.3; from center of
cloaca to Ist dorsal 45.4; to tip of tail 60.7.
First dorsal fin: vertical height 2.9; length of base 5.9.
Second dorsal fin: vertical height 2.6; length of base 5.6.
Interspace between: 1st dorsal and 2nd dorsal 0.0.
Pelwcs. anterior margin 14.4.
41
Teeth: 49°
Dise about 1.1 times as broad as long, so broadly and continuously
rounded in front that the anterior contour is not susceptible to angular
measurement; the tip of the snout projecting slightly, and blunted.
The pectoral margins only very slightly concave abreast of eyes and
spiracles, and broadly and continuously rounded thence rearward,
around to posterior corners, without definitely marked outer corners;
posterior corners rather abrupt; inner margins nearly straight.
Axis of greatest breadth about 24 (66%) of distance rearward from
tip of snout toward level of axils of pectorals.
Tail with very narrow lateral folds along posterior 24 (38-39%) of
its free length posterior to axils of pelvics; its length from center of
cloaca about 1.2 times as great to first dorsal and about 1.5 times as
great to tip as from center of cloaca to tip of snout.
Upper surface of disc, also upper surface and sides of tail, densely
set everywhere with rather coarse prickles curving rearward, or
minute thornlets, a very narrow band close along posterior edges of
pectorals being the only naked area.
Also a group of small recurved thorns along anterior part of rostral
ridge; four larger thorns in a line around inner side of each orbit; one
thorn in mid-line in nuchal region; six large thorns on scapular region,
two of these on either side with two in mid-line in pattern shown in
Figure 4; a line of six smaller thorns along mid-line of disc from
scapular region nearly to level of axils of pectorals, followed, thence
rearward, by 3+ irregular rows along anterior 4% of tail, succeeded by
two thorns in mid-line, to first dorsal fin. First and second dorsals,
and caudal membrane sparsely prickly; anterior lobes of pelvics
14 BREVIORA NO. 24
naked; posterior lobes rather densely prickly over inner and posterior
portions.
Lower surface of disc naked throughout. Lower surface of tail
densely set with prickles or thornlets similar to those of upper surface,
except that a very narrow median stripe is naked, both along anterior
24 of tail, and, again, rearward from level of origin of first dorsal fin.
Snout in front of orbits about 2/4 times as long as orbit; its length
in front of mouth about twice as great as distance between exposed
nostrils. Orbit about as long as distance between orbits and about 1.7
times as long as spiracle. Nasal curtain deeply fringed, with about 18
lobelets; expanded posterior (outer) margin of nostril fringed, also.
Mouth nearly straight, the lower jaw arched forward only a very little
centrally; its breadth about 1/5 (21%) as great as breadth of disc at
level of mouth, and only about 6 per cent as great as distance from tip
41
of snout to tip of tail. Teeth 49 with low triangular cusp, blunted at
tip, arranged in quincunx in juvenile male, probably also in female.
First pair of gill openings about 22 per cent as long as breadth of
mouth; distance between inner ends of first gills about 2.1 times as
long as between inner edges of exposed nostrils, and about 1.2 times
between inner ends of fifth gills.
Dorsal fins about alike in shape and equal in size, their bases con-
fluent, without intervening thorn or prickles. Caudal membrane,
posterior to second dorsal, about 24 as long as base of second dorsal.
Pelvic fins deeply concave outwardly; outer margin strongly
scalloped around the concavity with three marginal lobes on the one
fin, four on the other; but with the positions of the radial cartilages
only faintly indicated thence rearward; anterior lobe narrow, fleshy
with rounded tip; posterior lobe strongly convex; rear corner abrupt;
anterior margin of anterior lobe about 90 per cent as long as distance
from its point of origin to rear corner of posterior lobe; rear corners
extending back from about 14 of distance from level of axils of pectorals
toward first dorsal fin.
Firm rostral cartilage detectable by touch as reaching very nearly to
extreme tip of snout; tips of anterior rays of pectorals falling a little
short of level of tip of rostral cartilage.
Color. Upper surface of disc, tail and pectorals uniform dark ashy
gray, darkest on anterior lobes of pelvics, but without definite dark
markings anywhere. Lower surface of dise sooty chocolate to nearly
black, and noticeably darker than upper surface on head and around
1954 NEW FAMILY, GENUS AND SPECIES OF BATOID FISHES 15
outer belts of pectorals. A sub-triangular area in region of cloacareach-
ing forward about to pelvic girdle, and vaguely outlined, irregularly
interrupted areas on the inner parts of the pectorals rearward from the
gill region are of a somewhat paler sooty chocolate hue, perhaps partly
as a result of rough treatment in the trawl. Lower surfaces of pelvics
dark sooty, except pale on tips of anterior lobe; sides and lower surface
of tail of a very dark ashy-gray, except somewhat paler along a narrow
median stripe.
Size: The type (and only known) specimen being a juvenile male, it
gives no clue to the size to which this skate may grow.
Habits and range. The depth of capture (400-450 fathoms), added
to the dark color of its lower surface, marks fuliginea as a deep water
species. Present indications are that the upper boundary to its usual
range lies not far from 400 fathoms, else specimens of a skate made so
conspicuous by its dark lower surface would almost certainly have
been noticed among the catches of the many trawl hauls that have
been made in the Gulf at lesser depths. Nothing more than this is
known of its habits.
The locality of capture lies in the northwestern part of the Gulf,
some 100 miles off Galveston. It is an interesting question for the
future whether fuliginea is restricted to the Gulf, or whether it has
simply been overlooked in the open Atlantic.
BIBLIOGRAPHY
BrcELow, Henry B., and WILLIAM C. SCHROEDER
1948. New genera and species of batoid fishes. Sears Found., Jour. Mar.
Res., Vol. 7, No. 3, pp. 548-566.
1950. New and little known cartilaginous fishes from the Atlantic.
Bull. Mus. Comp. Zool., Vol. 103, pp. 385-408, 7 pls.
1951. A new genus and species of anacanthobatid skate from the Gulf
of Mexico. Jour. Washington Acad. Sci., Vol. 41, No. 3, pp. 110-
113.
195la. Three new skates and a new chimaerid fish from the Gulf of
Mexico. Jour. Washington Acad. Sci., Vol. 41, No. 12, pp. 3883-392.
CHANDLER, Asa C.
1921. A new species of ray from the Texas coast ... Proc. U. 8. Nat.
Mus., Vol. 59, pp. 657-658.
16 BREVIORA No. 24
GARMAN, SAMUEL
1877. On the pelvis and external sexual organs of selachians ... Proc.
Boston Soc. Nat. Hist., Vol. 19, pp. 197-215.
1881. Report on the selachians. Bull. Mus. Comp. Zool., Vol. 8, No. 11,
pp. 231-237.
1899. The fishes... Mem. Mus. Comp. Zool., Vol. 24, 431 pp. Atlas,
pls. 1-85, A-N.
1913. The plagiostomia. Mem. Mus. Comp. Zool., Vol. 36, xiii, 515 pp.
Atlas, 75 pls.
GILBERT, C. H.
1892. Descriptions of thirty-four new species of fishes... Proc. U.S.
Nat. Mus., Vol. 14, pp. 539-566.
1895. The ichthyological collections of the steamer “Albatross” .
Rept. U. 8. Comm. Fish. (1893), pp. 393-476, 16 pls.
Hott, E. W. L., and L. W. Byrne
1908. Second report on the fishes of the Irish Atlantic Slope. Sci. Invest.
Fisheries Ireland (1906), No. 5, pp. 1-26, 4 pls.
MUELLER, JOHANNES
1841. Fortgesetzte Untersuchungen iiber die Pseudobranchien. Arch.
Anat. Physiol., Jahrg. 1841, pp. 263-277.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. JANUARY 28, 1954 NuMBER 25
A NEW MIOCENE SPECIES OF PELUSIOS
AND THE EVOLUTION OF THAT "GENUS
By ERNEstT WILLIAMS
Among the reptilian remains from the island of Rusinga in Lake
Victoria, Kenya Colony, sent for determination to the British Museum
(Natural History) are the greater part of the carapace and a smaller
part of the plastron of an apparently new species of Pelusios.
Dr. W. E. Swinton, who suggested that I examine the unidentified
chelonian remains from British East Africa, has kindly consented that
I describe the new form. Accordingly I name it:
PELUSIOS RUSINGAE, new species
Type: Coryndon Museum Ru F3617—a_ partial carapace and
plastron.
Horizon: Miocene of Rusinga Island, Lake Victoria, Kenya Colony.!
Diagnosis: A Pelusios belonging to the adansonii-gabonensis section
of the genus, distinguished by the following combination of characters:
a very depressed shell (height included in length about four times);
the carapace expanded posteriorly; the vertebral region very shallowly
excavated, quite without keel; first vertebral scute much larger than
vertebral 2 and wider than long; vertebrals 2, 3, and 4 slightly longer
than wide; mesoplastra extremely narrowed medially, barely meeting.
The living species of Pelusios fall into two sections:
One, which is northern and western in distribution, comprises two
species, P. adansonii and P. gabonensis. This group is characterized
by having the anterior lobe of the plastron relatively long and the
abdominal scutes relatively short, so that the sulcus between the
abdominal scutes is included more than twice in the length of the
anterior lobe. Also the mesoplastra are more or less tapered medially,
so that the hyoplastra anteriorly and the hypoplastra posteriorly (or
the hypoplastra only) are longer medially than laterally, projecting
' For asummary of the geology and the Miocene fauna of Rusinga and adjacent areas see
Kent (1944).
2 BREVIORA NO. 25
into and filling up the interval left by the tapered margins of the
mesoplastra.
The other group within the living members of the genus is less
restricted in distribution. One of its species — P. subniger — overlaps
most of the range of the first group and in fact extends beyond that
range on the west to the Cape Verde Islands. On the east this same
species extends to Zanzibar, the Seychelles, Mauritius and Madagascar.
On the north, however, this species does not extend beyond British
{ast Africa into the Sudan range of P. adansonii.
The group typified by P. subniger is distinguished by having the
anterior lobe shorter and the abdominal scutes longer so that the
sulcus between the abdominal scutes is included less than twice in the
length of the anterior lobe, and by having the mesoplastra not tapered
and presenting straight transverse contacts with both hyo- and
hypoplastra.
The relationships of P. rusingae are clearly with the first of these
two living groups: the tapered mesoplastra clearly indicate this
position. From P. adansonii, however, P. rusingae differs (1) in the
more depressed shell, (2) in the absence of any vertebral keel, (3) the
first vertebral wider than long, (4) greater size. From P. gabonensis
it differs in (1) the posterior expansion of the shell, (2) the absence of
any trace of vertebral keel, (3) the second to fourth vertebrals longer
than wide. From both species it differs in the more extreme medial
narrowing of the mesoplastra. The table below summarizes the shell
characters of the two Recent and the fossil species (I utilize the data
of Loveridge, 1941, which I have, however, verified on other material).
P. adansonii
Sulcus between humerals
3-4 times as long as
that between pectorals.
Mesoplastra tapered
medially only poste-
riorly, thus a
transverse hinge with
the hyoplastra but an
oblique suture with the
hypoplastra.
A keel on the anterior
four vertebrals
throughout life.
P.. gabonensis
Sulcus between humerals
114-2 times as long as
that between pectorals.
Mesoplastra tapered
medially both anteriorly
and posteriorly, thus an
oblique suture with both
hyo- and hypoplastra.
A nodose keei in the
voung, lost in the
old.
P. rusingae
Unknown
Mesoplastra strongly
tapered medially
anteriorly and
posteriorly,
hardly meeting.
No keel, the
vertebral region
somewhat
depressed.
1954
Vertebrals about as long
as broad in adults.
Height in length
about 2.6 times.
Shell distinctly
broadened posteriorly.
Known maximum size:
185 mm.
EVOLUTION OF PELUSIOS
At least vertebrals |
to 3 broader than long
in adults.
Height in length
2.3 to 3.8 times.
Shell not broadened
posteriorly.
Kktnown maximum size:
259 mm.
Vertebral 1 wider
than long,
vertebrals 2-4
longer than wide.
Height in length
about 4 times.
Shell distinctly
broadened
posteriorly.
Estimated size:
245 mm.
P. rusingac thus contrives to combine some of the characters of both
the two living members of its group. It occurs, also, outside — south
and east — though not far outside, the present limits of its group.
Only P. subniger of the alternative group is known from Lake Victoria
today.
Three fossil species of Pelusios have been previously described:
P. rudolphi Arambourg from the Lower Pleistocene of Omo, founded
on a partial plastron and carapace (type in Paris Museum); P.
dewitzianus v. Reinach represented by fragments from the Middle
Pliocene of Wadi Natrun (type formerly in Munich, now destroyed);
and P. blanckenhorni Dacqué, a skull from the Lower Miocene of
Moghara (type in Berlin?). In addition and not previously recorded
there are abundant fragments (Nairobi Museum) and a complete shell
(British Museum No. R 5761) of P. sinuatus (a still living species of
the P. subniger group) from Bed I, Pleistocene of Olduvai.
The fossil P. stnuatus need not be compared with P. rusingae.
The Olduvai material is clearly referrable to the Recent species which
still occurs in this area.
P. rudolphi needs as little attention. The type (examined at the
Paris Museum) resembles closely old specimens of Recent P. sinuatus.
It may provisionally be accepted as ancestral to the P. sinuatus of the
later Pleistocene and of the Recent.
P. dewitzianus was originally described on quite inadequate material
which, however, was still sufficient to place it as a member of the
P. subniger group. It was redescribed from much better material by
Daequé (1912), who at the same time discovered that a supposed
Phocene species of Pelomedusa (P. pliocenica v. Reinach) was a
synonym of this species. P. dewitzianus as a member of the alternative
4 BREVIORA NO. 25
group requires no comparison with P. rusingace.
There remains, however, P. blanckenhorni, which is from a deposit
apparently equivalent in age and very similar in fauna to that of the
Lower Miocene of Rusinga Island, but 2000 miles distant. P. blancken-
horni and P. rusingae cannot be compared, since one is based on a
skull, the other on a shell. The skull of P. blanckenhorni, inadequately
described and figured only in dorsal view by Dacqué, seems similar to
that of P. gabonensis. It may, therefore, belong to the same group
within the genus as P. rusingac, and it is not impossible that the latter
is asynonym. But to hazard the identity of forms 2000 miles distant
from one another and represented by incomparable parts would be
without substantial basis.
Furthermore, a special element of doubt attaches to species be-
longing to this section of the Pelomedusidae which, as with P.
blanckenhorni, are founded solely upon the skull. It is a remarkable
fact that Pelusios and Pelomedusa, though quite distinct in shell
characters, have extremely similar skulls. The skulls of the two Recent
genera can be told apart only by characters which in many other
groups would be counted of specific value only. Reference of a fossil
skull, therefore, to either genus is a doubtful procedure unless there
is the confirmation of an associated shell. In the present case this
leaves us with the possibilities that P. blanckenhorni may be either
specifically identical with P. rusingae, or specifically different, or it
may belong to a different genus. This conclusion may appear as
absurd as it is unsatisfactory, but this is a dilemma not uncommon
in paleontology, and it is decidedly worthwhile to recognize and
emphasize the difficulty of evaluation of fossil species based on parts
not comparable. All that can be suggested as a method of decision,
which, while arbitrary, is still not devoid of reasonableness, is that
material from deposits of the same or equivalent ages and geographi-
cally close may be provisionally associated if any apparently valid
grounds for such association exist; but geographic distance or difference
in geologic age carry with them a presumption of distinctness which
must be countered by stronger arguments than those that — for the
moment — suffice in the other case.
P. rusingae and P. blanckenhorni (Qf this is really a Pelusios),
occurring in the Lower Miocene, are the oldest members of this genus.
It will be useful to consider them against the background of the early
history of the family of which they are a part.
The family Pelomedusidae is certainly very old; it probably stems
1954 EVOLUTION OF PELUSIOS 5
ultimately from the pleurosternids of the Upper Jurassic and the
Cretaceous. Unfortunately the form which has been suggested as the
oldest representative of the family, Platycheloides nyassae Haughton,
is incompletely known and doubtful as to age. It has small laterally
placed mesoplastra and is therefore not an obviously primitive form.
Mesoplastra meeting in the midline are certainly primitive for turtles
and Pelusios would therefore be more primitive in this respect, unless
the larger mesoplastra of P.lusios are a secondary development (see
below). The single known character in which Platycheloides differs
from Pelomedusa as ordinarily conceived — the absence of the median
plastral fontanelle — does not in fact separate it from that genus,
since, as I have been able to determine on British Museum specimens
from Uganda and the Sudan, the median fontanelle is sometimes
lacking in even small specimens of Pelomedusa. The beds from which
Platycheloides nyassae derives are Cretaceous in age, but to what part
of the Cretaceous they belong is not known. This African form is
therefore not certainly older than the better known pelomedusids of
the Upper Cretaceous of North and South America and Europe,
though it is probably as old. Widespread already in the Cretaceous,
the pelomedusids continued so in the early Tertiary with represent-
atives in North and South America, England, Italy (del Zigno, 1887),
Ygypt, Congo and India.
It is a curious fact that every one of these older members of the
family that are sufficiently known is pelomedusine in type rather
than pelusiine, that is: the mesoplastra are small and lateral elements,
as in Pelomedusa and Platycheloides, not large elements meeting in the
center of the plastron as in Pelusios and the pleurosternids. Nor is it
at all likely that this observation 1s an artifact resulting from a failure
to recognize as pelomedusids those with complete mesoplastra. A
pelomedusid with large complete mesoplastra is immediately dis-
tinguishable from a pleurosternid by the total absence of inframarginal
scutes.
The uniformity in the condition of the mesoplastra in the oldest
members of the family is an intimation that the pelomedusine type of
mesoplastra (small and lateral) may be primitive for the family and
that the pelusiine type (large and centrally meeting) may be second-
arily derived from the pelomedusine.
With this suggestion the known facts about Pelusios are fully
congruent. The members of the genus Pe/us‘os form a structural series
in regard to the size of the mesoplastra, P. rusingae having the most
6 BREVIORA NO. 25
reduced mesoplastra, P. gabonensis the next, P. adansonii next, and
the P. subniger group the most fully developed mesoplastra. A
structural series is always ambiguous unless the time dimension can
be added, but P. rusingae as the oldest shell belonging to the genus
seems to provide this time element.
The similarity of the skulls of Pelomedusa and Pelusios further
suggests relationship, and the existence of a species of Pelomedusa
(P. progaleata v. Reinach) anterior in time (Lower Oligocene) to the
earliest (Lower Miocene) Pelusios further supports the view that
Pelusios is a relatively late and specialized genus directly derived
from Pelomedusa.
P. rusingae is thus a fortunate discovery, offering a much needed
term in an evolutionary series — a series apparently affording an
example of the reversal of an evolutionary trend.
Acknowledgements: I am indebted to Dr. W. E. Swinton for the
opportunity of examining and describing this fossil, to Prof. C.
Arambourg for the privilege of examining the type of P. rudolphi at
the Paris Museum, to Dr. H. W. Parker for permitting me to examine
comparative material of the Recent genus in the Reptile Section,
British Museum (Natural History), and to M. Jean Guibé for similar
permission in Paris. The photographs in Plates 1 to 4 were made by
Peter Green and are reproduced by permission of the British Museum
(Natural History). This study is part of a series of researches made
possible by the grant of a Guggenheim Fellowship during the year
1952-53.
REFERENCES
ARAMBOURG, C.
1948. Contribution a l'étude géologique et paléontologique du bassin
du Lac Rodolphe et de la basse valée de |’?Omo. 2me Partie.
Paléontologie in Mission Scientifique de l’?Omo. 1932-1933.
Vol. 1, fase. 3, pp. 231-559.
Dacauk, E.
1912. Die fossilen Schildkréten Aegyptens. Geol. Paleont. Abhandl.,
vol. 14, pp. 273-333.
Havueuton, S. H.
1928. On some reptilian remains from the dinosaur beds of Nvassaland.
Trans. Roy. Soc. 8. Africa, vol. 16, pp. 67-75.
Kent, P. B.
1944. The Miocene beds of Kavirondo, Kenya. Quart. Jour. Geol. Soe.
London, vol. 100, pp. 85-116.
1954 EVOLUTION OF PELUSIOS
LOVERIDGE, A.
1941. Revision of the African terrapin of the family Pelomedusidae.
Bull. Mus. Comp. Zool., vol. 88, pp. 467-524.
Rernacu, A. VON
1903. Schildkrétenreste aus dem = aegyptischen Tertiir. Abhandl.
Senckenberg. Naturf. Ges., vol. 29, pp. 1-64.
ZIGNo, A. DEL
1887a. Chelonii scoperti nei terreni cenozoici delle prealpe Veneti. Mem.
R. Inst. Veneto, vol. 23, pp. 119-129.
1887b. Chelonio scoperto nel calcare nummulitico de avesa pressa Verona.
Ibid., vol. 23, pp. 185-145.
PLATE 1
Pelusios rusingae, dorsal view of type shell.
PLATE 2
Pelusios rusingae, ventral view of type shell.
Pelusios sinuatus (B.M.N.H.
Pleistocene of Olduvai.
PLATE 3
R. 5761), dorsal view of shell from Bed 1,
PLATE 4
Pelusios sinwatus (B.M.N.H. R. 5761), ventral view of shell from Bed 1,
Pleistocene of Olduvai.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. Fresruary 3, 1954 NuMBER 26
A PRELIMINARY LIST OF THE
EARTHWORMS OF NORTHERN NEW JERSEY
WITH NOTES
By H. Davirs
Dover, New Jersey
The literature of North American earthworms contains few
references to New Jersey and no paper has appeared describing worms
collected in that state. Moore (1895, p. 473) refers to three species
found within 30 miles of Philadelphia (Allurus tetraedrus, Bimastos
palustris and Sparganophilus tamesis), and Eaton (1942) mentions four
lumbricids found at Alpine, N. J. and Edgewater, N. J. This paper
is therefore presented as a contribution to the knowledge of the fauna
of New Jersey.
Collections of earthworms were made above a line drawn east and
west through Princeton, N. J. with a concentration in the area of
Morristown and Dover. Notes are given on the living conditions of
the various species where such are thought to add to our knowledge.
The term ‘clitellate’ is used in this note to describe worms with
recognizable features of clitellar development regardless of the stage.
Where the tubercula pubertatis only are present, ‘aclitellate’ is used,
while worms having no indications of clitellum or tubercula are con-
sidered ‘juveniles’.
Family LUMBRICIDAE
Genus ALLOLOBOPHORA Eisen 1874
ALLOLOBOPHORA ARNOLDI Gates 1952
Morristown, Mt. Kemble Road, in soil by stream; April 15, 1950,
2 clitellate specimens.
Andover, in rich loam in woods, April 12, 1953, 4 clitellate speci-
mens, many juveniles.
2 BREVIORA NO. 26
Dover, in garden soil, April and May 1953, 3 clitellate specimens,
many juveniles.
Morristown, James St., garden, May 19, 1953, 13 clitellate and
1 aclitellate specimens.
Mt. Freedom, in sandy ditch, May 19, 1953, 5 clitellate specimens.
Beatystown, in meadow, May 1953, 7 clitellate specimens.
Mt. Tabor, under stones in garden, June 8, 1953, 3 clitellate and
11 aclitellate (post sexual?) specimens.
This is the third record of a species which has hitherto been found
in Massachusetts and New Hampshire. The specimens correspond to
the diagnosis given by Gates except that the number of segments
covers a slightly wider range as the data from 21 specimens show:
139(2), 140(1), 141(2), 148(1), 1471), 153(1), 154(1), 160(1), 164(1),
174(1), 177(2), 181(1), 182(2), 183(1), 184(2), 187(1). In each of these
the anal segments were normal in appearance; however amputation is
prevalent in this species and some might prove to be amputees.
ALLOLOBOPHORA CALIGINOSA (Savigny) 1826
Dover, in garden soil (clay), June 8, 1953, 2 clitellate specimens.
Mt. Tabor, under stones in garden, June 10, 1953, 2 clitellate
specimens.
Pompton Plains, in marshy meadow, June 1953, 3 clitellate
specimens.
Paterson, in garden top soil, June 1953, 2 clitellate specimens.
These specimens are tentatively assigned to caliginosa, but do not
conform to the description given by Cernosvitov and Evans (1947,
p. 13), as Table I illustrates. In every case a grey-brown pigment is
present on the dorsal surface, the clitellum being a dull yellow.
Description of tubercula pubertatis:
Specimen No. 150a__band-like, indentation at xxxii on upper side.
150b band-like, indentation at xxxii on upper side.
157a_ band-like, tripartite origin, on xxxi-xxxill. Intersegmental
lines distinct.
157b L. As 157a.
R. Band-like, slightly indented at 31/32 and 32/33.
l6la
161b
162a
162b
Band-like, tripartite origin, on xxxi-xxxill.
OO
1954 EARTHWORMS OF NEW JERSEY
In each case the lateral portion of the tuberculum is translucent
while the median portion is opaque and appears as a band.
Table I
External characteristics of Allolobophora caliginosa ~
Number
Specimen of Tubercula Genital
No. Locality segments Clitellum pubertatis tumescences
150a Dover 133 27-335 31-33 310) oY ahs) av!
150b Dover 179 27-335 330-33 30 32 33 34
157a Mt. Tabor 167 27- 34 31-33 30 32 33 34
157b Mt. Tabor 156 27- 34 31-33 30 31 32 33 34
l6la Pompton
Plains 161 27- 34 31-33 30 32 33 34
161b Pompton
Plains 156 27— 34 31-33 33 34(R)
162a Paterson 159 27-— 34 31-33 30 32 33
162b Paterson 130 27— 34 31-33 BOL S2NonO4:
(amputee)
ALLOLOBOPHORA LIMICOLA Michaelsen 1890
Morristown, Glen Alpine Rd., in thick mud (pH 5.5) near branch
of Primrose Brook, May 27, 1952, 25 clitellate specimens, April
26, 1953, 8 clitellate specimens, June 6, 1953, 5 clitellate speci-
mens, many juveniles at each visit.
Dover, 2nd Street, marshy ground, June 8, 1953, 6 clitellate
specimens.
Beatystown, wet meadow, June 10, 1953, | clitellate specimen.
The Morristown location is undisturbed marsh and remains wet
throughout the year except during the winter freeze. Castings were
noted under logs and debris but were not apparent on the surface.
Both the Dover and the Beatystown locations are unimproved and
are saturated throughout the year. On the June 6 visit to the Norris-
town location, worms were observed in copula approximately 4 inches
underground.
This is the second record of this species in North America, Gates
(19538, p. 518) having found it in the Arnold Arboretum in Boston.
The specimens conform to the description given by Gates, the number
of segments corresponding very closely and illustrating the narrow
range of this species. Number of segments in 17 specimens: 104(2),
111(1), 112(1), 113(1), 114(3), 117(2), 121(2), 122(1), 123(2), 124(1),
129(1).
BREVIORA NO. 26
ALLOLOBOPHORA LONGA Ude 1885
Dover, sandy soil in garden, Nov. 1951, 3 clitellate specimens.
Morristown, in garden soil (clay), May 10, 1953, 1 clitellate
specimen.
Dover, in garden on 2nd St., June 8, 1953, 3 clitellate specimens.
Beatystown, damp soil in meadow, June 9, 1953, 1 clitellate
specimen, | aclitellate (post sexual?) specimen.
Genus BIMASTOS Moore 1893
BIMASTOS PALUSTRIS Moore 1895
Flanders, in wet moss in rocky stream near Route 31, Nov. 1951,
4 clitellate specimens.
Hardwick, in ditch in forest (running water), April 1952, 3
clitellate specimens.
Ironia, in wet moss by stream, Succasunna-Ironia Road, May
3, 1952, 1 clitellate specimen.
Swartswood, under moss on log by stream, May 29, 1952, 15
clitellate specimens in association with FL. tetraedra.
Mt. Freedom, in wet moss on marshy ground by stream, April
27, 1952, 7 clitellate and 1 juvenile specimens.
Shongum, Raynor Road, in very wet sandy loam by stream,
March 29, 1953, 4 clitellate specimens.
This species seems to have an affinity for running water and is
apparently more restricted in this respect than Evsenzella tetraedra
with which it is sometimes found. The above confirms Smith (1917,
p. 169) who states that B. palustris is found in the wet earth of rivers
and ponds and has been collected from New Jersey.
Spermatophores were noted on many specimens, the majority having
one pair and two specimens having two pairs. One specimen from
Swartswood has 53 segments, and one from Mt. Freedom 52 segments,
and in each case there was no indication of amputation, the anal
segment being normal in appearance. These numbers are smaller than
the 80 to 100 given by Smith (1917).
Genus DENDROBAENA Eisen 1874
DENDROBAENA MAMMALIS (Savigny) 1826
Morristown, Mt. Kemble Road, in moist soil by edge of stream,
May 1, 1951, 4 clitellate specimens.
1954 EARTHWORMS OF NEW JERSEY 5
This is the first record for this continent of this species. It is con-
sidered to be endemic in the British Isles and has also been collected
from a few localities in France. Its occurrence in North America is
not unexpected since it has been intercepted on plant materials im-
ported into this country (Gates, 1953, p. 530). The specimens con-
formed externally to the description given by Cernosvitov and Evans
(1947, p. 20).
DENDROBAENA OCTAEDRA (Savigny) 1826
Morristown, Glen Alpine Road, in rotting wood, June 30, 1952,
17 clitellate specimens in association with D. rubida.
Little is known of the distribution of this species in North America
although it has been collected in Massachusetts and Michigan in
addition to Newfoundland and Greenland.
DENDROBAENA RUBIDA (Savigny) 1826
Dover, under rotting grass cuttings in garden, Nov. 10, 1951,
2 clitellate specimens.
Ironia, in damp moss by Succasunna-Ironia Road, May 3, 1952,
2 clitellate specimens.
Morristown, Glen Alpine Rd., in rotting wood, June 30, 1952, 8
clitellate specimens and 1 juvenile with D. octaedra.
The records of this worm in North America refer only to collections
from New Hampshire, Massachusetts and Maine. However as the
writer has also collected it in Michigan, it may be widely distributed.
DENDROBAENA SUBRUBICUNDA (Eisen) 1874
This species is recorded by Eaton (1942) as occurring at Alpine,
N. J. It was not found in recent collecting.
Genus HISENIA Malm 1877
EISENIA FOETIDA (Savigny) 1826
This species is found plentifully in farm manure heaps in many
North Jersey localities and is sold for bait in the area. It was found
in every manure heap where a search was made and only one specimen
was collected in any other habitat. However, the current farming
practice of spreading manure daily instead of accumulating it, is
probably reducing the numbers of E. foetida which cannot be said to
be as “abundant” as Smith (1917, p. 165) implied.
6 BREVIORA NO. 26
EISENIA LONNBERGI Michaelsen 1894
Morristown, Mt. Kemble Road, in mud at bottom of stream,
April 1950, 2 clitellate specimens.
Stokes State Forest, Naponock Brook, in wet moss, April 12, 1952,
1 juvenile.
Ironia, under moss and in wet earth by stream, May 3, 1952,
2 clitellate and 4 juvenile specimens.
Shongum Lake, in wet soil by stream, May 1952, | clitellate and
4 juvenile specimens.
Dover, Millbrook Valley, in swampy ground, Aug. 1952, 5 clitel-
late and 4 juvenile specimens.
Great Swamp, Madison, in wet moss and debris by water (pH 6.0),
April 26, 1953, 2 juveniles.
In addition to the above, 4 clitellate and 9 juvenile specimens
were obtained from the Pennsylvania bank of the Delaware above
Montague, N. J. Some of these were completely submerged under
water in gravelly mud.
Table II
External characteristics of Eisenia lonnbergi
Number
of Tubercula Regener-
Condition segments Clitellum — pubertatis ation Remarks
aclitellate 77 — 26-28 56/57
clitellate 62 24-30 26-329 — Amputee
clitellate 110 323—30* 26-28 = Amputee
aclitellate 129 — 26, 27, 28 —
juvenile 76 = == —
juvenile 116 = == —
immature 48 = = 7/8 Apparent head
regenerate
clitellate 44 324-30 26-28 — Amputee
clitellate 79 ?24—30* 26, 27, 28 = Amputee
clitellate 51 24-30* 26-729 = Amputee
clitellate 142 224-30* 26-229 —
juvenile 133 == soe 101/102
juvenile 132 — — 47/48
juvenile 74 — — — Amputee
juvenile 118 == = —
juvenile 132 =a a —
juvenile 135 <= = =
juvenile 104 = = —
clitellate 125 24-30* 26-329 —
*Clitellum feebly developed.
eh eee in
1954 EARTHWORMS OF NEW JERSEY 7
There are few references to this worm and little has been published
regarding its habitat. It has been found in Georgia, Virginia, North
Carolina, Connecticut and Massachusetts. Evidently it is widely
distributed in northern New Jersey and is probably endemic. Its
habitat appears to be restricted to very wet locations in or along the
banks of rivers or streams.
The range in segment number of normal specimens from 125 to 142,
extends the previous record of 138 segments (Smith 1917, p. 164).
The incidence of amputation is very high in the specimens collected,
whether this is due to predatism or to autotomy is not known, but
none of the specimens autotomized during handling in collection or
preservation.
EISENIA ROSEA (Savigny) 1826
Stokes State Forest, in soil, May 30, 1950, 2 clitellate specimens.
Dover, in clay soil in garden, Nov. 17, 1951, 1 clitellate specimen.
Morristown, in marshy soil, (pH approximately 5.5), June 30,
1952, 1 clitellate specimen.
Swartswood, in mud at edge of stream, May 29, 1952, 1 clitellate
specimen.
Layton, in muddy ditch by road, April 3, 1953, 1 clitellate
specimen.
Andover, woods near Lackawanna R.R. (Cutoff), in rich black
loam, April 12, 1953, 2 clitellate specimens.
Table III
External characteristics of Eisenia rosea
Number Tumescences Tumescences
Specimen of Tubercula including including
No. segments -Clitellum pubertatis ab cd
44a, 112 25-32 29-331 26-31 12(L), 138(R)
44b 114 25-32 29-131 26-31 11, 12(L), 12(R)
61 109 25-31 29-— 30 26-31 als
55 128 26-32 29- 31 26-32 12(R)
122 p3l 26-32 29- 31 25-32 None
120 130 296-32." 30-930) V12() in oc) None
26-32
Further description of tubercula pubertatis:
No. 44a Tubercula bluntly elliptical from 28/29 to 431 without
marginal incisions. Intersected by furrow 29/30 only.
ied)
BREVIORA NO. 26
44b Shape as 44a but extending to 431. Slight indication of
intersegmental furrow 29/30 on L. side.
61 Tubercula elliptical, from 29/30 to 30/31 with incisions at
29/30 on both margins and distinct furrow across tubercula
at 29/30.
55 Tubercula somewhat ill defined ellipse from 28/29 to 31/32
intersected by distinct furrows at 29/30 and 30/31.
122 Single tuberculum on 29 and elliptical mass from 29/30 to
31/32. Furrow 29/30 very distinct. Slight incision on
median margin at 30/31 with trace of furrow.
120. ~=Tubercula elliptical from 29/30 to 31/32 with incisions in
both margins at 30/31. Indistinct furrow at 30/31.
The presence of tumescences in the clitellar region would appear to
identify these specimens as var. macedonica Rosa 1893, which is said
to be characterized ‘‘by the presence of small, mostly lightly coloured
glandular papillae along the border of the clitellum”’ (Cernosvitov and
Evans 1947, p. 23). However, this variety is ill defined in comparison
with the typical form and positive identification cannot be given.
Genus HISENIELLA Michaelsen 1900
EISENIELLA TETRAEDRA (Savigny) 1826 forma typica
Morristown, at roots of grass in stream, April 15, 1950, 1 clitellate
specimen.
Stokes State Forest, in mud under stone in stream, June 10, 1951,
1 clitellate specimen. In wet moss, Naponock Brook, April
12, 1952, 3 clitellate specimens.
Swartswood, under moss in stream, May 29, 1952, 2 clitellate
specimens in association with B. palustris.
Dover, in marsh near Orchard St. Cemetery, June 20, 1952, 18
clitellate specimens, many juveniles.
Great Swamp, Madison, under wet leaves, May 1952, 8 clitellate
specimens.
Buttzville, under wet leaves at edge of Pequest River, April
16, 1952, 8 clitellate and 3 juvenile specimens.
Morristown, Glen Alpine Road, in wet gravel at brook, March
and April 1953, 10 clitellate specimens.
One specimen collected at Morristown has the male pores on
segment ix with the clitellum correspondingly forward, apparently due
to hypomeric regeneration.
1954 EARTHWORMS OF NEW JERSEY 9
On many specimens, setae ab of segment xxii are genital, occasionally
on prominent lightly colored tumescences.
Genus LUMBRICUS Linnaeus 1758
LUMBRICUS RUBELLUS Hoffmeister 1843
Stokes State Forest, Big Flat Brook, under leaves, May 30, 1950,
1 clitellate specimen.
LUMBRICUS TERRESTRIS Linnaeus 1758
Throughout. that part of New Jersey with which we are concerned,
this worm is very common in lawns and gardens. It is also found in
meadows, particularly where the ground is marshy, and in ditches. It
is much in demand by local fishermen and is sold for bait in many
places.
Genus OCTOLASIUM Oerley 1885 —
OcTOLASIUM LACTEUM (Oerley) 1881
Buttzville, under rotting leaves at edge of Pequest River, Aug.
16, 1952, 1 clitellate specimen.
One clitellate specimen was also collected in May 1950 from gravelly
mud under water level from the Pennsylvania bank of the Delaware
above Montague, N. J. Both specimens have tumescences on ab of
Xxll (one specimen also on xxi, right).
Family GLOSSOSCOLECIDAE
Subfamily SPARGANOPHILINAE
Genus SPARGANOPHILUS Benham 1892
SPARGANOPHILUS EISENI Smith 1895
Morristown, Glen Alpine Rd., in mud under water (pH 5.5) in
bay of Primrose Brook, May 27, 1952, 28 clitellate specimens,
many juveniles. June 6, 1953, 2 clitellate specimens, many
juveniles.
Dover, Second St., in mud at water level of pond (pH of water
approximately 8.0), July 16, 1952, 10 clitellate specimens, many
juveniles. In stream feeding pond, June 8, 1953, 2 clitellate
specimens, 3 juveniles.
Castings were produced by this species at both localities. Early in
10 BREVIORA NO. 26
April, castings were formed on the surface of the mud, each casting
being approximately 3 in. in diameter and 3 in. high and generally
conical in shape. As time passed, the number of such castings became
greatly increased until in four weeks the whole area was covered. In
early May at the Morristown location, castings were observed under
water. These increased in size progressively and became supported
by grasses growing out of them, becoming eventually a beehive shape
with a height of 6 to 8 inches and projecting above water level. As
warmer weather arrived, the water level receded, leaving the casting
exposed. Examination of these castings revealed numbers of juveniles,
adult specimens being obtained only by digging in 4 to 6 inches of mud.
In many cases, both of clitellate and juvenile worms and particu-
larly at the Morristown location, the last 10. to 30 segments were
brownish in color, probably indicative of parasitic bodies accumulating
in the coelom. Specimens collected at Dover were noticeably shorter
than those at Morristown where one specimen possessed 258 segments
(165 to 225 according to Olson 1940, p. 9). Posterior regeneration
was noted in six specimens.
Family MEGASCOLECIDAE
Genus PHERETIMA Kinberg 1867
PHERETIMA AGRESTIS (Goto & Hatai) 1899
Maplewood, rotting leaf pile in private garden, June 1953, 2
clitellate specimens.
PHERETIMA LEVIS (Goto & Hatai) 1899
Maplewood, rotting leaf pile in private garden, June 1953, 3
clitellate specimens.
PHERETIMA sp.
Maplewood, rotting leaf pile in private garden, June 1953, 2
aclitellate and 1 clitellate specimens.
The last three specimens were athecal as well as anarsenosomphic
and could not be referred to either of the two above-mentioned species.
All the pheretimas were found in a garden which contains many
oriental shrubs. The species are known to have been present for at
least three years and may possibly have been imported directly from
the orient.
The leaf pile is under a large conifer and the protection thus afforded,
together with the heat generated by the pile may preclude severe
winter freezing.
The writer is indebted to Dr. G. E. Gates for the identification of
the specimens of Pheretima sp.
1954 EARTHWORMS OF NEW JERSEY 1]
DISCUSSION
Earthworms of twenty species have now been recorded from
northern New Jersey and of these fifteen are recorded from the state
for the first time. As might have been anticipated, the majority of
species are widely distributed peregrine lumbricids of Eurasian origin.
Dendrobaena mammalis was found for the first time outside of Europe
where it is endemic in the British Isles. This species has been inter-
cepted on plant materials imported into North America (Gates 1953,
p. 530), and its appearance, therefore, might have been anticipated.
Bimastos palustris and Eisenia lonnbergi are evidently endemic, and
relatively common in their restricted habitat along the banks of
streams. Neither of these is listed by Olson (1940) as occurring in
New York State and this may imply that northern New Jersey, corre-
sponding roughly to the extent of the ice cap in glacial times, is the
northern limit of their general distribution. The glossoscolecid
Sparganophilus eiseni is presumed to be endemic, although its original
source may well be south of New Jersey. That each of the endemic
species has a limnic habitat may be of significance as the lack of
competition and the ease of colonization thus afforded may have
enabled these species to quickly repopulate the barren areas left by the
retreating ice cap.
Allolobophora limicola, found in three locations, has just recently
been recorded from Boston (Gates, 1953), where it was thought to
have been introduced from Europe. As the three New Jersey localities
are respectively 10 and 16 miles apart, possibility of a single intro-
duction seems unlikely. Allolobophora arnold: Gates, recently de-
scribed and only known heretofore from two New England states, was
found to be relatively common. It is tentatively considered to be of
European origin. The species of Pheretima were not expected as no
attempt was made to search in ornamental gardens or greenhouses.
They are probably of East Asian origin and may have been imported
directly with exotic shrubs.
Of the anticipated species, Bimastos tenuis, Allolobophora chlorotica,
and Dendrobaena subrubicunda were not found in recent collections.
A. chlorotica was probably overlooked as it has been found in Rockland
County, New York, within a few miles of the New Jersey border.
Altogether, collections were made in 55 locations, some of which were
found to be heavily populated. Five species were taken from a Dover
garden and six species from a pond and adjacent swamp near Morris-
12 BREVIORA NO. 26
town. Allolobophora arnoldi, A. caliginosa, A. limicola and A. longa
were found together in one piece of marshy ground in a garden.
In northwestern New Jersey, comprising Warren and Sussex
counties, the terrain is characterized by dense woodland rising to
1500-1800 feet. The soil of these woods is largely glacial drift with
numerous rocky ledges and outcrops. Searching for earthworms in
such localities proved to be most unproductive, possibly due to the
fact that the soil drains and dries out very rapidly. However, Bimastos
palustris and Evseniella tetraedra were found in woodland ditches and
streams at isolated locations where the top soil was entirely without
worms. Similarly, the top soil of wooded areas in Morris County was
found to be largely without worms except where the ground was damp
as often denoted by the presence of skunk cabbage (Symplocarpus
foetidus).
SUMMARY
Twenty species of earthworms have now been recorded from
northern New Jersey, including sixteen lumbricids, one glossoscolecid
and three megascolecids. Thirteen of these arerecorded for the first time
from this state, Dendrobaena mammalis being reported for the first
time outside of Europe. Bimastos palustris and Eisenia lénnbergi are
considered to be endemic and are widely distributed in the northern
part of the state. Allolobophora limicola was found for the second
time outside Europe and in three localities. Three species of Pheretima
were present in a garden.
REFERENCES
CrerRnosviTov, L. and A. C. Evans
1947. Lumbricidae. No. 6 in Synopses of British Fauna. London,
Linnean Society, 1-36.
Eaton, T. H., JR.
1942. Earthworms from the North Eastern United States. Jour.
Washington Acad. Sci., 32, No. 8: 242-249.
Gatss, G. E.
1952. New species of earthworms from the Arnold Arboretum, Boston.
Breviora, 9: 1-3.
1953. On the earthworms of the Arnold Arboretum, Boston. Bull. Mus.
Comp. Zool. Harvard, 107, No. 10: 497-534.
1954 EARTHWORMS OF NEW JERSEY 13
Moore, H. F.
1895. On the structure of Bimastos palustris, a new oligochaete. Jour.
Morph., 10: 473-496.
Otson, H. W.
1940. Earthworms of New York State. Amer. Mus. Nov., No. 1090: 1-9.
SMITH, F.
1917. North American earthworms of the family Lumbricidae. Proc.
U.S. Nat. Mus., 52: 157-182.
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BREVIORA
Museum of Comparative Zoology
NuMBER 27
CAMBRIDGE, Mass. Fespruary 5, 1954
ANTERIOR REGENERATION IN A
SEXTHECAL SPECIES OF
LUMBRICID EARTHWORM
By G. E. GATES
Records of regeneration, in an anterior direction, at known levels,
by posterior pieces of lumbricid earthworms have been brought to-
gether in two recent contributions, the first (Gates, 1949) containing
all that had been found for Eisenia foetida (Savigny) 1826, the second
(Gates, 1953) those for the other species that had been studied.
Records also have been found of anterior regeneration by a number
of specimens, unwittingly used along with F. foetida, that must have
belonged to yet another species. Identification of this species is the
problem with which this note is concerned.
The records involved are available only because it had been found
to be ‘“‘a tedious operation cutting off a definite number of segments’’.
Accordingly, ‘the number of segments cut off was not counted at the
time”... but was calculated “after regeneration by the position of
the vasa deferentia! or segments containing the seminal receptacles”’
(Morgan, 1895, p. 452). The latter organs, now usually called sperma-
thecae, were said to be normally in segments “9-10-11” (¢dem, p. 455).
Such a characterization with reference to paired organs of earthworms
certainly can be understood to indicate presence of three pairs of the
spermathecae, located one pair each in segments ix, x and xi. But,
E. foetida has only four spermathecae, usually present, according to
various authorities, in ix—x though occasionally in x-xi. The variation
does not involve location of spermathecal pores which are on inter-
segmental furrows 9/10 and 10/11. In view however of the difference
in internal location might it be possible that two pairs of spermathecae
were so located as to require reference, in most cases, to three segments?
1 This means: segmental location of the external openings of the deferent ducts, i.e.. the
male pores. GEG
bo
BREVIORA NO. 27
In this connection location of spermathecae was determined in each
older individual (30 clitellate and 4 late juvenile) of the first entire
colony of E. foetida that was accessible. Some variation as to which
of two consecutive coelomic cavities any spermatheca got into as it
grew through the parietes was indeed found (Table below). Never-
Table 1
Segmental location of spermathecae in a colony of E. foetida
Segments Number of specimens
GaN OF. SAC SEW ce dle ete niet cee Dione 17
NOV ok tie pees ones ee Bae ee ce 2
102 by ees Veena Te A eee caeeeead oicatt: 5 pp cNeme 4
Dililiebe if 4 van Fas eee te Nahe rae eee 1
EG AAO. RAO bry it wes eye ay ee eee te ys: 3*
LAST ON By (Dens Ce eae aR PN Pod 1
1 ACO ped ba 0 noe Br Wee er ee ste ne cee eg ca 1 29
LO! THO SRO IG.) ee ees cae 1
POM OR MOAR tie 5. oe oe eel ae 1
ORO uO gL crak eye ater pets etary 1
ROS Rs pel OMe eee sce, ae See hes Ae 1 4
Spiral abnormality in metamerism of the
Spermathecal region... J. ge: oo ty ene 1 1
* On the left side spermathecae in ix and x, on the right side both sperma-
thecae in x.
Each worm had four spermathecae.
theless, only in four of the thirty-three metamerically normal worms
is it mecessary to refer to three segments to indicate spermathecal |
location, and in two cases all spermathecae are actually in one segment.
These results, together with previous findings, show it is unlikely that
twenty-two out of twenty-seven specimens of FE. foetida would have
had spermathecae in three segments. Furthermore, the method by
which location of spermathecae of the other five specimens was indi-
cated, e. g., ““-7-8’’, as if the first of three pairs’ had not been found,
also supports the interpretation of “normal” that was first suggested
above, i. e., presence of three pairs in three consecutive segments’.
2 Tn one case, ‘‘—6—-7?’’, in which the question mark seems to indicate uncertainty but as to
what, was not explained.
3 Seminal receptacles cannot be regarded as a lapsus calami for seminal vesicles. The latter
are four pairs, in ix—xii, in EH. foetida. Other species may have only three pairs but they are
not in three consecutive segments, ix, xi—xii.
1954 ANTERIOR REGENERATION IN AN EARTHWORM 3
Accordingly, it is concluded that Morgan’s specimens with sperma-
thecae in two segments were quadrithecal and, as he supposed, F.
foetida, but that the others were of some sexthecal form with sperma-
thecae in ix, x, xi. Three lumbricid species of this country are so
characterized: Allolobophora chlorotica (Savigny) 1826 and Lisenia
lénnbergi (Michaelsen) 1894 with spermathecal pores on 8/9-10/11,
Dendrobaena octaedra (Savigny) 1826 with the pores on 9/10-11/12.
E. lénnbergi, though at present within the same genus as foetida, has
a quite different habitus, is a native of the southern states, has never
been reported from the vicinity of the region where the worms in
question apparently were secured, has not been found in manure
heaps — the source of the material — and is unlikely to have been
involved. A. chlorotica normally has an obvious yellow or green color-
ation, as well as other characteristics that would immediately dis-
tinguish it at a glance from foetzda, and has only once been reported
from the vicinity of a manure pile. D. octaedra does have the same
red pigmentation that characterizes F. foetida, uniformly distributed
(instead of in transverse bands) as in many individuals of L. foetida,
has been found occasionally in manure piles, and does have much less
special glandularity in the region around the male pores (so that site
of the male pores would be less easily recognizable). Of the three
species, D. octaedra is the one most likely to have been inadvertently
accepted as FE. foetida. Nothing, however, has been known of re-
generation at levels in front of 15/16 in any species of Dendrobaena
and D. octaedra has not, apparently, been available to others who
have studied regeneration in earthworms. An unusually large octaedra
proportion of the population (in the manure heap that provided the
experimental material under consideration) seemingly is indicated by
such figures as are available: one of eleven specimens (Table 3, Morgan,
1895), five of sixteen (Table 10), six of eleven (Table 12, first half),
four of nine (Table 13), twenty-two of the twenty-seven cases in which
spermathecae were mentioned.
The number of records (22) of anterior regeneration that can now
be recognized as of octaedra is larger than for any other lumbricid
(Gates, 1953) except FE. foetida and two species which have not yet
been studied in this country, Allolobophora terrestris (Savigny) 1826
and Lumbricus rubellus Hoffmeister 1843. These records (Table below)
show that head regeneration is possible at each level from 1/2 to 8/9
inclusive. The five segment regenerate shows that equimeric regener-
ation can be expected at each level from 1/2 to 5/6 inclusive at least.
4 BREVIORA NO. 27
Failure to secure equimery at levels behind 3/4 indicates that con-
ditions, either of the external or internal environment or both, were
less than optimal for regeneration by octaedra, as well as for E. foetida’.
The results obtained from four worms that were deliberately cut
diagonally instead of transversely (note under table) may indicate
that head regeneration is also possible at levels back at least to 12/13
Table 2
Number of segments in head regenerates of Morgan’s
sexthecal earthworms
Number of segments Record quoted from
Level of regenerated Morgan, 1895,
amputation* 2 3 4 oy) pages
EL 2/3 1 = = = 447
EL 3/4 3 1 - = 447, 455
EL 4/5 2 = = 448, 456
EL 5/6 = 1 1 = 455
HL, 6/7 1 1 1 ~ 455, 456
EL 7/8 - 1 1 - 456
EL 8/9 1 1 1 1 455, 456
* A symmetrical homoeotic would not have been recognizable after operation.
EL Estimated level. Estimation made, after regeneration and presumably
after preservation, from position of male pores and/or location of
spermathecae.
When cuts were made diagonally “anterior segments obliquely amputated”
(p. 457), four to twelve segments were said to have been completed.
In three of these, which had spermathecae after regeneration in
“9-10-11”’ and in which 4, 8, and 12 segments had been ‘‘completed”’,
presumably no segments were completely excised. The other specimen
had a hypomeric regenerate presumably (indicated by location of the
spermathecae in ‘‘8-9-10’’). All of i-iii was then removed in addition
to parts of iv—x (‘7 segments completed’’).
After excision of a piece estimated to comprise 104 segments, and subsequent
regeneration (p. 455), there were still spermathecae in the first two
segments of the substrate. The estimate could have been correct,
regardless of species, only if the worm involved had been homoeotic
(+ 1 or more), or if spermathecae had been developed in the substrate
during regeneration. Nothing of the latter sort has ever been recorded
from the Lumbricidae.
4 The number of segments in the head regenerates of #. foetida was smaller than has been
obtained (Gates, 1949) and in view of this evidence from two different species the less favorable
conditions may have been in the external environment.
|
1954 ANTERIOR REGENERATION IN AN EARTHWORM
SUMMARY
Individuals of some sexthecal species were frequently used along
with EF. foetida in Morgan’s early studies of earthworm regeneration.
From the information available as to distribution, habitat, habitus,
etc., of the sexthecal species of this country, it is concluded that only
D. octaedra is likely to have been inadvertently taken for FE. foetida.
Nothing has been known of anterior regeneration in the genus Den-
drobaena and the records now attributable to D. octaedra show that it
is able to develop a head regenerate, in an anterior direction, at each
level back to 8/9, possibly to 12/13, and, in better conditions, complete
replacement of excised segments may be expected at least at all levels
from 5/6 anteriorly.
REFERENCES
Gatrs, G. E.
1949. Regeneration in an earthworm, Eisenia foetida (Savigny) 1826.
I. Anterior regeneration. Biol. Bull., 96: 129-139.
1953. On regenerative capacity of earthworms of the family Lumbri-
cidae. Am. Midland Nat., 50: 414 419.
Morean, T. H.
1895. A study of metamerism. Quart. Jour. Micros. Sci., 37: 395-476.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. Freeruary 8, 1954 NUMBER 28
CLEMMYDOPSIS BODA
A VALID LINEAGE OF EMYDINE TURTLES
FROM THE EUROPEAN TERTIARY
By Ernest WILLIAMS
In 1847 Hermann von Meyer gave the name L’mys turnauensis to a
small emydine turtle from the Upper (Sarmatian) Miocene of Turnau
in Steyermark, Austria. He published no description at that time,
but validated the name by a full description and figure in 1858.
Emys turnauensis was founded on an anterior fragment of carapace
(nuchal and both first peripherals, second peripheral of the left side,
both first pleurals, and parts of left pleurals 2, 3. and 4). Its most
evident peculiarity was the absence on the first pleurals of any indi-
cation of grooves for the costal scutes. The first vertebral thus ex-
tended across the entire front of the shell back of the anterior marginals
and the relatively broad nuchal scute. The second and third vertebrals
less obviously but unmistakably had a similar great lateral expansion.
Von Meyer commented: ‘‘Von allen mir bekannten Schildkréten mit
Grenzeindriicken zeichnet sich vorliegende durch den Mangel an
Seitenschuppen aus. ... Der Verlauf der Grenzeindriicke sonst ist in
dieser Schildkréte so regelmassig dass der Mangel an Seitenschuppen
unmoglich fiir eine zufillige Erscheinung oder fiir eine Abnormitit
gehalten werden kann: er wird der Species wirklich zugestanden haben,
und es wird sich eigentlich nur um Entscheidung der Frage handeln,
ob der giinzliche Mangel einer Schuppenart in einer Schildkréte zur
Errichtung eines eigenen Genus brechtigt oder nur zu den Kennzeichen
gehort, welche bei der Unterscheidung von Species in Anwendung
kommen. Mit der Beantwortung dieser Frage méchte ich um so mehr
bis zur Kenntniss der fehlenden Theile der Schildkréte, namentlich
des Bauchpanzers zuriickhalten, da die hervorgehobene Abweichung
im Hautskelet, so auffallend sie ist, mit einer Abweichung in der Zahl
oder Form der knéchernen Theile, wenigstens so weit diese verliegen
2 BREVIORA NO. 28
nicht verbunden sich zeicht; die knéchernen Theile sind vielmehr
Emys entsprechend gebildet. Zur Errichtung jedoch einer neuen
Species war wohl hinreichender Grund vorhanden.”’
Fig. 1. Clemmydopsis turnawensis (von Meyer). A. Type specimen, after
v. Meyer. B. Carapace, after Staesche. C. Plastron, after Staesche.
Von Meyer mentioned also that he had an anterior fragment from
Tertiary formations at Chaux-de-fonds, Switzerland, which might
belong to Emys turnauensis, but that he had never seen anything re-
sembling this species elsewhere in Switzerland.
In 1927 Anton Boda described and figured from the Lower Pan-
nonian (Lower Pliocene) beds near Sopron, Hungary, a complete
1954 CLEMMYDOPSIS 3
dorsal shell of a form which he recognized as similar in its vertebral
pattern to Emys turnauensis but which he referred to a new genus and
species as Clemmydopsis sopronensis. The new form differed from
Emys turnauensis in the shape of the neurals. In Clemmydopsis so-
pronensis neurals one to three were hexagonal, short-sided behind, and
neural four quadrilateral, while the first neural was oval, the second
7
Fig. 2. Clemmydopsis sopronensis Boda. A. Type carapace, after Boda.
B. Plastron, after Thenius.
to fourth hexagonal, short-sided in front, in Emys turnauensis. Boda
assigned his new form to the section of the Emydinae which includes
Geoemyda (Nicoria of authors), doing so on the basis of the character
of neural shape to which great taxonomic weight had been given by
Boulenger, Siebenrock and others.
In 1931, K. Staesche placed on record from the Sarmatian Miocene
of Steinheim in Wiirttemberg material very similar to the unique type
of Emys turnauensis but which he regarded as new, giving it the name
Clemmys steinheimensis. Staesche’s material was much more nearly
complete than that of either Boda or von Meyer. He had three speci-
mens, none individually perfect but together providing satisfactory
knowledge of both carapace and plastron.
Staesche described his material very fully and discussed the re-
lationship of his species to Emys turnauensis and Clemmydopsis so-
pronensis. He admitted the possible synonymy of his species with
4 BREVIORA No. 28
Emys turnauensis but considered that one minor difference might be
of specific value: the marginal scutes did not encroach upon the first
pleural of twrnauensis as they quite clearly did in steinheimensis. He
relegated both forms to the genus Clemmys, regarding the single point
of difference from Clemmys as usually understood — the absence of
the first and second costals — as of specific value only.
In the case of Clemmydopsis sopronensis, Staesche was not in doubt
as to the distinctness of the species, but he was quite doubtful of the
validity of the new genus. He argued that the difference in neural
shape appeared to preclude inclusion of stetnheimensis in Clemmydopsis
and that, therefore, if the character of the horny shields was counted
of much value, a new genus would need to be erected for steinheimensis.
He concluded: ‘‘Vermutlich diirfte daher wohl auch eine nahere
Verwandschaft zwischen Clemmys und Clemmydopsis bestehen, derart,
dasz der Gestalt der Neuralplatten (kurzeste Seiten hinten oder vorn
gelegen) keine so iiberragende systematische Bedeutung zukommen,
kann, wie dies Boda nach Siebenrock annimmt. Das Fehlen der 1.
und 2. Lateralschilder ist nicht als Genus — sondern nur als Artcha-
rakter zu bewerten. Dieses Merkmal tritt bei zwei verschiedenen
(Neuralplatten!) aber wohl verwandten Gattungen an der Wende von
Miociin und Pliocin auf, um alsbald wieder zu verschwinden. Man
k6nnte versucht sein in Dacque’schem Sinne von einer ModestrOmung
zu sprechen, denn ein besonderer Zweck diirfte dieser Einrichtung
kaum zugrunde liegen. Mit der Annahme einer niheren Verwand-
schaft beider Gattungen kann man aber auf diese Deutung verzichten.”
In 1934 T. Szalai in a list of the fossil turtles of Hungary synony-
mized Clemmydopsis Boda with Geoemyda Gray on the basis of the
similar neural shapes. rejecting as not significant at the generic level
the peculiarities of the horny shields.
Erich Thenius (1952) has followed Szalai’s generic assignment (pre-
ferring, however, the synonymous name Nicoria) in reporting new
finds of sopronensis from a new locality Brunn-Vésendorf near Vienna.
Thenius’ material is important in that it completes our knowledge of
sopronensis by providing (from dissociated specimens of all the plastral]
parts) the characters of the plastron, heretofore unknown.
Thanks to the kindness of the authorities of the Staatliche Museum
fiir Naturkiinde in Stuttgart and especially to the friendly cooperation
and diligence of Dr. Karl Dietrich Adam of that museum I have been
able to examine the type specimens of Clemmys steinheimensis Staesche.
There is little to add to Staesche’s accurate description and excellent
1954 CLEMMYDOPSIS 5
photographs. There are, however, some points of interest in regard
to the interpretation and systematic position of the fossils.
I wish first to suggest that steinheimensis may best be synonymized
with turnauensis. The two named forms are equivalent in age, not far
distant in locality and distinguished by a single quite trivial character
which may well be only an individual peculiarity of the unique ue
of turnauensis.
If this synonymy is correct we are dealing then with four occurrences
of only two forms. Yet these two forms have received from the very
few authors who have written about them four different generic as-
signments: “Emys’’, “Clemmydopsis, new genus”, “‘Clemmys’’, and
“Geoemyda ( = Nicoria)’’.
Of these generic names the first may be dismissed at once; it belongs
to a period in which almost all fossil emydine species and even some
forms not belonging to the family Testudinidae or the suborder
Cryptodira were placed in the genus E’mys.
More serious discussion must be accorded the other generic as-
signments, but it appears to me that previous authors have failed to
consider one important possibility: that the two forms, which are after
all chronologically consecutive, are phyletically related.
The absence of the first two costal scutes in the two forms turnau-
ensis and sopronensis is a quite extraordinary phenomenon apparently
not closely approached by any recorded aberration of the horny
shields. The similarities in detail shown by the two forms are fan-
tastically close if they are the result of parallelism only. There is a
real difficulty in the plural occurrence of so improbable an event. The
difficulty, however, ceases to exist if we assume the event happened
only once and if we explain the similarity of the two forms by direct
inheritance. The evolutionary and therefore the taxonomic dilemma
which the two forms have appeared to present is in all essentials solved
by the hypothesis of phyletic relationship.
There are, of course, real differences between the two species, and,
in fact, one of the differences — neural shape — has been considered of
high taxonomic value and is still used in the taxonomy of Recent
forms to discriminate genera.
Without question in the Recent emydine turtles the character of
neural shape tends to have utility in segregating natural groups,
although not without some instances of difficulty, but, granting to the
character the maximum of utility for Recent forms it must still be
used with discretion when dealing with fossil forms. It can never be
6 BREVIORA NO. 28
forgotten that neurals which are hexagonal, short-sided in front, are
primitive and that other types have evolved from that condition.
Inevitably then some of the ancestors of genera showing the modified
types of neurals would be placed — if that character were alone con-
sidered — in a more primitive genus. Quite certainly Geoemyda and
its relatives have evolved from a form with Clemmys type neurals.
There are indeed a number of examples of evolution in neural shape
displayed in the fossil record. The fossil tortoise Stylemys nebrascensis
of the Oligocene ef North America regularly has all the neurals after
the first hexagonal short-sided in front: the more highly evolved species
of the same genus froin the John Day beds (lowest Miocene) regularly
have one or more octagonal neurals. Hadrianus with primitive neurals
evolved into later subgenera of T’estudo with modified neurals. There
is also among Recent forms considerable intraspecific variation in
neural shape.
In the present instance turnauensis is temporally antecedent to
sopronensis and in the matter of neural shape structurally more primi-
tive. It is surely a possibility worth attention that the one has a
phyletic relationship to the otber.
Except for neural shape there are no known differences between
turnauensis and sopronensis which are not at the specific level or below
it, and, since none of fossils is quite perfect, it is possible in fact that
some of the recorded differences are matters of interpretation and
reconstruction and not real. I tabulate the differences below:
turnauensis sopronensis
1. gulars narrow, not overlapping 1. gulars broad, overlapping hu-
humerals merals
2. femoral scutes longer than 2. pectoral scutes longer than
pectorals femorals
3. no anal notch 3. a distinct though narrow anal
notch
4. first vertebral encroaching on at 4. first vertebral encroaching only
least the first, second and third on the first and second peri-
peripherals pherals
5. first neural rectangular-oval, 5. neurals 1-3 hexagonal, short-
neurals 2-8 hexagonal, short- sided behind, neural 4 quadri-
sided in front, all moderately lateral, neurals 5-8 hexagonal,
broad short-sided in front, all still
broader than in turnauensis
6. fifth vertebral and fourth costal 6. fifth vertebral and fourth costal
moderate in size quite small relatively to the
other scutes
1954 CLEMMYDOPSIS 7
These differences will serve — even if one or two should fail — to
distinguish the species.
Now, granting that the two species are directly related, to what
genus or to which genera should they be referred?
Admittedly, if the genus Clemmydopsis is recognized, it will be solely
on the pattern of the vertebrals and costals. It is, of course, possible
that some of the skeletal parts which are not known — skulls, limbs,
or vertebrae — might show striking differences from the related forms,
Geoemyda or Clemmys, but this is an untestable hypothesis, which it is
useless to consider. The question is then a simple one: Is a single
character — at once striking and superficial — adequate for generic
distinction?
There is unfortunately no objective way to evaluate such a question.
Characters involving quite radical differences are in some cases infra-
specific, while other characters which to the non-specialist are utterly
trivial separate suprageneric groups. That the rank of supraspecific
categories is wholly subjective or nearly so is a point generally agreed
upon. In practice it is necessary to steer a middle course between
those who would reduce genera arbitrarily on mnemonic grounds —
deploring the necessity of remembering so many names — and those
who tend to regard as generic any differences which are recognizable
immediately and as specific any differences which are determinable
with more difficulty.
In the present case there are several grounds for accepting — at
least for the present — Clemmydopsis as a valid genus.
1. The peculiar modification of the horny scutes, though a single
character, is also one without parallel in the normal characters or the
known variants of the chelonian order. Anomalies of the plates and
scutes of turtles have been studied intensively by Gadow, Newman.
Coker, Grant, Lynn and others using literally many hundreds of
specimens, yet nowhere does there appear to be any record of a scute
anomaly resembling that seen in these forms from the Miocene and
Pliocene of central Europe. The closest parallel is with two cases
cited by Grant, one in Testudo denticulata and one in Chrysemys picta
in which the vertebrals do touch the marginals. But even here the
similarity is verbal only, the marginals being enlarged and not the
vertebrals while the grossest asymmetries and distortions indicate the
abnormality of the condition.
2. There is currently recognized a Recent genus Notochelys with the
single species NV. platynota which differs from Clemmys primarily by
8 BREVIORA NO. 28
the intercalation of a small scute between the usual fourth and fifth
vertebrals. But this condition, while a population character in pla-
tynota, is a rather common variant in numbers of other members of
the Testudinidae. This variation is especially common in Homopus in
which a very substantial fraction of at least H. areolatus shows a
similar condition. Thus while there is a weak argument for Clemmy-
dopsis on the grounds of symmetry in generic discrimination, it may
in this case be countered by a demand for suppression of the generic
separation for N. platynota, which, while certainly a distinct species,
does not obviously merit more than subgeneric distinction.
3. However, the best ground for separating Clemmydopsis as a
genus is the impossibility of placing the two central European forms,
directly related though they seem to be, in any one currently recog-
nized genus. On the current definitions of emydine genera the Miocene
form belongs in Clemmys, where Staesche has already placed it, and
the Pliocene form in Geoemyda ( = Nicoria) to which Szalai and
Thenius have already referred it. This, however, is an inadmissible
solution since it would make the genus Geoemyda polyphyletic.
It is entirely possible, indeed probable, that Clemmys and Geoemyda
as at present defined are mere form. genera, but the true relationships
within the Clemmys-Geoemyda section of the Emydinae are still to be
analyzed and proinise to be difficult of analysis. Thus while it is
perhaps possible to look forward to a time when relationships will
become known, and larger genera may be recognizable within the
Emydinae, it is not now desirable to randomly unite emydine genera,
in so doing pretending to a knowledge which is not at hand. Rather
it is preferable to split rather finely at the moment, counting the
present as that period of analysis which must precede a synthesis.
We:need first to discover the correct phyletic sequences. We must
first recognize the family tree; afterward we may dispute the nomen-
clatorial rank of the several twigs and branches.
Clemmydopsis Boda should then be retained at present for an ap-
parently short-lived but uniquely specialized lineage of emydine turtles
from the later Tertiary of middle Europe. Whether or not it should
ultimately be retained as a genus in formal nomenclature, it is certainly
a “‘gens’’ in the sense of Vaughan 1905 (in Cain 1952) (a lineage or
phyletic line). ;
Examination of the type of Clemmys steinheimensis in Stuttgart was
made possible by the support afforded by a Guggenheim fellowship
1952-1953.
1954 CLEMMYDOPSIS 9
REFERENCES
Bopa, A.
1927. Clemmydopsis sopronensis, n.g., n.sp. aus der unteren pannonischen
Stufe von Sopron in Ungarn. Zentralbl. Min. ete., Abt. B, 1927,
pp. 375-383.
Cain, A.
1952. Geography, ecology and coexistence in relation to the biological
definition of species. Evolution, vol. 7, pp. 76-83.
Coker, R.
1910. Diversity in the scutes of Chelonia. Jour. Morph., vol. 21, pp.
1-75.
Gavow, H.
1899. Orthogenetic variations in the shells of Chelonia. im Arthur
Wiley’s Zoological Results, part 3, pp. 207-222.
IRANT, C.
1936. An extraordinary tortoise shell. Copeia, 1936, pp. 231-2382.
Lynn, W. G.
1937. Variation in scutes and plates in the box turtle, Terrapene carolina.
Amer. Nat., vol. 71, pp. 421-426.
Meyer H. von
1847. (Letter on various fossils.) Neues Jahrb. f. Min. ete., 1847, pp.
181-196.
1858. Schildkréten und Saugetiere aus der Braunkohle von Turnau in
Steyermark. Palaeontographica, vol. 6, pp. 50-55.
STAESCHE, K.
1931. Die Schildkréten des Steinheimer Beckens. A. Testudinidae.
Palaeontographica Suppl., vol. 8, part 2, pp. 1-17.
Szaual, T.
1934. Die fossilen Schildkréten Ungarns. Folia Zool. Hydrobiol., vol. 6,
pp. 97-142.
THENIUus, E.
1952. Die Schildkréten (Testudinata) aus der Unterpliocin von Brunn-
Voésendorf bei Wien. Neues Jahrb. f. Min. etc., 1952, pp. 318-334.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. Fresruary 8, 1954 NUMBER 29
ABSENCE OF MESOPLASTRA IN A
PELOMEDUSA
(TESTUDINES, PELOMEDUSIDAE)
By Ernest WILLIAMS
Mesoplastra (a pair of bones intercalated between hyo- and hypo-
plastra) are believed to be primitive features in turtles. They were
often present in Jurassic and Cretaceous genera, and two pairs of
mesoplastra were reported by Fraas (1913) in Proterochersis, one of
the Triassic genera. Mesoplastra have, however, been lost several
times independently: (1) in the Jurassic and Cretaceous plesiochelyids
and thalassemydids; (2) in Wealden Chitracephalus; (3) in all erypto-
dires; (4) in the chelids among pleurodires. In many of the groups in
which they are known, they were early reduced. Primitively they
extend quite across the plastron and meet centrally. In some Pleuro-
sternidae, however — Platychelys — and in some Baenidae (within the
genus Baéna, ct. Baéna riparia, Hay, 1908) they fail to meet in the
center.
To my knowledge, however, no instance has been recorded previ-
ously of the complete absence of mesoplastra in any form in which
they were typically present. Such an example is now furnished by a
specimen of Pelomedusa subrufa from Uganda, collected by Delme
Radcliffe and now 1905-5-19-1 in the collections of the Reptile Section,
British Museum (Natural History).
In Pelomedusa the mesoplastra are typically small and lateral. In
BM 1905-5-19-1, however, no mesoplastra are visible, and instead the
hyo- and hypo-plastra join for their full transverse extent and do so
quite symmetrically on the two sides just as in the Chelidae or the
Cryptodira. The carapace again joins the plastron perfectly without
affording any place for a trace or rudiment of mesoplastra.
2 BREVIORA NO. 29
Yet this specimen is clearly referable on “‘habituellen Merkmalen”’
to the family Pelomedusidae, the genus Pelomedusa, the species
subrufa (the single species of the genus).
The carapace is quite normal except for some minor asymmetries
and the fusion of vertebrals 3 and 4 to a single scute. The nuchal
scute is, as usual, absent. The first neural is rather characteristically
tapered in front. In normal fashion there are 7 neurals, and the
posterior pleurals meet behind the neurals separating them from the
suprapygal. In contrast to chelids the posterior peripherals are all
rather narrow and show no suggestion of posterolateral expansion.
The plastron is quite typical of the genus in the size and relations
of its horny seutes. There is the usual median fontanelle, though it is
small in this instance, as seems frequently to be the case in northern
representatives of the genus and species.
The skull is quite characteristically pelomedusid and very unchelid
in the temporal emargination from behind and in the marked pro-
jection of the opisthotic posterior to the squamosal.
All these features and the lack of mesoplastra are well shown in the
excellent photographs (Plates | and 2) taken by Peter Green of the
photographic staff of the British Museum (Natural History) and
reproduced by permission of the Trustees of that institution.
The absence of mesoplastra is quite certainly an individual variation.
Four specimens from Mt. Elgon, Uganda, otherwise very similar to
the individual without mesoplastra, all show these elements well-
developed and only slightly varying in size. All resemble the aberrant
specimen in the reduction of the median fontanelle of the plastron:
one, indeed, has the fontanelle completely closed.
The presence of mesoplastra is supposedly a family character of the
Pelomedusidae, but except in the genus Pelusios the mesoplastra are
always small and lateral, in effect vestigial, and the disappearance by
individual variation of a vestigial feature is not too surprising.
It is, however, of special interest in connection with the suggestion
recently made by me that the genus Pe/usios with well-developed
mesoplastra (meeting medially) has been derived directly from the
genus Pelomedusa with reduced mesoplastra by a secondary expansion
of these elements. If this suggestion be valid, then, taken in connection
with the present case, we see in Pelomedusa an interesting ambivalence
of evolutionary potentiality, one type of variation reversing a previ-
ously well-defined trend to restore (with some differences) an ancestral
condition and, on the other hand, the opposite type of variation
1954 ABSENCE OF MESOPLASTRA IN A PELOMEDUSA 3}
Figure 1. Plastra of various turtles to show presence or absence of
mesoplastra. Mesoplastra shaded. A. Pelomedusa without mesoplastra.
B. Normal Pelomedusa. C. achelyid (Hydromedusa). D. Pelusios. FE. Po-
docnemis. (B to FE after Boulenger.)
4 BREVIORA NO. 29
carrying to completion the former trend and realizing a more modern-
ized type of shell.
It does not seem advisable on the basis of a single, probably rare,
variation to revise the definition of the family Pelomedusidae to
include forms without mesoplastra. Such a variation would appear —
on the face of the present evidence — to have as little taxonomic
weight as the occasiona! men born without legs have for the definition
of the family Hominidae.
Yet there is this difference: a man without legs is not likely to be a
selectively valuable variant, but, as the majority of the bving forms
testify, a turtle without mesoplastra is not only perfectly viable but
highly successful. It cannot be assumed, therefore — given a forward-
looking evolutionary point of view— that the pelomedusids will
always and in all cases be forms possessing mesoplastra.
There is indeed a special caution pertinent for paleontologists. If
the present specimen had been found as a fossil, it would almost
certainly have been misallocated as to family. There is in fact no
assurance — the pelomedusids having once been world-wide — that
all past species have always had mesoplastra. The use of key charac-
ters in identification without regard to total habitus and neglecting
the ever surprising power of animals to vary and to undergo evo-
lutionary change will only lead to error.
The observations recorded here were made during a visit to the
European collections, made possible by a Fellowship from the Guggen-
heim Foundation. Access to the specimens mentioned was generously
granted by Dr. H. W. Parker, Keeper of Zoology, British Museum
(Natural History), and assistance was given by Mr. J. C. Battersby
and Miss A. G. G. Grandison of the Reptile Section.
REFERENCES
Fraas, E,
1913. Proterochersis, eine pleurodire Schildkréte aus dem Ieuper.
Jahreshefte Ver. Vaterlands Wiirtt. Naturkunde, Stuttgart, 1913,
pp: 13-30:
Bina, (Os 12
1908. Fossil turtles of North America. Carnegie Inst. Washington Publ.
No. 75, pp. 1-568.
LovERIDGE, A.
1941. Revision of the American terrapins of the family Pelomedusidae.
Bull. Mus. Comp. Zool., vol. 88, pp. 467-524.
= ef WWM Kea.
PLATE 1
Pelomedusa subrufa B.M. N.H. No. 1905-5-19-1. Internal view of plastron.
PLATE 2
Pelomedusa subrufa B.M.N.H. No. 1905-5-19-1. Dorsal view of skull and
shell.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. FEBRUARY 8, 1954 NUMBER 30
AESTIVATION IN A PERMIAN LUNGFISH
By ALFRED 8S. ROMER
Museum of Comparative Zoology, Harvard University
and
EVERETT C. OLSON
University of Chicago
It is generally recognized that the survival to modern times of the
three living genera of dipnoan fishes — [’piccratodus (Neoceratodus) of
Australia, Protopterus of Africa and Lepidosiren of South America
is due in great measure to adaptations which tide them over the
seasonal droughts characteristic of the tropical regions which they
inhabit. .\ primary adaptation lies in their possession of functional
lungs which enable them to utilize atmospheric oxygen, supplementing
the reduced oxygen content in stagnant waters during drought; indeed,
in Protopterus, the lung-breathing function has attained such domi-
nance that the fish is unable to survive if air-breathing is not possible.
But air-breathing in itself is not entirely satisfactory as a means of
drought survival — particularly if, as may often happen, the stream
or pond dries up entirely. The Australian lungfish, under such cireum-
stances, may wallow into the mud, and have some chance of escaping
desiccation or enemies. In the African and South American forms,
an additional adaptation is present —— the development of a technique
of aestivation. Members of these genera are rather elongate and eel-
like in shape, in contrast to Hpiceratodus and to primitive lungfishes.
On the approach of the dry season the fish burrows in coiled form into
the mud, which hardens about it to form a cocoon; an opening from
the surface allows the entrance of air, but the amount needed is small,
due to a great reduction of metabolic processes. The fish passes into
a “summer sleep’? comparable in great measure to the hibernation of
various northern animals — a phase which ends with the return of the
water and the resumption of normal aquatic life.
2 BREVIORA No. 30
Nothing has been known until now of the origin of lungfish aesti-
vation. The early lungfishes, as far as known, all had a fusiform body
shape fairly comparable to that of the modern E’piceratodus, and hence
were incapable of coiling in a burrow in the fashion of the elongate-
bodied Lepidosiren or Protopterus; but it is possible that burrowing
of some sort accompanied by phsyiological features of aestivation,
might have preceded change in body shape. The lungfish Gnathorhiza,
characteristic of the Clear Fork Permian of Texas, has been suspected
by both the present writers to be related to the Lepidosiren-Protopterus
group (Romer and Smith 1934, Olson 1951) but there is no strong
proof of relationship, and the body structure was unknown.
Some years ago Dr. H. J. Sawin and Mr. Adolph Witte, while
engaged in work for the Texas Bureau of Economic Geology were told
by a resident of Willbarger County, of a geologic occurrence which
local geologists had been unable to interpret. They visited the locality,
made collections of the material for the Bureau and informed others,
including the present writers, of the incident. In 1952 Mr. Witte took
a Harvard field party, including the senior author, to the site; obser-
vations and collections were made at that time and on a return visit
the following spring.
The locality lies in southern Willbarger County, Texas, on the Reed
Ranch, in Section 34, Block 3, H. & T. C. RR., Co. Survey. The area
has not been mapped in detail geologically but the horizon appears to
be a short distance above the Lueders limestones and hence in the
lower part of the Arroyo Formation of the Clear Fork Group, Lower
Permian. Gently sloping exposures here run for a hundred yards or so
close to the south bank of Minnie’s Creek. Between two thin impure
limestones lies a bed of red shale. Scattered over the surface are
numerous circular dises, somewhat like large checkers, with diameters
which are for the most part between 5.5 and 10 em. The material of
the dises is essentially the same as that of the shale bed from which
they are derived, but is slightly harder and somewhat more calcareous,
so that they tend to weather out of their enclosing matrix. On exca-
vation, these discs are seen to be segments cleaved from vertical
cylinders which are thickly spread through the shale, often only a few
inches apart. In no case was a complete cylinder seen; the greatest
length preserved in a specimen excavated and collected was 28 cm.
