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Full text of "Breviora"

HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



dloes not circulate 



BREVIORA 



MUSEUM OF COMPARATIVE ZOOLOGY 
AT 

HARVARD COLLEGE, IN CAMBRIDGE 



Numbers 1-66 
1952-1956 



CAMBRIDGE, MASS., U.S.A. 

1957 



<^ 



5° 

r 



CONTENTS 

BREVIORA 

Museum of Comparative Zoologv 

Numbers l-6fi 

1952 

No. 1. A new Panamanian tree frog. By Edward H. Tayi.or. 
3 pp., 1 pi. February 8. 

No. 2. A staurotypine skull from the Oligocene of South Da- 
kota. (Testudinata, Chelydridae). By Ernest Wil- 
liams. 14 pp., 2 pis. February 8. 

Xo. :>. Notes on siphonophores. 3. N ectopyramis spinosa n. sp. 
By Mary Sears. 4 pp. May 23. 

No. 4. A unique remopleuridid trilobite. By H. B. Whitting- 
TON. 10 pp., 1 pi. June 9. 

No. f), Tricholimnas condificius is probably a synonym of Tri- 
cholhnnas sylvcstris. (Aves, Rallidae). By James C. 
Greenway, Jr. 4 pp. August 29. 

No. (). ( Contributions toward a reclassification of the Formicidae. 
I. Tribe Platythyreini (Hymenoptera). By William 
L. Brown, Jr. 6 pp. August 29. 

No. 7. On the nomenclature of the Pacific gray whale. By Wil- 
liam E. Schevill. 3 pp. September 29. 

No. 8. A new species of the cyclostome genus Paramyxine from 
the Gulf of Mexico. By Henry B. Bigelow and Wil- 
liam C. ScHROEDER. 10 pp. October 21. 

No. 9. New species of earthworms from the Arnold Arboretum. 
Boston. By G. E. Gates. 3 pp. October 21. 

No. 10. On the earthworms of New Hampshire. By G. E. Gates. 
3 pp. December 29. 



1953 

X.). 11 Cliaracters and synonymies among the genera of ants. 
Part I. By William L. Brown, Jr. 13 pp. March 20. 

.\o. 12. A cave faunule from western Puerto Rico with a discus- 
sion of the genus Isolobodon. By Thomas E. Reyn- 
olds, Karl F. Koopman and Ernest E. Williams. 
7 pp., 1 pi. April 23. 

No. 13. Fossils and the distribution of chelyid turtles. 1. 
"Hydraspis" leifhii (Carter) in the Eocene of India 
is a pelomedusid. By Ernest Williams. 8 pp., 3 pis. 
April 23. 

No. 14. fieeord of a hermaphroditic horseshoe crab, Limulus 
polyphemus L. By John P. Baptist. 2 pp., 2 pis. 
May 28. 

Further notes on the earthworms of the Arnold Arbore- 
lum, Boston. By G. E. Gates. 9 pp. May 15. 

Notes on the races of Micrurus fronialis (Dumeril, 
Dumeril and Bibron). By Benjamin Shreve. 6 pp. 
May 28. 

A new Ordovician trilobite from Florida. By H. B. 
Whittington. 6 pp., 1 pi. May 28. 

Characters and synonymies among the genera of ants. 
Part IT. By William L. Brown, Jr. 8 pp. Septem- 
ber 23. 

Lower Cretaceous nautiloids from Texas. By Bernharp 
Kummel. 11 pp., 2 pis. September 23. 

Lower Triassic Salt Range nautiloids. By Bernhard 
Kummel. 8 pp., 2 pis. September 23. 

The ancestry of the family Nautilidae. By Bernhard 
Kummel. 7 pp., 1 pi. September 23. 

Notes on the ant, Leptothorax ohliquicanthus Cole (Hy- 
menoptera : Formicidael. By Robert E. Gregg. 3 pp. 
October 13. 

No. 23. A new species of Hypognatha from Panama. By Arthur 
M. Chickering. 8 pp. October 23. 



No. 


^:>. 


No. 


l(r 


No. 


17. 


No. 


18. 


No. 


19, 


No. 


20. 


No. 


21. 


No. 


22 



1954 

No. 24. A new family, a new genus, and tAvo new species of 
batoid fishes from the Gidf of Mexico. By Henry B. 
BiGELOw- and William C. Schroeder. 16 pp. Janu- 
ary 27. 

No. 25. A new Miocene species of I'clusios and the evolution ol' 
that genus. Bj'- Ernest Williams. 7 pp., 4 pis. Janu- 
ary 28. 

No. 26. A preliminary list of the earthworms of northern New 
Jersey with notes. By H. Da vies. 13 pp. February 3. 

No. 27 Anterior regeneration in a sexthecal species of lumbri- 
cid earthworm. By G. E. Gates. 5 pp. February 5. 