The cylinders, in general, weather out from the softer shales with a
smooth surface; in some instances portions of the surface show slicken-
sides. No upper terminal portions were found; apparently, at the top,
1954 AESTIVATION IN A PERMIAN LUNGFISH 3
the material of the cylinder blends laterally with the superposed
valcareous shale. The lower ends of the cylinders are generally
smoothly rounded; in some cases, however, a lower terminal portion
has a diameter rather less than the major portion of the ‘‘bore’’.
The great majority of the “checkers” fractured off from the cylinders
show no organic material on their surfaces. A fair number, however,
show fish remains. In some instances these appear to be scales and
other disarticulated elements of small palaeoniscoids Most of the
remains, however, are clearly dipnoan in nature. The hardness of the
shale and the relatively soft nature of the bone makes preparation
difficult, and reliance has been had mainly on the structures seen in
section on the disc surfaces or, in some cases, weathering of the
external surface of the cylinders. Bony scales and plates are frequently
seen in section and in a few specimens fracture has shown scales in
surface view, the largest about 20 mm. in greatest diameter. Numerous
dises show hollow ribs, neural or haemal arches and fin supports,
sometimes in articulated series. Preparation of the basal segment of
one of the cylinders (figured) has revealed a considerable portion of a
caudal fin of the sort typical of Paleozoic dipnoans. Vertebral arches
and fin elements are present, partly as bone, partly preserved as
impressions. The vertebral axis, presumably cartilaginous in life, is
represented by a vertically oriented band devoid of impressions.
The end of the tail, poorly preserved. was curved around the bottom
of the cylinder.
While much of the lungfish material suggests a considerable degree
of decay and maceration, it seems certain that in many instances we
are dealing with cross-sections of complete, or nearly complete lung-
fish bodies. In a few instances where “articulating” discs composing
the greater part of a cylinder have been collected, it is obvious that
we are dealing with successive sections of a single fish, larger, frequently
rounded, body sections in the upper and middle parts of the series and
caudal sections, typically lens shaped, at the lower end. In many
cases the internal skeletal materials are rather irregularly scattered
across the surface of the section; in others the arrangement of the ribs
and fin supports is essentially regular, and toward the disc margins
there can be made out body outlines, marked by scale sections with
a surrounding layer of darker matrix.
The only lungfishes known from the Clear Fork belong to the genus
Gnathorhiza, mentioned above; Sagenodus, the common Pennsylvanian
genus, possibly antecedent to Ceratodus and L[piceratodus, is not
4 BREVIORA No. 30
recorded in Texas above the Wichita group. But while it is probable
that the Reed Ranch lungfish is Gnathorhiza, no tooth plates have been
found there to make identification certain. A second find, however,
has produced a number of fragmentary teeth. These are unmistakably
the blade-like, shearing teeth of Gnathorhiza and are referable to the
species G. dikeloda rather than G. serrata. This locality was discovered
by the junior author and Dr. Nicholas Hotton IIT in 1949 and has
been revisited several times since then. The bed in which the cylinders
occur is located in north central Knox County, Texas, on the Waggoner
ranch. It crops out along a small tributary to Little Mustang Creek
in locality KI of the junior author (see Olson, 1951 p. 104). The age
is middle Vale of the Clear Fork Group, Lower Permian. The site is
approximately 700 to SOO feet higher in the Clear Fork section than
the Reed Ranch locality.
The physical features of the cylinders from the two sites differ in no
Important particulars. Those from the Vale locality occur in a deep
red shale which varies in thickness from two to three feet. Above and
below this bed are lighter colored shales that appear to have been
deposited under somewhat different conditions. The shale is best
exposed along a small arroyo, but crops out sporadically over an area
of about an acre. The cylinders are distributed in rather irregular
groups of ten to twenty in unevenly spaced areas not much more than
a yard square. The longest cylinder encountered measured 45 cm.,
but as in the ease of the Arroyo specimens no upper termination has
been identified. The vertical orientation shown in the specimen figured
is constant throughout the deposit.
Organic remains in the cylinders from the two localities differ
somewhat. The most common remains in the Vale cylinders are ribs,
skull plates and scales of Gnathorhiza but, although the ribs in some
instances suggest the general body shape of the fish, no case in which
the scales show the body outline has been observed. As at the Reed
Ranch locality, remains of lungfish are found only in a fraction of the
cylinders. Scales of small palaeoniscids and infrequent scraps of skulls
of some small captorhinid reptile occur in others. More abundant,
however, are partial vertebral columns and single vertebrae of the
small, worm-like amphibian, Lysorophus. In no case have these
columns been found in the coiled condition characteristic of Lysorophus
specimens throughout the Arroyo formation.
There can hardly be any question that the lungfishes from the two
localities belong to the same genus. Some question does exist, how-
1954 AESTIVATION IN A PERMIAN LUNGFISH 5
ever, with respect to specific identity. "T'wo species, G. serrata and
G. dikeloda, are known from the Vale formation, but only the first
has been found in the Arroyo (Olson, 1951). Tooth plates of G. serrata
are small, lower plates ranging from 10 to 12 mm. in length, whereas
those of G. dikeloda are between 25 and 32 mm. long. The skulls of
adult individuals of G. dikeloda appear to have been between 75 and
100 mm. long; this would suggest that the skulls of G. serrata, unknown
as yet, would have a range of between 25 and 35 mm. The fact that
the Arroyo and Vale cylinders are comparable in size indicates that
they were made by fish of about the same dimensions. There is,
however, the puzzling fact that no teeth of the larger species have
been found in the Arroyo in spite of very extensive exploration of its
exposures. Except at the Reed Ranch locality lungfish remains are
exceedingly rare in the Arroyo; only four or five occurrences have been
reported, and these have consisted of single teeth. It may well be that
(. dikeloda was in existence during the Arroyo times, but that it found
suitable habitat in few localities in the areas which are now exposed.
With increasing aridity during the Vale, the number of habitats and
the chance of discovery probably were greatly increased.
From the data given above, the nature of the phenomena described
will be as obvious to the reader as to the authors. We are here surely
dealing with evidence of aestivation of lungfish at an early, Paleozoic
stage in the history of the Dipnoi. The cylinders are aestivating
burrows, made by lungfishes in the muddy bottoms of drying lakes or
streams. That most of the cylinders are empty is to be expected; most
of the fishes which occupied them survived to the return of the waters
and swam away. The burrows were later filled by sediments fairly
similar to those which had already formed the mud banks into which
they had burrowed. Fortunately, however, the filling material at the
localities described was of a somewhat harder, more calcareous, nature
than that of the original mud; had they been just the same, differential
weathering would not have occurred and the burrows would probably
have escaped observation. The lack of definite upper ‘‘openings”’ of
the burrows is presumably due to the disturbance of surface materials
on the return of the water. The presence of lungfish remains in some
of the evlinders indicates that a certain percentage of the aestivating
population failed to survive until the return of the waters. The finding
of occasional actinopterygian remains in the burrows is due to the
presence of palaeoniscid minnows in the returning waters before the
burrows had been filled by sediments. This probably applies to the
6 BREVIORA NO. 30
remains of Lysorophus as well. This amphibian, like Gnathorhiza, had
both aestivating and free swunming phases. It characteristically was
coiled during aestivation. The fact that coiling has not been observed
in the burrows indicates that Lysorophus entered the open holes left
by the lungfish upon the return of the water, and died during the
non-aestivating phase of their existence.
The finding of Gnathorhiza teeth in the material from the Vale (KI)
locality, together with the absence of identifiable remains of any other
lungfish type in the Clear Fork beds makes it seemingly certain that
the burrows were made by that lungfish and that alone. This evidence
of the adoption of aestivating habits by Gnathorhiza strengthens the
suggestion, gained from tooth plates and skull pattern, that this genus
is possibly ancestral to the modern aestivators, Lepidosiren and
Protopterus. One major difference in the mode of aestivation between
modern forms and Gnathorhiza may, however, be noted. The Recent
genera are elongate, eel-like animals which coil their bodies in a flask-
shaped burrow. There has been no previous evidence of body shape
in Gnathorhiza. The Permiam burrows may be somewhat narrower in
their upper portions; they are, however, essentially cylinders rather
than ‘‘flasks”’, and in no case is there evidence of more than one
section through a fish body at a given level of a burrow. It thus seems
certain that the Gnathorhiza body was of a normal, non-elongate
fusiform shape; that the burrow was formed by a spiral rotation of the
body and tail with the fish’s long axis in a vertical position; and that
the aestivating fish was posed erect in the burrow, essentially “standing
on its tail” with the mouth at the top, close to the surface where air
was available.
EXPLANATION OF PLATE
Above, « series of exposed burrows at the Reed Ranch locality; a half-dollar
gives an indication of size. Center, a terminal section of a burrow developed
to show an included lungfish tail, its tip curved upward. The skeleton is partly
preserved in bone, partly as impressions which have been painted white.
At the left, a lateral view of the specimen; at the right, a view from the under-
side. Below, left, a burrow in place at the KI locality; length 45 cm. Belov,
right, an incomplete burrow from the Reed Ranch locality; length 25 em.
8 BREVIORA No. 30
The genus Gnathorhiza is known not only from the Texas beds, but
also from the late Pennsylvanian of Illinois. In Texas, however, it is
very rare in the Wichita formations of the Permian where Sagenodus,
common in the Pennsylvanian, is fairly abundant: on the other hand,
there is no trace of Sagenodus in the Clear Fork, where Gnathorhiza
replaces it. This local succession is in agreement with our beliefs as
to the climate of early Texas. Various lines of evidence suggest that
Clear Fork time, particularly that of deposition of the Vale and Choza
of that group, was one of increasing aridity with marked seasonal
variations in rainfall (cf. for example Olson, 1948). Accompanying this
change is a shift in the local dipnoan faune from Sagenodus, a presumed
non-aestivating form, to Gnathorhiza, better adapted for survival under
drought conditions.
LITERATURE CITED
Orson, E. GC.
1948. A preliminary report on vertebrates from the Permian Vale
formation of Texas. Jour. Geol., 56: 186-198.
1951. Fauna of Upper Vale and Choza. 38. Lung fish of the Vale
4. The skull of Gnathorhiza dikeloda Olson. Fieldiana, Geol., 10:
104-124.
Romer, A. S. and H. J. Siri
1934. American Carboniferous dipnoans. Jour. Geol., 42: 700-719.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. Marcu 3, 1954 NUMBER 31
NEW FRESHWATER GASTROPOD MOLLUSKS
OF THE AFRICAN GENUS LANISTES
Bye PAIN
London, England
The two new species and one subspecies described herein form part
of the collection of the Museum of Comparative Zoology at Harvard
College. I am deeply indebted to Dr. J. C. Bequaert and Dr. W. J.
Clench for the opportunity to examine and describe them.
LANISTES (LANISTES) KOBELTI, new species
Figures 3-4
1913. Lanistes sp.? Kobelt, Mon. Amp. in Martini and Chemnitz, Syst.
Conch. Cab., 1, Abt. 20, Neue Folge, pl. 30, figs. 44a.
Shell sinistral, fairly thin. Whorls strongly shouldered below the
suture, with a very slight ridge. Spire produced, apex much corroded.
Aperture ovate; parietal lip simple and only thinly calloused; colu-
mella thin, not reflected. Umbilicus very wide and deep, limited
behind by a very prominent, angular, thickened ridge, which forms a
slight angle at the base of the moderately concave columella. Growth-
lines irregular, interspaced with fine axial striae, and cut by very fine,
irregular, close-set spiral engraved lines in the holotype (very super-
ficial in the paratype). Color pale yellowish-brown, with numerous
irregularly spaced reddish-brown bands; interior of aperture pale
bluish-white banded with reddish-brown. Operculum corneous, of the
usual shape.
Measurements (in millimeters)
Length Width Aperture
Length Width
Holotype 23 24 15 11 Fig. 3
Paratype 23 23 15 11 Fig. 4
S 18 23 15 10
~
2 BREVIORA NO. 31
Locality. “Africa’”’ (probably West Africa); holotype M.C.Z. No.
173419; paratypes M.C.Z. No. 173420 and in T. Pain Coll.
This unusual little species was apparently figured by Kobelt (1913,
pl. 30), but he did not name it or mention it in the text. The umbilical
keel is very prominent, more so than in L. (L.) congicus Boettger.
The spire is also higher and the suture less flattened than in that
species. L. (L.) kobelti is, as the above table shows, very constant in
size and subject to but little variation in this respect.
LANISTES (MELADOMUS) CONNOLLYI, New species
Figures 1-2
Shell sinistral, perforate, fairly solid. Color dark reddish-brown,
without external bands. Whorls noticeably flattened and obtusely
shouldered above, rounded below, with a blunt angulation around the
umbilicus which is deep and moderately wide. Apex entirely corroded,
remaining whorls 3. Surface somewhat malleated, with fine and regular
growth-lines; spiral sculpture obsolete. Aperture ovate vertical; lip
simple, thin, edged with pale blue. Columella thin, evenly curved,
not angular at the base. Parietal lip bluish-white and only thinly
calloused. Color dark reddish-brown, without external bands; interior
of the aperture marked with faint reddish-brown lines. Operculum
unknown.
Measurements (in millimeters)
Length Width Aperture
Length Width
Holotype 40 36 31 22 Fig. 2
Paratype 45 40 O2N9 26 Fig. 1
- 43 39 30.5 24.5
“s dd 50 38.5 30
45 44 33 PAs)
Localities. Southern Rhodesia: Victoria Falls, holotype ex Connolly
Coll., M.C.Z. No. 173421, and paratypes, M.C.Z. Nos. 173422 and
179845; also at Congo Museum, Tervuren, and in T. Pain Coll.
Northern Rhodesia: River Kafue at N’kana, near the Congo border,
paratype, collected by C. von Hirschberg, M.C.Z. No. 173423.
Additional specimens of this species, referred to Lanistes ellipticus
von Martens by the late Major Connolly, are from the mouth of the
Chobe River, Southern Rhodesia (Holub) and from the Kafue River,
Mumbwa District, Northern Rhodesia (Pitman). (See Connolly, M.,
1954 NEW FRESHWATER GASTROPOD MOLLUSKS 3
1938, Ann. South African Mus., 33, p. 556.)
L. connollyi is easily confused with L. ellipticus v. Martens, which
occurs in the same general territory. It differs in its less upright
elliptical form, decidedly shouldered whorls more or less flattened
below the sutures, the deeper and wider umbilicus and the pronounced
blunt angulation around the umbilicus.
LANISTES (MELADOMUS) OVUM MWERUENSIS, new subspecies
Figures 5-7
Shell small, fairly thick. Color uniform greenish-brown, without
trace of bands. The spire is much corroded; remaining whorls 3,
rapidly increasing, the last considerably inflated. Sutures impressed,
but little flattened. Umbilicus narrow but deep. Surface with fairly
prominent growth lines, interspaced with fine wavy striae, especially
on the body-whorl. Aperture ovate, lip simple, interior pale blue.
Columella thin, curved, joined by a moderately thick callus ef bluish-
white color. Operculum as in typical L. (M.) ovum.
Measurements (in millimeters)
Length Width Aperture
Length Width
Holotype 30 30 19.5 16 Fig. 5
Paratype 30 32 21 U5 Fig. 6
“ 25 25 19 13.5 Fig. 7
Localities. Belgian Congo: Lake Mweru (or Moero) at Kilwa,
holotype, collected by Dr. L. van den Berghe, M.C.Z. No. 109842,
and paratypes, M.C.Z. No. 185683; also at Congo Museum, Tervuren.
and in T. Pain Coll.; Lake Mweru at Katenge near Kilwa, paratypes,
M.C.Z. No. 152483 (received from Dr. M. Wanson); Lake Mweru at
Pweto, paratypes, M.C.Z. No. 152346 (received from Dr. M. Wanson);
Lake Mweru at Moba (Chamfubu River) near Pweto, M.C.Z. No.
185682 (received from Dr. M. Wanson).
Only the holotype (Fig. 5) and one of the paratypes (Fig. 6), both
from Kilwa, appear to be full-grown. The paratype in Figure 7, also
from Kilwa, is immature. Most of the other paratypes are even
younger, the smallest being only 9 mm. in length
Apparently a dwarf race of L. (.W.) ovum Peters, possibly confined
to Lake Mweru. When adult it resembles the much larger L. (W.)
ovum procerus v. Martens, but has always a much thicker and more
solid shell. Immature shells are always of a uniform olive-green color
4 BREVIORA NO. 31
often with traces of malleation on the body-whorl. So far as known,
Lanistes ovum is the only species of the genus recorded from Lake
Mweru, where it is said to live in the swamps near the coast.
Figs. 1-2, Lanistes (Meladomus) connollyz, new species: 1, paratype, N’kana;
2, holotype. Victoria Falls. Figs. 3-4, Lanistes (Lanistes) kobelti, new species,
Africa: 8, holotype; 4, paratype. Figs. 5-7, Lanistes (Meladomus) ovum
mweruensis, new subspecies, Kilwa: 5, holotype; 6 and 7, paratypes. All figures
slightly over natural size.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. Marca 12, 1954 NuMBER 32
FOSSILS AND THE DISTRIBUTION OF
CHELYID TURTLES
2. Additional Reputed Chelyid Turtles on
Northern Continents:
Palaeaspis conybearti (Owen) — a Pelomedusid.
By ERNeEstT WILLIAMS
In the previous paper of this series a re-examination of the descrip-
tion of ‘‘ Testudo”’ leithii Carter from the Eocene of India demonstrated
that this fossil, so long considered a chelyid, belongs instead to the
related but quite different family Pelomedusidae.
The family Chelyidae of side-necked turtles (Pleurodira) is at
present restricted to South America east of the Andes, and to the
Australian region. The Indian fossil was one of several forms which
were thought to complete the discontinuous distribution of the family
by showing its former presence outside the Recent range of the family.
The present paper discusses another of these forms. I am deliberately
including every case known to me which has been seriously proposed
since the modern conception of the Chelyidae arose.
In 1870 the genus Palaeaspis was proposed by Gray for Emys
conybearti Owen, a fossil turtle from the London Clay Eocene. This
generic name has been ignored by all subsequent authors. Gray
placed his new genus in the Chelyidae next to the genus Hydromedusa
with the following generic definition: ‘‘Shell depressed; nuchal shield
large and quadrangular, included within the anterior marginal shield.
First vertebral shield very small, oblong and transverse, at hinder base
of nuchal: second ard third vertebral hexangular, as long as broad.
Sternum broad, truncated in front and notched behind, transversely
concave in the middle and bent up at the ends.”
Examination of the type specimen of Emys conybearii Owen
(B.M.N.H. 39449) at the British Museum reveals that Gray’s interpre-
2 BREVIORA NO. 32
tation of the dorsal shields is a perfectly possible one (Fig. 1). The
first shield behind the first marginals, which meet in the mid-line, is
divided transversely and on the analogy of Hydromedusa could be
interpreted as a nuchal succeeded by a small first vertebral. This
Fig. 1. Emys conybearii Owen. Dorsal view of type shell (after Owen).
1/6 natural size.
transverse division of the first median shield has been ignored in later
references to Emys conybearii, although it was mentioned and discussed
by Owen in the original description and was clearly shown by him in
his plate XXVIITA, which is a much restored but essentially accurate
representation of the specimen.
E. conybearti Owen was synonymized by Lydekker and Boulenger
(1887) with the earlier named Emys delabechei Bell, probably justly,
but the latter name was based upon an imperfect shell which does not
show the region which is critical in E. conybearii. With the example of
Hydromedusa in mind, it is not possible without further evidence to
dismiss the condition shown in EF. conybearii as an individual varia-
tion. Further, since the bony areas under the first vertebral are
1954 FOSSILS AND DISTRIBUTION OF CHELYID TURTLES 3
unusually elongate (as in chelyids and Pelusios), the first neural
especially being very elongate, there is greater probability of an
unusual condition of the horny scutes. Palaeaspis is probably a valid
genus.
Fig. 2. Emys laevis Bell. Dorsal view of type shell (after Owen and Bell,
reconstructed). 1/3 natural size.
Fortunately, as Lydekker and Boulenger were able to state, further
preparation of the type of F. conybearii after Owen’s description of it
revealed the presence of small lateral mesoplastral bones as in
Podocnemis and Pelomedusa. The existence of these bones I have been
able to verify on the type specimen. The form is therefore a pelome-
dusid, not a chelyid.
The case of Palaeaspis is instructive. On the characters of the
dorsal shell it was entirely plausible for Gray to place this form as a
chelyid. The only other known example of a transversely divided
first vertebral! occurs in a chelyid genus. The elongation of the first
vertebral and of that region of the bony shell is rather character-
istieally chelyid (though occurring also in Pelusios). If it were not for
1 I prefer to interpret the situation in both Hydromedusa and Palaeaspis in this rather than in
the conventional fashion.
4 BREVIORA NO. 32
the later discovered typically pelomedusine mesoplastra, this genus
would still be regarded as chelyid and thus would, it seems likely, be
used as such in zoogeographical speculation. Palacaspis may. thus
serve as a special caution against the use of imperfect fossils as critical
zoogeographical data.
.
e
Fig. 3. Emys delabechei Bell. Dorsal view of type shell (after Owen and
Bell). 1/5 natural size.
Comparison of the types of Emys conybearti Owen, Emys laevis Bell
(B.M.N.H. 37209) (Fig. 2), Emys delabechei Bell (Geological Survey
of Great Britain No. 6634) (Fig. 3), and a comparison of these with
the figure of Platemys bowerbanki Owen (Fig. 4) (the type of which
appears to be lost) leads to the belief that all the pelomedusids of the
London Clay are synonymous. Emys laevis has already been syn-
onymized by Lydekker and Boulenger with Platemys bowerbanki
under the name Podocnemis bowerbanki. They have also synonymized,
as already mentioned, Emys conybearvi with Emys delabechei, again as a
species of Podocnemis. Only the type of Emys conybearit approaches
completeness, but all four specimens, though differing in size, are
1954 FOSSILS AND DISTRIBUTION OF CHELYID TURTLES 5
congruent in every other determinable aspect, and unfortunately in
none of them do we know the gular-intergular pattern so important
in pelomedusine taxonomy. Emys laevis in which a portion of the first
neural is present shows that the latter is as strikingly elongate as in
B
Fig. 4. Platemys bowerbanki Owen. A. Dorsal view of type shell. B. Ventral
view of type shell. (Both after Owen and Bell.) 1/4 natural size.
Emys conybearti. While Emys laevis does not exhibit the transversely
divided first vertebral of Emys conybearii, the transverse sulcus may
in this smaller and younger specimen have been placed more anteri-
orly, on that portion of the first neural which is not preserved (see
Fig. 2). Alternatively, the transverse division of the first vertebral
in the type of P. conybearii may be an abnormality.
Assuming that the four types of Owen and Bell represent a single
form, there is still one further complication. Gray in 1870 in a single
publication proposed three distinct generic names for three of the
four forms here regarded as synonymous: Palemys with the type
Emys laevis Bell (p. 51); Palaeochelys with the type Platemys bower-
banki (p. 64); and Palaeaspis with the type Emys conybearti (p. 73).
6 BREVIORA NO. 32
Palaeochelys Gray 1870 is preoccupied by Palaeochelys H. v. Meyer
1847, but the other two names appear to be available. Since neither
name has any sanction deriving from usage — both having been
universally ignored since their erection — I choose Palaeaspis as the
generic name of the pelomedusine of the London Clay, on the ground
that the type species on which this name is founded has as its type the
most complete and best specimen of the four here discussed.
The oldest specific name for any of the group is bowerbanki Owen
and the correct name for the single species involved will then be
Palaeaspis bowerbanki (Owen). The genus may be rediagnosed as
follows:
PALAEASPIS Gray 1870
Type. Emys conybearti Owen.
New diagnosis. A flattened pelomedusid resembling Podocnemis
expansa in shell characters and adult size but without a vertebral keel
and with a very elongate first neural (ca. 214 times as long as wide)
and with the first vertebral divided transversely to form two successive
scutes.
BIBLIOGRAPHY
Gray, J. E.
1870. Supplement to the Catalogue of Shield Reptiles in the Collection
of the British Museum. Part I. Testudinata. London. 120 pp.
LypEKKER, R., and G. A. BoULENGER
1887. Notes on Chelonia from the Purbeck, Wealden and London Clay.
Geol. Mag., ser. 3, vol. 4, pp. 270-275.
Owen, R.
1858. Monograph of the Fossil Reptilia of the London Clay ... Part I.
Supplement I. Chelonia (Emys). Pal. Soc. Monographs, 1858,
PDs i190:
Owen, R., and T. BELL
1849. Monograph of the Fossil Reptilia of the London Clay ... Part I.
Chelonia. Pal. Soc. Monographs, 1849, pp. 1-76.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, MASS. May 14, 1954 NUMBER 33
SYSTEMATIC AND OTHER NOTES ON
SOME OF THE SMALLER SPECIES OF
THE ANT GENUS RHYTIDOPONERA MAYR
By Wiuuiam L. Brown, JR.
Museum of Comparative Zoology
Harvard University
The following notes concern a few species belonging to the
imopressa and metallica groups of the genus Rhytidoponera Mayr.
These forms have been included by most recent authors in
Chalcoponera Emery, but I have shown (Brown, 1953, Breviora,
No. 11, p. 2) that Emery’s group cannot be separated generically,
or even subgenerically, from Rhytidoponera. The present prob-
lem consists simply of recognizing the specific groupings in ques-
tion by means of their external characters, chiefly utilizing the
workers for study objects, and making the appropriate synonymie
reductions. The salient characters will be given briefly and
informally with a view toward rendering identification more
rapid and certain than has heretofore been possible.
Cabinet investigation is based upon material in the Museum
of Comparative Zoology at Harvard University, most of which
was gathered in Australia by Prof. W. M. Wheeler, or through
his correspondents and contemporary colleagues, and by myself
during the course of a recent year-and-a-half stay in Australia
under the auspices of the United States Educational (Ful-
bright) Foundation and of the Parker Fellowship Fund of
Harvard University.
Other papers on various groups of Rhytidoponera, intended
to follow this pattern, are under preparation.
2 BREVIORA No. 33
7ROUP IMPRESSA
This closely interrelated series of forms comprises R. ampressa
(Mayr) and its erstwhile varieties with their synonyms. R.
aenescens Kmery of New Guinea may also belong in this group,
but no specimens have been examined for the present work. The
Australian fauna contains three species of the group as far as
known, all similar in general details of structure, and all of
similar size. The body is slightly larger than in most members
of the more common metallica group, and generally more slender.
The head is only slightly longer than broad, and the oceipital
lobes or ‘‘ears’’ are much less perfectly developed than in
metallica, so that in direct dorsal cephalic (‘‘full faece’’) view,
the oceipital border appears approximately straight or even
feebly convex. In lateral view, the weakly developed occipital
lobes are seen as lamellate flanges, the sharpness of curvature of
which affords a good species eriterion (see below). Most char-
acteristic of the group is the gently but distinctly constricted
or ‘‘saddled’’ alitrunk as seen from the side. This formation
is due to a coneavity or gradually rounded stepping in the
region of the posterior mesonotum and anterior propodeal dorsum.
The legs and antennae are longer and more slender than in
metallica, in keeping with the more rapid locomotion of the
imopressa 2roup species.
Distribution, as known, runs from New Guinea down along
the mountain chains paralleling the eastern Australian coast.
The normal habitat is rainforest or, in the southern part of the
range in Victoria, in moist or wet selerophyll (euealypt-domi-
nated) forest. The nests are usually built in rotten logs or sticks
lying on the forest floor, but in areas where rotten wood is not
available, R. chalybaea at least will nest directly in the soil or
beneath rocks.
When the nest is breached, the workers seatter rapidly and
hide; some workers will feign death, but not as readily nor for
so long a time as in the ease of metallica. The colonies are small,
generally containing 30 to 250 workers and one dealate queen,
but mature nests will often produce large broods of winged males
and females. Sexual brood production oceurs during the end
of the rainy season, and circumstantial evidence indicates that
a normal nuptial flight commonly occurs during the first part of
1954 . ANT GENUS RHYTIDOPONERA MAYR 3
the dry season, even though rainy seasons are virtually opposite
in the northern and southern parts of Australia. Isolated nest-
founding queens are found in conditions resembling those of
the usual ant nest-formation pattern throughout the respective
dry seasons in the different parts of the range. The impressa
eroup species, along with just a few other forms (probably
including R. aspera and R. croesus), are exceptional among
Rhytidoponera groups in the ‘‘normality’’ of their production of
both sexual forms uninidally and with apparently clear-cut perio-
dicity, and their method of nest foundation through normal,
fertile, dealate females. Most other species of the genus appear to
deviate from this pattern considerably, and in these forms, whieh
often seem to lack entirely a distinguishable female caste, the
method of nest foundation is still unknown.
In studying fairly abundant material of this group from a wide
range of localities, [ could distinguish only three species in the
continental Australian fauna, one of which also occurs in New
Guinea. The three Australian species, at least from the present
material, appear to replace one another from north to south
in the narrow eastern ‘‘wet strip’’ with very slight geographical
overlap. Applying the prior names for these populations, we find
R. purpurea Emery in New Guinea and in the rainforests of the
Cairns-Atherton Tableland area of North Queensland, R. v-
pressa (Mayr) beginning on the southern part of the Atherton
Tableland and occurring at various intermediate points south-
ward to the vicinity of Brisbane, in southeastern Queensland,
and R. chalybaea Emery, starting probably in the region just
north of Brisbane on the mountaintops and ranging south at
least to Melbourne in south central Vietoria. Though the ehar-
acters are rather slight for the separation of purpurea from
impressa, they appear to be discontinuous together, and I have
little doubt that they delimit two distinct species.
R. chalybaea, on the other hand, seems to be distinguishable
from impressa only on the basis of coloration, and a rather ten-
uous and eabinet-labile metallic coloration at that. At the present
writing, the possibility still exists that these two forms are mere
geographical variants of the same species, although at Montville,
in the Blackall Range, about 50 or 60 miles north of Brisbane,
typical blue chalybaea were found nesting under rocks on the
4 BREVIORA No. 33
cleared top of the ridge, while only a few hundred feet below
in the rainforest, I consistently took only the brown typical
impressa. One series, however, coming from the environs of
Brisbane, showed the brown color of impressa with just the
feeblest hint, at high magnifications, of bluish metallescence. I
do not attach special significance to this series because it is an
old collection and may have undergone a change in the structural
and pigmental coloration. All collections from the small remain-
ing part of southeastern Queensland to the south of Brisbane
have proven to be of the typical blue chalybaea, and I can testify
that the population on Tambourine Mountain, only some 40 miles
south of Brisbane, is uniformly of blue color. It is obvious that
the populations inhabiting the scattered rainforest patches close
to Brisbane require further study in order to settle this question
finally, and it is fortunate that these are so readily accessible.
RwYTIDOPONERA CHALYBAEA Emery. New status
Rhytidoponera impressa var. chalybaea Emery, 1901, Ann Soc. Ent. Belg.,
45: 51, worker. Type locality: New South Wales.
Ectatomma (Rhytidoponera) cyrus Forel, 1910, Rev. Suisse. Zool., 18: 13,
worker, female. Type locality: Ballina, New South Wales, (W. W.
Froggatt). New Synonymy.
Rhytidoponera (Chalcoponera) cyrus, Emery, 1912, Deutsch, ent. Zeitschr.,
p. 81, discussion of worker characters and relationships.
Rhytidoponera (Chalcoponera) aspera var. scabrior Crawley, 1925, Ann.
Mag. Nat. Hist., (9) 16: 590, worker. Type locality: Lismore, New
South Wales (Deuquet). New Synonomy.
This, the southern representative of the impressa group in
Australia, is metallic blue or purplish-blue in color, with the
gaster matching the head and alitrunk, but usually slightly
duller. Legs and antennae light to sordid ferruginous, contrast-
ing strongly with the body color, especially so in life. The pat-
tern of fine striolation on the second gastric (second postpetiolar )
segment varies widely in direction even in uninidal series. Occip-
ital lobe or flange seen from the side narrowly rounded, forming a
near right angle, as contrasted with the more broadly rounded
and lower flange of purpurea; there is, however, some slight
variation in the flange of chalybaca, and rare specimens isolated
from their nest series may approach the condition seen in pur-
1954 ANT GENUS RHYTIDOPONERA MAYR 5
purea. Seen from the side, the anterior and dorsal faces of the
petiolar node usually meet in a blunt angle, though here again
occasional specimens may approach the more sharply angulate
condition of purpurea. In differentiating from the very similar-
appearing purpurea, fresh series are most easily told by the
different coloration and the lateral occipital lobe conformation.
Even old cabinet specimens are readily separable when one gets
an eye for these characters.
Emery (1912, loc. cit.) demonstrated the true relationship of
Forel’s cyrus, originally falsely compared, but he did not carry
his observations to the formal synonymy they indicate. From
Emery’s remarks, it appears that the cyrus cotype he had from
Forel was only a small worker of chalybaea with the usual
characteristics of such specimens. A cotype of Crawley’s variety
scabrior in the Museum of Comparative Zoology agrees well with
the remainder of the chalybaea series at my disposal; Crawley
knew this group so poorly that he attached his variety to an
entirely different species with similar coloration, R. aspera
(Roger), which is discussed farther below.
Localities for material in Museum of Comparative Zoology :
VICTORIA: Ferntree Gully, under stones (C. Barrett, W.
Brown). Slope of Warburton Range immediately above War-
burton, ca. 400 M., tree-fern gully remnant, in rotten log
(Brown). Millgrove, in rotten logs (F. E. Wilson, Brown).
NEW SOUTH WALES: Kurrajong (W. W. Froggatt). Na-
tional Park (W. M. Wheeler, A. M. Lea). Lismore; Armidale
(C. F. Deuquet). Coff’s Harbour, tree-fern gully near coast,
rotten log (Brown). Hornsby; Sutherland (Wheeler). Dorrigo
(W. Heron). QUEENSLAND: National Park, MacPherson
Range (H. Hacker). Toowong (Wheeler). ‘‘ Brisbane’’ (Whee-
ler). Stanthorpe (E. Sutton). Tambourine Mt., rotten logs in
rainforest, ca. 400 M.; Montville, Blackall Range, ca. 500 M.,
under stones on ridge top, grassy cleared rainforest area
(Brown). Winged forms are in the nest in September and
October in Victoria, in May in the Blackall Range.
XHYTIDOPONERA IMPRESSA (Mayr)
Ectatomma impressum Mayr, 1876, Jour. Mus. Godeffroy, 12: 92, worker,
female. Type locality: Gayndah, Queensland.
6 BREVIORA No. 33
Rhytidoponera (Chalcoponera) impressa, Emery, 1912, Deutsch. ent.
Zeitschr., p. 77, worker, in key.
When fully colored, this distinctive species is deep reddish
brown to chocolate brown, with conspicuous light reddish-fer-
ruginous legs and antennae; a faint trace of coppery reflection
may oceasionally be present on the dorsal surfaces of the body,
but this is negligible in the material I have seen. I ean find no
other characteristics but the general body color to distinguish
this form from chalybaea (q.v.). The color and the rectangularly
reunded lateral occipital lobe in lateral view will serve to sep-
arate impressa from purpurea, even where the two forms occur
only a few miles apart at Millaa Millaa and Malanda on the
Atherton Tableland. During several days’ collecting at Malanda,
I found only purpurea there, whereas at Millaa Millaa, which
has not been systematically collected for the genus, only im-
pressa has so far been found.
The Millaa Millaa series was taken by P. J. Darlington, while
all of my own collections were made in rainforest on the slopes
and at the foot of the Blackall Range, where this species is com-
mon in rotten logs and sticks lying on the forest floor, often in ex-
ceedingly dense shade. At Kondalilla Falls and along the Obi-Obi
River, I found something like twenty nests of impressa, about
half of which had numerous males and females together in the
winged condition and mostly fully colored (May, 1951). Dealate
nest. queens were found easily in most colonies opened, one to a
colony. No single isolated females were found during this
period.
Probably Brisbane is just about at the southern hmit of this
species, which must occur in most suitable rainforest patches and
perhaps also in gallery forest along almost the whole of the east
Queensland ‘‘wet strip’’ to as far north as the southern edge
of the Atherton Tableland. Millaa Millaa is about 2500 feet
(760 M.) above sea level, and is rainforest country on rich
voleanic soil, much of which has been cleared to grassland for
grazing. Malanda, a little farther north and occupied by BR.
purpurea, is similar country lying at an altitude of about 2400
feet. No barriers of any consequence fall between the two locali-
ties.
~)
1954 ANT GENUS RHYTIDOPONERA MAYR
RHYTIDOPONERA PURPUREA (Emery). New status
Ectatomma impressum var. purpureum Emery, 1887, Ann. Mus. Civ. Stor.
Nat. Genova, (2) 5: 444, worker, female. Type locality: Hatam, New
Guinea (Beceari).
Ectatomma (Rhytidoponera) impressum var. splendidum Forel, 1910, Rev.
Suisse Zool., 18: 12, worker. Type locality: Kuranda, near Cairns,
Queensland (R. Turner). New Synonymy.
This species differs from R. chalybaea, to which it is at first
sight very similar, in the very narrow, broadly rounded lateral
occipital flange or lobe as seen in lateral view. The difference
is Slight, and can best be appreciated through comparison of series
of both forms. The angle between the anterior and dorsal faces
of the petiolar node is usually well-marked, and even sharp, but
there is enough variation in this character among all impressa
group species to render it of little discriminatory value.
The color of the head and alitrunk varies somewhat by nest
series, but in fresh specimens is basically a rich, shghtly reddish
purple, usually with elusive greenish or bluish highlights. The
gaster has a slightly contrasting metallic coloration of a pre-
dominantly bluish-green hue, often somewhat dull compared to
the purple of head and alitrunk; the gastric color seems constant
and affords a good distinguishing character even in most old
cabinet specimens. The legs and antennae are considerably
darker than in chalybaea, and appear more nearly black,
especially to the naked eye in living specimens, though tending
to fade to light brownish in old cabinet specimens: the size
averages slightly larger.
The above description is taken from long series of specimens
obtained at Kuranda, Queensland, in rotten logs in rainforest
by myself, as well as from older series from the same locality
taken by Wheeler and T. Greaves, and from the Cairns district
by F. P. Dodd. The species is also common along the Black
Mountain logging road, on the western side of the MacAlister
Range north of Kuranda. I found it somewhat less common at
Malanda, in rainforest at 2400 feet (730 M.) altitude on the
Atherton Tableland, also in rotten logs, although one isolated
nest-founding female was taken here in the ‘‘ peat’® about the base
of an epiphytic fern growing on the trunk of a rainforest tree
about seven feet above the ground. I visited these localities
8 BREVIORA No. 33
during October and the beginning of November, 1950, which is at
the end of the dry season in this district. During this time, many
dealate females were found isolated with small larval broods,
and others were found as queens of established nests, but no
winged forms or sexual pupae in obvious stages were seen;
Wheeler, however, took the winged forms at Kuranda in October.
I have seen one specimen from New Guinea, collected in the
Rawlinson Mountains and sent by Dr. H. Kutter under the name
‘‘var. purpurea Emery.’’ While an old and somewhat discolored
specimen, this example does not seem to possess any notable
features by which it can be separated from the Kuranda series
of splendida. I am expressing this lack of difference by synony-
mizing Forel’s variety, but I hope that a future study of further
New Guinea material will test the synonymy thoroughly. R.
purpurea should be expected to occur in the seattered rainforest
patches following the mountains up the eastern side of Cape
York Peninsula.
GROUP METALLICA
The group of species related to R. metallica (Fred. Smith) is
in considerable taxonomic confusion, due chiefly to the high
degree of geographie variability shown by R. metallica itself over
a wide range and to the circumstance that certain very similar
species have been included in metallica as varieties. Most of the
forms have been described two or three times under different
names and with the briefest and often most irrelevant of charac-
terizations. Such descriptions, which variety-deseribers econ-
sidered it their privilege to publish in keeping with the humble
status of the variety, are the bane of ant taxonomy, and they
show little reflection of the true taxonomic situation. During the
1930’s, Clark added to the complications aftlicting this group by
describing a number of doubtful species based chiefly on variable
characters, such as color of metallescence, but apparently without
considering sufficiently the degree of variation even in his limited
series. In a subsequent paper, I shall try to unravel some of the
more difficult complexes in this group to the extent of the material
currently available to me, but at this time I am ready to deal with
two of the less common species with relatively uncomplicated and
obvious synonymy and relationships,
1954 ANT GENUS RHYTIDOPONERA MAYR 9
The metallica group species are among the smallest Rhytido-
ponera, and are distinguished by having an evenly convex ali-
truneal profile and prominent posterolateral occipital lobes, or
‘“ears,’’ which often cause the occipital border to be broadly and
rather deeply excavate as seen from direct dorsal cephalic view.
Coloration is often brilliantly metallic blue, green or purple, but
some species lack metallescence completely, and in others it is
very variable. Antennal scrobes are lacking or merely suggested,
and are not ever so strong as in the victoriae complex ; however,
the victoriae complex is hardly more than a subgroup of the
metallica group. The petiolar node is thick but erect, with steep
anterior and posterior faces. The treatments of R. aspera and
R. croesus do not require further general comment.
RHYTIDOPONERA ASPERA (Roger)
Ponera metallica var. aspera Roger, 1860, Berlin. ent. Zeitschr., 4: 308,
worker. Type locality: Australia.
Rhytidoponera (Chalcoponera) aspera, Emery, 1912, Deutsch. ent. Zeitschr.,
p. 77, worker, in key.
Rhytidoponera (Chalcoponera) arnoldi Forel, 1915, Arkiv. f. Zool., 9 (16):
14, worker. Type locality: Healesville, Victoria (H. A. Arnold). New
Synonymy.
This ant is similar to metallica, but is larger and stouter, and
is of a brillant metallic green color (blue or purplish in some old
dried specimens) with contrasting reddish ferruginous legs, an-
tennae and mandibles. The second gastric (second postpetiolar )
segment is largely smooth and shining, but has some fine, super-
ficial, diverging oblique striae anteriorly, mainly toward the sides
above. Forel’s characterization of arnoldi fits very well with the
specimens at hand, and these in turn agree with the essentials
of Roger’s description. Fortunately, the color, size and details of
gastric sculpture leave very little doubt about the identity of
this species and its synonym, as it is the only species combining
all these characteristics in its genus.
This species is not common in collections, I find, and I have
myself met with it only once in the field, at Upper Ferntree Gully
Station, Victoria, where strays were found running up the trunk
of a large manna gum by the station driveway. These workers
were very conspicuous, but I did not succeed in finding the nest,
10 BREVIORA No. 33
nor did I see them at the same place on other visits. Small series
or strays have also come to me from various Victorian localities :
Portland (H. W. Davey) ; Belgrave (F. E. Wilson) ; Melbourne
(?) (H. Edwards), and I have a pin labelled ‘‘Fernshaw,’’ a
locality unknown to me and possibly outside Victoria. A single
dealate queen was found on the same pin with two R. chalybaea
dealates, collected by H. Hacker at National Park, MacPherson
Ranges, in southeastern Queensland. If this record is eorreet, the
species must range through the moister districts of eastern New
South Wales, though no records are available from that state.
The locality records indicate a habitat in eucalypt forest of the
intermediate to high rainfall types. I have seen a single specimen
collected at Auckland, New Zealand, from poles originating at
an unknown Australian port, but the species appears not to have
colonized New Zealand.
tHYTIDOPONERA CROESUS Emery
Rhytidoponera croesus Emery, 1901, Ann. Ent. Soe. Belg., 45: 50, worker.
Type locality: New South Wales.
Rhytidoponera (Chalcoponera) croesus Emery, 1912, Deutsch. ent. Zeitschr.,
pp. 77, im key, and 81, worker.
Rhytidoponera (Chalcoponera) fastuosa Santschi, 1916, Bull. Soe. ent.
France, p. 174, all eastes. Type locality: Australia (Duechaussoy),
probably New South Wales. New Synonymy.
Chalcoponera victoria (sic) var. andrei Wheeler and Chapman, 1925,
Philippine Jour. Sei., 28: 60, worker. Type locality: cited in error as
from the mountains of Victoria; recte Dorrigo, New South Wales (W.
Heron). New Synonymy.
R. croesus workers and females are recognizable by means of
the low, thick, anteriorly rounded petiolar node, the posterodorsal
border of which slightly overhangs the posterior face. The color
is also distinctive, being basically bright ferruginous, but over-
lain more or less completely and heavily with deep, brilliant
purple metallescence. In the extreme form, represented by what
Santschi deseribed as fastwosa, most of the head, alitrunk, petiole
and gaster is densely purple in color, whereas intermediates like
the types of the original croesus may have only the alitrunk
thoroughly saturated with purple, while the head, the gaster, or
both remain more or less pure ferruginous like the appendages.
Single nests, excepting tenerals, are usually fairly uniform
1954 ANT GENUS RHYTIDOPONERA MAYR 11
through the series, but a collection containing series from several
nests shows all degrees of intergradation, even from close locali-
ties within the restricted distribution in eastern New South
Wales and southeastern Queensland. Santschi’s description is
confused by his comparison with croesus, which species he credits
to Forel as author. It seems that his comparison is really made
against cyrus Forel (chalybaea Emery, see above), and in this
connection his description makes more sense. The variety
andre: was described due to a highly improbable series of errors,
involving mislabelling of the specimens and misreading even of
these labels. Dr. J. W. Chapman has reviewed this situation and
produced three worker cotypes. These are faded purple speci-
mens, and their character and kind of mounting shows that they
are part of an old series Wheeler had received from Dorrigo,
New South Wales, from W. Heron, the collector. ‘* Victoriae’’
of someone’s temporary label was misread as *‘ Victoria,’’ but no
specimens of this species have yet been recorded from Victoria.
The supposed introduction into Luzon must also be regarded as a
doubtful record, since the relevant specimen is apparently
missing.
Material in Museum of Comparative Zoology: from eastern
New South Wales: Bulli Pass (W. M. Wheeler). Dorrigo (W.
Heron). Lismore (C. F. Deuquet). Queensland: Tambourine
Mt. (A. M. Lea; Brown) running on rotten tree trunk in dense
rainforest at 1100 feet. Stanthorpe (E. Sutton). National Park,
MacPherson Range (H. Hacker). &. croesus is apparently re-
stricted to the wet subtropical forests of eastern New South Wales
and extreme southeastern Queensland.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. JuLY 20, 1954 NuMBER 34
A REVIEW OF THE COXALIS GROUP OF
THE ANT GENUS STICTOPONERA MAYR
By Wiriam L. Brown, JR.
Museum of Comparative Zoology
Harvard University
The genus Stictoponera is a chiefly Oriental and Papuasian
assemblage of typical ectatommines. In my revision of the tribe
Ectatommini, to be published elsewhere, I have enlarged Sticto-
ponera to include the species from the same general region for-
merly placed in Wheeleripone Mann and Rhopalopone Emery.
With the new composition, Stictoponera includes three or four
rather vague and intergradient species-groups, of which the larg-
est and most familiar comprises species with a low, paniform
petiolar node and a distinet tooth on the dorsum of the posterior
coxa. These species are the ‘‘typical’’ members of Stictoponera
related to the genotype, S. coxalis (Roger). The species formerly
called Wheeleripone albiclava Mann perhaps belongs in the
coxalis group, but it is aberrant in sculptural detail and is some-
what intermediate to other groups in the shape of the node; it
will not be discussed here.
The coxalis group is most in need of revision at the species
level, and the notes offered here amount to a revision of those
species already described of which recognizable material is now
available to me. The majority of this material is in the Museum
of Comparative Zoology, Harvard University, and is due mainly
to the fine collecting of Dr. J. W. Chapman in the Philippines
and the late Eric Mjoberg in northern Borneo. Dr. Chapman also
has furnished considerable ecological information and other aid
concerning many of the species, and it was he who inspired the
present work and encouraged it continuously.
P) BREVIORA No. 34
Since a key to the species is not yet possible because of doubts
about several species not seen by myself, I can offer only the
outline of characters as follows:
A. Second postpetiolar (IV abdominal) segment coarsely
longitudinally costate, at least on the basal half.
1. Eyes placed in the posterior quarter of the sides of the
head: S. posteropsis Gregg.
2. Eyes placed between the middle point and the posterior
quarter of the sides of the head: S. coxalis (Roger), S.
costata Emery, S. biroi Emery.
B. Second postpetiolar (IV abdominal) segment not costate.
1. Eyes placed anterior to middle of sides of head (ac-
cording to original deseriptions) : S. crassicornis Forel,
S. spiralis Karawajew.
2. Eyes placed at or slightly behind the middle of the sides
of the head: S. bicolor Emery, S. binghamu Forel, S.
laevior Forel, S. menadensis Mayr.
In addition to the species listed above, with their synonyms,
I have also seen two apparently undescribed species of group
B,2.
STICTOPONERA LAEVIOR Forel
Ectatomma (Stictoponera) laevius Forel, 1905, Mitt. Naturh. Mus., Ham-
burg, 22: 7, worker. Type locality: Tjibodas, Java.
Stictoponera laevior var, avia Forel, 1912, Notes Mus. Leyden, 34: 96, all
eastes. Type locality: Nongkodjadjar, Java. New Synonymy.
A small, slender shining species with sparse foveation. The
species was described from a unique, obviously teneral; var. avia
refers to fully pigmented specimens. I have seen one worker
from Poentjak, Java, no collector cited.
STICTOPONERA MENADENSIS Mayr
Ectatomma (Stictoponera) menadensis Mayr, 1887, Verh. zool.-bot. Ges.
Wien, 37: 539, nota, worker.
Stictoponera stylata Menozzi, 1925, Philippine Jour. Sci., 28: 440, pl. 1, figs.
a, b, worker. New Synonymy.
Stictoponera menadensis var. obscura Santschi, 1932, Mem. Mus. Hist. Nat.
Belg., (hors série) 4 (5): 11, worker. New Synonymy.
This species is of medium size, shining, with full adult color
dark reddish-brown to piceous, gaster black. Mesonotum with a
1954 ANT GENUS STICTOPONERA MAYR 3
smooth longitudinal median strip free of foveation, extending
in some specimens for a short distance onto the posterior portion
of the pronotum; the latter otherwise always rather closely
foveate. This species is apparently the common lowland and
foothill Stictoponera in the East Indies and the Philippines. It
oceurs on Java and Sumatra together with S. bicolor, a very
closely related species, but no intergrades between the two have
yet been reported from these localities. I have seen no specimens
of menadensis from mainland Asian loealities, and records from
the mainland are searce in the literature. From the present data,
S. menadensis appears to be a peripherally distributed form in
the process of being replaced by S. bicolor, the latter spreading
from southeastern Asia outwards into the archipelagic regions.
The types of S. menadensis and the var. obscura come from
Celebes, and Santschi’s deseription mentions no characters dif-
ferentiating his variety from normal menadensis populations.
The type of S. stylata came from Mt. Makiling, Luzon, and is
present in the collection of Dr. J. W. Chapman. Unfortunately,
Dr. Chapman’s collection was somewhat damaged during the
period when it was hidden in the mountains of Negros while the
owner was interned in prison camp during the late war, and
some of the specimens of Stictoponera became separated from
their labels, among them the stylata type. Dr. Chapman and I
have, however, segregated a small group of specimens in the
collection by elimination of possibilities, and some specimen of
this group is the type.of stylata. Since all the specimens in this
group are clearcut menadensis, and since menadensis is the com-
mon species on Mt. Makiling, we concluded that the above syn-
onymy is required. Menozzi’s description offers no characters
in opposition to this decision, and the description itself could be
the basis of the synonymy. Obviously, Menozzi did not have
a clear idea of the identity of the typical menadensis, and merely
described it under a new name. In addition to the Mt. Makiling
record, the collections of Dr. Chapman and the Museum of
Comparative Zoology contain series from the following Philippine
localities: Sarai, Paete, Laguna (Luzon) (R. C. MeGregor). Mt.
Manapla, Negros Occidental (F. del Rosario); Lake behind
Dumaguete; Cuernos Mts., several collections, 1800 to about
3600 feet, Negros Oriental (J. W. Chapman). Momungan and
4 BREVIORA No. 34
Lanao, Mindanao (Chapman and D. Empeso). Los Banos,
Luzon (F. X. Williams). Samar I. (McGregor et al). Kolam-
bugan, Mindanao (C. 8. Banks). There are also series from
Borneo: Mt. Tibang, 1400 M., and Pajan (E. Mjoberg). Ku-
ching (Hewitt). Sumatra: Wai Lima, Lampongs (Karny). In the
Philippines, nests are built in rotten logs and tree fern stems and
under moss on rocks. Dr. Chapman informs me that this, the
most common Stictoponera in the Philippines, is usually seen
running over foliage or resting in the axils of large leaves in wet
foothill ravines.
STICTOPONERA BICOLOR Emery
Ectatomma (Stictoponera) bicolor Emery, 1889, Ann. Mus. Civ. Stor. Nat.
Genova, 27: 493-494, worker.
Ectatomma (Stictoponera) bicolor var. minor Forel, 1900, Jour. Bombay
Nat. Hist. Soe., 13: 316, worker.
Stictoponera menadensis subsp. minor, Brown, 1948, Psyche, 54: 264,
(teneral) worker.
Stictoponera bicolor, Brown, 1950, Wasmann Jour. Biol., 8: 245-246, worker,
synonymization of var. minor.
Since this form was described, not a few authors, including
myself, have been confused by it one way or another. It is very
close to S. menadensis, and has the same head shape, with prom-
inently projecting posterior occipital lobes or ‘‘ears’’ and a cor-
responding, deeply concave, border between them as the head is
viewed in full face. The eyes are rather large and situated well
posterior to the middle of the sides of the head. The size is as in
menadensis, with some series averaging a trifle smaller, but the
color of the alitrunk is hghter and brighter, varying shades of
orange-ferruginous. The head varies from about the same color
as the alitrunk to piceous, the darker color being more common.
The median smooth strip of the menadensis mesonotum is re-
placed in bicolor by fine, indefinite, more or less opaque longi-
tudinal rugulation, and the sculpture throughout is usually
stronger, closer and less shining. S. bicolor may or may not
possess minute propodeal denticles; these appear to be an allo-
metric character of the sort that grades through within and be-
tween series. The tenerals are straw-colored, and appear very
different from fully-pigmented individuals in the same nest
series.
1954 ANT GENUS STICTOPONERA MAYR 5
Like S. menadensis, the present species has often been mis-
identified. It is still possible, of course, that menadensis and
bicolor are mere geographical color representatives of one species,
but from the present material they can still be separated. The
record of bicolor from the Philippines is based on a male of un-
certain species; no authentie bicolor records are yet known for
these islands.
I have reviewed material from the following localities: Indo-
ehina: Coxan and Dong Mo (F. Silvestri). Hainan I.: Dwa Bi;
Ta Han; Loi Molia; Nodoa (J. L. Gressitt). Sumatra: Wai Lima,
Lampongs (Karny). Java: Pemalang (lL. G. E. Kalshoven).
Hong Kong: no further locality (Ris). Emery described this
species from material taken in various Burmese localities, both
upecountry and in Tenasserim.
STICTOPONERA BINGHAMII Forel
Ectatomma (Stictoponera) binghamii Forel, 1900, Jour. Bombay Nat. Hist.
Soe., 13: 137, worker, female. Type locality: Burma [Pegu Yoma?].
Stictoponera borneensis Emery, 1901, Ann. Mus. Civ. Stor. Nat. Genova, 40:
662, nota, worker. Type locality: Sarawak. New Synonymy.
This species resembles S. menadensis, but lacks the prominent
lateral occipital ears and has a normal full adult coloration of
rich ferruginous red. The eyes are at or close behind the middle
of the sides of the head, and the posterior occipital border is
straight to gently concave in different specimens and according
to the view. Small teeth are present on the propodeum of most
individuals. The insect is rather strongly shining, the foveation
tending to be less dense than in menadensis, and the middle of
the pronotum has a variable smooth, shining area free of foveae.
The petiolar node seen from above is approximately as broad as
long, length being favored in larger specimens. The second post-
petiolar segment is very smooth and shining, foveation very in-
distinct and shallow and virtually confined to the sides. The
antennal funiculi are rather thick, the median segments broader
than long even in the largest specimens.
At first I had separated binghamiu from borneensis, though
the two were obviously closely related, but I now find that
specimens referable to both were taken by Dr. Chapman on the
same date at the same locality near his camp in the Cuernos
6 BREVIORA No. 34
Mts., Negros Oriental, Philippines; these probably represent a
single nest series, and in any ease, it is apparent from a study of
the full series, including those in Dr. Chapman’s personal reserve
collection, that the large and small forms are only allometrie
extremes of one form. I have also seen material referable to
the same species from Tutu River, North Borneo (Mjéberg) and
a specimen from ‘‘Pedada-B., Lampongs, Sumatra,’’?’ unknown
collector, and I feel sure that Forel’s Burmese and Emery’s Bor-
nean types are merely the small and large extremes of one
variable species, though I have not examined type material. Dr.
Chapman tells me that he found this species nesting in rotten
logs in ravines in forest.
STICTOPONERA CRASSICORNIS Forel New status
Ectatomma (Stictoponera) binghami subsp. crassicornis Forel, 1912, Zool.
Jahrb. Syst., (suppl.J 15: 51, worker. Type locality: Indrapura, Su-
matra.
Forel states that the eyes are anterior to the middle of the
sides of the head, which if true would separate this form from
binghamu very decisively. The other characters cited, however,
indicate considerable similarity, and Forel may well have been
mistaken about the eyes. The description of S. spiralis, the next
species following, also claims a similar position for the eyes.
Among all the series available to me, I have seen no Stictoponera
specimens with the eyes in front of the middle of the sides of
the head. In any ease, it is very unlikely that crassicormis can be
a race of S. binghami, since the known distribution of the latter
straddles the Sumatran type locality of crassicornis. Provisional
specific rank is indicated for crassicornis until the type can be
re-examined,
STICTOPONERA SPIRALIS Karawajew
Stictoponera spiralis Karawajew, 1925, Konowia, 4: 79-81, worker.
This species, also described from Sumatra, reads very much
like NS. crassicornis in what seem to be the significant features, and
it is possible that the two names are synonymous.
STICTOPONERA BIROI Emery
Stictoponera birdi Emery, 1902, Term. Fiizetek, 25: 154, worker.
S. biroi, the only species of the genus so far recorded from
New Guinea, appears to be quite distinct from the Indomalayan
forms on the basis of its original description.
1954 ANT GENUS STICTOPONERA MAYR t
STICTOPONERA POSTEROPSIS Gregg
Stictoponera posteropsis Gregg, 1952, Psyche, 58: 77-80, figs. 1, 3a, 3ce,
female.
This very aberrant species has large eyes situated almost at the
extremes of the posteriorly salient occipital corners. The type,
a dealate female, came from Sumatra, but I have seen a few
additional worker specimens from the Cuernos Mts., Negros
Oriental (Dr. Chapman), taken in rotten logs in forest ravines.
Some of these workers are larger even than the female type,
and their color is dark piceous instead of ferruginous red.
STICTOPONERA COSTATA Emery
Ectatomma (Stictoponera) costatum Emery, 1889, Ann. Mus. Civ. Stor. Nat.
Genova, 27: 494, worker. Forel, 1900, Jour. Bombay Nat. Hist. Soe.,
13: 316, 317, worker. Bingham, 1903, Fauna Brit. India, Hym., 2:
83, worker.
Ponera rugosa Fr. Smith, 1857, Jour. Proce. Linn, Soc. London, Zool., 2:
66, worker; name preoccupied by Le Guillou, 1840. New Synonymy.
Stictoponera costata Emery, 1901, Ann. Mus. Civ. Stor. Nat. Genova, 40:
662, worker.
Stictoponera costata var. unicolor Forel, 1901, Rev. Suisse Zool., 9: 335,
worker, male. New Synonymy.
Stictoponera rugosa var. parva Forel, 1913, Zool. Jahrb. Syst., 36: 6, worker.
New Synonymy.
?Stictoponera costata var. simalurensis Forel, 1915, Tijdschr. v. Ent., 58:
23, worker. New Synonymy, with doubt.
Stictoponera costata var. pinealis Wheeler, 1929, Boll. Lab. Zool. Portici,
24: 31. New Synonymy.
Stictoponera wallacet Donisthorpe, 1932, Ann. Mag. Nat. Hist., (10) 10:
447, nom. pro Ponera rugosa Fr. Smith. New Synonymy.
The key reference to this form is that of Emery for 1901 (loc.
cit.) in which he cites variation in size among specimens from
Sumatra and Mentawei (the costata type locality is in Burma)
and suggests, but does not declare, the synonymy with Ponera
rugosa of Smith. He also cites differences between costata and
a type of coxalis he received from the Roger Collection in Berlin,
the latter differing ‘‘from costata in the much less coarse sculp-
ture of the whole body, and especially of the abdomen, and in the
more slender antennae, with the flagellar segments less thick, the
third to the fifth longer than thick (thicker than long in S.
costata).’’ I am able to confirm and amplify Emery’s differentia-
8 BREVIORA No. 34
tion of these two species, as will be seen below under S. cozalis.
In the series I refer to S. costata, stemming from ten localities
ranging from Malaya to Borneo and Mindanao, there is a large
amount of variation in size, color, and degree of development of
sculpture; the variation in sculpture of the first gastrie (post-
petiolar) segment is particularly notable. The prevailing color
in North Bornean specimens is rich ferruginous red, but certain
specimens from Mindanao are piceous, and the female type of
pinealis Wheeler, from Penang, is deep reddish with the gaster
black. The variety wncolor Forel, based largely on a Bornean
male, was never really differentiated from the types of costata.
The various color conditions are constant within some nests, but
not in others; and considering the long teneral period shown
by ants of this genus, the relative conspicuousness of tenerals
under the collector’s eye, and the chances of color changes in
preservative and cabinet, I can attach no great importance to
eolor by itself. In the present material, color, size, variation
in the shape of the lateral occipital ‘‘ears’’ or lobes, and sculpture
are discordant geographically, so that it seems not possible to
recognize objective geographical races based on these characters.
Emery mentions, as stated above, that certain Sumatran series
varied in size, and it seems lkely that Forel’s variants parva
and simalurensis are merely small costata. The Bornean speci-
mens show significant size variation even between relatively close
localities. Essentially, the sculpture of costata, except for the
very coarsely and regularly costate second gastric (second
postpetiolar or IV abdominal) segment, is in the form of large,
rather deep punctures, foveae or pits, with more or less distinet
and smooth, shining spaces between them. On the first gastric
segment, the punctures tend to be more elongate, and the spaces
between them, particularly toward the posterior edge of the
diseal surface, begin to form more or less definite longitudinal
ridges or costae. In some specimens, such as those from Mt.
Penrissen and Mt. Tibang, in Borneo, the spaces are broad and
shining, and the punctures definitely prevail over the rather
weakly suggested costae, while in others, such as most Bornean
specimens and the pinealis type, the costae are closer and sharper
on the first gastric segment, at least posteriorly, and the pune-
tures are closer together and more elongate. The sculpture of
1954 ANT GENUS STICTOPONERA MAYR 9
the remainder of the body follows that of the gaster more or less
closely, but differences are harder to see and describe. In spite
of these rather distinct differences between extremes, I find that
Borneo alone supphes a full set of intergrades which is enough
to obseure,any possible taxonomic split based on this character. It
therefore seems that Emery was correct in maintaining costata
as a single taxonomic unit despite the variation he saw. His
reasonable suggestion that F. Smith’s rugosa, from Sarawak,
equalled costata is also accepted here, and since the name rugosa
is preoccupied, and Donisthorpe’s nomen novum wallacer later
than costata, the name costata will remain in use.
I have studied series in the Museum of Comparative Zoology
from the following localities: North Borneo and Sarawak: Mt.
Penrissen; Mt. Tibang; 8. Slau; Brooketon; Pajan; 8S. Saranibo;
Baian River (E. Mjoberg). Mindanao, Lanao district: Maria
Christina Falls (J. W. Chapman). Momungan (D. Empeso)
Malaya: Penang Island (F. Silvestri), type of var. pinealis.
Sumatra: Langkat, E. Coast (Jourin).
STICTOPONERA COXALIS (Roger)
Ponera coxalis Roger, 1860, Berl. ent. Zeitschr., 4: 308, worker.
Ectatomma (Stictoponera) coxale Forel, 1900, Jour. Bombay Nat. Hist. Soe.,
13: 316, worker. Nec Bingham, 1903, Fauna Brit. India, Hym., 2:
84, fig. 44, worker.
Stictoponera coxalis Emery, 1901, Ann. Mus. Civ. Stor. Nat. Genova, 40:
662, worker.
Roger described this species very incompletely for modern
needs from specimens collected by Nietner in Ceylon. Emery’s
statement of the differences between cowxalis and costata has
already been translated above under S. costata. I have been able,
through the courtesy of Prof. M. Beier, of the Naturhistorisches
Museum, Vienna, to examine a worker (here designated as lecto-
type, and so labelled) and a dealate female from the type series
of coxalis, that Roger had early sent to Gustav Mayr. As Emery
mentions, the sculpture throughout is considerably finer than in
costata; it is also denser and consists more predominantly of
longitudinal costulation or coarse striation instead of the large,
predominating punctures of costata, although somewhat smaller
punctures are still clearly present and often conspicuous. Under
10 BREVIORA No. 34
lower magnifications, coralis tends to appear more opaque gen-
erally than does costata. On the first gastric segment, there are up
to two or three irregular, but rather close and fine longitudinal
costae for every one seen in the average costata specimen. Also,
as mentioned by Emery, the funiculi are notably more slender
in coxalis, with the third through the fifth (I would inelude the
sixth and possibly also the seventh) funicular segments slightly
longer than broad. The same segments are broader than long,
though somewhat variable in proportions, in the costata samples
I have seen.
I agree with Emery’s separation of the two species on the pres-
ent evidence ; obviously, however, the two are very closely related.
To my knowledge, S. coxalis remains known only from the type
series from Ceylon, though various authors have followed Bing-
ham in confusing this species with 8S. menadensis and S. bicolor.
When better collections are available from southern India, it may
be necessary to re-examine the relationship of coralis and costata.
Note on ‘‘Stictoponera sauteri’’
The name Stictoponera sauteri (Chapman and Capeo, 1952,
Check list of the ants of Asia, Monogr. Inst. Sci. Tech., Manila,
1: 30) is a combination proposed in error, and actually refers to
Ectomomyrmec sauteri, a species described earlier by Forel. This
species has no connection with Stictoponera; the Check List com-
bination was purely an unintentional clerical slip.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. JULY 28, 1954 NUMBER 35
NEW OR REDESCRIBED PELOMEDUSID
SKULLS FROM THE TERTIARY OF
AFRICA AND ASIA
(TESTUDINES, PELOMEDUSIDAE)
1. Dacquemys paleomorpha, new genus, new species from the
Lower Oligocene of the Favum, Eevpt.
C=) « De
By ERNEST WILLIAMS
In 1912 E. Daeque described a skull from the Lower Oligocene
of the Fayum of Egypt which he referred, with doubt, to Stereo-
genys libyca Andrews. The skull, which is the property of the
Staathches Museum fur Naturkunde in Stuttgart, has survived
the war. Thanks to the kindness of the authorities in Stuttgart,
and especially of Prof. F. Berckhemer and Dr. Karl Dietrich
Adam, I have been able to re-examine it.
Re-examination confirms the excellence and accuracy of
Daequeé’s figures, but it also leaves no doubt that he failed to
sufficiently appreciate the uniqueness of the specimen in certain
regards. It is unquestionably a new genus of pelomedusid, which
may be named and defined as follows:
DACQUEMYS, new genus
Diagnosis. A pelomedusid belonging to the Podoenemis group
within the family and possessing the enlarged ‘‘carotid canal’’
and basioccipital-quadrate contact characteristic of that group
but differing from the others of the group and from all other
members of the family in the extensive squamosal-parietal con-
tact.
Genotype. Dacquemys paleomorpha, new species.
The genotype is at present the only species.
bo
BREVIORA No. 35
DACQUEMYS PALEOMORPHA, hew species
Type. No. 12645 in the paleontological collection of the Staat-
liches Museum fur Naturkunde, Stuttgart, a skull without
mandible.
Type locality. Diueh, Fayum of Egypt.
Horizon. Lower Oligocene.
Specific diagnosis. With the characters of the genus.
Description. A moderately elongate skull (estimated condylo-
basal leneth 72mm.) of rather heptagonal shape with a com-
pletely covered temporal region and the orbits far forward,
these and the nares concealed in dorsal view. No forehead
eroove. Frontals enter orbit broadly. Interorbital width rela-
tively great. Well defined grooves bounding a nearly equilater-
ally triangular interparietal scute. Parietal scutes, well indicated
by grooves on the bone, large and meeting for a considerable
distance posterior to the interparietal scute. Jugal excluded from
parietal by the postorbital which is relatively large. Squamosals
and parietals in very broad contact. Temporal roof in conse-
quence carried posterior to the cavum tympani. Squamosal in
two planes: (1) a dorsal nearly horizontal arched surface joing
the parietal paramedially and bordering laterally on the quad-
rate; (2) a ventral nearly vertical surface posterior to the quad-
rate, bearing a well-marked groove for the depressor mandibuh,
and curving medially to meet the backward prong of the
opisthotie which, as typically in pelomedusids, projects somewhat
further posteriorly than the more dorsally placed backward
process of the squamosal. Quadrate fully closed behind the
columella. A narrow nearly horizontal opening in the cavum
tympani leading posteriorly into a posterodorsal squamosal
antrum. Below this is the tear-shaped columellar foramen
through which the columella dorsally and the eustachian tube
ventrally enter the middle ear. In front of the columellar open-
ine (on one side only) is a very shallow depression which may
represent the precolumellar fossa frequently present in pelo-
medusids and chelyids. The side of the temporal region ap-
parently deeply emarginate, and the quadratojugal displaced
dorsally. The orbits small, circular, far anterior, smaller than the
cavum tympani. The maxilla rather narrow vertically. Indica-
tions of grooves marking off a ‘‘subocular’’ scute sensu Sieben-
Se
1954 DACQUEMYS PALEOMORPHA
rock, 1902, such as is present in some living species of Podocnemis.
The premaxillae perhaps forming a hook; if so the hook eroded
in the specimen. Nares much broader than high. Two strone
ridges on the triturating surface of the maxilla, the ridges uniting
anteriorly well short of the premaxilla. A deep premaxillary
fossa, broken through in the specimen, implying a_ strongly
hooked mandibular symphysis. The maxillae united behind the
premaxillae for a significant distance in an incipient secondary
palate. The vomerine region filled with crystal; probably no
vomer was present. The ectopterygoid processes typically pleuro-
diran, moderately developed, and stronely inelined backward.
The hypertrophied ‘‘carotid channel’? smaller than in many
Podocnemis but clearly specialized in the fashion peculiar to this
eroup within the pelomedusids. The basisphenoid broad, sub-
pentagonal, a lone oblique lateral suture between quadrate and
basioceipital. The occipital condyle missing in the specimen.
Discussion. It is very easy to show that the skull here named
Dacquemys paleomorpha cannot belong to the genus Stereogenys.
The type of Stereogenys is S. cromeri (U. Eocene of the Fayum)
(Andrews, 1906), and the type of that species is a skull that
differs more from Dacquemys than Dacquemys does from some
of the species of Podocnemis sensu lato. I tabulate the major dif-
ferences below.
Stereogenys cromert
Skull very broad, nearly as broad as
long
Squamosal not in contact with parie-
tal
Orbits exposed dorsally
Secondary palate greatly developed,
formed by a long median suture of
the palatines, the maxillaries not
meeting medially or meeting for a
short distance only
Internal choanae therefore well be-
hind the orbits
Maxillary triturating surface with-
out ridging
Pterygoids (fide Andrews 1906) con-
ceal basisphenoid ventrally
Dacquemys paleomor pha
Skull moderately elongate, one and
one half times as long as wide
Squamosal in broad contact with
parietal
Orbits not exposed dorsally
Only an incipient secondary palate
formed by the median union of the
maxillaries behind the premaxil-
laries
internal choanae nearly at posterior
level of orbits
Maxillary with two ridges
ventrally in
3asisphenoid exposed
normal fashion
4 BREVIORA No. 35
The differences here summarized and also the great general dif-
ference in habitus go far beyond the amount that may be ad-
mitted within a genus of turtles. Daequé was indeed aware of
these differences (ef. p. 310 his paper) but he seems to have
beheved that the generic concept in turtles was founded on shells
alone, and therefore that if a skull could be associated with a
shell which appeared to be referable to a certain genus, the whole
animal must receive the shell name, no matter how aberrant the
features of the skull. This is, of course, incorrect. Not only is
the type of Stereogenys cromeri the highly peculiar skull and
not only are the shells which are referred to this genus so
referred on grounds which are not conclusive, but even if this
were not so, the generic concept in turtles as elsewhere is based
on the total animal and not any arbitrarily chosen part of it.
Generic¢ distinction rests on the total grade of specialization.
(Juite as surely as it cannot belong to Stereogenys, the skull
here discussed cannot belong to Podocnemis or to any other pelo-
medusid genus in which the skull is known,
From Shweboemys (Swinton, 1939) it differs in much the
same ways that it differs from Stereogenys. From Podocnemis in
a restricted sense (type P. erpansa and including only South
American species), the Oligocene Egyptian skull is distinguished
by the small, strictly lateral orbits and the very broad interorbital
width as well as by the squamosal-parietal contact and the
absence of a forehead groove.
From both EHrymnochelys (type and only lvine species FE.
madagascarensis) and Peltocephalus (type and only living
species P. dumeriliana) it is distinguished by the double rather
than single maxillary ridge, by the better developed secondary
palate, by the shallowness or absence of a precolumellar fossa as
well as by the squamosal-parietal contact and probably also by
the lateral emargination of the temporal region.
From Pelomedusa, Pelusios and Carteremys (Williams, 1953),
it is sufficiently distinguished by the covered temporal region and
especially the squamosal-parietal contact, but from at least
Pelusios and Pelomedusa it differs in the characters which it
shares with the Podocnemis group of genera, to which it must,
indeed, be referred — the enlarged ‘‘carotid channel’’ and the
hasioecipital-quadrate contact ete. From Bothremys (Hay, 1908 )
1
1954 DACQUEMYS PALEOMORPHA
it differs as radically as from Stereogenys e.g. in lacking the
extraordinary pits in the maxillae so characteristic of Bothremys,
in the much narrower skull, in the absence of a vomer, ete.
Other genera of fossil pelomedusids are known, but in none
of these is the skull known, and there is no plausible reason for
ascribing this skull to any of them. But while it is thus easy to
show that to place the Egyptian skull in any deseribed genus
would be erroneous or probably erroneous on present knowledge,
it is impossible to be sure that it does not belong to one of previ-
ously described Fayum species based on shells. I incline, indeed,
to the opinion that 1t does belong to one of these species.
From the Lower Oligocene beds of the Fayum from which
Dacquemys derives, three species of pelomedusid turtles have
been described: Podocnenis fajumensis Andrews, Ntereogenys
libyca Andrews, and Pelomedusa progaleata v. Reinach.
If Pelomedusa progaleata (material re-examined by Daequé,
1912) is correctly assigned and there appears to be sufficient rea-
son to think that it is, D. paleomorpha cannot be synonymous
with it, since the skulls in the two genera are so very different
that they do not belone even in the same subsection of the
Pelomedusidae.
Unless the Dacquemys skull belongs to some variety of shell
still undiscovered, decision must then lie between Podocnemis
fajumensis and Stereogenys libyca. This task is difficult, requir-
ing a rather complex and tenuous argument.
The argument may be stated in this fashion :
The shell of Podocnemis fajwmensis differs so little from that
of Recent Erymnochelys madagascarensis that, considering the
shell alone, the distinctness of the fossil species might be ques-
tioned. The skull of the fossil form might therefore be expected
to be very like that of the Recent species, and in faet the Dac-
quemys skull is more similar to the skull of Erymnochelys or of
related Peltocephalus than to any other known pelomedusid
skulls. Since it seems evident that skulls in the Podocnemis
sidebranch of the Pelomedusidae are subject to much more radical
modification than the shells, which tend to be conservative and
relatively difficult to distinguish, it is a possibilty worth con-
sidering that the Dacqué skull may belone to ‘* Podocnemis’’
fajumensis.
6 BREVIORA No. 35
In the alternative case of Stereogenys libyca the shell which is
the type of this species is clearly very similar to that referred to
S. cromert. The argument that the shells referred to S. cromeri
really belong to that species is plausible even though not con-
elusive. Andrews (1906, p. 298) remarks: ‘‘ Although in no case
have the carapace and plastron been found associated with the
skull in such a manner as to leave no doubt that they belong to
the same individual, nevertheless the shell... may be regarded
with reasonable certainty as belonging to the present species. In
the first place, this form of shell, like the skull, is the commonest
occurring in the Qasr-el-Sagha beds, and in the second place it
differs widely from the shell of Podocnemis, the only other Pleu-
rodiran genus found at this horizon.’’ Accepting Andrews’ in-
terpretation of the situation, if NStereogenys libyca is really
closely related to S. cromer?, it should have a skull to some degree
resembling that of S. cromeri. The Daequé skull is so very dif-
ferent from that of S. cromeri that it is very unlikely to be the
skull of S. libyea.
Thus the Daequé skull is unlikely to be that of S. libyca and
might be that of P. fajumensis. But to assert the truth of the
latter suggestion confidently would be a bold venture. In the next
paper of this series I shall present additional evidence based on
a skull from the Miocene of Moghara that this suggestion may
possibly be true, but even with this addition the evidence will still
be incomplete. It remains an unfortunate fact that with none
of the pelomedusids of Egypt is there an unequivocal association
of a skull and a shell. I have therefore described the species
paleomorpha as new, preferring to indicate the present state of
the evidence: that we have here a very distinet type of skull
which may belong to either of two distinct shell types or possibly
to a third unknown one.
Andrews (1906, p. 295) has described from the same beds with
Dacquemys paleomorpha, P. fajumensis and 8. libyca an anterior
portion of mandible which he has referred to P. fajumensis. Wis
comment on this element is as follows: ‘‘This mandible differs
from that of P. madagascarensis in having a broader symphysis ;
the biting surface is also broader and is partially divided in two
by a median ridge.’ Figure 1 is a photograph of this mandibular
fragment made by Peter Green and published by permission of
1954 DACQUEMYS PALEOMORPHA fi
the Trustees of the British Museum. It will be seen that the
ridging of this mandible would fit well with the maxillary rideine
of Dacquemys paleomorpha, but there is no strone hook at the
mandibular symphysis such as seems to be implied by the deep
premaxillary form of the skull. In this case, as in the previous
one, only discovery of associated material can verify the reference
to genus and species,
Fig. 1. Dorsal view of mandible referred by Andrews to Podocnemis
fajumensis. BM (NH) R 3346, from the Lower Oligocene of the Fayum,
Egypt. xl. Photograph reproduced by permission of the Trustees of the
British Museum.
The photographs of the skull of Dacquemys paleomorpha were
made by the Museum fur Naturkunde in Stuttgart and I am very
erateful to the authorities there for the privilege of reproducing
them.
[0 6)
BREVIORA No. 35
PAPERS CITED
ANDREWS, C. W.
1906. <A descriptive catalogue of the Tertiary vertebrata of the Faytim,
Egypt. London. 324 pp.
DacquE, E.
1912. Die fossilen Schildkréten Aegyptens. Geol. Palaeont. Abhandl.,
vol. 14, pp. 275-337.
Islwe, (O12
1908. The fossil turtles of North America. Carnegie Inst. Publ. No. 75.
568 pp.
SIEBENROCE, F.
1902. Zur Systematik der Schildkrétengattung Podocnemis Wagl. Sitz.
Ber. Akad. Wiss. Wien, math. nat. Kl, Abt. 1, vol, Mi;
pp. 157-170.
Swinton, W. E.
1939. A new fossil fresh-water tortoise from Burma. Rec. Geol. Survey
India, vol. 74, pp. 548-551.
WILLIAMS, BH.
1953. Fossils and the distribution of chelyid turtles. 1. ‘‘ Hydraspis
leithii’’ in the Eocene of India is a pelomedusid. Breviora, No.
133, jones Jletsy
Pilate 1. Type skull of Dacquemys paleomorpha, new genus, new species.
Staatliches Museum fiir Naturkunde, Stuttgart No. 12645. Above: ventral
view; below: dorsal view. Photographs reproduced by courtesy of the
Staatliches Museum fiir Naturkunde. x1.
BREVIORA
Museum of Comparative Zoology
PRESENT KNOWLEDGE OF THE SNAKE
ELACHISTODON WESTERMANNI
REINHARDT
By Cart GANS
Carnegie Museum, Pittsburgh, Pa.
and
ERNEST EK. WILLIAMS
Museum of Comparative Zoology, Cambridge, Mass.
INTRODUCTION
In 1863 Johannes Reinhardt described a peculiar snake belong-
ing to the new genus and species Hlachistodon westermanni, the
specific name referring to the collector, William Westermann,
who obtained the initial specimen in Runepore (Rangpur, see
Fig. 1), northeast Bengal. Reinhardt’s single specimen was
characterized by certain specializations of the integument, by
an almost complete absence of teeth, and by certain of the verte-
brae bearing hypapophyses which penetrated the esophagus.
Comparison with the African snake Dasypeltis, long known to be
specialized for egg-eating by a similar series of modifications,
was inevitable, and Reinhardt came to the conclusion that Hlachi-
stodon was also an egg-eater. Differences between Elachistodon
and Dasypeltis centered about minuscule grooved teeth and an
enlarged row of dorsal scales in the new genus. The presence of
the grooved teeth on the posterior aspect of the maxillae sug-
gested the placement of the form in the composite section
Opisthoglypha, but Reinhardt pointed to the close similarity
between this form and the aglyphous Dasypeltis as evidence for
the artificiality of this category.
2 BREVIORA No. 36
Since 1863 only four additional specimens have been reported
and there has been httle additional contribution to the morphol-
ogy or natural history of the species.
A recent study (Gans, 1952) has demonstrated a number of
previously undiscussed modifications in Dasypeltis and has
shown that this genus is much more highly adapted to its very
special mode of life than had previously appeared to be the
case.
In view of this new knowledge of Dasypeltis it was felt that a
re-examination of the status of Hlachistodon would be especially
NE Pay
Jalpaiguri
Rangpur,
Fig. 1. The distribution of Blachistodon westermanni. Crosses mark the
localities where specimens were found.
appropriate at this time. Unfortunately, however, the existing
specimens are too few to permit a detailed investigation such as
was carried out for Dasypeltis. It has, indeed, been possible
for one of us (Williams) to make an examination of the specimen
in the British Museum and of the osteological preparations from
that specimen figured and referred to by Maleolm Smith (1943,
p. 403), but the information so derived is suggestive rather than
adequate, and it is very evident that further specimens will be
needed for thoroughgoing anatomical studies.
1954 ELACHISTODON 3
This paper, therefore, has two objects. First, it is intended to
provide a recapitulation of the existing data on Llachistodon in
the light of the findings in the genus Dasypeltis, and secondly
it is desired to reawaken interest in Hlachistodon in the hope of
obtaining the additional specimens upon which the more intensive
studies might be based.
HISTORICAL RESUME
The initial specimen was rather fully described by Reinhardt
(1863, p. 198), whose careful investigation of its internal anat-
omy was undertaken because the enlarged vertebral scales of the
new form had left him in doubt as to its taxonomic placement.
He prepared the tooth-bearine elements of the right side of the
skull and one of the mandibles, and cheeked the condition of
the vertebral hypapophyses and their relation to the esophagus
without dissecting out the vertebrae or separating them. He
found teeth on both palatine and pterygoid as well as teeth and
minuscule fangs on the maxilla and described the shape and
appearance of the dentary. In comparing Elachistodon with
Dasypeltis he pointed out that in both there were two types of
penetrant hypapophyses, though both types in the new form were
less well-developed. He figured the palatal and mandibular
elements of both genera (see Fig. 4).
The stomach and gullet of the type specimen of /. westermanni
were found to be filled with an amorphous congealed mass whieh
analysis indicated might be either milk or ege fluid. Though he
did not find shell fragments mixed in this mass, Reinhardt
speculated on the feeding habits of the animal and appears to
have leaned toward the theory that the species was an egg-eater.
Reinhardt also described in detail the squamation of Mlachisto-
don, mentioning the very pecuhar deep pit on the posterior
nasal as well as similarities to Dasypeltis. Further details on
these and other structural points will be presented in the dis-
cussion of the morphology of EF. westermanni. Reinhardt placed
the new genus in the family Rachiodontidae (sic).
The next to refer to Elachistodon was Gunther (1864, p. 444)
who mentioned it on Reinhardt’s authority only. He suggested
that the genus should be separated as a distinct group of
colubrids.
4 BREVIORA No. 36
Blanford (1875, p. 207) reported the second specimen from
Bihar, Purnea (see Fig. 1), and his general description agreed
with that of Reinhardt. He suggested that the genus be referred
to the Dipsadidae.
Cope (1886, p. 494) considered only the familial assignment
of the genus, placing it in the subfamily Dasypeltinae with
Dasy peltis.
Boulenger in the Fauna of British India (1890, p. 362) placed
the genus in the Opisthoglypha, Dipsadinae, next to Dipsas. He
stated that he had not been able to examine specimens but from
the key (pp. 356-857) it appears that the assignment was based
on the grooved teeth, the enlarged vertebral shields and the
elliptically vertical pupil. He also referred to esophageal teeth
capped with enamel.
Sclater (1891, p. 48) in lsting the snakes in the Indian Mu-
seum recorded a specimen from Bengal which seems to have
been that earher recorded by Blanford. He placed it in the
Dipsadinae.
In the Catalogue of the Snakes of the British Museum, volume
3, Boulenger (1896, p. 263) erected the subfamily Elachistodon-
tinae of the Colubridae with the single genus Elachistodon. He
regarded this as the opisthoglyphous analogue of the aglyphous
Rachiodontinae. His description of the subfamily mentions ptery-
goid teeth.
The next record is that of Wall (19138, p. 400) who reported
a new specimen from the Jalpaiguri district (see Fig. 1) and
published a detailed comparison of the three specimens then
known. He did not give the sex of the third specimen (it is, how-
ever, the female deseribed again by M. Smith, 1943) but did cast
doubt on the presence of pterygoid teeth. His later ‘ Handlist’
(1923, p. 878) did not contain any new data.
In a serialized discussion of the snakes of northern Bengal
and Sikkim, Shaw, Shebbeare and Barker (1941, p. 65) list the
specimens known to them. In addition to the original Rangpur
specimen in the Museum at Copenhagen they cite a second from
Purnea (the Blanford specimen) in the Indian Museum at that
time. The specimen obtained by Travers at Jalpaiguri (Wall,
1913) was stated to be in the Bombay Museum (it is now in
the British Museum), while the then whereabouts of two addi-
1954 ELACHISTODON 5
tional specimens obtained by Travers at Baradighi! appeared
to be unknown.
Maleolm Smith (1943, p. 403) redeseribed the genus and
removed it from the Elachistodontinae te the Dasypeltinae. He
presented the first published figures of the modified vertebrae and
also refigured the palatal bones. Pterygoid teeth are not men-
tioned in his text or shown in his figure.
Smith and Bellairs (1947, p. 362) mention only that the
Harderian gland is enlarged in this form as in Dasypeltis.
MORPHOLOGY
General
The following condensed diagnosis is designed to aid identifica-
tion of specimens. For this reason it follows the summary pre-
sented by Smith (1943, p. 404) with only minor modifications.
Head fairly distinet from neck ; eye large with vertically ellipti-
eal pupil (round fide Reinhardt); a large pit in the posterior
part of the nasal shield. Body moderately elongate, feebly com-
pressed. Tail short, subeaudals paired.
Internasals as large as prefrontals; frontal large, longer than
its distance from the end of the snout; nasal large; 1 small pre-
ocular, the loreal below it entering orbit; two postoculars; two
long anterior temporals; 6 or 7 supralabials, the third and fourth
touching the eye; 2 pairs of sublinguals. Seales smooth, in 15
rows, 19 on the neck, the vertebral series much enlareed, hexag-
onal. Anal single. Ventrals 208-217. Caudals 59-65.
Above, dark olive brown to blackish, the vertebral scales yel-
lowish-white, except at their outer margins, forming a leght
vertebral stripe extending the whole length of the body; sides
spotted or flecked with the same color; whitish below, the outer
margins of the ventrals and adjacent rows of scales edged with
brown; a yellow stripe along the top of the head from the snout
to the angle of the mouth, passing above the eye; an angular bar
or spot on the nape; lips yellow.
Q 784 mm. (670 + 114 mm.). Ventrals 217. Caudals 59. Rung-
pore, Bengal. Copenhagen Museum (Reinhardt).
1 We have been unable to find Baradighi on any map, but the Indian Consular
Office in New York has very kindly informed us that there is a railroad station
by that name about 30 miles north of Calcutta.
6 BREVIORA No. 36
? 800 mm. (670 + 130 mm.) fide Smith, (762 mm. [635 + 127
mm.] fide Wall). Ventrals 213 (210). Caudals 62 (64). Near
Mal, Jalpaiguri District. Formerly Bombay Museum, now
British Museum (Natural History).
222 mm. (186 +- 35.7 mm.), Ventrals 208. Caudals 63. Bihar,
Purnea. Indian Museum (Blanford).
Fig. 2. EF. westermanni. Oblique view of the head of the British Museum
specimen to show the expansion lines between the lower labials at the angle
of the mouth.
Squamation of Head and Neck
The most significant integumentary similarities between EHlach-
istodon and Dasypeltis are the structure and arrangement of the
eular and cheek scales. This similarity was already pointed out
by Reinhardt who, however, did not know the function of the
parallel structures. The fact that in Dasypeltis these gular and
cheek specializations have been definitely shown to be elaborate
adaptations for egg-eating suggests that the possession of similar
structures by Elachistodon is correlated with similar habits.
In both Elachistodon and Dasypeltis there are two pairs of
sublinguals closely joined and lacking any evidence of the median
eroove that in most snakes provides for expansion. In Dasypeltis
~
1954. ELACHISTODON
there are no true gular scales at all, while in Llachistodon there
is but one row of these. In both of these forms the sublinguals
are so rigidly tied in that expansion of the chin region can only
take place along the angle of the mouth during ingestion. In
Dasypeltis this expansion is made possible by the presence of
extremely distensible skin between the scales of this region. The
last labials have also been rearranged to form the first scales of the
lowest lateral rows. The same condition is also the case in
Elachistodon, and the resulting lines of expansion are clearly
seen in Figure 2. Such a specialization, the value of which is
probably solely restricted to egg-eating, would tend to furnish
additional evidence for the idea that other described modifica-
tions of Elachistodon are adaptations to this habit.
A possibly significant point of difference between HLlachistodon
and Dasypeltis is the presence in the form of a large pit on the
posterior part of each nasal shield. The function of similar pits
in the Crotalidae as well as in certain species of Pythonidae has
been determined to be that of a thermosensitive range-finder
(Noble and Schmidt, 1937), and it is known that similar strue-
tures of as vet uninvestigated function exist in other forms. In
Elachistodon nothing is known of the function of the pits.
Squamation of the Posterior Body
The enlargement of the vertebral series of dorsal scales char-
acteristic of Hlachistodon is a feature also found in a large num-
ber of dipsadine and other snakes but not in Dasypeltis. Little
is known about the function of this enlargement although Peters
(MS 1952, p. 27) has speculated on its relation to the arboreal
habits of the dipsadines. He believes that in conjunetion with
the extreme lateral flattening of the body it might impart a
stiffening effect. He offers the analogy of an I-beam, and suggests
that the enlargement of the vertebrals might permit the unusual
horizontal rigidity and extension observed in chmbing members
of the Dipsadinae. We cite the suggestion here without passing
judgment upon its appheability to Hlachistodon, which shows
but feeble lateral compression.
It is also to be noted that the scales of Elachistodon are smooth
while those of Dasypeltis are strongly keeled. This character is,
however, of little value at the generic level or above, in view of
Co
BREVIORA No. 36
the number of cases in snakes in whieh it shows intrageneric¢
variation.
Elachistodon exhibits no trace of the size reduction, angling
and serration exhibited by the second to sixth rows of dorsal
scales in Dasypeltis. Except for the vertebral row all the dorsals
are uniform in size and character.
Soft Tissues of the Head
No published observations are available on gum structure.
Examination of the British Museum specimen indicated that the
eum ridges are probably covered with a somewhat less convoluted
mass of tissue than are those of Dasypeltis.
Smith and Bellairs (1947, p. 362) have noted that the Harder-
ian gland is notably enlarged in Elachistodon paralleling the
econ ition in Dasypeltis.
Fig. 3. H. westermanni. A, Palatal bones; 6, anterior hypapophysial
vertebrae; C, posterior hypapophysial vertebrae. Abbreviations: e, ecto-
pterygoid; mz, maxillary; pal, palatine; pt, ptergygoid. After Malcolm
Smith (1943).
Skull
To the best of our knowledge no complete skull has ever been
prepared. The palatal elements have been figured by Smith
(1943) (see Fig. 3, A). A check of the material by one of
1954 ELACHISTODON i)
us (Willams) found the figures accurate in all particulars inelud-
ing the fact that pterygoid teeth (described and illustrated by
Reinhardt in the Copenhagen specimen) are not to be seen.
Comparison with the homologous elements in Dasypeltis in-
dicates few differences beyond the presence of the relatively
enlarged grooved teeth on the posterior aspect of the maxilla in
Elachistodon. This portion of that bone is also somewhat more
strongly developed and there are general though minor differ-
ences in the relative proportions of the various structures. The
union between palatine and maxillary may be somewhat weaker
than the corresponding articulation in Dasypeltis. The wine-
shaped process of the palatine is much abbreviated though this
process is shown as greatly extended in Reinhardt’s drawing.
These bones are, however, very fragile, and it may be possible
that some of the apparent differences may be traced to the
method by which these elements were cleaned. The similarities
seen lead us to the conclusion that the palatal elements of
Elachistodon are as rigidly joined as those of Dasypeltis.
The dentitional formulae are as follows: Maxilla edentulous
except for two small teeth followed by two larger grooved teeth ;
palatine with four minute teeth in its center; pterygoid — ac-
cording to Reinhardt — with three extremely minute teeth. Wall
(1913) and Smith (1943) — both treating of the specimen from
Jalpaiguri — do not refer to pterygoid teeth. The dentary has
eight to twelve teeth diminishing in size posteriorly.
Reinhardt has figured and described the anterior end of the
mandible for both Llachistodon and Dasypeltis. The appearance
of the element in Hlachistodon is very strange, and since the fig-
ure of the same element in Dasypelt's bears very little resem-
blanee to actual specimens seen by us, we do not care to comment
further in this matter (see ie. 4).
Vertebral Column and Esophagus
There are certain rough similarities in the appearance of the
anterior vertebrae (hypapophysials) in the two forms. In both
there are two basic types of hypapophyses — anterior and _ poste-
rior, but in Dasypeltis some of the anterior hypapophyses undergo
considerable ontogenetic variation so that it is best to distinguish
anterior, intermediate, and posterior types. It is not known
10 BREVIORA No. 36
whether similar ontogenetic variation is found in Hlachistodon.
For the latter genus, therefore, these comments must of necessity
be restricted to the modified vertebrae of the adults.
In Dasypeltis the anterior hypapophyses are little modified.
They are of the ‘‘normal’’ squarish shape with only their ventral
edges enlarged into sled-like runners. The ege glides along these
runners during ingestion, but the runners do not serve any cut-
ting or sawing function. The ege is pierced by the elongate,
spiniform, forward pointed hypapophyses of the posterior series,
which penetrate the esophagus in all specimens (Gans, 1952, p.
236). After the shell has been broken, the heavy rounded boss-
hike hypapophyses of the intermediate series transmit a crushing
force to the ege shell and fold this, while the egg contents are
being squeezed into the stomach through the esophageal valve.
The forces involved in this action are sizable, and for this reason
the articulating surfaces of the neural arches are greatly ex-
panded, the pre- and postzvgapophyses being laterally displaced
and extended by spiniform processes. This offers additional sur-
face for the action of the dorsal musculature which is kept from
lateral slippage by the confining inclination of the surfaces of
attachment.
The intermediate and some of the anterior hypapophyses are
penetrant in adults of Dasypeltis, but this character shows con-
siderable variation in juveniles. There is also evidence that some
seasonal variation of this character may occur in adult specimens
as well.
In Elachistodon nothing appears to be recorded about the
hypapophysial vertebrae immediately posterior to the cervicals,
and these have never been figured. Perhaps it may be assumed
that they are of ‘‘normal’’ appearance, possibly performing the
same function as do the homologous structures in Dasypeltis.
There is no record as to the number of these ‘‘ unmodified’’ verte-
brae, though Smith (19438, p. 403) states that the modified
hypapophyses start opposite the tenth ventral shield,
The amount of skeletal preparation which has been done for
this region is very limited. Reinhardt (1865, footnote to p. 202)
stated that he had not skeletonized but rather examined the
vertebral column in situ. Smith figures two short sections of
three vertebrae each (Fig. 3, B and C). One of us (Williams)
er es
SS ~
Fig. 4. Plate from the original description by Reinhardt. A, B, C, Dorsal,
ventral and side views of the head of EH. westermanni; D, palatal bones of
EB. westermanni; FE, dentary of EF. westermanni; F, palatal bones of
Dasypeltis scaber; G, mandible of Dasypeltis scaber.
12 BREVIORA No. 36
has examined these vertebrae, and the following statement is
based on this examination,
The more anterior of the prepared and figured hypapophyses
are very strange in appearance. Bosses are present as in the
intermediate hypapophyses of the African form, but they are not
as closely integrated with the base of the centrum. They are
instead mounted on a pedicel beyond which they extend both
anteriorly and posteriorly. The vertebrae bearing these hypa-
pophyses are relatively undifferentiated. No extension or inelina-
tion of the zygapophysial surfaces is apparent.
In some ways the anterior hypapophyses of Elachistodon are
reminiscent of stages in the ontogeny of the intermediate hypa-
pophyses in Dasypeltis, though the mounting on a pedicel sug-
eests that they may be specialized in a different direction. Rein-
hardt (1868, p. 203) stated that there were nine such anterior
hypapophyses, while Smith (1943, p. 403) counted eighteen
all of which projected through longitudinal slits into the esopha-
cus. Both authors believed that these are cutting instruments,
which hardly seems hkely in view of the findings in Dasypeltis.
Compared to the posteriormost hypapophyses in Dasypeltis
which are such highly effective tools, these same elements in
Elachistodon seem even less modified than the anterior hypa-
pophyses. In Llachistodon the hindmost hypapophyses are block-
shaped spines whose major specialization hes in the facet that
they are directed forward rather than backward. They most
nearly resemble the hypapophyses of several of the species of
Klaphe recently mentioned as being specialized for egg-eating
(Gans and Oshima, 1952, p. 15), but they also resemble those
hypapophysials of very young Dasypeltis that are transitional
between the intermediate and the posterior series.
Reinhardt speaks of thirteen of the posterior type hypa-
pophyses in Elachistodon, with only the first ten penetrating the
esophagus, while Smith mentions eight of these structures, none
penetrating the esophagus. Smith’s observation would of course
be in strong contrast to the findines in Dasypeltis, and the con-
tradiction in observations here makes it quite clear that the egg-
eating habits of this form will have to undergo a separate analysis
to determine the extent to which parallelism in function exists,
and to explore the differences.
1954 ELACHISTODON ile
In spite of this, it may be stated that all indications exist that
egos are opened by Llachistodon in a manner similar to that
employed by Dasypeltis and Elaphe climacophora, i.e. by exerting
a foree concentrated by the processes of the vertebrae.
A comment may be made on Smith’s Figure C (our Fig. 8 C).
The third figured hypapophysial carries a hypapophysis with
what appears to be a shovel-shaped tip. While the drawing: is
entirely accurate, its two-dimensional nature does not do the
subject justice, as this tip is actually deformed into a hook in
the horizontal plane in a most peculiar manner. The asymmetry
and other features of the structure make it obvious that this is
merely an individual aberration of no adaptive or other sig-
nificance. Similar variations occur on the hypapophyses of a
number of other oophagous and non-oophagous species of snakes
examined by us.
Finally it may be well to mention that the hypapophyses in
Elachistodon are formed of very dense bone only and do not
consist of enamel as stated by various authors. While it has not
been possible to undertake histological studies for this form,
detailed and repeated investigations have shown this to be true for
Dasypeltis (Gans, 1952; Gans and Oshima, 1952), and no evi-
dence exists which might permit the contrary conelusion in
Klachistodon.
BIOLOGY
No specimen of Hlachistodon has ever been kept in captivity
and there exist no notes on habits or habitats in the wild. With
the exception of Reimhardt’s specimen which had ege yolk (?)
in its stomach, no other observations on stomach contents have
been published. The concensus of the literature (based on no
direct evidence) is that the form eats ‘‘eges but not exclusively.”’
The presence of grooved teeth and the nasal pit suggest that
the species may feed also on lve and homoiothermous prey.
The only other item of interest concerns breeding habits. The
type was a gravid female, containing seven (40 x 11 mm.) eges
(Reinhardt, 1868, p. 210). None of these contained embryos
although they were already covered with a relatively heavy shell.
This would indicate that the species is oviparous as is also true
for Dasypeltis.
14 BREVIORA No. 36
Comparison of Elachistodon and Dasypeltis
The following tables sum up the more important similarities
and differences — at present known — of the two genera.
Points of Similarity
Sublingual shields fused, mental groove absent; special pro-
vision for expansion of the skin alone the angle of the mouth
and in the cheek region; Harderian gland much enlarged;
palatal elements modified, largely edentulous, with teeth present
only on the posterior aspects of maxilla and palatine ; ? upper jaw
elements rigidly tied together; teeth on posterior aspect of
dentary only; a loose articulation between dentary and com-
pound bone. Vertebral hypapophyses modified, some penetrat-
ine the esophagus; anterior hypapophyses generally rounded,
with sledge-like runners or bosses; posterior hypapophyses de-
veloped as forward-pointing spines ; oviparous.
Points of Difference
Dasypeltis Elachistodon
Nasal pit absent Nasal pit present
No gulars One row of gulars
Scales strongly keeled Scales smooth
Vertebral scale row subequal to dor- Vertebral scale row enlarged
sals
Some lateral rows with scales re- All lateral rows with scales of equal
duced in size, inclined and with size
serrate keels
Maxillary teeth minute and equal
Intermediate hypapophyses heavy,
rounded, boss-like, closely applied
to centra
Posterior hypapophyses bear sharply
pointed spines and _— penetrate
esophagus
EK. liptieal pupil
No pterygoid teeth
1-2 enlarged grooved teeth on rear
of maxilla
Bosses of hypapophyses separated by
centrum, i.e.
constriction from
mounted on pedicel
Posterior hypapophyses much _ less
strongly developed, ? no penetra-
tion
? Round pupil (fide Reinhardt)
? Pterygoid teeth (fide Reinhardt)
1954 ELACHISTODON 15
Inspection of the above lists reveals that the similarities be-
tween the two forms are primarily those of struetures which in
Dasypeltis are known to be connected with ege ingestion. Many
of the differences are at a level generally associated with a generic
separation, while only the nasal pit and the grooved fangs might
indicate a higher level of difference.
It has been suggested that Hlachistodon may be a less special-
ized form than Dasypeltis but derived from a common ancestral
stock that may have been opisthoglyph or even proteroglyph.
According to this, greater specialization in’ Dasypeltis has
brought about the total loss of the fanes as well as the additional
modifications exhibited by this genus. Analysis of the relation-
ships of the genus ELlachistodon may indeed yield evidence shed-
ding light on the general problem of the origin of the opistho-
elyphs.
Beyond the two genera which have been compared here, analy-
sis will have to be carried forward for the various other forms
known or suspected to be specialized egg-eaters before it will be
possible to determine whether we are dealing with a single series
or a set of parallel modifications in separate lines.
It is to be borne in mind that we have here reported for the
most part the observations of others and have had little opportun-
ity to check these against actual specimens. It may well be that
careful examination of a large series of specimens would foree
a revision of our understanding of the morphology of Hlachisto-
don. This is particularly likely in view of the high amount of
variation exhibited by the much better known Dasypeltis. The
many points of difference noted from the scanty literature testify
only too clearly that a similarly high amount of variation is
present in Llachistodon. It would obviously be dangerous to
speculate at this time on the larger problems offered by the Indian
eenus.
In view of this dearth of knowledge and the high interest of
the Indian genus, the authors wish to urge local zoologists or
anyone whose work may take him into the range of this form
to secure us specimens. Any such material may be sent to
either of our institutions. Credit will of course be given in any
subsequent reports.
Acknowledgments. Our thanks are due to a considerable num-
16 BREVIORA No. 36
ber of people with whom phases of the problem were discussed.
We wish to express our thanks also to Dr. H. W. Parker, Mr.
J. C. Battersby and Miss A. G. G. Grandison of the British
Museum (Natural History) for assistance there and for permis-
sion to examine their specimen of EHlachistodon. We are indebted
also to the authorities of the University of Michigan Museum of
Zoology for the use of library facilities while in Ann Arbor.
BIBLIOGRAPHY
BLANFOoRD, W. T.
1875. Note on (i) Blachistodon westermanni, (i) Platiceps semifascia-
tus, and (iii) Ablepharus pusillus and Blepharosteres agilis.
Jour. Asiat. Soc. Bengal, vol. 44, pt. 2, pp. 207-209.
BOULENGER, G. A.
1890. Reptilia and Batrachia. in The Fauna of British India, including
Ceylon and Burma. London, xviii + 541 pp.
1896. Catalogue of the snakes in the British Museum (Natural His-
roO7
tory). London, vol. 3, xii, + 727 pp.
Cops, E. D.
1886. An analytical table of the genera of snakes. Proc. Amer. Philos.
Soc. 1886, pp. 479-499.
GANS, C.
1952. The functional anatomy of the egg-eating adaptations in the
snake genus Dasypeltis. Zoologica, vol. 37, pt. 4, pp. 209 244.
GANS, C. and M. OSHIMA
1952. Adaptations for egg eating in the snake Hlaphe climacophora
(Boie). Amer. Mus. Novitates, No. 1571, 16 pp.
GUNTHER, A.
1864. The reptiles of British India. London, xxvi + 452 pp.
Nosug, G. K. and A. SCHMIDT
1937. The structure and function of the facial and labial pits of
snakes. Proc. Amer. Philos. Soc., vol. 77, no. 3, pp. 263-288.
PHTERS, J. A.
1952. The snake subfamily Dipsadinae in South and Central America.
University of Michigan Doctoral Dissertation, 369 pp.
REINHARDT, J.
1868. En ny Slaegt af Slangefamilien Rachidontidae. Oversigt. K.
Danske Vidensk. Selskab., pp. 198-210.
1954
=
ELACHISTODON 1
ScLATER, W. L.
1891.
List of the snakes in the Indian Museum. Caleutta, x + 79 pp.
SHAW, G. E., E. O. SHEBBEARE, and P. E. BARKER
1941.
The snakes of northern Bengal and Sikkim. Jour. Bengal Nat.
Hist. Soc.; vol: 16) pp. a7-67.
SmirH, M. A.
1943.
Reptilia and Amphibia. in The Fauna of British India. London,
vol. 38 (Serpentes), 583 pp.
SmituH, M. A. and A. p’A. BELLAIRS
1947.
WALL, F.
1913.
1923.
The head glands in snakes with remarks on the evolution of
the parotid gland and teeth of the Opisthoglypha. Jour. Linn.
Soc., vol. 41, pp. 351-368.
A rare snake, Elachistodon westermanni from the Jalpaiguri
District. Jour. Bombay Nat. Hist. Soc., vol. 22, pp. 406-401.
A handlist of the snakes of the Indian Empire. Part. Til. Jour.
Bombay Nat. Hist. Soc., vol. 29, pp. 864-878.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. Avueust 18, 1954 NUMBER 37
ON THE EVOLUTION OF AN ORIENTAL
HKARTHWORM SPECIES, PHERETIMA
ANOMALA MICHAELSEN 1907
By G. E. GAres
Most individuals of this aptly named species are referable to
three formae, though large collections usually have contained
one to several individuals regarded as intermediates. F. centralis,
in most localities much the rarer of the three, differs significantly
from the generic pattern only in location of the male pores on xx.
F. typica, the first to be found, is athecal and additionally distin-
guished from centralis by extra pairs of testes and male funnels
in y-ix, frequently even supernumerary gonads in those segments,
as well as by extra pairs of mushroom-shaped glands in one or
more of segments xxi-xxiv. F’. insolita is thecal, each organ of the
battery normal and the pores on 5/6-7/8 as in centralis, but lacks
the mushroom glands and the male genital terminalia.
According to the first attempt at explaining the origin of these
differences, f. typica and f. insolita, are respectively male and
female secondarily evolved from the normally hermaphroditie
centralis. In spite, however, of the presence of testes in unusually
large numbers, sperm are not matured in the supposed males. On
the contrary, sperm are produced by the supposed females even
though discharge of the matured gametes from the body seems to
be impossible.
According to a second explanation, the two divergent forms
result from selective inhibition of secondary sex organs, during
post-hatching development, by metabolic products released from
two different types of protozoan parasites. Before an opportunity
could be found to study results of activities of the parasites that
2 BREVIORA NO. 37
obviously are present in such large numbers in the aberrant forms
of this and also other species of Pheretima, data from other
sources began to indicate the necessity for another explanation.
P. anomala has now been found in several widely separated
localities in India to which it has been transported from its
proper home somewhere to the east of the Irrawaddy River. Only
f. typica was secured at each of those localities. Presumably
then, this form breeds true. This assumption is also supported by
the incidence of the various formae in certain localities in Burma,
as for instance in the Kamaunegthwe River region of Tavoy district.
In two collections from that area, taken two months apart, cen-
tralis, typica, nsolita and the intermediates were represented by
0-+-0, 377-+-69, 114-2, and 31+8 specimens respectively. The in-
termediates, in most of these cases, would be referable to a less
strictly defined insolita as they were distinguished only by the
possession of one or more of the mushroom glands characteristic
of centralis and typica.
F. insolita has not been found as yet in complete isolation, but
its incidence is high in some localities. One collection from Taungyi,
in which centralis, typica, insolita and intermediates were repre-
sented by 7, 32, 293 and 3 specimens respectively, also suggests
a possibility of a true-breeding form. Here again the intermedi-
ates were referable to a less strictly defined insolita.
F. centralis never has been found in isolated colonies. It usu-
ally can be secured in most localities in Burma if search for it is
continued long enough. Incidence always has been low, the great-
est, according to the records now available, about 20 per cent, in
a Karenni collection (from Koopra) in which the formae were
represented, in order as above, by 21, 5, 67 and 9 specimens. Geo-
eraphically, Karenni quite possibly is nearer the region where
anomala may have arisen.
Throughout the Burmese portion of its range, P. anomala now
seems to have been largely segregated into true-breeding forms.
This could have been brought about, in earthworms, through re-
placement of sexual by parthenogenetic reproduction. As sperm
are not produced by f. typica and since it lacks the organs for
reception of male gametes from another individual, it can be
assumed to reproduce parthenogenetically. In f. insolita, though
sperm are produced, they cannot be discharged during copulation
1954. EVOLUTION OF AN ORIENTAL EARTHWORM 3
and few individuals are likely to have an opportunity to copulate
with those that can. No good evidence has been found for self
fertilization. Presumably then, these supposed females, like the
‘“‘males’’, usually do not reproduce sexually but parthenogeneti-
eally.
Evidence for copulation of individuals of different formae
cannot now be presented due to destruction of the records during
World War II. Copulation between individuals of typica cer-
tainly would appear to be futile. Cross copulation between indi-
viduals of isolita and typica would have no result for the typica
worm, and the imsolita partner ordinarily would receive into its
spermathecae only secretions from the unusually well developed
prostate glands of the typica individual. The opaque and non-
iridescent coagulum often present in spermathecae of clitellate
specimens of insolita may have been prostatic secretion received
from a typica worm though the same non-iridescent opacity can
eventuate from resorption of sperm in the seminal fluid received
from another sort of partner.
Copulation in the cross insolita x centralis would not even result
in transfer of prostatic secretions to the spermathecae of the
normally sexual centralis individual which would then have to
reproduce, if at all, by self fertilization (unlikely) or by partheno-
genesis! In the latter case, offspring of the centralis type would
be expected. The imsolita partner, normally parthenogenetic,
would however receive centralis sperm and unless parthenogene-
sis had become obligatory could be expected to reproduce sexually !
Copulation between two individuals of centralis can be expected
to result only in centralis offspring but in view of the rarity of
this form throughout most of Burma the cross can be expected
there only infrequently.
A majority of the so-called intermediates resemble insolita ex-
cept for the presence of one or more mushroom glands. Such
anarsenosomphie worms can be expected to reproduce in the same
way as insolita. The intermediates with male genital terminalia
on one or both sides of the body can be expected to reproduce in
the same way as centralis.
Of the 99 intermediates from the collections that were made
throughout Burma during 1928-1932, 96 were assumed, because
of their structure, to show relationships to both ‘nsolita and cen-
4 BREVIORA No. 37
tralis, the only forms which can be expected, according to the
preceding discussion, to produce offspring from a hybrid cross.
Of the 96, 71 were without male terminalia like insolita and 25
had male terminalia as does centralis. This looks like a good
approximation to the expected ratio in the Fy generation if ‘‘in-
solita’’ is dominant to ‘‘centralis’’. The Mendelian relationship,
if such it be, is however recognizable only because the mushroom
olands of centralis have been carried over into the insolita pheno-
type. The centralis phenotype likewise was aberrant as prostates
were variously located in segments xix-xxi, or only their ducts
were present, or only one prostate was present, while mushroom
olands always were lacking in xvii-xix and when present were in
the segment where prostates should be expected.
The three remaining intermediates were thought to show rela-
tionships to typica and centralis. Possibly three of the 528
specimens that were identified as typica during the same period
should also have been included. Each of these six worms had
testes and male funnels in some or all of v-ix as in typica but
three had a single normal centralis (or insolita) spermatheca and
a fourth had a rudimentary one concealed within the parietes. The
external aperture of one of the normal spermathecae was on 8/9
and this is the only instance that has been recognized in anomala
of a pore at that posterior level. Offspring cannot be expected
of a cross between typica and centralis, according to a previous
portion of this discussion, and the mass of material collected sub-
sequent to 1932 in the hope of clarifying the relationship was
destroyed during the war. However, it can be said that an occa-
sional individual of typica showed in seminal vesicles and/or testis
sacs rather dubious evidence of having produced a few sperm.
Hence, rarely, a cross between typica and centralis or even wmsolita
may be possible. As the latter two are both thecal, presence of
spermathecae would provide no clue as to which of the two forms
had been involved in the hybrid parentage. If the hybridization
is possible, the six cases just cited would seem to indicate that
‘‘typica’’, i.e., extra testes and male funnels, is dominant over
‘“spermathecae’’ i.e., ‘‘centralis’’ or ‘‘insolita’’.
P. anomala must have arisen, presumably somewhere to the east
or southeast of Burma, from an ancestor that may have had mush-
room glands, or an equivalent, but which did have the male geni-
1954 EVOLUTION OF AN ORIENTAL EARTHWORM 5
tal terminalia in xviii, as throughout most of the genus Pheretima.
The evidence provided by aberrant individuals of this and other
species agrees in indicating that transfer of capacity to develop
prostate glands, from one segment to another, whether anteriorly
or posteriorly, takes place in a single step rather than by the
much more gradual sort of migration that has been assumed in
the classical oligochaete phylogeny. An early, if not the first
step, in the evolution of anomala was then the establishment of a
mutation for transfer of prostate developing capacity from xviii
to xx. No difficulty is to be expected for the male deferent ducts
in reaching the new segment through which they must now open
to the exterior, as numerous specimens of insolita have shown that
the ducts can grow back as far as xxx behind which level male
pores very rarely have been recorded in any family of earthworms.
F. insolita also demonstrates that the male deferent ducts do not
acquire an external aperture in absence of the prostates. Union
of male deferent and prostatic ducts, regardless of the segment
in which the glands are located, suggests that the former, on
reaching the prostatic segment, are attracted towards the growing
glands. Union presumably takes place at parietal level. Subse-
quently the prostatic duct ectal to the junction becomes much
elongated, carrying the region of that junction deep into the
coelomic cavity. In aberrant individuals without prostate glands
but with well developed muscular prostatic ducts there is no dis-
tinct level of demarcation between the latter and the deferent
duct. The gradual transition from one to the other suggests that
premature union of the ectal end of the deferent duct with the
ental end of the prostate rudiment, instead of at the side, may
have had something to do with absence of the prostate gland.
Shortly after establishment of the translocation mutation, ap-
pearance of another one may be assumed, this time for permissive
or facultative parthenogenesis. Due to the ability of a single
individual to reproduce when a copulatory partner was unavail-
able, the species may have been able to colonize new areas much
more rapidly than would have been possible otherwise. Certainly,
P. anomala has spread through most of Burma, probably to a
greater extent than any other species of the genus.
From the steady increase in number of earthworm species, in
various families, that are being found to have uniparental repro-
Gar BREVIORA No. 37
duction, a parthenogenetic mutation appears to be fairly common.
Though recognized in anomala hitherto only in association with
additional mutations to be mentioned below, conditions in other
species indicate that they can be independent of each other. Thus,
in forms like P. diffringens (Baird) 1869, which is now widely
spread throughout the United States, reproduction usually seems
to be parthenogenetic in spite of presence of a complete battery
of normal spermathecae and of normally developed male genital
terminalia.
If now a third mutation, ‘‘athecal’’, inhibiting development of
the spermathecae, were to arise in anomala at the appropriate
time, establishment of a true-breeding line would be expected.
As the atheeal individual could not receive sperm from a copula-
tory partner, all of its own offspring would be mutants. Offspring
of a normal copulatory partner would be in part athecal also and
if the mutation is dominant its rapid spread through the popu-
lation should be possible.
The athecal mutation also seems to be standard as the condition
has appeared in individuals of species belonging to various fam-
ilies but with especially annoying frequency in the genus Phere-
tima, depriving the taxonomist of a whole set of the very organs
most useful for his species identifications. The mutation has not
been found, in other species of the genus, in association with
extra testes. To obtain f. typica from a centralis ancestor, yet
another mutation, ‘‘testis’’, is postulated. This too may prove to
be standard as extra testes anterior to the usual gonad series
already have been found in species of two other families. If this
spawning of hypothetical mutations in a single line is to end here
it is necessary to conclude that mutation ‘‘testis’’ is not of the
apparently simple type of mutation ‘‘athecal’’. In addition to
bringing about development of extra gonads and adding two or
more pairs of mushroom glands to the centralis series it inhibits
production of sperm by any of the gonads.
To get f. insolita from an ancestor lke centralis, a mutation,
‘‘aprostatie’’, for inhibition of development of prostates, is re-
quired. This condition also appears throughout prostate-possess-
ing families of earthworms but has been noticed more often in
the genus Pheretima. Unless another mutation is to be postulated
it must be assumed that ‘‘aprostatic’’ also expresses itself by
1954. EVOLUTION OF AN ORIENTAL EARTHWORM 7
inhibition of development of the mushroom glands. For such an
assumption a certain amount of evidence is provided by aberrant
individuals in other species of the genus.
Mutations ‘‘athecal’’ and ‘‘aprostatic’’ have not yet been found
in the same individual of anomala but the two conditions have
appeared simultaneously in individuals of other species in the
genus, thus depriving the sorely tried taxonomist of the last sets
of organs required for species identifications. Association of the
two mutations already has been found in at least one of the dozen
species of Pheretima that have been accidentally introduced into
this country.
Another hypothetical mutation, ‘‘aseptal’’, has long been estab-
lished in anomala, as well as in many other species of the genus.
This partially or completely aborts, during embryonic develop-
ment or post-hatching growth, the transverse partition separating
the coelomic cavities of segments vili and ix. Abortion rather
than inhibition is indicated by persistent rudiments of varying
size while incomplete or delayed penetrance is shown by occasional
individuals in which a considerable portion of the septum is still
recognizable or in which the partition even has become muscular.
Yet another mutation, ‘‘uniporal’’, which has been involved in
the ancestry of most species of Pheretima, results in union of the
paired oviducts within the parietes so as to open to the exterior
by a single, median pore. Here again, incomplete or delayed
penetrance is suggested by the failure of the oviducts, in an ocea-
sional individual, to unite ectally thus resulting in a reversion
to the ancestral condition with paired female pores.
Postulating a mutation capable of bringing about addition of
five pairs of testes, along with five pairs of male deferent ducts —
in one step — ean be avoided in two ways at least. One alterna-
tive would be to assume that a mutation for addition of one or
two pairs of testes at the anterior end of the existing series was
repeated an appropriate number of times. Repetition of four
different one-step-at-a-time mutations now seems to be required
in the phylogeny of another genus currently under examination.
Another alternative is provided by the widely accepted evolution
of the Oligochaeta from marine Polychaeta during which diffuse
germinal tissue was strictly concentrated into one intrasegmental
location, within the septa close to the nerve cord, but in several
8 BREVIORA NO. 37
consecutive segments. Nine or ten pairs of gonads almost seem
to be required in oligochaete phylogeny by the families of Micro-
drili. Anlage of such gonads laid down early in embryonic devel-
opment of anomala, instead of being aborted throughout most of
the series as usual, could be permitted by mutation ‘‘testis’’ to
develop into a macroscopic discoidal stage. Origin of super-
numerary gonads in some of the segments can be attributed to
early embryonic fragmentation such as breaks a first nephridial
rudiment into early components of the enteronephric excretory
system in this same genus. Development of male funnels is how-
ever induced by the adjacent growing gonads. If the induction
effect is adequate, anlage not only become plicate funnels but
develop ducts which may even become continuous with those of
the posterior segments.
SUMMARY
Evolution of an advanced species from a more generalized
generic type and segregation within that species of three true-
breeding formae is attributed to the establishment of single effect
and multiple effect mutations. Hach of these postulated mutations
is of a standard sort, required by conditions in aberrant individu-
als appearing in species of several genera, or by phyletic develop-
ments that obviously have taken place in genera of different
families.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. NOVEMBER 28, 1954 NuMBER 38
ONNIA (TRILOBITA) FROM VENEZUELA
By H. B. WauHtrtTineton
In 1938 Leith described a Middle Ordovician fauna from the
Venezuelan Andes which included the graptolite Dicranograptus,
a eryptolithinid trilobite, and fragmentary brachiopods and pele-
eypods. Recently I received several specimens of the trilobite
collected from the original locality by Dr. G. R. Pierce, of the
Creole Petroleum Corporation, and sent to me by Dr. V. D.
Winkler and Mr. Kx. F. Dallmus, also of the Creole Petroleum
Corporation. I am deeply indebted to these gentlemen, and to
the Creole Petroleum Corporation for presenting the specimens
to the Museum of Comparative Zoology, and for permitting me
to publish this note on them. Dr. Karl M. Waagé, Yale Uni-
versity, kindly loaned the original specimen described by Leith.
The new material enables a more complete description to be
given, and shows that the species is related to contemporary
British, central and southern European, and North African
forms, rather than to North American ones. The terminology
is that previously employed (Whittington, 1940), with the
additions of Stauble (1953).
Family TRINUCLEIDAE Emmrich, 1884
Subfamily CRYPTOLITHINAE Whittington, 1941
Genus ONNIA Bancroft, 1933
ONNIA TERRYI (Leith, 1938)
Peel hiorcaelers
Holotype. Peabody Museum of Natural History, Yale Uni-
versity, no. 15317, from Caparo-Bellavista Series, “‘ first hill out-
side the Capar|r]o River floodplain, about three-quarters of a mile
south of the river, along the trail between Santa Barbara (State
2 BREVIORA NO. 38
of Zamora) and Mueuchachi (State of Mérida)’’ (Leith, 1938,
p. 338).
Additional Material. From the same locality near the Caparo
River, State of Barinas (—Zamora), collected by G. R. Pierce
in 1951-1952. The fossils are contained in a soft, grey-white,
iron-stained, micaceous siltstone, and preserved with some dis-
tortion as moulds.
Age. Middle Ordovician (Leith, 19388). Later collections have
not included additional species that might throw further heht
on the precise horizon.
Description. The holotype shows the mould of the inner sur-
face of the upper lamella of the fringe, but the new material
includes moulds of the inner and outer surfaces of both lamellae
of the fringe, as well as moulds of the pygidium and thorax,
and enables amplification of Leith’s description (1938, pp. 341-
342, fig. 2).
Glabella clavate, expanding forward; occipital ring narrow,
eonvex, long median spine, the shallow occipital furrow with
deep appendiferal pits laterally. Immediately in front of the
cecipital furrow the glabella appears to be exceptionally inflated
where it is crossed by the occiput (Pl. 1, figs. 2, 5), and in front
of the occiput glabellar furrows may have been present, as in
typical eryptolithinids. The distortion and crushing of the
specimens (PI. 1, figs. 1-3, 5) gives, however, a variable appear-
ance to this portion of the glabella. Axial furrows with anterior
pits. Cheek lobes descend steeply to fringe ; cheek frame straight
and transversely directed, fringe frame directed backward and
outward so that posterolateral part of fringe is wide and projects
posteriorly.
Fringe with outer surface of upper lamella sloping forward
anteriorly, but anterolaterally and laterally convex, so that there
is a depressed region adjacent to the cheek-lobe. Lower lamella
slopes outward, anteriorly flexed at girder. Long genal spine
directed backward and slightly outward. Girder a broad ridge
anteriorly, less conspicuous anterolaterally and laterally, where
a lower ridge (in the outer surface of the lower lamella) of
almost equal width runs between the two rows of pits (I, and
I.) immediately inside the girder, and a narrower, still lower
ridge separates the next inmost rows (Iz and I;; Pl. 1, fig. 2).
1954 ONNIA (TRILOBITA) FROM VENEZUELA 3
Near the genal angle these inner ridges die out, and the girder
runs into the channel in the under-side of the genal spine (PI. 1,
fig. 4). Pits of Ey smaller than those of E,, laterally and antero-
laterally in a continuous row, but anteriorly becoming irregular,
and about 10 pits from the midline Ey bifureates and Es. is
formed in this anterior region (PIL. 1, fig. 1). The pits of Es
are smaller and more numerous than those in Es, and are not
radially arranged with them. E, and I, are the largest pits in
the fringe, in regular rows disturbed only at the genal angele,
and anteriorly and laterally radially arranged with each other
and the rows inside. Is complete in front of the glabella, I.
commencing at about the 4th pit of I, from the median line, Iy
commencing anterolaterally, and additional rows laterally. In
the inner, posterior part of the fringe concentric arrangement
is replaced by quincuncial, while radial arrangement is con-
spicuous laterally and anterolaterally, especially in the de-
pressed inner region. Low concentric ridges separate the pits of
Ky and E,, E, and I,, 1, and Is, especially laterally and antero-
laterally. In this same region radial ridges separate the pits
of E, and the rows inside it, and such ridges separate the pits of
the internal region anteriorly. Numbers of pits in left side of
fringe in holotype and original of Plate 1, figure 1, are respec-
tively: Es, 37 and 32; E,, 31 and 29; I,, 30 and 29; I., 32 and
30. No ornament is preserved on the moulds of the cheek lobes
or glabella.
Four segments of the characteristic thorax poorly preserved
as external mould (PI. 1, fig. 6). Pygidium (PI. 1, figs. 6, 7)
subtriangular in outline, low axis reaching to tip, pleural regions
flattened, with narrow, ridge-lke border and steeply-descending
margins. First axial ring and pleural furrow distinct, remainder
of axis not subdivided, but two or three additional pleural fur-
rows and interpleural ridges may be discerned running straight
outward and backward.
Discussion. This species is clearly a eryptolithinid (Whitting-
ton, 1941, pp. 23-25) but is distinguished from Cryptolithus by
the following: (a) There are two complete rows of pits outside
the girder, and a third row anteriorly. (b) Characteristic radial
ridges do not separate the pits in the outer row.
The pits of Es are smaller than those of E,, especially later-
4 BREVIORA No. 38
ally, and here the upper lamella is convex upward, with a de-
pressed region adjacent to the cheek lobe. These same characters
distinguish the Venezuelan species from species of Crypto-
lithoides (Whittington, 1941), and in addition the cephalon of
Cryptolithoides has a different outline, and the pits inside the
outer two rows are irregularly arranged laterally. While the
Venezuelan material shows some features suggestive of Broeg-
gerolithus (see Whittington, 1941, p. 24; Bancroft, 1949, pp.
298-299, Pl. 9, figs: 4; 5, 6, 8, Pl. 11, fig. 38); but has more rows
of pits in the fringe and lacks the strong radial ridges in Kj,
it is much more like species of Onnia (Whittington, 1940, Pl. 3;
Lamont, 19485 Pl i; Bancroft: 1949. p: 299, (Pl. 9 fhicsaO alae
Pl. 10, fig. 16). These latter have Ey of small pits, E, and I,
of larger pits, and the region within I, showing a strong radial
arrangement, particularly in the depressed lateral region. The
girder in Onnia is strong anteriorly, but laterally concentrie
ridges of almost equal strength may run between E, and Eps,
and/or I, and I, (e.g. Whittington, 1940, Pl. 3, fig. 5; Lamont,
1948, Pl. 1, figs. 4,5). O. terryi shows a comparable development
of ridges in the outer surface of the lower lamella in the interior
region (Pl. 1, figs. 2, 4). O. terry: differs from other species of
Onnia in the presence of Es anteriorly, but this single feature
scarcely seems to warrant the erection of a separate genus. The
development of additional external rows of pits anteriorly occurs
in the probably older genus Salterolithus (Bancroft, 1949, p. 292,
Pl. 9, figs. 1, 2), but species of this genus have fewer rows of
pits internal to the girder.
Onnia is known from Middle and Upper Ordovician rocks in
central and northern Britain, Bohemia, and as far south as North
Africa (Termier and Termier, 1950, Pl. 187, figs. 8-11). The
occurrence of Onnia in Venezuela suggests the possibility of a
faunal province extending from central Europe and North
Africa to northern South America in Ordovician times. Such
a province may have passed through what is now Florida, for
Colpocoryphe exsul Whittington, 1953, from early Middle Ordo-
vician rocks of Florida, also has central European-North Afri-
ean affinity. If such a province existed, it is notable that it
appears to lie almost entirely east and south of the belt of
Upper Cambrian and early Ordovician ‘‘ Atlantic faunas’’ re-
cently portrayed by Wilson (1954, fig. 4), and that the rocks
are mainly elastic.
1954 ONNIA (TRILOBITA) FROM VENEZUELA D
REFERENCES
BANCROFT, B. B.
1949. Upper Ordovician trilobites of zonal value in southeast Shrop-
shire. Proce. Roy. Soc., London, Ser. B., vol. 136, pp. 291-315,
pls. 9-11.
LAMONT, A.
1948. B. B. Bancroft’s geological work. 2. Upper Ordovician of the
Cross Fell inlier. Quarry Manager’s Jour., vol. 31, pp. 416-418,
466-469, pl. 1.
LeitH, E.
1938. A Middle Ordovician fauna from the Venezuelan Andes. Amer.
Jour. Sei., 5th ser., vol. 36, pp. 337-344, pl.
TERMIER, G. and H.
1950. Paléontologie Marocaine, vol. 2. Actual. Sci. Industr. no. 1095,
Paris.
STAuBLE, A.
1953. Two new species of the family Cryptolithidae. Natur. Canad.,
vol. 80, pp. 85-119, 201-220, figs. 1-24.
WHITTINGTON, H. B.
1940. On some Trinucleidae described by Joachim Barrande. Amer.
Jour. Sci., vol. 238, pp. 241-259, pls. 1-4.
1941. The Trinucleidae — with special reference to North American
genera and species. Jour. Paleont., vol. 15, pp. 21-41, pls. 5, 6.
1953. A new Ordovician trilobite from Florida. Breviora, no. 17,
pp. 1-6, pl. 1.
WILSON, JAMES L.
1954. Late Cambrian and early Ordovician trilobites from the Marathon
Uplift, Texas. Jour. Paleont., vol. 28, pp. 249-285, pls. 24-27.
TXPLANATION OF PLATE
Onnia terryi (Leith, 1938). Caparo-Bellavista Series, Middle Ordovician,
near the Caparo River, State of Barinas, Venezuela.
Fig. 1. Rubber cast from external mould of cephalon, showing upper
lamella of fringe, X 4, MCZ 5183a. Extremities of third external row of
pits indicated by ‘‘ E3.’’
Fig. 2. Mould of internal surface of glabella and cheek lobes, external
surface of lower lamella of fringe, X 3, MCZ 5183b. Position of first (Ey)
and second (E») external row of pits, girder (g), and of first (11), second
(Io), and third (Ig), internal rows shown.
Fig. 3. Holotype, original of Leith (1938, fig. 2), internal mould of gla-
bella, cheek lobes and inner surface of upper lamella of fringe, X 2, Peabody
Museum, Yale University, 15317. Extremity of third external row indicated
by SSRiss72
Fig. 4. Mould of external surface of right cheek lobe, lower lamella of
fringe and genal spine, X 3, MCZ 5183c.
Fig. 5. Internal mould of cephalon showing upper lamella of fringe.
X 214, MCZ 5183d.
Fig. 6. Rubber cast from external mould of pygidium and four poorly
preserved thoracic segments, X 3, MCZ 5183e.
Fig. 7. Rubber cast from external mould of pygidium, X38, MCZ 5188f.
Ripe.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, MAss. NUMBER 39
NOVEMBER 24, 1954
NEW OR REDESCRIBED PEIGOMEDUSID
SKULLS FROM THE TERTIARY OF AFRICA
AND ASIA
(TESTUDINES, PELOMEDUSIDAE)
2. A Podoenemide Skull from the Miocene of Moghara, Egypt.
By Ernest WILLIAMS
In 1952-1953 while visiting the British Museum (Natural
History) as a Guggenheim Fellow I was privileged to examine
an undescribed skull from the Moghara Miocene of Egypt. The
skull evidently belonged to a member of the genus Podocnemis
sensu lato or to a close relative of that genus; the critical recogni-
tion marks of this section of the Pelomedusidae — the enlarged
‘carotid’? channel, and the contact of basioceipital and quadrate
—were readily visible. Further examination left no doubt of
the novelty of this fossil as compared with any previously known
podoenemide! skull, and suggested interesting problems in regard
to its proper correlation with a shell name and in regard to its
phyletie position.
The British Museum Moghara skull is unfortunately im-
perfect in many respects. The snout is missing so that important
characters and relations here cannot be checked. Major parts of
the postorbitals and jugals are broken away, as are both squa-
mosals, the quadrate of one side, parts of the parietals and the
occipital condyle.
Many structural details of taxonomic and phyletie importance
are therefore not ascertainable: the presence or absence of a
complete temporal roof, the presence or absence of a vomer, the
position of the foramina incisiva, the relations of the internal
laminae of the premaxillae and maxillae to each other and to
1I use the term ‘‘podocnemide”’ as a convenient and informal designation for
a subsection of the family Pelomedusidae typified by the genus Podocnemis.
2 A PODOCNEMIDE SKULL FROM MOGHARA 1954
the echoanal margin, the presence or absence of an anterior
process of the palatines dividing the choanal opening, the pres-
ence or absence of a median pit in the palatal surface of the
premaxillae and of a hooked process anteriorly on the premaxil-
lae.
The following points may, however, be made out: The skull
must have been rather broad, certainly very little longer than
broad. The interparietal scale marked off by grooves on the
skull roof is broad also, but tapers posteriorly, so that the parietal
scales must have met behind it. There is no hint of a forehead
groove but perhaps too little of the interorbital region is pre-
served. The orbits are visible in dorsal view.
There were two triturating ridges on the posterior portion of
the palatal surface of the maxilla —a low, rough, median ridge
and, parallel to it, a still lower, rougher, ridge at the margin of
the choanae. There was no extreme development of a secondary
palate.
Grooves on the postorbital bar indicate the presence of a
‘*suboeular’’ seute in the sense of Siebenrock (1902).
The ‘‘earotid’’ channel is fully enlarged in podoenemide
fashion.
The ectopterygoid processes are large, blunt and almost wholly
lateral in orientation. The basisphenoid is a conspicuous triangu-
lar element, the anterior apex blunted. There is a narrow
basioccipital quadrate contact, more longitudinal than trans-
verse.
The cavum tympani is large and lacks a precolumellar fossa.
These determinable characters are few indeed on which to
hazard an estimate of the relationships of the Moghara form.
One method of appraisal, however, is to tabulate the more
diagnostic of these characters against the characters of other
known podoenemide skulls. Tabulating first against the living
podoenemide species (Table 1), we find that very little is learned
except that the British Museum Moghara skull is not exactly like
that of any modern podocnemide. If, now, we compare the
British Museum Moghara skull with the previously known fossil
skulls of podoenemide type (Table 2), the impressive fact is the
close agreement, in cited characters, of the British Museum skull
and Dacquemys paleomorpha. We need in fact to inquire what
characters assure us that we are not dealing with Dacquemys.
3
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1954
MOGHARA
A PODOCNEMIDE SKULL FROM
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NO. 39 BREVIORA 5
It is unfortunate that the unique generic character of Dac-
quemys — the long squamosal-parietal suture — cannot be veri-
fied or denied for the Moghara specimen. This region has been
broken away.
There are, however, differences which may or may not imply
generic distinctness. The British Museum skull is broad; that of
Dacquemys is elongate. The orbits are visible dorsally in the
Miocene specimen; they are completely concealed in Oligocene
Dacquemys. The two triturating ridges are low and rough in
the Moghara specimen, the inner parachoanal ridge especially so ;
the same ridges are high, smooth and sharply defined in Dac-
quemys.
The suggestive point about these differences is the fact that in
each case the Moghara skull is closer in these characters to
Erymnochelys (=Podocnemis madagascarensis of Boulenger )
than to Dacquemys. The skull of Erymnochelys madagascarensis
is relatively broad, the orbits are exposed dorsally, there is but
one low ridge on the triturating surface of the maxilla. But in
regard to the last character there is some roughening of the
parachoanal border of the maxilla in Hrymnochelys; the condi-
tion in that genus could be explained as a further carrying
through of a trend initiated in the British Museum skull. In
fact the British Museum skull might on all its characters be
interpreted as an intermediate between Dacquemys and Erymno-
chelys, perhaps somewhat closer to Hrymnochelys.
If we assume the reality of this intermediate phyletic station
of the British Museum Moghara skull, it is then probable that
this skull belongs with the common podocnemide shell of the
Moghara deposit, the shell type named by Andrews Podocnemis
aegyptiaca, which has every shell character of Hrymnochelys
and is on shell characters barely, if at all, distinguishable from
Erymnochelys madagascarensis. | shall hereafter refer to Podoc-
nemis aegyptiaca Andrews as Erymnochelys aegyptiaca (An-
drews).
There is only one other podoenemide shell type known from
Moghara — the form named by Fourtau (1920) ‘* Podocnemis’’
bramly:. This form differs from Hrymnochelys aegyptiaca and
from all Erymnochelys in the larger size of the intergular scale
which separates the gulars as in the Recent South American
6 A PODOCNEMIDE SKULL FROM MOGHARA 1954
podoenemide species (Podocnemis sensu stricto and most speci-
mens of Peltocephalus). If we could assume that P. bramlyi is
a precursor of Peltocephalus then it would not be too anomalous
for our Moghara skull to belong to this species, rather than to
FE. aegyptiaca. In the living species Peltocephalus dumeriliana
(in which the gular-intergular pattern is typically very similar
to that of P. lramlyi) the skull has definite, strong similarity
to that of Erymnochelys or of Dacquemys and thus also to the
Moghara skull.
I know of no grounds for decision between the two alternatives
thus presented. The British Museum skull may as _ plausibly
belong to P. bramlyi as to FE. aegyptiaca. We have too little of
the Moghara skull, and in addition we are probably too close to
the branching off point of Peltocephalus and Erymnochelys to
expcet wide differences in skull structure between these two, then
nascent, genera. But, though we must thus remain undecided as
to the species allocation of the Moghara skull, I think that one
positive statement of some importance may be made. The Mog-
hara skull — on whichever fork of the phyletic tree it belongs —
is a structural intermediate between the Recent genera, Pelto-
cephalus or Erymnochelys, and the Oligocene Dacquemys.
This, indeed, is the principal suggestion that I wish to make:
that there is a phyletic relationship between Peltocephalus,
Krymnochelys and Dacquemys of the sort diagrammed below
(Gigs db)e
Peltocephalus — Recent of Erymnochelys — Recent of
dumeriliana S. America madagascarensis Madagascar
"Podocnemis" bramlyi Erymnochelys — Miocene of
Miocene of Egypt aegyptiaco Egypt and
E. Africa
Dacquemys — Oligocene of
paleomorpha -—_ Egypt
\
? Dacquemys
fajumensis
Fig. 1 Presumed phyletic position of the Egyptian Miocene podocnemides
No. 39 BREVIORA ra
It cannot be demed that this suggested phylogeny rests upon
a rather elaborate structure of inferences, any or all of whieh
may be wrone; nothing else, however, is possible in the current
state of the evidence.
It may be useful, here, to compare this suggestion with the
previous phyletic scheme for ‘‘Podocnemis’’ which was set up
by Daequé and has been accepted by Zangerl. The points of
agreement will be found to be more numerous than at first seems
the case. We are agreed that there are two lines within the pelo-
medusid species grouped as Podocnemis by Boulenger, Andrews
and others —an African and a South American line. | differ
with Dacqué, Zangerl and others in that I separate from the
South American group the species dumeriliana (type of Pelto-
cephalus) and regard the few resemblances of the latter species
to the other South American forms — similarity in gular pattern
and in cervical articulations —as convergent only. I further
postulate a considerable evolution in skull structure within the
line which we may still, for convenience, call African — an evolu-
tion involving so much morphological divergence that I recognize
three genera within this lineage.
Of these two respects in which I differ with the Dacqué-Zanger|
phylogeny, the recognition of African affinities in Peltocephalus
dumer.diana is the more solidly founded. I shall argue this point
at leneth in a future discussion of the rank, characters and varia-
tion of the livine species placed by Boulenger in Podocnemis.
The second conception — that of the Dacquemys-Erymnochelys-
Peltocephalus series — is as vet hypothesis only, and the grounds
for this view have been stated here.
It needs to be mentioned that Dacqué (1912) described another
skull from Moehara which he named Sternothaerus blancken-
horm. The deseription is very brief and the only figure is a dorsal
view. The skull was also quite imperfect, lacking the temporal
and occipital regions,
If it belongs to Sternothaerus (—Pelusios) — an assignment
neither contradicted nor specially supported by the figure — it
has, of course, nothing to do with the skull whieh has here been
described. However, in dorsal view there would be few differ-
ences in such imperfect fragments as these two skulls from
Moghara; | think, nevertheless, that the interorbital width is
ereater in the skull which has been here discussed than in the
8 A PODOCNEMIDE SKULL FROM MOGHARA 1954
type of blanckenhorni. Beyond this, we must rely on Dacqué’s
description of differences, and his most significant remark ap-
pears to be his observation: ‘‘ Bemerkenswert erscheint, dasz die
Maxillaria die Choane auf der Schadelunterseite offenbar ganz
uberdecken.’’? This remark would appear to imply a strongly
developed secondary palate, a striking difference from the skull
here noticed; maxillary ridges are also unmentioned. It must be
commented that a strong secondary palate would be remarkable
also in the genus Pelusios and resembles more closely conditions
in the Stereogenys-Shweboemys series (which will be discussed
in the concluding paper of this series), but in any event ‘‘Sterno-
thaerus’’ blanckenhorni would appear to have no bearing on
the problems raised by the skull in the British Museum.
I am indebted to Dr. W. E. Swinton for the privilege of
examining and describing the British Museum skull from Mog-
hara. The photographs of this specimen are published with the
permission of the Trustees of the British Museum; they were
made by Peter Green of the British Museum staff.
PAPERS CITED
ANDREWS, C. W.
1906. A descriptive catalogue of the Tertiary vertebrata of the Fayam.
London, 319 pp.
Dacauk, E.
1912. Die fossilen Schildkréten Aegyptens. Geol. Palaeont. Abhandl.,
vol. 14, pp. 275-337.
FourtTau, R.
1920. Contribution a 1’étude des vertébrés miocénes de l’Egypt. Cairo
(Government Press), pp. 29-34.
SIEBENROCK, F.
1902. Zur Systematik der Schildkréten-Gattung Podocnemis Wagl.
Sitz. Ber. K. Akad. Wiss. Wien., Math.-Nat. K1., vol. 3, pp. 1-14.
WILLIAMS, E.
1954. New or redescribed pelomedusid skulls from the Tertiary of
Africa and Asia (Testudines, Pelomedusidae). 1. Dacquemys
paleomorpha, new genus, new species, from the Lower Oligocene
of the Fayum, Egypt. Breviora, no. 35, pp. 1-8.
ZANGERL, R.
1948. The vertebrate fauna of the Selma formation of Alabama. Part
Il. The pleurodiran turtles. Fieldiana: Geol. Mem., vol. 3,
pp. 28-56,
PIA Hs
British Museum skull from Moghara, dorsal view. Posterior end at top of
page.
PLATE 2.
British Museum skull from Moghara, ventral view. Posterior end at top of
page.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. November 29, 1954 NuMBER 40
SOME MOLLUSKS FROM THE
CONTINENTAL SLOPE OF NORTHEASTERN
NORTH AMERICA*
By ArtHurR H. CuARKE, JR.
The continental slope area of northeastern North Ameriea
has always been a relatively unknown and unexplored region.
Only meager information has been on record in regard to the
kind and abundance of its fauna. More knowledge is desirable,
not only for academic interest, but also that proper evaluation
may be made with the ultimate aim of supplementing man’s ever-
increasing need for additional sources of food.
With these objects in mind, in 1952 and 1953 the Woods Hole
Oceanographic Institution began a survey of the bottom fauna
existing on the continental slope area. An eighty-three foot
trawler, the Cap’n Bill IT was.equipped for the task, and bottom
trawlings were made along the slope from a point east of Cape
Charles, Virginia, to a point southeast of Nova Scotia in various
depths ranging approximately from 100 to 700 fathoms. Otter
trawls were used carrying one and one-half inch mesh at the
cod end, and measuring thirty-five feet, fifty feet, and sixty-
five feet initially across the mouth. The work was under the
direction of Mr. W. C. Schroeder of the Woods Hole Oceano-
eraphic Institution, ,to whom we are indebted for the fine array
of mollusks collected.
Mollusks were not the primary concern of the survey, and
the large size of the mesh prevented retention of the smaller
specimens, though many of the larger species were obtained.
These were submitted to Dr. W. J. Clench, Curator of Mollusks
at the Museum of Comparative Zoology to be added to the
research collection,
1 Contribution No. 722 from the Woods Hole Oceanographic Institution.
i)
BREVIORA No. 40
After identification of the mollusks collected, a search of the
literature was made so that the data gained from this survey
could be compared with previous records. Two papers by A. E.
Verrill (1884 and 1885) were found listing the bathymetric
ranges of all of the then known western Atlantic species of
marine mollusks found below 60 fathoms.
Verrill’s records have apparently been generally overlooked
by many of the more recent compilers of molluscan lists. His
data, as obtained primarily from material gathered by the U.S.
Fish Commission steamers Fish Hawk and Albatross from 1880
to 1884, indicate much more extensive bathymetric ranges for
a great many species than have subsequently been given. W. H.
Dall, in his ‘‘Preliminary Catalog .. .’’ (1889) omitted much
of Verrill’s data, and although Dall frankly stated that his
catalog was incomplete, this oversight has been perpetuated by
later writers. It is urged that future investigators refer to
Verrill’s most complete and excellent list as a supplement to
modern lists.
In the following tabulation of species collected by the Cap’n
Bill IT, the minimum depth range indicated by the data is given.
For instance, if a particular species was found at two stations
where the depths of trawling ranged from 200 to 250 fathoms
and from 300 to 350 fathoms respectively, the minimum range
would be 250 to 300 fathoms. A distinction is also made between
living and dead specimens, and bathymetric ranges are given
for each, if not identical. For comparative purposes, the broad-
est bathymetric and geographic ranges indicated by the litera-
ture are also included, and information sources are cited. When
veographie ranges are extended, these localities are defined in
terms of miles from a position on land. When not extended,
approximate location only is indicated.
By examination of the following list of 98 species, it will be
seen that these new data have extended the previously recorded
bathymetric ranges of seven species (marked *) and the geo-
eraphie ranges of six species (marked 7).
List of Mollusea Collected
YOLDIA THRACIAEFORMIS (Storer) *+
One large dead specimen at station 72. Bathymetric range:
400 fms. dead (this survey) ; 29-182 fms. dead (Verrill 1884).
(eX)
1954 MOLLUSKS FROM NORTHEASTERN NORTH AMERICA
Geographic range : 240 mi. east of Sandy Hook, New Jersey (this
survey) ; Gulf of St. Lawrence to Massachusetts (Johnson 1934) ;
West Greenland (Thorson 1951).
PECTEN VITREUS (Gmelin)
Three living and two dead specimens at station 110. Three
living specimens at station 208. Bathymetric range : 250-320 fms.
(this survey); 50-800 fms. (Dall 1889). Geographic range:
Aretie Ocean (Dall 1889); off Newfoundland and Nova Scotia
(Johnson 1934) ; south of Nova Scotia (this survey ) ; off Martha’s
Vineyard (Johnson 1934) ; east of Sandy Hook, New Jersey (this
survey) ; west Florida [Gulf of Mexico] (Dall 1889); Europe;
Mediterranean Sea; Africa (all Verrill 1884); west Greenland
(Thorson 1951).
ANOMIA ACULEATA Miller
Living specimens attached to shells and rocks at stations 14,
19, 38, 53, 110, 131, 173, and 195. Bathymetric range: 145-320
fms. (this survey) ; 4-640 fms. (Verrill 1884) ; 0-80 fms. (Dall
1889). Geographic range: Arctic Ocean to Cape Hatteras, North
Carolina (Dall 1889); south of Nova Scotia to east of Cape
Henlopen, Delaware (this survey) ; Europe (Dall 1889).
A. very variable species. The normal imbricated or sealy sur-
face sculpture is entirely lacking in many individuals and in
some entire large lots of specimens.
ARCTICA ISLANDICA (Linné)
Three single valves at station 1. Bathymetric range: 33 fms.
dead (this survey) ; 6-90 fms. (Johnson 1934). Geographic range :
east of Montauk Point, Long Island, New York (this survey) ;
Arctie Ocean to Cape Hatteras, North Carolina (Dall 1889).
ASTARTE SUBAEQUILATERA Sowerby
Three small living specimens at station 86. Bathymetric range:
230 fms. (this survey) ; 22-410 fms. (Johnson 1934). Geographic
range: Labrador to Florida (Johnson 1934); south of Nova
Seotia (this survey).
PANOMYA ARCTICA (Lamarck) +
One very large, apparently freshly dead specimen at station
38. Bathymetric range: 190 fms. dead (this survey) ; 20-506 fms.
4 BREVIORA No. 40
dead: 300 fms. alive (both, Verrill 1884). Geographic range:
Arctic Ocean to Georges Bank; cireumpolar (both, Johnson
1934) ; 170 mi. east of Barnegat Bay, New Jersey (this survey).
The specimen cited measures as follows: length 109 mm., height
74 mm., width (valves together) 48 mm. A similar specimen,
alive and nearly as large, was taken nearby by the Caryn in 1949
(39°57’N., 70°38’W.), thereby confirming the fact that the
species exists alive in this area.
CALLIOSTOMA BAIRDI Verrill and Smith
One dead specimen at station 83. One living specimen at sta-
tion 146. Bathymetric range: 87 fms. dead; 65 fms. alive (both
this survey) ; 56-640 fms. dead; 64-192 fms. alive (both, Verrill
1884). Geographic range: south southeast of Nantucket (270 mi.
east of Barnegat Bay, New Jersey) ; east of Cape Henlopen, New
Jersey (both, this survey) ; south of Martha’s Vineyard to Flor-
ida Keys (Johnson 1934); Caribbean Sea (Verrill 1889).
The living specimen is very beautiful. Patches of lavender are
seen through the thin iridescent periostracum which appears
golden between the spiral rows of tubereules on the upper whorls
and golden peach on the lower whorls. The inner nacre of the
aperture shines with iridescent pink and green. The cireular
operculum is multispiral, typical of the family.
SOLARIELLA OTTOI (Philippi)
One dead specimen at station 72. Three living specimens at
station 211. Bathymetric range: 400 fms. dead; 600 fms. alive
(both, this survey) ; 64-1555 fms. (Dall 1889). Geographic range:
south of Martha’s Vineyard and Georges Bank (Johnson 1934) ;
east of Sandy Hook, New Jersey (this survey); New Jersey;
Virginia (both, Dall 1889); Florida Keys; West Indies (both,
Johnson 1934); St. Thomas, Virgin Islands (Dall 1889).
Natica cLAUsA Broderip and Sowerby
One very large dead specimen (23 mm. long) at station 182.
Bathymetric range: 280 fms. dead (this survey) ; 13-1255 fms.
dead; 238-843 fms. alive (both, Verrill 1884); 16-1537 fms.
(Johnson 19384). Geographic range: Arctic [Ocean] (Verrill
1884) ; Labrador to off North Carolina (Johnson 1934) ; south-
1954 MOLLUSKS FROM NORTHEASTERN NORTH AMERICA 5
east of Cape Sable, Nova Scotia (this survey) ; Europe (Verrill
1884) ; Arctic and Bering seas to San Diego, California; Japan
(both, La Roeque 1953).
POLINICES HEROS (Say)
One dead specimen at station 144. Bathymetric range: 150
fms. dead (this survey) ; 0-238 fms. (Dall 1889). Geographic
range: Gulf of St. Lawrence to North Carolina (Johnson 1934) ;
east of Chincoteague Island, Virginia (this survey).
POLINICES TRISERIATA (Say) *
One very large, dead specimen (33 mm. long) at station 144.
One dead specimen at station 200. Bathymetric range: 100-150
fms. dead (this survey); 0-63 fms. (Dall 1889). Geographic
range: Labrador to Cape Hatteras (Dall 1889) ; south of Georges
Bank to east of Chincoteague Island, Virginia (this survey).
CAPULUS UNGARICUS (Linné)
One small living specimen attached to Pecten vitreus at station
110. Bathymetric range : 320 fms. alive (this survey) ; 1-458 fms.
(Johnson 1934). Geographic range: south southeast of Cape
Sable, Nova Scotia (this survey) ; Greenland to Florida (John-
son 1934) ; Europe (La Roeque 1953).
APORRHAIS OCCIDENTALIS (Beck) *
Living specimens at stations 95 and 177. Dead specimens at
stations 38, 83, 86, 95, 164, 176, 182, and 184. Bathymetric range :
340-360 fms. alive, 90-365 fms. dead (both, this survey) ; 115-349
fms. alive; 3414-1000 fms. dead (both, Verrill 1884). Geographic
range: Gulf of St. Lawrence to off North Carolina (Johnson
1934) ; southeast of Nova Scotia to south of Nantucket (this
survey); west Greenland (Thorson 1951).
BuccINuM cILIATUM Bruguiere
One small living specimen at station 184 (southeast of Cape
Sable, Nova Scotia). Bathymetric range: 265 fms. alive (this
survey ) ; 26-471 fms. (Johnson 1934). Geographic range: Arctic
[Ocean] (Verrill 1884) ; Labrador to Cape Cod, Massachusetts
(Johnson 1934); southeast of Cape Sable, Nova Scotia (this
survey); Europe (Verrill 1884).
6 BREVIORA no. 40
NEPTUNEA DECEMCOSTATA (Say) *+
Living specimens at stations 88, 95, 110, 165, 173, 182, and
184. Dead specimens at stations 38, 72, 86, 104, 164, 187, and
195. Bathymetric range: 270-3860 fms. alive; 225-660 fms. dead
(both this survey) ; 41-86 fms. alive; 6-322 fms. dead (both, Ver-
rill 1884). Geographic range: Cireumboreal (La Roeque 1953) ;
Nova Scotia to Massachusetts Bay and Georges Bank (Johnson
1934) ; southeast of Nova Scotia to 240 mi. east of Barnegat Bay,
New Jersey (south of Nantucket) (this survey).
Great variation of form exists in this species. Some specimens
are attenuated to the extent that the length is 21% times the great-
est width. Other specimens show a length of only 114 times the
width. Variation is also seen in degree of sculpture, length of
aperture in relation to overall length, ete. The possibility of the
existence of one or more eclines will be investigated.
NEPTUNEA DESPECTA TORNATA (Gould) *}
One small, dead specimen at station 159. Bathymetric range:
690 fms. dead (this survey); 10-471 fms. (Johnson 1934).
Geographic range: Gulf of St. Lawrence to off Martha’s Vine-
yard (Johnson 1934) ; 120 mi. east of Atlantic City, New Jersey
(this survey).
CoLus ISLANDICUS (Gmelin)
Dead specimens at stations 5, 22, 23, 29, 84, 144, 201, and 225.
Bathymetric range: 85-415 fms. dead (this survey) ; 20-1650 fms.
(Dall 1889). Geographic range: Labrador to Norway (Johnson
1934); Aretic Sea to South Carolina (Dall 1889), south of
Georges Bank to east of Chincoteague Island, Virginia (this
survey ).
COLUS STIMPSONI STIMPSONI (Morch)
Two living specimens at station 152. One dead specimen at
station 201. Bathymetric range: 330 fms. alive; 85-87 fms. dead
(both, this survey); 1-471 fms. (Johnson 1934). Geographic
range: Arctic Sea to [Cape] Hatteras, [North Carolina] (Dall
1889) ; south of Georges Bank to east of Cape Henlopen, New
Jersey (this survey).
CoLUS PUBESCENS (Verrill)*
Living specimens at stations 10, 13, 17, 53, 72, 139, 182, and
1954 MOLLUSKS FROM NORTHEASTERN NORTH AMERICA 7
184. Dead specimens at stations 8, 10, 14, 17, 19, 35, 49, 83, 95,
104, 114, 131, 182, 139, 144, 164, 173, 184, and 225. Bathymetric
range: 130-400 fms. alive; 90-415 fms. dead (both, this survey) ;
18-179 fms. dead; 192-640 fms. alive (both, Verrill 1884). Geo-
graphic range: Gulf of St. Lawrence to North Carolina (Johnson
1934) ; Nova Scotia to South Carolina (Dall 1889) ; southeast of
Nova Scotia to east of Chincoteague Island, Virginia (this sur-
vey).
PLEUROTOMELLA AGAssIzI Verrill and Smithy
One dead specimen at station 211. Bathymetric range: 600
fms, dead (this survey); 39-1309 fms. alive; 1608 fms. dead
(both, Verrill 1884). Geographic range: 105 mi. southeast of
Nantucket (this survey); south of Martha’s Vineyard to the
West Indies (Johnson 1934).
SCAPHANDER PUNCTOSTRIATUS (Mighels and Adams)
One living specimen at station 109 and one living specimen at
station 189. Bathymetric range: 240-305 fms. alive (this sur-
vey) ; 46-1255 fms. alive; 1362-1467 fms. dead (both, Verrill
1884). Geographic range: Gulf of St. Lawrence to West Indies
(Johnson 1934); southeast of Nova Scotia (this survey); Bar-
bados; Norway (both, Dall 1889).
RosstA SUBLAEVIS Verrill
One living specimen at station 69. Bathymetric range: 490
fms. (this survey); 45-640 fms. (Johnson 1934). Geographic
range: Newfoundland to 32°33’15”’N. (Johnson 1934) ; south of
Georges Bank (this survey).
ILLEX ILLECEBROSA (Lesueur)
Two living specimens at station 68 and one living specimen at
station 74. Bathymetric range: 450 fms. (this survey) ; 0-1022
fms.; beaks 1091-1917 fms. (both, Verrill 1884). Geographic
range: south of Georges Bank (this survey) ; Greenland to Cape
Hatteras (Johnson 1934).
CHIROTEUTHIS LACERTOSA Verrill
One living specimen at each of the following stations: 68, 69,
and 74. Bathymetric range: 490-500 fms. (this survey) ; 435-
2369 fms. (Johnson 1934); arms, 2949 fms. (Verrill 1884).