.\o. 28. Clemmydopsis Boda a valid lineage of emj^dine turtles 
from the European Tertiary. By Ernest AYilliams. 
9 pp. February 8. 

No. 29. Absence of mesoplastva in a Pelomednsa (Testudines. 
Pelomedusidae). By Ernest Williams. 4 pp.. 2 pis. 
February 8. 

No. 30. Aestivation in a Permian lungfish. By Alfred S. Romer 
and Everett C. Olson. 8 pp., 1 pi. February 8. 

No. 31. New freshwater gastropod mollusks of the African 
genus Lanistes. By T. Pain. 4 pp. March 3. 

No. 32. Fossils and the distribution of chelyid turtles. 2. Addi- 
tional reputed chelyid turtles on northern continents : 
Palaeaspis conyhearii (Owen) — a pelomedusid. By 
Ernest Williams. 6 pp. March 12. 

No. 33. Systematic and other notes on some of the smaller 
species of the ant genus Rhytidoponera Mayr. By 
William L. Brown, Jr. 10 pp. May 14. 

No. 34. A review of the coxalis group of the ant genus Sticto- 
ponera Mayr. B}^ William L. Brow^n, Jr. 10 pp. 
July 20. 

No. 35. 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 Ernest 
Williams. 8 pp., 1 pi. July 28. 



No. 36. Present knowledge of the snake EJachistodon wester- 
manni Reinhardt. By Carl Gans and Ernest E. 
Williams. 17 pp. August 6. 

No. 37. On the evolution of an oriental earthworm species, 
Pheretima anomala Michaelsen 1907. By G. E. Gates. 

8 pp. August 18. 

No. 38. Onnia (Trilobita) from Venezuela. By H. B. Wnrr- 
TixGTON. 5 pp., 1 pi. November 28. 

No. 39. New or redescribed pelomedusid skulls from the Terti- 
ary of Africa and Asia (Testudines, Pelomedusidae). 
2. A podocnemide skull from the Miocene of Moghara. 
Egypt. By Ernest Williams. S pp., 2 pis. Novem- 
ber 24. 

No. 40. Some moUusks from the continental slope of northeast- 
ern North America. By Arthur H. Glarke, Jr. 11 
pp. November 29. 

No. 41. Gomments on the classilication of rodents. By Albert 

E. Wood. 9 pp. December 17. 

1955 

No. 42. A new salamander of the genus Parvimolge from Mexico. 
By George B. Rabb. 9 pp. February 28. 

No. 43. Speed-induced skin folds in the bottle-nosed porpoise. 
Tursiops truncatus. By Frank S. Essapian. 4 pp.. 

9 pis. April 17. 

No. 44. A new Murex from Matanzas, Cuba. By William J. 
Clench. 2 pp., 1 pi. April 8. 

No. 45. Palaeotaricha oligocenica, new genus and species, an 
Oligocene salamander from Oregon. By Richard 
VAN Frank. 12 pp., 3 pis. June 13. 

No. 46. Cave-fossil vertebrates from Camaguey, Cuba. By Karl 

F. KooPMAN and Rodolfo Ruibal. 8 pp. June 24. 

No. 47. A new species of whiptailed lizard (genus Cnemido- 
phorus) from the Colorado plateau of Arizona, New 
Mexico, Colorado, and Utah. By Charles H. Lowe. 
Jn. 9 pp. July 12. 



No. 48. Notes on Aineric-an earthworms of the family- Luni- 
bricidae. I-II. By G. E. Gates. 12 pp. October 10. 

No. 49. Three new shark records from the Gulf of Mexico. By 
Heney B. Bigelow, "W. C. Schroeder, and Stewart 
Springer. 12 pp. November 18. 

No. 50. New fro^s of the genera Asteruphrys and Oreophryne 
from New Guinea. By Arthi^r Loveridge. 5 pp. 
November 30. 



1956 

No. 51. A small mustelid from the Thomas Farm Miocene. By 
Stanley J. Olsex. 5 pp. January 27. 

No. 52. Remarks on some Miocene anurans from F^lorida, with 
a description of a new species of Hyla. By Walter 
Auffenberg. 11 pp. April 6. 

No. 5;l Food-finding by a captive porpoise [Tursiops trun- 
catus). By William E. Schevill and Barbara Law- 
rence. 15 pp. April 6. 

No. 54. A revision of the genus Braclupneles (Scincidae), with 
descriptions of new species and subspecies. By AV alter 
C. Brown. 19 pp. June 6. 

Rediscovery otilyla dorsalis and Lechriodus papuanu^ 
in New Guinea. By Arthur Loveridge. 4 pp. June 6. 

Notes on the Jamaican and Cajonan Island lizards of the 
genus Celesius. By Penny Norseen Cousens. 6 pp. 
June 15. 

Three new species of ^limetidae lAraneae) from Pan- 
ama. By Arthur M. Guickering. 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 Chamacleo fi-om Rhodesia and new 
race of Mahuya from Kenya Colony. By Arthur 
Loveridge. 4 pp. September 12. 