8 BREVIORA no. 40
Geographic range: south of Georges Bank (this survey) ; Nova
Scotia to West Indies (Johnson 1934).
MASTIGOTEUTHIS AGASsIzi. Verrill*
One living specimen at station 102. Bathymetric range: 600
fms. (this survey) ; 640-1050 fms. (Johnson 1934). Geographic
range: Gulf of Maine to North Carolina (Johnson 1934) ; south-
east of Nova Scotia (this survey).
ALLOPOSUS MOLLIS Verrill}
One living specimen at station 69 (south of Georges Bank)
and one living specimen at station 90 (southeast of Nova Scotia).
Bathymetric range: 330-490 fms. (this survey) ; 238-1346 fms. ;
fragment, 1735 fms. (both, Verrill 1884). Geographic range:
110 mi. southeast of Cape Sable, Nova Scotia; south of Georges
Bank (both, this survey) ; Nantucket to Chesapeake Bay, eastern
Atlantie (both, Johnson 1934).
BATHYPOLYPUS ARCTICUS (Prosch)
One living specimen at station 69 and one living specimen at
station 72. Bathymetric range: 450-490 fms. (this survey) ; 28-
843 fms. (Johnson 1934). Geographic range: south of Georges
Bank (this survey) ; Bay of Fundy to 32°N. (Johnson 1934).
GRANELEDONE VERRUCOSA (Verrill)
One living specimen at station 69. Bathymetric range: 490
fms. (this survey) ; 466-1255 fms. (Johnson 1934). Geographic
range: south of Georges Bank (this survey); Nova Scotia to
Delaware Bay (Johnson 1934).
Location and Depth of Stations Cited
Station Number North Latitude West Longitude Depth in Fathoms
1 40°50’ 71°09’ 33
5 39°35’ 71°57’ 240-260
8 39°36’ 71°52’ 405-410
10 39°46’ 71°35" 395-405
13 39°55’ 71°27’ 180-190
14 39°56’ 71°26’ 150
yi 39°55’ (ie Ni 290-330
19 39°57’ ile ie 175-180
22 39°53’ 70°53’ 300-330
1954
102
104
109
110
114
131
132
139
144
146
152
159
164
165
173
176
177
182
184
187
189
195
200
201
208
211
225
MOLLUSKS FROM NORTHEASTERN NORTH AMERICA
39°52!
39°52’
40°02"
40°04’
39°57"
391.55"
40°08’
40°10’
40°12’
40°17’
39°59’
39°59’
42°23’
42°46’
42°41’
42°45’
42°35!
42°40’
42°20’
42°17’
40°46’
38°13’
38°08’
37°38"
37°45!
38°33!
38°39’
39°26’
42°43’
42°42’
42°40’
42°33"
42°32’
42°28’
42°23’
42°15’
42°18’
40°34’
40°46’
40°45’
40°09"
40°00’
39°53"
70°51"
70°43’
70°24"
Uae
69°37’
69°26"
68°19’
68°16’
68°08’
67253)
69°32’
69°35"
64°58"
63°22"
63°33"
63°47’
64°03’
64°08"
65°03’
65°06’
66°40’
73°40’
73°45’
74°14’
74°09"
Tioml Sy
73°05’
72°10’
63°50’
63°47’
64°10’
64°17’
64°19’
64°31’
64°52’
64°58"
65°05’
67°02’
66°48’
66°51’
68°24’
68°49’
70°40’
375-420
415-440
105-110
190-225
415-560
165-180
400-510
490
400-450
450-500
87-90
82-85
230-245
340-350
300-330
330-340
560-600
350-380
305-320
320-360
290-300
145-160
205-275
120-130
150
63-65
330-400
690-720
310-335
360-370
240-270
280-320
360-420
280-305
265-295
660-705
220-240
290-300
100-105
85-87
250-340
600-670
345-355
10 BREVIORA no. 40
REFERENCES
DALL, W. H.
1889. A preliminary catalogue of the shell-bearing marine mollusks
and brachiopods of the southeastern coast of the United States,
with illustrations of many of the species. Bull. U. S. Nat. Mus.
37: 1-221, 74 pls.
GouLD, A. A.
1870. Report on the Invertebrata of Massachusetts. Edited by W. G.
Binney. Boston. Pp. i-v + 1-524, 12 pls.
JOHNSON, C. W.
1934. List of marine Mollusea of the Atlantic Coast from Labrador
To Texas. Proc. Boston Soe. Nat. Hist., 40, no. 1: 1-203.
La Rocqur, AURELE
1953. Catalogue of the Recent Mollusca of Canada. Nat. Mus. Canada
129: i-ix + 1-406.
Nicou, DAvip
1951. Recent species of the veneroid pelecypod Arctica. Jour. Wash-
ington Acad. Sci., 41, no. 3: 102-106.
THORSON, GUNNAR
1951. The Godthaab expedition 1928. Meddel. om Grenland, Copen-
hagen, 81, no. 2: 1-117.
TRYON, G. W., JR.
1879. Manual of Conchology (1), 1. Cephalopoda. Acad. Nat. Sei.,
Philadelphia, pp. 1-316, 112 pls.
VERRILL, A. E.
1884. Second catalogue of Mollusca recently added to the fauna of
New England and adjacent parts of the Atlantic, consisting
mostly of deep sea species, with notes on others previously
recorded. Trans. Conn. Acad., 6, no. 1: 139-294, 5 pls.
1885. Third eatalogue of Mollusca recently added to the fauna of
New England. ... Trans. Conn. Acad., 6, no. 2: 395-452, 3 pls.
Jab
MOLLUSKS FROM NORTHEASTERN NORTH AMERICA
1954
‘SUOTZEIS PoyST] FO WOTPVIO] SuULMOYS vorv odojs [eyUaUTZUOD Jo dey,
069 | 202 |
4U0Q $abs0a5
uojsog
4404 MAN
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. DECEMBER 17, 1954 NuMBER 41
COMMENTS ON THE CLASSIFICATION
OF RODENTS
By ALBert E. Woop
Biology Department, Amherst College
There is no order of mammals where there are so many un-
certainties in the current classification as there are in the rodents.
The rodents have generally been subdivided into three groups,
the Sciuromorpha, Myomorpha and Hystricomorpha, although
there have been several attempts (notably Miller and Gidley,
1918, and Winge, 1924) to break away from the tripartite
division. The morphologic criteria on which these three groups
are based permits living rodents to be marshalled into them
with greater or lesser success. But the resulting problems remain
very prominent, and well-nigh insuperable when the paleontolo-
oie history of the order is taken into account. For example, if
the Hystricomorpha are a natural group, how ean their present
distribution (South America, Africa, southern Europe and
south Asia, with scattered recent Latin American immigrants
to North America) be explained in view of the complete absence
of Tertiary hystricomorphs in North America? Moreover, the
morphology of the various genera and families of fossil rodents
does not permit these forms to fit into a three-fold classification
anywhere near as well as do the living members of the order.
As long as we deal only with the living members of the order,
the criteria for subordinal classification can be sorted out
fairly well. However, studies of fossil rodents have shown that
sometimes the structures on which the classification has been
based were independently derived by diverse groups. For
example, Wood (1937) showed that the sciuromorph type of
zygoma and masseter was developed in the Oligocene genus
Titanotheriomys, a form that by no stretch of the imagination
2 BREVIORA No. 41
could be considered to be ancestral to anything included among
the recent sciuromorphs.
As a result of general dissatisfaction with the three-fold classi-
fication, there have been a number of attempts to arrive at
alternative arrangements. Those of Miller and Gidley (1918)
and Winge (1924) developed so many additional problems, while
solving a few, that they are not generally followed at present.
Wood (1937) basically followed the three-fold arrangement,
but proposed a fourth subdivision, the Protrogomorpha, to inelude
the primitive rodents that have not acquired the features of the
jaw muscles that characterize the members of the three classic
suborders. All members of this suborder are extinct except
for Aplodontia, the ‘‘Mountain Beaver’’ of parts of California,
Oregon, Washington and British Columbia.
Simpson (1945) made a valiant attempt to squeeze all rodents
into the three-fold classification, though pointing out in many
places that the scheme does not work well.
In an extensive review of the early Tertiary rodents of North
America, Wilson (1949) followed Simpson’s classification but
raised many queries and made a number of very significant sug-
gestions, particularly as to the probability of a close relationship
between the heteromyids and cricetids, heretofore generally con-
sidered to be sciuromorphs and myomorphs, respectively.
Simpson (1950) and Wood (1950) suggested that the South
American ‘‘hystricomorphs’’ should perhaps be separated as a
distinct suborder, though without working out the details. Wood
(1955), and Wood and Patterson (in press) are formally carry-
ing out such a separation, with detailed discussion as to the
reasons for the change.
Lavoeat (1951), in a review of certain Oligocene faunas of
France, agreed with this point of view, and showed that the
European Eocene and Oligocene Theridomyidae and Pseudosciu-
ridae could not be ancestral to either the Old World or New
World hystricomorphs of the present day.
For many years, Stehlin worked on a monograph of the evolu-
tion of rodent dentition, which was finished after his death
by Schaub (Stehlin and Schaub, 1951). This deals exclusively
with the dentition, and develops the thesis that rodents with
five-crested teeth are all descended from animals with the basic
tooth pattern represented by the European Eocene-Oligocene
1954 COMMENTS ON THE CLASSIFICATION OF RODENTS 3
theridomyids. Although this monograph contains a brief taxo-
nomic section, there is no attempt to present a comprehensive
revision of the classification of the order. There are, however,
a large number of suggestions and comments which, while not
building up any particular classification, must nevertheless be
given serious consideration by anyone interested in the super-
generic arrangement of the rodents. I cannot agree with all of
Schaub’s conclusions, and am therefore attempting to explain
some of these differences in detail here. The taxonomic results
of these differences of opinion are being published elsewhere as
a suggested classification of the rodents (Wood, 1955).
Schaub has recently (1953a) published a revised classification
of the ‘‘Hystricomorph’’ rodents. In this paper and others
(1951, 1953b; Stehlin and Schaub, 1951), he also makes com-
ments on the ‘‘Sciuromorpha’’ and ‘‘Myomorpha’’. Basically,
his idea is that these last two groups are wastebaskets, made up
of superfamilies that have had an entirely independent history,
and which cannot profitably be grouped in suborders. Although
he is not explicit, he gives the impression that he feels that, in
one or two eases, there might be some profit in grouping two of
the superfamilies into a suborder. With this part of his thesis
I feel a great deal of sympathy if not complete agreement,
although the classification proposed in Wood, 1955, does not
completely follow Schaub in doing away with these suborders.
The rodents representing the initial North American Eocene
radiation (the Ischyromyoidea), the middle Tertiary to Recent
Aplodontoidea, and the Sciuridae certainly seem fairly closely
related. At least they all retain primitive features of teeth, of
the zygomasseteric region, or of both, and they may well be kept
in a common suborder, for which the name Sciuromorpha may
be retained. The other superfamilies usually called sciuromorphs
seem to be quite distinct from these forms, and may belong
in rather diverse parts of the order.
The families that Wood places in the Myomorpha, however,
are a group about whose relationships there is little or no cer-
tainty. There are some suggestions of mutual relationship, and
some indications that Schaub is correct in thinking that they
have nothing in common with each other. If they are not related,
this would be the end of the Myomorpha, and the superfamilies
would have to be either left out of all suborders or, since this is
4 BREVIORA No. 41
illegal in formal taxonomy, each placed in its own suborder.
It is therefore obvious that Schaub is correct in his contention
that the classical concepts of the Sciuromorpha and Myomorpha
will have to be abandoned, and either new and perhaps more
restricted definitions developed for these suborders, or the sub-
ordinal concept will have to be largely abandoned for this
order. It is still too early to reach a definite choice between these
alternatives.
But on one of Schaub’s major points I find myself in disagree-
ment. One of the basic tenets of Stehlin and Schaub’s invalu-
able monograph is that the Theridomyidae are ancestral to all
forms that have a five-crested tooth pattern or that are derived
from a five-crested ancestor. Schaub has therefore proposed a
suborder, Pentalophodonta, to include all forms with five-crested
teeth, or that are derivable from a five-crested ancestry. How-
ever, it seems no more reasonable to assume that a given tooth
pattern has not arisen independently several times than to as-
sume that a given zygomasseteric pattern has not so arisen.
Indeed, all the current indications from the study of rodent
evolution are that extensive parallelism is the rule rather than
the exception in this order. Of Schaub’s Pentalophodonta, the
South American forms are surely a natural group, the Suborder
Caviomorpha of Wood (1955), and the Infraorder Nototrogo-
morpha of Schaub (1958a). But these forms must have been
derived from rodents that entered South America from North
America, as is now generally agreed (Schaub, 1953a; Simp-
son, 1950; Wood, 1950). There is also agreement that this entry
must have been no later than late Hocene. Schaub (1958a, p.
393) argues that this ancestral stock could have been theridomy-
ids that crossed North America to reach South America in the
Eocene, Paleocene, or even earlier.
It is impossible to prove that a form did not live in an area
or at a time where it is not represented by fossils. But no
rodents of any kind are known from South America before the
Lower Oligocene Deseadan, at which time they have become quite
abundant and are beginning to be diversified, although clearly
stemming from a common origin. Because of the rapid diversi-
fication of the caviomorph rodents beginning with the Deseadan,
it seems almost certain that, in this deposit, we are observing
an explosive radiation in the process of exploding. A backward
1954. COMMENTS ON THE CLASSIFICATION OF RODENTS 5
extrapolation of the post-Deseadan rate of evolution would indi-
eate that this explosion must have begun not very long prior
to the deposition of the Deseado beds. Although such an extra-
polation is dangerous, and although we obviously cannot be
certain of the reasons for such an evolutionary explosion, by far
the most reasonable interpretation, in view of what is known of
the general Tertiary history of South America, is that the
explosion began with the arrival of rodents in South America.
Here was a virgin field, with no highly developed competition,
and, once the rodents were established, they rapidly took over all
enawing niches. If this explanation is not correct, it is exceed-
ingly difficult to understand the rapid post-Deseadan develop-
ment of the caviomorphs. Hence it seems very questionable that
rodents could have been present in South America for any
appreciable time before the Deseado. If this postulate is ac-
cepted, and if their North American origin be granted, most
authors consider it more probable that the caviomorphs were
derived from known North American types (such as the Par-
amyidae or Sciuravidae) than from the European Theridomyi-
dae, still unreported from any of the richly fossiliferous North
American deposits.
An additional objection to Schaub’s suggestion of a Paleocene
or even earlier passage of the theridomyids to South America
is the complete absence of any rodents from the record prior to
the latest Paleocene, when only the single genus Paramys is
known. Presumably there must have been rodents prior to their
earliest appearance in the record, and they may have been in
existence for some time, although there is no valid criterion for
fixing the length of time necessary to differentiate the rodents
from their ancestors. This time may therefore be fixed as long
(extending back into the Cretaceous) or short (reaching only
the Middle Paleocene), depending on the individual student’s
opinion as to the probable rates of evolution involved and the
significance of the morphologic differences between Paramys
and whatever group is postulated as the ancestor. Moreover,
all the earlest rodents, both in North America and in Europe,
are referable to the Paramyidae, and to the most primitive
stages of that family. The only non-paramyid lowermost Eocene
genus known is the pseudosciurid Adelomys, which Wood, in his
6 BREVIORA No. 41
monograph of the Paramyidae, will indicate was derived from
the paramyids.
The absence of theridomyids from North America is, of course,
negative evidence. It is possible that this family may have lived
in eastern North America where there are no continental Hocene
deposits. But there is no evidence that the family was so repre-
sented, and it seems highly probable that the rodents that reached
South America went by way of what is now Mexico and Central
America, and hence were probably derived from the western part
of the continent.
Therefore, while admitting that all this is negative evidence,
it seems most probable that a representative of one of the
known North American Eocene groups reached South America
late in the Eocene, to give rise to the South American radiation.
A further point of disagreement with Schaub is in the phylo-
genetic position of the Paramyidae. Wood (1946) referred to
the specialization of the Paramyidae, which indicated that they
could not have the fundamental ancestral position generally
given them. With this point of view, Schaub is in complete
agreement (1953b, p. 35), considering that the Sciuridae have
the most primitive tooth pattern in the order. Wood’s 1946
opinion was based on preliminary investigation of the paramyids,
which includes many rather specialized genera. Further study
of Lower Eocene forms, however, has caused a revision of this
opinion, and I now feel that the Upper Paleocene and Lower
Eocene paramyids are the most primitive known rodents, al-
though it is certain that the earliest sciurids have made little
or no advance over them in the structure of the cheekteeth. The
data for this conclusion have not been published, but are in-
cluded in a monograph on the Paramyidae, now nearing comple-
tion. Therefore, the North American Paramyidae could have
been ancestral to the South American radiation, although the
intermediate steps have not been traced in detail. If the para-
myids were the ancestors, Schaub’s Suborder Pentalophodonta
immediately becomes a composite group, the two infraorders
having nothing in common.
Schaub places the Theridomyoidea, Castoroidea and Hystri-
coidea in the Infraorder Palaeotrogomorpha. But Lavocat
(1951) has presented evidence, derived from structures other
1954 COMMENTS ON THE CLASSIFICATION OF RODENTS (
than the cheekteeth, that the theridomyoids cannot be related
to the hystricoids, so that this infraorder would also seem to be
composite. Schaub places a number of other groups in the
Palaeotrogomorpha, particularly the Spalacidae and Rhizo-
myidae. This seems perhaps somewhat less likely than that they
belong in the Myomorpha, if there is any such group. Certainly
there is not enough known of the history of either family to
justify placing them with confidence in any particular category.
Schaub’s new proposals for classification of rodents are very
useful and thought-provoking. The fact that there are disagree-
ments with them is not surprising. The current status of rodent
phylogeny and classification is such that anyone can point out
inconsistencies in anybody else’s classification. The rodents are
a sufficiently large and complex group so that no one person
can be familiar with the entire order. Progress will be made only
if various students attack the problem, even though it will result
in presenting solutions that others can show obviously to be
incorrect. Wood’s current (1955) classification has been pro-
posed with this object in view, in the hope that there will be as
many criticisms, both constructive and destructive, as possible,
since this is the only way to progress.
Schaub has made one further suggestion that very probably
will be incorporated in the ultimate classification of the order.
This is the idea that many of the superfamilies are entirely
separate from all others, presumably having an independent
history since the Eocene. There very clearly was a sudden
diversification of the rodents, at or near the end of the Eocene,
which apparently resulted from a rather sudden change from
enawing based on the use of the temporal muscle to that based
largely on the use of the masseter, with consequent enlargement
and modification of the latter. The present evidence suggests,
but does not prove, that this occurred a great many times inde-
pendently. It is very probable that these lines all developed from
different groups of Eocene rodents, and that, therefore, each of
these lines will deserve separate taxonomic treatment. Generally
speaking, such superfamilial lines could be grouped into sub-
orders only artificially. Although the present evidence suggests
the probability of such a bushy phylogeny (rather than a tree
with half a dozen main branches), it is perhaps more conserva-
8 BREVIORA No. 41
tive to struggle with an attempt to group rodents into suborders
for a while longer, while keeping in mind the possibility (or
probability) that a classification with entirely independent
superfamilies will ultimately prove to be the only one that will
fit in with the facts of rodent paleontology, as these gradually
come to light. Certainly it will be some time before an adequate
classification of the order is compiled.
REFERENCES
Lavocat, R.
1951. Révision de la faune des mammiféres oligocénes d’Auvergne et
du Velay. Editions ‘‘Sciences et Avenir,’’ Paris, 153 pp.
Miuurr, G. S., and J. W. GIDLEY
1918. Synopsis of the supergeneric groups of rodents. Jour. Wash.
Acad. Sei., 8: 431-448.
ScHAUB, S.
1951. Review of: H. G. Stehlin et S. Schaub, ‘‘Die Trigonodontie der
simplicidentaten Nager.’’? Mammalia, 15: 204-210.
1958a. Remarks on the distribution and classification of the ‘‘ Hystri-
comorpha.’’ Verh. Naturf. Ges. Basel, 64: 389-400.
1953b. La Trigonodontie des Rongeurs simplicidentés. Ann. Paléont.,
39: 29-57.
SIMPSON, G. G.
1945. The principles of classification and a classification of mammals.
Bull. Amer. Mus. Nat. Hist., 85: XVI + 350 pp.
1950. History of the fauna of Latin America. Amer. Scientist, 38:
361-389.
STEHLIN, H. G., and 8S. ScHAUB
1951. Die Trigonodontie der simplicidentaten Nager. Schweiz. Pal.
Abh., 67: 1-385.
WIitson, R. W.
1949. Early Tertiary rodents of North America. Carnegie Inst. Wash.
Publ., 584: 67-164.
WINGE, H.
1924, Pattedyr-Slaegter, vol. 2. Rodentia, Carnivora, Primates. Copen-
hagen, H. Hagerups Forlag, 321 pp.
1954 COMMENTS ON THE CLASSIFICATION OF RODENTS 9
Woop, A. E.
1937. The mammalian fauna of the White River Oligocene by W. B.
Scott and G. L. Jepsen. Part II. Rodentia. Trans. Amer. Phil.
Soc., 28: 157-269.
1946. Early Tertiary rodents of the Family Paramyidae. Bull. Geol.
Soe. Amer., 57: 1245 (Abstract).
1950. Porceupines, palaeogeography, and parallelism. Evolution, 4:
87-98.
(In Press.) <A revised classification of the rodents. Jour. Mammalogy,
36.
Woop, A. E., and B. PATTERSON
(In Press.) The rodents of the Deseadan Oligocene of Patagonia and
the beginnings of the South American rodent radiation. Chicago
Nat. Hist. Mus.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. FEBRUARY 28, 1955 NUMBER 42
A NEW SALAMANDER OF THE GENUS
PARVIMOLGE FROM MEXICO
By Grorce B. Rapp
Museum of Zoology, University of Michigan
In the Museum of Comparative Zoology there is a specimen of
a Mexican plethodontid salamander superficially similar to in-
dividuals of Pseudoeurycea cephalica. Examination indicates
that it actually represents an undescribed form of the genus
Parvimolge Taylor, which may be known as
PARVIMOLGE PRAECELLENS, new species
(Figure 1)
Type. An apparently immature female, Museum of Compara-
tive Zoology No. 24701, collected at Hacienda el Potrero, near
the city of Cordoba, Vera Cruz, Mexico, on December 24, 1940,
by Archie F. Carr.
Diagnosis. A small plethodontid salamander related to Parvi-
molge townsendi by virtue of syndactylous feet, with the par-
tially free digits having pointed tips, and by the presence of
conspicuous though small glands about the middorsal line. It
differs from P. townsendi in larger body size, greater number of
teeth, and in having small nostrils. It differs from the only other
species assigned to this genus, richardi, not only in these charac-
ters but also in having conspicuous dorsal glands.
Description. General aspect of body robust. Length of tail
approximately six-sevenths of distance from snout to posterior
end of vent. Head length (snout to center of gular fold) con-
tained about four times in snout-vent distance; maximum head
No. 42
BREVIORA
‘UU GQ yHOgB YYSus, [e}oO} [eNpYV
‘suayja0anid abjowimwwg FO odsy, *T ono
1955 A NEW SALAMANDER OF THE GENUS PARVIMOLGE 3
width contained in same about five times. Head width three-
fourths the snout-gular fold distance and nearly equalling snout-
posterior angle of jaw length. Width of an eyelid less than
minimum distance between eyelids. Horizontal diameter of eye
greater than one and a half times snout-anterior angle of eye
distance. Internarial distance four-sevenths the interorbital
measurement. Viewed dorsally, the snout is gently rounded,
though short in relation to anterior angles of eyes. Canthus
rostralis not sharply defined, the area between anterior mid-
dorsal line of head and edge of upper jaw being a gradual slope.
Nostrils small with nasolabial grooves forking on weakly de-
veloped nasolabial protuberances. Snout not projecting beyond
tip of lower jaw. Lower jaw semi-oval in outline, ventrally
viewed. Lateral extensions of gular fold meet grooves extending
from posterior angle of eye and continue dorsally to join the
posterior ends of a V-shaped groove originating parietally. Epi-
branchial ridge extending beyond axillary groove. A deep
furrow, perhaps partially due to preservation, follows the mid-
dorsal line from scapular region onto tail. The fairly thick skin
of dorsum has about fifty small but conspicuous glands (pre-
sumably of the ‘‘poison’’ type). These glands appear to be
arranged in four irregular longitudinal rows, two of them very
close to midline. On the neck the middorsal rows fork and
curve laterally. There are a few conspicuous glands on the tail.
Maxillary-premaxillary teeth, counting both sides, total 75.
They are practically all of uniform size, those in the premaxil-
lary position not piercing the lp. Eighty-seven mandibular
teeth. Prevomerine teeth 20 on right and 23 on left side mostly
in double rows which arch very slightly posteriad in approaching
the midline from initial points lateral to the small choanae.
Distance between prevomerine series at midline is equal to three
times the diameter of a choana. Parasphenoid teeth in two ob-
long patches, each of which has about 70 teeth arranged in diag-
onal rows. Anteriorly the groups are separated by slightly
more than one choanal diameter, posteriorly by about three
diameters. Tongue large and free, the fleshy sublingual fold
anterior to it well developed.
The moderately chunky body has twelve costal spaces. It is
difficult to ascertain whether there are twelve or thirteen costal
ae BREVIORA No. 42
grooves because of the contorted condition of the specimen. There
are about twenty-two caudal grooves indicated posterior to the
vent. Tail quadrangular in cross-section, slightly broader dorsally
than ventrally. Throughout most of its length, it is deeper than
broad.
Limbs slender and moderately long, hind limb shghtly larger
and longer than forelimb. Limbs fail to meet by about one
costal space when pressed to sides of the body. Hands and feet
well developed but not expansive, with slight pads ventrally.
The somewhat slender, terminally free digits have pads which
are discrete from the palmar pads. Innermost and outermost
digits of hand and foot completely enclosed in web. Terminal
phalanges of second and fourth toes and second finger free of
web, while third toe and finger have almost two phalanges free.
Third and fourth toes and second and third fingers noticeably
pointed distally, while the second toe appears more rounded.
Toes in order of decreasing length: 3, 4, 2, 5, 1; fingers in same
order: 3, 2, 4, 1.
In alcohol, the ground color of dorsal surfaces of head, body,
tail, and limbs is brownish-black. Ventral surfaces show a faded
version of this color with some areas being almost completely
unpigmented (gular fold, anterior part of chin, and palms and
soles). Underside of tail hght brown. Jaw margins appear
mottled due to lack of pigment in spots. Bluish-white spots and
blotches on lower sides of body and on tail were possibly white
in life. A distinct but broken bar of this color on the head
between the eyes.
Measurements (in millimeters). Snout-posterior end of vent,
c. 35; tail, c. 30; snout-gular fold, 9.4; snout-angle of jaw, 7.5;
snout-anterior angle of eye, 1.6; horizontal diameter of eye, 2.6;
head width, 7.0; internarial distance, 2.0; interorbital distance,
3.5; arm, 8.9; leg, 9.4.
Remarks. Before commenting on relationships I wish to place
on record some information on P. townsendi not obtainable from
previous accounts (Dunn, 1922 and 1926; Taylor, 1944). The
major source of the data is a series of ten specimens collected
at the same time and place as the type of P. praecellens. Varia-
tional data for the eight alcoholic adults (MCZ 24712-19) are
given in Table I. The males have two or three enlarged premax-
1955 A NEW SALAMANDER OF THE GENUS PARVIMOLGE 5
illary teeth, prominent mental glands, and well developed naso-
labial protuberances, with more truncate snouts and with larger
external narial openings than the females (average 0.8 mm.
versus 0.2 mm.). The parasphenoid teeth groups are separated
in half of the eight specimens. Counts of prevomerine tooth
rows averaged 5.8 on a side. One male, not included in the aver-
age, has these teeth in an irregular patch of ten on each side.
UMMZ 111305, an immature specimen (snout-vent, 15 mm.; tail,
13 mm.) from this series, has fewer teeth (prevomerine, 3-5;
maxillary-premaxillary, 34; mandibular, 47) and fewer glands
(31) than any of the adults of Table I. A female, UMMZ 63944,
taken at Cordoba, Vera Cruz, on June 19, 1927, is larger than
any recorded specimen, measuring 24 mm, from snout to vent,
with a tail of the same length. The ovarian eggs are large, about
0.7 mm. in maximum diameter.
Many minor variations in color were found. The ground
color was diverse shades of brown, usually lighter ventrally, and
sometimes dorsally, than on the sides. The dark bars described
by Dunn are most pronounced on the tail, with the most econ-
spicuous bar or ring occurring at the weakly marked basal con-
striction of the tail. A herringbone or V pattern can be made
out dimly in some specimens held under water; it is apparently
the result of concentration of dark pigment in the costal grooves.
Most of the specimens have an incomplete light bar between the
eyes well indicated, but two do not. Three have some evidence
of an additional incomplete anterior interocular bar. In two the
rostral region appears as a brilliant white square, due to absence
of dark pigment, which is shghtly reduced in this area in the other
specimens. The limbs, throat, and venter have scattered light,
probably pigmentless, spots. Light streaks on the sides are well
developed in some, practically absent in others. In one specimen
the middorsal area is light brown, contrasted with dark brown
dorsolaterally, which in turn is set off by a double row of light
streaks or spots on the lower sides. In the same specimen there
are striking dorsolateral white patches (apparently pigment-
less areas) in the nuchal region.
The tenth specimen from the Potrero series, an adult male
measuring about 23 mm. from snout to vent, was stained and
cleared (UMMZ $-1556). The testes of this specimen were
6 BREVIORA No. 42
relatively enormous and were in the bilobate condition which I
have found to characterize the larger, and presumably older,
classes of males in Chiropterotriton and Pseudoeurycea. The
more interesting skeletal features are described here. Skull
broadly elliptical, maximum width at the otic capsules three-
fourths of the premaxillary-occipital condyle distance; frontals
and parietals ossified completely to midline; facial lobe of maxilla
large, forming with nasal bone a posterior border of nasal cap-
sule; nasal fairly prominent, angling forward medially, about
two and a half times as long as broad; prefrontal two-thirds the
length and one-half the width of nasal, not participating in the
margin of the nasal capsule; basally fused frontal processes of
premaxilla diverge considerably forming a wide fronto-premaxil-
lary fontanelle (a tenth of otic capsular width) ; these processes
laterally compressed anteriorly; more posteriorly and dorsally
they gradually twist until they appear horizontally flattened
where the ends articulate with grooved projections of the frontal
bones; two large premaxillary teeth, twenty-three teeth on each
maxilla, twenty-eight teeth on each of the well ossified mandibles,
eight prevomerine teeth on a side; no median suture between
prevomers; parasphenoid teeth in two patches. Skull measure-
ments (mm.) made with ocular micrometer: otic width, 2.9;
premaxillary-condylar length, 4.4; prefrontal length, 0.5, width,
0.15; nasal length, 0.75, width, 0.8; fronto-premaxillary fonta-
nelle width, 0.8. Hyoid structures as described by Tanner
(1952), but second ceratobranchials definitely osseous or calcified,
which is also the condition in the only cleared individual avail-
able of Lineatriton lineola (UMMZ S-1594). Phalangeal for-
mula of foot, 1, 2, 3, 3, 2; of hand, 1, 2, 3; 2; eight carpals, ulnare
and centrale partially fused on both hands, on left hand all
ossified to some extent but on right hand radiale and carpal im-
mediately distal to it are cartilaginous. At least seven, probably
nine tarsals, four ossified on each side. Atlas weak, not ossified
middorsally ; only the second vertebra with a noticeable neural
ridge. Long, proximally forked ribs present from second to
fourteenth vertebra, fifteenth having both transverse processes
but no rib. Sixteenth vertebra is the sacral, followed by two
transitional postsacral or precaudal vertebrae. Twenty-three
caudal vertebrae, the anterior ones with a fin-hke process ex-
1955 A NEW SALAMANDER OF THE GENUS PARVIMOLGE a
tending forward from the ventroposteriorly directed haemal
process to the anterior edge of the vertebra. Short transverse
processes, more laterally angled in the first few caudal vertebrae,
more anteriorly directed farther caudad.
This account of the skeletal features is at variance with Tay-
lor’s deseription (1944: 223) on several points, notably the pres-
ence of the prefrontal and the ossification of the carpals and
tarsals. The latter character may be variable, but the condition
in this specimen does not support contrasting the cartilaginous
carpals and tarsals of Parvimolge with the osseous ones of
Thorius. This is especially true in view of the fact that Hilton
(1946, 1948) has recorded them as cartilaginous in 7’. pennatu-
lus, and since the single cleared specimen referable to T.
dubitus that I have examined likewise has entirely cartilaginous
ones. However, I belheve that most of the discrepancies between
the two descriptions of P. townsendi may be ascribed to different
types of material used, and it should also be borne in mind that
the foregoing account is based on a single specimen.
TABLE I
Quantitative Characters in Mexican Parvimolge
snout- dorsal premax: mandibu-
vent tail costal body maxillary lar
length length spaces glands teeth teeth
townsendi R 20-23 20-26 32-42 45-58 54-69
males (4) M 22.0 22.5 3.7 BOO) 50.0 61.5
townsendt R 19-22 16-21 30-41 48-58 00-73
females (4) M 21.0 OEG) 4.2 38.0 53.0 64.0
praecellens
(female) 35 30 i 50 75 87
R, range; M, mean; tooth counts are totals of both sides; measurements in
mm.; costal spaces refer to number between adpressed limbs.
8 BREVIORA No. 42
Relationships. Parvimolge praecellens is closely related to P.
townsendi, as indicated by the foot structure, coloration, con-
spicuous dorsal glands, and shape of tail. A third form, richardi,
described with reservations as a Parvimolge (Taylor, 1949,
1952), lacks the conspicuous glands. This fact, apparent differ-
ences in details of the foot, and the enormous disjunction in
range — Vera Cruz to Costa Rica—imply that richardi is
preferably not to be associated with townsend and praecellens.
Determining the nature of the affinity between P. praecellens
and townsendi necessitates much more material. However, the
more numerous teeth, longer legs, and small nostrils of praecel-
lens are probably correlated with its greater size. These charac-
ters and the smaller size of the dorsal glands appear to indicate
that praecellens is less specialized. The morphological and eco-
logical relationship may be of the same general type existing
between sympatric large and small species in Psewdoeurycea and
Chiropterotriton. Indeed, in the latter genus, C. dimidiata shows
seemingly paedomorphie features very like those found in town-
send,
That Parvimolge is allied to Lineatriton as inferred by Tanner
(op. cit.) is supported by the distinctive osseous character of the
second ceratobranchials found to be common to P. townsendi and
L. lineola. Nevertheless, the derivation of townsendi and prae-
cellens was probably from a less specialized form, perhaps in the
Pseudoeurycea cephalica group or its ancestral stock. The foot
shape in very young specimens of cephalica, the presence of ir-
regularly. protruding poison glands on the dorsum in some forms
of the group, and the guanophore-spotted, brown to black ground
color in the group suggest this. The syndactylous foot structure
could have become established by a paedogenetic process in
Parvimolge stock as it differentiated from some such ancestor.
Acknowledgments. I wish to thank Arthur Loveridge for the
privilege of examining most of the specimens described in this
paper. Charles F. Walker and James E. Mosimann have read
the manuscript and given useful suggestions. The drawing was
executed by Harold J. Walter. The abbreviations MCZ and
UMMZ respectively designate the Museum of Comparative Zool-
ogy and the University of Michigan Museum of Zoology.
1955 A NEW SALAMANDER OF THE GENUS PARVIMOLGE 9
LITERATURE CITED
Dunn, E. R.
1922. A new salamander from Mexico. Proc. Biol. Soc. Washington, 35:
5-6.
1926. The salamanders of the family Plethodontidae. Smith College,
Northampton, Mass. viii + 441 pp.
Hiuton, W. A.
1946. Skeletons of Mexican and Central American salamanders of the
family Plethodontidae. Jour. Entomol. and Zool., 38, No. 1: 1-8.
1948. The carpus and tarsus of salamanders. Jour. Entomol. and Zool.,
40, No. 1: 1-13.
TANNER, W. W.
1952. A comparative study of the throat musculature of the Plethodon-
tidae of Mexico and Central America. Univ. Kansas Sci. Bull.,
347 Pt. Ll, No. 10: 583-677.
TAYLOR, E. H.
1944. The genera of plethodont salamanders in Mexico, Pt. I. Univ.
Kansas Sci. Bull., 30, Pt. I, No. 12: 189-232.
1949. New salamanders from Costa Rica. Univ. Kansas Sci. Bull., 33,
Pt. I, No. 6: 279-88.
1952. The salamanders and caecilians of Costa Rica. Univ. Kansas Sci.
Bull., 34, Pt. II, No. 12: 695-791.
i b%
ea 1
CE ah Pe
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, MASs.
April 7, 1955
NUMBER 43
SE RED-INDUCED SKIN FOLDS IN THE
BOTTLE-NOSED PORPOISE,
LOR STORRS LRUNCALY Ss“
By FRANK 8. ESSAPIAN
Marineland Research Laboratory
While engaged in a study of the behavior of bottle-nosed por-
poises, Tursiops truncatus (Montagu), at Marine Studios,
Marineland, Florida, over a period of three to four years, the
author has observed on hundreds of oeeasions a remarkable
modification of the skin contour of these animals whenever they
resorted to rapid movement about the tank (Essapian 1953,
p. 399). The cireular tank, where the porpoises are on exhibit,
is seventy-five feet in diameter and about fourteen feet deep
at the center. It houses an average of ten animals.
Generally lost to view due to swiftness of action, this trans-
formation of the skin contour takes place when an animal aceel-
erates its speed in excess of its normal rate, or, when swimming
rapidly, comes to a sudden stop. It is then that transverse skin
folds may stand out in relief on part of an animal, or may
extend over its entire length, from head to tail. The pattern of
these formations depends on the speed and sex of the swimming
animal, but is not restricted to any age group. The folds may
be seen even in an animal only a day or two old.
The skin folds generated by a sudden burst of speed are by far
the more common of the two categories, and are usually oe-
casioned when the animals engage in pursuit of one another, are
frightened and trying to escape, or when racing for a food fish
tossed into the tank. The duration of folds is then conditioned
1 Contribution No. 751 from the Woods Hole Oceanographic Institution.
2; BREVIORA No. 43
by the factors which motivate the animals’ movements. In the
case of a brief dash, for example, when an animal exerts itself
to snatch a fish, the skin folds may last only a second, or even a
fraction of a second (Pls. 1 and 2). However, in the ease of
prolonged rapid movement around the tank, the animals may
intermittently exhibit skin folds for a period of one or two or
more seconds. Often an animal at play or during the mating
season may suddenly leap out of water, exhibiting folds at the
peak of acceleration, just prior to its emergence from water,
and again after it has reentered the water (Pls. 3 and 4). At
this point it may be of interest to note that the fat adult females
tend to exhibit fewer folds, but greater in size and at less regular
intervals, than are produced in the adult male or in the younger
slimmer animals when swimming at comparable speeds. The
same female in Plate 4 is also shown in Plate 5; here this animal’s
folds are shown in a transitory stage, just prior to full accelera-
tion.
The skin folds of the second category occur at less frequent
intervals and are of very brief duration. These take place
when an animal swimming rapidly without exhibiting any folds
comes to an abrupt halt. Then the folds may suddenly be dis-
played (Pl. 6). The duration of folds of this sort is indicated
by the fact that this entire sequence occupies only twelve
frames of 16 mm. film at 16 frames per second. This would
indicate a total elapsed time of three quarters of a second for
the formation and disappearance of folds. Plate 6 is the fourth
frame, showing that these folds were fully formed in one quarter
of a second.
One feature in the production of skin folds is of particular
interest. In an animal maintaining a high and sustained rate
of speed the folds are stationary and do not progress wave-
fashion. At times, when extremely high speeds are achieved by
adult animals, the skin folds tend to slope posteriorly. Observa-
tions also indicate that the folds are more likely to occur on
ventral and lower lateral surfaces.
While the superficial layer of the porpoise’s skin is extremely
thin, the blubber is of considerable thickness and is tough and
fibrous. In a Tursiops truncatus fifteen days old and weighing
thirty pounds, the skin, including blubber, is approximately half
JG}
ee
1955 BOTTLE-NOSED PORPOISE TURSIOPS TRUNCATUN
an inch thick. In an adult it is nearly one and a half inches
thick on the ventral surface. Although normally not apparent
to the view or touch, the porpoise’s skin is, nevertheless, pliant
and loose on the body. This slackness of the skin in a live animal
held out of water in a sline is shown in Plate 7.
Discussion
It is tempting to speculate upon the causes of this phenomenon.
The answer may lie in the possibility that these folds are caused
by the unequal pressure of water upon the body of the animal,
in which case the production of folds would appear to be beyond
the muscular control of the animal. Clarke and Ruud (1954, p.
144), in discussing different types of marks used in whale mark-
ing, Say: ‘*... whenever a whale is moving naturally the envelope
of blubber is always sliding, more or less, over the contracting
and relaxing muscle beneath, so that a mark with stoppers
which is set partly in blubber and partly in muscle would almost
certainly be worked out.’’? The Marineland porpoises while
swhnmine normally do not display such looseness of skin as
is here ascribed to the great baleen whales. However, this
similarity in the flexibility of skin in such divergent species sue-
gests that all animals of the order Cetacea may share this
characteristic.
These speed-induced folds are to be distinguished from those
caused simply by flexure, such as the ones shown in Plates 8
and 9.
Racovitza (1903, pp. 438-44, pl. 3, fig. 17) mentions a group of
four Hyperoodon (bottle-nosed whales) of which one individual
is sketched emerging from water and exhibiting an exaggerated
likeness of skin folds on its trunk. Unfortunately the author has
failed to explain whether this particular animal was swimming
rapidly at the time or whether this deformity in the skin contour
Was apparent even when the animal had raised itself partly out
of water. The author furnished no further explanation beyond
expressing his astonishment and a conviction that the animal
didn’t appear to be sick or emaciated and that the projections
on its trunk appeared to be composed of soft material. This
animal didn’t differ from the three other individuals in any
other respect.
4 BREVIORA No. 43
At sea, in conditions of greater freedom of movement, the
porpoises may exhibit skin folds for more protracted periods
than is feasible in the confines of a tank. In any case, these
folds should be of interest to hydrodynamicists.
The author wishes to express his deep gratitude to Mr. William
E. Schevill for his encouragement, as well as for his helpful
suggestions and criticisms of the manuscript. The author is also
thankful for the use of a motion picture camera and film which
was made available by Mr. Schevill under a contract between
the Office of Naval Research and the Woods Hole Oceanographic
Institution.
REFERENCES
CLARKE, RopertT and JOHAN T. RuuD
1954. International cooperation in whale marking: The voyage of the
‘“‘Bnern’’ to the Antarctic 1953. Norsk Hvalfangst-Tidende,
43(3): 128-146.
ESSAPIAN, FRANK 8.
1953. The birth and growth of a porpoise. Natural History (New
York), 62(9): 392-399, ills.
RacovitTza, E. G.
19038. Cétacés. Résultats du voyage du S. Y. Belgica. 1897-1899.
Rapports Scientifiques, Zoologie. 142 pp., 15 text figs., 4 pls.
All photographs by courtesy of Marine Studios; the porpoises shown are
all Tursiops truncatus on exhibition at Marineland, Florida. Plate 7 is from
a photograph by Bob Neelands; all the others are by the author.
‘puosas B® JO QQG/ b/A 4B poydeasoyoyg
“YANOUL Joy WE Wes St poztes ysnl
Sey oYS YS oY], “WOr4Badja008 Jo yvod oy} ye splog sAvldstp ‘pro syyzuour woo4yysta Ayoyeurxorddy Sue WP Te]
"“puoves Bae) Ga/
F° OSG/TL F woe peyduaxsoz,oyq US & OZlos OF FNOGeE ‘UOLZEALOLavUpP FO 04V —
= = z < 3 L4pnpe uy
Pl. 3. A female, approximately eighteen months old, about to leap out
of the water. Note also the folds on the head of the pursuing five-year
old male.
Photographed at F/3.5 at 1/500 of a second.
Pl. 4. An adult female displays folds as she reenters the water at com-
pletion of a leap.
Photographed at F/3.5 at 1/250 of a seeond.
*puodoas GEe/aA iB paydeasoqoyg
“SOABM .1lOJBM odB TMS
94} “T 93R[q UL SY “WOr}BLe[ad0R [Ng
ATOPIM SJIQIYXO F 9}V[Q UL sR [BUIIUB OURS OY, *G “Tq
oy} Youe I xt OF ONP SI Wd} ) puew JYST, Aepnso1I1
0} 10ottd ‘Mel ayy Jo opsue oY} JB SULUULSaq Sp[OJ ULyS po
6. An adult male displays folds as he comes to an abrupt stop in a
vertical position (second porpoise in background shows no folds). En-
larged from 16 mm. film.
Photographed at 16 frames per second.
‘SpuvpoaN qog Aq ydeisojoyg
“ULYS VY} FO SSoUISOO] ollla.1}xo oY} SAvTdstp SUT[S B UL pozJIT opeuLaT [Nps oaAllT VW °*L “Id
Pl.
8. Throat folds caused by downward flexion of head (esp. lower jaw).
That these differ from the occasional speed folds is shown by the per-
manent creases between them (seen also on the unflexed throat of the
porpoise in Plate 7). Note also the characteristic pigmentation.
Photographed at F/3.5 at 1/250 of a second.
Pl. 9. Folds in the neck-shoulder region caused by turning the head side-
ways. This is the same animal as in Plates 4, 5, and
Photographed at F/3.5 at 1/250 of a second.
ee) Denman _—- : oo es 27 - a a. ——- ve — we
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, MAss. APRIL 8, 1955 NUMBER 44
A NEW MUREX FROM MATANZAS, CUBA
By Wiruu1AmM J. CLENCH
Through the kindness of C. J. Finlay of Varadero Beach, and
E. H. Monroe of Cardenas, Cuba, we have received a series of a
new Murex which was obtained from shell traps set in 100 fathoms
in Matanzas Bay, Cuba.
Murex (MUREX) FINLAYI, new species
Figures 1-3
Description. Shell reaching 938 mm. (384 inches) in length,
rather solid in structure and moderately spinose. Whorls ten
and moderately convex. Nuclear whorls glass-like and smooth.
Color brownish-yellow to brownish-cream with a few spiral threads
of a darker brown which follow the crests of the cords. Spire
moderately extended. Suture irregular and deeply indented.
Aperture subecircular and shghtly oblique. Parietal lip glazed,
adherent to the body whorl and fairly thick. Palatal lp erenu-
lated and slightly thickened. Siphonal canal greatly extended,
sometimes nearly half the length of the shell, usually curved
upward and to the left when viewed dorsally. Two or three
previously formed stages of the siphonal canal remain as seale-
like spines. The sculpture consists of three equidistant varices,
each supporting a rather large shoulder spine and maybe two or
three smaller spines between the shoulder and the base of the
shell. The varices on each whorl are more or less aligned with the
varices on the whorl above. Between the varices there are two
to four axial ridges which are strongly nodulose. Spiral seulp-
ture consisting of numerous thread-like cords. The varices are
formed by two arched plates, the plates on the aperture side being
somewhat smaller, the outer (first formed) leaving a flange or
2 BREVIORA No. 44
web-like ridge as a crest on the varix. This is usually broken
away on the early varices or remains as a series of small blade-
like spines.
Measurements
length width whorls
87.0 mm. 30.0 mm. 10 Holotype
93.0 31.0 10 Paratype
85.0 33.0 10 Paratype
82.5 34.0 10 Paratype
Types. Holotype, Museum of Comparative Zoology no. 189939
from shell trap, Matanzas Bay, Cuba, in 100 fathoms. Paratypes
from the same locality in the Museum of Comparative Zoology
and the collections of C. J. Finlay and E. H. Monroe.
Remarks. This species appears to be rather closely related to
both Murex antillarum Hinds and Murex beawi Fischer and
Bernardi. From M. antillarwm it differs by having fewer and
much larger axial costae, having fewer spines, and in possessing
the webbing along the varices and the greatly extended siphonal
eanal. Krom WM. beau it differs by having fewer and much larger
axial costae, shorter spines and not having the extensive webbing
which characterizes M. beaui. This present new species may also
be related to MW. aguayoi Cleneh and Farfante but it differs from
aguayo. by being larger, having the diffused brownish-yellow
coloration, fewer spines, larger axial costae and by having the
webbing on the varices.
The development of the webbing and the greatly extended
siphonal canal probably allows these mollusks to exist on a rather
soft muddy bottom.
The subgenus Murex in which this new species is included, is
avery complex group and of very wide distribution in nearly all
tropical and south temperate seas.
Murex finlayi Clench. Figs. 1 and 3, paratypes. Fig. 2, holotype. Matanzas
Bay, Cuba in 100 fathoms. (All slightly enlarged.)
BREVIORA
Museum of Comparative Zoology
Pp S\
June 13, 1955 NUMBER 45
CAMBRIDGE, Mass.
PALAHOTARICHA OLIGOCENICA, NEW
GENUS AND SPECIES, AN OLIGOCENE
SALAMANDER FROM OREGON.
By RicuHarp vaAN FRANK
In 1951 Mr. George R. K. Moorhead, a member of the Salem
(Oregon) Geological Society, collected a fossil urodele in a small
slab of shale from a locality situated about three miles southeast
of Eugene, and about 1, mile north of Goshen, Oregon. The
specimen was sent by Dr. Herman Clark of Willamette Univer-
sity to the Museum of Comparative Zoology for study.
The strata in which the fossil was found are designated by
Vokes et al. (1951) as post- Fisher and Eugene (Oligocene )
plant-bearing tuffs. These strata are described by the authors as
‘‘a sequence of apparently water-laid coarse tuffs with inter-
bedded thin layers of carbonaceous gray shale... [containing |
an abundant flora... [of which fifteen species] were determined
by Dr. Roland Brown,! who assigned a late Oligocene age to the
containing strata.’’ Dr. Ralph W. Chaney of the University of
California has written me that he too considers the strata to be
younger than EKugene or Fisher, and is terming the strata the
Willamette formation. On the other hand, Dr. Clark writes that
he and Dr. Ewart M. Baldwin of the University of Oregon prefer
to assign them to the Eugene formation, although conclusive
evidence is lacking. At any rate, all concerned seem confident of
the Upper Oligocene age of the deposit. It is the gray shale just
mentioned which contains the salamander.
The specimen is about 120 mm. in total length, and lacks a
centimeter or so of the tail, the right mandible, most of the ribs,
the left scapula and humerus, and parts of the hyoid apparatus.
It is blaek in color, apparently highly carbonized. The skull is
land listed by Vokes et al. (1951)
2 BREVIORA NO. 45
erushed dorsoventrally, and the vertebral column is twisted so
that it lies on its side with respect to the skull.
The dorsal surface of the skull when first prepared seemed
to be an uneven but continuous structure. Since depressions
representing the orbits and the cavum internasale were visible,
and the surface was much darker than the matrix, and had the
outline of a skull, this dorsal material was thought to be bone in
a bad state of preservation. Futile attempts to understand the
skull in the light of this interpretation (the sides of the brain-
case, moreover, appeared to be pressed out and in contact with
the maxillae), suggested that research in experimental paleontol-
ogy might be useful. In this pursuit a skull of a Recent Taricha
granulosa was decalcified with dilute hydrochloric acid, and
then flattened between two glass plates and allowed to dry. The
sides of the braincase were still no closer to the maxillae than
before. This increased my growing suspicion that the material
exposed on the dorsal surface of the fossil skull was not bone, and
prompted exploratory probing through the surface. The opera-
tion revealed undeniable bone underneath. The removal of most
of the matrix from the unexposed side then followed. I believe
that the formerly exposed surface was mineralized skin, which
had roughly followed the contours of the skull — not revealing,
however, any evidence of the taxonomically important fronto-
squamosal arches. The same dark material is present elsewhere,
as revealed by further preparation, and is separated from the
bone by about 1 mm. of matrix. Additional evidence of the
preservation or influence of soft parts can be seen (PI. 1, fig. 1) in
the thoracic region, where the matrix is unusually dark (and
hard) ; and above the anterior half of the vertebral column, where
a possible dorsal crest is preserved.
The specimen is recognized as a urodele by the presence of
the ypsiloid cartilage, the long, broad parasphenoid reaching to
the foramen magnum, the atlas with an odontoid process, and
haemal arches on the caudal vertebrae. The absence of paired
tooth rows, and the presence of two longitudinal vomerine tooth
rows, diagnose it as a salamandrid. Paroccipital processes,
frontosquamosal arches, and an unpaired premaxilla are char-
acters of this fossil as well as, in Recent salamandrids, Cynops
Tschudi, 1838, Diemictylus Rafinesque, 1820, Euproctus Gené,
1838, Hypselotriton Wolterstorff, 1934, Pachytriton Boulenger,
1955 PALAEOTARICHA OLIGOCENICA 3
1878, Taricha Gray, 1845, and several species of Triturus' Rafin-
esque, 1815. The fossil genera Archaeotriton von Meyer, 1860,
Heliarchon von Meyer, 1863, Oligosemia Navas, 1922, and Poly-
semia von Meyer, 1860, likewise all seem to have frontosquamosal
arches and paroccipital processes, but information on the pre-
maxillae is lacking. The phalangeal formula of the Oregon
specimen is 1-2-3-2 (manus) and 1-2-3-3-2 (pes), as also in
Cynops, Hypselotriton, Pachytriton, and Taricha. Cynops, Hyp-
selotriton, and Diemictylus (the only Recent American sala-
mandrid genus other than Taricha) can be excluded from identi-
fication with the fossil because of their relatively long, narrow
skull, inconspicuous dorsal opening of the cavum internasale, and
failure of the ascending processes of the premaxilla to extend
beyond the anterior border of the cavum. Diemictylus can be
further ruled out by its paired longitudinal ridges extending
from the nasals to the posterior border of the parietals, and an
abrupt angle in the arches posterior to the frontal-squamosal
suture. Pachytriton is excluded by its long, narrow skull; a ridge
(joining the maxillae across the premaxilla) situated between,
and concentric with, the outer tooth row and the anterior border
of the vomer;? and the very small dorsal cavum internasale
opening. Huproctus is ruled out by its phalangeal formula of
2-2-3-2, 2-2-3-3-2, an elongated skull with a spatulate anterior
end, concave lateral maxillary borders, and the paroccipital
processes extending farther back than the occipital condyles.
The five or six species of Triturus that have complete fronto-
squamosal arches are excluded by their phalangeal formula,
2-2-3-2, 2-2-3-3-2, and the elongated skull. Of all the known
salamandrid genera there is no doubt that Taricha is the closest
to the fossil. Skull, body and limb proportions are similar, and
other important characters are: the frontosquamosal arches ;
paroccipital processes; large openings of the cavum internasale,
the dorsal one bordered anteriorly and laterally by the premax-
illa. All these similarities to the Recent Taricha species (torosa,
granulosa, rivularis, and sierrae), and the fact that the fossil
comes from the present geographic range of one of them (granu-
losa), may justify assigning the fossil to that genus. But some
las used by Wolterstorff and Herre (19385).
2 yomeropalatine or prevomer.
4 BREVIORA NO. 45
very important characters distinguishing the fossil from the
Recent species lie in the nature of the vertebrae: characters as
fundamental as some upon which several genera, known from
vertebrae only, have been based. The pelvis is also peculiar, and
the tarsal basale count is different, as described below. With
these differences, and the belief that if the animal were alive,
neoherpetologists would distinguish it generically because of the
spinal column, I feel that erecting a new genus for the fossil is
justified.
Class AMPHIBIA
Order CAUDATA
Family SALAMANDRIDAE
PALAEOTARICHA, new genus
Genotype: Palaeotaricha oligocenica, new species.
Diagnosis: Precaudal neural spines high, with laterally ex-
panded, broad, rectangular, sculptured, flat tops, which are in
contact with each other fore and aft. Fourteen presacral verte-
brae. Pelvis with relatively large, knob-like, laterally directed
prepubie processes. Tarsal basale formula 1, 2, 8, 4+5. Maxillae
extending eaudad, united by suture with quadrates.
PALAEOTARICHA OLIGOCENICA, hew species
Holotype: University of Oregon Museum of Natural History
no. F'-5405, a nearly complete specimen.
Collector: Mr. George R. K. Moorhead, August 1, 1951.
Horizon: Upper Oligocene plant-bearing tuffs, not older than
Eugene and Fisher formations, the Willamette formation of Dr.
Ralph W. Chaney (in litt.).
Locality: About 3 miles (5 km.) southeast of Eugene, Lane
County, west central Oregon; southeast 14, see. 14, T. 18 8, It.
3 W: ‘‘about 550 feet east of the Southern Pacifie railroad tracks
on the south side of a small hill in a cut made for the newly con-
structed highway near the east end of the overpass over the
railroad. This is about 400 ft. north of lat. 44 deg. and 38000 ft.
west of long. 123 deg... . in the Eugene quadrangle’’ (Dr.
Herman Clark, in litt.).
Diagnosis: As for the genus.
co
1955 PALAEOTARICHA OLIGOCENICA
Description: Total length (as preserved), 120 mm.; original
length, ca. 135 mm.; length of skull (anterior end to foramen
magnum, dorsal surface), 15 mm.; width of skull (across paroe-
cipital processes), 13 mm.; length and also width of ischiopubis,
6 mm.; length of limb bones: humerus, 9.5 mm.; radius, 7 mm. ;
ulna, 7.5 mm.; digit 3, manus, 6.5? mm.; femur, 9 mm.; tibia,
5.5 mm.; fibula, 6 mm.; digit 3, pes, 6 mm.
The paroceipital processes are large and are produced laterad,
abutting the squamosals from behind. The frontosquamosal
arches are complete and robust. The dorsal aperture of the cavum
internasale is long and wide, and is bordered anteriorly and
laterally by the ascending processes of the premaxilla. The
vomerine opening of the cavum internasale is large (probably in
part artificially) ; it hes wholly within the vomer. The anterior
border of the vomer is an are of a circle (roughly), with no sharp
breaks in its periphery. No teeth are preserved, but the vomerine
tooth rows are evident though crushed and incomplete.
The vertebrae are relatively shorter than in Taricha. There
are 14 presacral vertebrae (13 in Taricha). Even the atlas has an
expanded top, which, however, narrows anteriad. Instead of the
dorsal surface of the atlas sloping downward anteroventrally as
in Taricha, in Palaeotaricha oligocenica it is horizontal for most
of its length, then there is a short ventrally directed slope toward
the anterior end, and then a sudden vertical drop to the condylar
facets (Pl. 1, figs. 2, 3). The neural spines of the caudal verte-
brae lack the expanded tops.
Five! thoracic ribs of the left side are exposed in good condi-
tion (Pl. 1, fig. 1; Pl. 3, fig. 1). All but the caudalmost have
conspicuous uncinate processes, which, in successively posterior
ribs, become shorter and more distal.
The great length of the scapula suggests that the suprascapula
was ossified (or calcified). There are four digits in the manus.
The phelangeal count is 1-2-8-2, which, together with the carpal
basale for mula (1+2, 3, 4), is the same as in Taricha.
The pelvie girdle has floated free and, with the limbs, is fairly
well preserved as a unit. The pelvic girdle is apparently a highly
ossified and codssified structure. Zones of weakness may perhaps
exist anteriorly between left and right halves, and between the
1Some o tae things that look like ribs in Plate 1, figure 1 are phalanges of the
left maaus.
6 BREVIORA NO. 45
pubes and ypsiloid cartilage. The latter is heavily ossified (or
calcified ; there is no obvious difference in preservation between
it and any other part of the skeleton). About 5 mm. of it are
preserved, of which 2 mm. are proximal to the bifurcation. The
prepubie processes are rounded and robust. They project later-
ally and apparently shghtly posteriad in P. oligocenica, but
anterolaterad in Taricha, where they are also less demarcated
from the pubes. There are five digits in the pes; the phalangeal
count is 1-2-3-3-2. The tarsal basale formula is 1, 2, 3, 4+5,
whereas the Taricha specimens have 1-2, 3, 4, 5.
On the whole the skeleton gives the appearance of having great
strength, with the girdles and limb elements all well ossified, and
the top-heavy neural spines.
Discussion: The elongation of the maxillae seems to be a
primitive character of Salamancricae. In all the fossil genera
of which the skulls are known the maxillae articulate with the
quadrates: Archaeotriton, Brachycormus von Meyer, 1860, Heli-
archon, Oligosemia, Palaeopleurodeles Herre, 1941, and Poly-
semia (all Miocene; Oligosemia also Oligocene). In three of the
more primitive Recent salamandrics, Pleurodeles Michahelles,
1830, Salamandrina Fitzinger, 1826, and Tylototriton Anderson,
1871, the maxillae may very nearly or actually touch the quad-
rates. In the Palaeotaricha oligocenica specimen, although the
maxillae clearly articulate (fuse?) with the suspensoria, the in-
distinct quadrate-squamosal sutures make it difficult to see exactly
which suspensorial element is involved with the maxillae. I have
assumed that the quadrate is involved, because of the condition
in the forms just cited. Other than the maxillae, the P. oligo-
cenica skull seems to fall within the variation of the skulls of the
several Taricha species. In T. granulosa the vomerine tooth rows
generally diverge slightly, from the anterior to the posterior
ends; in the other species they are parallel or very slightly
divergent for most of their length, then suddenly they are
strongly divergent for the remaining 14 or 14. There is, how-
ever, a good deal of individual variation and intergradation be-
tween these tooth-row patterns. The available skulls of 7. sverrae
and T. rivularis (three of the former, four of the latter) seem
to be rather similar, and to stand apart from the skulls of the
other two species. For one thing, the skull of rivularis and
sierrae tends to he somewhat deep and domed, in contrast to the
1955 PALAEOTARICHA OLIGOCENICA
~]
more-or-less flattened and widened roof of the others. 7. rivularis
and sierrae tend to have the dorsal opening of the cavum inter-
nasale especially long and wide. In anterior view it is seen as a
notch whose bottom lies ventral to the dorsal border of the nares,
which is not the case in torosa or granulosa; nor does it extend
so far forward in the latter two species. The above characters
are more pronounced in rivularis than in sierrae, but the fol-
lowing character is equally developed in the two. The ventral
(vomerine) opening of the cavum internasale is greatly elongated
—roughly the size and shape of the dorsal opening — and ex-
tends beyond the anterior border of the vomer into the pre-
maxilla. In the other species this aperture is usually cireular or
slightly elliptical, and is wholly situated within the vomer. In
about ten alcoholic specimens each of rivularis and sierrae, and
many more of torosa and granulosa, this difference is corrobo-
rated. In one skull of granulosa, however, there is a very small,
extra, opening into the cavum internasale anterior to the vomer-
premaxilla border—a feature that would be very difficult to
note in an alcoholic specimen. It would be very interesting to
know whether the elongated aperture is an evolutionary advance
(ef. Palaeotaricha) or is primitive (ef. primitive extant genera of
salamandrids, which have the premaxilla paired, and some have
the aperture extremely large).
A full description of the skull of 7’. torosa is given by Bolkay
(1928) (under the name Diemyctylus |sic| torosus; his specimen
could be a granulosa, judging by the vomerine tooth rows), as
well as descriptions of the skulls of most of the Recent sala-
mandrid species.
The condition in the vertebrae —the expanded tops — of
Palaeotaricha oligocenica is not only different from that in
Taricha, where the neural spines are high but narrow, but quite
different from any other urodele vertebrae that I can find. Among
Recent salamandrids Tylototriton comes closest in this respect,
fide Herre (1949) and corroborated in the specimens available
tome. But in Tylototriton the expansion is much less prominent,
and the width decreases anteriad; the tops of the spines are,
however, sculptured, and they do contact in series, as in P.
oligoccnica. The closest approach of all seems to be in the
Miocene Chelotriton Pomel, 1853, which has a broad top on the
neural spine also. But (at least in the specimen figured by Herre,
8 BREVIORA NO. 45
1949, p. 225, fig. 8) the anterior half of the spine comes to a
rounded point, and the posterior end is bifurcated by a wedge-
shaped notch. Thus it seems that the Chelotriton vertebra has
merely a greatly expanded neural spine, and is not fundamentally
different from that of most salamandrids (e.g. Taricha, Pl. 1,
fig. 5). Now arises the question (considering the absence of
visible sutures between the vertical and horizontal portions of the
P. oligocenica neural spines) of whether we have a structural
series of increasingly expanded neural spines — Taricha, Tyloto-
triton, Chelotriton, Palaeotaricha — or whether the latter is not
in this series, the spinal expansions being dermal plates plastered
on top (as suggested by the sculpturing). I favor the latter view:
the expansions look lke dermal plates; and, of course, the series
just given has only descriptive validity.
The uncinate processes of the P. oligocen’ca ribs are longer
than in the Recent Taricha specimens at hand, but these strue-
tures are quite variable in individual salamanders. In three
specimens of Taricha g. granulosa I have seen, one has uncinate
processes on ribs 2 and 3 on the right side; on rib 2 only, on the
left. Another has uncinates on ribs 2 to 4 on both sides; the
third specimen has them on 2 to 7 on both sides. One specimen of
Cynops pyrrhogaster has uncinates on ribs 2 to 6 on the left side,
on 2 to 7 on the right; another has no uncinates at all. And so
it goes. In all these cases the larger and more fully ossified the
individual, the greater the number of uncinate processes. In
nearly every specimen I have seen there are incipient uncinates
on the first one or two ribs distal to the last one that has a distinct
uncinate process.
The tarsal basale formula in P. oligocenica — 1, 2, 3, 4+5 —
contrasts with 14-2, 3, 4, 5 in the Recent Taricha, Chioglossa
Barboza du Bocage, 1864, Salamandra Laurenti, 1768, Mertensi-
ella Wolterstorff, 1934, and Pleurodeles Michahelles, 1830, while
Salamandrina with four toes on all feet apparently has 1, 2, 3, 4,
and all other genera have 14-2, 3, 4-++5, fide Bolkay (1927), and
in agreement with additional material seen by me.
It is interesting to note that the carpus, as compared with the
tarsus, is apparently much more nearly constant throughout the
Salamandridae. As figured by Bolkay (1927) and corroborated
and extended by the specimens available to me, the carpus has
1955 PALAEOTARICHA OLIGOCENICA 9
basales 1+2, 3, 4, prepollex basale,! centrale, radiale, inter-
medium, and ulnare, in all genera. The latter two may or may
not be fused, but usually are: in one specimen of Triturus vul-
garis the intermedium of the right manus is separate; the left
is fused with the ulnare.
While the observations of Hilton (1948) agree in general with
those of Bolkay and me, Hilton illustrates various cases of fusion
of carpal and tarsal elements, and extra elements (such as the
non-fusion of tarsal basales 4 and 5 in a specimen of Cynops
pyrrhogaster). In the hight of Hilton’s data, the difference be-
tween the Palaeotaricha and the Taricha tarsus may not be sig-
nificant. Yet it must be pointed out that all of the 16 specimens
of Taricha at hand have the same tarsal and carpal structures.
The large size and knob-like shape of the prepubic process of
P. oligocenica also seem to be unique. When these processes
exist in the Recent specimens at hand, they are always smaller,
more pointed, and directed at least partly anteriad. Nor do any
of these Recent pelves have the fusion of the left and right halves
to the extent seen in P. oligocenica, where scarcely even a raphe
remains of a former suture.
From all appearances Palaeotaricha is not ancestral to Taricha,
but is its nearest relative. I do not feel that Palaeotaricha helps
determine the further affinities of Taricha, for the one character
that might seem to indicate relationship with other genera — the
elongated maxillae in Palaeotaricha —is merely primitive and
not unexpected.
Material examined: | have been fortunate in being able to
examine skulls and skeletons of every genus of Recent Sala-
mandridae, with the single exception of Pleurodeles, and to see
alcoholic specimens of nearly every species of all genera. It must
Le emphasized, however, that except for 16 specimens of the three
species of Taricha, in no case have I seen more than two skeletons.
Hence I do not feel that my knowledge of intraspecific variation
is sufficient for more than tentative authority on the above com-
parative skeletal studies.
Note on the stereophotography
The ventral view of the P. oligocenica skull (Pl. 2, A) was
1 Bolkay follows another usage in calling this element carpal basale 1; and what
he calls 2 is here called 142. Likewise he designates the prehallux basale as
tarsal basale 1, and 142 as 2.
10 BREVIORA No. 45
made by Dr. Donald Baird, of this Museum, and myself, with a
binocular (stereoscopic) microscope, and a Leica 3C camera
fitted with a Leitz Focaslide and Leitz Micro-Ibso. The ocular
lens of the latter was inserted alternately into the two tubes of
the microscope, and exposures were made in the two positions
of the apparatus, at an interocular distance of 2144 inches. The
latter refers to the positions of the Focaslide, which was used
because it was found that the side telescope of the Micro-Ibso was
not sufficiently accurate for the depth of focus required. This
method may be theoretically superior to the tilt method usually
used, since it takes care of matters of interocular distance and
angles more easily. But for subjects as large as the present ones
considerable trial-and-error choice of microscope lenses was neces-
sary to get the entire image into view.
All the other figures were made with a Kodak 2-D 5” x7”
view camera fitted with a 135 mm. lens, and mounted on a
pivoted arm. One picture of each stereoscopic pair was taken
vertically, the other at 7° from the vertical. Exposures were
20 seconds at f. 22, using panatomie film, and illuminated by
two 100- and one 200-watt bulb with reflectors, each about 2 feet
from the subject.
ACKNOWLEDGMENTS
I am very grateful to Dr. Donald Baird for his help and the
use of his equipment in the photography; Mr. Benjamin H.
Banta, Stanford University, for the gift of a number of speci-
mens; Mr. Charles M. Bogert, American Museum of Natural
History, for the loan of material; Dr. Ralph W. Chaney, Uni-
versity of California (Berkeley), for stratigraphic information
on the fossil locality, and for permission to use his unpublished
term, ‘‘ Willamette formation’’; Dr. Herman Clark, Willamette
University (now retired), for sending the Palaeotaricha speci-
men, and for providing and permitting me to quote his informa-
tion on the fossil locality; Dr. Max K. Hecht, Queens College,
New York, for much valuable help; Mr. Arthur Loveridge,
Museum of Comparative Zoology, for permission to examine and
skeletonize a number of specimens; Dr. Richard M. Ritland,
College of Medical Evangelists, for the use of material from his
personal collection; Dr. Jay M. Savage, Pomona College, for
valuable help; and above all to Dr. Ernest E. Williams, Museum
of Comparative Zoology, for first suggesting that I undertake
this study, and for much help and continued advice.
1955 PALAEOTARICHA OLIGOCENICA ol
BIBLIOGRAPHY
BoOLKAY, STEPHAN JOSEF
1927. Materialen zu einer vergleichenden Morphologie der Carpal- und
Tarsalelemente der Unterfamilie Salamandrinae. Glasnik Zemal-
jskog Muzeja u Bosni i Hercegovini, vol. 39, pp. 59-67.
1928. Die Schidel der Salamandrinen, mit besonderer Riicksicht auf
ihre systematische Bedeutung. Zeitschr. Anat. Entwicklungs-
gesch., vol. 86, pp. 259-319.
GOLDFUSS, GEORG AUGUST
1831. Beitriige zur WKenntniss verschiedener Reptilien der Vorwelt.
Nova Acta Acad. Leop. Carol., vol. 15, pp. 61-128.
Herre, Wour
1934. Die systematische Stellung von Varicha torosa Eschholz. Blitter
fiir Aquarien- und Terrarienkunde, vol. 45, pp. 250-252.
1935a. Die Schwanzlurche der mitteleocinen (oberlutetischen) Braun-
kohle des Geiseltales und die Phylogenie der Urodelen unter
Kinsechluss der fossilen Formen. Zoologica, vol. 33, art. 4, pp.
1-85.
1935b. Uber Oligosemia spinosa Navas, einen fossilen Schwanzlureh aus
dem spanischen Tertiir. Palaeont. Zeitschr., vol. 17, pp. 91-105.
1941. Palaeopleurodeles hauffi noy. gen. noy. spec., ein fossiler Schwanz-
lurch aus dem Mioziin Stiddeutschlands. Zool. Anz., vol. 134, pp.
1-17.
1949. Neue Tatsachen zur Stammesgesechichte der Schwanzlurche.
Zool. Jahrb., Abt. Syst., vol. 78, pp. 217-236.
1950a. Sehwanzlurche aus dem Paleociin von Walbeck. In Neue Ergeb
nisse und Probleme der Zoologie (Klatt-Festschrift). Leipzig,
Geest und Portig, pp. 286-301.
1950b. Der derzeitige Stand unseres Wissens tiber die fossilen Urodelen,
zugleich einige kritische Bemerkungen iiber Boomgardia_ sala-
mandriformis vy. Huene. Neues Jahrb. Geol. Paliiont., Monatsh.,
1950, pp. 19-25.
llinwon, WILLIAM A.
1948. he carpus and tarsus of salamanders. Jour. Entomol. Zool.,
vol. 40, pp. 1-13.
LAUBE, GUSTAV C.
1901. Synopsis der Wirbelthierfauna der bohm. Braunkohlformation
und Beschreibung neuer, oder bisher unvollstandig bekannter
Arten. Abhandl. deutsch. naturwiss.-med. Ver. BOhmen ‘‘ Lotos,’’
vol. 2, art. 4, pp. 1-80.
Be BREVIORA NO. 45
Mryrr, HerRMANN VON
1860. Salamandrinen aus der Braunkohle am Rhein und in Béhmen,
Palaeontographica, vol. 7, pp. 47-73.
1863. MHeliarchon furcillatus, ein Batrachier aus der Braunkohle von
Rott im Siebengebirge. Jbid., vol. 10, pp. 292-298.
Vokes, H. E., Parkn D. SNAVELY, JR., and Donaup A. MYERS
1951. Geology of the southern and southwestern border areas of the
Willamette valley, Oregon. U.S. Geol. Sury. Oil and Gas Invest.
Map. OM 110.
WoLTERSTORF, WILLY and WoLr HERRE
1935. Die Gattungen der Wassermolehe der Familie Salamandridae.
Arch. Naturgesch., new ser., vol. 4, pp. 217-229.
EXPLANATION OF PLATES
PLATE 1
Kigs. 1, 2, 4. Palaeotaricha oligocenica, holotype, University of Oregon
Museum of Natural History, no. F-5405. Fig. 1. Entire specimen (except
end of tail). Skull seen ventrally, spinal column from right side. Fig. 2,
A, A’. Stereograph of vertebrae 1 to 3, left side. Fig. 4. Vertebrae 10 to
12, left side. The neural spines of all precaudal vertebrae are bent sinistrad.
Figs. 3, 5. Vertebrae of a Taricha torosa. Fig. 3. Vertebrae 1 and 2. Fig.
5. Vertebrae 9 and 10. od, Odontoid process. Fig. 1x1; figs. 2-5 x 6.
PLATE 2
Palaeotaricha oligocenica, skull of holotype. A, A’, stereograph of dorsal
view. B, B’, ventral view. a, Atlas; deci, dorsal opening of cavum internasale;
en, external naris; f, frontal; fsa, frontosquamosal arch; fv, foramen vesti-
buli; hy, parts of hyoid apparatus; in, internal naris; lj, lower jaw; m,
maxilla; n, nasal; nld, nasolacrimal duct?; oc, occipital condyle; op, occi-
pito-petrosal; os, orbitosphenoid (the white dot may be the optic foramen) ;
p, parietal; pam, palatine part of maxilla; pm, premaxilla; poc, paroccipital
process; prf, prefrontal; ps, parasphenoid; pt, pterygoid; q, quadrate; s,
squamosal; vei, vomerine opening of cavum internasale; v, vomer. All figures
x2.
PLATE 3
Palaeotaricha oligocenica, holotype. Fig. 1. Right pectoral girdle and
limb, and ribs 3 to 7. Forearm and manus in dorsal view, other parts in right
side view. be 1+2, Carpal basales 1+2; be {44, carpal basaleg vh4; h,
humerus; me 1, metacarpal 1; ppb, prepollex basale; 7, radius; ra, radiale;
16, rib 6; s, scapula; uv, ulna; ul+i, ulnare + intermedium.
Fig. 2. Pelvie girdle and limbs. Pelvis and left limb in ventral view, right
limb (left side of photograph) in dorsal view. bt 4+4, Tarsal basales 4-++5;
c, centrale; f, femur; fe, fibulare; fi, fibula; 7, intermedium; phb, prehallux
basale; prp, prepubie process; ft, tibia; ti, tibiale; y, ypsiloid process. Both
figures x 244.
PLATE 1
wo
D)
PLATE
PLATE :
Oo
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. June 24, 1955 NUMBER 46
CAVE-FOSSIL VERTEBRATES FROM
CAMAGUEY, CUBA
By Karu F. Koopman
Department of Biology, Queens College, Flushing, New York
and RopoLtro RurBaL
Division of Life Sciences, University of California, Riverside
Of all the Cuban provinces, Camaguey is the least known in
regard to its former and present vertebrate fauna. With this fact
in mind the junior author, in January of 1952, visited the Sierra
de Cubitas in Camaguey with the purpose of locating cave-fossil
material.
The Sierra de Cubitas is a range of low (maximum altitude of
750 feet) Tertiary and Cretaceous limestone hills running north-
west to southeast along the northern half of Camaguey. The
hills rise abruptly from the flat serpentine Camaguey savanna,
and are covered by one of the few remaining tracts of tropical
forest in the province. Two caves were visited by the junior
author. Both are located on the south slope of the hills, at the
Trinchera de Martin Castillo (the site of a battle during the
war against Spain), which is about seven kilometers west of the
village of Banao. The first cave was the Cueva del Indio (re-
ferred to as Cave 1 henceforth in the text), an extensive cave that
is on occasion visited by excursionists from the city of Camaguey.
The second cave lies a short distance east of Cave 1, and will be
designated as Cave 2, since it bears no known name.
The present entrance to Cave 1 is a broad, high-domed opening
which is continuous with the large chamber from which the
deeper passages of the cave extend. Due to the ample entrance
the entire chamber is well within the twilight zone. The floor of
the chamber has been excavated for bat guano. The entrance to
the cave is higher than the chamber floor, and consists of rocks
fallen from the eroded cave front. The red soil of the chamber
2 BREVIORA No. 46
was damp, but at higher levels near the entrance the soil was
dry. However, during rains, water enters the cave and material
has been deposited at different levels near the entrance. All of
the fossil material collected was found as surface litter deposited
at different points near the cave entrance. Material was ex-
tremely abundant, any handful from the surface yielding many
bones and snail shells. Collections were made at random from the
many pockets where the litter consisted almost exclusively of
bones and snail shells.
In the case of Cave 2 erosion has cut back so that it has exposed
to direct sunlight a former high dome, about seventy feet high)
with massive stalactites and stalagmites along the sides. In this
eave, digging for bat guano had only ceased two years previous
to the time of the visit and consequently the entire floor of this
shallow cave is covered with the mounds of red litter that were
screened out by the guano collectors and the material washed out
from these mounds by the rain. The abundance of bones in this
eave can only be described by saying that there are millions. To
walk in any part of this cave is to walk upon a floor of bones
and snail shells. Collecting in this cave was essentially the same
as in Cave 1, but of a greater volume. The material was scooped
up at random from the mounds and from the floor. Due to the
little time available no adequate investigation was done in any of
the more protected parts of the cave where material was still un-
disturbed by the guano collectors.
The snail shells collected were kindly identified by Dr. W. J.
Clench of Harvard University. Eight species were found and
according to Dr. Clench all the species are at present common
in Camaguey.
MATERIAL, COLLECTED
Class MAMMALIA
Order INSECTIVORA
Nesophontes micrus. Represented by one maxilla, eight man-
dibles, three femora, and one humerus from Cave 1, and by two
partial skulls, eight mandibles, and three femora from Cave 2.
This species was first described from the provinces of Matanzas
(Allen, 1917b), and later from Oriente (Anthony, 1919) and
Isla de Pinos (Allen, 1918). In addition there is material in the
Museum of Comparative Zoology from Las Villas and Pinar
1955 CAVE-FOSSIL VERTEBRATES FROM CUBA a
del Rio, and in the United States National Museum from Habana
province. The present material from Camaguey completes the
record and demonstrates that the species existed throughout
Cuba.
Solenodon cubanus. Represented by a portion of one mandible
and six isolated teeth from Cave 1. At least two individuals are
represented in the material collected. This species is known
today as a living animal only from Oriente (8. poeyi is con-
sidered a subspecies, following Aguayo, 1950). It has previously
been found fossil in both Oriente (Allen, 1918) and Pinar del
Rio (Aguayo, 1950). It apparently also occurred in the Sierra de
Trinidad in the province of Las Villas up until the middle of
the nineteenth century (Gundlach, 1895). The present material
agrees well with fresh specimens from Oriente, but differs by
somewhat larger size and robustness. Since the species was dif-
ferentiated into two subspecies in Oriente, and the animals from
the Sierra de Trinidad appear to have been distinct in color from
the Oriente forms, it is possible that the Camaguey material may
represent a different subspecies. However, the material available
is not adequate to determine this.
Order CHIROPTERA
Macrotus waterhousei. Represented by two mandibles from
Cave 1, and a portion of one skull and three mandibles from
Cave 2. This species was previously known from Isla de Pinos
and all the provinces except Camaguey. Apparently a common
and widely distributed form.
Monophyllus cubanus. A single mandible from Cave 2. A
single periotic bone from Cave 2 is also allocated to this species.
All recent records of this bat appear to be from Oriente, but
Gundlach (1868, pp. 48, 49) records it from Rangel in Pinar
del Rio.
Brachyphylla nana. Represented by two mandibles from Cave
1, and nine partial skulls and 18 mandibles from Cave 2. This
species was previously known from all the provinces of Cuba
except Camaguey and Isla de Pinos. A common and widespread
species.
Artibeus jamaicensis. Represented by five partial skulls and
two mandibles from Cave 1, and eight partial skulls and twenty-
4 BREVIORA No. 46
six mandibles from Cave 2. Presumably known from all the
provinces of Cuba and from Isla de Pinos, though we have been
unable to find any definite record for Matanzas. Several speci-
mens of this species were shot in Cave 1. A very common and
widespread species.
Phyllops falcatus. Represented by two partial skulls and three
mandibles from Cave 1, and ten partial skulls and twelve man-
dibles from Cave 2. Previously recorded from Oriente, Las Vi-
llas, Matanzas, and Pinar del Rio, but probably occurs through-
out Cuba and Isla de Pinos. None of the fossil material from
Camaguey represents P. vetus, still known only as fossil from
Oriente (Anthony, 1919).
Erophylla sezekorm. Represented by two mandibles from
Cave 1, and two rostra and seven mandibles from Cave 2. Pre-
viously known from Habana, Las Villas, and Oriente, it is prob-
ably rather widespread in Cuba.
Phyllonycteris poeyi. Represented by one mandible from Cave
1, and by ten rostra and fifteen mandibles from Cave 2. Previ-
ously known from Pinar del Rio, Habana, Matanzas, Las Villas,
and Oriente, and is thus a widespread species.
Natalus primus. Represented by a single mandible and a
fragmentary rostrum from Cave 2. This species was first de-
seribed as a fossil from Oriente by Anthony (1919), but it is also
known from Las Villas (Koopman, unpublished). Our record is
the first from anywhere else and the first rostral material, the
other finds consisting entirely of mandibles. Known only as fossil,
this species was apparently rare but probably fairly widespread.
Tadarida brasiliensis. (We follow G. M. Allen, 1911, in con-
sidering 7. muscula a subspecies of T. brasiliensis.) Repre-
sented by two rostra and two mandibles from Cave 2. Previously
known from all the provinees of the Cuban mainland. Evidently
a common and widespread form.
Tadarida molossa. Represented by one rostrum and one man-
dible from Cave 2. These appear to be the first Cuban specimens
of this species with definite locality data, T. macrotis, a synonym,
having been described only from ‘‘the interior of Cuba”’ (Allen,
1911). Apparently a very rare or local form.
The geographic data for the bats were obtained from Miller
(1904), Allen (1911), Peterson (1917), Anthony (1919) and
from the specimens in the collections of the Museum of Compara-
1955 CAVE-FOSSIL VERTEBRATES FROM CUBA 5
tive Zoology at Harvard, the American Museum of Natural His-
tory in New York City, and the United States National Museum
in Washington, D. C.
Order RODENTIA
Boromys offella. Represented by one mandible and two iso-
lated teeth from Cave 2. Originally described as fossil from
Oriente (Miller, 1916) it has since been recorded from Las Villas
(Allen, 1918), Pinar del Rio (Museum of Comparative Zoology
specimens), and Isla de Pinos (Allen, 1918). It apparently was
formerly a widespread species.
Boromys torrer. Represented by one mandible from Cave 1
and by two rostra, fifteen mandibles, and three isolated teeth from
Cave 2. Ten auditory bullae from the latter deposit are also
tentatively referred here. This species has previously been found
in Oriente (Anthony, 1919), Las Villas (Museum of Comparative
Zoology), Matanzas (Allen, 1918), and Pinar del Rio (Museum
of Comparative Zoology), and also on Isla de Pinos (Allen, 1918).
Evidently this was formerly a widespread and common species.
Capromys (Mystateles) nana. Represented by a single tooth
from Cave 1, and by one rostrum, four mandibles, and six isolated
teeth from Cave 2. This species is known as a living animal only
from the Cienega de Zapata (Allen, 1918 and 1942), and as a
fossil was only known from Matanzas (Allen, 1917a), Las Villas
(Allen, 1918), and Oriente (Anthony, 1919). Apparently orig-
inally rather widespread.
Capromys (Mystateles) prehensilis. A single rostral fragment
from Cave 2 is best allocated to this form. The species occurs
in Pinar del Rio, Habana, Matanzas, and Las Villas, and is repre-
sented by a subspecies on Isla de Pinos (Mohr, 1939). A wide-
spread form in western and central Cuba.
Capromys (Geocapromys) columbianus. Represented by one
mandible and five isolated teeth from Cave 1, and by three man-
dibles and five isolated teeth from Cave 2. This species was
originally described from cave-fossil material from Las Villas
(Chapman, 1892), and it has since been recorded from Oriente
(Anthony, 1919), Matanzas, Pinar del Rio, and from Isla de
Pinos. Apparently once a common and widespread form.
Rattus. Represented by many partial skulls, mandibles, and
6 BREVIORA No. 46
posteranial skeletal parts from Cave 1 and Cave 2.
Mus. Skeletal remains from both Cave 1 and Cave 2. Not as
abundant as the Rattus remains.
A great deal of posteranial and some cranial material, particu-
larly of rodents, remains unidentified.
Class AVES
The bird remains collected at Cave 1 and Cave 2 were not
identified.
Class REPTILIA
Order SQUAMATA
Anolis lucius. A few dentaries and maxillary elements from
Cave 1 are referable to this species. In Cave 2 only a single
parietal of this species was found. This is, at present, a common
lizard inhabiting the limestone walls at the entrance of the caves.
Anolis equestris. Represented by the proximal portion of a
lower jaw and portion of one maxilla from Cave 2. This species
was observed inhabiting the forest outside of the caves.
Lewocephalus sp. A large dentary that seems to represent a
species of Letocephalus was collected at Cave 1. Part of a maxilla
from Cave 1, and a dentary and a maxilla from Cave 2 are also
referable to this genus.
Tarentola americana. This rare gekkonid species is represented
by a single dentary from Cave 2.
Alsophis angulifer. A few vertebrae from Cave 2 are referable
to this species of snake.
Some cranial and posteranial reptilian material from Caves 1
and 2 remains unidentified.
Class AMPHIBIA
Order ANURA
Hyla septentrionalis. Cranial and postcranial elements of this
common tree frog were the most abundant non-mammalian re-
mains to be found in Cave 1 and Cave 2. Some of the bones
found appeared to be more rugged and larger than those of the
two skeletons prepared for comparison.
1955 CAVE-FOSSIL VERTEBRATES FROM CUBA 7
DISCUSSION
Most of this collection probably represents the result of ow)
predation. However, it is probable that some of the material does
not represent owl pellet remains but rather bones washed into the
cave, the remains of animals that took refuge in the cave, or
remains of native kitchen middens.
None of the material was stratified, all of it having been inter-
mixed by the diggings for bat guano. It is therefore impossible
to establish the relative age of the fossils. The presence of two
introduced European species — Rattus and Mus — proves that at
least part of the material collected is post-Columbian in age.
However, the presence of species now extinct in Cuba — Neso-
phontes, Natalus primus, Boromys, and Capromys (Geocapro-
mys) columbianus — or extinet in Camaguey — Solenodon and
Capromys (Mystateles) nana — suggests an early pre-Columbian
age for some of the material. There is no evidence of difference
in age or type of deposition between Cave 1 and Cave 2.
The collections from these caves also establishes the presence of
a number of species previously not recorded from Camaguey.
BIBLIOGRAPHY
AGUAYO, C. G.
1950. Observaciones sobre algunos mamiferos cubanos extinguidos.
Bol. Hist. Nat. Soc. Felipe Poey, 1: 121-134.
ALLEN, G. M.
1911. Mammals of the West Indies. Bull. Mus. Comp. Zool., 40: 173-
263.
1917a. An extinct Cuban Capromys. Proc. New England Zool. Club,
6: 53-56.
1917b. New fossil mammals from Cuba. Bull. Mus. Comp. Zool., 61:
1-12.
1918. Fossil mammals from Cuba. Bull. Mus. Comp. Zool., 62: 131-148.
1942. Extinet and vanishing mammals of the Western Hemisphere.
Lancaster, Pennsylvania.
AntHony, H. E.
1919. Mammals collected in eastern Cuba in 1917, with descriptions
of two new species. Bull. Amer. Mus. Nat. Hist., 61: 625-643.
CHAPMAN, F. M.
1892. Notes on birds and mammals observed near Trinidad, Cuba,
with remarks on the origin of West Indian bird life. Bull.
Amer, Mus. Nat. Hist., 4: 279-330.
8 BREVIORA No. 46
GUNDLACH, J.
1868. Revista y catalogo de los mamiferos cubanos. Repertorio Fis.
Nat. de la Isla de Cuba, Habana, 2: 40-56.
1895. Notes on Cuban mammals. Proc. Linn. Soc. N. Y., 7: 13-20.
MILLER, G. 8.
1904. Notes on the bats collected by William Palmer in Cuba. Proc.
U.S. Nat. Mus., 27: 337-348.
1916. Bones of mammals from Indian sites in Cuba and Santo Do-
mingo. Smithsonian Misc. Coll. 66, (12): 1-10.
Morr, E.
1939. Die Baum- und Ferkelratten-Gattungen Capromys Desmarest
(sens. ampl.) und Plagiodontia Cuvier. Mitt. Hamburgischen
Zool. Mus. Inst., 48: 48-118.
PETERSON, O. A.
1917. Report upon the fossil material collected in 1913 by the Messrs.
Link in a cave in the Isle of Pines. Ann. Carnegie Mus., 11:
359-361.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. Juny 12, 1955 NUMBER 47
A NEW SPECIES OF WHIPTAILED LIZARD
(GENUS Cnemidophorus) FROM THE
COLORADO PLATEAU OF ARIZONA, NEW
MEXICO, COLORADO, AND UTAH
By CuHarues H. Lowe, JR.
Department of Zoology, University of Arizona
In a previous paper it has been shown that the holotype of the
form of Cnemidophorus described by Burger (1950) as C. sacki
stictogrammus represents a distinct species and is not a member
of the sympatric species C. sacki to which it was assigned. In the
present paper C. sacki mnotatus also described by Burger (op.
cit.) in the same study is similarly shown to be a different species
from C. sacki; it is Cnemidophorus velox described by Springer
(1928) as C. gularis velox. Both of these forms described by
Burger (stictogrammus, and innotatus = velox) are sympatric
with C. sacki in Arizona.
Neither Burt (1931) nor Burger (1950) properly assigned
Springer’s (1928) material. Springer (op. cit.) designated no
specimen as type and his four original specimens were cotypes.
Burger (op. cit.) accepted M.C.Z. No. 37208 as lectotype of velox
by Burt’s (1931:124, 127) designation; this specimen is also
accepted here as the type specimen. However, Burger continued
the error of the identity of this specimen with the statement
(1950:3) that ‘‘It is an example of C. inornatus,’’ and proceeded
to redescribe velox as C. sacki innotatus. M.C.Z. No. 37208 is
not an example of C. inornatus (= perplexus auct.) as is shown
by analysis of characters of scalation ; it is an example of C. velox
Springer. Burger (1950:4), in redeseribing C. velox as C. sacki
innotatus, states that ‘‘Springer (1928) may have had specimens
of this lizard [innotatus] on hand when he described velox; in
9 BREVIORA No. 47
some respects his description fits it better than wnornatus.’’ This
is quite true.
During the past few years I have collected topotypes of C.
velox from Oraibi, Arizona, and Pueblo Bonito (in Chaco Can-
yon National Monument), New Mexico; both of these localities
were spoken of by Springer (1928) as localities from whence
came his type series of four specimens of C. gularis velox (Oraibi,
3; Pueblo Bonito, 1). The topotypic material of C. velox is
species-identical with a series of topotypes of C. sacki innotatus
EE — INORNATUS
N=25
oo KANAB, 16 ~— VELOX
140 150 160 170 ‘180 190
DORSAL SCALES (OCCIPUT TO RUMP)
Comparison of C. inornatus Baird and Girard and C. velox Springer. The
white rectangles enclose two standard errors of the mean. One white plus
black rectangle encloses one standard deviation. The differences between
the two species are highly significant. The differences between the two
populations of C. velox are not significant. Data from Tables 1 and 2.
Burger that have also been collected for this study in the vicinity
of Kanab, Utah, the type locality. Moreover, this species (C. velox)
has been collected together with C. sacki at several localities in
Arizona, as further discussed below.
In Table 1 are presented results of statistical analysis of char-
acters of scalation of C. sacki, C. inornatus, and C. velox. Table
2 presents a comparison of C. inornatus with samples of C. velox
from the two type localities involved: (a) the type locality of
1955 NEW SPECIES OF WHIPTAILED LIZARD 3
Springer’s (1928) C. gularis velox (Oraibi, Navajo County,
Arizona), and (b) the type locality of Burger’s (1950) C. sacki
imnotatus (vicinity of Kanab, Kane County, Utah). Both of
these type localities are in Pinyon-Juniper Woodland. Only one
other species of lizard in the genus Cnemidophorus is known to
occur in the vicinity of Oraibi or Kanab; this is C. tigris which is
represented by the northern subspecies C. tigris septentrionalis
Burger which is a valid and especially distinctive race. From
analysis of scalation and of color pattern it is clearly evident
that C. gularis velox Springer (1928) = C. sacki innotatus Bur-
ger (1950).
The question now arises as to whether velox is or is not con-
specifie with sachi (— gularis) as now understood. This question
has been unequivocally answered by having found velox and
sacki to be sympatric over a wide area where the ranges of the
two overlap in Arizona. At such localities each maintains its
own respective genetic differences without evidence of gene
exchange. One is able to shoot adults of both species (velox and
sacki) while standing in the same spot at localities where they
are sympatric (e.g., Pinal Mountain, Gila County; Prescott
Mountains, Yavapai County; Oak Creek, Coconino County).
See Table 1 for certain similarities and differences in morpho-
logical characters of these two species.
Cnemidophorus velox is a whiptail of Woodland and Coniferous
Forest habitats. It is the common striped whiptail of the so-called
Colorado Plateau of northern Arizona and New Mexico, southern
Utah and Colorado. It exceeds in both altitude and north lati-
tude the ecologic distribution of C. sacki in Arizona and New
Mexico. C. sacki does oceur (marginally) in Coniferous Forest
and occurs in Pinyon-Juniper Woodland. Thus on outlying
mountain masses south of the main body of the Colorado Plateau,
the two species are found to be sympatrie in Woodland and
Coniferous Forest habitats in central Arizona, near the northern
limit of the geographie distribution of C. sackt and the southern
limit of C. velox.
The much larger C. velox is also specifically distinct from the
small C. inornatus (= perplexus auct.). These two species re-
semble one another superficially in color and pattern, particularly
in the total absence of light spots on the body of both juvenile
and adult; small individuals of the two species have been most
4. BREVIORA No. 47
commonly confused. The highly significant difference in the
dorsal sealation of these two species is shown in Table 2 and
Figure 1. The species rather than subspecies relationship of the
two is also clearly evinced by the fact that the much larger
bodied form (C. velox) is the one with the northward distribution
(on the Colorado Plateau) while the smaller sized form (C. in-
ornatus) is a predominantly Mexican population ranging only
into the southern United States. This relationship of surface-
volume ratio to environmental temperature is opposite to that
which is to be predicted on the basis of the Bergmann Principle
applied to ectotherms (see Fitch, 1940 :123; Cowles, 1945). Also
C. inornatus is a species primarily of grasslands and plains and
does not reach the higher elevations attained by C. velox in Conif-
erous Forest habitats (7000-8000 ft.) in the United States.
The following diagnosis of C. velox Springer (1928:102) is
based upon the type specimen, topotypes recently collected, and
additional series recently collected in Colorado, Utah, Arizona,
and New Mexico.
CNEMIDOPHORUS VELOX Springer
Plateau Whiptail
Type specimen. No. 37208, Museum of Comparative Zoology.
Collected in August, 1928 by Stewart H. Springer. The type
locality is here restricted to Oraibi, Navajo County, Arizona.
See discussion below.
Diagnosis. A moderately-sized species of Cnemidophorus char-
acterized by: (1) a maximum snout-vent length of approximately
85 mm.; (2) 6 or 7 longitudinal light body stripes, with the
seventh (vertebral) stripe, when occurring, less distinct than the
remaining stripes; (3) total absence of hght spots in the dark
fields on the body; (4) ground color of the upper surfaces of the
body black to blackish-brown; (5) ventrum immaculate whitish
_very faintly tinged with bluish; (6) distal portion of tail light
bluish in adults and bright blue in juveniles; (7) 171.1 + 1.3
(160-184) scales on the midline from occiput to rump; (8)
73.1 + 0.63 (63-85) scales around midbody, excepting the en-
larged ventrals; (9) 7.7 + 0.16 (5-10) scales between paraverte-
bral stripes at midbody; (10) mesoptychial scales conspicuously
enlarged and abruptly differentiated from the adjacent granu-
1955 NEW SPECIES OF WHIPTAILED LIZARD 5
lar scales of the gular fold; (11) postantebrachial scales not
greatly enlarged (not enlarged as in C. sacki) to 4 or more times
the size of adjacent scales.
Distribution. On the so-called Colorado Plateau of northern
Arizona and New Mexico, southern Utah and southwestern
Colorado. The range extends southward on outlying mountain
ranges to at least central Arizona and to central or southern
New Mexico. The relationships to C. velox of a group of ap-
parently non-spotted, small, mountain-dwelling populations of
northern Mexico is currently being investigated.
Comparisons. C. velox is distinguished from C. sacki, with
which it is sympatric, by characters of color pattern and body
size as well as sealation (see Table 1). C. velox is smaller, non-
spotted, has a black to blackish-brown ground color, and a light
bluish tail (adults) that is bright blue in juveniles; C. sacki is
larger, spotted, has a light brown to dark brown ground color,
and a brownish tail.
C. velox, while superficially resembling C. inornatus in color
pattern, is clearly distinguished from inornatus in body size and
scalation (see Tables 1 and 2). C. velox is considerably longer,
larger, and with a greater number of dorsal scales.
Discussion. Smith and Taylor (1950:184) give the type speci-
men of C. gularis velox Springer as Butler University No. 848,
thereby following Burt (19381:124, 127) while at the same time
accepting and using Burger’s (1950:3) nomenclature (C. in-
ornatus with C. gularis velox a synonym) based on the type
specimen in question as Harvard M.C.Z. No. 37208. The problem
is not so simple as stating or assuming that Butler Univ. No. 848
is now M.C.Z. No. 37208.
Mr. Arthur Loveridge of the Museum of Comparative Zoology
has kindly checked details of the record concerning M.C.Z. No.
37208. He has recently informed me that the M.C.Z. has had no
recorded dealings with Butler University, that the museum
register records this specimen as having been received from one
J. Piatt on January 11, 1934, and that it is the holotype of
Cnemidophorus gularis velox Springer, collected at Lee’s Ferry,
Arizona, August, 1928, by Stewart H. Springer. The data on the
specimen label gives the same information, including ‘‘Gift from
J. Piatt, Indiapolis[sic].’’ The specimen in question (No.
37208) is recorded as the holotype of Cnemidophorus gularis
6 BREVIORA No. 47
velox Springer in the second list of type material in the M.C.Z.
(Barbour and Loveridge, 1946 :92) ; this action, with type locality
given as Lee’s Ferry, Arizona, antedates the action of Smith and
Taylor (1950) who restrict the type locality to Pueblo Bonito,
New Mexico. While it is possible that M.C.Z. No. 37208 could be
Butler Univ. No. 848, there is nothing in the record to so indicate.
As far as can be now determined, there is no record at Butler
University for the final disposition of this specimen or any of the
others in the type series of C. gularis velox Springer. Dr. N. EH.
Pearson, Head of the Department of Zoology of Butler Univer-
sity, informed me on December 1, 1954, that he was unable to
find either a record or specimen used by Springer in describing
Cnenidophorus gularis velox.
In a paper entitled ‘‘An annotated list of the lizards of Lee’s
Ferry, Arizona,’’ Springer (1928) described Cnemidophorus
gularis velox and recorded the collection of his four specimens
(ecotypes) as follows: Oraibi, Arizona, 3; Pueblo Bonito, New
Mexico, 1. My study of M.C.Z. No. 37208 shows it to be a speci-
men of C. velox described by Springer, and not a specimen of
C. inornatus (= perplexus auct.) as is stated by Burger (1950:3).
I consider the holotype of C. velox (= C. gularis velox)
Springer to be M.C.Z. No. 37208, the type locality of which is
here changed to Oraibi, Navajo County, Arizona, rather than
Lee’s Ferry, Coconino County, Arizona (fide Springer, 1928).
Ample topotypes are now available from Oraibi and vicinity.
Burger (1950) concurs in the citation of M.C.Z. No. 37208 as the
type of C. g. velox Springer. The later action by Smith and
Taylor (1950:184) in their restriction of the type locality to
Pueblo Bonito, San Juan County, New Mexico, is not acceptable.
Dr. Hobart Smith has recently concurred in this opinion (in
litt.).
ACKNOWLEDGEMENTS
I wish to express my appreciation of Dr. Doris M. Cochran
of the United States National Museum, Dr. Raymond B. Cowles
of the University of California, Los Angeles, Dr. Howard K.
Gloyd of the Chicago Academy of Sciences, Mr. Arthur Lover-
idge of the Museum of Comparative Zoology, and Dr. T. Paul
Maslin of the University of Colorado, for the loan of specimens
1955 NEW SPECIES OF WHIPTAILED LIZARD a
under their care, and to Dr. J. T. Marshall of the University of
Arizona for the collection of specimens important to this study.
LITERATURE CITED
Barsour, T. and A. LOVERIDGE
1946. First supplement to typical reptiles and amphibians. Bull. Mus.
Comp. Zool., 96:(2) : 57-214.
Burger, W. L.
1950. New, revived, and reallocated names for North American whip-
tailed lizards, genus Cnemidophorus. Chicago Academy of Sci-
ences, Nat. Hist. Misc., 65: 1-9.
Burt, C. E.
1931. A study of the teiid lizards of the genus Cnemidophorus with
special reference to their phylogenetic relationships. Bull. U. 8.
Nat. Mus., 154: 1-286.
Cowtes, R. B.
1945. Surface-mass ratio, paleoclimate and heat sterility. Amer. Nat.,
79: 561-567.
Fircu, H.
1940. <A biogeographical study of the ordinoides Artenkreis of garter
snakes (genus Thamnophis). Univ. Calif. Publ. Zool., 44(1):
1-150.
SmiruH, H. M. and HE. H. TaYtor
1950. An annotated checklist and key to the reptiles of Mexico exclu-
sive of the snakes. Bull. U. S. Nat. Mus., 199: 1-258.
SPRINGER, S.
1928. An annotated list of the lizards of Lee’s Ferry, Arizona. Copeia,
169: 100-104.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. OcroBER 10, 1955 NuMBER 48
NOTES ON AMERICAN EARTHWORMS OF
THE FAMILY LUMBRICIDAE. J-II.
By G. E. GatEs
Information as to variation of taxonomically important char-
acters in our little known endemic lumbricids hitherto has been
lacking. Some now can be recorded through the kindness of
Dr. G. E. Pickford, Prof. Walter Harman, Ottys Sanders, and
others. The author’s thanks are extended to all those who have
supplied or loaned material.
I
EISENIA LONNBERGI (Michaelsen) 1894
Mt. Carmel, New Haven, Conn. In wet soil at bank of river,
Oct. 28, 1928, 1-0-3. Dr. G. E. Pickford.
Cockaponsett, Conn. Near Indian caves, Nov. 9, 1941, 0-0-1.
Dr. G. E. Pickford.
Cockaponsett State Forest, Conn. In wet soil (pH ca. 5.5)
under water-logged logs at edge of lake (water level low
leaving muddy shore with stumps), Oct. 6, 1940, 0-1-2.
Dr. G. E. Pickford. (Clitellate specimens were copulating
when taken. )
Meshomasic State Forest, Conn. By stream (pH ca. 5.0),
April 28, 1935, 0-0-3. In soil (pH ca. 5.0) in bank of
stream below Hypochilus colony, Nov. 4, 1935, 0-0-1. Dr.
G. E. Pickford.
Woodbridge, Conn. In marshy soil of alder copse near Bald-
win’s Road, Wehawaug River valley, Oct. 21, 1928, 1-0-2.
Same locality, in marshy soil (pH ca. 6.2), Oct. 16, 1928,
2 BREVIORA No. 48
1-0-1. At edge of small pond, June 14, 1931, 0-0-1. Dr.
G. KE. Pickford.
Tallulah Falls, Georgia. In ravine, in sandy loam in seep of
stream, July 27, 1931, 1-0-1383. J. M. Valentine per Dr.
G. EK. Pickford. (‘‘Abundant’’ according to collector.)
Mt. Vernon, Virginia. From nearby swamp, April 22, 1917,
0-0-1 (macerated). Walter D. Webb Jr., per Prof. E.
Carpenter. (Two other Virginia specimens, kindly loaned
by Prof. Harman, have been examined.)
External characteristics. Length, 77-104 mm. (Conn.), -130
mm. (Georgia). Diameter in clitellar region which is flat on
ventral side, 6-8 mm. Body becomes transversely rectangular in
cross section posteriorly. Segments, 114-134 (cf. Table). Pig-
mentation in dorsum, dark, rather nondescript, at least in present
condition, possibly lighter in vili-xi near mD. Prostomium epilo-
bous ; tongue short, open or closed (3 specimens). Setae begin on
ii (33) on which all are usually present, located at four corners
of the body posteriorly where AB < or ca. = CD, BC < AA,
AA ca. = DD. Nephropores recognizable only on a few of the
segments, first seen on iv, usually in or close to CD on iv-xiii,
between levels of female and male pores on xiv-xvi, elsewhere
in median part of BC or dorsal to D but without regular alterna-
tion or segmental symmetry, occasionally in DD on one side
even in xiv-xvi. First dorsal pore on 4/5 (32), 6/7 (1).
Spermathecal pores on 8/9-10/11 (380 specimens), 8/9-11/12
(1), just lateral to mD, each on a small tumescence and in a
longitudinal groove so that four small tubercles seem to be
present on each intersegmental furrow. Female pores at eq/xiv
just lateral to B. Male pores each at bottom of a slight trans-
verse depression, at eq/xv and much nearer B than C.
Clitellum red (formalin preservation), saddle-shaped, reach-
ing ventrally nearly to B or to tubercula pubertatis, 23/24-
eq/xxx (1), xxiii-xxx (5), xxiv-xxx (22). Tubercula pubertatis
longitudinally placed, rather broad bands just lateral to B, on
XXVi-Xxvili (31), occasionally continued onto anterior half of
xxix and/or posterior half of xxv but there gradually narrowed,
markedly protuberant (several) and then sometimes with b setae
on median face, the b setae on several other specimens slightly
1955 AMERICAN EARTHWORMS OF THE FAMILY LUMBRICIDAE I-II 3
displaced mesially. Epidermis in AA of clitellar segments about
as thick as in genital tumescences.
Genital tumescences present (or recognizable?) only on three
worms, around c-d of viii (1), of xi-xii (1), around a-b of x (1),
of xx-xxx (1, ex copula), of xxili-xxx (1, ex copula), of xxiii (1).
The a and 6 setae of xx-xxx (1), xxi-xxx (1), xxii-xxx (8), xxiii-
rox CLO) xxix, (11), exxiv-xxix’ (1) xxivexxx ()) xxvi-
xxviii (1) are deeply retracted into the parietes and the margin
of each follicle aperture is protuberant as a slight but obvious
annular tumescence. The a and Db setae of x (3) and/or the ¢
and d setae of x (7) likewise are retracted and follicle apertures
have an annular tumescence. Male pore tumescences slight,
median to the pore, usually not reaching either intersegmental
furrow, occasionally quite unrecognizable.
Internal anatomy. Septa 12/13-14/15 muscular and increas-
ingly thickened posteriorly.
Oesophageal wall shghtly thickened posteriorly in x but with-
out any trace of sacs. Calciferous glands large, in xi-xii, not
quite reaching insertion of 12/13. Intestinal origin in xv (33).
Gizzard in xvii-xvill, some special muscularity in xix in at least
one of the worms. The typhlosole begins abruptly in region of
XXIV-XXVli or may be recognizable from ca. xx as a flat horizontal
band gradually thickened and heightened posteriorly. The shape
in cross section behind xxviii is fora time like that of an inverted
T but the ventral margin gradually becomes more rounded. The
typhlosole is high, nearly reaching floor of gut and when the
ventral portion is cut off readily separates into two lamellae, the
space between with only a few delicate fibres and a little granular
matter. The end, rather abrupt, normally is in region of ¢-exv
(cf. Table I).
The subneural trunk, adherent to nerve cord, usually is fairly
large and filled with blood but when empty may be unrecogniz-
able. A large vessel passes up from each extra-oesophageal trunk
in vicinity of 9/10. Last hearts in xi (383), usually small and
empty. Hearts of x often smaller than those of ix. Hearts of vi
or vi-vii sometimes also are small. Nephridia have been found only
from iii posteriorly (lacking in 11?). Nephridial duets apparently
pass into parietes in region of B.
Seminal vesicles three pairs, in ix, xi-xii, those of xii the larg-
4 BREVIORA No. 48
est, those of ix the smallest though still of fair size. Male deferent
ducts appear to be slightly thickened in the entalmost portion
and rather sinuous but without definite loops on posterior faces
of septa just behind the funnels. Spermathecal ducts slender,
usually longer than the ampullae which are in ix-xi or ix-xii.
Ovisaes small, shghtly lobed in each adult.
Glandular tissue is present on the parietes median to A (and
also lateral to B) in a band that extends from xxx to or well
towards xxii and mesially nearly to the nerve cord. Similar tis-
sue is present on the parietes laterally in xi-xii of one worm but
none is present over the male pore tumescences in any of these
specimens.
Follicles of the retracted genital setae are markedly protu-
berant into the coelom, thick and opaque, only the ental end of a
shaft visible at inner end of a follicle. Such follicles (¢ and d)
were found in x of several specimens on which annular tumes-
cences of the apertures had not been noticed prior to dissection.
Juveniles. Glandular tissues on the parietes were unrecogniz-
tuberant into the coelom, especially in the middle of the series.
Abnormality. Male funnels of x though plicate and iridescent
in one sexual worm are only about half the size of those in xi.
Each spermatheca of ix of one worm has two ampullae both of
which contain sperm.
The ampullae of the spermathecae in xii of the octothecal worm
are smaller than those in the other segments and are transparent
though other spermathecae are distended and with spermatozoal
iridescence.
Brown bodies. Ovoidal, spheroidal and discoidal bodies of
various sizes, in a posterior amputee, fill coelomic cavities of
several segments just in front of the anus. These bodies some-
times contain setae and apparently differ from the usual
‘‘brown’’ bodies only in being white.
Habitats. E. lonnbergi, in Connecticut, according to Dr. Pick-
ford (in lit.) inhabits wet places, commonly along with Hiseniella
tetraedra (Savigny) 1826 and ‘‘is not found in cultivated soils,
fields, forest soils, nor in stagnant boggy ground but rather in
banks of streams and rivers or in marshy backwaters.’’ Presence
in soil with a pH of 4.5 also was mentioned by Dr. Pickford.
1955 AMERICAN EARTHWORMS OF THE FAMILY LUMBRICIDAEI-II 5
Infe history. Reproduction presumably is biparental ordi-
narily as evidence of sperm maturation and/or reception was
recognized in all but one of the adults. The exceptional worm
(April 28) obviously had not yet copulated nor matured sperm
though the clitellum was fairly well developed (presexual clitel-
late). Marked iridescence in distended spermathecal ampullae
and on male funnels, lke the large soft seminal vesicles, in the
other two April and the June worms, just as clearly indicated
sexual maturity and copulation. Iridescence on male funnels and
in spermathecae of July specimens (Georgia) was much less
marked. Spermathecal ampullae of two of those worms were
filled with a watery fluid or pink jelly in which there was no
iridescence, and ovisacs were occupied by a brown granular
debris. The slight iridescence on male funnels and in sperma-
thecae of some of the fall worms (Oct. 10, 21, 28) and especially
the brown debris filling the seminal vesicles again indicated a
postsexual state. Two ex copula individuals (Oct. 6) and another
(Nov. 4) that had recently copulated show that reproduction may
occur in the fall also. Slight iridescence on the male funnels
showed that some sperm already had been matured in the aclitel-
late (Oct. 6) worm which had not yet copulated. Whether a
summer period of sexual inactivity separates spring and fall
breeding seasons remains to be determined.
Parasites. Nematodes were present in the coelomic cavities of
x-x1 of one worm.
Fairly large cysts, presumably of gregarines, are present in
seminal vesicles of several worms where they are quite obvious
as soon as the specimen is opened. Other cysts are present in the
parietes of several specimens, some even recognizable from the
exterior with which they seem to be in communication by a
minute aperture. Large, transparent cysts in coelomic cavities
of last few segments (several worms) have two hemispheroidal
opaque bodies each with a single nucleus. Smaller opaque cysts
are numerous in the same region.
Ovoidal bodies with an opaque center and a transparent pe-
riphery are attached each by a slender and relatively very long
stalk to the parietes of segments behind xix (one worm).
Remarks. The coloration in vivo, according to Dr. Pickford
(in lit.) is a peculiar, dark grey, irregularly interrupted by
6 BREVIORA No. 48
lighter areas and so characteristic as to enable easy recognition
of the species in the field. Pigment may have been lysed at
maturity in the dorsum of vili-xii in one worm, just as happens
in Kisena foetida (Savigny) 1826, and may have been leached
in others by the formalin. The coloration in some of the Georgia
specimens was about the same in the ventrum as in the dorsum
and was not externally distinguishable from the darkening of
unpigmented worms that takes place, in certain conditions, in
formalin.
The clitellum of one ex copula worm is markedly tumescent,
perhaps more so than on any other specimen. The tumescence
of the clitellar region of the copulatory partner is however only
very slight.
Whether glandular tissue on parietes lateral to B in elitellar
region is associated with GS follicles and tumescences or with
tubereula pubertatis is unknown.
An anal segment had been reconstituted after amputation (or
autotomy) in 16 specimens. Regeneration of a tail in this species,
as in Lumbricus terrestris L., may not take place.
In the evolution of HL. lonnbergi a pair of spermathecae opening
on 7/8 has been lost. Two pairs opening on 9/10-10/11 have
been acquired. The genetic factors responsible for addition of a
pair of spermathecae in xii of the octothecal mutant presumably
have been involved in the evolution of various lumbricid species.
II
EISENIA CAROLINENSIS Michaelsen 1910
Helodrilus (Hisenia) carolinensis Michaelsen 1910, Ann. Mus. Zool. St.
Petersburg, 15, p. 15. (Type locality, presumably Fayetteville, North
Carolina. Type, in Hamburg Mus., intercepted in a plant shipment at
Hamburg. )
Eisenia pearsei Stephenson 1933, Proc. Zool. Soc. London, 1932, p. 935, fig.
28. (Type locality, Durham, North Carolina. Types, 13, in Brit. Mus.)
Eisenia carolinensis, Cernosvitov, 1942, Proe. Zool. Soc. London, 111, (B),
p. 222. (14 specimens, 5 immature, from Durham, N. C., in Brit. Mus.)
!
1955 AMERICAN EARTHWORMS OF THE FAMILY LUMBRICIDAEI-II 17
Chapel Hill, North Carolina, April 19, 1932, 0-0-1. J. M.
Valentine per Dr. G. E. Pickford.
Highlands, North Carolina. In peaty leaf mould (pH 4.5)
among tree roots under leaves near path on way to Pri-
meval Forest, July 26, 1931, 0-0-1. In pocket of sandy
black soil (pH 5.0) under dead leaves by stream in woods
on path to Primeval Forest, July 25, 1931, 0-0-2. In leaf
mould and sandy loam (pH ca. 5.0) near path leading up
to Whiteside Mountain at altitude of ca. 4500 ft., August
21, 1932, 0-0-1. Dr. G. E. Pickford.
Linville, North Carolina. In earth and leaf mould (pH be-
tween 5.0 and 6.0) by stream in ravine, near road at 4000
ft. and 14 miles from Blowing Rock, July 31, 1931, 1-0-3.
Dr. G. E. Pickford.
Indian Gap (now Newfound Gap, in Great Smoky Moun-
tains), Tennessee. In leaf mould and loamy soil (pH be-
tween 5.0 and 6.0) among tree roots and rotting wood, at
ca. 5000 ft., July 30, 1931, 0-0-2. Dr. G. E. Pickford.
Chilhowie, Virginia. In pocket of loamy soil among grass
roots at bank of stream by bridge at ca. 3500 ft., about
1014 miles on road to Whitetop Mountain, July 21, 1931,
0-1-4. Dr. G. EK. Pickford.
(Several other Virginia worms, loaned by Prof. Harman,
also have been examined. )
Tickfaw River Valley, Louisiana, 0-7-0-0. (Mus. Comp. Zool.
No. 2061.)
External characteristics. Length, 80-99 mm. (normal worms
only). Diameter, 5-7 mm. Segments (122?) 135-142 (cf. Table).
Pigment unrecognizable in alcoholic and most formalin material.
A light red characterizes dorsum of region in front of xv in
freshly preserved material (Harman’s), and at parietal incisions
circular muscle layer looks red. Body four-sided posteriorly, the
dorsal surface slightly wider than the ventral, the sides slightly
eoncave. Prostomium epilobous (28 specimens), tongue open
and usually quite short. Setae begin on 11 (28) on which all
usually are present, one pair at each angle of the body posteri-
orly where CD < or ca. = AB, BC < AA < DD < UC.
Nephropores first recognizable on ili, usually close to or dorsal
to D on iii-vii, viii or ix, slightly lateral to B on xv-xvi (16)
and thence posteriorly often at that level but occasionally in
8 BREVIORA No. 48
DD, the location often not the same on opposite sides of the same
segment. First dorsal pore on ?5/6 (2), 5/6 (25).
Spermathecal pores on 9/10-10/11 (27), close to mD, each in a
quite small tumescence. Female pores equatorial, just lateral to
B, on xiv. Male pores equatorial, on xv, each in a transverse
slit-like depression in median portion of BC.
Clitellum red (formalin preservation) or white (one alcoholic),
saddle-shaped, reaching ventrally nearly to B, on (xxili)xxiy-
xxxi (3), (coil) xxiv-xxxi(xxxm), (1), sodli-ood (3), xxix
/2 (1), xxili-xxxii (1), xxiv-xxxi (6), xxiv-xxxii/2 (1), xxiv/2-
xxxli/2 (2), xxv-xxxi (1). Epidermis in AA of clitellar segments
thickened but not as much as in genital tumescences. Tubercula
pubertatis lateral to B, band-like but not as wide as in E. lonn-
bergi, on XXvil-xxix (25), perhaps occasionally extending slightly
onto xxx and/or xxvi.
Genital tumescences conspicuous and transversely elliptical,
always including both a and b setae, on ix (1), xi (3), xii (3),
xvi (2). xvii (21), xvi (5), xx (7), xxx (16), xoqi (C1) Socal)
ExIV (3), XXV-xxk (27) xxx 4); ocx (2!) | ocx (ace eon
xxiv (1). Each may have a greyish translucent central area.
Posterior tumescences are not delimited laterally from the clitel-
lum. Male pore tumescences restricted to xv (7), extending
onto xvi but not onto xiv or only very slightly (5), extending
well into xiv and xvi (7).
Internal anatomy. Septa 12/138-14/15 muscular and increas-
ingly thickened posteriorly. Longitudinal muscle band at mD
and peritoneum apparently without pigment.
Oesophagus with no sacs in x. Calciferous glands as in EL.
lonnbergi Michaelsen 1894, large, moniliform, in xi-xi, sepa-
rated by a deep constriction, the posterior gland often not reach-
ing insertion of 12/13. Gizzard in xvii-xviil, muscularity occa-
sionally extending slightly beyond insertion of 16/17 or 18/19.
Intestinal origin in xv (18). Typhlosole begins in region of xxii-
xxvil, abruptly or rather gradually as a flat band that shortly
becomes vertical, reaching a height of 144-2 mm. and to or nearly
to floor of gut, thickly lamelliform, the interior with only a few
delicate fibres and some granular debris. The highest anterior
portion, as in lénnbergi, may have a deep groove on the ventral
face. The end normally is in region of xevii-ev (cf. Table IT).
Subneural trunk quite small, adherent to nerve cord, some-
1955 AMERICAN EARTHWORMS OF THE FAMILY LUMBRICIDAEI-II 9
times quite unrecognizable. Last hearts in xi (18), the hearts
of x-x1 usually slenderer than the ones in ix, none found in vy.
Nephridial ducts pass into parietes in the b gap.
Seminal vesicles three pairs, in ix, xi-xii (18, ineluding the
juvenile), those of xi sometimes (5) smaller to much smaller
than the others. Male deferent ducts apparently shghtly thick-
ened in the entalmost portion which is in 2-4 very short loops
crowded close together on the posterior face of the septum be-
hind the funnel, recognizable (as in lénnbergi) on parietes into
xv. Spermathecae with short ducts not confined to parietes,
ampullae in ix-x (18, including the juvenile).
Follicles of the genital setae (those in copulatory tumescences)
enlarged, obviously protuberant into coelom but not as long (nor
as muscular?) as in lonnbergi. Glands, atrial and such as are
present on the parietes in lonnbergi and species of Bimastos are
lacking.
Abnormality and Variation. Spermathecal ducts of the Chapel
Hill worm are confined to the parietes and each ampulla is deeply
constricted by the septum, the portion anterior to the septum of
quite different appearance from that behind. Seminal vesicles of
ix are lacking. Genital tumescences are present 0N XXXi-xXXX1l1 as
well as on 1x and in addition were developed around c-d of xii.
Genital tumescences in front of xv and behind xxx otherwise
were only on worms from Linville and Chilhowie.
Regeneration. Many of the worms clearly are posterior ampu-
tees (cf. Table Il) and two others may be. No regeneration,
except reconstitution of an anal segment, had taken place al-
though the amputation in some cases probably had not been
recent:
life history. Reproduction presumably is biparental as sperm
had been matured and/or received in each adult. Sperma-
tophores (at 25/26 in BC, on xxviii at mV) and brilliant iri-
descence in spermathecal ampullae presumably indicated recent
eopulation by a July 31 worm (Linville) in which no iridescence
was recognizable on male funnels. Funnel iridescence was bril-
hant on the April (Chapel Hill) and four July 21 (Chilhowie)
specimens. Male deferent ducts were iridescent in the April
worm which is the only one with maximal clitellar tumescence.
The single aclitellate individual (July 21, Chilhowie) with dis-
10 BREVIORA No. 48
coloration in dorsum of elitellar segments, slight iridescence only
at periphery of male funnels and empty spermathecal ampullae
clearly is postsexual. Spermathecal ampullae were empty in two
worms (Linville) and iridescence was lacking in opaque matter
in ampullae of another as well as in transparent watery content
of two, in all of which funnel iridescence was not brilliant though
still unrestricted. Ovisacs of one of the worms were filled with
a brown granular debris. The peak of reproductive activity in
these worms presumably was passed at time of preservation. The
breeding season presumably extends from April well into July.
Parasites. Nematodes were present in the coelomic cavities of
x-xi of one worm. Large and medium-sized cysts, opaque or
translucent, probably of gregarines, were numerous in coelomic
cavities of the last ten to fifteen segments in several of the pos-
terior amputees.
Remarks. The red coloration of the clitellum appears to be a
formalin induced artifact as it was lacking in the single alcoholic
specimen.
Nephropores of one worm were lateral to B on the right side
of xxxiv-xxx1x and in DD on the left side of xxxy-xxxix.
The spermathecal battery of the ancestral lumbricid presum-
ably comprised two pairs opening to the exterior on 7/8-8/9. In
the evolution of carolinensis two pairs, opening on 9/10-10/11
have been acquired, and all four of the original ones have been
lost. As this is two more than have been eliminated in the an-
cestry of lonnbergi, in that respect carolinensis is more advanced.
Seminal vesicles of primitive lumbricids presumably were in
four segments, ix-xii. Those of x frequently have been lost in
the family but in Hisenia they still have not disappeared in some
‘‘formae’’ of a species. Presence of vesicles in x of one (or
more?) of the types of pearse?, is about all that remains to dis-
tinguish it from carolinensis, and scarcely warrants specific
status, if, indeed, any taxonomic recognition.
E. carolinensis and E. lénnbergi are the only lumbricids
endemic in America that have retained spermathecae. No evi-
dence was found in available material of either species that would
indicate elimination of those organs is under way.
1955 AMERICAN EARTHWORMS OF THE FAMILY LUMBRICIDAEI-II 11
Table I
Typhlosole termination and segment number
in Hisenia lonnbergi
Typhlosole Atyphlo-
ends in solate Number of
segment segments segments Locality Remarks
60-61* a 62 Georgia Amputee
61-64* 0 64 Georgia Amputee
72 4 76 Virginia Amputee
74 4 78 Georgia Amputee
74 7 81 Amputee
78 10 88 Amputee Juvenile
81 6 87 Georgia Amputee
82 7 89 Amputee
84 3 87 Amputee
84 8 92 Amputee
86 13 99 Amputee? Juvenile
100 3 103 Amputee
100 16 116
101 13 114
103 9 107+ Broken
104 5 109 Georgia Amputee
. 105 8 113 Georgia Amputee
100-106* 10 116 Amputee
107 5 112 Georgia Amputee
108 13 121 Ex copula
q q 124 Damaged
110 14 124 :
110 alt 127
111 17 128
114 18 132 Ex copula
114 3 117 Amputee
115 19 134
* Typhlosole ends very gradually rather than abruptly as in other worms.
Posterior ends had been broken off from worms not included above (not
available), except for one juvenile, presumably of this species, which
has 100 segments.
12 BREVIORA No. 48
Table II
Typhlosole termination and segment number
in Misenia carolinensis
Typhlosole Atyphlo-
ends in solate Number of
segment segments segments Locality Remarks
65* 6 71 Amputee
73 23 96 Juvenile. Amputee?
79 21 100 Amputee
80 19 101 Amputee
88 26 114 Amputee
82-89T Hl: 100 Chapel Hill Amputee
90 32 122 Amputee
93 7 100 Amputee
97 25 122 Indian Gap Amputee?
98 44 142 Linville
100 37 137 Highlands
103 32 135 Chilhowie
105 24 129 Chilhowie Amputee?
* Unusually low in seven preceding segments. |
} Ends gradually rather than abruptly as in other worms.
Posterior ends had been broken off from the specimens not listed above
at time of collection and anal pieces were unavailable.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. NOVEMBER 18, 1955 NUMBER 49
THREE NEW SHARK RECORDS FROM
THE GULF OF MEXICO
By Henry B. Bicktow, W. C. SCHROEDER
and STEWART SPRINGER* —
The trawlings recently carried out by ‘‘Oregon’’ of the U. S.
Fish and Wildlife Service in the Gulf of Mexico have yielded two
specimens of Himopterus pusillus (Lowe) 1839, one of Centro-
phorus granulosus (Bloch and Schneider) 1801, and one of
Dalatias licha (Bonnaterre) 1788, sharks that had long been
known in the eastern side of the Atlantic, but which had not
been reported previously from the Gulf, though one specimen
of Dalatias has been taken on the northern edge of Georges Bank.
As the Gulf of Mexico specimens of F#. pusillus and of C.
granulosus are the first of their kinds to be reported from the
western side of the Atlantic or tributary waters, brief accounts
follow to emphasize such of their diagnostic characters as may
not be immediately apparent from the accompanying illustra-
tions (Figs. 1, 2), and to justify our identifications of them.
Genus H}1TMOPTERUS Rafinesque 1810
Type species Squalus spinax, Linnaeus 1758
Whitley (1939, p. 266) has recently revived the genus Acan-
thidium Lowe 1839, type species Centrina mgra Lowe 1834
which, in 1839, Lowe renamed Acanthidium pusillum, thinking
it separable generically from Hitmopterus Rafinesque 1810, type
species Squalus spinax Linnaeus 1758. But the differences in
fin-characters cited by Whitley do not seem to us sufficient to
justify this separation. And the specific name niger is preoccu-
1 Contribution No. 787. Woods Hole Oceanographic Institution.
2 BREVIORA No. 49
pied in Htmopterus by mgrum (Cloquet) 1820,* p. 93 and various
subsequent authors, proposed as a substitute for spinax Lin-
naeus 1758.
ETMOPTERUS PUSILLUS (Lowe) 1839
Study material. Male, 445 mm. long, U. S. Nat. Mus. No.
157835, and female of 458 mm., Mus. Comp. Zool. No. 39572,
northern part of Gulf of Mexico, ‘‘Oregon’’ Station 1281, Lat.
29°13’ N., Long. 87°54’ W., in 250 fathoms. Also female 278 mm.
long, Madeira (Mus. Comp. Zool. No. 1026), and female of 167
mm., off Equatorial West Africa, Lat. 6°08’ S., Long. 11°24’ E.,
Mus. Comp. Zool. No. 38002, received through the kindness of
Dr. Max Poll.
The Gulf of Mexico specimens agree so closely with those with
which we have compared them, from Madeira and from tropical
West Africa that we have no hesitation in referring them to the
same species.
The morphological feature the most sharply diagnostic for
pusillus, among its genus-mates of the Equatorial and North At-
lantic, of the Mediterranean and of the Gulf of Mexico, is that
the denticles on the sides of its trunk are low, truncate, the great
majority with concave crowns, but an occasional denticle flat, or
perhaps even weakly convex, on a conspicuously 4-radiate base.”
In these respects they contrast with the slender, bristle-like den-
ticles of E. spinax (Linnaeus) 1758 and of E. schultz (Bigelow,
Schroeder and Springer 1953, fig. 9G), with the stouter, more
thorn-like denticles of EH. hillianus Poey 1861 (Bigelow and
Schroeder 1948, p. 489, fig. 92B), of E. polli (Bigelow, Schroeder
and Springer, 1953, fig. 7 C) and of EF. princeps (see Bigelow,
Schroeder and Springer, 1953, figs. 8 E, 8 F), and with the more
conical claw-like denticles of E. virens (Bigelow, Schroeder and
Springer 1953, fig. 10 D, E).
The color, also, of E. pusillus is distinctive. It recalls HE. prin-
ceps, Collett 1904, of higher latitudes in both sides of the At-
lantic, and E. schultzi Bigelow, Schroeder and Springer 1953,
of the Gulf of Mexico, in the uniformly dark slaty to black-
1Garman (1913, p. 223) eredits the earliest ee of niger in this connection tu
Gunner 1763, but we have not been in a position to verify this citation.
2 Our earlier characterization of them as a whole (Bigelow, Schroeder and
Springer 1953, p. 240) as “flat or weakly convex” was an evident misstatement.
THREE NEW SHARK RECORDS FROM THE @QULF OF MEXICO 3
1955
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4 BREVIORA No. 49
ish hue of its trunk not only below, but above as well (except
for the pale intraocular spot mentioned below), and in the lack
of definitely outlined paler and darker areas on its sides. But
the outer part of its pectoral fins, with the rear part of its two
dorsal fins are white and more translucent, which is not the case
elther in princeps, in schultzi, or in any other Etmopterus for
that matter, that is known yet from the North Atlantic, from the
Mediterranean, or from the Gulf of Mexico. This very conspicu-
ous feature of pusillus is shown on Lowe’s (1848, Pl. 6) original
illustration of it (as Acanthidium pusillwm) ; also on the colored
illustration by Braganza (1904, Pl. 2, fig. 2, as H. pusillus).
Other features that in combination are diagnostic for pusillus
are: that the upper margin of its caudal fin is only about as long
as from the tip of the snout to the level of the second pair of gill
openings; that the distance from the rear end of the bases of its
pelvic fins to the origin of the lower side of its caudal fin is shorter
than from the tip of the snout to the level of the first pair of gill
openings; that the rear end of the base of its first dorsal fin is
much nearer to a perpendicular at the axils of the pectoral fins
than to a perpendicular at the origin of the pelvic fins; that the
margins of its pectoral fins are not deeply fringed normally as
they are in E. schultzi (Bigelow, Schroeder and Springer 1953,
fig. 9 D) ; that the anterior edge of the first pair of gill openings
is not concave enough to expose the tips of the gill-folds as it is
in E. princeps (see Bigelow, Schroeder and Springer 1953, fig.
8 D) ; and that the upper surface of the head is marked between
the eyes with a pale yellowish spot, as it is in E. polls, in E.
schultzi, in E. virens, and in E. spinax, (where there is also a
white spot ‘‘above the hind part of each orbit,’’ Garman 1913,
p. 224), but not in EF. princeps.
Two species of Etmopterus, schultzi and virens, Bigelow,
Schroeder and Springer 1953, had been known previously in the
Gulf. The discovery of pusillus there now increases the local list
to three. And a fourth member of the genus (hillianus Poey
1861) is to be expected there for it has been found widespread
in Cuban waters and in the West Indian region, and has been
recorded from as far to the northward as the offing of Chesa-
peake Bay.
E. pusillus, originally reported from Madeira, has since been
1955 THREE NEW SHARK RECORDS FROM THE GULF OF MEXICO 5
recorded from various localities in the eastern side of the Atlantic
between Equatorial West Africa (Lat. 6°08’S, see above) and
the coast of Portugal, the Cape Verde Islands, the Canaries and
the Azores. Earlier reports of it from the West Indian region,
listed elsewhere (Bigelow and Schroeder 1948, p. 493), seem
actually to have been based on F. hillianus.
The discovery that pusillus occurs in the Gulf of Mexico shows
that its range parallels that of Centrophorus granulosus (p. 9),
also that of C. uyato Rafinesque 1810, which had been known only
from the Mediterranean and from the eastern side of the Atlantic
in low and mid-latitudes until 1951-1952, when ‘‘Oregon’’
trawled 2 specimens of it in the Gulf (Bigelow, Schroeder and
Springer, 1953, p. 227).
Tanaka (1912, pl. 22; p. 88) also, has referred to pusillus the
Japanese shark that was earlier described and pictured by Piet-
schmann (1907, p. 395; 1908, p. 654, pl. 1, fig. 2, pl. 2, fig. 2)
as E. frontimaculatus. But while the latter agrees with pusillus in
the nature of its dermal denticles (Pietschmann, 1908, p. 657, text
figs. 1, 2), it differs from pusillus in a more rearward position of
the first dorsal fin (relative to the positions of the pectoral and
pelvic fins), in a relatively longer interspace between the pelvic
and caudal fins, and in color pattern, with the lower surface of its
head, its belly, and a definitely outlined area above and anterior
to each pelvic fin much darker than the upper part of its sides.
A further difference is that the lower surface is as densely
clothed with denticles as are the sides and back in pusillus
whereas it is largely naked in the Japanese species.
Genus CENTROPHORUS Miller and Henle 1837
Type species Squalus granulosus Bloch and Schneider 1801
CENTROPHORUS GRANULOSUS (Bloch and Schneider) 1801
Study material. Female 922 mm. long from northern part of
Gulf of Mexico, Lat. 29°15’N., Long. 88°18’W., ‘‘Oregon’’ Sta.
1246, 200-210 fathoms, U. 8. Nat. Mus. No. 157833; also skin of
a male, 855 mm. long, eastern Atlantic, exact locality not known,
Mus. Comp. Zool. No. 662.
NO. 49
BREVIORA
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1955 THREE NEW SHARK RECORDS FROM THE GULF OF MEXICO ic
The most evident diagnostic features that set apart C. granu-
losus (with C. machiquensis Maul 1955) from the other two
species of its genus (squamosus Bonnaterre 1788 and wuyato
Rafinesque 1810) that are known from the North Atlantie prov-
ince are: that the inner corner of its pectoral fins is greatly
extended and narrowly pointed (Fig. 2); that the interspace
between its two dorsal fins is longer than the head (to origin
of pectoral fins) by a distance nearly as long as the eye; that
the denticles on the sides of the body are low, block-like with
the ridges on the outer surface converging rearward, closely
spaced in quincuncial arrangement, but not overlapping (Fig.
2, D) ; and that the upper teeth are erect and nearly symmetrical
all along the central part of the jaw (Fig. 2, B). In granulosus
also, as in squamosus, the cutting edge of the lower teeth is par-
tially serrate, but so finely so that this feature is visible only on
eareful examination under a lens (Fig. 2, C). In wyato, on the
contrary, the lower teeth are smooth-edged, like the uppers.
Proportional dimensions, in per cent of total length, of female,
922 mm. long, northern part of Gulf of Mexico, U. S. Nat. Mus.
No. 157833.
Trunk at origin of pectoral. Breadth 9.7, height 10.8.
Snout length in front of. Outer nostrils 2.8; mouth 9.1; eye 4.9
Eye. Horizontal diameter 4.9.
Mouth. Breadth 7.9.
Nostrils. Distance between inner ends 3.6.
Spiracles. Distance between inner ends 7.0.
Labial furrows, upper. Length 5.4; distance between inner ends
6.4.
Gill openings, lengths. 1st 2.6; 2nd 2.7; 3rd 2.8; 4th 3.0; 5th 3.3.
First dorsal fin. Vertical height 5.4; length of base 11.1; base
rearward from anterior beginning of spine 8.5; diagonal from
emergence of spine to rear end of base 7.1; free rear margin
of fin 6.9.
Second dorsal fin. Vertical height 5.0; length of base 7.0; base
rearward from anterior beginning of spine 5.7; diagonal from
emergence of spine to rear end of base 5.4; free rear margin
of fin 4.5.
Caudal fin. Upper margin 20.3; lower anterior margin 12.3.
Pectoral fin. Outer margin 12.9; inner margin 13.2; greatest
width 7.6.
8 BREVIORA no. 49
Distance from snout to. 1st gill opening 16.8; to origin of Ist
dorsal spine 34.8; to origin of 2nd dorsal spine 69.2; upper
caudal 79.7; pectoral 22.2; pelvics 58.7.
Interspace between. Anterior beginning Ist dorsal spine to 2nd
dorsal spine 34.4; rear tip of 1st dorsal fin to rear tip of 2nd
dorsal 29.5; 2nd dorsal and caudal 6.0; pelvics and caudal 13.4.
Distance from origin to origin of. Pectoral and pelvics 37.6;
pelvies and caudal 18.1.
Head, to origin of pectoral fins, about 28 per cent of trunk to
origin of upper side of caudal fin; snout moderately rounded, its
length in front of snout about 1% of head to origin of pectorals;
eye about 22 per cent as long as head; distance from level of
front of eyes to tip of snout about as long as eye; spiracle about
14 as long as eye, its anterior edge only slightly posterior to rear
corner of eye. Nostrils a little less than 14 as long as distance
between nostrils, approximately transverse, the outer ends pos-
terior to level of tip of snout by a distance a little shorter than
distance between nostrils. Anterior nasal flap short, narrowly
triangular. Distance between nostrils a little less than 1% as
great as from tip of snout to mouth. Mouth very low-arched, its
gape when closed occupying about 4+ of breadth of head at level
of outer corners of mouth. Distance between inner ends of nasal
furrows about 1.8 times as long as between nostrils. Longest gill
opening (5th) is almost as long as distance between nostrils.
Teeth Tas. of shapes illustrated in Figure 2, B; uppers
with the post-functional (outermost) row partly lost, followed by
a nearly vertical functional row and then by an oblique backward
pointing row that will be next in service; lowers with two
rows visible from outside the mouth, one of which is fune-
tional, 16 teeth pointing to the right hand side, of which one is
a median tooth, and 15 teeth pointing to the left hand side;
cutting edge of some of the lowers with microscopically fine and
somewhat irregular serrations (Fig. 2, C) but others merely
somewhat irregular, a difference probably due to wear. Dermal
denticles on sides of body sessile, block-like, sub-quadrate, with
the more acute corner directed rearward, closely spaced in quin-
cuncial pattern but not overlapping, the crown weakly convex
or flat, its anterior 14 to 24 with (usually) 5-7 low ridges, con-
verging rearward (Fig. 2, D), the outermost pair the longest.
1955 THREE NEW SHARK RECORDS FROM THE GULF OF MEXICO )
Denticles at mid-level of side below first dorsal fin averaging
about 0.9 mm. long.
Length of base of first dorsal fin, measured from point of
emergence from skin of anterior side of first dorsal spine, about
1.3 times as long as base of second dorsal fin, similarly measured.
Interspace between rear end of base of first dorsal and point
of emergence of second dorsal spine longer than head (to origin
of pectorals) by a distance about equal to length of eye and
about 71% times as long as between nostrils. Exposed portion of
first dorsal spine about 11% times as long as that of second spine,
the second spine reaching about mid-way along the free anterior
margin of the fin. Interspace between rear end of base of second
dorsal fin and origin of upper side of caudal fin about as long
as base of second dorsal fin measured from point of emergence,
from skin, of anterior edge of second dorsal spine.
Upper edge of caudal fin about as long as from tip of snout to
level of fourth pair of gill openings, its lower edge with obtuse
subterminal notch and rounded lower anterior lobe. Interspace
between origin of lower edge of caudal fin and rear ends of bases
of pelvic fins about %5 as long as from tip of snout to level of
dth pair of gill openings. Origin of pelvic fins (in female)
anterior to a perpendicular at emergence of anterior edge of
second dorsal spine, by an amount about equal to interspace
between origin of lower edge of caudal fin and rear ends of bases
of pelvic fins. Pectorals with inner corner greatly extended and
narrowly pointed, reaching rearward (when laid back) nearly to
a perpendicular from rear base of first dorsal fin.
Sharks referable with certainty to granulosus, not to wyato
which Rey (1928, p. 486) considered synonymous with it, had
previously been recorded in the Mediterranean, off the coast of
Portugal, and at Madeira (specimen in British Museum; Giinther
1870, p. 421). The capture of a typical granulosus in the Gulf
now shows that its Atlantic range parallels that of C. uyato
Rafinesque 1810, which was recently found to oceur in the Gulf
(Bigelow, Schroeder and Springer 1953, p. 227). The third
species of Centrophorus that is known from the Atlantic (squam-
osus Bonnaterre 1788) has so far been reported only off the
coast of Portugal and northward to southwestern Iceland and
the vicinity of the Faroes.
10 BREVIORA No. 49
Pietschmann (1908, pp. 663-667) credits granulosus to Japan
also, from his comparison of two Japanese specimens with gran-
ulosus from the Mediterranean. But the eventual decision,
whether the northwestern Pacifie Centrophorus of this general
character is indeed indistinguishable from the Atlantic-Mediter-
ranean granulosus, is best postponed until a larger number of
specimens have been examined, critically, with this question in
mind.
Genus DALATIAS Rafinesque 1810
DALATIAS LICHA (Bonnaterre) 1788
Study material. Female 845 mm. long from northern part of
Gulf of Mexico, Lat. 28°25’N., Long. 86°02’W., ‘‘Oregon’’ Sta.
1275, 225 fathoms, U.S. Nat. Mus. No. 157834.
The specific identification of this specimen as D. licha is so
evident, from its close agreement in bodily form, fin characters,
teeth, and denticles with the Georges Bank specimen described
and pictured previously (Bigelow and Schroeder 1948, p. 502,
figs. 96, 97) that no further account seems called for here.
As the only previous record for this shark in the western
Atlantie was this Georges Bank example, the capture of one in
the northern part of the Gulf of Mexico widely expands its known
range.
In the eastern side of the mid and north Atlantic the known
range of D. licha extends from Equatorial West Africa (Rio de
Oro) to the Irish Atlantic slope, including the Mediterranean.
And it is doubtful whether the representatives of the genus that
have been reported from South Africa, from the New Zealand-
Australian region and from Japan can be separated specifically
from D. licha of the Atlantic. (For discussion, see Bigelow and
Schroeder 1948, p. 501).
REFERENCES
BIGELOW, HENRY B., and W. C. SCHROEDER
1948. Fishes of the Western North Atlantic. No. 1, Mem. Sears Foun-
dation for Marine Research, Part 1, pp. 59-576, text figs. 6-105.
1955 THREE NEW SHARK RECORDS FROM THE GULF OF MEXICO 11]
BIGELOW, HENRY B., W. C. SCHROEDER, and STEWART SPRINGER
1953. New and little known sharks from the Atlantic and from the
Gulf of Mexico. Bull. Mus. Comp. Zool., vol. 109, no. 3, pp. 213-
276, text figs. 1-10.
Boon, M. E., and I. G. SCHNEIDER
1801. Systema Ichthyologiae iconibus ex illustratum. lx + 584 pp.;
110 pls.; 2 vols.
BONNATERRE, P. J.
1788. Ichthyologie in:Tab. Encyc. Méthod. Trois régnes de la Nature.
lvi + 215 pp., pls. A, B + 1-100. Paris.
BRAGANZA, CARLOS DE
1904. Esqualos obtidos nas Costas de Portugal. Resultados das Invest.
Sci. ... Yacht ‘‘Amelia.’’ Ichthyologia. II; 107 pp., 2 pls.
Lisbon.
CLOQUET, HIPPOLYTE
1820. Le sagre, spinax niger. Dict. Sci. Nat., vol. 1, suppl., p. 93.
CoLLeEtTt, R.
1904. Diagnoses of four hitherto undescribed fishes from the depths
south of the Faroe Islands. Videns. Selskabs Forhand. Christi-
ania, 1904, no. 9, 7 pp.
GARMAN, SAMUEL
1913. The Plagiostomia. Mem. Mus. Comp. Zool., vol. 36, xiii + 515
Dp. 47 pls.
GUNTHER, ALBERT
1870. Catalogue of fishes of the British Museum, vol. 8, xxv + 549 pp.
JORDAN, D.S., and B. W. EVERMANN
1896. The fishes of North and Middle America. Bull. U. S. Nat. Mus.
No. 47, Part 1, lx + 1240 pp.
LINNAEUS, C.
1758. Systema naturae, 10th Ed., vol. 1, 824 pp., Holmiae.
LOWE, R. T.
1834. A collection of fishes made in Madeira. Proc. Zool. Soe. London
(1833), Part 1, 1834, pp. 142-144.
1839. A supplement to a synopsis of the fishes of Madeira. Proc. Zool.
Soc. London, Part 7, 1839, pp. 76-92.
1843. <A history of the fishes of Madeira ...196 pp., 27 pls., London.
MAUL, G. E.
1955. Five species of rare sharks new for Madeira. ... Notul. Natur,
Acad. Nat. Sei. Philadelphia, No. 279, 13 pp., 3 pls.
MULLER, JOHANNES, and F. G. J. HENLE
1837. Ueber die Gattungen der Haifische und Rochen ... Arch. Natur-
gesch., Jahrg. 3, vol. 2, pp. 394-401, 434.
12 BREVIORA No. 49
PIETSCHMANN, VIKTOR
1907. Zwei neue Selachier aus Japan. Anz. Akad. Wiss. Wien. Vol. 44,
pp. 394-396.
1908. Japanische Plagiostomen. Sitzber. Akad. Wiss. Wien, math. Nat.
KIS vol Liv; pt. 1 pp. 637-7405 plss daa:
Pory, FELIPE
1858-1861. Memorias sobre la historia natural de la isla de Cuba, vol.
2, 442 pp., 19 pls.
RAFINESQUE, CONSTANTINE SAMUEL
1810. Carratteri di alcuni nuovi generi e nuovi specie di animali e
piante della Sicilia. pp. iv, 105, 20 pls. Palermo.
Rey, Luis Lozano
1928. Fauna Iberica. Peces. Vol. 1, 690 pp., 20 pls. Inst. Nac.
Ciencias, Madrid.
TANAKA, SHIGEHO
1912. Figures and descriptions of the fishes of Japan. Vol. 5, pp. 71-
86 + [1], pls. 21-25; vol. 6, pp. 87-108, pls. 26-30.
WHITLEY, GILBERT
1939. Studies in ichthyology no. 12. Ree. Australian Mus. Vol. 20, No.
4, pp. 264-277.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. NOVEMBER 30, 1955 NUMBER 50
NEW FROGS OF THE GENERA
ASTEROPHRYS AND OREOPHRYNE FROM
NEW GUINEA
By ArrHur Loveripcr
During the past year the Museum of Comparative Zoology has
received from Mr. Kenneth R. Slater some interesting microhylids
including Asterophrys rufescens (Macleay), valvifera (Barbour),
turpicola (Muller) ; Nenobatrachus rostratus (Méhely) ; Meto-
postira ocellata Méhely and Cophixalus oxyrhinus (Boulenger),
taken at Omati, near Port Moresby, Papua.
There was also a Jarge and handsome frog which keyed down
to Asterophrys boettgert (Méhely) in the synopsis to that genus
furnished by H. W. Parker in his ‘‘Monograph of the Mierohy-
lidae’’ (1934:60. Brit. Mus.). However, A. boettgeri comes from
Halmahera Island in the Moluceas lying to the northwest of Dutch
New Guinea, while Omati is situated in the southeast of New
Guinea. After Dr. E. E. Williams and I had re-examined the
Omati specimen and come to the conclusion that it was probably
new, it was submitted to Dr. Parker for favor of his confirmation.
Meanwhile, in the hope that the collector might be able to secure
more material, publication has been deferred for nearly a year.
Mr. Kenneth Slater, after whom I take pleasure in naming it,
now informs me that there is little hope of his obtaining a series
for he will soon be returning to Australia.
ASTEROPHRYS SLATERI sp. nov.
Holotype. Museum of Comparative Zoology No. 28205, an adult
2, from Omati, near Port Moresby, Papua. Collected by Kenneth
R. Slater between January and April, 1954.
Diagnosis. Apparently closely related to A. boettgeri (Méhely )
from which it differs in having a much longer (not shorter) snout;
2, BREVIORA NO. 50
an interorbital space that is one and one-third times as broad as
(instead of equal to) an upper eyelid; tibio-tarsal articulation
of adpressed hind limb reaching eye (instead of nostril) ; some
details of coloration; and size, being 52 mm. (instead of 39 mm.;
a difference that may be sexual). Possibly it is only racially dis-
tinct.
Description. Anterior palatal ridge curved, smooth. Snout
subacuminate, longer than the horizontal diameter of the eye,
which is equal to the distance from nostril to orbit; canthus ros-
tralis rounded ; interorbital space one and a third times as broad
as an upper eyelid ; tympanum moderately distinct, its horizontal
diameter about half that of the eye; fingers long, the first much
shorter than the second, which is slightly shorter than the fourth,
the third being much the longest, all terminating in large sub-
triangular disks which are larger than those of the toes; both
fingers and toes free of web; inner metatarsal tubercle indistinct ;
tibio-tarsal articulation of the adpressed hind limb reaches the
eye.
Skin of head, except on occiput, pustular, especially noticeable
on upper eyelids; some glandular folds on the occiput, scapular
region and dorsum, the latter also displaying a few scattered
tubercles; forelimb, more particularly the upper arm, studded
with tubereles. Below, smooth.
Color. Above, purplish brown spotted with black and flecked
with white (on body tubercles) ; on the occiput and back are four
or five, more or less W-shaped, white lines corresponding to the
elandular folds; posteriorly an obsolete, hair-like, hght vertebral
line is barely distinguishable ; thighs heavily blotched with black,
the elongate blotches separated by irregular white lines; on the
tibia these light lines suggest obsolete crossbars.
Below, pale brownish spotted with darker brown and variegated
with indistinct white vermiculations; these coalesce to form
indistinct crossbars on the underside of the forelimbs, larger,
broader, and pure white on the hind limbs; lower jaw with two
downward-pointing, white, triangular spots on either side of the
symphysis, while a less conspicuous pair is situated near the
angle of the jaw.
Size. Holotype 2. Length of head (from end of snout to back
of tympanum), 18 mm.; length from snout to anus, 52 mm.;
length of hind limb (from anus to tip of longest toe), 80 mm.
1955 NEW FROGS FROM NEW GUINEA 33
Diet. The stomach and intestinal contents, examined by my
colleague Dr. W. L. Brown, were so finely masticated as to be
undeterminable except for some fragments of small shells, the leg
of an ant, and the chitinous plates of some arthropod.
Parasites. The stomach wall and other organs carried numerous
encysted nematodes ; one threadworm (preserved ) was free in the
body cavity.
OREOPHRYNE PARKERI sp. nov.
Cophixalus geislerorum Loveridge (not of Boettger), 1948, Bull. Mus. Comp.
Zool.; 101: 423.
Holotype. Museum of Comparative Zoology No. 12964, an adult
2, taken from running water in sago-palm forest at Matapan,
Australian New Guinea. Collected by E. A. Briggs about Jan-
uary, 1923.
Paratype. British Museum No. 1955.1.1.17, with same data as
type.
Diagnosis. In Parker’s synopsis to the genus (1934:160) these
frogs key down to the anthonyi-birot section but are not very
closely related to either species. Oreophryne anthony? (Boulen-
ger), of which we have a cotype, 1s a more robust frog than the
eravid holotype of parkeri, from which it differs shightly in almost
every character, the most conspicuous being the coloring of the
underside and:
Eye separated from tympanum by a distance equal to 114 times the horizontal
diameter of the latter. Total length of adult 45 mm.. . . . anthonyi
Eye separated from tympanum by a distance equal to only 144 the horizontal
diameter of the latter. Total length of gravid 9 30mm... . . . parkeri
O. parkeri agrees with biroi, of which we have a series, in size
and many other respects, but differs sharply in tympanic char-
acters.
Tympanum scarcely distinct, very small, 4 to 4% the eye diameter; disk of
third digit nearly % the eye diameter; ratio of tibia length to head width
at commissure of mouth 1.1 to 1.3 (4 ex.) . . . dbiroi
Tympanum very distinct, large, 24 the eye diameter ; disk of third digit nearly
2% the eye diameter; ratio of tibia length to head width at commissure of
mouth 1.37 to 1.43 (2 Ox Ry Gat. soa, <euee te ee ea Danke
The head of parkeri, as Dr. R. G. Zweifel has pointed out to
me, is relatively small compared with those of other Oreophryne.
4 BREVIORA No. 50
Description. Palate with a row of three transversely elongate
tubercles followed by a denticulated dermal ridge in front of
pharynx. Snout truncate, subequal to the horizontal diameter of
the eye, which is about equal to seven-eighths its distance from
the nostril; canthus rostralis rounded; loreal region slightly
oblique, distinctly concave; interorbital space more than one and
a half times as broad as an upper eyelid ; tympanum very distinct,
almost two-thirds the diameter of the eye to which it is very close ;
fingers long, the first much shorter than the second, which is sub-
equal to the fourth, the third being much the longest, its disk
almost two-thirds the diameter of the eye, all fingers terminating
in large subtriangular disks which are somewhat larger than those
of the toes; toes webbed at the base, the third slightly shorter
than the fifth, inner metatarsal tubercle very indistinct, outer
absent ; tibio-tarsal articulation of the adpressed hind limb reaches
the shoulder.
Skin of head smooth, except for a slightly raised median line
and the suggestion of a supratympanic fold; between the shoul-
ders a )(-shaped glandular fold. Belly smooth (? preservation).
Yolor. Above, brown, uniform except for some dark crossbars
on the limbs and light areas in groin. Below, brown, paling pos-
teriorly towards groin; thighs flecked with white, tibiae largely
white.
Size. Holotype @. Length of head (from end of snout to back
of tympanum), 7 mm.; length from snout to anus (which was 28
mm. prior to dissection), now 30 mm.; length of hind limb (from
anus to tip of longest toe), 37 mm.
Diet. The stomach of the holotype contained the remains of
scores of a shiny black Iridomyrmex ant (smaller than either
batesi or scrutator), a group that is known to forage in trees;
also a single head of one of the arboreal Podomyrma ants.
Remarks. In 1948, when reporting on the ‘‘ New Guinean Rep-
tiles and Amphibians in the Museum of Comparative Zoology’’
(Bull. Mus. Comp. Zool., 101: 303-480), I erroneously referred
these two frogs to Cophixalus geislerorum Boettger. More recent-
ly, when Dr. R. G. Zweifel of the American Museum of Natural
History was working over our material, he redetermined them as
Oreophryne of no known species. I then submitted them to Dr.
H. W. Parker who replied that a clavicle is present though not
reaching the scapula, so that they are unquestionably Oreophryne,
1955 NEW FROGS FROM NEW GUINEA 5
also eleutherognathine.
I am also indebted to Dr. Parker, after whom this new species
is named, for supplying me with the ratios of tibial length to head
width at commissure which, in the cotypes of O. anthonyi is 1.14
to 1.38, with an average of 1.25. Parker also observes that, ac-
cording to Boettger, Nieden and Vogt, C. geislerorum is without
a tympanum; the contrary view was stated by van Kampen and
so got included in the monograph.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. January 27, 1956 Numpser 51
A SMALL MUSTELID FROM THE THOMAS
FARM MIOCENE
By STANLEY J. OLSEN
INTRODUCTION
For the past ten years the Museum of Comparative Zoology
has conducted field work at the Thomas Farm (Florida) for at
least one season each year. During the latter part of this
decade an intensive search has been made with the view of
increasing our knowledge of the microfauna of this unique
Miocene deposit. Although minute vertebrate fossils have been
collected from every part of the excavation, by far the most
productive area has been the boulder bar described by Dr. T. E.
White in 1942. The material— mammal, bird and reptile —
occurs here in the matrix between the boulders of Ocala lime-
stone comprising the bar, and the latter have served to prevent
the crushing and breakage elsewhere characteristic of the quarry.
Three large mustelids have been described from this area by
White: Mephititaxus ancipidens 1941, Aelurocyon spissidens
1947 and Oligobunis floridanus 1947. The isolated upper molars
referred to by White in 1942 (M.C.Z. Nos. 3639, 3640) as mus-
telids, have been re-identified as deciduous teeth of a canid. The
lower jaws, without teeth, mentioned by him (M.C.Z. Nos. 7029,
7030) are conspecific with the specimen here described.
This individual, although incomplete, marks the first occur-
rence of associated mandible and tooth in a small mustelid from
this locality.
2 BREVIORA NO. 51
Family Mustreipsar
Subfamily MustTELINAE
MIoMUSTELA(?) sp.
Referred material. M.C.Z. No. 7016 (Fig. 1), incomplete left
mandible with M, present and alveoli for C, P;, Pe, P3, and Ms
(incomplete).
Horizon and locality. Arikareean Miocene. Thomas Farm, Gil-
christ County, Florida.
Ui dls
Miomustela(?) sp. Labial, occlusal and lingual views. M.C.Z. 7016. X 3.
Characters. Closest to Miomustela madisonae (Douglass 1908 ;
Hall 1930) among previously known forms but differing as fol-
lows: M, with metaconid smaller, all other crown elements 25
per cent larger.
Description. The mandible is short and shallow and the num-
ber of premolars is reduced to three. All of the premolars are
1956 A SMALL MUSTELID FROM THE THOMAS FARM MIOCENE 3
double rooted and their arrangement indicates a crowded tooth
row. M, has a large well-defined protoconid with a pronounced
cleft between the protoconid and paraconid. The metaconid is
about one-half the size of the paraconid. The talonid is basined,
with a well-defined rim and the hypoconid retains its individual-
ity. The hypoconulid is small but distinct. The paraconid-
protoconid shear is nearly parallel with the long axis, making
the protoconid-metaconid width narrow as compared with the
tooth length. My is single rooted. The ascending ramus of the
jaw starts just posterior to M,, causing Mz to be tilted upward
and forward. The masseteric fossa extends forward to the an-
terior rim of the alveolus of Ms. Mental foramina are present
below the centers of P; and Py.