No. 60. A new species of Agriognatha from Jamaica, B.W.I. 
By Arthur M. Chickering. 7 pp. September 12. 



No. 


55. 


No. 


56. 


No. 


57. 


No. 


58. 


No. 


59. 



No. 61. On regeneration by earthworms of a species of the lum- 
bricid genus Dendrohaena Eisen 1874. By G. E. 
Gates. 6 pp. September 14. 

Xo. 62. A third leaf -nosed species of the lizard genus Anolis 
from South America. By James A. Peters and Gus- 
tavo Orces-V. 8 pp. October 3. 

No. 63. New bathyal Isopoda from the Caribbean with observa- 
tions on their nutrition. By Robert J. Menzies. 10 
pp. October 11. 

No. 64. Rare species of Copepoda, Calanoidea, taken from the 
Tzu Region. By Otoiiito Tanaka. 8 pp. October 11. 

No. 6d. a new species of Agriognatha (Araneae, Argiopidae 
from Panama. By Artiiitr M. Chickering. 7 pp. 
November 20. 

No. 66 The Caninae of the Thomas Farm Miocene. By Stan- 
ley J. Olsex. 12 pp. November 27. 



INDEX OF AUTHORS 

BREVIORA 

MUSEUM OF COMPARATIVE ZOOLOOY 
Numbers 1-66 
1952-56 

No. 

AUFFENBERG, WALTER 52 

Baptist, John P. 14 

BiGELOW, Henry B. and William C. Schroedeb 8, 24 

BiGELOw, Henry B., W. C. Schroeder and Stewart 

Springer 4!> 

Brown, Walter C. 54 

Brown, William L., Jr 6, 11, 18, 33, 34 

Chickering, Arthur M 23, 57, 60, 65 

Clarke, Arthur H., Jr 40 

Clench, William J 44 

Cousens, Penny Norseen 56 

Davies, H 26 

EssAPiAN, Frank S 43 

Cans, Carl and Ernest B. Williams 36 

Gates, G. E. y, 10, 15, 27, 37, 48, 61 

Greenway, James C, Jr 5 

Greenway, J. C, Jr. and E. Mayr 58 

Gregg, Robert E. 22 

KoopMAN, Karl F., and Rodolfo Ruibal 46 

KooPMAN, Karl F., Thomas E. Reynolds and Ernest E. 

Williams 12 

Kummel, Bernhabd 19, 20, 21 

Lawrence, Barbara and Willla.m E. Schevill 53 



LovERiDGE, Arthur 50, 55, 59 

Lowe, Charles H., Jr 47 

Mayr, E. and J. C. Greenway, Jr 58 

Menzies, Robert J . 63 

Olsen, Stanley J. 51, 66 

Olson, Everett C. and Alfred S. Romer 30 

Orces-V., Gustavo and James A. Peters 62 

Pain, T 31 

Peters, James A. and Gustavo Orces-V 62 

Rabb, George B 42 

Reynolds, Thomas E., Karl P. Koopman and Ernest E. 

Williams 12 

Romer, Alfred S. and Everett C. Olson 30 

RuiBAL, RoDOLFO and Karl F. Koopman 46 

Schevill, Willlv-m E 7 

ScHEViLL, William E. and Barbara Lawrence 53 

ScHROEDER, WiLLiAM C. and Henry B. Bigelow 8, 24 

Schroeder, W. C, Henry B. Bigelow and Stewart 

Springer 49 

Sears, Mary 3 

Shreve, Benjamin 16 

Springer, Stewart, Henry B. Bigelow and W. C. 

Schroeder 49 

Tanaka, Otohito 64 

Taylor, Edward H 1 

VAN Frank, Richard 45 

Whittington, H. B. 4, 17, 38 

Williams, Ernest 2, 13, 25, 28, 29, 32, 35, 39 

Williams, Ernest E. and Carl Gans 36 

Williams, Ernest E., Karl F. Koopman and Thomas E. 

Reynolds 12 

Wood, Albert E 41 



E V I O R A 

Miiseiiim of Coimparative Zoology 

Cambridge, Mass. February 8, 1952 Number 1 

A NEW PANAMANIAN TREE FROG 
By Edward H. Taylor 

Department of Zoology, rriiversity of Kansas, Lawrence, Kansas 

While studying ( "entral American frogs in the Museum of ( "om- 
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 ( "entral America, I conclude that the species is new 
and I present herewith its description. 

HyLA SHREVEl Sp. nOV. 

Tiipr. M.C.Z. No. 2(i7()9 cf . La Loma, Chiriquicito, Republica de 
Panama; K. R. Dunn and ('. Duryea, collectors. 