MEASUREMENTS (IN MILLIMETERS) OF MANDIBLE & TOOTH
LENGTH OF TOOTH ROW AS INDICATED BY ALVEOLI
(ANTERIOR MARGIN OF P TO POSTERIOR MARGIN OF M,)- --- ------
METACONID
PARACONID
SEYCOCONUEID, © at oe ee) iS
DISCUSSION
Plesictis (including Mustelavus) agrees with M.C.Z. No. 7016
in the general structure of M,, but Plesictis has the metaconid
and paraconid of equal size, and a jaw that is longer (with four
premolars) and lighter in build, the ascending ramus arising
well behind Ms (Teilhard de Chardin 1914; Viret 1929; Clark
1937; Simpson 1946; Hall 1951; and original material).
Agreement of our specimen with Miomustela is closer than with
Plesictis — sufficient at least to justify tentative generic as-
signment. Minor differences between the type of Miomustela and
M.C.Z. No. 7016 are an apparently lower paraconid and talonid
4 BREVIORA No. 51
in the type. I believe that these are due to the unworn condition
of the described specimen, compared with the worn cusps of
the type. This is based on study of the amount of wear under-
gone by M, in a large series of Martes americana, the essen-
tially similar cusp arrangement shared by this genus and Mio-
mustela permitting a valid comparison. This series shows that
such differences may indeed be due to wear.
The species represented by this fragment is clearly distinet
from M. madisonae but, considering the fragmentary nature of
available specimens, I refrain from proposing a new species.
Further material may, in fact, show that a new generic assign-
ment is necessary.
Acknowledgments. I am indebted to Professor A. S. Romer
for the use of the material and to Professor Bryan Patterson for
his helpful criticism. Also to Dr. J. LeRoy Kay for the loan of
the type of Miomustela. The figure was drawn by Miss Patricia
Washer.
REFERENCES
CLARK, J.
1937. The stratigraphy and paleontology of the Chadron formation
in the Big Badlands of South Dakota. Ann. Carnegie Mus., 25:
261-350.
Douauass, E.
1903. New vertebrates from the Montana Tertiary. Ann. Carnegie
Mus., 2: 145-199.
FILHOL, H.
1879. Etude des mammiféres fossiles de Saint-Gérand le Puy (Allier).
Bib. Ecole Hautes Etudes, Sect. Sci. Nat., 19: 1-252.
Haun, EB. R.
1930. Three new genera of Mustelidae from the later Tertiary of
North America. Jour. Mam., 11: 146-154.
1951. American weasels. Univ. Kansas Publ., Mus. Nat. Hist., 4: 1-466.
SIMPSON, G. G.
1946. Paleogale and allied early mustelids. Amer. Mus. Nov., 1320:
1-14.
TEILHARD DE CHARDIN, P.
1914. Les carnassiers des Phosphorites du Quercy. Ann. Paléont., 9:
103-192.
1956 A SMALL MUSTELID FROM THE THOMAS FARM MIOCENE 5
VIRET, J.
1929. Les faunes de mammiféres de 1’Oligocéne supérieur de la
Limagne Bourbonnaise. Ann. Univ. Lyon, (n.s. 1), sci. med.,
47: 5-305.
WHitr, T. E.
1941. Additions to the Miocene fauna of Florida. Proc. New England
Zool. Club, 18: 91-98.
1942. The Lower Miocene mammal fauna of Florida. Bull. Mus. Comp.
Zool., 92: 1-49.
1947.
Additions to the Miocene fauna of North Florida. Bull. Mus.
Comp. Zool., 99: 497-515.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. April 6, 1956 NUMBER 52
REMARKS ON SOME MIOCENE ANURANS
FROM FLORIDA, WITH A DESCRIPTION
OF A NEW SPECIES OF AYLA
By WatterR AUFFENBERG
Up to now, the only reference to Miocene anurans from Flor-
ida is by Tihen (1951), who described a new species of Bufo and
allocated other remains to Rana sp. All of these fossils were
collected at Thomas Farm, Gilchrist County, Florida. While
washing a considerable amount of matrix from this same deposit
in search of fossil snakes, a rather large number of amphibian
remains were found, and these form the basis of this report. The
new material permits addition of three more genera to the fauna
of the Miocene, and also contributes to a better understanding
of the relationships of the previously described Bufo.
Buro PRAEVIuS Tihen
This toad was described on the basis of fragmentary elements
of both the axial and appendicular skeletons. The relationships
of B. praevius were not discussed at length by Tihen, although
he stated that the ilium is somewhat comparable to that in B.
terrestris and B. valliceps. Considerable additional material
from the same deposit contributes to our knowledge of its prob-
able relationships. Of particular interest is the fact that there
are now available cranial elements, generally considered to be
quite diagnostic between species of toads. This is especially true
of the orientation and degree of development of the various cra-
nial crests. Information regarding these structures is now avail-
able for praevius.
The systematic relationships of many of our North American
toads have been the subject of considerable controversy during
2 BREVIORA No. 52
the last few years. The present paper does not deal with the
status of the species or subspecies of Recent Bufo, and the latest
checklist (Schmidt, 1953) has been followed throughout for tax-
onomy.
As comparative material, a number of Recent toads have been
skeletonized and examined: Bufo t. terrestris (23), B. t. amert-
canus (1), B. w. woodhouser (2), B. w. fowleri (8), B. valliceps
(2), B. punctatus (1), B. compactilus (1), B. simus (1), B. gran-
ulosus (1), B. quercicus (4), B. perplexus (1) and B. pelticeph-
alus (1).
The ilium is considered by many workers as being of consider-
able value in distinguishing many genera of fossil anurans, and
is the element on which the description of B. praevius rests.
Sixty-two additional ilia of praevius have been collected at the
type locality ; these have been deposited in either the collections
of the University of Florida (UF), or those of the Museum of
Comparative Zoology (MCZ).
One of the most important characters of the ilium is the shape
and degree of development of the dorsal prominence. In praevius
this prominence is fairly low, similar to the same structure in B.
terrestris, valliceps and woodhousei. On the other hand, B. com-
pactilis and cognatus (latter fide Tihen, op. cit.) have a spine-like
prominence, being high, with a narrow base. This fact appar-
ently indicates a closer relationship between praevius and the
terrestris group than with compactilis and/or cognatus. In B.
simus, punctatus, pelticephalus, quercicus, perplexus and granu-
losus the prominence is not roughened or knob-like, but forms a
smooth, well-defined, rather sharp dorsal edge.
Though the ilial prominence appears, in general, to be useful
in separating natural species groups in the genus Bufo, changes
in shape and proportions with growth are complicating factors
that must be recognized. In an examination of the shape of the
prominence in a series of ilia of Bufo praevius now available, it
is obvious that this character is more variable than indicated in
Tihen’s original series. As in modern B. terrestris, this structure
may vary from very low and smooth in small specimens, to con-
siderably higher and roughened in larger individuals.
Tihen stated that the ilial shaft of praeviws seemed to be less
compressed than in other toads with which he compared it; but
he also indicated that the degree of compression was not beyond
1956 MIOCENE ANURANS FROM FLORIDA 33
the range of variation found in the living forms. The diagnostic
importance of this character is considerably lessened by the fact
that a comparison of the degree of compression in the larger
series of ilia of praevius now available and a series of ilia of mod-
ern terrestris shows little, if any, difference between the two
species.
Another character of apparent diagnostic importance is the
eurvature of the ilial shaft, which according to Tihen is less than
that in other species of Bufo which he had examined, though not
beyond the range of variation of modern species. This character,
as well as many others, shows considerable ontogenetic change.
However, the present series tends to confirm Tihen’s observation,
1e., that the ilial shaft in praevius is, for the most part, less
eurved than in most other species.
In none of the ilia is there a process on the antero-ventral edge
of the acetabular expansion such as Tihen observed in one speci-
men (MCZ 1933). He states that although the ilium may be
aberrant, it might also represent still another species of Bufo.
A similar element has not turned up and it is highly probable
that this specimen is an aberrant one.
Three fragmental urostyles were also available to Tihen, who
stated that the crest was probably lower in praevius than in most
modern species. Although a number of fragmental urostyles
have been collected, only two elements have been found complete.
In these, as well as in the urostyles of two specimens of Recent
valliceps, the height of the crest is equal to, or slightly less than
the width of the articular surfaces. This crest is practically al-
ways higher than the greatest width of the anterior articular sur-
faces in Recent specimens of B. t. terrestris, t. americanus, w.
woodhousei and w. fowler.
A considerable number of presacral vertebrae are available for
the fossil species. They do not appear to be ‘‘heavier’’ than those
of modern species, but are seemingly identical with those of B.
terrestris and B. woodhousei. A number of sacral vertebrae are
also available, One sacral was found to be fused to the preceding
element, forming a very symmetrical unit. This unit is clearly
referrable to a bufonid, most likely representing an aberrant
specimen of Bufo praevius.
Of particular interest in the diagnosis of modern species of
toads is the shape of the cranial crests. Fortunately, cranial ele-
4 BREVIORA NO. 52
ments of the fossil species are now available. These include three
frontoparietals (UF 9892), two temporals (UF 9893), three
bones of the occipital complex and a number of squamosal stems
and mandibles (UF 9894). Thus it is possible to reconstruct the
major cranial ridges of the fossil form. In the following discus-
sion the terminology used by Sanders (1953) is used throughout,
since the system seems reasonable and practical.
The available frontoparietals of the fossil species are some-
what fragmentary, but indicate a number of important facts.
They are cancellous and provided with well-developed crests.
Furthermore, the shape and development of these crests clearly
show that its relationships lie with the terrestris-woodhousei
species complex. From above, the parietal ridges of praevius are
slightly diverging posteriorly. The fragmentary elements are not
large enough to determine the length and size of the frontal
ridges, although such ridges probably existed, as based on an
examination of the anterior ends of the fragments. The parietal
ridges are low anteriorly, gradually increasing in height pos-
teriorly, much in the manner of those of woodhousei. From
above, these ridges are relatively narrower than in terrestris or
houstonensis (the latter fide Sanders, op. cit., fig. 2), but like
those in woodhousei. From the side they are slightly arched, as
in valliceps, not straight as in woodhousei, terrestris or houston-
ensis. The otoparietal ridge is well-developed and rounded, being
somewhat intermediate between that in houstonensis and w. fow-
leri. It is not as high and narrow as in valliceps. It is better de-
veloped than in terrestris, and of a different shape. The angle
formed between the otoparietal and parietal ridges is approxi-
mately 90°, not greater, as in valliceps. The occipital groove, if
present, is very inconspicuous, The otoparietal plate is not much
wider than its ridge, as in houstonensis. However, it should be
pointed out that this character, given considerable weight by
Sanders (op. cit.), varies with age and possibly with locality in
at least B. terrestris. The largest part of the temporal plate is
cancellous, provided with a well-developed, but rounded temporal
ridge. The tympanic and supratympanic ridges are definitely
well developed, but to an unknown degree due to the fragmentary
nature of the available elements. The parietal spur is well devel-
oped, directed backward and medially. It is not club-like as in
t. americanus, but is similar to that in B. woodhousei. The shape
1956 MIOCENE ANURANS FROM FLORIDA 5
and position of the cranial crests in B, praevius and three closely
related modern species are shown in Figure 1.
From the foregoing description of additional remains of Bufo
praevius it is highly possible that its relationships lie with the
eastern toads. This has been intimated by Tihen in regard to the
ial prominence, and is very much strengthened by the shape and
Fig. 1. Shape of the cranial crests in four species of Bufo. A, Bufo
houstonensis (after Sanders, op. cit.); B, Bufo t. terrestris; C, Bufo prae-
vius; D, Bufo w. fowleri.
development of the cranial ridges. A few of the characters, such
as the height of the urostyle crest and the arching of the parietal
ridge are somewhat suggestive of B. valliceps. The importance of
the suggested close relationship of praevius to the terrestris-
woodhousei-houstonensis complex is that it indicates that this
group was already well established by early Miocene.
6 BREVIORA NO. 52
RANA sp.
This genus has already been reported from the Thomas Farm
by Tihen (op. cit.). Two additional ilia are now available (UF
5919). These remains are too fragmentary to make a species
identification possible, although the shape of the posterior edge
of the dorsal crest is highly suggestive of the pipiens ‘‘group’’,
including palustris, pipiens, sylvatica, clamitans, ete. They are
not as readily referrable to the group including heckscheri, cates-
beiana and grylio.
ScAPHIOPUS cf. HOLBROOKI
In the fossil record, Scaphiopus is known from Pliocene (Tay-
lor, 1941) and Pleistocene (Tihen, 1954) deposits of North
America. Its occurrence in the early Miocene of Florida is thus
of considerable interest.
The genus is frequently divided into two subgenera: Scaphi-
opus (ineluding h. holbrooki, h. hurteri and couchi) and Spea
(ineluding multiplicatus, bombifrons and hammond). Skele-
tons of all species except multiplicatus have been examined.
Included in the fossil material from Thomas Farm are four
fragmentary ilia (UF 9896), one maxilla (UF 9897), three frag-
mentary frontoparietals (UF 9898) and one presacral vertebra
(UF 9899), all of which appear to belong to this genus.
The ilium of the subgenus Scaphiopus is quite diagnostic.
There is no ilial crest, and the dorsal prominence is absent or but
slightly developed. When present it may exist as a small,
rounded protuberance, directed dorso-laterally, about half way
between the base and the end of the acetabular expansion. In the
subgenus Spea the prominence is usually ridge-like, directed
more dorsally, and contributing to the height and length of the
dorsal portion of the acetabular expansion. In the fossil ilia the
dorsal prominences are very small and rounded, identical with
those found in the subgenus Scaphiopus. The disposition of the
maxillary teeth, as well as the shape of the entire element, are
also comparable to those of the subgenus Scaphiopus. There is
no frontoparietal boss, such as is present in some members of the
genus. On the basis of the available fossil remains, the Miocene
form is not distinguishable from the modern species, Scaphiopus
holbrook. ‘
1956 MIOCENE ANURANS FROM FLORIDA fi
Whether or not the Miocene Scaphiopus was identical to the
species holbrooki cannot be absolutely determined at this time.
However, the fossil form is clearly referrable to the subgenus
Scaphiopus. Whether or not the subgenus Spea had differenti-
ated by this time, or whether the former is ancestral to the latter
is, of course, unknown. Further collecting in other Tertiary de-
posits of middle North America may clarify many of our con-
cepts regarding the ancestry of this group.
Fig. 2. Comparison of the fossil microhylid ilium from the Thomas Farm
locality (4) with that of a recent specimen of Microhyla carolinensis (B),
from Gainesville, Florida. The line represents 1 mm.
MICROHYLA sp.
Of considerable interest is the fact that one fragmental ilium
from the Thomas Farm (UF 5144) apparently represents the
genus Microhyla. Its very small size and well-developed dorsal
prominence (which is triangular from the side and without any
protuberances or roughened areas of any kind) are highly sug-
gestive of this genus. In addition, the acetabular cup is very
similar to that in Wicrohyla, in that the edges of the cavity are
considerably raised above the surface of the acetabular expan-
8 BREVIORA NO. 52
sion. Although broken, the ventral portion of the acetabular
expansion was apparently narrow and directed downward as in
this genus. That this element does not represent a young speci-
men of Bufo is indicated in the shape of the dorsal prominence.
In small specimens of Bufo this prominence is weakly developed,
being low and long, usually with a roughened area on, or very
near the dorsal edge. The genus Microhyla has not been reported
as a fossil. Additional material, both fossil and modern, is neces-
sary before specific identification of the fossil form will be pos-
sible (see Fig. 2).
In addition to the remains already mentioned, three ilia were
found which are referrable to either Hyla, Pseudacris or Acris.
They are placed in the genus Hyla mainly for reasons of conven-
ience. Neither Acris nor Pseudacris have been found as fossils.
Hyla has been reported from the Pleistocene of North America
(Brattstrom, 1953). The fossils under consideration appear to
represent a new species. I wish to name this form for Dr. Cole-
man Goin, who has contributed materially to our knowledge of
modern amphibians in life history studies, ecology and taxonomy.
HYLA GOINI nov. sp.
Diagnosis. A Miocene Hyla with a relatively high ilial shaft ;
ilium without a dorsal crest; ilial prominence rounded, well de-
veloped, with the protuberance located rather low on the promi-
nence, projected dorsally and laterally, its anterior edge practi-
cally even with, or slightly behind the anterior edge of the acet-
abulum; ventral portion of the acetabular expansion broad at its
base; acetabulum somewhat sub-triangular.
Holotype. MCZ 2277; the distal 7 mm. of a right illum, col-
lected by Walter Auffenberg, March, 1954 (Figure 3).
Horizon and Type Locality. Hawthorne formation, Lower
Miocene, Arikareean; Boulder Bar, Thomas Farm, Gilchrist
County, Florida.
Referred Material. UF 9900; three fragmental ila from the
same locality and horizon.
The ilia of the genera Hyla, Acris and Pseudacris are easily
separated from those of Rana and Eleutherodactylus by the ab-
sence of a dorsal crest. From Microhyla, Bufo and Scaphiopus
they are readily distinguished by the well-developed dorsal prom-
1956 MIOCENE ANURANS FROM FLORIDA 9
inence, directed dorsolaterally to laterally, and by the slender-
ness of the shaft itself.
The ilia of Hyla goint have been compared with the same ele-
ment in the following genera and species: Hyla cinerea (12), H.
squirella (6), H. femoralis (2), H. crucifer (2), H. versicolor
E
Fig. 3. Interspecific variation in the ilia of Hylidae. 4A, Hyla femoralis,
4 miles E. Gainesville, Florida; B, Acris gryllus dorsalis, Biven’s Arm,
Alachua County, Florida; C, Hyla c. cinerea, Gainesville, Alachua County,
Florida; D, Pseudacris ormata, 7 mi. northeast of Gainesville, Florida; £,
Hyla crucifer bartramiana, Camp Olena, Columbia County, Florida; F, MCZ
2277, type ilium of Hyla goini nov. sp., L. Miocene, Thomas Farm, Gil-
christ County, Florida. The lines represent 1 mm.
10 BREVIORA No. 52
(1), H. septentrionalis (1), Pseudacris mgrita (4), P.- ornata
(3), P. brimleyi (1), Acris gryllus (3).
The dorsal prominence of Acris gryllus is oval and on a rather
long base, located in front of, to very slightly behind, the an-
terior edge of the acetabulum. In addition, the ventral portion
of the acetabular expansion is shorter at its base than in H. goini.
In all the hylids that I have examined, with the exception of
crucifer, and to some extent femoralis, the dorsal prominence is
oval, usually with a long base, and with the protuberance of the
prominence located considerably higher than in goin. In speci-
mens of H. crucifer, and some of femoralis, the protuberance of
the dorsal prominence is relatively small, rounded, not too high,
and on a fairly small base. However, the prominence in crucifer
is anterior to the anterior edge of the acetabulum, and the lower
portion of the acetabular expansion is considerably longer and
narrower than in goini. In femoralis the protuberance is higher
on the dorsal prominence and the acetabular expansion is also
somewhat higher. In Pseudacris nigrita, P. brimleyt and P.
ornata the dorsal prominence is rounded, never as oval as in most
species of Hyla that I have seen. Its base is relatively short and
the entire prominence is usually anterior to the anterior edge of
the acetabulum. However, goin is considerably larger than any
modern Pseudacris. It is apparently the size of an average speci-
men of Hyla cinerea. It also differs from modern species of
Pseudacris that I have examined, in the shape of the acetabular
expansion which is longer at its base. In addition, the somewhat
subtriangular acetabulum apparently separates it from most spe-
cies of this genus.
In many respects, the ilia of Hyla and Pseudacris are very sim-
ilar. This is to be expected if the genera are really as closely
related to one another as is generally supposed. The fossil form,
goini, is clearly allied to one or both of these genera on the basis
of its ilial shape. As a representative of the Hylidae in the Lower
Miocene it could, conceivably, be ancestral to both genera. The
fact that goini shows characters found in both modern genera
may be indicative of an ancestral position, or Simply a reflection
of the close relationship which evidently exists between Hyla and
Pseudacris. In any ease, goini can be reasonably placed in the
genus Hyla on the basis of its size and the fact that all of its char-
acters are duplicated in one form or another in this genus. Ad-
1956 MIOCENE ANURANS FROM FLORIDA 3 lak
ditional specimens representing other parts of the skeleton are
needed before its exact status can be clearly fixed. Figure 3 com-
pares the ilia in certain hylid frogs, selected to show the typical
shape within each species, with the type of H. goini.
ACKNOWLEDGMENTS
For advice and eriticism of this paper I wish to thank Drs.
A. B. Grobman, P. Brodkorb, E. E. Williams and A. S. Romer.
Acknowledgment is due Mr. L. Ogren, University of Florida,
who has helped me on innumerable occasions in the field, in iden-
tifying fossil remains, and in preparing comparative skeletal
material. I also wish to thank Mr. R. Highton for the loan of a
number of anuran skeletons.
LITERATURE CITED
BratTTstRom, B.
1953. The amphibians and reptiles from Rancho La Brea. Trans. San
Diego Soc. Nat. Hist., 11(14) :365-92.
Cooke, C.W.
1945. Geology of Florida. Bull. Florida Geol. Sury., 29:1-339.
SANDERS, O.
1953. A new species of toad with a discussion of morphology of the
bufonid skull. Herpetologica, 9(1) :25-47.
Scumipt, K. P.
1953. <A checklist of North American amphibians and reptiles. Amey.
Soe. Ichthyologists and Herpetologists. Univ. Chicago Press:
Chicago, Illinois.
Tayuor, E. H.
1941. Extinct toads and salamanders from middle Pliocene beds of
Wallace and Sherman Counties, Kansas. Bull. Kansas Geol.
Surv., 38(6) :177-96.
TIHEN, J.
1951. Anuran remains from the Miocene of Florida, with the descrip-
tion of a new species of Bufo. Copeia, 1951 (3) :230-35.
1954. A Kansas Pleistocene herpetofauna. Copeia, 1954 (3) :217-21.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. APRIL 6, 1956 NUMBER 53
FOOD-FINDING BY A CAPTIVE PORPOISE
TURSIOPS TRUNCATU SY:
By Wiu1amM FE. ScHEVILL AND BARBARA LAWRENCE
Introduction 1
Experimental arrangements 2
Passive location 4
Active or echolocation 4
Kcholocation to determine presence of fish 6
Kcholocation to distinguish between alternate feeding
places if
Passive versus active location 8
Night tests 9
Vision 10
Discussion 12
Acknowledgments 14
References 14
INTRODUCTION
It soon appears on acquaintance with porpoises that these ani-
mals are well endowed with hearing and sight, and that they use
both these senses in their normal environment. A captive Twur-
siops truncatus was confronted with the problem of finding un-
familiar food under varying and sometimes rather complicated
circumstanees, all unlike the normal hunting of his previous
wild free existence. Experiments to find out which sense was
relied on most and in what ways it was most useful have shown
a wide and not unexpected variation in a single individual. Like
man, the animal used all the clues he could get. Sometimes he
listened, sometimes he looked, and sometimes he was so busy
doing whatever he had last done that he missed perfectly obvious
clues.
1 Contribution No, 832 from the Woods Hole Oceanographic Institution.
2 BREVIORA NO. 53
Although there was such variation in his responses to similar
circumstances that many experiments produced conflicting evi-
dence, there was a certain pattern which makes possible a number
of conclusions. When he was finding his food he was most eager,
swift, and accurate in coming to the sound of a slap on the water.
When he had no such clue he would find the fish, evidently by
echolocation, if it was in a region where he expected to find food.
His final searching for the fish was by eye, though he could not
clearly distinguish his preferred butterfish from other offerings.
His willingness to use clues, almost we might say his ability to
notice them, depended on his memory and other psychological
factors as well as hunger.
EXPERIMENTAL ARRANGEMENTS
Our laboratory on an island at Woods Hole was a pond 34 m.
long, 23 m. wide, and 2.5 m. deep, cut off from the sea by a stony
beach about 30 m. wide. Our subject was an old bull Tursiops
truncatus, 203.6 kg. in weight and 267 em. long, brought here
from Florida especially for this experiment. Some injury had
damaged his right eye, which we never observed him using, and
towards the end of our work his left eye started to cloud over.
Whatever the damage was, it did not affect his hearing, which
was sharply directional. Temperamentally, he differed from our
earlier subject (Lawrence and Schevill 1954), being bold and
aggressive, and for this reason was often seen at the surface.
Puzzling situations near at hand often caused him to thrust his
head above the water, or ‘‘souse out’’ in this way to look, and
when further off he would ‘‘pitchpole’’ straight up for a third or
more of his length. He learned fast, and while his evident pre-
ference for doing as he pleased would make any formal analysis
of his responses rather meaningless, this independence gave a
much more varied and accurate picture of his inclinations and
abilities than we could have obtained otherwise. Since he pre-
ferred going hungry to being foreed into situations he disliked,
the tests we devised were as simple as were consistent with get-
ting reliable answers. Under these circumstances, it soon became
clear that in many cases failure on his part to respond was due
not to inadequacy of his senses, but to lack of hunger, temporary
unwillingness to approach the feeder, or some other non-sensory
reason.
1956 FOOD-FINDING BY PORPOISE 3
The porpoise was ordinarily fed from a punt 3.6 m. long
moored against the bank, usually at right angles, but sometimes
parallel to it. In some experiments a small dinghy served as a
second feeding station. The position of the feeder in the boat
varied. The porpoise was fed dead butterfish (Poronotus triacan-
thus) about 8 to 23 em. long, held in the water by hand, and was
called by an acoustic signal made by slapping the water, or by
hammer strokes on a partially immersed iron pipe, or by tones
(ranging usually between 500 and 30,000 cycles per second) from
an audio oscillator through an underwater sound projector; the
pipe and oscillator signals were always remote from the feeding
station.
Most of our work was done during daylight, so that we could
watch where the animal was and how he responded under differ-
ent conditions. It was not difficult to keep track of him because he
frequently showed at the surface. Because porpoises see well
both above and under water, we had to be sure than an appar-
ently acoustic response was not in reality visual. This was the
easier because his constant swimming kept the water stirred up
and very murky. Transparency by Secchi disk from the surface
was rarely as great as 70 em. and often less than 45 em.; the very
unusual maximum was 85 em. on 19 September. At 11 o’clock
on a typical sunny day measurements made by diving after dark-
adaptation gave a Secchi disk reading of 23 em. from the diving
mask at a depth of 2.4 m., and 61 em. at the surafce. Under-
water visibility will scarcely exceed the Secchi disk reading, and
will be appreciably less for less bright objects.
This, and observations of the porpoise’s behavior when he was
finding fish by eye, led us to believe that we are conservative in
saying that through the water he could not possibly have dis-
cerned with any clarity objects as much as 1 m. away from him.
As a further check we repeated most of our tests at night.
We selected the pond because of its great freedom from noise-
making animals; the beach protected us from most of the noises
of the sea outside. Our listening gear included an AX58C
Rochelle salt crystal hydrophone and a WHOI sound level meter
(Suitcase), and was sensitive enough to pick up very plainly the
noise of fine beach sand strewed into the water 20 m. from the
hydrophone.
4 BREVIORA NO. 53
PASSIVE LOCATION
One of the most conspicuous traits of our porpoise was the
accuracy and confidence with which he promptly made his way
to the place where the water had been slapped. There was no
uncertainty or hesitation in his response to such a signal ; no mat-
ter where he was in the pond, he always came with alacrity when
he was thus called unless he was not hungry or there were other
obvious psychological reasons for his refusing to do so. While
failure to reward response to a remote signal often made him ig-
nore subsequent calls, this was never the case with a slap on the
surface. As often as we repeated the slap he would return quickly,
though we never called him more than four or five times in suc-
cession without giving him a fish.
Not only was he prompt in his response to this slap, but, when
conditioned to it, he chose it in preference to other clues. If
(as described on p. 8) a fish was slapped in one place and held
in the water in another, he would choose to go to the place of
the sound instead of to the fish, until he learned this clue was un-
reliable. His apparent reliance on this signal alone sometimes
led him to miss a nearby fish entirely.
The porpoise’s hearing was sharply directional and his esti-
mates of range (distance) were very close, as is shown by his
repeatedly homing directly on a single slap (with no fish in the
water) to well within 20 em. from ranges often as great as 20 to
25 m. through turbid waters and without coming up to look on
the way.
ACTIVE OR ECHOLOCATION
The evidence that our porpoise was echolocating his fish was
accumulated over eleven weeks of close observation of his be-
havior, when an increasing knowledge of what to expect from
him under different circumstances made it possible to understand
and to check his various capabilities with a fair degree of
accuracy.
Early in our work we noticed that our porpoise usually made
a characteristic sequence of ‘‘ereaks’’ as he came in for a fish,
and that in the last meter or so these sounds were matched to
horizontal movements of his head. By ‘‘creaks’’ we mean a series
1956 FOOD-FINDING BY PORPOISE 5
of impulsive clicks made at widely varying repetition rates
(from less than 10 to more than 400 per second), the slower ones
sounding like knocks and the faster ones like snarls or whines.
They have also been called ‘‘barks’’, ‘‘snores’’, ‘‘rusty hinge’’,
or ‘‘rasping and grating sounds,’’ ete., by various authors, such
as McBride and Hebb 1948, Kritzler 1952, and Wood 1954, and
have been heard from several odontocetes. Some acoustic details
of Tursiops calls have been given by Kellogg, Kohler, and Mor-
ris 1953. The other common odontocete sound, the whistle-like
squeal, was evidently not employed in echolocation, and appears
to be primarily communeative. The acoustie details of these
sounds will be reported elsewhere.
Before long we noticed that when he was creaking he almost
always swam directly to a fish held quietly in the water. When
he was not creaking he would not do so. Though this suggested
echolocation rather persuasively, we had to be sure that we were
not inadvertently giving him other clues and that he was not
using sight. His remarkably good hearing and his evident reliance
on passive auditory clues made it especially important to make
sure he was not coming because he heard us put a fish in water.
Very occasionally it seemed highly likely this was just what he
was doing, though we ourselves could never detect any sounds,
even with our extremely sensitive listening gear. Responses at
these few times were discounted. At other times when we were
doubtful, we checked his behavior by dipping fish or fingers in
and out or dabbling at the surface. This almost never brought
him, nor could we get him to come unsignaled to a fish’s nose in
the water, even though he was coming accurately to a whole
fish. Repeated checks also eliminated movements in the boat
or the position of the feeder as clues.
The possibility that he was using sight to guide him to a fish
had also to be investigated, and we did this in two ways. First
of all we devised a series of daytime experiments which, because
of the murkiness of the water (see Secchi disk readings), elimi-
nated any possibility that he was using vision to locate his fish
from a distance. These experiments are described in some detail
below. After we had learned how he reacted to these different
situations we repeated the tests at night, with similar results,
which are also described below.
6 BREVIORA NO. 53
Two rather different sets of circumstances stimulated him to
rely on echolocation alone in finding his food. Sometimes he
used this means to discover whether or not a fish was waiting for
him, and sometimes this was the way he distinguished between
alternate feeding places.
ECHOLOCATION TO DETERMINE PRESENCE OF FISH
His reliance on echolocation to tell him whether or not a fish
was waiting showed in a number of ways. Often, in the absence
of an expected signal, he would circle the pond, creaking only as
he passed the feeding station. This was anywhere from 1 to 5
meters away, and occasionally farther. If a fish were in the
water as he passed, he would turn and swim directly to it; if
there were no fish he would go on by. If a fish were slipped as
silently as possible (and we believe inaudibly) into the water
after he had passed when he was starting to circle away, he
would usually turn and come back. We tried this many times on
seven different days. Of these the first is especially significant.
It was early in our work and was our first attempt to bring him
to a fish without a signal, he being then conditioned to come
only to the slap of a fish on the water. At first, although he
passed nearby creaking, he did not come to the fish, but later
that morning he was attracted some of the time. Five days later
we tried again, and he had apparently learned not to wait for a
signal. By now he would come in for a fish held silently in the
water if he was creaking as he passed by; his decision to come
on in seemed usually to be made at a distance of less than 5 but
occasionally as much as 15 m. This was repeated a number of
times during the next four days, and again two months later.
Our evidence that the porpoise was not seeing the fish before
deciding to swim to it was partly his excessive distance when he
turned toward it and partly the fact that he often had his bad
eye towards the punt as he passed.
As well as using echolocation to find fish when he suspected
fish should be available, he also used this means to confirm the
reliability of a signal. While he almost never refused to come to
the sound of a fish slapped on the water, various circumstances
on several occasions made him uncertain about other signals.
Early in his training he learned that an oscillator note or pipe-
banging in various parts of the pond meant that he would get a
1956 FOOD-FINDING BY PORPOISE 7
fish at the feeding station, and, like our earlier porpoise, on
being signaled with no fish in the water, he would make his way
to the proper place to get it. Sometimes, however, he ignored the
remote signal until there was a fish in the water as well. With
this to convince him he would come unhesitatingly. This reaction
was especially clear on four occasions when we had made changes
which he did not like in the feeding station. These were all times
when he was in good health and hungry, and the remote signal
was one we knew he could hear. Until he was accustomed to the
new arrangement, no matter how often we called, he always waited
till there was a fish in the water before responding to the signal.
The first time we had to signal six times with a fish in the water
before he began to come on signal only. The other three occasions,
at a much later date, were at successive feedings on two days. The
first morning it took twelve fish, that afternoon three, and the
following morning seven before he would come with no fish in
the water. At these times he was creaking as he swam and for
the most part responded to the signal by starting towards the
feeding station, but with no fish in the water he usually turned
away at 214 meters or more, though sometimes he circled as near
as a meter and a quarter. When he was thus relying on echo-
location to tell him of the presence of a fish, he rarely troubled to
look above the surface. Later, when he was coming on the remote
signal with no fish held in, he often soused out of the water on
the way in, eyeing the situation from a distance of ten or more
meters.
His ability to arrive at a fish he could not see was further
demonstrated one afternoon when his left eye failed. At that
time, on a remote signal, he repeatedly swam directly to fish held
anywhere over a 514 m. radius.
ECHOLOCATION TO DISTINGUISH BETWEEN
ALTERNATE FEEDING PLACES
To test his use of echolocation, two feeders slapped fish simul-
taneously on opposite sides of the punt and then one held a fish
in the water while the other held a hand out over the water as if
feeding. The distance between slaps was about 2 m., and a net
projecting 2.5 m. from the end of the punt and hanging to the
bottom of the pond prevented the porpoise from circling close
8 BREVIORA No. 53
to investigate by eye. We alternated feeding in irregular fashion
and the feeders often exchanged places in order to eliminate any
other clues. The first time we tried this he came to the correct
side 24 times and the wrong side 11, the next time he made 45
correct responses and 13 wrong. His behavior was similar on
both occasions. He never ignored a summons and always came
directly and fast, creaking as he approached deep. The Secchi
disk reading was 61 em., and he was at the very least 2.5 m. from
the fish when he had to decide which side of the net to go. In
contrast to his behavior with a remote signal, he made no optical
checks on the way in, though sometimes when feeding was slow
he pitchpoled out, looking at us. We cannot say how often his
wrong responses were caused by the net interfering with his
sound patterns. We suspected this on some occasions, while
other errors probably are evidence that his echolocation was not
perfect.
PASSIVE VERSUS ACTIVE LOCATION
Experiments to find out what kinds of clues were most success-
ful or preferred produced interesting results. One rather simple
but very instructive one, here called the A-B experiment, con-
sisted of slapping at A and putting a fish in the water at B or
vice versa, with the distance between A and B farther than he
could see. This, with minor variations, we tried many times on
each of fifteen different occasions, and though his responses
varied, they made a very neat pattern. The first time we tried it,
and when we returned to it after a period of other work, he
would always swim, creaking, directly to the point of slap,
search diligently there, and usually swim away unrewarded. If,
in leaving, he found the fish, he would thereafter search at the
point of slap and then swim directly to the fish no matter how
we varied the relation of these two places to each other. Finally,
it was possible to destroy his confidence in the slap as a worth-
while clue, and then for the most part he would go directly to
the fish. When he swam directly to the fish there was no possi-
bility that memory could have guided him, because the fish might
have been anywhere over a six meter stretch. Nor could vision
have helped, because he was never nearer than 614 meters when
we signaled, and usually a great deal farther away. When he
1956 FOOD-FINDING BY PORPOISE y
looked first at the point of slap and then went on to find the fish,
not pausing on the way, the distance between the two places
(minimum 1.3 m., usually more, sometimes as much as 5.2 m.)
seemed to preclude the possibility of his being guided by sight,
the more so because in this secondary finding of a fish it made no
apparent difference whether he approached with his bad or his
good eye towards it.
In one variation of the A-B experiment we fixed the points
2m. apart. If we slapped at A with the fish in the water at B
or vice versa, he went directly to the fish; if we slapped with no
fish in the water, he searched at the point of slap but never in-
vestigated the alternate place as he left unrewarded ; if we let him
search at the point of slap and then eased a fish quietly into the
water at the other place, he would immedately go to it. On a
number of other occasions we tried this third modification, vary-
ing the place where we put the fish, and he always creaked his
way to it. This is not entirely conclusive, as there is a small but
unlikely possibility that he might have heard the fish put in,
though our efforts to check this led us to believe that this was
not passive location (ef. below, p. 11).
NIGHT TESTS
All of the experiments described above were carried out dur-
ing the day. We also tried most of these same experiments on
dark nights and found the results closely paralleled our daytime
observations. On two nights when we tried to get him to take
fish unsignaled, he came, apparently directly, making about ten
successful runs each time, though he sometimes swam past, creak-
ing, without coming in, and sometimes appeared to search near
the fish without taking it. On two other nights we tried the A-B
experiment, the first time feeding ten fish which he readily
found, though it was too dark for us to see if he made a pre-
liminary search at the point of slap. During the second and
more prolonged A-B experiment he repeatedly came to the fish,
not the point of slap, especially when he came from far down
the pond. Occasionally he searched first at point of slap, and
twice he missed the fish entirely. The last night feeding was
partly from the dinghy in the middle of the pond. Two slaps
informed him that fish were to be had in the vicinity ; thereafter
10 BREVIORA NO. 53
on remote signal he came directly and accurately seven times
in a row to a fish held anywhere in a radius of 6 m. Later, when
we fed from the punt, he responded to the correct side 6 times, to
the wrong side 2, and did not respond at all 2 other times.
VISION
While sound was important to our animal, we have good evi-
dence that he relied greatly on vision as well, both above and
beneath the surface of the water. A bad if not completely blind
right eye made him left-sided in his approach to things he wished
to see. This was a convenient check when we could not see his
eye rolled towards what interested him.
When on arrival from Florida he was dumped into the pond,
his first check of his new surroundings was optical. As soon as
he hit the water he swam off fast and silently, sousing high out
of the water and blowing frequently, with his good eye towards
the shore. It was three minutes before we heard him utter a single
sound, and nearly two minutes more before he spoke up as loudly
and persistently as he did for most of the rest of his stay. It
was hard not to think that he was looking over his new sur-
roundings, perhaps searching for a break in the beach. His
obvious awareness of things on the shore showed in many ways.
During his first two weeks in the pond there were often clusters
of people working at different places along the bank. At such
times he often blew near them, rolling a little on his right side
so that his left eye cleared the surface. Soon he took his sur-
roundings more for granted and his inspections of the shore
were less frequent, though he quickly noticed changes. Possibly
also with the passage of three or four weeks he became more ac-
ecustomed to his blind eye; at all events he rolled more often on
an even keel, and without bringing his good eye out of the water.
While this kind of check on his surroundings was not impor-
tant in helping him find his food, it did have a bearing on his
behavior at feeding time. Too many people on the bank near the
feeding station made him shy, and he would come in deep and
depart hastily. The presence of someone in the boat from which
we fed aroused his interest, and he would blow nearby, looking.
Sometimes it even seemed as if he reacted differently to different
feeders.
1956 FOOD-FINDING BY PORPOISE 11
When we actually called him to eat with the well-understood
slap of a fish on the water, he rarely troubled to make an optical
check on the way in as he did with a remote signal, but swam
directly from wherever he was, to collect his morsel. At other
times when he was less sure of the summons, or when we were
slow sending signals, he rolled high, looking towards the feeder, or
soused out to see what was going on. On occasions when he was
more than ordinarily curious, he would pitchpole out of water
as far as his flippers, with his good eye looking ventrad towards us.
In addition to keeping track of things above the water, under
ordinary circumstances he relied on vision to a great extent in
his final accurate taking of a fish from the feeder. Often we could
see his eye rolled forward towards the fish. As with our earlier
animal, space permitting he would turn over on his side when
close, and in the eleven weeks we fed him he only once took a
fish with his right eve up. When he lingered at the end of the
boat waiting for a fish it was always with the left eye up. When
we fed him in a sort of narrow stall 1.2 m. wide so that he did not
roll over as he approached, he swung his head from side to side
farther to the right than to the left, so that his left eye was in
position to scan both sides of the stall as well as the end of
the punt.
Fish put in on his blind side or above his head did not attract
his attention unless they were splashed. Fish put in nearby and
directly in front of him he also took in more fumbling fashion,
and on at least two oceasions actually bumped into them before
seizing them. On the other hand, anything within his range of
vision quickly caught his attention. When a fish was moved 15
em. or so above the water he would follow it with his eye and
when it was held lower would put his snout out to snatch it.
Repeatedly, we found a difference in his fish-taking when visi-
bility was especially poor or he himself not seeing well. This
usually meant that he would begin his search farther from the
fish, nodding his head more widely as he approached slowly, and
would be more hesitant about taking the fish, sometimes fumbling
and dropping it. A few times, when the fish were small, he missed
them entirely. This was,true not only when the water was murky,
but also when we held the fish deep.
These indications that vision is important in his ultimate locat-
12 BREVIORA NO. 53
ing of the fish were borne out by his behavior on two separate
occasions when the salinity in the pond was down and the water
dirty. At these times it appeared that his left eye also was not
seeing well. On the first occasion, for three days his fish-finding
was less accurate and his search wider than was usual. Once he
even bit at the corner of the punt, though the fish was less than
50 em. away. The other occasion was one afternoon when his
left eye failed; then, he approached with it shut, and would lie
left side up at the end of the punt, not noticing the fish when it
was moved above the surface, but coming to grab it clumsily
when it was put in the water. Once when he swam past a fish in
the water, he worked his way back to it slowly with very exag-
gerated head noddings and took the fish awkwardly deep in his
mouth. Though his sight recovered after this, he never seemed
to see out of his left eye as well as in the beginning. A whitish
spot began to form and he would partly close his eye against a
low sun, whether because it hurt or dazzled was impossible to tell.
At these times his fish-taking was again less accurate, and con-
trasted with a greater ease when the sun was not shining directly
in his eye.
While taking the fish was easier if he could rely on seeing it,
he apparently could not discriminate between objects very well.
Squid and flattened tin cans don’t resemble butterfish much, but
even after he had found out he did not lke the first two he re-
peatedly took them in his mouth when they were offered instead
of fish. In the same way he would bite at floating vegetation
near the boat, at bits of rope, or even at a rusty pail. Size seemed
easier for him to tell, and he not infrequently appeared to in-
spect and then leave small butterfish, about which he was not
enthusiastic.
DISCUSSION
In this study we took a gregarious free-ranging animal that in
nature is almost always found at least in small groups and often
in immense herds, and placed it in solitary confinement. Whereas
wild porpoises hunt live food that occurs in schools, ours was
obliged to take single dead fish from the hand. Moreover, the
single fish was usually close to a boat or a bank, instead of in the
more open water to which wild Tursiops are accustomed.
1956 FOOD-FINDING BY PORPOISE 13
Perhaps the most striking result of our work is the great varia-
tion observed in the way our single animal sought his food. His
primary reliance on passive auditory clues could have been due
to the small size of his target and to the confused echo patterns
in the pond. Nevertheless, evidence accumulated that he often
echolocated the food we offered him, thus supporting the wide-
spread supposition (for example, Kellogg, Kohler, and Morris
1953) that this was how cetaceans hunted. The sounds the
porpoise made at these times were faint; indeed, only the very
loudest were audible to a submerged man, and, in fact, were
picked up by our sensitive listening gear only because we at last
had a porpoise in a really quiet place. Thus we learned that the
supposed taciturnity of solitary porpoises (Lawrence and Schevill
1954, pp. 229-231) is rather a relative matter; it appears that
they merely speak very softly. The noisy listening conditions of
our previous experiment had led us into error when we reported
(op. eit., p. 229) ‘‘the complete absence’’ of evidence for echoloca-
tion although we cited some, unrecognized, at the bottom of page
227.1 The only evidence we had been aware of was in McBride’s
posthumous note (in press) on net avoidance (what our porpoise
taught us about this will be reported in another paper).
To demonstrate whether an animal is using echolocation, the
most definite way is to show that acoustic intereference affects
performance. Thus Griffin and Galambos (1941) and Griffin
(1953) by deafening bats and nocturnal birds showed that these
animals then collided with obstacles that, undeafened, they had
avoided. It is of course necessary to make sure that other senses,
such as smell (evidently not available to cetaceans) or sight, have
been excluded. Furthermore, it must be shown that sounds suit-
able for echolocation are produced. This last point is abundantly
proved for porpoises (e.g., Wood 1952, 1954). The role of sight in
our porpoise’s food-finding has been discussed under Vision. We
did not deafen our animal or interfere with his sound production.
Therefore, our evidence for echolocation by the porpoise is
essentially that he consistently found fish when we could convince
ourselves that no other clue (sight or sound not made by the
porpoise himself) was available.
1 On page 414 of an article published while this paper was in press, Griffin
(1956. Hearing and acoustie orientation in marine animals. Deep-Sea Re-
search, 3, Suppl. (1955), pp. 406-417) suggests that just such an improved
signal-to-noise ratio might reveal evidence of echolocation by porpoises.
14 ; ' - BREVIORA No. 53
The porpoise’s performance seemed poorer on targets behind
him, particularly at the longer ranges. In general, the creaks
with the higher repetition rate were heard at the shorter ranges,
but this orderly arrangement was usually confused, perhaps
because of additional targets. At close range the creaks were
timed to a horizontal sweeping of the head (nodding when on his
side). These observations may be interpreted as indicating direc-
tionality, presumably in his sound production. We have not
investigated this arresting possibility further, except to consider
that perhaps the pneumatic cephalic sinuses may modify the
radiation of sound from the larynx.
Keholoecation was evidently not a perfect method for our por-
poise. Perhaps the fault lay in the special conditions in the pond,
where the presence of multiple reflections from the stones in
the banks and bottom must have confused the echoes. The
primary target was a small fish; behind it was the punt, and
behind that was the shore. We noticed that when being fed from
the small dinghy (with so much less boat in the water to return
an echo), he ordinarily came right to the fish with less hesitation
than when feeding at the punt. These are indications that echolo-
cation did not give him clear and unequivocal information
(human users of this technique will sympathize).
ACKNOWLEDGMENTS
This work was supported by an Office of Naval Research con-
tract with the Woods Hole Oceanographic Institution. The por-
poise was generously supplied by Marine Studios, Marineland,
Florida. We are most grateful also to the Naushon Trustees for
hospitably permitting us to use and modify the pond and its
immediate surroundings. And of course we were greatly depend-
ent on the ever-ready support of many of our colleagues at Woods
Hole, without whose help the maintenance of the pond and
porpoise would have been impossible. We with to thank Drs.
R. H. Backus and C. P. Lyman and Prof. D. R. Griffin for
critically reading the manuscript.
REFERENCES
GRIFFIN, D. R.
1953. Acoustic orientation in the oil bird, Steatornis. Proc. Nat. Acad.
Sei., 39 (8): 884-893.
1956 FOOD-FINDING BY PORPOISE 15
GRIFFIN, D. R., and GALAMBOS, R.
1941. The sensory basis of obstacle avoidance by flying bats. Jour.
Exper. Zool., 86: 481-506.
Kewioaa, W. N., KoHLER, R., and Morris, H. N.
1953. Porpoise sounds as sonar signals. Science, 117 (3036) : 239-243.
KRITZLER, HENRY
1952. Observations on the pilot whale in captivity. Jour. Mamm. 33
(3) : 321-334.
McBripE, ARTHUR F.
(In epee Evidence for echolocation by cetaceans. Deep-Sea Research,
McBripe, A. F., and Hess, D. O.
1948. Behavior of the captive bottle-nose dolphin, Tursiops truncatus.
Jour. Compar. Physiol. Psychol., 41 (2): 111-123.
LAWRENCE, B., and SCHEVILL, W. E.
1954. Tursiops as an experimental subject. Jour. Mamm. 35 (2): 225—
232.
Woop, F. G., JR.
1952. Porpoise sounds. A phonograph record of underwater sounds
... Tursiops truncatus and Stenella plagiodon. Published by the
Marineland Research Laboratory.
Woop, F. G., JR. :
1954. Underwater sound production and concurrent behavior of captive
porpoises, Tursiops truncatus and Stenella plagiodon. Bull. Mar.
Sei. Gulf and Caribbean, 3 (2): 120-133.
BREVIORA
Museum of Comparative Zoology
~ June 6, 1956
CAMBRIDGE, Mass.
NUMBER 54
A REVISION OF THE GENUS BRACHYMELES
(SCINCIDAE), WITH DESCRIPTIONS OF
NEW SPECIES AND SUBSPECIES
By Watter C. Brown!
INTRODUCTION
The genus Brachymeles was erected by Duméril and Bibron
(1839) to accommodate a single species, B. bonitae, of this
unique group of Philippine skinks. Subsequently, B. bicolor
was described by Gray (1845) as the type of the genus Senira;
B. gracilis and B. schadenbergi were described by Fischer (1885) ;
and Taylor described B. elerae and B. burksi (1917), B. suluensis
and B. vermis (1918), B. boulengert (1922) and B. pathfinderi
and B. wright’ (1925). In 1922 Taylor correctly assigned B.
suluensis to the synonymy of B. gracilis and also pointed out that
B. boulengeri is closely related to the latter. It now appears that
B. boulengeri is probably best regarded as a geographically
isolated population which is only subspecifically differentiated
from B. gracilis. It differs only in minor characteristics from
the nominate form and does not overlap in range with any other
population of this species as it does with B. schadenbergi. A re-
examination of the type of B. bonitae, for which assistance I am
deeply indebted to Dr. Jean Guibé, indicates that B. burks? is
1 Menlo College and Natural History Museum, Stanford University
bo
BREVIORA No. 54
conspecific with it. Two additional species are described in the
present paper.
The members of this genus are secretive, burrowing lizards
that exhibit to a high degree elongation of the body and reduction
of limbs, eyes, and ears — specializations that in general charac-
terize other strictly burrowing skinks. The most highly special-
ized forms have rarely been encountered by field zoologists. As
a result, several of the species are represented by only a few
individuals and nothing of their inter-island variation is known,
assuming for the moment that their range, as in the instance of
B. bonitae, includes more than one island. Contrarily, at least
five of these highly specialized species are known only from type
localities, and the possibility also exists that many of these popu-
lations have very restricted ranges and are represented by rela-
tively small numbers. Substantiating evidence can be pointed
out in the instance of B. tridactylus. Four examples of this
species were collected in the mountains of southern Negros
Oriental, some 20 kilometers west of Bais, during a field period
of about two weeks in January and March, 1955, whereas no
specimens were uncovered in the Cuernos de Negros area of the
same mountain range, about 50 kilometers to the south, although
this area was much more thoroughly investigated over a nine
month period from July, 1954 to March, 1955. The author is of
the opinion that this species does not occur in the Cuernos de
Negros area. Other species, as B. gracilis and B. schadenbergi are
widespread within the archipelago, and sufficiently large samples
are available from several of the islands that well differentiated
populations can be recognized. These are regarded as distinct
subspecies.
No member of the genus has been recorded from outside of the
Philippine Archipelago. However, in view of its wide range
within this group of islands, including the Sulus and Palawan,
both of which are in close proximity to Borneo, it is not im-
probable that the genus may occur in North Borneo, at least.
Several of the more specialized species can be readily dis-
tinguished on the basis of the degree of reduction of the limbs
and elongation of the body. Greater difficulty exists in the recog-
nition of valid species and subspecies of the pentadactyl forms.
Part of the difficulty is due to the fact that certain of the differ-
entiating characters are not easily quantified, and other, fre-
1956 REVISION OF THE GENUS BRACHYMELES 3
quently used, key characters may hold for the separation of
populations of certain species on a given island, but have been
found not to be of value when one is concerned with populations
of the same species occurring on a different island. It is hoped
that the extent of our present knowledge of the variability of
these species may have overcome this difficulty.
In the course of the present study 266 specimens have been
examined. The name of the institution in whose collections
cited specimens are deposited is, in most instances, abbreviated
as follows:
C.A.S., California Academy of Sciences, San Francisco,
California
C.N.H.M., Chicago Natural History Museum, Chicago
T1linois
M.C.Z., Museum of Comparative Zoology, Cambridge, Massa-
chusetts
N.H.M.S.U., Natural History Museum of Stanford Univer-
sity, Stanford, California
S.U., Silliman University, Dumaguete City, Negros Oriental
SYSTEMATIC DISCUSSION
BRACHYMELES Duméril and Bibron
Brachymeles Duméril and Bibron, 1839, Erpt. Gen., 5: 776 (type species:
Brachymeles bonitae Duméril and Bibron, 1839, by monotypy).
Senira Gray, 1845, Cat. Lizards Brit. Mus.: 98 (type species: Senira bicolo)
Gray, 1945, by monotypy).
Brachymelus Agassiz, 1846, Nomen. Zool. Index Uniy.: 51 (emendation).
Diagnosis and definition. Pterygoid and palatine bones not in
contact mesially, both without teeth; maxillary and mandibular
teeth, moderate, conical ; body elongate ; limbs reduced or absent ;
eye small; lower eyelid scaly; external ear opening small or
absent.
The following key will serve to distinguish the known species
of the genus Brachynieles.
1. Limbs absent....:.. 2 qh: Meatidte Spc eee eee B. vermis
Limbs present .... a. Bg cont EE A Te Va bie Maes
2. Limbs NTE GAL Fpaaks Sao Ws tion MT. Me, Yee AS a Oe
Either fore or hind abe or both a ese fan 5 digits
Digits entirely absent or a single vestigial claw present .. B. bonitae
YG CO bo
Co
4 BREVIORA No. 54
Two or more digits present (frequently represented only as
Clawedeviestises)n= sen saree er eweny: gle te Baad ott oe eee 4
4. Limbs with 2 digits ......... ... B. samarensis sp. nov.
Timi swat hem onrest hanya Calo ts epee ie eee sine 5
5. Limbs with 3 digits ...... Kaas. Scat chee B. tridactylus sp. nov.
Limbs with more than 3 digits . RR Ne 5 OE se oc 6
6. Fore limbs with 5 digits; hind limbs with 4....._.... B. pathfinderi
Fore and hind limbs with 4 digits ................ 1 Me ae |
7. Midbody scale rows -22-; scale rows between the parietals and
thebaserotithertallSS-85> tae ocee tase rehe ee ce ere teaenene B. elerae
Midbody scale rows -28-; scale rows between the parietals and
the basorotswhe sbaill il OOS oa ee eee ere eee B. wrighti
8. Limbs short, length of the hind limb about 9 to 12 per cent of
the snout-vent length for mature individuals; number of scale
rows along the vertebral line between the parietals and the base
Othe talliQ2 OAs re ete ce ee eee een oe B. bicolor
Limbs moderate, length of the hind limb about 16 to 25 per cent
of the snout-vent length for mature individuals; number of scale
rows along the vertebral line between the parietals and the base of
the tail very rarely as great as 75 (generally 63 to 73) .......... 9
9. Size at maturity 75-129 mm. snout-vent length for 28 specimens;
supranasals large in contact (30 of 45 specimens examined) or
narrowly separated; number of midbody scale rows 26-32 (mean
= 27.9+ 0.265) for 49 specimens; number of middorsal scale
rows between the parietals and the base of the tail 67-73 (mean
= 69.1 + 0.214) for 49 specimens; venter and lower lateral sur-
faces very light with no or very few dark spotted scales. ......
B. schadenbergi
Size at maturity 57.5 to 95 mm. for 72 adult specimens; supra-
nasals moderate, generally rather widely separated, in contact in
only 2 out of 50 specimens examined; number of midbody scale
rows 24-28 (mean = 26.0 + 0.104) for 81 specimens; number of
middorsal scale rows between the parietals and the base of the
tail 63-69 (mean = 65.6 + 0.132) for 100 specimens; lower
lateral surfaces and frequently venter with numerous dark
Spotted scales (except in B. gracilis boulengeri) ........ B. gracilis
BRACHYMELES VERMIS Taylor
Brachymeles vermis Taylor, 1918, Philip. Journ. Sci., 138:255 — Bubuan Id.,
Sulu Archipelago.
Material examined. Jolo Id. 5 (C.A.S. 60720-22, 60857-58) ;
Bubuan Id., Sulu Arch. 1 (C.A.S. 62489). ;
1956 REVISION OF THE GENUS BRACHYMELES a
Diagnosis. Habitus very slender; snout to vent length 64-76
mm. for 3 mature specimens; limbs absent; no ear opening; no
postnasal; 22-24 seale rows around the middle of the body (6
specimens) ; 104-109 scale rows along the middorsal line between
the parietals and the base of the tail (6 specimens).
Range. Known from Bubuan, Jolo and Sulu Islands, Sulu
Archipelago.
BRACHYMELES BONITAE Duméril and Bibron
Brachymeles bonitae Duméril and Bibron, 1839, Erpet. Gen., 5:777 — Manila,
Luzon Island.
Brachymeles burksi Taylor, 1917, Philip. Journ. Sci., 12:275 — Sumagui,
Mindoro Island.
Material examined. Holotype (examined by Dr. Jean Guibé) ;
Mindoro Id. 3 (C.A.S. 62064, C.N.H.M. 22525, 8.U. R-20) ;
Luzon Id. 3 (C.A.S. 61376-77, 62578) ; Kalotkot Id. 2 (C.A.S.
60556-57) ; Polillo Id. 3 (C.A.8. 62278-79, 62575).
The single specimen (No. 1151, now C.A.S. 62578) referred to
B. bonitae by Taylor (1917, 1922), with which he compared B.
burkst when describing that species, is somewhat aberrant,
whether compared with the type of B. bonitae or B. burksi; it
differs primarily in the much greater number of scale rows along
the middorsal line between the parietals and the base of the tail,
113 instead of 100 to 106 for 11 specimens of B. bonitae (holo-
type 104). The fusion of the first pair of lower labials with the
mental, illustrated by Taylor, is characteristic of the type of
bonitae and also of 6 of the 9 specimens of B. burksi examined
for this character. Whether or not the greater number of mid-
dorsal scale rows is actually an individual aberration or char-
acterizes a more or less isolated population of B. bonitae cannot
be determined at this time.
Diagnosis. Habitus very slender; snout to vent length 57-82.5
mm. for 6 mature specimens; limbs reduced to vestiges without
evidence of toes or with a single claw, length of hind limb about
3.5 to 4.3 per cent of the snout-vent length (4 specimens) ; no
ear opening; no postnasal; 22-23 scale rows around the middle
of the body for 10 specimens; 100-106 scale rows along the mid-
dorsal line between the parietals and the base of the tail. One
specimen (C.A.S. 62578) from Los Banos, Luzon, is far out of
6 BREVIORA NO. 54
this range, exhibiting 113 middorsal scale rows, but as noted
above, is tentatively referred to this species.
Range. Known from Luzon, Mindoro, Polillo and Kalotkot
Islands in the northern part of the archipelago.
BRACHYMELES SAMARENSIS sp. nov.
Holotype. C.N.H.M., No. 44472, a juvenile, collected by G. N.
Rysgaard, at Guiuan, Samar Island, Philippine Islands, January
10, 1945.
Diagnosis. A slender Brachymeles with very short limbs, pos-
sessing only two reduced digits on both the fore and hind limbs;
midbody scale rows -22-; scale rows along the vertebral line be-
tween the parietals and the base of the tail -86-.
Description. A Brachymeles of very slender habitus; head
little wider than the body, tapering anteriorly to the rounded
snout; rostral large, in broad contact with the frontonasal ;
nostril in a small nasal; no postnasal; supranasals widely sep-
arated in the midline; prefrontals of moderate size, widely sep-
arated ; 5 supraoculars, anterior two in contact with the frontal ;
6 superciliaries; frontoparietals in contact; interparietal large,
round-pointed posteriorly; parietals in contact posterior to the
interparietal; a pair of nuchals which are narrower than the
parietals; two frenals, anterior slightly longer and wider than
the posterior ; first upper labial largest, fourth beneath the orbit ;
no external evidence of ear; number of scale rows around the
middle of the body -22-, number of scale rows along the vertebral
line between the parietals and the base of the tail -86-; limbs
very short, length of hind limb about one fifteenth the snout-
vent length; digits reduced to 2 clawed stumps on each foot.
Measurements of holotype. Snout to vent 43.5 mm.; length
of hind limb 3 mm.
Color (in preservative). Dorsal and lateral surfaces are dark
yellowish brown, each scale generally being dark spotted pos-
teriorly ; the venter is only slightly hghter.
Relationship. It would appear to be more closely related to
B. elerae than to B. bonitae for, although the extent of reduc-
tion of the limbs and digits is somewhat intermediate, the elonga-
tion of the body as measured by the number of middorsal scale
rows between the parietals and the base of the tail is essentially
1956 REVISION OF THE GENUS BRACHYMELES 7
the same as that of B. elerae, 86 as compared to 83-85 for 2
specimens of the latter. The number of scale rows between the
parietals and the base of the tail for 11 specimens of B. bonitae
is 100-106.
Range. Known only from the type locality.
BRACHYMELES TRIDACTYLUS sp. nov.
Holotype. N.H.M.S.U. No. 18354, a probably mature female,
collected 1 to 3 kilometers northwest of Mayaposi spring, about
20 kilometers west of Bais, Negros Oriental, at an altitude of
about 2000 feet on January 12 to 17, 1955, by Mr. Filomeno
Empeso.
\—
ieee
Fig. 1. Dorsal view of head of Brachymeles tridactylus.
Paratypes. N.H.M.S.U. Nos. 18355-56 and M.C.Z., No. 54258,
collected at the same locality as the holotype.
Diagnosis. A slender Brachymeles, with short limbs, possessing
three reduced, clawed digits on both the fore and hind lmbs;
scale rows 22-24; scale rows along the middorsal line between the
parietals and the base of the tail 95-103.
Description. A Brachymeles of very slender habitus; head
not or scarcely wider than the body, tapering anteriorly to a
narrow, truncate margin of contact with the frontonasal; nostril
in a minute nasal; supranasals not in contact in the midline for
any of the present series; prefrontals moderate, rather widely
separated; 4 supraoculars, anterior two in contact with the
frontal; 5 superciliaries; frontoparietals narrowly or moderately
8 BREVIORA No. 54
separated in the midline; interparietal large, rather pointed
posteriorly ; parietals meeting posterior to the interparietal; a
distinct pair of nuchals in only one specimen; two frenals, nearly
equal, or the anterior slightly shorter and wider; first upper
labial largest, fourth beneath the orbit; no external evidence of
ear; number of scale rows around the middle of the body 22 to
24; number of scale rows along the vertebral line between the
parietals and the base of the tail 95 to 103; limbs much redueed,
the length of the hind limb about one sixteenth to one twentieth
the snout-vent length; digits reduced to three clawed stumps on
each foot.
Measurements of holotype. Total length 106 mm.; snout to
vent 65 mm.; axilla to groin 49 mm.; snout to forelimb 13 mm. ;
leneth of hind limb 4 mm.
The largest specimen measures 78 mm. from snout to vent.
Color (freshly preserved material). Dorsal and lateral surfaces
are nearly chocolate brown to dark slate brown; the venter is
somewhat lighter in shade. Actually the basal part and center
of each scale is pigmented, the overlapping lateral and distal
edges are not.
Relationship. This species is apparently most closely related
to B. bonitae from which it differs primarily in less reduced
limbs and the presence of three stump-like, clawed digits on each
limb. B. bonitae has lost all the digits or retained a vestige of
only one at the tips of the minute limbs.
Range. Known only from the type locality.
BRACHYMELES ELERAE Taylor
Brachymeles elerae Taylor, 1917, Philip. Journ. Sci., 12:273 — Philippine
Tslands.
Material examined. Luzon Id. 2 (C.A.S. 61499-500).
Diagnosis. Habitus very slender; snout to vent length 69-71
mm. for 2 mature specimens; limbs greatly reduced with vestiges
of 4 clawed toes on both fore and hind limbs; length of hind
limb 7.2 to 9.3 per cent of the snout-vent length (4 specimens) ;
no ear opening ; no postnasal ; -22- scale rows around the middle
of the body; 83-85 scale rows along the middorsal line between
the parietals and the base of the tail.
Range. Known definitely from Balbalan, Mountain Provinee,
Luzon Island.
1956 REVISION OF THE GENUS BRACHYMELES 9
BRACHYMELES WRIGHT! Taylor
Brachymeles wrighti Taylor, 1925, Philip. Journ. Sci., 26:106 — Trinidad,
northern Luzon Island.
Material examined. Holotype (M.C.Z. 26589: examined by Mr.
Arthur Loveridge).
Diagnosis. Habitus slender; snout-vent length 180 mm. for
one specimen; limbs greatly reduced with 4 clawed toes on both
fore and hind limbs, length of hind limb 7.5 per cent snout-vent
length (1 specimen) ; ear opening questionable because of injury ;
no postnasal; -28- scale rows around the middle of the body;
-102- scale rows along the middorsal line between the parietals
and the base of the tail.
Range. Known only from the mountains of northern Luzon
Island.
BRACHYMELES PATHFINDERI Taylor
Brachymeles pathfindert Taylor, 1925, Philip. Journ. Sci., 26:104 — Glan,
Cotabato Province, Mindanao.
Material examined. Holotype (M.C.Z. 26581: examined by Mr.
Arthur Loveridge) ; Mindanao 2 (M.C.Z. 26582-83).
Diagnosis. Habitus slender; snout to vent length 58-61 mm.
for 3 mature specimens; limbs greatly reduced with vestiges of
five clawed toes on the fore limbs and four on the hind limbs,
length of hind limb 13.6-17.2 per cent of snout-vent length (3
specimens) ; ear opening minute; no postnasal ; 22-23 scale rows
around the middle of the body ; 59-67 scale rows along the ver-
tebral line between the parietals and the base of the tail.
Range. Known only from the type locality.
BRACHYMELES BICOLOR (Gray )
Senira bicolor (part), Gray, 1845, Cat. Lizards Brit. Mus.: 98 — Philippine
Islands.
Material examined. Uolotype and one additional specimen
(examined by Mr. J. C. Battersby).
Diagnosis. Habitus slender; snout to vent length 155 mm. for
2 mature specimens; limbs reduced with 5 clawed toes present on
each foot, length of hind limb 9.7 to 11 per cent of snout-vent
\
length (2 specimens) ; -28- rows of scales around the middle of
10 BREVIORA No. 54
the body; 92-94 rows of scale along the vertebral line between
the parietals and the base of the tail.
Range. Harly records are given as from the Philippine
Islands. More definite information as to the distribution of this
species can not be stated at this time.
BRACHYMELES GRACILIS (Fischer )
Several populations of this species exhibit rather conspicuous
and constant differences in color pattern and in certain instances
in sealation or other characters. The Negros population does
not exhibit the narrow, dorso-lateral light stripe which in gen-
eral characterizes other known populations. The venter and
lower lateral surfaces of the Polillo population are very much
lighter in color than is true of the Mindanao-Sulu population.
The dark spotting of the venter is somewhat intermediate for
examples from the Negros and Bohol. The postnasal is small and
does not contact the second upper labial in the Mindanao-Sulu
population, but is larger and in contact with the second upper
labial in most examples of the other known populations. Also as
pointed out by Taylor (1922), the ear opening is better developed
in B. g. boulengert than in B. g. gracilis.
The population from Bohol Island is in closer agreement with
regard to color pattern, distinctness of the dorso-lateral stripe
and the size of the postnasal shield with the population of the
northern islands (Luzon and Polillo) than it is with that of
Mindanao to the south, and is referred to the subspecies bouw-
lengert Taylor which was described from Polillo Island. If this
interpretation is correct, intervening islands of Leyte and Samar
are in all probability occupied by this subspecies. The differ-
ences in the number of midbody scale rows and the number of
middorsal scale rows between the parietals and the base of the
tail exhibited by populations of different islands are not sig-
nificant (Table 1).
BRACHYMELES GRACILIS GRACILIS (Fischer)
Eumeces (Riopa) gracilis Fischer, 1885, Jahrb. wiss. Anst. Hamburg, 2: 85 —
Mindanao Island. ;
Brachymeles suluensis Taylor, 1918, Philip. Journ. Sei., 13:254 — Bubuan
Island, Sulu Archipelago.
a oorererererererercreaeee_ES_ee _E_ EO
1956 REVISION OF THE GENUS BRACHYMELES aha
Material examined. Basilan Id. 2 (C.A.S. 60365-66) ; Minda-
nao Id. 32 (N.H.M.S.U. 18596; C.A.S. 15567; C.N.H.M. 52637,
52642-52, 52654-70, 52800).
Diagnosis. Habitus rather slender ; snout to vent length 57.5-82
mm. for 10 mature specimens ; length of hind limb 15.3 to 19.2 per
cent of the snout to vent length for 10 adults ;"postnasal short,
not in contact with the second upper labial in any of 30 speci-
mens examined for this character; 24-26 scale rows around the
middle of the body for 36 specimens; 64-69 scale rows along the
middorsal line between the parietals and the base of the tail for
36 specimens ; narrow dorsolateral light stripe generally present,
extending anteriorly to the posterior edge of the orbit and
posteriorly to the groin or nearly so; lower lateral surfaces dark
spotted; midventral region distinctly dark spotted (24 out of
30 specimens), the dark spots generally occupying the basal
and central portion of each seale.
Range. Known definitely from the Sulu Arehipelago and
Mindanao.
SRACHYMELES GRACILIS BOULENGERI Taylor
Brachymeles boulengert (part) Taylor, 1922, Philip. Bur. Sei. Pub. No.
17:246 — Polillo Island.
Material examined. Polillo Id. 6 (C.A.S. 62272-77) ; Luzon Id.
2 (C-A:S* 61096-97) ; Bohol Id. 24 (S.U. R-353; N.H.M-S.U,
18271-76, 18707-11, 18714-20; M.C.Z. 54252-3).
Diagnosis. Habitus rather slender; snout to vent length 61-92
mm. for 19 mature specimens; length of the hind limb 17.9-23.8
per cent of snout-vent length for 19 mature specimens; postnasal
long, in contact with the second supralabial in 24 of 26 specimens
examined ; 24-28 scale rows around the middle of the body (mean
= 26.1 + 0.175) for 27 specimens; 63-66 scale rows along the
middorsal line between the parietals and the base of the tail
(mean = 64.2 + 0.198) for 27 specimens; narrow dorso-lateral
stripe (in life near to Monkey skin or Vassar tan, Maerz and
Paul, 1930, pls. 6 and 10) generally present (27 out of 28
specimens), extending anteriorly onto the supraorbital region
and posteriorly to the groin (this stripe is generally more dis-
tinct than in B. g. gracilis) ; lower lateral surfaces dark spotted ;
venter with moderate dark spotting or with such spotting absent.
12
BREVIORA
Table 1
Comparison of subspecies of Brachymeles gracilis
No. 54
B. g. gracilis
(Mindanao and
Basilan )
B. g. boulengeri
(Polillo)
B.g. boulengeri
(Luzon)
B. g. boulengeri (°)
(Bohol)
B. g. taylori
(Negros)
Bb. g. taylori
(Mindoro )
No. of midbody
scale rows
R = 24-26
WSS PO Se OLD
N = 36
R = 24-26
M = 25.7 5 0.305
IN-==.6
Ra 26026
INS 2
R = 24-28
M = 26.2 = 0.158
Ne 2
Dorsal seale rows
between parietals
and base of tail
R = 64-69
M = 66.4 + 0.201
Ni 36
R = 64-66
Me 6522/2103 67,
INE ==6
Ri = 63,65
NGS=2,
R = 63-66
M = 64.0 + 0.199
IN = 2 ill
R = 64-68
M = 65.6 = 0.150
N = 49
Ri == 66
N=)
Dorso-lateral
stripe present
Yes
Yes
No
No
R=range, M—=mean, N = number of specimens
Color (freshly preserved material). The following notes were
based on material which had been in preservative only a day
or two. Six to 8 middorsal scale rows are Natal Brown to
Chestnut (Maerz and Paul, pl. 7), not uniform but each seale
with a darker brown blotch at the base. A dorso-lateral stripe
(generally one and a half scale rows in width anteriorly and up
to one and two half scale rows posteriorly) is somewhat variable
usually near Monkey skin or Army brown (Maerz and Paul, pls.
5 and 6). Five to 7 lateral rows exhibit prominent, dark brown
spots; the venter is Rose tan (Maerz and Paul, pl. 12), oceasion-
ally with small brownish spots. Four or 5 rows on the chin,
posterior to the mental, are dusky or brownish.
1956 REVISION OF THE GENUS BRACHYMELES 13
Range. Known from Polillo, Luzon and Bohol Islands. The
latter population is assigned to this subspecies with some reser-
vations, pending the time when collections are available for com-
parison from the islands of Samar and Leyte.
BRACHYMELES GRACILIS TAYLORI subsp. nov.
Brachymeles boulengeri (part) Taylor, 1922, Philip. Bur. Sei. Pub. No.
17:246.
Holotype. N.H.M.S.U., No. 18615, a male, collected by W. C.
Brown and D. 8. Rabor on low ridge on the north side of the
Maite River, about 13 kilometers west of Dumaguete on Cuernos
de Negros, Negros Oriental, July 4, 1954.
Paratypes. Negros Id. 126 (N.H.M.S.U. 17954-55, 18611-14,
18616-24, 18626-28, 18630-31, 18633-37, 18664-80, 18694-700, low
Fig. 2. Dorsal view of head of Brachymeles gracilis taylor.
ridge north of the Maite River, Cuernos de Negros, about 3-7
km. west of Luzuriaga, Negros Oriental, July, 1954 to February,
1955; 18597-18604, 18659-63, high ridge north of Maite River
and on north slope of north peak of Cuernos de Negros, about
6-8 km. west of Luzuriaga, Negros Oriental, February and
March, 1955; 18638-58, 18691-93, 18728, 18748-50, ridge, south
of Maite River, east slope of Cuernos de Negros, about 4 to 7
km. west of Luzuriaga, Negros Oriental, November, 1954, to
March, 1955; 18686-88, east side of Cuernos de Negros, Negros
Oriental; 12027, 12224, 12226, Dumaguete environs, Negros Ori-
14 BREVIORA No. 54
ental, August, 1940; 18751, 18729-32, lowlands 2 to 6 km. west
of Dumaguete, Negros Oriental, July and August, 1954; 18689,
14 km. 8.W. of Palimpinon, Ocoy River Valley, Negros Oriental,
August, 1954; 18605, 18681-85, 18690, 18752, Mayaposi environs
and hills to the north and west of Mayaposi, about 20-30 km.
west of Bais, Negros Oriental, January to March, 1955; 18727,
Ilaya sitio, 3-4 km. south of Caliling, Negros Occidental, Decem-
ber, 1954; S.U. R-117-18, 242-44, 246, 251, 262, 267, 272-73, 275-
78, Cuernos de Negros area, Negros Oriental; M.C.Z. 54254-6,
low ridge north of Maite River, Cuernos de Negros, 3 to 7 km.
west of Luzuriaga, Negros Oriental, 1954.
Diagnosis. Habitus rather slender; snout to vent length 62-95
mm. for 32 mature specimens; length of hind limb 16 to 22.5 per
cent of snout-vent length for 28 mature specimens; postnasal
long, in contact with the second supralabial in 37 of 45 speci-
mens examined for this characteristic ; 24-28 scale rows around
the middle of the body for 49 specimens; 64-68 scale rows along
the middorsal line between the parietals and the base of the tail
for 49 specimens; dorsolateral light stripe not present; lower
lateral surfaces and generally venter dark spotted.
Color (freshly preserved material). The dorsum and the
upper lateral surfaces vary from light to dark brown, forming a
pattern of dark and light, narrow, longitudinal stripes or nearly
uniform. The lower lateral surfaces vary from near Rosetan to
Blush or darker (Maerz and Paul, pl. 12), frequently with secat-
tered to numerous small brown spots.
Range. Known from Negros Island and probably Mindoro.
BRACHYMELES SCHADENBERGI Fischer
The Negros population exhibits constant differences in certain
characteristics for a sufficiently large series of specimens to jus-
tify its recognition as a subspecies which is distinct from the
typical one of Mindanao Island. The 3 available examples from
Jolo and 2 specimens from the mountains of northern Luzon are
close to the Negros population in the number of middorsal seale
rows between the parietals and the base of the tail, and the
presence of a more or less distinet, dorsolateral light stripe, and
are tentatively referred to this subspecies. The single specimen
from Palawan (C.A.S. 15571) lacks the light stripe and would
1956 REVISION OF THE GENUS BRACHYMELES 15
appear to agree most closely with the Mindanao population.
[f larger samples do indeed prove the Luzon, Negros and Jolo
populations to be the same from a taxonomie standpoint, and
that of Palawan, Basilan, Mindanao, Leyte and Bohol to be
taxonomically identical, this distributional pattern, which is not
wholly in accordance with logical routes of dispersal in terms of
what is known of the geological history of the archipelago, sug-
gests three possible interpretations: (1) chance colonization of
widely separated islands, across intervening sea barriers, by two
distinct subspecies; (2) more or less isolated populations of a
polytypie species exhibiting chance convergence with respect
to the presence or absence of a dorsolateral stripe; (3) two
population groups of sibling species, very similar morphologi-
eally, which have maintained disjunct distributions within this
archipelago. The first interpretation is accepted with some reser-
vations in this paper.
3RACHYMELES SCHADENBERGI SCHADENBERGI (Fischer )
Lumeces (Riopa) schadenbergi Fischer, 1885, Jahrb. wiss. Anst. Hamburg, 2:
87 — Mindanao Island.
Material examined. Basilan Id. 15 (C.A.S. 60305-10, 60312,
60438-40, 60493-98) ; Mindanao Id. 7 (C.N.H.M. 22528-29, 52638-
41, 61963-64) ; Leyte Id. 3 (N.H.M.S.U. 18701; C.N.H.M. 42779,
42792); Bohol Id. 5 (N.H.M.S.U. 18702-06); Palawan Id. 1
(C.A.S. 15571).
Diagnosis. A moderately large Brachymeles with rather well
developed limbs; number of midbody scale rows 26-28 (mean =
26.5 = 0.161 for 25 specimens) ; a light dorso-lateral stripe ab-
sent; the second pair of chin shields separated by three scales.
Color (freshly preserved material from Bohol Island). Dor-
sum and upper surface of limbs are blackish brown; lateral
surfaces are near Melon, Baby rose, Crabapple, Burmese gold
or Burnt orange (Maerz and Paul, pls. 2 and 3). The venter is
whitish, lightly tinted with the above shades.
Range. Known from the Sulu Archipelago, Mindanao, Leyte
and Bohol Islands (probably also Palawan Island).
16 BREVIORA NO. 54
Table 2
Comparison of subspecies of Brachymeles schadenbergi
Dorsal scale rows
No. of midbody between parietals Dorso-lateral
scale rows and base of tail stripe present
B.s. schadenbergi R = 26-28 R = 67-71 No
(Mindanao and M= 26.5 +0161 M=68.8 = 0.250
Basilan ) N95 ING ==25
B. s. schadenbergi R = 28 R = 68,68,70 No
(Leyte) N =3 N=3
B. s. schadenbergi eS R= 1027 C4) No
(Bohol) N15 Ne
B. s. schadenbergi (2?) R = 28 Ri ==70 No
(Palawan ) N= Nis
B.s. talinis R = 28-32 R= 67-70 Yes
(Negros) N= 3 OF S=1022, 716 M6 822101193
N= ING 2
B. s. talinis (2?) R= 29:30532 R 72;72;73 Yes
(Jolo) NG 3 N-=.3
B. s. talinis (2) R = 26,28 R — 69,70 Yes
(Luzon) Ni 2 N ==(2
R=range, M=mean, N = number of specimens
BRACHYMELES SCHADENBERGI TALINIS subsp. noy,
Holotype. N.H.M.S.U. No. 18358, a female, collected by F.