Diagnoftis. \ diminutive frog (25.2 mm.) with a br(nid, rather flat 
head ; skin smooth ; outer fingers with a trace of web, and well-developed 
discs; canthus rostralis wanting; nostrils nearly terminal; tympanum 
large directed strongly upward, its diameter at least half of eye length; 



2 BREVIORA NO. 1 

eye directed obliquely forward; \'onierine 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 ti/pe. 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 (3 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 of 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 ecjually 
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 
\ -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; discs 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 l)etween second and third; approximately 
half webbed between the third and fourth and two fifths webbed be- 
tween the fovu-th and fifth, the web forming a slight ridge to discs 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 l)etween upper and lower surface of thighs, followed 



1052 NEW TREE FROG 3 

l)y 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 l)rown 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 zetcki 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 (^ instead of ^s oi 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. 



NO. 1 



«Ov- 




Hyla shrevci up. n.uv. AI.C'.Z. Xu. 20709, La Lonia, Chiruiuicitu, Republicu 
de Panamd. E. R. Dunn and C. Duryea, collectors. (Type d^ X 2.) 1. Dorsal 
view. 2. Lateral view of head. 3. Enlargement of nuptial spines on inner 
digit. 



/Uc'^i- /K 



r 



B R E V I 

Mesemm of Comparsitive Zoology 

Cambridge, Mass. February 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 Xenochelys 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 Staurotypus-Wke. 
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 Xcnochclys 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, Xcnochclys is a good structural inter- 
mediate between Dermatcmys 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 Xcnochclys. He quite explicitly cited Staurotypus 
and Claudius as "the nearest living relatives of Xcnochclys" (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 scnsu 
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 concomitantlv 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- 

' 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, Staurotj/pus salvhiii, and Stcrnothenis 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 Stauroiypus or 
Stcrnotherus. 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 Staurotypvs and Sternothents. The depth of the 
premaxilla is markedly greater than in Stavrotypvs. The orbit is rela- 
tively smaller than in Siavroiypvs 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 V3 and ^-^ as in Staurotypus. 

In lateral view the prootic is seen to project far foi-ward 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. x\ similar forward projection of prootic is seen in Stauroiypus 
(and in Graptcmys) but is far less extreme. 

The "crista praetemporalis" which is the feature exaggeratedly de- 
veloped in the fossil and in Stauroiypus and Graptcmys 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 Graptem-ys 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 Staurotypus, 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 Staurotypus 
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 Stcrnothcrxis, 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 Stauroiypiis, 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 Staurofj/pus- and Stertlother^ls, not as in Claudius. 

The sutures bounding the frontals are somewhat obscured by breaks, 
but it is extremely probable that as in other (helydridae 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 Sternothcrus (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, Macrodeviys, 
Staurotypus, Claudius, Kinosternon, Stcryiotherus), 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 

I A (liscimsion of the antitoniy of Claudiut is in preparation. 



6 BREVIORA NO. 2 

to refer Adelochclys (= 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 tliree 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 Chdydra, 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) 



1952 A STAUROTYPINE SKULL 7 

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., Terrapcne, Cvora) have a beak quite similar to that of chelydrids. 
Most testudinids (but not Mcdaycmys) 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 steepl}^ 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 
prema,xillary 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 diflference 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 Staurotypus 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 Staurotypus. 
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 9 

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 Xenochclys 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, Gafsachclys 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 Hay 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 Baptcmys (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 staurotypinesandkinosternines, 
and in Hoplochelys the shell is tricarinate, as it is also in one species of 
Baptemys. The hiunerus 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 Baptcmys 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 Xenochelys 
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" dccheni 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" miirchisoni Bell and four other named species of "Chelydra" 
and Chelydropsis earinata Peters record the rather widespread oc- 
currence of the subfamily. The remarks above for "C." dccheni apply 
also to "C." murchisoni and the other European forms referred to 
"Chelydra." 

The carapace of Chelydropsis earinata 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 II 

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 Chdydropsis is available. 

From the Miocene of North America tliree chelydrines have been de- 
scribed. Two are known from skulls only: Chelydrops strida Matthew 
and Macrodemys schmidti Zangerl. Both of these are from Nebraska, 
M. schmidti from the Middle Miocene, Chelydrops strida from the 
Upper Miocene. Both are certainly related to Recent M. temminddi, 
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. tcm- 
minckii 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 hcckmani, 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. temminckii. Gilmore (1923) has described from the San 
Pedro Valley of Arizona, either Pliocene or Pleistocene, a Kinosternon 
which is said to differ from Recent A', flavcscens mostly in size. 

Two species of Chdydra and one of Macrodemys 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 Macrodemys temminckii. 

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 Xenochdys formosa and Chelydrops strida. Dr. A. S. Romer and 
IVIr. 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 ^fO. 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 Zangerl 

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 lemminckii (Holbrook) Zangerl 
Early Pliocene (S. Dakota) North America 

Pleistocene: 

Macroclemys floridana Hay 

Pleistocene (Florida) North America 
Chelydra laticarinata Haj- 

Pleistocene (Florida) North America 
Chelydra sculpt a Hay 

Pleistocene (Florida) North America 
Kinosternon arizonerise Gilmore 

Pleistocene (Arizona) North America 
and additional North American records for M. temtninckii 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 HistorjO- London. 311 pp. 