Empeso, December 14, 1954 on the low ridge north side of the
Maite River, 5 to 6 km. west of Luzuriaga, Negros Oriental.
Paratypes. N.H.M.S.U. Nos. 12225 collected Dumaguete en-
virons, Negros Oriental, 1940; 18359, low ridge north of the
Maite River, Cuernos de Negros, about 5-6 km. west of Luzuriaga,
Negros Oriental, Dec., 1954; 18363-64, 18366, high ridge north of
the Maite River and on north slope of north peak of Cuernos
de Negros, about 6-8 km. west of Luzuriaga, Negros Oriental,
Dee., 1953, March, 1955; 18360-62, ridge south of the Maite
1956 REVISION OF THE GENUS BRACHYMELES V7
River, east slope of Cuernos de Negros, about 6-7 km. west of
Luzuriaga, Negros Oriental, Nov., 1954, March, 1955; M.C.Z.
94257, high ridge north of Maite River, Cuernos de Negros,
about 6-8 km. west of Luzuriaga, Negros Oriental; S.U. R-261,
ridge south of the Maite River, east slope of Cuernos de Negros,
about 6-7 km. west of Luzuriaga, Negros Oriental; C.N.H.M.
22527, Mt. Canlaon, Negros Oriental.
Diagnosis. A large Brachymeles with rather well developed
limbs, differing from the typical subspecies in the generally
greater number of midbody scale rows 28-32 (mean = 30.4 +
0.670 for 11 specimens) ; a moderately wide (two and two half
seale rows at the shoulder), dorso-lateral light stripe, reddish in
life, sometimes fading in older specimens; second pair of chin
shields generally wider, separated by 1 scale in 8 specimens, 2
in 1 specimen and 3 in 2 specimens as compared to 3 in a sample
of 23 specimens of the nominate subspecies examined for this
character,
Fig. 3. Dorsal view of head of Brachymeles schadenbergi talinis.
Color (freshly preserved material). Dorsum is dark reddish
brown, the lateral margins of the scales tending to be dull red-
dish producing an indistinct, ined pattern. This dorsal brown
band is rather uniformly six and two half scale rows in width
and is bordered on either side by a dull red stripe one and two
half rows in width. (These fade to white or whitish tan in
preservative.) These stripes begin just posterior to the eye and
extend to the region of the hind limbs or the base of the tail.
18 BREVIORA No. 54
They are most uniform and conspicuous in juveniles, tending to
bécome somewhat paler and irregular, especially on the posterior
part of the body for some adults. The red stripes are bordered
ventrally by one to three or four rows of brownish or brown
spotted scales. Lower lateral surfaces grayish white to light
tan, frequently diffused with orange-yellow and with scattered
brown spots especially in the midventral line and the region of
the hind limbs. The labials are brownish ; mental, rostral, nasals,
supranasals and postnasals grayish.
Range. Known definitely at the present time from Negros
Island (probably also Luzon and Jolo Islands, see p. 14)
The name talinis in the native dialect refers to mountains such
as Cuernos de Negros which this subspecies inhabits.
ACKNOWLEDGMENTS
I am greatly indebted to Dr. Jean Guibé, Muséum National d’Histoire
Naturelle, Paris, and Mr. Arthur Loveridge, Museum of Comparative Zool-
ogy, Harvard University, and Mr. J. C. Battersby, British Museum (Natural
History) for their kindly assistance in re-examining the types of Brachy-
meles bonitae, Brachymeles wrighti, Brachymeles pathfinderi and Brachymeles
bicolor; to Dr. Robert F. Inger, Chicago Natural History Museum and Dr.
Joseph R. Slevin, California Academy of Sciences for permitting me to
examine material of this genus in the collections of their respective institu-
tions. I also wish to thank Dr. Inger for his kindness in suggesting that I
describe at this time a new species from Samar Island which is represented
by a unique specimen in the collections of the Chicago Natural History
Museum.
This study was begun while the author was serving as a Fulbright lec-
turer at Silliman University, in the Philippine Islands, under the auspices
of the United States Educational Foundation, and it was completed while
working as a research associate at Stanford University supported by a grant
from the National Science Foundation. Illustrations were prepared by Mr.
Walter Zawojski, Stanford Research Institute.
LITERATURE CITED
BOULENGER, GEORGE ALBERT
1887. Catalogue of the lizards in the British Museum. Ed. 2. London,
vol. 3, i-xii + 575 pp., pls. 1-40.
DUMERIL, ANDRE MARIE CONSTANT, and GABRIEL BIBRON
1839. Erpetologie general. Paris, vol. 5, i-vili + 855 pp., pls. 43-58.
1956 REVISION OF THE GENUS BRACHYMELES 19 |
FISCHER, JOHANN GUSTAV
1885. Ichthyologische und herpetologische Bemerkungen. Part IV:
Uber eine Kollection von Amphibien und Reptilien von Mindanao.
Jahrb. wiss. Anst., Hamburg, vol. 2, pp. 80-81.
GRAY, JOHN EDWARD
1845. Catalogue of the specimens of lizards in the collection of the
British Museum. London, i-xviii + 289 pp.
MAeERZ, A. and M. REA PAuL
1930. <A dictionary of color. New York, MeGraw Hill Book Co., i-vii +
207 pp., pls. 1-56.
TAYLOR, EDWARD HARRISON
1917. Brachymeles, a genus of Philippine lizards. Phil. Journ. Sci.,
vol. 12, pp. 267-277, pl. 1.
1918. Reptiles of the Sulu Archipelago. Phil. Journ. Sci., vol. 13, pp.
233-267, pls. 1-3.
1922. The lizards of the Philippine Islands. Publ. 17, Phil. Bur. Sci.
Manila, pp. 1-269, pls. 1-23.
1925. Additions to the herpetological fauna of the Philippines, IV.
Phil. Journ. Sci., vol. 26, pp. 97-111.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. JUNE 6, 1956 NUMBER 55
REDISCOVERY OF HYLA DORSALIS
AND LECHRIODUS PAPUANUS
IN NEW GUINEA
By ARTHUR LOVERIDGE
Last year I had occasion to describe a new microhylid discov-
ered by Mr. Kenneth R. Slater at Omati, Papua. More recently
we have received part of a collection made by Mr. Slater during
a trip up the Aramia River. This river les somewhat to the
north of Katow (i.e. Binturi River, Papua), a locality from
which Macleay (1877, Proe. Linn. Soe. New South Wales, 2, pp.
135-138) described a number of amphibians, among them Litoria
dorsalis Macleay, now known as:
Hyua porsauis (Macleay)
This hylid, so far as I know, has not been taken again during
the period of almost eighty years that has since elapsed. Among
the Slater material is a 20 mm. gravid 2° Hyla (M.C.Z. 28389)
that agrees so closely with Macleay’s brief description of the
holotype of dorsalis (whose length is given as ‘‘9 lines,’’ i.e. 19
mm., its hind limb as ‘‘16 hnes,’’ i.e. 33.8 mm. as against 35
mm. in ours) that I have no hesitation in assigning our frog to
dorsalis, which must be one of the smallest members of the
genus Hyla.
Slater’s specimen differs only in the absence of a light verte-
bral lLne from snout to anus. On either eyelid are two tiny
tubercles that, if constant in the species, may well have been
overlooked by Macleay.
i)
BREVIORA NO. 55
LECHRIODUS PAPUANUS (Roux)
The same collection contained half a dozen frogs (M.C.Z.
28382-7) bearing a distinct resemblance to Cornufer, but I am
indebted to Dr. L. D. Brongersma of Leiden for pointing out
that they have arciferal shoulder girdles and more or less dilated
sacral diapophyses, so that he would refer them to Lechriodus
papuanus (Roux) which was described in 1927 from a single
juvenile ?, 23 mm. in leneth, taken near Lake Sentani on the
northern coast of Netherlands New Guinea. In the thirty years
that have intervened since the holotype was taken by Dr. P.
Wirz, no second specimen has been recorded so far as I know.
No material was available to Parker when he monographed
the Leptodactylidae of Australasia (1940, Novit. Zool., 42, pp.
1-106) and he had but two examples of L. fletcheri (Boulenger)
of which Roux made papuanus a subspecies. Parker (pp. 25, 27)
treated papuanus as a full species but the dermal skinfolds by
which he separates the two forms are so highly variable that it
is questionable if they really do so. If Brongersma and I are
correct in assigning these frogs to papuanus, the 7 -shaped
interorbital fold figured by Roux is either irregularly transverse,
more often \_y/-shaped; the ‘‘curved, convergent dorso-lateral
folds’’ are often rather ill-defined and may be represented by a
few odd plicae extending no further than the scapula region.
In only one frog do they approach the distinctness suggested by
Roux’s figure; they are quite indistinguishable from our
fletcheri 2 2 from New South Wales in this respect. As I find
little but color to separate the two, it would seem that Roux
was justified in regarding papuanus as only a race of fletcheri.
If I were called upon to distinguish the two forms I would do
so as follows:
Color above pale brown with traces of a blackish brown line
from end of snout along the canthus to the eye and from eye
over tympanum to a point above elbow; lower lip dark
brown; no interorbital skin fold (on either of our 9 2 from
New South Wales). Range: New South Wales; Queensland ;
and (fide Parker) British New Guinea. .... . ... /f. fletcheri
Color above light gray, a white, finely black-edged, shieldlike
patch on snout which may merge with two similar, but
smaller, anteorbital patches; from eye over tympanum to a
1956 HYLA DORSALIS AND LECHRIODUS PAPUANUS 3
point above elbow a broad or narrow blackish brown line
which may break up into a series of black specks; lower lip
white; an indistinct interorbital skin fold (4 ¢¢, 2 92°
from Aramia River). Range: British and Dutch New Guinea Sy Los
f. papuanus
Our knowledge of f. papuanus, hitherto derived from a single
immature 2, is naturally extended by the Aramia River series
of adult ¢ ¢ and @ ?; on the latter the following description
is exclusively based.
Description. Vomerine teeth in two confluent or semiconfluent
series extending in an almost unbroken undulating line right
across the roof of the mouth behind the level of, and beyond, the
choanae. Head slightly broader than long, i.e. the breadth equals
the distance from end of snout to angle of jaw; snout oval to
obtusely rounded; nostril slightly nearer the front of the lip
than it is to the orbit; interorbital space narrower than an
upper eyelid ; upper eyelid with several small, sometimes scarcely
noticeable, tubercles ; a rather indistinct, transverse or\__/shaped
interorbital fold; from the eye a supratympanie fold, which may
be broken up, extends posteriorly to a point above the shoulder ;
tympanum vertically oval, its horizontal diameter about two-
thirds that of the eye; from behind the eyelids a pair of more
or less )(-shaped glandular skin folds (frequently reduced to a
pair of short lines) extend to, or beyond, the scapular region
of the otherwise largely smooth dorsum.
Fingers free, their tips somewhat swollen but not dilated,
without horizontal grooves, first and second subequal in all six
specimens; subarticular tubercles large, metacarpus with a large
supernumerary tubercle; toes webbed, from first to fifth the
following phalanges free (except for a lateral seam) 2, 2, 3, 4,
214, their tips shghtly dilated and with or without a circum-
marginal groove, third toe longer than the fifth; subarticular
tubercles prominent, a conspicuous inner metatarsal two-thirds
the length of the first toe, no outer metatarsal tubercle ; tibio-
tarsal articulation of an adpressed hindlimb extends almost to
the nostril, or to the nostril, or end of snout. Skin of throat
and belly smooth.
4 BREVIORA NO. 55
Color. Above, tip of snout with a conspicuous white, finely
black-edged, shieldlike patch that may be disconnected with a
similarly light patch in front of each eye, or fork posteriorly and
merge with the anteorbital patches; from eye towards flank a
more or less conspicuous black line which breaks up into black
flecks on the flank immediately posterior to the forearm; dorsum
pale gray, with or without a few fine black lines; supra-anal
area pale; circum-anal area black, becoming lighter on the’ but-
tocks toward the tibio-tarsal joint; fore- and hindlimb more or
less faintly crossbarred. Below, white, uniform except that the
chin and jaws are grayish in é6 6, while in both sexes the
lower posterior aspect of forearms, anterior aspect of tibia, and
soles of feet, are black.
Size. Length from snout to anus of largest ¢ (M.C.Z. 28385),
49 mm., of @ (M.C.Z. 28382), 52 mm.; length of head from tip
of snout to back of tympanum in ¢, 18 mm.; in ¢?, 20 mm.;
length of hind limb from anus to tip of longest toe in ¢, 77
mm.; in @, 86 mm. In size, therefore, papuanus differs little
from fletchere.
Stomach contents. That of the ¢ was found by my colleague
Dr. W. L. Brown to consist of a chrysomelid beetle, a termite’s
head, and remains of ants referable to the genus Odontomachus.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. JUNE 15, 1956 NuMBER 56
NOTES ON THE JAMAICAN AND CAYMAN
ISLAND LIZARDS OF THE
GENUS CELESTUS
By Penny NORSEEN COUSENS
The collections of the Museum of Comparative Zoology afford
a representative sample of the Celesti of Jamaica and the Cay-
man Islands! and, additionally, include types of 5 of the 8 forms
recognized in the last revisions of Chapman Grant (1940b and c).
Renewed examination of this material has confirmed the distinct-
ness of certain forms but has placed in question the status of
certain others. It is hoped to return to the study of this genus
in these and other islands in the future but as an aid to students
of this difficult group it has been thought desirable to place on
record at this time the new conclusions as to the status of the
Jamaican and Cayman forms.
The characters studied were: habitus; color pattern; dorsal,
ventral and midbody seale counts, numbers of chin shields, and
number of subdigital lamellae on the fourth digit ; general body
measurements; presence or absence of a keel and number of
striae on dorsal scales. Head scalation was checked but not
recorded on at least one individual of each species. No internal
characters were used. It was impossible to sex the specimens
except in the case of females carrying young, but it is doubtful
if there is any sexual dimorphism in the species studied.
Only four species appear to be valid, as follows:
1 One species, the giant form Celestus occiduwus Shaw, is unrepresented. How-
ever, omission of this species from the present discussion appears to be justified
in as much as its right to consideration as a still surviving form is very improb-
able. It has not been collected in over 100 years. Grant’s suggestion that
hewardii is a subspecies of this form cannot be seriously regarded.
2 BREVIORA NO. 56
Key to the Jamaican and Cayman forms of Celestus:
1. Adpressed limbs meet or fail to meet by less than length of fore-
arm. Dorsal color pattern consisting of broken bars or rings 2
Adpressed limbs fail to meet by greater than length of fore-
arm. Dorsal color pattern not ringed .. é ; a 3
2. 19 or fewer subdigital lamellae on fourth toe of hind foot.
Throat mottled . ee ee Oe .......... hewardti Gray 1845
Range: Central and western Jamaica and Portland Point.
21 or more subdigital lamellae on fourth toe of hind foot.
Throat not mottled .......... ca _.. duquesneyi Grant 1940a
Range: Portland Point only.
Co
89-116 dorsal scales, occipital to point above vent .............. 4
125-139 doreal seales, occipital to point above vent ..............
tange: known only from Mandeville in south central Jamaica
and Fern Gulley in north central Jamaica. : ,
4. Tail dorsally with the same coror and pattern as back ............
BS Ae HS ate MRS a errr eee Meer eer Carte een crusculus crusculus Garman 1187
Range: all of Jamaica, at least in the lowlands.
Tail dorsally lighter than back and without pattern ..............
ees ane A ENE CR CSR SLOT: 6 Cheon Ea ena crusculus maculatus Garman 1888
Range: Cayman Brae and Little Cayman.
DISCUSSION
As the key indicates, and as Grant (1940b) has previously
emphasized, the four species fall into two groups — one long-
legged, the other short-legged. This may be a quite natural
erouping, in these islands at least.
In the long-legged group the two species, although similar in
general body proportions, differ in several characters. The rela-
tively widespread C. hewardii is a stockier, blunt-snouted lizard,
its dorsum with dark brown broken bars on a light brown back-
ground, its throat mottled, 12-19 lamellae on fourth toe, 4 pairs
of chin-shields. C. duquesneyi, confined to Portland Ridge, is a
more slender lizard with a pointed snout, its dorsal pattern similar
to hewardii but lighter (specimens collected in same year), back-
ground tan with light brown bars, the throat immaculate, 21-22
lamellae on fourth toe, 5-7 pairs of chin-shields. Unquestionably
C. duquesneyi is a distinct species. M.C.Z. 45181 from Portland
1956 JAMAICAN AND CAYMAN CELESTUS 3
Ridge was misidentified as hewardw but proved easily separable
from a hewardu with the same locality data despite the loss in
45181 of the blue tail so much stressed by Grant as diagnostic
for duquesneyt.
In the second, short-legged group the series of C. barbouri
studied included the type and 13 specimens collected by Bar-
bour,’ all from Mandeville, plus a single specimen from Fern
Gulley. One character, dorsal scale count, readily separates C.
barbourt from C. crusculus.2 This difference is very marked and
shows no overlap. Crusculus has the lowest dorsal scale count of
any Jamaican Celestus (mean 101.52, range 89-116); barbouri
has the highest (mean 132.27, range 127-139). At Mandeville,
where the two species are sympatric, crusculus shows a very low
mean of 97.64 (range 96-101) and also the strongest divergence
of color pattern. The dorsal pattern of barbouri consists of a
series of dark brown chevrons with the V’s pointing caudad.
Crusculus shows a variety of patterns: plain tan dorsum with
dark brown lateral bands extending the length of the body; a
checkerboard dorsal pattern; checkerboard blending into chev-
rons; chevrons with lateral bands; chevrons only, this pattern
almost indistinguishable from barbouri in several specimens. At
Mandeville, the last type does not occur and the chevrons-with-
lateral-stripes pattern occurs only once. These pattern types were
established to aid in grouping specimens — gradations and modi-
fications are common.”
C. crusculus 1s a variable species also in dorsal, ventral, and
midbody seale counts. There is, however, no correlation between
scale counts and pattern, and the two Jamaican subspecies pro-
posed by Grant (1940a) on such supposed correlations are there-
fore here regarded as invalid, C.c. cundalli with the type locality
Mandeville being untenable on any ground. The Mandeville
population cannot be distinguished from that at Kingston, the
1 These were labeled by him as impressus Cope. ‘The status of this name is very
much in doubt and its disposition will depend on a new evaluation of the types
in the Philadelphia Academy.
2 Ventral seale counts are, as usual, fully parallel to dorsal scale counts and
not really an independent variable.
3 Another color difference has been pointed out to me by Garth Underwood.
The side of the neck in barbouri bears rather large pearly spots as compared with
smaller punctations in erwsculus.
4 BREVIORA No. 56
type locality of crusculus. C.c. molesworthi is more nearly justi-
fiable. Specimens from the area ascribed to this latter sub-
species, Boston Bay, Ecclesdown and Buff Bay in northeastern
Jamaica, show exceptionally high dorsal counts. However a total
of five localities shows a count higher than the average for the
species as a whole: Boston Bay (mean 109.4) and Eecclesdown
(mean 106.5), on the fringe of the tropical rain forest on the
eastern side of the island which may act as an isolate; Balaclava
(mean 110.5), about 15 miles northwest of Mandeville; Bogwalk
and Mona, close to Kingston. The coefficients of difference (C.D.)
were calculated for Ecclesdown-Boston Bay and Balaclava com-
pared to both Mandeville and the species as a whole:
Boston Bay-Eeclesdown Balaclava
Mandeville (CAD Saar CD S2i32
entire species CD 2676 CD y= 101
As is to be expected, as compared to the entire species, the C.D.
is not significant. The Eeclesdown-Boston Bay and Balaclava
populations are sufficiently different from the Mandeville popula-
tion as determined by the C.D. that in the opinion of some they
might well be considered subspecies. Subspecific rank has not
been accepted for these populations for two reasons: the samples
are so small that there is no probability that the sample mean
equals the population mean; furthermore, considered for the
Jamaican populations as a whole the distribution of high or low
dorsal counts seem to be random. The midbody scale counts are
also higher in the northeast populations, but again samples are
small, and high and low mean counts are merely the extremes
of a cline in which there is overlap between even the southeast
and northeast populations.
Celestus maculatus of Cayman Brae and Little Cayman, re-
tained by Chapman Grant as a full species, does not differ from
C. crusculus in scale counts or measurements but does differ in
one detail of color —the coloring of the dorsal side of the tail.
In C. maculatus the tail is light tan or ivory in alcohol and so
without pattern; in C. crusculus of Jamaica the dorsal color and
pattern are continued on to the tail. In other respects the colora-
tion of C. maculatus falls within the range of Jamaican crusculus,
and its low scale counts resemble the Mandeville population of
the latter. The Cayman population must be regarded as con-
1956 JAMAICAN AND CAYMAN CELESTUS dD
specific with Jamaican populations but because of the tail color
difference it may be recognized as a distinct subspecies, C. crus-
culus maculatus.
SUMMARY
1. Four species of Celestus occur on Jamaica.
2. They may be arranged in two groups: the hewardu group
(long-legged group) made up of C. hewardi and C. duquesneyi.
and the crusculus group (short-legged group) comprising C.
erusculus and C. barbourt.
3. C. hewardu, C. crusculus and C. barbouri are sympatric at
Mandeville.
4. C. hewardu and C. duquesneyi (sympatric at Portland
Ridge) can be distinguished by habitus, coloration, lamellar count
and number of chin shields.
5. C. barbouri and C. crusculus can be distinguished by dorsal
seale count.
6. C. crusculus is extremely variable and composed of two
subspecies: C. crusculus crusculus occurring on Jamaica and C.
crusculus maculatus occurring on Cayman Brae and Little Cay-
man. There may be a third subspecies C. crusculus molesworthi,
distinguishable by high dorsal scale counts and found in north-
eastern Jamaica. Its validity is, however, not yet firmly demon-
strated.
I wish to express my thanks to Dr. Ernst Mayr and Dr. E. E.
Williams for their generous and valuable advice, to Mr. Benja-
min Shreve who made the scale counts and measurements on the
specimen of Celestus barbourt from Fern Gulley, and to Mr.
Garth Underwood for useful criticism of the manuscript.
LITERATURE CITED
CopE, H. D.
1868. An examination of the Reptilia and Batrachia obtained by the
Orton Expedition to Ecuador and the Upper Amazon with notes
on other species. Proce. Acad. Nat. Sci. Phila.: 96-140.
6
GARMAN, S.
1887.
1888.
GRANT, C.
1940a.
1940b.
1940e.
Gray, J. E.
1845.
BREVIORA No. 56
On West Indian Geckonidae and Anguidae. Bull. Essex Inst., 19:
17-24.
Reptiles and batrachians from the Caymans and from the
Bahamas. Bull. Essex Inst., 20: 103-116.
Notes on the reptiles and amphibians of Jamaica, with diagnoses
of new species and subspecies. Jamaica Today: 151-157.
The Reptiles in Lynn and Grant: The herpetology of Jamaica.
Bull. Inst. Jamaica, Sci. Ser., 1: 1-65.
The herpetology of the Cayman Islands. Bull. Inst. Jamaica, Sei.
Ser., 2: 1-65.
Catalogue of the specimens of lizards in the collection of the
British Museum. London, 289 pp.
ae Rice in
i
ny “an A
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. JUNE 29, 1956 NUMBER 57
THREE NEW SPECIES OF MIMETIDAE
(ARANEAE) FROM PANAMA
By ArtHur M. CHICKERING
Albion College, Albion, Michigan
In 1947 I published a brief paper attempting to bring our
knowledge of the Mimetidae of Panama up to date. At that time
six species of Gelanor were recognized and two of these were
regarded as new. Five species of Mimetus were also recognized
at that time and two were considered new to science. As a result
of two field trips into various parts of Panama since 1947 (1950
and 1954) a considerable number of specimens belonging to this
family have been sorted from my collections. Three new species
belonging to the two genera mentioned above have been selected
from these later collections. These are described in this paper
in accord with my usual procedure. I regret that I cannot yet
be more certain regarding the correct association of males and
females in several of the known species. The types are being
deposited in the Museum of Comparative Zoology.
It is again a pleasure to acknowledge my indebtedness to the
following persons for their continued encouragement in the
pursuit of my studies: Dr. A. S. Romer and Dr. P. J. Darlington,
Jr., Director and Curator of Insects, respectively, in the Museum
of Comparative Zoology at Harvard College. Without the priv-
ileges which have been extended to me for many years in this
institution the continued progress of my studies would have been
much more difficult.
3) BREVIORA NO. 57
Genus GELANOR Thorell, 1870
GELANOR DEPRESSUS Sp. Nov.
(Fig. 1)
Female holotype. Total length 4.16 mm. Carapace 2.145 mm.
long; 1.69 mm. wide opposite second coxae where it is widest;
narrowed to .91 mm. just behind PLE; about 1.04 mm. tall
opposite first coxae where it is tallest; rises gradually from PE
to highest point, then descends fairly steeply to posterior border ;
with median thoracic pit about midway of the posterior declivity.
Eyes. Eight in two rows as usual; heterogeneous; lateral eyes
on a moderately prominent tubercle. AME separated from one
another by slightly more than their diameter, from ALE by 9/7
of their diameter. PME separated from one another by about
7/10 of their diameter, from PLE by 2.6 times their diameter.
Laterals contiguous to one another. Seen from above, anterior
External Anatomy of Gelanor and Mimetus
Fig. 1. Gelanor depressus sp. nov.; epigynum, ventral view.
Figs. 2-3. Mimetus saetosus sp. nov.; epigynum, ventral and posterior views.
respectively.
Fig. 6. Mimetus variegatus sp. nov.; epigynum, ventral view.
1956 NEW SPECIES OF MIMETIDAE 3
row quite strongly recurved, posterior row gently procurved.
Seen from in front, anterior row gently recurved, measured by
eenters. Central ocular quadrangle wider in front than long in
ratio of 13 : 11, wider in front than behind in ratio of 3 : 2.
Height of clypeus equal to 9/14 of the diameter of AME. Ratio
of eyes AME : ALE ; PME : PLE = 14:9 : 11 : 10 (long
diameters used when differences exist).
Chelicerae. Vertical, parallel, somewhat narrowed at their
bases; bases united for about one fifth of their length; basal seg-
ment .975 mm. long; with several long slender spines but none
notably enlarged; basal boss lacking. Fang groove with the
usual row of six or seven stiff spines but with no visible teeth;
retromargin with a single low tuberculate tooth near middle.
Mazxillae. Convergent; moderately slender; almost contiguous
distal to lip; palp inserted into basal third.
Iip. Clearly united to sternum with no sternal suture visible;
only shghtly longer than wide near middle.
Sternum. Scutiform m general; moderately convex; widest
between first coxae but nearly as wide between second coxae;
longer than wide in ratio of about 11 : 8; with numerous long
slender bristles; posterior end extended between fourth coxae
and terminating in a sharp point; with fourth coxae separated
by a little less than the width of one of them; first and second
coxae robust, third and fourth slender.
Legs. 1248. Width of first patella at ‘‘knee’’ .352 mm., tibial
index of first leg 7. Width of fourth patella at ‘‘knee’’ .22 mm.,
tibial index of fourth leg 9.
(All measurements in millimeters)
Fmora Patellae Tibiae Metatarsi Tarsi Totals
its 3.770 1235 3.510 3.640 2.015 14.170
Ze 3.510 1.100 2.600 2.510 1.430 15.0
oe 1.950 .660 1.690 1.690 .750 6.740
4, 2.405 .700 1.700 1.755 .750 7.310
Spines. First leg: femur dorsal 0-1-1, prolateral and retro-
lateral only one near distal end, ventral 0; patella 0; tibia only
with definite spines on prolateral surface as follows: a series of
4 BREVIORA NO. 57
seven robust spines beginning near proximal end and extending
to near distal end and with seven series of shorter spines follow-
ing the robust spines and distributed as follows: 2,3,3,3,4,4,5 ;
metatarsus with only definite prolateral spines as follows: a
series of four robust spines beginning at proximal end and with
fourth near middle and four series of shorter spines following
the robust spines and distributed as follows: 5,7,8,19. Second
leg: femur and patella as in first except femur dorsal 1-1-1; tibia
only with definite spines as follows on prolateral surface: with
five robust spines, the first weaker and near proximal end, the
fifth about one fifth from distal end and with shorter spines
following the robust spines arranged as follows: 0,1,4,4,5; meta-
tarsus also only with definite spines as follows: on prolateral sur-
face one moderately long and robust proximal spine and near this
there is a very long and robust spine with another similar spine
about one fourth of the length of the segment from the base;
immediately following the first long robust spine there is a series
of six shorter curved spines, and following the second long spine
there is a series of 18-20 shorter curved spines diminishing to
mere bristles distally. Third leg: femur only dorsal 0-1-0; patella
only one dorsal distal spine, long and slender; tibia only dorsal
distal 1 and prolateral 0-1-0; metatarsus only dorsal distal 1
(very slender) and prolateral 0-1-0-0-0. Fourth leg: femur 0;
patella as in third; tibia only dorsal 0-1-0-1-0; metatarsus ap-
parently only prolateral 0-1-0-0-0. Palp: femur apparently only
with a pair of dorsal distal spines; patella with a single long
slender dorsal distal spine; tibia with a group of three or four
distal spines, lacking ventrally ; tarsus dorsal 0-1-0-0-0, prolateral
and retrolateral 0-1-1-0. Palpal claw with one short and one long
tooth. Tarsal claws three with spurious claws as usual. The
‘‘drum’’ has been seen on the dorsal side of some tarsi about one
fourth of the length of the segment from the distal end.
Abdomen. Moderately high, well rounded laterally; slightly
broader than long near middle. Anal tubercle and spinnerets as
usual in the genus. Colulus well defined. Tracheal spiracle well
defined and just anterior to base of colulus. Genital groove well
defined just posterior to epigynum.
Epigynum. Quite tuberculate ; posterior surface with a pair
of chitinized areas separated by a shallow slit; ventral surface
1956 NEW SPECIES OF MIMETIDAE 5
with a pair of moderately deep depressions separated by a septum
Gee):
Color in alcohol. Legs: third and fourth pairs nearly uni-
colorous yellowish; first pair darker yellowish with femora red-
dish dorsally and patellae also somewhat reddish especially at
proximal ends and tibiae reddish at distal ends. Chelicerae a
clear yellowish with fangs brownish. Lip and maxillae yellowish
with reddish streaks. Carapace yellowish with reddish flecks just
behind PE and a pair of irregular spots near apex and two pairs
of irregular reddish spots further posterior. Sternum generally
yellowish but with a small reddish dot opposite each second
coxa and a large irregular spot covering most of the posterior
half. Abdomen: anterior third of dorsum yellowish with a few
bright red flecks at the base; about one third from the base are
two irregular bright red spots, each with two small white dots
nearly enclosed by the red color; behind these spots there is a
series of nine or ten very narrow red transverse lines; the venter
is yellowish lateral to the epigynum but reddish centrally behind
this organ and with numerous oblique red ventro-lateral lines.
Type locality. Female holotype is from Barro Colorado
Island, C. Z., June, 1950. There are no paratypes.
Genus MIMETUS Hentz, 1832
MIMETUS SAETOSUS sp. nov.
(Figs. 2-3)
Female holotype. Total length 3.71 mm. Carapace 1.95 mm.
long; 1.43 mm. wide between third coxae where it is widest;
narrowed to .845 mm. just behind PLE; with no longitudinal
thoracic groove but with a shallow depression about midway
along the posterior declivity ; rises sightly from PLE to opposite
interval between first and second coxae and then after a gentle
arch descends steeply to posterior border; with twelve spines
in three rows behind PE, the middle row of only two and with
each lateral row with five, the posterior one on each side being
offset laterally.
Eyes. Kight in two rows; heterogeneous; LE on prominent
tubercles; AME at base of a tuberele which extends considerably
6 BREVIORA NO. 57
dorsal from the eye level, with a spine at tip of each cone as in
M. rusticus Chickering. AME separated from one another by
two thirds of their diameter, from ALE by 1.3 times their
diameter. PME separated from one another by slightly more
than their radius, from PLE by nearly 2.2 times their diameter
(long diameters used when eyes are not circular). Laterals
separated only by a line.
Chelicerae. Vertical, parallel, quite robust, quite convex in
front; basal segments united at bases for about one fourth their
leneth which is .88 mm.; without basal boss; with numerous
short slender bristles in front and several longer ones; with one
fairly robust spine in medial position near distal end overlapping
its mate on opposite side. Fang as usual in the genus. Retro-
margin of fang groove with a single tuberculate tooth near
medial end; promargin with a row of ten robust spines.
Mazillae. Slender, slightly convergent; palp inserted near
base; serrula extends only along outer distal corner.
lip. Wider than long in ratio of about 5 : 4; reaches well be-
yond middle of maxillae; no definite suture observed.
Sternum. Seutiform; longer than wide in ratio of 13 : 16;
widest between first coxae; slightly convex; with numerous long
slender bristles; posterior end rounded, not extended between
fourth coxae which are separated by about two thirds of their
width. First coxae considerably swollen; third coxae the smallest.
Legs. 1248. Width of first patella at ‘‘knee’’ .286 mm., tibial
index of first leg 8. Width of fourth patella at ‘‘knee’’ .198 mm.,
tibial index of fourth leg 9.
(All measurements in millimeters )
Femora Patellae Tibiae Metatarsi Tarsi Totals
il. 2.860 780 2.730 Dono 1.250 9.895
2 2.340 .650 1.950 1.820 1.105 7.865
3h. 1.300 455 1.040 .855 840 4.490
4, 1.820 .528 1.690 12235 845 6.118
Spines. First leg: femur dorsal 0-1-1-1-1-1-0 with irregularities
in placement, prolateral 0-0-1-1-1, retrolateral only one robust
spine near distal end but with a somewhat oblique row of about
1956 NEW SPECIES OF MIMETIDAE 7
a dozen short stiff spinules in the proximal fifth, ventral a nearly
median row of about fourteen long slender spines together with
another row of similar but less regular spines in a prolateral
position ; patella only dorsal spines 1-1 together with several very
slender spines irregularly placed; tibia dorsal 0-1-1-1-0, pro-
lateral robust spines 0-1-1-1-1-1-1-1-0 together with short spines
between the robust spines and distal to the last in numerical
series as follows: 3-2-4-3-4-4-5, retrolateral robust spines 0-1-0
together with numerous long slender spines, ventral with only
numerous long slender spines; metatarsus dorsal only a series of
long slender spinules, prolateral robust spines 1-1-1-1-1-1-0 with
the expected short curled spines intervening between the robust
spines and distal to the last of these in six series distributed as
follows: 3-4-5-6-7-9, retrolateral with a row of 9 long slender
spines, ventral with another row of long slender spinules, hardly
more than bristles. Second leg: femur essentially as in first except
for the presence of the proximal prolateral row of short. stiff
spinules presumably used in opposition to the retrolateral row of
similar spinules on the first femur; patella as in first ; tibia dorsal
0-1-1-1-0 plus several long slender spines somewhat irregularly
placed, prolateral 0-1-1-1-1-1-0 plus smaller spines between robust
spines and distal to the last of these but these are less numerous
and less conspicuous than in first, retrolateral and ventral as in
first; metatarsus dorsal as in first, prolateral robust spines
1-1-1-1-0 with shorter spines terminally bent in four series dis-
tributed as follows: 3, 4, 6, and 11. Third leg: femur essentially
as in first; patella as in first; tibia dorsal O-1p-0-0, prolateral
Q-1-0-0, retrolateral 0-1-0, ventral O-1p-0-1p-0; metatarsus dorsal
0, prolateral 0-1-0-0, retrolateral 0-1-0, ventral 0-2 (irregular )-0.
Fourth leg: femur and patella essentially as in first; tibia dorsal
Q-1r-0-1-0, prolateral 0-1-0-0, retrolateral 0-1-0 together with sev-
eral other slender spines irregularly placed, ventral 0-1p-0-1-0
together with several long slender and weak spines irregularly
placed ; metatarsus essentially as in third. Palpal claw pectinate
with a single row of 5 or 6 slender teeth in proximal half.
‘
Abdomen. 2.08 mm. long; 2.34 mm. wide between ‘‘shoulder
humps’’ which are conspicuously developed. With numerous long
slender stiff bristles, almost spines. Colulus well developed ;
tracheal spiracle a short distance anterior to base of anterior
8 BREVIORA NO. 57
spinnerets and with a fairly well developed chitinous lip. Spin-
nerets as usual in the genus.
Epigynum. With a moderately well chitinized plate; moder-
ately protuberant; somewhat broader than long; with a narrow
notched lip along posterior border (Figs. 2-3).
Color in alcohol. Legs and palps yellowish with many reddish
brown spots and irregular rings. Chelicerae a rich reddish
brown. Carapace in general yellowish with brownish markings;
the largest of these begins at the clypeus and extends backward
and narrows irregularly to a blunt point near the posterior
border; three small irregular brownish spots occur on each
lateral side. Sternum: with three pairs of small brownish spots
and a single similar median spot at the posterior end. Abdomen:
yellowish with many small brownish spots.
Type locality. Female holotype from Barro Colorado Island,
C. Z., July 18, 1954. No paratypes.
MIMETUS VARIEGATUS Sp. nov.
(Figs. 4-6)
Male holotype. Total length 2.795 mm. Carapace 1.3 mm. long ;
1.17 mm. wide opposite interval between second and third coxae
where it is widest; narrowed to .58 mm. wide just behind PLE;
with no longitudinal median thoracic groove but with a broad
shallow depression about midway along the posterior declivity ;
rises slightly from PE to opposite interval between second and
third coxae and then descends steeply to posterior border; ap-
parently with twelve slender spines in three rows on cephalic
area posterior to PE.
Eyes. Eight in two rows; heterogeneous; all except PME on
prominent tubercles. AME separated from one another by nearly
1.5 times their diameter, from ALE by about one third as much.
PME separated from one another by their radius, from PLE by
their diameter. Laterals separated from one another by a line.
Seen from above, posterior row slightly procurved; seen from
in front, anterior row slightly procurved. Central ocular quad-
rangle wider in front than behind in ratio of about 4 : 3, slightly
wider in front than long. Ratio of eves AME : ALE : PME :
1956 NEW SPECIES OF MIMETIDAE 9
PLE = 5.5 : 4 : 4.5 : 4.5. Height of clypeus equal to about
14/11 of the diameter of AME.
External Anatomy of Mimetus
Figs. 4-5. Mimetus variegatus sp. nov.; tibia and tarsus of male palp;
two views.
Chelicerae. Vertical and parallel; rather long and slender;
basal segment .65 mm. long; united at their bases for about one
sixth of their length; without basal boss; with two slender spines
on front surface and one more robust spine in medial position
near distal end overlapping its mate on opposite side. Fang
stout at base, strongly bent at middle, slender in distal half.
Fang groove moderately distinct and without ordinary teeth
but promargin has a row of 7 or 8 stout spinules (examination
made on a paratype to avoid injury to the holotype).
10 BREVIORA NO. 57
Mazillae. Slender, somewhat convergent; reach considerably
beyond lip; with palp inserted into base; with serrula only at
outer distal corner.
Lip. About nine tenths as long as wide; with a well developed
sternal suture; reaches well beyond middle of maxillae.
Sternum. Broad, scutiform; almost as wide as long; strongly
convex ; widest between second coxae but nearly as wide between
first coxae; with a moderate supply of long slender bristles;
posterior end rounded and not extended between fourth coxae
which are separated by about 7/10 of the width of one of them ;
first coxae most robust; third and fourth nearly equal in size.
Legs. 1243. Width of first patella at ‘‘knee’’ .198 mm., tibial
index of first leg 5. Width of fourth patella at “‘knee’’ .132
mm., tibial index of fourth leg 7.
(All measurements in millimeters )
Femora Patellae Tibiae Metatarsi Tarsi Totals
ile 2.990 .682 3.055 3.510 1.365 11.602
os 2.405 .520 lal 2.2785 1.000 8.312
oF 1.625 300 975 1.054 616 4.550
4. 1.820 390 1.430 1.430 .616 5.686
Palp .682 198 .264 --—- .682 1.826
Spines. First leg: femur dorsal 1-1-1-1-1, prolateral 0-1-1-1-0,
retrolateral a row of short, somewhat robust spines extending
through about three fifths of the length of the segment beginning
at proximal end, together with a single long spine near distal end,
ventral with several stiff bristles but hardly spines; patella dorsal
1-1; tibia dorsal 0-1-1-1-0 plus several weak spines hardly more
than bristles between the robust spines, prolateral O-1-1-1-1-
1-1-1-0, retrolateral 0-0-1-0, ventral 0; metatarsus dorsal nine
in the proximal four fifths of varied size and length, prolateral
1-1-1-1-1-0-0 with a few small intervening spines among the
robust spines (in contrast to the expected condition), retrolateral
with numerous weak spines but none robust of the usual sort,
ventral 0. Second leg: femur dorsal as in first, prolateral devoid
of long robust spines but with a row of very short stiff spinules
extending through proximal third of segment and probably used
in opposition to the similar spinules on the retrolateral surface
1956 NEW SPECIES OF MIMETIDAE ital
of first femur, retrolateral apparently with only one near distal
end, ventral 0; patella essentially as in first ; tibia dorsal 0-1-1-1-0
with several weak spinules and bristles irregularly distributed
among the more robust spines, prolateral 0-1-1-1-1-1-0 with some
irregularity together with spinules irregularly distributed
among the more robust spines, retrolateral 0-0-1-0-1 (weak), ven-
tral 0; metatarsus dorsal 0, prolateral 1-1-1-0-0 with short
spinules between the robust spines and distal to the last, retro-
lateral with a series of 10-12 spines of diminishing lengths toward
the distal end where there are only bristles. Third leg: femur
dorsal apparently 1-1-1-1-1 but with numerous spinules, else-
where 0; patella only dorsal 0-1, tibia dorsal 0-1-0-0, prolateral
the same, retrolateral 1(weak)-1-0, ventral 1p-1p-0; metatarsus
dorsal O-1-0-0, prolateral essentially the same, elsewhere 0.
Fourth leg: femur and patella essentially as in third; tibia
dorsal 0-1-0-1-0, prolateral and retrolateral 0-1-0, ventral 0;
metatarsus only dorsal 0-1-0-0 with prolateral essentially the
same. A comparison of the spines, especially on legs one and
two, shows that they differ from those regarded as typical of the
genus especially because of the great reduction of the number,
size, and regularity of occurrence of the small spines in between
the larger ones of tibiae and metatarsi. Palp: femur with only
one dorsal distal spine; patella with two long slender dorsal
spines; tibia with a row of five or six long slender spines or
spinules curving along near distal border.
Palp. Of moderate length; tibia and patella short; tarsus
moderately compleated ; with structural features best shown in
Figures 4-5. Base of eymbium with a lobule which suggests that
of M. verecundus Chickering but is, nevertheless, quite distinct
from that species.
Abdomen. Of moderate height; about 9/11 as wide as lone
at level of ‘‘shoulder humps’’; 1.43 mm. long; widest at level of
‘shoulder humps’’ which occur slightly anterior to middle ; with
numerous long slender spinules on dorsal and dorsolateral sur-
faces. Anal tubercle and six spinnerets as usual in the genus.
The colulus appears to be reduced. Tracheal spiracle near base
of spinnerets with a poorly developed chitinous lip.
12 BREVIORA NO. 57
Color in alcohol. Legs generally light yellowish with many
grayish and brownish spots together with dark rings, more
marked on first two pairs. Hach chelicera has a small dark spot
in front near base. Carapace: the eyes are ringed with reddish
pigment; there is a median dorsal grayish stripe extending from
the PE backward and diminishing to a termination between an
irregular V-shaped dark mark which extends forward from the
depression at the center of the posterior declivity. The sternum
is a clear yellowish. Abdomen: the dorsum from opposite the
posterior border of the ‘‘shoulder humps’’ to base is richly
variegated with black, yellow, and red marks of a great variety
of shapes; behind the ‘‘shoulder humps’’ the dorsum is whitish
with narrow reddish transverse procurved lines connected lat-
erally with black elongated spots; the venter is provided with
several median dark spots and dark lateral bars.
Female allotype. Total length 2.925 mm. Carapace 1.49 mm.
long; 1.17 mm. wide opposite third coxae where it is widest;
narrowed to .65 mm. wide just behind PLE. Otherwise essen-
tially as in male.
Eyes. AME separated from one another by 7/5 of their diame-
ter, from ALE by about 2/5 of their diameter. PME separated
from one another by about 5/9 of their diameter, from PLE by
slightly more than their diameter. Laterals separated only by a
line. Seen from above, posterior row shghtly procurved; seen
from in front, anterior row the same, measured by centers.
Central ocular quadrangle wider in front than behind in ratio
of about 4 : 3, wider in front than long in ratio of about 8 : 7.
Ratio of eyes AME : ALE : PME : PLE = 10:8 : 8 : 8.5.
‘Height of clypeus somewhat greater than diameter of AME.
Otherwise essentially as in male.
Chelicerae. Basal segment .814 mm, long. The spinules along
the promargin of the fang groove are more conspicuous than in
the male. Otherwise essentially as in that sex.
Mazillae, Lip, and Sternum. Essentially as in male.
Legs. 1243. Width of first patella at ‘‘knee’’ .231 mm., tibial
index of first leg 6. Width of fourth patella at ‘“‘knee’’ .154 mm.,
tibial index of fourth leg 8.
1956 NEW SPECIES OF MIMETIDAE 13
(All measurements in millimeters )
Kemora Patellae Tibiae Metatarsi Tarsi Totals
le 2.860 SPAS 2.925 2.730 1.300 10.540
2 2.405 645 2.145 1.885 1.040 8.120
3 1.495 390 1.105 1.040 15 4.745
4. 2.080 455 1.495 1.300 .780 6.110
Spines. Small spines between the robust spines on the first
and second legs are much more conspicuous and more regular
in occurrence than in the male. First leg: femur dorsal 1-1-0-0-
1-1, prolateral 0-1-1-1-0, retrolateral with a row of 16-18 short
stiff stridulation spinules in proximal third and with one robust
spine near distal end, ventral with a double row of long slender
spinules; patella only dorsal 1(weak)-1; tibia dorsal 0-1-1-1-0,
prolateral with O0-1-1-1-1-1-1-1-0 robust spines and short curved
spines between the more robust spines usually four in each series
increasing in length distally, retrolateral 0-0-1-0-1 (weak), ven-
tral 0; metatarsus dorsal a row of 12-15 lone slender spinules,
hardly spines, prolateral 1-1-1-1-1-0, retrolateral 0 but with
numerous bristles, ventral only numerous bristles. Second leg:
femur dorsal 0-1-0-0-1-1, prolateral only with a row of short
stiff spinules to oppose the corresponding similar spinules on
first femur, ventral essentially as in first, retrolateral apparently
only a single spine near distal end; patella as in first; tibia
dorsal 0-1-1-1-1, prolateral 0-1-1-1-1-0 and with a series of shorter
weaker curved spinules following each of the larger robust spines,
retrolateral only 0-1-0, ventral 0; metatarsus dorsal only a row
of bristles or slender spinules, prolateral 1-1-1-0-0 with three
series of shorter weaker curled spines following the robust spines
as follows: 4, 5, and 12 or 13, retrolateral only a row of bristles
or weak slender spines, ventral the same. Third leg: femur only
dorsal 1-1-0-0 and numerous stiff bristles; patella as in first ; tibia
apparently only dorsal 0-1-0-0; metatarsus apparently only dor-
sal and prolateral 0-1-0-0. Fourth leg: femur dorsal 1-1-1-0-0,
elsewhere 0 exeept for the frequent weak long slender spines,
hardly more than bristles; patella essentially as in first; tibia
dorsal 0-1-0-1-0, prolateral and retrolateral 0-1-0, ventral 0-1p-
0-1p-0; metatarsus only prolateral 0-1-0-0 and retrolateral 0-1-
0-0-0.
14 BREVIORA NO. 57
Abdomen. Except for sexual differences essentially as in male.
Epigynum. Similar to that of M. verecundus Chickering but
with distinct differences; massively tubercular; with a small
shallow depression at each anterior lateral corner (Fig. 6).
Color in alcohol. Similar to that of male exeept that the
variegated abdominal area extends over a large part of the dor-
sum behind the ‘‘shoulder humps.’’ Considerable variation in
color has been noted among the paratypes.
Type locality. Male holotype and female allotype from the
Canal Zone Experiment Garden, August, 1954. Two male para-
types from near Pedro Miguel, C. Z., August, 1954 and Canal
Zone Experiment Garden, C. Z., August, 1954. Female paratypes
from near Chiva, C. Z., July, 1954; Summit, C. Z., July, 1950;
Arraijan, R. P., July, 1950.
BIBLIOGRAPHY
CHICKERING, ARTHUR M.
1947. The Mimetidae (Araneae) of Panama. Trans. Amer. Micros.
Soc., 66: 221-248.
CAMBRIDGE, O. P. and F. P. CAMBRIDGE
1889-1905. Avachnida-Araneida. Vols. I-Il. Jn: Biologia Centrali-Ameri-
cana. Dulau & Co., London.
> i ie oe ee fe tat ae
a eens: >
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. JUNE 29, 1956 NUMBER 58
SEQUENCE OF PASSERINE
FAMILIES (AVES)
By EK. Mayr ann J. C. GREENWAY, JR.
At the XI International Ornithological Congress at Basel,
Dr. Stresemann proposed that a committee be appointed which
would recommend to the editors of ornithological journals a
standardized sequence of the families of Passerine birds. Dr.
Stresemann pointed out that some editors follow Hartert, others
Sharpe’s Handlist, others the sequence of the AOU, and still
others the recently proposed sequence of Mayr and Amadon
(1951). If a reader wants to find a given family in an article, he
may have to look either at the beginning or in the middle or at the
end depending on the particular sequence adopted by the
respective editor.
As a result of Dr. Stresemann’s suggestion a committee was
appointed by the president of the Congress with the following
members :
Dr. G. C. A. Junge, Chairman
Rijksmuseum Nat. Hist., Leiden
Prof. J. Berlioz
Museum d’Hist. Nat., Paris
Prof. Dr. G. Dementiev
Zool. Mus. University, Moscow
Prof. E. Mayr
Mus. Comp. Zool., Cambridge, (Mass. )
Mr. R. E. Moreau
Edward Grey Institute, Oxford
Dr. F. Salomonsen
Zool. Museum University, Copenhagen
Prof. Dr. E. Stresemann
Zool. Museum University, Berlin
bo
BREVIORA NO. 58
In the discussion, it was pointed out that more and more
editors adopt the sequence of already published volumes of
Peters’ Checklist, and the editors of Peters’ Checklist were re-
quested to submit to the committee the sequence which they had
been planning to use. Dr. Mayr expressed the views of the editors
by saying that they would very much like to follow a standard-
ized and universally adopted sequence, in view of the fact that
there are no decisive arguments available in favor of any of
the previously proposed sequences. He stated that the editors
of Peters’ Cheeklist would be willing to adopt whatever sequence
the committee would vote for.
REPORT OF THE EDITORS OF PETERS’ CHECKLIST
In order to facilitate the work of the appointed committee,
the editors of Peters’ Checklist submitted to that body some
tabulated historical material on sequences adopted by earlier
authors. In the introduction to this report the following sub-
ject matter was discussed :
The task of the student of avian classification is particularly
difficult beeause too little is as yet known about the paleontology
of the songbirds to be of help in devising a sound classification.
It is evident for this and more basie reasons that any linear order
must be arbitrary. Three considerations usually guide those who
attempt to find a suitable sequence :
(A) To follow as closely as possible the traditional arrange-
ments, except where subsequent work has shown conclusively
that a change is advisable ;
(B) To place families near each other which are presumably
closely related ;
(C) To place the more primitive families near the beginning
and the more advanced families near the end.
The following comments may be made on these three principles.
(A) Traditional arrangements. In nearly all recently proposed
sequences it is acknowledged that the larks and the swallows are
aberrant in various ways. These two families are, therefore,
usually disposed of at the beginning of the system. The remain-
ing bulk of songbirds is usually grouped into three major assem-
blages: (1) Old World Insect-eaters and relatives (Campephagi-
dae, Pyenonotidae, Sylviidae, Timaliidae, Turdidae, Muscicapi-
1956 SEQUENCE OF PASSERINE FAMILIES (AVES) 3
dae, ete.), (2) New World Insect-eaters and finches, (3) crows,
birds of paradise, and associated families. All the more pecular
and isolated families, as well as the Old World Nectar-eaters,
are grouped rather irregularly within this broad framework.
The sequence of the three major assemblages is by no means
standardized, and to achieve a universal (or at least general)
acceptance of one should be particularly the object of the com-
mittee.
The sequence 1, 2, 3 is that of Shufeldt (1904), Sharpe’s Hand-
list, Hartert (in reverse here), the official Checklist of Australian
Birds of 1926, Chasen’s Handlist (1935), Mathews’ Systema
(1930), and more recently of the Checklist of Japanese Birds
(1942), Berlioz (1950), Mayr and Amadon (1951) and Biswas
GL952)).:
The sequence 1, 3, 2 was suggested first by Wallace (1874),
as far as we can discover. This arrangement was based on the
reduction in the number of primaries. As early as 1885 Sharpe
remarked that it was difficult to follow it in a linear sequence.
However, it was followed in certain sections of the Catalogue of
Birds in the British Museum, but in others the arrangements of
Sundevall (1872) and Garrod (1876) were adopted. We cannot
find that the exact order of the ‘‘Catalogue of Birds’’ has been
followed by anyone. This same order (1, 3, 2) was adopted by
Stejneger in 1885. Evans (1899) used this order in the Cam-
bridge Natural History, Selater used it in 1930, and Stresemann
again in the Handbuch der Zoologie (1954).
Finally, the sequence 3, 1, 2 was adopted by Wetmore and
Miller (1926), and has been the order used in the American
Ornithologists’ Union Checklist (1931) and in works following
the A.O.U. Checklist.
It should be added parenthetically that neither Fuerbringer
(1888) nor Gadow (1893, 1898) made any attempt to classify the
Oscine Passeres into families or subfamilies.
(B) Mutual relationship. To determine exact interrelation-
ships of these families is in many cases very difficult, if not
impossible. The system of songbirds abounds in artificial aggre-
eates such as the ‘‘finches’’ or the ‘‘shrikes’’ of old, groups
which may have little in common, except, for instance, the shape
of the bill. In the improvement of the grouping considerable
progress has been made in recent years such as the breaking up
4 BREVIORA NO. 58
of the artificial assemblage ‘‘finches’’ into carduelines, buntings
(Emberizidae) and cardinals, or the assembling of the vireos,
wood warblers, tanagers, and related families into a single
ageregate. There are still many conventional groupings such as
Paridae - Sittidae - Certhiidae which are presumably quite arti-
ficial. Yet, it would serve no useful purpose to break up these
well-known sequences, until additional information sheds new
light on the relationship of these families.
(C) Phylogenetic sequence. A satisfactory classification into
‘‘primitive’’ or ‘‘specialized’’ families is virtually impossible
for the exceedingly similar groups of songbirds. The convenience
of placing the rather undifferentiated ‘‘Old World Insect-
eaters,’’ and particularly the very generalized Campephagidae
and Pyecnonotidae near the beginning, is now accepted by the
great majority of classifiers. But whether to follow the sequence
1, 2, 3, or 1, 3, 2—that is, whether to consider the finches or
the crow and bird-of-paradise group relatively more specialized
— depends entirely on the criteria used. If adaptation to a seed
diet or loss of the tenth primary are considered characters of
great importance, then the sequence 1, 3, 2 is certainly best.
However, if the cerebralization of the Corvidae and the extraor-
dinary courtship habits of the birds of paradise and bower birds
are considered indications of evolutionary significance among
songbirds, then the sequence 1, 2, 3 is to be preferred.
The recent realization that the cone-billed birds (‘‘finches’’)
form an artificial, polyphyletic assemblage has weakened the sup-
port for sequence 1, 3, 2. Not only should the true finches (in-
cluding Carduelinae) be removed from the New World Ember-
izidae-Cardinalinae assemblage (Tordoff 1954) but likewise the
Kstrildidae should be removed from the Ploceidae (Steiner
1955). Possibly the Estrildidae are related to the cardueline
finches, both groups showing a curious mixture of primitive and
advanced characters. The loss of the outermost primary which
has happened in many families of birds independently and irreg-
ularly is, like all loss-characters, a very unsafe taxonomic cri-
terion. All this militates against the sequence 1, 3, 2.
However, weighty arguments can also be advanced against the
sequence 1, 2, 3. To consider cerebralization a most important
character (Portmann) is somewhat anthropomorphic, as Wet-
more (1951) has rightly observed. Furthermore, high cerebral-
1956 SEQUENCE OF PASSERINE FAMILIES (AVES) 5
ization has also occurred among Non-Passeres (Psittaci), and the
relationship Corvidae-Paradisaeidae-Ptilonorhynchidae is by no
means unequivocally established. In many purely morphological
respects (bill, wing) the families of this group are undoubtedly
more generalized than are some of the finches, and particularly
the nectar-eating birds.
There is, thus, no decisive criterion that would permit coming
to an unequivocal conclusion. It should be added that it would
be shortsighted to accept a sequence which would satisfy Euro-
pean and American ornithologists only, but not those in the
remainder of the world. The placement of the birds of paradise
and bower birds may be immaterial to the student of European
or of North American birds, but no world list of birds can be
prepared which does not pay as much attention to the classifica-
tion of these families as to that of the finches or thrushes.
It appears to the editors of Peters’ Checklist that the 1, 2, 3
sequence is the one most widely used in the ornithological litera-
ture of the world. Since they know of no decisive reason for
changing it, they have adopted a sequence which agrees roughly
with the order 1, 2, 3.
ACTIONS OF THE COMMITTEE
The editors of Peters’ Checklist requested that the members
of the Committee express their preference between the sequences
1,2, 3and 1, 3, 2 and 3, 1, 2, and they used this opportunity to ask
for suggestions concerning the placement of individual families.
The result was that the members of the Committee (Berlioz,
Dementiev, Junge, Mayr, Moreau, Salomonsen, and Stresemann )
voted unanimously in favor of the sequence 1, 2, 3. They also
submitted questions concerning the family rank of certain genera
and groups of genera and suggested a number of modifications.
Before these are discussed, two matters of policy, which will
euide the editors of Peters’ Checklist, may be stated.
(A) There are a number of natural groups among the Oscines
such as the thrush-flyeatcher group, the ‘‘New World finches,’’
and others delineated in Mayr and Amadon, 1951. In a linear
listine unrelated groups of familes must often of necessity be
placed next to one another. This does not imply that such
adjacent families are considered related, but is merely the un-
6 BREVIORA NO. 58
avoidable consequence of having to present a three-dimensional
phylogenetic tree cut up into a one-dimensional linear sequence.
As a result the Bombycillidae, for instance, are found next to
the Cinclidae, the Pachycephalinae next to the Paridae, the
Meliphagidae next to the Emberizinae, ete. Family group head-
ings will be used in Peters’ Checklist in order to minimize the
effect of such misleading, but unavoidable groupings.
(B) The editors consider it unwise in such an essentially uni-
form group as the Oscines to recognize many families containing
only a single species. They prefer to place somewhat isolated
species in separate subfamilies, near the families with which the
genera are traditionally associated. This they intend to do for
such genera as Hypositta, Dulus, and Pityriasis. There are
many additional genera of songbirds which could be treated with
equal justification as separate families, such as Lamprolia, Ticho-
droma, Promerops, and others. Such a multiplication of families
would not be constructive in the absence of sound anatomical
support for the splitting.
The following comments concern the placing of individual
families and deal with questions raised by committee members.
1. Alaudidae. Should this family, currently lsted near the
beginning of the Oscines, be transferred nearer to the Emberizi-
dae, a position which it held in some of the older classifications?
Answer: This is not advisable. The Alaudidae are a very
peculiar family. They differ from all other Acromyiodean Pas-
seres by having not only the front but also the back of the tarsus
scutellate and in having the pessulus rudimentary. This indi-
eates that the larks may not be closely related to any of the
other families. Since they are not specialized to any great
extent they are probably best placed near the head of the list.
Two functional characters, the heavy bill in some of the seed-
eating genera, and the reduction in the number of primaries,
cannot be considered evidence for relationship to the finches.
2. Grallinidae and Artanudae. Should these families be trans-
ferred from a place near the Laniidae to the neighborhood of the
Callaeidae?
Answer: Yes. It is advisable for two reasons to keep together
all peculiar Australian families, the relationship of which is”
obscure. It is probable that these families had their origin in the
Australian region and that they are distantly related. It is also
1956 SEQUENCE OF PASSERINE FAMILIES (AVES) 7
desirable for purely practical reasons to have all purely Aus-
tralian families near each other. In view of a superficial similar-
ity, it was once believed that the Artamidae might be related to
the Vangidae. However, there is no anatomical evidence avail-
able in favor of such an association and much zoogeographical
and biological evidence which contradicts it.
3. Bombycillidae. Should this family be transferred from a
position near the Sturnidae to one nearer to the Laniidae?
Answer: Yes. The Bombycillidae are presumably more closely
related to some of the families in group 1 than to those in
group 3.
4. Ptilogonatidae and Dulidae. Should not these two groups
be retained as families?
Answer: They are better placed as subfamilies for the reasons
stated above under B.
dD. Pityriasis. Does the inclusion of this genus in the Prionopi-
dae reflect true relationship ?
Answer: Perhaps not, but with the available evidence it is not
possible at present to make a satisfactory decision on relation-
ship. There is no support for the belief that it might be related
either to the Starlings or to the Shrikes (in the restricted sense).
Since it is not advisable for the reasons stated above under B to
separate the genus in a monospecific family, it will be best to list
it as a subfamily in the Prionopidae where the genus has been
listed traditionally.
6. Estrildidae. Should they be combined with the Ploceidae
or be treated as a separate family ?
Answer: Steiner (1955) has listed much evidence indicating
that the Estrildidae deserve family ranking, indeed that they
may not even be closely related to the Ploceidae.
7. Turdidae, Sylvidae, Muscicapidae, Fringillidae, Ploceidae,
Emberizidae. Should not all these be retained as families?
Answer: For the reasons stated by Hartert, the first three
should be combined in a single family. Fringillidae and Plocei-
dae should be retained as families while the Emberizinae should
be retained as a subfamily of the New World finches. The oldest
family group name proposed for the New World finches is ap-
parently Emberizoidea (Suschkin 1925). The name of the family
then would be Emberizidae.
8 BREVIORA No. 58
The final sequence of the families of Oscine Passeres which
emerged from these discussions was submitted to the committee,
which agreed with it, except that Prof. Berlioz stated that he still
preferred a placement of the Alaudidae near the Emberizidae.
The editors of Peters’ Checklist still feel that such an arrange-
ment has less to recommend it, for the anatomical reasons stated
above. The sequence approved by the committee is as follows:
Sequence of Oscine Families
Alaudidae
Hirundinidae
Motacillidae
Campephagidae
Pyenonotidae
Trenidae
Laniidae
Prionopidae
Vangidae
Bombyeillidae
Bombycillinae
Ptilogonatinae
Dulinae
Cinclidae
Troglodytidae
Mimidae
Prunellidae
Muscicapidae
Turdinae (incl. Zeledonia )
Timaliinae (incl. Chamaea)
Paradoxornithinae
Polioptilinae (incl. Rhamphocaenus and Microbates )
Sylviinae (incl. Regulus, Leptopoecile, Lophobasileus )
Malurinae
Museicapinae
Monarchinae
Pachyeephalinae ry
Paridae
Sittidae
Sittinae
Hyposittinae
Neosittinae
Certhiidae
Dicaeidae
1956 SEQUENCE OF PASSERINE FAMILIES (AVES) )
Nectariniidae
Zosteropidae
Meliphagidae
Emberizidae
Emberizinae
Cardinalinae = Richmondeninae
Tanagrinae = Thraupinae
Tersininae
Coerebinae
Parulidae
Drepaniidae
Vireonidae (incl. Vireolanius + Cyelarhis )
Ieteridae
Fringillidae
Fringillinae
Carduelinae
Estrildidae
Ploceidae
Sturnidae
Oriolidae
Dieruridae
Callaeidae
Grallinidae
Artamidae
Cracticidae
Ptilinorhynchidae
Paradisaeidae
Corvidae
LITERATURE CITED
A.O.U. (AMERICAN ORNITHOLOGISTS’”? UNION )
1931. Check-list of North American birds. 4th ed., 526 pp., Lancaster,
Pa.
BERLIOZ, J.
1950. Jn Grassé (ed.), Traité de Zoologie, vol. 15, Oiseaux, 1164 pp.,
Paris (Systematies pp. 845-1055).
Biswas, B.
1952. A check-list of genera of Indian birds. Records Indian Mus.,
50, pt. 1, pp. 1-62 (in author’s copy).
CHASEN, F.N.
1935. A handlist of Malaysian birds. Bull. Raffles Mus., Singapore,
no. 11, 389 pp.
10 BREVIORA No. 58
Evans, A. H.
1899. Birds. The Cambridge Natural History, 9, 635 pp., London.
FUERBRINGER, M.
1888. Untersuchungen zur Morphologie und Systematik der Vogel. 2
vols., 1800 pp., 4°, Jena and Amsterdam.
Gapbow, H.
1893. In H. G. Bronn’s Klassen und Ordnungen des Thierreichs ete.
Vol. 6, 4 Abth., II, Systematischer, 301 pp. (pp. 279, 301).
1898. A elassification of Vertebrata, ete. 82 pp., London.
GARROD, A. H.
1876-1877. On some anatomical peculiarities which bear upon the major
divisions of the passerine birds. Part 1, Proc. Zool. Soe. London,
1876, pp. 506-519; Notes on the anatomy of passerine birds, pt.
2, 1877, pp. 447-453, pt. 3, t.c., pp. 523-526; pt. 4, 1878, p. 143.
HARTERT, E.
1903-1922. Die Vogel der palaarktischen Fauna. 3 vols., Berlin.
MATHEWS, E. M.
1927-1930. Systema avium Australasianarum. 2 vols., London.
Mayr, E. and D. AMADON
1951. A classification of recent birds. Amer. Mus. Novit., no. 1496,
pp. 1-42.
ORNITHOLOGICAL SOCIETY OF JAPAN (A Special Committee ed. )
1942, A handlist of the Japanese birds. 238 pp., Tokyo.
PETERS’ CHECKLIST (PETERS, J. L.)
1931-1951. Check-list of birds of the world. 7 vols. (series not completed),
Cambridge, Mass.
PORTMANN, A.
1946-1947. Etudes sur la cérébralisation chez les oiseaux. Alauda, 14, pp.
NEPANS UM Too, Melby (Cate 5 ees, jo, IMGIlelAl (Gone, &)).
R.A.O.U. (Royal AUSTRALIAN ORNITHOLOGISTS’ UNION, CHECKLIST
COMMITTEE )
1926. Official checklist of the birds of Australia. 212 pp., Melbourne.
ScLATER, W. L.
1930. Systema avium Aethiopicarum. Pt. 1, pp. 1-304 (1924); pt. 2,
pp. 305-922 (1930), London.
SHARPE, R. B.
1885. Catalogue of birds of the British Museum. 10, p. 1.
1956
SEQUENCE OF PASSERINE FAMILIES (AVES) 1]
SHARPE, R. B.
1899-1909.
A hand-list of the genera and species of birds. 5 vols., London.
SHUFELDT, R. W.
1904.
STEINER, H.
1955.
An arrangement of families and the higher groups of birds.
Amer. Nat., 38, pp. 833-857.
Das Brutverhalten der Prachtfinken, Spermestidae, als Aus-
druck ihres selbstandigen Familiencharacters. Act. XI Cong.
Int. Orn. (Basel), pp. 350-355.
STEJNEGER, L. H.
1885.
Birds. Jn Standard Natural History (J. S. Kingsley ed.),
Boston.
STRESEMANN, E.
1934.
SUNDEVALL,
1872-1873.
Aves, Handbuch der Zoologie. Kitkenthal and Krumbaeh ed., 7
(2), 898 pp., Berlin and Leipzig.
Ops
Methodi naturalis avium disponendarum tentamen. 2 pts., 4°,
Stockholm.
SUSCHKIN, P. P.
1925,
‘ToRDOFF, H.
1954.
The Evening Grosbeak (Hesperiphona), the only American
genus of a Palaearetie group. Auk, 42, pp. 256-261.
A systematic study of the avian family Fringillidae based on
structure of the skull. Mise. Publ. Mus. Zool., Univ. Michigan,
no. 81, 42 pp.
WALLACE, A. R.
1874.
On the arrangement of the families constituting the order
Passeres. Ibis, pp. 406-416.
WETMORE, A. and W. W. MILLER
1926,
1951.
The revised classification for the fourth edition of the A.O.U.
check list. Auk, 43, pp. 337-346.
A revised classification for birds of the world. Smithsonian
Mise. Coll., 117, no. 4, pp. 1-22.
saad
Vide hm oP 1 4
oy if Eid AL 4 :
M
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. SEPTEMBER 12, 1956 NUMBER 59
A NEW SUBGENUS OF CHAMAELEO FROM
RHODESIA AND NEW RACE OF MABUYA
FROM KENYA COLONY
By ArrHurR LOVERIDGE
Irifty years ago when Boulenger (1906, Ann. Mag. Nat. Hist.,
(7), 18 p. 346, fig.) described Rhampholeon marshalli from
Mashonaland, he remarked on the only occurrence of this genus
south of the Zambezi as being ‘‘of very great interest.’’ In as-
signing marshalli to Rhampholeon, Boulenger was undoubtedly
guided by his own key (1887, Cat. Lizards Brit. Mus., 3, p. 438)
to the genera of chameleons.
More recently Parker (1942, Bull. Mus. Comp. Zool., 91, p. 82)
examined marshalli for certain osteological characters, i which
respect he found it agreed with other continental species of
Rhampholeon, a group I have since suggested should be regarded
as only a subgenus of Brookesia.
Dr. V. FitzSimons (1943, The Lizards of South Africa, p.
172, pl. xxi, fig. 5) also treated marshalls as the only South Afri-
can representative of Rhampholeon. He personally collected
topotypes in Chirinda Forest, Selinda Mountain, as well as an
extensive series from Vumba Mountain, also in Southern Rho-
desia. Thirty-one of these specimens of marshalli are now in the
Museum of Comparative Zoology and were examined by me when
making a synopsis of the continental African Brookesia (sub-
genus Rhampholeon). At that time (1951, Bull. Mus. Comp.
Zool., 106, p. 182, footnote) I rejected marshalli as a Rhampho-
leon, despite its bicuspid claws, referring it to Chamaeleo on
account of its prehensile tail.
Actually the bicuspid claws (present or absent in Rhampho-
leon) constitute the only character in which marshalli differs
from the forms assigned to Chamaeleo; thus it bridges the gap
2, BREVIORA NO. 59
between Chamaeleo and the subgenus Rhampholeon of Brook-
esia.' With a view to inviting attention to the intermediate status
of this peculiar little chameleon, I suggest it should be made the
type of a subgenus of Chamaeleo, viz.
BicusptIs subgenus new
Type. Khampholeon marshalli Boulenger; known only from
Southern Rhodesia.
Diagnosis. General appearance and soft nasal protuberance
resembling that of Chamaeleo rather than Rhampholeon. Seales
on soles smooth; claws bicuspid; tail prehensile, half to two-
thirds the length of head and body. The tail is included in the
length of head and body from 1.6 to 2.1 times, with an average
of 1.86 times for our entire series of 31 marshalli. The largest
of the series, a topotypic gravid 2 (M.C.Z. 44445), measures
103 (68 + 35) mm.
The only Brookesia (subgenus Rhampholeon) with a tail any-
thing like as long proportionately as that of marshalli, is B. k.
kersten (Peters), the claws of whose forefeet have a secondary
cusp — though a secondary cusp is lacking on the claws of the
forefeet of its northern representative B. k. robecchi (Bou-
lenger).
MABUYA BAYON!I KENIENSIS subsp. nov.
When the Museum of Comparative Zoology received a pair of
typical Mabuya bayonii Bocage from Chitau, Bihe District,
Angola, in 1936, it was immediately apparent that its Hast
African representatives should be separated, if only on the
basis of their strongly tricarinate dorsal scalation. The matter
was deferred until such time as a revision of all African Mabuya
could be undertaken, or a decision reached as to whether bayonu
itself should be treated as a race of gravenhorstu Dumeril and
Bibron of Madagasear. As time for any such thorough investiga-
tion is lacking, I propose the name Mabuya bayonw kenvensis
subsp. nov.
Holotype. Museum of Comparative Zoology No. 29662, an
adult ¢, from the northern Uaso (Guaso) Nyiro, Sotik, Kenya
1As Angel (1942, Mem. Acad. Malgache, 36, pp. 154, 178) designated no type
for his genus Hvoluticauda, I suggest that Brookesia nasus Boulenger be so
regarded.
1956 LIZARDS FROM KENYA AND RHODESIA 3
Colony. Collected by the Smithsonian-African Expedition, 1909.
Paratypes. A specimen from Mount Kenya (U.S.N.M. 40710),
another from Wambugu (U.S.N.M. 40781), a third from Lake
Sergoit (Sirgoit: U.S.N.M. 42024), and three others (U.S.N.M.
40947-8 and M.C.Z. 29663) with the same data as the type.
Unquestionably the juvenile from a salt marsh on the Loita
Plains, recorded by Angel (1922) is referable to this Kenya
race, as are also four listings of bayoniti by me (19238; 1924;
1929; 1937). Less certain as to which race it should be assigned
is Oscar Neumann’s specimen from ‘‘Sero’’ (? Ssera, Lake
Tanganyika) referred to Mabuia bayoni (sic) by Tornier, 1896
(Die Kreichthiere Deutsch-Ost-Afrikas, Berlin, p. 42); this
formed the basis of subsequent listings by Tornier (1897; 1900)
and Nieden (1913), who added a series taken between Lake
Victoria and Nguruman (i.e, Neurumani).
For the loan of the paratype material in the United States
National Museum, as well as an Angolan b. bayonii (U.S.N.M.
26389) for comparative purposes, | am indebted to Dr. Doris
M. Cochran.
Diagnosis. Dorsals tricarinate, whereas in b. bayoni they are
strongly quinquecarinate. Certain other characters may prove
to have an average difference, but this cannot be demonstrated
until more material of typical bayoni is available.
Description. Based on the holotype (variations of paratypes
are placed in parentheses). Supranasals in contact behind the
rostral; centre of nostril in advance of, even though slightly,
the vertical of the suture between rostral and first labial; post-
nasal in contact with first labial only (except on right side of
U.S.N.M. 40710, and both sides of U.S.N.M. 40781, where it
touches the second labial also) ; anterior loreal in contact with
first and second labials (or occasionally second only); supra-
oculars 3, the first and second being fused into a single shield
(3-4 in paratypes) ; supraciliaries 4-5 (3-5), first largest; sub-
ocular narrowed inferiorly, reaching the lip between the fifth
and sixth upper labials (in all) ; lower eyelid with a transparent
disk that is subequal to, or shghtly larger than the ear-opening,
which has 2 (2-3) more or less acuminate lobules projecting from
its anterior border; frontonasal in contact with the first (as it is
fused with the second) supraoecular (though usually not in con-
4 BREVIORA NO. 59
tact with the first) ; frontoparietal single, larger than the inter-
parietal, which separates the parietals completely; nuchals
multicarinate.
Midbody seale-rows 36 (34-36), dorsals strongly tricarinate;
preanals not or but shehtly enlarged; scales on soles not or but
shghtiy pointed; subdigital lamellae smooth; toes of the ad-
pressed hind limb meet the finger tips (or fail to meet, or extend
to wrist) of the backward-pressed forelimb.
Color. Above (pale or dark) olive brown; dorsum with several
longitudinal series of black (and white) flecks (or ocelli); a
cream (or white) dorsolateral line extends backwards from the
supraocular region to some distance along the tail; from the
white labials a lateral line extends along the flanks on to base
of tail. Below, white, uniform.
Size. Total length of type ¢ (M.C.Z. 29662), 148 (62 + 86)
mm.; of paratype @ (M.C.Z. 29663), 160+ (82+ 787) mm.,
tail-tip missing.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, MAss. SEPTEMBER 12, 1956 NUMBER 60
A NEW SPECIES OF AGRIOGNATHA FROM
JAMAICA, B. W. I.
By ArtHuur M. CHICKERING
In connection with my study of the genus Tetragnatha
Latreille, 1804 from Panama I have also had the opportunity to
study several species believed to belong to the rather poorly-
known genus Agriognatha O. P. Cambridge, 1896. Apparently
there is at least one species in Panama not included among those
recognized by F. P. Cambridge in 1903 and not described by any
later author. It is expected that a description of this new species
will be published shortly.
For some time prior to her death in 1953, Miss E. B. Bryant
of the Museum of Comparative Zoology at Harvard College had
been engaged in a study of a fairly representative collection of
spiders from Jamaica, B. W. I. This collection had come from
several sources but it had been assembled largely through the
interest of Mr. C. Bernard Lewis, Curator of the Science Museum,
Institute of Jamaica, Kingston, Jamaica. The collection is now
in my possession ond it is hoped that considerable time can we
devoted to its study during the next few years.
Among the new Tetragnathinae selected by Miss Bryant for
description was the species here described in accord with my
usual procedure. I have thought it appropriate to name the
species in honor of Miss Bryant who for many years gave un-
stintingly of her time and energy to the care of the large collec-
tion of spiders in the Museum of Comparative Zoology at Har-
vard College. So far as I know, this is the only species of
Agriognatha reported from the island of Jamaica.
>; BREVIORA NO. 60
The holotype and allotype will be deposited in the collection of
the Museum of Comparative Zoology at Harvard College.
Genus AGRIOGNATHA O. P. Cambridge, 1896
fo)
The genus Agriognatha O. P. Cambridge, 1896 was based upon
a male from Costa Rica. In 1897 Simon described a species from
St. Vincent as Cyrtognatha serrata, apparently following Key-
serling who established the genus Cyrtognatha in 1881. F. P.
Cambridge had Simon’s C. serrata for study and regarded it as
congeneri¢ with A. bella (O. P. Cambridge) from Costa Rica. He
considered it probable that Cyrtognatha Keyserling was the same
as Agriognatha O, P. Cambridge. He also noted that the name
Cyrtognatha was preoccupied and he placed all species regarded
by the authors as belonging to either genus in the genus Agrio-
gnatha although he had doubts regarding the proper position
of A. lepida (O. P. Cambridge). Petrunkevitch (1911) placed
all five species then known in the genus Cyrtognatha. Roewer
(1942) has retained this practice. I have not seen Cyrtognatha
globosa Petrunkevitch, described from a female taken in the
San Lorenzo River region of Panama but it seems unlikely that
this species belongs in the genus Agriognatha as the latter is
now understood. It now seems to the author of this paper that
the two genera under consideration here are distinct but that
all species recognized by Miss E. B. Bryant (1940, 1945) from
Cuba and Hispaniola can safely be placed in the genus Agrio-
gnatha. This also seems certain for the species from Jamaica
deseribed in this paper.
AGRIOGNATHA BRYANTAE SP. NOV.
(Figures 1-5)
Male holotype. Total length 3.965 mm. (chelicerae not ex-
tended anterior to head region). Carapace 1.745 mm. long, 1.30
mm. wide opposite second coxae where it is widest; rather sharply
narrowed opposite first coxae; median thoracic pit rather shallow
and broad (considerably different from that seen in specimens
from Panama).
Eyes. Hight in two rows as usual in the genus; LE extend
slightly beyond border of head. Viewed from above, anterior
1956 AGRIOGNATHA FROM JAMAICA 3
row strongly recurved, posterior row moderately recurved ;
viewed from in front, anterior row moderately recurved, posterior
row slightly procurved, both measured by centers; central ocular
quadrangle wider in front than behind in ratio of 9 : 8, as wide
in front as long. Ratio of eyes AME : ALE : PME :PLE=9:
8.5 :8.5:7. AME separated from one another by 11/9 of their
diameter, from ALE by 5/3 of their diameter. PME separated
from one another by nearly three fourths of their diameter, from
PLE by about 7/4 of their diameter. Laterals separated from
one another by a little less than the radius of PLE. Lateral eyes
on a moderately prominent tubercle. Height of elypeus equal
to diameter of AME.
Chelicerae. Very divergent, nearly horizontal; in place of
the basal boss there is a prominent tubercle from which a ridge
extends nearly to the base of the fang; basal segment of each
chelicera 1. 235 mm. long and, therefore, nearly two thirds as
long as carapace. Fang long, slender, somewhat sinuous, and
with a prominent cusp on the inner surface just proximal to the
middle. The fang groove has a robust non-bifureate tooth near
the base of the fang on the promarein together with three small
teeth; the retromargin has four small teeth as shown in Figure 1.