FucHS, Erika 

1938. Die Schildkrotenreste 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. Mas., 
Vol. 62, pp. 1-8. 

Glaessner, M. F. 

1933. Die Tertiarschildkroten Niederosterreichs. Neuea Jahrb. Min. 
Geol. Pal., Abt. B, Vol. 69, pp. 353-387. 

Hat, O. p. 

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. 

Hoffman, C. K. 

1890. Schildkroten in Bronn's Klassen und Ordnungen des Tierreicha. 
Leipzig. 442 pp. 

Laube, G. C. 

1 900. Neue Schildkroten und Fische aus der bohm Braunkohlenf ormation : 
Abhandl. deutsch. naturwiss.-med. Ver. Bohmen "Lotos", Vol. 2. 
no. 2, pp. 37-56. 

Matthew, 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. Schildkrotenreste aus den osterreichischen Tertiar-ablagerungen. 

Denkschr. math-naturwiss. Kl. Akad. Wiss. Wien, Abt. 2, Vol. 9, 

pp. 1-22. 
1869. Zur Kenntniss der Wirbelthiere aus den Miocanschichten von 

Eibiswald in Steiermark. I. Die Schildkrotenreste: Denkschr. 

math-naturwiss. Kl. Akad. Wiss. Wien, Vol. 29, pp. 111-124. 



14 BREVIORA NO. 2 

POMEL, A. 

1854. Catalogue m^thodique et descriptif des vert^br^s fossiles d^cou- 
verts dans le bassin hydrographique superieur de la Loire et surtout 
dans la valine de son affluent principal I'Allier. Paris. 193 pp. 

Reinach, a. von 

1900. Schildkrotenreste in Mainzer Tertiarbecken und in benachbarten 
ungefahr gleichalterigen Ablagerungen. Abhandl. Senckenberg. 
naturf. Ges., Vol. 24, pp. .3-135. 

SlEBENROCK, F. 

1897. Das Kopfskelet der Schildkroten. Sitz.-Ber. Akad. Wiss. Wien, 

Abt. 1, Vol. 106, pp. 24.5-328. 
1907. Die Schildkroten Familie Cinosternidae m. monographisch 

bearbeitet. Sitz.-Ber. Akad. Wiss. Wien, Abt. 1, Vol. 116, pp. 

527-599. 
1909. Synopsis der rezenten Schildkroten 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. 

Teppner, W. 

1915. Ein Chelydra-Resi von Goriach. Mitt, naturwiss. Ver. Steiermark, 
Vol. 51, pp. 474-475. 

Williams, 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 Schildkrotenschadels. Bull. Geol. 
Inst. Univ. Upsala, Vol. 19, pp. 89-114. 



1952 A STAUROTYPINE SKULL 15 




PLATE 1 

Staurotypus salvinii M.C.Z. 4989: Dorsal, ventral, anterior and lateral views 
of skull. X about 1 }4- 



16 



BREVIORA 



NO. 2 




PLATE 2 

Xenochelys formosa Princeton 13686: Dorsal, ventral, anterior and lateral 
views of referred skull fragment. About natural size. 



BREVIORA 

Miiseiuinni of Comparative Zoology 

Cambridge, Mass. May 23, 1952 Number 3 

NOTES ON SIPHONOPHORES 

3. Nectopyramis spinosa n. sp. 
By Mary Sp^ars^ 

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 Vogtia pentacantha 
Kolliker, I', 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, PI. 25, figs. 5-7) as Hippopodius (?) 
cuspitaia, 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. 



NO. 8 



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. Nectopyraviis, 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 



GG PASLEY DEL 




V. %^^ 



1 V '.-.J^*;;^; 






•ct^- 







B 

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 Nedo-pyramis, as may be the case 
among other prayids, we have two species originally described by 
Bigelow (1911a; 1911b), N. theiis and A'', diomcdeae, both of which had 
branched somatocysts, but simple radial canals and two species, 
N. cuspitata Moser and A^ 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 8IPHONOPHORES 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. 

SI 50. 2000 m. wire. 1 nectophore. 
St. 3677^^. 5°28'S., 130°39'E. 23. III. 1929. 

SI 50. 2000 m. wire. 2 bracts. 
St. 3920^\ 1°12'N., 62°19'E. 9. XII. 1929. 

SI 50. 1000 m. wire. 1 bract. 
St. 3964'''. 25°19'S., 36°13'E. 15. I. 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 N c dopy r amis . 
The nectosac opens into the base of the triangle and like A' . 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 
hydroecium. Oa two of the specimens, two threadlike branches are 
given off dorsally and appear distally to have a globular connection. 



4 BREVIORA NO. 3 

BIBLIOGRAPHY 

BiGELOW, H. B. 

1911a. Biscayan plankton collected during a cruise of H.M.S. "Research" 

1900. Pt. XIII. The Siphonophora. Trans. Linn. See, London, 

Zool., 2nd ser., 10(10): 337-357, pi. 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 pis. 

1931. Siphonophorae from the Arcturus Oceanographic Expedition. 
Zoologica. N. Y., 8(1): 525-592, figs. 185-220. 

BiGELOw, H. B., and M. Sears 

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 1®' de Monaco. Res. Camp. Sci., Monaco, 87: 1-64, 1 
double plate. 

1934. Siphonophores calycophorides de I'Oc^an Atlantique tropical el 
austral. Bull. Mus. Hist. Nat. Belg. 10(6): 1-87, 15 figs. 

1936. Siphonophores calycophorides (suite) et physophorides provenant 
des campagnes du Prince Albert 1®' de Monaco. R^s. Camp. Sci.. 
Monaco, 93: 1-35, 2 double plates. 

Leloup, E., and E. Hentschel 

1935. Die Verbreitung der calycophoren Siphonophoren im Slidatlan- 
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 pis., 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 

Museem of Comparative Zoology 

Cambridge, Mass. June 9, 1952 Number 4 

A UNIQUE REMOPLEURIDID TRILOBITE 
By H. B. Whittington 

LNTRODUC'TION 

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. 

SYSTEMATK " DESCRIPTION 
Family REMOPLEURIDIDAE Hawle and Corda, 1847 

Genus HyPODICRANOTUS gen. nov. 

Type species. Rcmopleuridcs 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 Ili/podicranofu.s stridtulu.^- ditfers from 
that of the type species of Rnuophurids (\\ hittington, 1950, pp. 540- 



BREVIORA 



NO. 4 



543, PI. 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 Rr m o pie u rides; (c) 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. H ijpodicranotus striatuliis (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. ?h = 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 Ili/podicranottts 
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 ('NU^rK RE.MOPI.KnUDIl) THILOHITK 3 

The axial and pleural furrows in Ili/ijodirranofu.s are shallow, there are 
no axial spines, and no segment has the pleural spines exceptionally 
elongated. The pygidium of Ili/podirranotus is poorly known, but 
appears to be rectangular in outline, longer than wide, with the axis 
much shorter than that of Rcmophiiridcs. The dorsal exoskeletons of 
other remopleuridid genera may readily be distinguished from that 
of Hypodicranotus or Rcnwplcuridr.s. 

Hypodicranohis 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 Missom-i and 
Illinois, the Viola limestone of Oklahoma, and from the Ordovician 
strata at Silliman's Fossil Mount, 15afhn Island. 



HvPODRRAXoTrs sTRiATi'Hs (Walcott, bS75) 

Plate 1, figs. 1-10; text-figure 1. 

Lrctofi/pc (selected Raymond, 1925, explanation of Plate 3). Mus. 
Comp. Zool. No. 1()16, 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 Ccrauriis 
plcutrxaufhrmus' Green, 1832; Calliops ntU'urphalus (Hall, 1847); 
Lconaspi.s ? froifoncnsis (Hall, 1847); Diiicdiithtt.sjji.s ? parnila (Walcott, 
1S79); Illnrinis cf. a iiirrlcfuiiis liillings, 1859; Isotdus gigas DeKay, 
1824; Flr.ricdli/iinni' .sciutrid (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). 

Parah/pes. 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: 
( 'ephalon : 

Length (sagittal) 10.4 

(exsagittal) 18.2 

Width at genal angle 16.4 

Height at second glabellar furrow 6.5 

Maximum width across palpebral lobes 11.2 

Length of eye lobe 7.2 



4 BREVIORA .\'0. 4 

Thorax : 

Width axis first segment 10.2 

" " ninth " ,approx 4.3 

" first segment (across tips of pleurae) 13. G 

" seventh segment (across tips of pleurae) 7.7 

Length (sagittal) complete thorax, approx 13.0 

Dimensions of hypostome, Mus. C/omp. Zool. Xo. 1()17, in milli- 
metres : 

Length (sagittal) 5.2 

(exsagittal) 20.2 

Maximum width (at about half the length) 7.4 

Width of middle bofly 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 TNigTE REMOFLErRiDii) tkilobitp: 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 doul)lure (PI. 1, fig. S; 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 (PI. 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 
(PI. 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 doubhn-e is flexed up sharply. Mus. Comp. Zool. 
No. 1618 (PI. 1, fig. 4) shows that the doubkn-e 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 
(PI. 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. G) 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 anfl 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, G. 