Maxillae. Essentially parallel but with distal third slightly
curved outward and considerably widened; shghtly more than
twice as long as lip.
Lip. Considerably wider at base than long; sternal suture
gently procurved ; sternal tubercles prominent at ends of sternal
suture.
Sternum. Cordiform ; moderately convex ; slightly longer than
wide between second coxae where it is widest; moderately ex-
tended between all coxae; terminates in a blunt point between
bases of fourth coxae which are separated by slightly more than
one third of their width.
Legs. 1243. Width of first patella at ‘‘knee’’ .220 mm., tibial
index of first leg 5. Width of fourth patella at ‘‘knee’’ .165 mm.,
tibial index of fourth lee 7.
4 BREVIORA NO. 60
Femora Patellae Tibiae Metatarsi Tarsi Totals
(All measurements in millimeters )
It. 3.770 810 3.770 4.192 1.365 13.907
2. 2.990 745 2.600 2.900 1.105 10.340
3. 1.560 390 910 1.170 .640 4.670
4. 2.210 455 1.820 2.210 780 7.479
Palp 726 185 198 aes .682 1.791
Many true spines as well as hairs on all legs. It is considered
unnecessary to record all spines, hence, only those which seem
to be more or less distinctive are emphasized here. First lee:
there is a row of four short but fairly robust ventral spines near
the base of the femur together with a row of seven very short
but fairly robust retrolateral spines extending through the prox-
imal two thirds of the segment and also a long robust retrolateral
spine near the distal end of this segment; the patella has the
characteristic retrolateral chitinous ridge; the metatarsus has
a row of about 25 short ventral robust spines extending nearly
throughout the length of the segment. The second leg is essen-
tially like the first in respect to the characteristics noted with the
specialized robust retrolateral spine near the distal end of the
femur arising from a raised base. The expected prolateral
trichobothria on both third and fourth femora are present but
are much less conspicuous than in the Central American species
thus far observed.
Palp. Complicated; somewhat difficult to describe because of
numerous associated apophyses. The paracymbium appears to
have a somewhat characteristic terminal enlargement. Near the
tip of the cymbium there is a small pit, regarded as a type of
sense organ. The same type of presumed sense organ has been
noted on related genera and species. See Figures 2 and 3 for
details of palpal structure.
Abdomen. Unnotched at anterior end which is_ bluntly
rounded ; 2.34 mm. long; 1.06 mm. wide near middle; with spir-
acle near base of spinnerets; with a well defined colulus, oval in
outline; with genital fold (Fig. 5) only slightly posterior to
openings of book lungs.
Color in alcohol. Chelicerae brown; legs yellowish brown with
many irregular darker areas especially on dorsal surfaces.
Carapace: with a narrow brown marginal stripe from opposite
1956 AGRIOGNATHA FROM JAMAICA 5
PE to opposite third coxae where it becomes a wide stripe which
continues to posterior border; also with an irregular broad
median dark stripe from PME to median thoracic pit. Abdomen :
a broad serrated median dark gray stripe extends from base to
spinnerets; on each side of this there is an irregular light stripe
with scattered silvery spangles; the lateral sides are dark gray
with oblique extensions dorsally; the venter is yellowish with a
row of small silvery spangles on each side.
Female allotype. Total length 4.94 mm.; including the cheli-
cerae 5.46 mm, Carapace 1.95 mm. long, 1.48 mm. wide opposite
interval between second and third coxae where it is widest ; other
features essentially as in male.
Eyes. Central ocular quadrangle about as wide in front as
long. Ratio of eyes AME : ALE : PME : PLE =9:10:9:8.
AME separated from one another by 10/9 of their diameter, from
ALE by slightly less than twice their diameter. PME separated
from one another by 10/9 of their diameter, from PLE by twice
their diameter. Laterals separated from one another by about
1 of the diameter of ALE. Height of clypeus equal to slightly
more than 7% of the diameter of AME. Other features essentially
as in male.
Chelicerae. Robust ; only she¢htly divergent and porrect; basal
segment 1.04 mm. long; fang regularly curved and without
special features; promargin of fang groove well marked, with
three teeth; retromargin of fang groove with four teeth (Fig.
+); some variation in placement and relative sizes of the teeth
has been noted and must be expected among paratypes.
Maxillae, Lip, and Sternum. Essentially as recorded for the
male.
Legs. 1248. Width of first patella at ‘‘knee’’ .264 mm., tibial
index of first leg 7. Width of fourth patella at ‘‘knee’’ .198 mm.,
tibial index of fourth leg 10.
Femora Patellae Tibiae Metatarsi Tarsi Totals
(All measurements in millimeters )
ite 3.380 845 3.140 3.412 1.365 12.142
2, 2.600 .780 2.190 2.632 1.040 9.242
3. 1,495 435 845 975 715 4.465
4, 2.210 487 1.495 1.820 .780 6.792
6 BREVIORA NO. 60
All legs with stout spines and hairs but the spines seem not to
be specialized as in the male. The trichobothria on the third and
fourth femora essentially as recorded for the male.
Abdomen. Very gibbous dorsally just behind the middle but
this swelling seems to be completely lacking in some female para-
types; 3.575 mm. long; 2.080 mm. wide opposite the gibbosity
where it is widest ; genital groove as shown in Figure 5.
Color in alcohol. Chelicerae much lighter than in male with
mottled gray markings in front in basal half. Otherwise essen-
tially as in male.
Type locality. Male holotype and female allotype from Hard-
war Gap, Jamaica, B. W. I., June 27, 1954. Several paratypes
of both sexes from the followimg localities: Blue Mountains,
August, 1934 (Darlington) ; St. Andrews, Clydesdale, July, 1950
(Bengry); Hardwar Gap, June, 1954'; Hanover, Askenish,
trail to Dolphin’s Head, June 24, 1954.
BIBLIOGRAPHY
BRYANT, ELIZABETH B.
1940. Cuban spiders in the Museum of Comparative Zoology. Bull.
Mus. Comp. Zool., vol. 86, pp. 249-532, pls. 1-22.
1945. The Argiopidae of Hispaniola. Bull. Mus. Comp. Zool., vol. 95,
no. 4, pp. 359-418, pls. 1-4.
CAMBRIDGE, O. P. and F. P. CAMBRIDGE
1889- Arachnida-Araneida. Vols. I and II in: Biologia Centrali-
1905. Americana. Dulau & Co., London.
KEYSERLING, GRAF EUGEN VON
1881. Neue Spinnen aus Amerika. Verhandl. zool. bot. Ges. Wien, vol.
ol, p. 276.
PETRUNKEVITCH, ALEXANDER
1911. A synonymic index-catalogue of spiders of North, Central, and
South America, etc. Bull. Amer. Mus. Nat. Hist., New York,
vol. 29, pp. 1-809.
Arachnida from Panama. Conn. Acad. Arts and Sciences, vol.
27, pp. 51-248, 157 figs.
—
is)
ho
Or
1Since the completion of this paper two females and one adult male believed to
belong to this species have been collected at Hardwar Gap, Jamaica, in July,
1955, by Dr. Allan F. Archer who has kindly loaned these specimens for examina-
tion,
~
1956 AGRIOGNATHA FROM JAMAICA
ROEWER, C. F.
1942. Katalog der Araneae. Vol. 1. Bremen.
Simon, E.
1897. On the spiders of the Island of St. Vincent. Part III. Proe. Zool.
Soe. London, pp. 860-890.
External Anatomy of Agriognatha bryantae sp. nov.
Fig. 1. Male chelicerae, from in front.
Figs. 2-3. Tarsus of male palp; lateral and ventral views.
Fig. 4. Female cheliceral teeth.
Fig. 5. Genital furrow of the female.
beset ait hin: STA techie “ela
Gey, kate aR ales
AN EN rt aioe peaianay
7 D u Py rom a
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ry
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a are
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. SEPTEMBER 14, 1956 NUMBER 61
ON REGENERATION BY EARTHWORMS OF A
SPECIES OF THE LUMBRICID GENUS
DENDROBAENA EISEN 1874.
By G. E. Gatrs
Regeneration, either cephalic or caudal, after natural or ex-
perimental amputation, for any species of Dendrobaena has not
been recorded hitherto, though several of the head regenerates
attributed by Morgan to Eisenia foetida (ef. Gates, 1953, 1954)
probably were D. octaedra. Some data as to regeneration in
another species of the genus can now be presented.
The worms, presumably all of an athecal morph frequently
called Bimastos tenwis in the past, were clitellate and were
secured early in the summer in Michigan. Amputation was
without anesthesia. Amputees were kept in damp leaves at 22° C
for the periods indicated in the tables.
The author’s thanks are extended to Prof. Murchie for making
this material available for study.
Anterior regeneration in D. rubida (Savigny, 1826)
All regenerates (cf. Table I) are hypomeric. Metamerism in
most regenerates deviates more or less from normal. Regenera-
tion of a head clearly is possible at all levels back to and including
13/14. The regenerate at 17/18 lacks normal anal or buccal seulp-
turing distally and, since nephropores are unrecognizable, pro-
vides no external indications as to whether it is of cephalic or
caudal organization.
Abnormal metamerism and hypomery at anterior levels, in
certain other earthworm species studied by the author, resulted
from unfavorable conditions either in the external or internal
environments. Accordingly, metamerically normal regenerates at
i)
BREVIORA NO. 61
all levels to 15/14 and equimery at least to 5/6 can be expected
in D. rubida when conditions are more favorable.
The level at which the indeterminate monstrosity was devel-
oped is elose to, if not actually in, a region where regenerative
capacity in E. foetida (ef. Gates, 1949-1950) is bipotential.
Monstrosities such as the one under consideration often are pro-
dueed in morphogenetic regions of dual capacity. However, the
data now available as to the results of posterior amputation in
D. rubida seareely warrant anticipation of heteromorphiec tail
regeneration in that species. Presumably then, a regenerate de-
veloping at 17/18 in optimal conditions will be normally cephalic.
Especially noteworthy, even in these few instances, is absence
of any indication of decline in number of segments regenerated
as level of amputation moves posteriorly (cf. Gates, 1949, p.
SWE
TABLE I
Head regeneration in Dendrobaena rubida
i at Re £ 2 L
EP VERSE TG iy at in wae 2 2
1 2/3 ii 2 19
2 3/4 1 3 19 Metamerism still indistinct in regen-
erated left halves of iv-vi.
3 5/6 3 4 32 Regenerate metamerism nearly nor-
mal.
4 5/6 4 4 32 Regenerate metamerism nearly nor-
mal but 6/7 abnormal in regen-
erated right halves of vi-vii.
5 6/7 3 4 32 Metamerism nearly normal.
} 6/7 4 -3 19 Metamerism probably was develop-
ing abnormally.
it 7/8 2 4 37 Metamerism not quite normal. Re-
generated ventral half of viii ab-
normal.
REGENERATION IN DENDROBAENA 3
a ! n & fe 3 Nn
a2 Big bea 8 OB) Pate %
iat WAS epi ice dsber eGR Wes Ue F
ReGen ce lone e002 seh ay ote g
WM ay te
9 7/8 3+ 3 19 Proximal portion of regenerate too
short for a normal segment.
10 8/9 4 3 19 Metamerism apparently developing
abnormally in regenerated right
halves of ix-x.
11 8/9 + + 32 Regenerate segments ii-iv not quite
normal.
12 8/9 4 4 32 Segment iv too large. Excised dor-
sal half of ix had been regenerated.
13 8/9 4 -3 19 Metamerism apparently normal.
14 aly aS fe ieee oc: 32 Regenerate metamerism abnormal.
15 13/14 5 2 =3 32 Metamerism in regenerated left
halves of xiv-xvi probably develop-
ing abnormally.
16 13/14 5-6? —3 30 Metamerism rather indistinct in re-
generated portions of xiv-xv, ven-
tral half of xiv and a small portion
of xv.
17 17/18 4-6? —3 32 Regenerate terminating distally in
rounded knob like a rudimentary
prostomium but towards the ven-
tral rather than the dorsal side.
Exeept as indicated otherwise above, excisions appear to have been transverse
and along intersegmental furrows.
Stages.
1. Prostomium and buceal invagination as yet unrecognizable.
2. Prostomium and mouth developed.
3. Intersegmental furrows demarcate regenerate into segments.
—3, Furrows still indistinct.
Pigment present but obviously different from that of substrate.
5. Setae and nephropores recognizable.
6. Pigment now like that of substrate, external stigmata of regen-
eration, except for typical metameric anomalies, unrecogniz-
able.
4 BREVIORA NO. 61
Posterior regeneration in D. rubida
An anus of more or less normal appearance had been acquired,
at end of a July-August period of nineteen days, by the six
worms from which a posterior portion of the intestinal region
(ef. Table IIL) had been excised. Healing probably had been
enteroparietal. The new anal region had not yet been delimited
from the last substrate segment, by development of an interseg-
mental furrow, in Nos. 4-6 where little or no indication of re-
organization is externally recognizable. A very small anal re-
gion, in No. 2, is demarcated by an interseemental furrow and
again there are no other external indications of reorganization.
A terminal portion of No. 1, presumably comprising only the
last segment, at time of preservation was being reorganized.
Completion of the process apparently under way probably would —
have resulted in development anteriorly of eight new setal fol-
licles with setae and appearance of an intersegmental furrow
demareating a terminal anal portion from a metamere with the
usual stigmata of regeneration. Reorganization, instead of re-
generation, would then have been recognizable externally only
if the original nephropores had been retained in the smaller
daughter segment or if some of the pigment had escaped lysis.
Reorganization, accordingly, had been most drastic and had
taken place most rapidly at the anteriormost level of amputation.
Evidence as to tail regeneration has been sought in several
hundred individuals of D. rubida that have been available from
various states and from other countries. Many of these worms
certainly are posterior amputees and some of the others, obvi-
ously brevicaudate, very probably are. No tail regenerates were
found. Externally recognizable indications of reorganization
were noted but rarely, and in each case there could have been
produced, in addition to the anal region, only one or two new
segments.
Homomorphie tail regeneration, from a growth zone of rapid
segment production (Gates, 1948) obviously does not, usually,
follow posterior amputation in this species.
1956 REGENERATION IN DENDROBAENA By)
TABLE II
Results of posterior amputation in D. rubida
ey A ne L.
ze ssa 03 BB 2
Le 2a aSde =
ae eam e % a
=) Ha az e 2 Pe
i 42/43 ? Terminal substrate segment has lost much
pigment, setae and nephropores.
2 54/55 ? An intersegmental furrow delimits small
white anal region from last substrate
segment.
3 65/66 12 Same as in No. 2.
| 80/81 6 Anal region not delimited from last sub-
strate segment which still has setae and
nephropores.
5 85/86 fi Anal region not delimited from last sub-
strate segment where some of the setae
still are present.
6 87/88 10 Anal region not delimited.
Time allowed for regeneration, 19 days.
Discussion
Homomorphic tail regeneration does not necessarily follow
immediately after amputation and may not begin until nine
months later. The process gets under way in some earthworm
species, regardless of time of amputation, only when the ‘‘inter-
nal environment’’ permits. Differences in that environment may
well be responsible for discordant results obtained by different
investigators who have used the same species. As the importance
of the unknown factors of the internal environment of earth-
worms seems not to have been appreciated, the following in-
stance is worthy of record.
Individuals of Eisenia foetida, in the author’s earlier studies
(Gates, 1949-1950), after removal of the posterior portion at
6 BREVIORA No. 61
levels behind 40/41, almost always regenerated promptly and
in all seasons of the year. Several dozen specimens of that species,
apparently in good condition, from three localities, were deprived
of their tails in the region between 40/41 and 50/51 on three
occasions in the last five years, to provide material for a school
demonstration. All of the worms survived the operation. Not
one showed any indication of formation of new segments during
a period of several weeks though the external environment was,
so far as could be determined, the same as before.
Summary
Cephalic regeneration in an anterior direction can be expected,
in D. rubida in optimal conditions, at all levels back to 17/18,
with equimery back to 5/6. Caudal regeneration in a posterior
direction, from a growth zone of rapid segment formation, can-
not be expected ordinarily if at all. Instead, the terminal sub-
strate segment may be reorganized into an anal region and one
or two metameres with some of the stigmata of regeneration, the
reorganization possibly being more drastic and more rapid
when at more anterior levels.
REFERENCES
GATES, G. EH.
1948. On segment formation in normal and regenerative growth of
earthworms. Growth, 12: 166-180.
1949-1950. Regeneration in an earthworm, Hisenia foetida (Savigny) 1826.
T-I1l. Biol. Bull., $6:129-139; 98:36-45; 99:425-438.
1953. On regenerative capacity of earthworms of the family Lumbri-
cidae. Am. Midland Nat., 50: 414-419.
1954. Anterior regeneration in a sexthecal species of lumbricid earth-
worm. Breviora, 27:1-5.
fe ae
A bl dea %
“
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. OcTOBER 3, 1956 NUMBER 62
A THIRD LEAF-NOSED SPECIES OF THE LIZARD
GENUS ANOLIS FROM SOUTH AMERICA
By JAMES A. PETERS
Brown University, Providence, Rhode Island
and
GusTAvo OrcEkEs-V.
Escuela Polyteenica Nacional, Quito, Ecuador
It gives us considerable pleasure to be able to announce the
discovery of another specimen of the leaf-nosed anoles. It is the
first time that leaf-noses have been known to occur outside of
the Amazon Basin. The locality from whence it came is on the
Pacific side of the Andes in the hot and humid **‘Choco’’ portion
of northwestern Ecuador. While the new individual is obviously
closely related to the holotype of the species described by Myers
and Carvalho (1945), the differences are sufficiently great that
we feel justified in calling it a new species.
Since the original specimen was collected, the Escuela Poly-
tecnica has sent several other collectors into the area and the
senior author made several trips to the type locality during the
summer of 1954 when he obtained comparative material of other
species of the genus Anolis,’ but no one has yet been able to
obtain additional representatives. Nothing is known as yet as to
the habits of these unusual animals or the use to which the
rostral appendage is put.
1 Field work in Ecuador during 1954 by the senior author was supported by
a grant from the Penrose Fund of the American Philosophical Society.
2 BREVIORA NO. 62
ANOLIS PROBOSCIS, new species
Type. Museum of Comparative Zoology No. 54800, a mature
male from the neighborhood of Cunuco, a small town at 1200
meters elevation, five kilometers northwest of Mindo, on the
south bank of the Rio Mindo, a northern tributary of the upper
Rio Blanco, in Pichincha Province, Ecuador, collected by An-
tonio Proaho, during April 27-29, 1953.
Diagnosis. This new species is distinguished from all other
species but one in the genus Anolis by the presence of an elon-
gated, fleshy appendage, which is about as long as the head, on
the snout. The only other species with a similar appendage is
Anolis phyllorhinus Myers and Carvalho, from which it differs
in having a serrated edge on the rostral appendage, 10-11 upper
labials to a point below the center of the eye, 9-10 lower labials,
and a median dorsal row of scales which is produced into a
serrated dorsal erest, as well as other characters mentioned below.
Description. Snout with an elongate, fleshy appendage pro-
jecting anteriorly, with a distinct downward curve; length of
appendage approximately equal to length of head, measured
from tip of rostral to ear opening. Appendage arises approxi-
mately half way between eye and nostril, on midline; strongly
serrate alone upper margin to level of rostral, serration less
marked but still serrate to tip, both above and below; nine
scales wide at level of rostral, tapering to three at tip; rather
fleshy and rounded at snout, tapering to thin and vertically com-
pressed at tip. Scales on dorsum of head smooth, pavimentose,
irregular. No distinct depression in frontal region, frontal ridges
almost obsolete, faintly indicated from middle of supraorbital
semicircles to frontal area, where they end at level of anterior
margin of eye. Canthus rostralis not strongly angulate but
rather rounded, with 5-6 canthal scales from nostril to eye; area
between canthi oceupied by irregular, smooth scales, 8-9 in a
straight line between canthi at level of origin of nasal appendage
and approximately the same number between the posterior ends
of the canthi, due to gradual increase in size of scales in frontal
area. Scales of supraorbital semicircles large, somewhat scari-
fied, slightly angulate; not in contact on midline, separated by
two rows of smaller, pavimentose scales. Supraorbital scales in
contact with semicircles, no granular scales separating them;
1956 LEAF-NOSED ANOLE FROM SOUTH AMERICA 3)
three or four scales in center of supraorbital area are markedly
larger than others, but no rows or series are formed. Inter-
parietal large, considerably larger than ear opening; separated
from supraorbital semicircles by two rows of scales.
Rostral completely horizontal below appendage, extending
well beyond lip line. Nostril sunk in a single scale. Two seales
between canthus rostralis and upper labials anteriorly, four or
five posteriorly, no distinguishable horizontal rows of loreal
scales. Last scale in canthus rostralis followed by a single super-
ciliary, which is in turn followed by the granular area of eyelid.
Granules of eyelid continuous with smaller scales of supraocular
region; these two areas are not separated by a series of super-
ciliaries. Single row of three or four enlarged subocular seales
from loreals to granular postocular region and between the
eranules of eye and the upper labials; terminal scale of this row
enters the labial row on both sides, and on one side extends to
the lip line. Temporal region with many small, pavimentose
scales, not arranged in rows or series. Ear opening quite small.
Eleven upper labials on left side, ten on right to below center of
eye, one additional labial behind eleventh on left, one right sub-
ocular enters labial row, followed by granular scales. First labial
on both sides wedged between rostral and second labial, lower
edge only partially on lip line.
Mental almost completely divided on midline, suture incom-
plete anteriorly only. Nine lower labials on left, ten on right.
Seven sublabials on each side, anteriormost in contact with lower
labials, posterior four or five separated from labials by smaller
scales; sublabials reduced in size posteriorly, gradually merge
with other chin scales, which are small, numerous, and pavi-
mentose or occasionally granular.
30dy very shehtly compressed, vertebral angle comparatively
sharp. Dorsal crest of raised denticulations runs down spine
from head onto tail, with 85 spines from back of head to level
of posterior insertion of hind limbs. In several instances adjacent
denticulations are fused, with sutures still visible. Seales on
dorsum of body pavimentose, quite small, smooth, no enlarged
paravertebral rows. Lateral scales approximately equal in size
and appearance to dorsals. Granular scales, i.e., scales with raised
centers rather than a flat surface, are present on neck, shoulders,
4. BREVIORA NO. 62
and above hind legs. Ventral scales considerably larger than
dorsals and laterals, smooth, for most part in imbricate rows.
Seales on throat quite granular except on moderately developed
evular fold, where the scales are larger and quite imbricate. Im-
bricate scales of belly pass into pavimentose or even granular
scales at level of pelvis, and remain so to anus. No enlarged
preanal scales; no femoral or preanal pores.
Seales on all parts of limbs and feet subequal, pavimentose or
slightly granular. Seales on dorsum of digits expanded, much
wider than long, extend across dorsal surface of digit; smaller
scales on lateral edges of digits, which are dorso-laterally flat-
tened ; expanded scales on ventrum of digits pavimentose at base
of toe, lamellar on proximal part of toe. Lamellar counts are
shown in Table 1.
Tail very strongly compressed, vertebral angle sharp, with a
prominent dorsal crest. Seales at base of tail smooth, pavi-
mentose, and irregularly arranged; gradually become larger,
squarish, and arranged into vertical series, but still pavimentose
at level of posterior end of hemipenial sheath. Two large, smooth
scales form base of postanal hemipenial pockets. At this point
the scale rows begin to become more prominent and imbricate,
forming whorls; lowermost rows of scales begin to have keels at
about ¥ of length of tail, keels invade lateral rows until at 1%
length of tail all rows are keeled; keels increase in prominence
until at end of tail there are four pronounced edges formed by
keels on four rows of scales.
TABLE 1
Fore Foot Hind Foot
Lett SAUNA) 4-11-19-29-18
tight 5-11-17-17-10 4-12-14-30-18
Seale formulae for digits in holotype. All enlarged scales on the venter
of each digit were counted, beginning with the first distinctly enlarged,
pavimentose scale at the base of the toe and continuing to the tip of the
expanded lamellar portion. Scales on the most distal, curved phalanx, which
arises dorsally from the lamellar portion of the toe, are not included.
Dorsal ground color (in aleohol) a dull gun-metal blue, with
1956 LEAF-NOSED ANOLE FROM SOUTH AMERICA 5
irregular, parallel, horizontal black streaks along vertebral mar-
ein. These streaks vaguely line up and form a pair of lateral
bands between the limbs. Dark brownish-black spot at shoulder.
Limbs and lateral surfaces spotted with hght yellowish-white ;
limbs vaguely barred with darker blue; skin in interstices be-
tween scales of dorsum of digits light, giving appearance of
bandine on fingers and toes. Dorsum of head unicolor, as back ;
temporal region and lips hghter, with faint reddish or purplish
tinge and a marked light spot over the ear opening. Ventral
surfaces generally lighter, with vague reddish tints on chest and
chin; belly stippled with light spots. Venter of limbs heavily
spotted with white proximally, becoming totally white on foot.
Tail alternately barred with gun-metal blue and blackish, bands
approximately equal in width.
Measurements (in millimeters). Total length (not including
rostral appendage), 171; body length (rostral to vent), 74;
tail length, 97; head length (rostral to ear opening), 23; head
width at widest point, 11; length of rostral appendage, 23;
length of fore limb, 31; length of hind limb, 42.
Remarks. Since only a single male of A. proboscis is known,
the possibility exists that the specimen belongs to a previously
deseribed Ecuadorian form, known either solely from females or
from juvenile males, both of which might lack the rostral append-
age. Since the species occurs in a very distinctive biotic region,
the Choco of Eeuador, Colombia, and Panama, it is also possible
that it might be the first Ecuadorian representative of a Choco
species. There are seventeen species which have been described
from type localities within the biotic area, Of the sixteen that
are currently considered valid species, eight (chloris Bler.,
culaemus Bler., festae Peracca, gracilipes Blgr., granuliceps
Bler., latifrons Berthold, maculiventris Blgr., and peraccae
Bler.) have been previously recorded from Eeuadorian localities.
While in the Choco area of Heuador in 1954, the senior author
collected four specimens of A. breviceps Blgr., three individuals
of A. peraccae Bler., and one of A. festae Peracca, at Hacienda
Equinox, which is 30 kilometers northwest of Santo Domingo de
los Colorados. A. breviceps had not been known from Eeuador.
Seven additional species have been recorded in the literature as
taken from Eeuadorian localities within the Choco area, although
6 BREVIORA NO. 62
their type localities are elsewhere. Two other taxons, aequatori-
alis Werner and irregularis Werner, must also be considered,
since they were described from Eeuador with no added informa-
tion, and no additional specimens have been taken to establish
a range for the species.
There are sixteen species, including albi Barbour, bitectus
Cope, breviceps Blgr., fasciatus Blgr. (with its synonyms elegans
Blgr. and irregularis Werner), festae Peracea, fraseri Giimther
(with its synonym devillei Blgr.), gracilipes Bler., granuliceps
Blgr., latifrons Berthold (with its synonym princeps Bler.),
lemniscatus Blgr., lemurinus Cope, macrolepis Bler., maculi-
ventris Bler., notopholis Bler., palmeri Blgr., and peraccae
Blgr., for which both males and females are known, and for
which available specimens or descriptions provide many and
obvious differences from A. proboscis. Females only are known of
the species anton Blgr. and ventrimaculatus Bler., but the
descriptions given for the types shows that these species are
quite different from A. proboscis. Only males of aequatorialis
Werner, chloris Bler., eulaemus Bler., rosenbergi Bler., and
tropidogaster Cope have been described in the literature, but
here again each appears to be quite distinct. The sex was not
given in the original descriptions of binotatus Peters, gemmosus
O’Shaughnessy, and lionotus Cope, but they are sufficiently well
deseribed to eliminate them as available names for A. proboscis.
Myers and Carvalho (1945, p. 7) felt that the relationships of
their new species were with the punctatus group of the genus
Anolis, and this is undoubtedly true also of proboscis. They
discussed A. punctatus, A. boulengeri, A. transfasciatus and
A. nasofrontalis as probable members of the species group. They
had no specimens of A. boulengeri available, and the specimens
called A. punctatus were only tentatively identified as such, They
had, however, excellent material of A. transfasciatus Amaral, a
species they considered sufficiently similar to A. punctatus to
suggest that the former might well be placed eventually as a
subspecies or synonym of the latter.
A single specimen of A. boulengeri O'Shaughnessy, collected
in Santiago-Zamora Province, Ecuador, in the region between
the Rio Pastaza and the Rio Santiago, is catalogued as No.
45776, in the Museum of Comparative Zoology. It provides an
~]
1956 LEAF-NOSED ANOLE FROM SOUTH AMERICA
opportunity to compare data for this species with the notes made
on other species by Myers and Carvalho. The specimen has, on
its snout, a substantial protuberance which does not extend to
any distance beyond the end of the jaw. This swelling is without
doubt the primitive condition that led eventually to the elonga-
tion observed in phyllorhinus and proboscis. It is shghtly more
prominent than that of A. transfasciatus and presumably also
of punctatus. In addition to the presence of keels on the ventrals,
A. boulengeri differs from both leaf-nosed species and trans-
fasciatus in the presence of keels on the enlarged supraocular
scales. The suboculars and the lowest row of loreals form a
straight series in both boulengert and phyllorhinus, while in
transfasciatus and proboscis the subocular series turns upward
around the eye, not forming a linear series with the loreal row.
There is no middorsal scale row in boulengeri. Two rows of
scales, shehtly larger than the other dorsals, occupy the vertebral
line. There is no midventral row of scales. All dorsal, lateral,
and ventral scales have keels which can be seen if examined
under sufficiently high magnification. This is not true of pro-
boscis or phyllorhinus, and apparently not true of transfasciatus.
It appears, then, that A. boulengeri is quite distinet from all
these species, and presumably, from A. punctatus as well. Verifi-
cation of the latter assumption must await direct comparison of
the two species.
Cope (1876, p. 165) deseribed Scytomycterus laevis, from the
Huallaga River, in eastern Peru, which had the rostral plate
produced into a flexible appendage. Both Anolis phyllorhinus
and Anolis proboscis are quite distinct from this species, for
they have the snout projection composed of several rows of small
scales, while A. laevis has an appendage composed of a single
scale, the rostral. Boulenger (1885, p. 11) synonymized Scyto-
mycterus Cope with the genus Anolis, where it has remained
since.
LITERATURE CITED
BOULENGER, G. A.
1885. Catalogue of the Lizards in the British Museum. Vol. IT :i-xiii,
1-497, 24 pls.
8 BREVIORA NO. 62
Coreg, E. D.
1876. Report on the reptiles brought by Professor James Orton from
the Middle and Upper Amazon, and Western Peru. Journ.
Acad. Nat. Sci. Philadelphia, (2) 8:159-188.
Myers, G. S. and A. L. pk CARVALHO
1945. A strange new leaf-nose lizard of the genus Anolis from Ama-
zonia. Bol. Mus. Nae. Brasil, Zool., No. 43:1-14, 9 figs.
Anolis proboscis Holotype ¢@ (M. C. Z. 54300).
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. OcroBER 11, 1956 NUMBER 63
NEW BATHYAL ISOPODA FROM THE CARIBBEAN
WITH OBSERVATIONS ON THEIR NUTRITION
By Rosert J. MENZIES
Marine Zoologist
Lamont Geological Observatory 1
(Columbia University)
Palisades, New York
INTRODUCTION AND ACKNOWLEDGMENTS
In this paper 5 new species including 1 new genus of marine
Isopoda are described. All were collected by the R/V VEMA,
Cruise 8, 1955-56 with a single trawl using the closing-opening
Epibenthie Traw] (Menzies, in press) from bathyal depths (680
fms.) in the Caribbean Sea at 16° 59.1’ N. Latitude and 79°
07.9’ W. Longitude (South of Jamaica). The substratum con-
sisted of pteropod and foraminiferal ooze.
In addition to isopods, the trawl contained polychaete worms,
a few ophiuroids and several amphipods.
The specimens have been deposited in the Museum of Compara-
tive Zoology (abbreviated MCZ) whose publication of this manu-
seript is appreciated.
The research was supported by a Rockefeller Foundation
Grant RF 54087 for research in deep sea biology.
ECOLOGY AND NUTRITION
The ‘‘stomach’’ contents of Acanthocope spinosissima Menzies
(this paper, p. 2) which packed the gastric mill to swelling con-
sisted foremost of bacteria-size and shape particles and some
globular bodies, possibly protozoans. Next in abundance were
1Lamont Geological Observatory, Contribution No. 219, Biology Prograin No. 7.
iw)
BREVIORA NO. 63
the parapodial setae of some species of polychaete annelid, some
demospongiae spicules, a few empty small foraminiferal tests,
some diatom tests and a few tintinnoid loricae. These observa-
tions suggest that Acanthocope spinosissima is both carnivorous
and a detrital feeder with possibly some selectivity in its nutri-
tion.
SYSTEMATIC DESCRIPTION
ACANTHOCOPE SPINOSISSIMA Nn. sp.
Figure 1
Synonyms. None.
Diagnosis. Front coneave. Dorsum with three pairs of spines
on somites 5-7 inclusive. First antenna with 10 articles, first
swollen and spinulate at margins, second to fourth subequal in
length, fifth longest, about 3 times the length of fourth. Pleotel-
son long spinulate apical spine. Uropoda uniramous, peduncle
and ramus subequal in length, maxilliped with 7 coupling hooks.
Coxal plates and lateral margins of body spinulate. Mandibular
palp with one or two articles, last article much elongated.
Measurements. Holotype, length 8 mm., width 2 mm.
Type. Deposited in the MCZ, Cat. No. 12665.
Distribution. Known only from type locality.
Composition and affinities. Of the known species, A. spinosis-
sima resembles A. spinicauda Beddard (1886) most. It differs
from that species most obviously in having its pleotelsonal spines
curved cephalad rather than caudad.
The type of this genus is A. spinicauda Beddard and although
Beddard shows the uropoda as having five articles this is surely
an error as is the apparent lack of fusion of the hinder peraeonal
somites. The genus is close to Hurycope Sars (1864) and
Storthyngura Vanhoffen (1914). It differs from both in having
uni- and not biramous uropoda and in having a markedly pointed
(spinelike apex) pleotelson and not a rounded or bifureating one.
The following species appear to belong to the genus:
Acanthocope spinicauda Beddard
Acanthocope acutispina Beddard
Acanthocope atlantica (Beddard )
Acanthocope intermedia (Beddard )
All are from bathyal or abyssal depths of the sea.
1956 NEW ISOPODA FROM THE CARIBBEAN 3
Sy
7
—— SI
ae —_
Figure 1
Figure 1. Acanthocope spinosissima N. SP
holotype female length 8 mm.,
A, toto; B, maxilliped; C, first peraeopod; D, first antenna; E, left mandi-
ble; F, seventh peraeopod.
4 BREVIORA NO. 63
Figure 2
9
Figure 2. Dendrotion hanseni, n. sp., holotype male, length 3.5 mm. A,
toto; B, first pleopod; C, maxilliped; D, pleotelson; HE, gnathopod; F, left
mandible.
1956 NEW ISOPODA FROM THE CARIBBEAN 5
DENDROTION HANSENT, Nn. Sp.
Figure 2
Synonyms. None.
Diagnosis.’ Lateral processes spinulate. Third peraeonal
somite with one pair of dorsal spines; fourth somite with two
pairs of dorsal spines; seventh somite and pleotelson with one
pair of dorsal spines, pleotelson with 10-11 spinelike setae on
lateral margins followed by a fringe of smaller setae around
apex. Maxilliped with two coupling hooks. Apex of male first
pleopod with four setae.
Measurements. Holotype male, length 3.5 mm., width 0.8 mm.
Type. Deposited in the MCZ, Cat. No. 12666.
Distribution. Known only from type locality.
Composition and affinities. The genus contains only two spe-
cies, the type, D. spinosum G.O. Sars (1897), which was collected
from the Hardanger Fjord at Mosterhaven, Norway, at 150
fathoms and the new one herein described. Previously the genus
had been known only from Norway.
Genus NEOANTHURA, n. g.
Synonyms. None.
Diagnosis. Anthuridae with piercing and sucking mouth parts.
Kyes lacking. First antenna with seven articles; second antenna
with six articles comprising flagellum. Maxilliped composed of
a single lanceolate article. All six pleonal somites clearly marked
dorsally, statocyst lacking. Pleon not indurated. First two pairs
of peraeopoda subchelate, others less so but with carpo-propod
over-ridden by merus.
Type species. Neoanthwra coeca, new species.
Composition and affinities. This genus contains only the type
species. It is most closely related to Paranthura Bate and West-
wood (1866) from which it differs principally in having a uni-
and not a triarticulate maxilliped.
NEOANTHURA COECA, Nn. sp.
Figure 3
.Y F h if
Synonyms. None.
Diagnosis. Mandibular palp triarticulate, lack setae. Apex of
6 BREVIORA No. 63
Figure 3
Figure 3. Neoanthura coeca, n.g., n. sp., holotype female. A-A’, toto; B,
maxilliped; C, first antenna; D, second antenna; H£, uropod and telson.
2)
——_— ee
1956 NEW ISOPODA FROM THE CARIBBEAN i
Hieure 4
Figure 4. Antarcturus annaoides, n. sp., holotype male. 4, lateral view ;
B, penis; C, gnathopod; D, seventh peraeopod; £, inner view of apex of
uropod.
8 BREVIORA NO. 63
pleotelson rounded with only three setae. Margins of uropodal
rami smooth with few setae. Frontal margin of cephalon pointed.
Maxilliped with seven setae. Body smooth, glistening, a pair of
setae present at lateral margins of first two peraconal somites.
Measurements. Holotype female, length 3.75 mm.; width 0.25
mm,
Type. Deposited in the MCZ, Cat. No. 12667.
Distribution. Known only from type locality.
Composition and affinities. Unique.
ANTARCTURUS ANNAOIDES n.Sp.
Figure 4
Synonyms. None.
Diagnosis. An Antarcturus with dorsum of body and pleotel-
son without spines. Eyes present. Coxal plates without lateral
spines. Pleotelson with a pair of spines at: anterolateral border
and a similar pair at posterolateral angles as well as a horn-like
spine on dorsum at midline of posterior border. Dactyls present
on peraeopoda one and seven. Second antenna exceeds two times
the leneth of the body. Ventral surface of body without spines.
Measurements. Male holotype, length 13 mm., width 2.0 mm. ;
second antenna 25 mm. long.
Type. Deposited in MCZ, Cat. No. 12668.
Distribution. Known only from type locality.
Composition and affinities. The species’ closest relative appears
to be Antarcturus anna (Beddard 1886), described in the ‘‘Chal-
lenger’’ report from 600 fathoms off the Rio de Ja Plata. It differs
from that species in lacking a marginally serrated pleotelson
and in lacking armed coxal plates.
[ANIRELLA CARIBBICA, 0. Sp.
Figure 5
Synonyms. None.
Diagnosis. Front with median setiferous horn. Pleotelson with
three spines at lateral border, and four on dorsal surface. First
peraeonal somite with three dorsal spines. Second peraeonal
somite with 7 spines on dorsum, third with 9; fourth with 9;
fifth to seventh with 2. Mawyilliped with two coupling hooks. First
antenna with 13 articles.
1956 NEW ISOPODA FROM THE CARIBBEAN 9
Figure 5
Figure 5. Janirella caribbica, n. sp. A, type toto; B, dactyl first peraeo-
pod; C, maxilliped; D, dactyl seventh peraeopod; Z, first antenna.
10 BREVIORA NO. 63
Measurements. Holotype female, length 5.5 mm.; width 2.4 mm.
Type. Deposited in the MCZ, Cat. No. 12669.
Distribution. Known only from type locality.
Affinities. This species, being dorsally spined, is most closely
related to Lanirella nanseni Bonnier (1896), a species collected
by the ‘‘Caudan”’ from 950 meters depth north of the Azores
at 40°38’ KE. Longitude and 44°17’ N. Latitude. It differs mark-
edly from that species in having 3, not 4 spines at each pleotel-
sonal border and 3 rather than 1 spine on the dorsum of the
first peraeonal somite.
LITERATURE CITED
BEDDARD, F. E.
1886. Report on the Isopoda collected by H.M.S. ‘‘Challenger’’ dur-
the years 1873-76, pt. 2. ‘‘Challenger’’ Reports, Zoology, vol.
7, 175 pp: Zo! pis:
BONNIER, J.
1896. Edriophthalones, in Res. Sci. Campagne du Caudan dans le
Golfe de Gascogne. Ann. Univ. Lyon, vol. 26, pp. 527-689, pl. 33.
MENZIES, R. J.
1956. New abyssal tropical Atlantic isopods with observations on their
biology. In press. Am. Mus. Nat. History.
Sars, G. O.
1897- An account of the Crustacea of Norway. I, Isopoda. pp. 1-270.
1899. 100 pls. Bergen.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, MAss. OcToBER 11, 1956 NUMBER 64
RARE SPECIES OF COPEPODA, CALANOIDEA,
TAKEN FROM THE IZU REGION
By OvToHito TANAKA
Department of Fisheries, Kyushu University, Japan
INTRODUCTION
The present paper is part of a report on the pelagic copepods
of the Izu Region, the Pacific Coast of Middle Japan. About
three hundred and twenty species have been described from that
region and a list of the species has been published in ‘‘ Records
of Oceanographic Works in Japan,”’ vol. 1, no. 1, 1953. How-
ever, owing to various difficulties, the descriptions and figures
have not as yet been published.
The author wishes to express his thanks for help received from
Dr. A. Fleminger, U. 8S. Fish and Wildlife Service, a fellow
worker with whom he has had occasion to discuss various copepod
problems during the past year, and Dr. Elisabeth Deichmann,
Curator of Marine Invertebrates, in the M.C.Z., Cambridge,
Mass. It is due to their efforts that he has now the pleasure of
seeing this contribution in print.
RYOCALANUS gen. nov.
Diagnosis. Body elongate; head separate from first thoracic
segment; fourth and fifth segments separate. Rostrum stout,
one-pointed. Abdomen five-jointed in male. First antenna 24-
jointed ; eighth and ninth joints fused; male with right antenna
transformed into grasping-organ; joints 23 and 24 fused, hinge
between joints 22 and 23. Second antenna as in Drepanopsis
Wolfenden, with three setae on inner margin of second joint of
exopodite; first basal joint with a brush of hairs in posterior
aspect. Mandible with slender palp; endopodite two jointed,
BREVIORA NO. 64
a $
Figure 1. Ryocalanus infelix n. sp. a, Complete animal, dorsal view; b,
head, lateral view; c, last thoracic segment and abdomen, lateral view;
d, rostrum; e, grasping antenna; f, second antenna; g, first leg and proximal
outer margin of endopodite; %, second leg; i, third leg; j, third joint of
exopodite of fourth leg.
1956 RARE SPECIES OF COPEPODA, CALANOIDEA 3
shorter than exopodite. First maxilla as in Mimocalanus Farran,
but exopodite, second basal joint and lobes slender. Second
maxilla as in Pseudocalanus Boeck; fifth lobe largest. Maxilli-
ped as Drepanopsis Wolfenden except that distal two joints are
furnished with well developed setae on outer margin. Swim-
ming legs as in Spinocalanus Giesbrecht and Monacilla Sars
except that distal joint of exopodite of first leg with a spine on
outer margin at about mid-length of joint. Fifth pair of legs of
male asin Pseudocalanus. Type species, Ryocalanus infelix n. sp.
RYOCALANUS INFELIX sp. nov.
Description. Male: Length, 2.18 mm. ; cephalothorax, 1.93 mm. ;
abdomen, 0.25 mm.; head separate from first thoracic segment ;
last two thoracic segments separate; cephalothorax elongate,
ovate; head contracted anteriorly; last thoracic segment pro-
duced into an acute spine on each side, ventrally directed ; ventral
margin of segment furnished with fine spinules; rostrum one
point and acute, in lateral view with a notch on posterior surface
near distal end.
Abdomen five-jointed; segments and furca with proportional
length of 32, 18, 11, 7, 14, 18 = 100; first abdominal segment
about half as long as wide, with fine spinules on distal corner
of right side; second to fourth segments with spinules on distal
margin; furcal ramus wider than long, with five setae, outermost
seta short and slender; dorsal surface of ramus with minute
spinules; appendicular seta short.
Left first antenna damaged, with distal joints missing; joints
eight and nine incompletely fused on anterior margin; joints with
following proportional lengths :
Joint 1 2.93 4 9 6 fis 10 11 12 13 14—* 15
PAD) A Sad) ay By ayy 1! if 5 6 6 6 6 6
Right antenna a grasping organ, extending to distal margin
of second thoracic segment; joints eight and nine completely
fused; joints 18 to 22 inflated; joint 20 with a strong seta on
posterior margin; joint 21 with fine denticles and a seta on pos-
terior margin; joint 22 highly chitinized on posterior margin
and with a short distal seta; joint 23 and 24 incompletely sep-
arated, forming a comb-like projection with 19 denticles on
posterior margin; posterior margin with a minute seta at junc-
4 BREVIORA NO. 64
Figure 2. Ryocalanus infelix n. sp. k, mandible; Jl, first maxilla; m,
second maxilla; n, maxilliped; 0, fifth pair of legs.
1956 RARE SPECIES OF COPEPODA, CALANOIDEA 5
tion with joint 24; joint 25 with five setae and an aesthetask;
joints 12-19 with hairs on posterior margin.
Second antenna with exopodite about as long as endopodite;
exopodite seven-jointed, second joint with three marginal setae;
distal joint of endopodite with eight plus six setae. Mandible
with exopodite slightly longer than endopodite; exopodite five-
jointed ; endopodite two-jointed; distal joint of endopodite with
ten setae; second basal joint with three setae; cutting edge with
eight teeth. First maxilla well developed; outer lobe with four
long and two short setae ; exopodite with eleven setae; endopodite
with seven plus six plus four setae; second basal joint with five
setae; third inner lobe with four; second inner lobe with five
setae; first inner lobe with 14 spines. Second maxilla normal ;
first lobe with four setae, second to fifth lobe, each with three
setae ; sixth lobe with two setae, distal one short, endopodite with
six setae; first basal joint with an outer marginal seta. Maxilli-
ped slender ; outer marginal seta on fourth and fifth joint of endo-
podite long; second joint of endopodite the largest.
First leg with three-jointed exopodite and one-jointed endopo-
dite; outer edge spine on joints of exopodite long, third joint
with four inner marginal setae and two outer marginal spines,
distal half of outer margin hollowed ; endopodite with three mar-
ginal setae, two apical setae and a process on proximal outer
margin; second basal joint with row of spinules on inner distal
margin at base of endopodite; exopodite of right leg with an
abnormal structure, inner marginal setae on second and third
joints of exopodite converted into strong spines. Second leg with
three-jointed exopodite and two-jointed endopodite; joints of
exopodite — especially second and third — densely covered with
fine spinules on posterior surface ; third joint with five inner mar-
ginal setae ; posterior surface of joints of endopodite with groups
of spinules; terminal spine of exopodite with 23 teeth connected
with what appears to be a thin lamella; outer margin of second
joint, and proximal section of third joint of exopodite with
acicular spines and a row of slender, curved spines (as in Figure
1h); this arrangement of spinules observed also in exopodite of
third and fourth legs; basal joint with groups of minute spinules ;
third leg with three-jointed exopodite and endopodite; joints of
exopodite and endopodite with spinules on posterior surface as
6 BREVIORA NO. 64
By
Figure 3. Calocalanus gracilis n. sp. a, Complete animal, dorsal view;
b, head, lateral view; c, last thoracic segment and abdomen, lateral view;
d, first leg; e, second leg; f, third leg; g, fourth leg; h, fifth leg.
1956 RARE SPECIES OF COPEPODA, CALANOIDEA 7
in second leg; terminal spine of exopodite with 25 teeth; third
joint of endopodite with six setae in all. Fourth leg with three-
jointed exopodite and endopodite, and of similar structure as
third leg; terminal spine of exopodite finely serrate.
Fifth pair of legs with five joints on each side; left lez much
longer than right; terminal joints of each leg with two unequal
apical spines; right leg with an outer edge spine on third joint.
Material. One adult male from a vertical haul, 1410 m.-0 m.,
near Izu, Suruga Bay, Japan (November 1938).
Material deposited. Kyushu University.
Remarks. The species appears to be allied to the genus Auta-
nepsis, described by Wolfenden (1911, Report Deutsche Siidpolar
Kixpedition), in the structure of the swimming legs, but the
mouth parts are entirely different. The grasping antenna of the
present species differs from those found in the members of the
tribe Heterarthrandria which have a movable articulation be-
tween the joints 18 and 19. The structure of the first to fourth
legs resembles those of the Pseudocalanidae, yet the peculiar
structure of the grasping antennae has not—as far as I am
aware — been reported in the species hitherto described. At the
present moment I therefore place this curious species in a new
genus, under the name Ryocalanus infelix, in memory of my
only son who lost his life in his early boyhood,
CALOCALANUS GRACILIS Nn. sp.
Description. Female: Length, 0.62 mm. Body slender, cephalo-
thorax 38.5 times as long as wide; abdomen contained 4.8 times in
length of cephalothorax. Abdomen with three segments, segments
and rami with proportional lengths of 40, 8, 32, 20— 4100;
genital segment as long as wide; furcal rami as long as wide.
Terminal joints of first antennae missing; length of the re-
maining joints as follows:
Joint 1 2 3 4 5 6 7 8-9) 10) PAE 2 aS enya
eee es) yep) een eer les rmd beta AO) ays) ear BA7/
Swimming legs small; first leg with two-jointed exopodite and
one-jointed endopodite; first basal joint with small spines on
inner distal margin near the middle. Second to fourth leg each
with three-jointed exopodite and three-jointed endopodite. First
joint of exopodite of second leg with four small spines on outer
8 BREVIORA No. 64
margin at base of outer edge spme. Third and fourth legs with
joints of endopodite devoid of groups of spines on posterior sur-
face ; leaf-like spines of second and third joints of exopodite also
absent. Fifth pair of legs three-jointed; terminal joint about as
long as first joint, with an end-spine of similar length as the
three joints taken together, and with a minute spine on outer
distal corner.
Material. One adult female from a vertical haul, 50-0 m., near
Izu, Suruga Bay, Japan (August 1937).
Material deposited. Kyushu University.
Remarks. Resembles C. tenuis Farran in dorsal view, but the
small size and the structure of the swimming legs separate it from
the latter which was originally described from the Bay of
Biscay.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass.
~ NovEMBER 20, 1956 Numper 65
A NEW SPECIES OF AGRIOGNATHA (ARANEAE,
ARGIOPIDAE) FROM PANAMA
By ArtHur M. CHICKERING
Albion College, Albion. Michigan
I. P, Cambridge (1903) recognized two species of Agriognatha
from western Panama and another from Costa Rica although he
was in some doubt as to the proper position of the Panamanian
A. lepida (O. P. Cambridge). To my knowledge, no species of
Agriognatha from Panama have been recognized since that time.
No mention of the genus is made by Kraus (1955) in his recent
paper on the spiders of El Salvador.
During my recent studies on the genus Vetragnatha Latreille,
1804 in Panama numerous specimens of what I am compelled to
regard as a new species of Agriognatha O. P. Cambridge, 1896
have been sorted out of my collections. This species is described
in accord with my usual procedure, and the types will be de-
posited in the Museum of Comparative Zoology at Harvard
College. A single male of another species from El Volean, R. P.
has also appeared in my material. Unfortunately, this specimen
has lost both palpal tarsi and, hence, is regarded as unworthy
of a description at the present time. This latter species appears
to be close to A. bella O. P. Cambridge from Costa Rica.
Genus AGRIOGNATHA O. P. Cambridge, 1896
In a recent paper (1956) describing a new species of Agrio-
gnatha from Jamaica, B. W. I., I have given reasons for retaining
this generic name rather than returning to Cyrtognatha as
toewer (1942) has done. In view of the uncertainty regarding
the relationship between the genus Cyrtognatha Keyserling, 1881
2 BREVIORA NO. 65
and the genus Agriognatha O. P. Cambridge, 1896 I think it is
still desirable to retain the name given here.
External Anatomy of Agriognatha insolita sp. nov.
Fig. 1. Eye group of male from in front.
Fig. 2. Chelicerae and cheliceral teeth from in front.
Figs. 3-4. Male palpal tibia and tarsus; lateral and ventral views, re-
spectively.
Fig. 5. Female cheliceral teeth.
Fig. 6. Genital groove of female.
AGRIOGNATHA INSOLITA sp. nov.
(Figures 1-6)
Male holotype. Total length 5.59 mm. including chelicerae ;
exclusive of the chelicerae 5.2 mm. Carapace 1.95 mm. long;
1.17 mm. wide opposite interval between second and third coxae
where it is widest; considerably narrowed opposite first coxae;
median longitudinal thoracic groove well marked.
Eyes. Eight in two rows; viewed from above, both rows re-
curved with anterior row strongly so; viewed from in front, an-
1956 A NEW SPECIES OF AGRIOGNATHA FROM PANAMA 3
terior row gently recurved, posterior row gently procurved, both
measured by centers. Central ocular quadrangle slightly wider
in front than behind, slightly longer than wide in front. Ratio
of eyes: AME ALE: PME; PDB == 11: 10210259) Avie
separated from one another by about three-fourths of their
diameter, from ALE by shghtly less than 1.2 times their
diameter. PME separated from one another by about four-fifths
of their diameter, from PLE by 1.8 times their diameter. Lat-
erals separated from one another only by a broad line (Fig. 1).
Chelicerae. Very divergent; nearly horizontal, Basal segment
1.625 mm. long and, therefore, about four-fifths as long as
the carapace. The fang is moderately sinuous and without special
features. Near the base of the fang on the promargin of the fang
groove there is a robust bifurcate tooth and three other small
teeth more proximal in position; the retromargin has four some-
what larger teeth (Fig. 2).
Maxillae. Nearly parallel; somewhat broadened distally ; about
three times as long as broad in middle; moderately concave along
lateral borders; slightly more than twice as long as lip.
Lip. About as long as wide at base; sternal suture irregularly
procurved; with pronounced sternal tubercles at ends of sternal
suture.
Sternum. Essentially cordiform; as wide as long; moderately
convex; moderately scalloped opposite each coxa; moderately
extended between all coxae with second and third coxae relatively
widely separated ; continued by a narrow sclerite between fourth
coxae which are only slightly separated.
Legs. 1243. Width of first patella at ‘‘knee’’ .264 mm., tibial
index of first leg 5. Width of fourth patella at ‘‘knee’’ .214 mm.,
tibial index of fourth leg 8.
Femora Patellae Tibiae Metatarsi Tarsi Totals
il 4.420 .910 4.615 5.200 1.365 16.510
a 3.380 .750 2.762 .250 .975 iatalyy
ae 1.625 410 975 1.285 .700 4.945
4. 2.730 .020 2.080 2.470 .885 8.685
Palp 520 .220 242 —_— .770 oe
(All measurements in millimeters )
4 BREVIORA NO. 65
With many true spines on all legs. Those which seem to be
most distinctive occur as follows. First leg: near the base of
the femur on the ventral surface there are three short robust
spines in a short row; on the retrolateral surface there is a row
of nine or ten spines several of which seem to be modified for
some special use with the most distal one long and robust; the
metatarsus has a series of about twenty-five short robust ventral
spines beginning at the base of the segment and extending
through about three-fourths of its length and diminishing in size
distally. Second leg: essentially as in first with the special
metatarsal spines less marked. The third femur has a row of
five conspicuous trichobothria on its prolateral surface in the
proximal fourth; the fourth femur has a row of eight similar
trichobothria on its prolateral surface in the proximal third.
Palp. Complicated and somewhat difficult to describe because
of the presence of numerous transparent tarsal apophyses. It
appears to be rather typical of those described by Miss Bryant
from Cuba and Hispaniola (1940, 1945). The paracymbium is
long and slender with a terminal enlargement (Figs. 3-4).
Abdomen. Unnotched at base dorsally ; only slightly extended
over the carapace ; 3.51 mm. long; .91 mm. wide; nearly uniform
in width to the rounded posterior end.
Color in alcohol, Chelicerae reddish brown; elsewhere essen-
tially like that of female except that there are fewer of the silvery
spangles on the abdomen.
Female allotype. Total length including chelicerae 8.125 mm. ;
exclusive of the chelicerae 7.80 mm. Carapace 2.47 mm. long;
1.625 mm. wide opposite interval between second and third coxae
where it is widest ; with the cephalic region only slightly raised ;
the shallow thoracic median pit is shaped quite differently from
that usually present in the genus Tetragnatha and is extended
forward in a narrow point.
Eyes. Eight in two rows as usual; with laterals on a rather
prominent tubercle; viewed from above, posterior row slightly
recurved, anterior row strongly so; viewed from in front, pos-
terior row gently procurved, anterior row moderately recurved ;
both measured by centers. Central ocular -quadrangle only
1956 A NEW SPECIES OF AGRIOGNATHA FROM PANAMA 5
slightly wider behind than in front; longer than wide behind
in ratio of 11: 10. Ratio of eyes AME: ALE: PME: PLE =
10: 9: 10: 8. AME separated from one another by 1.1 of
their diameter, from ALE by 2.3 times their diameter. PME
separated from one another by 1.2 times their diameter, from
PLE by 2.2 times their diameter. ALE separated from PLE
by one-fourth of the diameter of PLE (some workers might
regard them as being contiguous because the corneas are nearly
so, but the margins of the corneal hypodermis are separated
as indicated). Height of clypeus equal to .9 of the diameter
of AME.
Chelicerae. Moderately well developed; only slightly diver-
gent; basal segment 1.865 mm. long; fang regularly curved,
without cusps; fang groove well marked; promargin of fang
groove with only three teeth of moderate size plus two minute
denticles; retromargin with four teeth also of moderate size but
slanted irregularly (Fig. 5).
Mazillae. Slightly divergent, with outer distal corners con-
siderably extended ; longer than lip in ratio of about 9: 4; longer
than wide in ratio of about 18: 7.
Iip. Only slightly broader at base than long. Sternal suture
gently procurved; with pronounced sternal tubercles at lateral
ends of suture.
Sternum. Essentially as in male.
Legs. 1243. Width of first patella at ‘‘knee’’ .330 mm., tibial
index of first leg 5. Width of fourth patella at ‘‘knee’’ .242 mm.,
tibial index of fourth leg 8.
Femora Patellae Tibiae Metatarsi Tarsi Totals
1s 4.875 1.105 5.200 5.720 1.592 18.492
2. 3.510 .910 3.250 3.880 1.175 12.725
Bi 2.000 520 975 1.560 athe 5.830
4. 3.185 85 2.405 2.730 1.105 10.010
(All measurements in millimeters )
All legs with spines and hairs. The third femur has a row of
six prominent trichobothria on its anterior surface in the proxi-
mal third of the segment; the fourth femur has a similar row of
about fifteen prominent trichobothria.
6 BREVIORA No. 65
Abdomen. Only slightly gibbous in anterior half; 5.525 mm.
long; only slightly extended posterior to spinnerets; squarely
truncate at anterior end; genital groove as shown in Figure 6.
Color in alcohol. Legs: generally yellowish with tibiae and
metatarsi darker; irregular black streaks beneath patellae and
distal ends of femora; grayish areas are frequent on several
segments. Mouth parts are generally light yellowish brown with
the fang darkest of all. Sternum light yellowish brown. Carapace
light yellowish brown with darker markings. Abdomen: cardiac
region very hight yellowish; remainder of dorsal and dorsolateral
areas light yellowish with many small silvery spangles; mid-
ventral area light yellowish with a narrow row of silvery spangles
on each side; farther laterally the venter is free of the silvery
spangles.
Type locality. Both male holotype and female allotype are
from Barro Colorado Island, C. Z., August, 1950. Male and
female paratypes are in the collection from: Barro Colorado
Island, C. Z., July, 1936; February, 1936 (Gertsch) ; August,
1939; May, 1946 (T. C. Schneirla) ; June-August, 1950; July-
August, 1954; Arraijan, R. P., August, 1936.
1956 A NEW SPECIES OF AGRIOGNATHA FROM PANAMA 7
BIBLIOGRAPHY
BRYANT, ELIZABETH B.
1940. Cuban spiders in the Museum of Comparative Zoology. Bull.
Mus. Comp. Zool., Vol. 86, pp. 249-532, pls. 1-22.
1945. The Argiopidae of Hispaniola. Jbid., Vol. 95, pp. 359-418,
pls. 1-4.
CamBripgGR, O. P. and F. P. CAMBRIDGE
1889-
1905. Arachnida-Araneida. Vols. I and II in: Biologia Centrali-
Americana. Dulau & Co., London.
CHICKERING, A. M,
1956. A new species of Agriognatha from Jamaica, B. W. I. Breviora,
Mus. Comp. Zool., No. 60, pp. 1-7, 5 figs.
KEYSERLING, GRAF EUGEN VON
1881. Neue Spinnen aus Amerika. Verh. zool. bot. Gesell. Wien, Vol.
31 p26.
Krauts, Orro
1955. Spinnen aus El Salvador (Arachnoidea, Araneae). Abh.
Senckenberg. Naturf. Gesell., No. 493, pp. 1-112.
Rorewenr, C. F.
1942. Katalog der Araneae. Vol. 1. Bremen.
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. NOVEMBER 27, 1956 NUMBER 66
THE CANINAE OF THE THOMAS FARM MIOCENE
By STANLEY J. OLSEN
Florida Geological Survey
INTRODUCTION
The generic relationships of the various members of the
Caninae from the Florida Miocene have been uncertain ever since
Dr. E. H. Sellards first described a new species of Mesocyon
from the Griscom Plantation in northwest Florida (Sellards,
1916). Our knowledge of the dogs of this epoch in Florida has
been greatly increased by the work of Dr. Theodore E. White
at the Thomas Farm between 1940 and 1946 (White, 1941, 1942,
1947) and by others who have worked at the site since then.
The Thomas Farm locality has been described in those papers
dealing with the earlier vertebrate finds (Simpson, 1932; White,
1942).
In carrying out this study all specimens studied by Simpson
and by White have been examined, together with a cast of the
type of M. iamonensis Sellards. In addition, much unstudied
Thomas Farm material in the Museum of Comparative Zoology
(M.C.Z.), University of Florida, and the Florida Geological
Survey (F.G.8.), collected subsequent to the work of Simpson
and White, has been available.
The types, or casts of the types, of the following species have
been at hand for comparison or have been examined: Mesocyon
hortulirosae Schlaikjer, M.C.Z. 2882; M. geringensts Barbour and
Schultz, Univ. Neb. 1902; M. drummondanus D-uglass, Carnegie
Museum 792; Tomarctus brevirostris Cope, Amer. Mus. Nat.
Hist. 8302, A.M.N.H. 13836 (type of T. hippophagus Matthew) ;
T. optatus Matthew, A.M.N.H. 18916; Cynodesmus thomsoni}
1 White (1941, p. 95) refers this species to Tomarctus.
9 BREVIORA No. 66
Matthew, A.M.N.H. 12874, and C. thodides Princeton Univ.
10412.
As a result of the study presented here, it is evident that the
Thomas Farm Caninae are referable to two species representing
two genera. These are Cynodesmus iamonensis (Sellards) and
Tomarctus canavus (Simpson). Tomarctus canavus, the smaller
of the two, and its synonyms, will be discussed first. Complete
redescriptions are unnecessary, as they have been given in the
original discussions.
TOMARCTUS CANAVUS (Simpson)
Figures 1, 4
Cynodesmus canavus Simpson, 1932, p. 19, fig. 4; White, 1941, p. 91.
Tomarctus canavus (Simpson), White, 1942, p. 8, pls. 2, 6.
Tomarctus thomasi White, 1941, p. 94, pl. 14, fig. 3; 1942, p. 8.
Nothocyon insularis White, 1942, p. 7, pl. 1, fig. 3; 1947, p. 502, fig. 2,
D, E.
In Simpson’s type description of Cynodesmus canavus (F.G.S.
V-5260) no mention is made of a comparison between this speci-
men and Tomarctus, although the type of Cynodesmus nobilis
(F.G.S., V-5255) was compared with this related genus. The
talonid of M, of the type of canavus, though considerably worn,
retains enough of the characteristics to definitely identify it as
Tomarctus (ef. Figure 4). In canavus the talonid of M, ex-
hibits a ridge between the hypoconid and the entoconid that is
characteristic of Tomarctus. This species also has two subsidiary
cusps anterior to the hypoconid and entoconid respectively, the
latter being the more pronounced. These cusps and the cross
ridge are not present in Cynodesmus or Mesocyon. P* in the
series exhibits a parastyle varying from incipient to small but
definite. The lower premolars are less crowded than those of
T. thomsoni. On comparing Tomarctus canavus with the speci-
mens of Tomarctus brevirostris Matthew (1924) from the Snake
Creek, it was found that the proportions and cusp arrangement
were nearly the same, but the specimens of 7. brevirostris were
somewhat larger. White (1942) correctly interpreted canavus as
having the generic characters of Tomarctus rather than of Cyno-
desmus, and referred it to that genus.
1956 CANINAE OF THOMAS FARM
MCZ
Figure 1. Tomarctus canavus, labial, occlusal and lingual views.
3628. X1.
4 BREVIORA No. 66
Tomarctus thomast White was based on a partial left maxilla
bearing P4-M?. White originally compared this species only with
T. optatus and T. brevirostris. Later (1942; p. 8), he stated that
thomasi was slightly larger than canavus with relatively larger,
more crowded teeth. The differences between M.C.Z. 3628 and
3712 (the specimens that White evidently compared), slight in
any case, disappear when the whole series is examined. The
species 7. thomasi cannot be maintained.
Nothocyon insularis was also based on a fragmentary maxilla.
White placed the species in Nothocyon because of the spacing of
the first and second molars, the quadrangular M2, and the coni-
cal hypocone of M!, relying largely upon the obtuse posterolat-
eral angle of M? to exclude the species from Tomarctus. The
spacing of M1-2 does not appear to differ markedly from that
observed in other members of the series. The hypocone of M}
| MCZ | MCZ ¢ MCZ FRI MCZ 3 MCZ * | MGZ +1 Moz +|Mcz %*|MCZ # == MCZ #
3629 | 3712 3924 4242 ©|4333 © |4334 Glaser ao eee
| 204 | 19.7 | 219 | 22.1 | 10 | 200 | 220 | 19.6 |
3% ESSENTIALLY UNWORN C- CRUSHED, oe
+ SLIGHTLY WORN NOT ENTIRELY RELIABLE
++ MODERATELY WORN
FP WELL WORN
is slightly more conical but this is not a profound difference ;
M.C.Z. 3641, when unworn, may have been entirely comparable
in this respect. M? is more quadrangular than in the other two
available examples of the tooth. The difference is due to the
structure of the external cingulum. This may be regarded as an
individual peculiarity. In the description of a referred lower
jaw (White 1947, p. 502), he observed that there was little
difference between the alveolar length of this specimen and of
those referred to 7. canavus, but that it was more slender
throughout, the teeth smaller and shorter crowned, the condyle
shorter and smaller, and set at an oblique angle to the horizontal
ramus, indicating a broad skull for this form. Experience with
Thomas Farm material has shown that exact jaw angles, lengths,
depth of rami, ete., are not too reliable, due to the distortion and
1956 CANINAE OF THOMAS FARM 5
cracking that this material has undergone during the process of
fossilization. The measurements as given on the opposite page
demonstrate the variability in depth of ramus.
As may be seen from the analysis of the dental measurements
(Table 1), the teeth referred to insularis by White are not
significantly smaller than those referred by him to canavus and
thomast. As he noted, ‘‘the heel of the carnassial shows the ridge
between the hypoconid and entoconid that is characteristic of
Tomarctus.’? There is, in fact, no reason for separating N.
insularis from T. canavus.
CYNODESMUS IAMONENSIS (Sellards)
Figures 2, 3
Mesocyon? tamonensis Sellards, 1916, p. 88, pl. 11, fig. 11; pl. 18, fig. 1;
Simpson, 1930, p. 160.
Cynodesmus ? iamonensis (Sellards) Simpson, 1932, pp. 14-19.
Cynodesmus nobilis Simpson, 1932, p. 17, figs. 1-3; White, 1941, p. 91.
Paradaphoenus nobilis (Simpson) White, 1942, p. 5, pl. 2, fig. 1; pl. 3.
Paradaploenus tropicalis White, 1942, p. 5, pl. 1, fig. 2; pl. 4.
Parictis bathygenus White, 1947, p. 500, fig. 2A (in part, not including
type).
This species has been buffeted about somewhat as regards its
generic assignment. Sellards, who knew it only from the type, a
maxillary fragment with P4-M*,! was quite justified, on that
basis, in his tentative assignment to Mesocyon, this genus being
similar to Cynodesmus in the structure of these teeth. Simpson,
on the basis of more complete material, including the critical
M,, referred it to Cynodesmus under the name of C. nobilis.
White placed nobilis in Paradaphoenus on the grounds that the
entoconid and hypoconid of M, are of nearly equal size, whereas
in Cynodesmus thooides, the type species of the genus, the ento-
conid is smaller than the hypoconid. However, Paradaphoenus
possesses M*® (Wortman and Matthew 1899, p. 129), which
zamonensis does not, and for this and other reasons such an
assignment is not possible. In the present state of our knowledge
of the earlier Caninae, Simpson’s reference to Cynodesmus is
the only practicable one. Comparison of Thomas Farm material
1The type of MW. iamonensis was first listed by Sellards as Florida Geological
Survey no. 5082, later changed to V-319, and was then sent to the U. S. National
Museum and given the permanent number, U.S.N.M. 88386.
6 BREVIORA No. 66
TABLE |
STATISTICAL DATA ON LINEAR DIMENSIONS OF PERMANENT TEETH OF
TOMARCTUS CANAVUS
14.0—16.2 15.0% .34 834 .23 5.5+1.6
7.0— 9.0 79+ 32 i (- aed 9.8% 28
114-128 (2m 47+ 09 39+ 7
7.0 - Tie .20 et Per: 18
Hines [233 lS 33t oat
6.9 - gt: Bot 23 10.6 + 3.1
=
9.3- 11.8 10.5+ .29 aes .20 ae 1.9
4.7- 5.6 5.1t 08 .26+ 06 Cuilas fie
16.3 — 18.5 oy ar Bit
6.7 - 8.3 lgetiee: 47+ .09
9.1 —10.2 Gee 2 50t
5.7 - 6.9 eee 53+
L
L
WwW
L
Ww
TABLE 2
STATISTIGAL DATA ON LINEAR DIMENSIONS OF PERMANENT TEETH OF
CYNODESMUS IAMONENSIS
14.5 - 20.0 ‘ia.0t 04 1st 03
7.8 -10.0 9.04 08 30+ 06
9.9-138 IZoO=.. ot 7 cote
12.2 —15.5 14.5+ 02 1.00+ 02 6.88 t1.15
6.8t 13 set g.ist 140
: L KOx7a==eO2 103+ .02 9.60% 1.65
10.9 — 12.8 120 sain 55+ . 4.56% 1.02
5.8- 6.8 SSO Lia. 465+ 1.03
18.3 — 21.2 20.0+ 27 Ay daa 8.39 + 1.65
s4t 1 4et. 498+ 98
agt sot 8.05+ 201
7.55 + 1.89
= z 7
| | i
=o5 p20
v
>|
= £205
= Tr
1956 CANINAE OF THOMAS FARM 7
~-
SS
> =
Figure 2. Cynodesmus iamonensis, labial, occlusal and lingual views.
MCZ 3714. X 1.
8 BREVIORA No. 66
with C. thodides shows a close agreement in structure and pro-
portions, I* is rather large relative to I1-2, P? is rather large
relative to P*, and the heel of M, is basined. This last feature
definitely rules out all possibility of reference to Mesocyon, in
which M, has an unbasined talonid consisting of a large, bluntly
pointed, anteroposteriorly elongate hypoconid and a low internal
cingulum. With wear, the hypoconid in Mesocyon grinds out a
circular area in the center of M!, a feature not encountered in
the zamonensis material.
C. iamonensis is readily distinguished from C. thodides; PS
have lower, less pointed paracones, the protocone of P# is rela-
tively smaller, the parastyle of M1 smaller and less projecting,
the hypocone of M1! more expanded posteriorly and the anterior
border of the posterior narial opening opposite M?, rather than
immediately posterior to M2, as in thodides. As White noted
(1942, p. 6), the entoconid of M, is small in the type of thodides;
it is, in fact, subordinated in the internal cingulum, although
this is raised to rim a definite basin. In C. iamonensis the ento-
conid of M, is usually distinct, but in M.C.Z. 3965 it is com-
pletely submerged in the cingulum, which is somewhat higher
than, but otherwise identical to, that of the type of thodides.
Nothing need be added to Simpson’s comparisons (1932, p. 18)
with other species referred to Cynodesmus.
Simpson (1932, p. 19) regarded the Thomas Farm specimens
as specifically distinct from tamonensis — the type of which is
from the Griscom Plantation — on the basis of larger size, hypo-
cone of P* more projecting internally, M? ‘‘relatively larger,
more oblique, hypocone basin stronger and projecting more
postero-internally.’’ The present series shows that the difference
in size is not significant and that M?, in particular, is a highly
variable tooth; specimens at hand connect Simpson’s paratype
of nobilis (F.G.S. V-5259) with the type of camonensis. The
geological evidence now available indicates that Griscom Planta-
tion and Thomas Farm are of the same age. In 1930 (pp. 160-161,
fig. 5) Simpson described some fragmentary canid material from
the Franklin Phosphate Company mine as Mesocyon iamonensis.
This may be referable to Mesocyon but certainly not to iamonen-
sis. Simpson did not mention these fragments explicitly in his
later paper, but they are obviously the basis for his record of
1956 CANINAE OF THOMAS FARM 9
-METAGONID PROTOCONID
yy
ne
PROTOCONID “ LENTOCONIDY.
PARAGONID | woe ~~
PARACONID
PARACONE
Bs 851/11 HI
LEFT My
OSCLUSAL VIEW METACONE
LEFT My
LINGUAL VIEW
Lert mM!
OCCLUSAL VIEW
CYNODESMUS IAMONENSIS
Figure 3. Tooth characters of Cynodesmus iamonensis. X 1%,
Lingual
cSubsidiar
aa Cusp J
OCCLUSAL VIEW
LEFT My
LINGUAL VIEW
~Metaconule
LEFT Mm}
OCCLUSAL VIEW
TOMARCTUS CANAVUS
Figure 4. ‘Tooth characters of Tomarctus canavus. Key characters indi-
cated by arrows. X 1%.
10 BREVIORA No. 66
? Temnocyon sp. from the Franklin Phosphate Company (1932,
p. 14). No additional material representing this species has yet
come to light.
Paradaphoenus tropicalis White was stated to be one-seventh
smaller than nobilis, with the protocone and metacone of M2
larger and the hypocone reduced. As regards size, analysis of
the measurements shows no significant difference. M? is de-
cidedly variable as to total size and also as to details of cusp
proportion and structure; the differences mentioned by White
disappear in the larger series.
The paratype of Parictis bathygenus White, a maxillary, is not
morphologically distinguishable from iamonensis, and it is of in-
terest in this connection that White noted a very close resem-
blance to Mesocyon, which as already remarked, is not unlike
Cynodesmus in the structure of the posterior upper cheek teeth.
A specimen referred to Parictis in the Chicago Natural History
Museum, no. P-27157, shows conclusively that the Thomas Farm
specimen has nothing to do with this group of canids.!
Cynodesmus tamonensis is a variable species, both as to size
and as to details of dental structure, more so than Tomarctus
canavus.
CONCLUDING REMARKS
It must be emphasized that this is a revision of the species
present in the Thomas Farm deposit only. Whether or not these
species are valid can be determined with certainty only by study
of all material referable to the genera in question, a task beyond
present opportunities. Nomenclature aside, it is possible to state
with confidence that only two species of the Caninae are repre-
sented in the Thomas Farm deposit. It is curious that two true
dogs very similar in size should have coexisted in nearly equal
abundance. The differences in the structure of the posterior
cheek teeth (cf. Figs. 3, 4) may have reflected some difference
in habit, but this would at present be an uncertain inference.
The available skulls are so crushed and poorly preserved that
very little can be gained from them; the only impression re-
ceived is that the face anterior to P+ may have been a little longer
in C. 1amonensis.
1 The type mandible is not referable to Parictis either, a point that will be
taken up in a subsequent contribution.
1956 CANINAE OF THOMAS FARM all
Knowledge of the postcranial skeletons, however, may in the
future reveal differences that cannot now be appreciated.
ACKNOWLEDGMENTS
My foremost acknowledgment and very grateful appreciation
is to Professor Bryan Patterson for his counsel and aid in pre-
paring this paper. I wish to thank also Dr. Herman Gunter,
Dr. A. S. Romer, Dr. G. G. Simpson and Mrs. R. Nichols for
making available material in their respective institutions. Drs.
C. L. Gazin, C. B. Schultz, and J. LeRoy Kay have aided me
greatly by sending casts of type specimens for comparison.
The assistance given by Dr. R. Bader in presenting the statistical
data has been especially valuable. The figures and charts were
drawn by Miss Patricia Washer and Mr. Kenneth Highsmith.
REFERENCES
Barpour, E. H. anp C. B. SCHULTZ
1935. A new Miocene dog Mesocyon geringensis. Bull. Nebraska State
Museum, 1: 407-418.
Corr, E. D.
1884. The Vertebrata of the Tertiary formations of the West. Book
I, Report United States Geological Survey of the Territories,
F. V. Hayden, in charge, 3: i-xxxv 11-1009.
EYERMAN, J.
1894. Preliminary notice of a new species of Temnocyon and a new
genus from the John Day Miocene of Oregon. Amer. Geol., 14:
320-321.
MatrHew, W. D.
1924. Third contribution to the Snake Creek fauna. Bull. Amer. Mus.
Nat. Hist., 50: 59-210.
SCHLAIKJER, E. M.
1935. Contributions to the stratigraphy and paleontology of the
Goshen Hole area in Wyoming. IV. New vertebrates and the
stratigraphy of the Oligocene and early Miocene. Bull. Mus.
Comp. Zool., 76: 97-189.
Scort, W. B.
1895. The Mammalia of the Deep River beds. Trans. Amer. Phil. Soe.,
18: 55-185.
SELLARDS, E. H.
1916. Fossil vertebrates from Florida: A new Miocene fauna: New
Pliocene species; the Pleistocene fauna. Florida Geol. Survey,
8th annual report: 77-160.
12 BREVIORA NO. 66
SIMpson, G. G.
1930. Tertiary land mammals of Florida. Bull. Amer. Mus. Nat.
Hist., 59 art. 3: 149-211,
1932. Miocene vertebrates from Florida. Bull. Florida Geol. Survey,
10: 3-41.
Wuits, T. E.
1941. Additions to the Miocene fauna of Florida. Proc. New England
Zool. Club, 18: 91-98.
1942. The lower Miocene mammal fauna of Florida. Bull. Mus. Comp.
Zool., 92: 1-49.
1947. Additions to the Miocene fauna of North Florida. Bull. Mus.
Comp. Zool., 99: 497-515.
WorTMAN, J. L. AND W. D. MarrHew
1899. The ancestry of certain members of the Canidae, the Viverridae
and Procyonidae. Bull. Amer. Mus. Nat. Hist., 12: 109-139, figs.
1-10, pl. 6.
TOMARCTUS CANAVUS
My My My My
FIGURED SPECIMENS | an TEROPOSTERIOR GREATEST ANTEROPOSTERIOR GREATEST
Ly DIAMETER TRANSVERSE DIAMETER TRANSVERSE
SYNONYMS
DIAMETER DIAMETER
canaovus FG
olz
a
<
a
nm
a
°o
a
°
.canavus MGZ 3629
conavus MCZ 4334
=
a
=
conavus MGZ 3924
a
conavus MGZ 3628
a
thomasi MGZ 3712
“N
oa
CYNODESMUS IAMONENSIS
°
. nobilis MCZ 3633
9
nobilis MCZ 4521
~
tropicalis MCZ 3966
~
tropicalis MCZ 3965
x
tropicalis MCZ 3714
C.nobilis FGS V5255
P. nobilis MCZ 4330
P. nobilis MGZ 3724
Harvard MCZ Librar:
DOES NOT CIRCULATE
eit ar ae oe -
rs:
Eeaeod
Sao
ald? Wie Dre Pata Se ne