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 
(PI. 1, figs. 1, 2, 4). Similar but larger structures are present on the 



1952 T'NIQI'E REMOPLETRIDin TRILOBITK 7 

free cheeks, the genal spine, and the base and outer part of the long 
lateral spine (Pi. 1, fig. 1). There is a tiny median tubercle on the 
occipital ring, closer to the fui-row 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 
(PI. 1, figs. 1-4, 6, 8). The lines tend to run in curves convex inwards 
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 pleiu^ae, 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 l)ody 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 ^^'alcott (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 P^oerste (1920, p. 222, PI. 22, figs. 18 A-C). 
Raymond (1921, p. 31, PI. 9, fig. 7) described a hypostome from the 
Middle Trenton of Trenton, Ontario, and referred it to H. striahdus, 
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 Hi/podicranotus is //. missouriensis (Foerste, 1920, pp. 
220-222, PI. 21, fig. 17; PI. 22, figs. 17A, 17B; Bradley, 1930, pp. 246- 
247, PI. 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. striatidus. The characteristic hypostome of Hypodicranotvs 
also occurs in the Trenton of Duck Greek Quarry, near Green Bay, 
Wisconsin (U. S. Nat. Mus. Nos. 72181, 87087), the Prosser limestone 



8 BREVIORA \0. 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 Cri/ptolifhoidcs ulrichi Whittington, 1941, Trinodus sp., 
Robergia sp., and an asaphid. The specimen figured by Roy (1941, 
p. 155, fig. 114), as Rcmopleurides 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 Ili/podicranotus striatulus, apart from the 
hypostome, is like that of other remopleuridids of Middle Ordovician 
age, e.g. the Irish specimens (\^hittington, 1950, pp. 540-543, PI. 69, 
figs. 5-10, PI. 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, PI. 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 difi'erent 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, PI. 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 
Ili/podirrduotits-. 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 passible movement of the thorax and pygidium in the 
vertical plane, relative to the cephalon, must have been severely 



1952 rXUilK KKMOPLKIKIDII) IKII.OHITK 9 

limited. The mode of artieulutiou of the thorax prechides any eoii- 
siderable movement relative to the cephalon in the horizontal plane. 
There is ample room for the appendage.s 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, etc. Was HiiporUcran- 
ofus a burrowing, crawling, floating, or swimming form? No definite 
answer is possible but I am inclined to think of it as either floating or 
swimming. 

Referexces 
Bradley, J. H. 

1930. P'auiia of the Kimmswick Limestone of Missouri and Illinois. 
Contrib. Walker Mus., vol. 2, no. 6, pp. 219-290, pis. 2.3-30. 

FOER.STE, A. F. 

1920. The Kimmswick and Plattin Limestones of Northeastern Missouri. 
Denison Univ. Bull., J. Sei. Lab., vol. 19. pp. 17.5-224, pis. 21-23. 

K.^y. G. M. 

1937. Stratigraphy of the Trenton Group, l^ull. Geol. Soe. Am., vol. 
48, pp. 233-302, pis. 1-10. 

Raymond, P. PI 

1921. A Contribution to the Description of the Fauna of the Trenton 
Group. Geol. Surv. Canada, Mus. Bull. 31, Geol. Ser. 3S, pp. 1-64. 
pis. 1-11. 

1925. Some Trilol)ites of the Lower Middle Ordovician of PJastern North 
America. liull. Mus. Comp. Zool., vol. (i7, no. 1, pp. 1-1 SO, pis. 
1-10. 

Ross, R..J. 

1951. Stratigraphy of the Garden City Formation in Northeastern 
rtah, and Its Trilobite Faunas. Peabody Mus. Nat. Hist., Vale 
Fniv., Bull. 0, pp. 1-161, pis. 1-36. 

Roy, S. K. 

1941. The Fijper Ordovician Fauna of Frobisher Bay, Baffin Land. 
Mem. Field Mus. Nat. Hist., Geo)., vol. 2, pp. 1-212, 14() fijis. 

Walcott, C. D. 

1875. New Species of Trilobite from the Trenton Limestone at Trenton 
Falls, N. Y. Cincinnati Quart. .J. Sci., vol. 2, j^p. 347-349, fig. 27. 

Whittington, H. B. 

1950. Sixteen Ordovician Genotype Trilol^ites. ,J. Paleont., vol. 24, 
no. 5, pp. .531-565, pis. 68-75. 



EXPLAXATIOX OF PLATE I 

H ypodicranotus striatulus (Walcott, 1S7.5). Trenton limestone, probably 
Denmark member of the Sherman Fall formation, Trenton Falls, Oneida 
County, New York, all X 3, except figure 10, X 17,V2, Mus. Comp. Zool. 
collections. 

Figs. 1-3, 5, 8. Lectotype, Xo. 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, Xo. 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, Xo. 1537, dorsal view. 

Fig. 7. Paratype, No. 1617, ventral view, presumed original of Walcott, 
1875, p. 347, fig. 27a, and Raymond, 1925, PI. 3, fig. 5. 

Fig. 9. Xo. 3269, dorsal view of interior of hypostome. 



BREVIORA 



Unique Remopleuridid Trilobite. Plate 1 




E V I 

Museuim of Comparative Zoology 

Cambridge, Mass. August 29, 1952 Number 5 

TRICHOLIMNAS CONDITICIUS 

IS PROBABLY A SYNONYM OF 

TRICHOLIMNAS SYLVESTRIS 

(Aves, Rallidae) 

By James C. Greenway, Jr. 

Comparison of the type of Tricholimnas conditicius Peters and 
Griscom (Proe. New England Zool. Club, vol. 10, 1928, pp. 99-103) 
with a series of Tricholimnas 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 conditicins was found pre- 
served in alcohol in the Museum of ( 'omparative 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 wa& 
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 (^harles 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 cvcnjthing 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 [iVpaiang 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 hut 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.) "3 — Fuhca — 
common about our fresh water ponds." It is quite unhkely that he 
would consider Tricholimnas to be a sea bird. Nor is it likely that he 
would mistake the short, decomposed tail feather of TricJwUmnas 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 prol)able 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 conditicius 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 tlu-oat 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 syhestris available. Since G. M. Allen, in the 
original description indicated that the skeleton was obviously that of 



4 BREVIORA NO. 5 

an immatiire bird, perhaps it may reasonably be assumed that this 
character is due to age. 

Measurements, in millimeters, are as follows: 

"conditicius" sylvestris lafresnayanus 

cf 9 

Wing 132 135-139 133-142 185 

Tail 58 58-63 61-65 109 

Bill 45 54-56 47-50 52 

Tarsus 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. 



E V I 

Mmsemnn of Coinnparative Zoology 

Cambridge, Mass. August 29, 1952 Number 6 

CONTRIBUTIONS TOWARD A 
RECLASSIFICATION OF THE FORMICIDAE 

I. Tribe Platythyreini (Hymenoptera) 

By William 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 {Proholomyrmex, Escherichia and Eubothro- 
ponera) must be included. Proholomyrmex 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, Psilohethylus, 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 



2 BKEVIORA 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. Proholomyrmex 
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 Proholomyrmex boliviensis and the species of Plaiythyrea, 
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 Proholomyrmex to 
represent a direct derivative of Plaiythyrea modified for a highly 
cryptobiotic existence in the same way, as I also believe, that the 
proceratiines are only modified ectatommines. If Plaiythyrea and 
Proholomyrmex 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 Proholomyrmex, 
and may even be synonymous and representative of an ergatoid or 
other intermediate female caste of one of the Ethiopian Proholomyrmex 
species. 

A fourth genus that must be included in the Platythyreini is Evbo- 
throponera, in spite of its unfortunate name. In his original diagnosis 
of this Australian group, Clark stated, "... certainly close to Boihro- 
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 Eiihoihroponera species 
described to date except E. hrunnipes 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 Platythyrca, 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. \Miat 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. crihrinodis (Gerstaecker) very definitely has short 
palpi, formula 3, 2. In P. schuUzei 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- 

Australian) 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 

Australia) Eubothroponera Clark 

Distinct erect pilosity limited to mouthparts and gastric ape.x. (Tropico- 
politan; occurring naturally or as tramp species in some warm temperate 
regions) 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 Probolomyrmex (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., Fasc. 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, nee 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): E. dentinodis Clark (p. 9,, fig. 1, 
nos. 6, 6a); E. micans Clark (p. 10, fig. 1, nos. 7, 7a); E. bicolor Clark 
(p. 11, fig. 1, nos. 8, 8a); he also included in the genus Forel's Pachy- 
condyla (Bothroponcra) 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, pi. 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 E. septentrionalis (p. 34, 
fig. 17) from Queensland. 

A seventh and last species was added by Clark when he described 
E. brunnipes (1938, Proc. 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. 



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. holiviensis 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. ciL, 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 bremrostris 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. 



E V I O R A 

Museiim 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 
RhachianccU's glaucus (Cope). In 1937 van Deinse and Junge published 
an important paper of which the main part is a thorough comparative 
study of subfossil Kschrichtius rohusius 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 Rhachiaiuctcs glaucu.s 
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 antl 
valuable osteological work, which has demonstrated the generic 
identity of Eschrichtius Gray 1864 and Rhachianrctrs Cope 1869 
(= Agapliclus Cope 1868). But they also argue for specific identity, 
and thus unite RJiacliianccics glaucus (Cope) 1868 with Eschrichtius 
rohusfus (Lilljeborg) 1861. Moreover, they argue that Dudley's scrag 
whale (1725), named Balacna gihbosa 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 gibbosvs 
(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 Oceanographir Institution. 



2 BREVIORA NO. 7 

Additional and better material of E. 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 estaVjlished; 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 Balacna 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 scrag whale (Balacna 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 Balarnoptrra 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 glancus until better material of Eschrichtius 
/•o6t/5/i/5 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 glancus 
(Cope). 
This status may be summarized thus: 
Genus 

Eschrichtius Gray 1864 (p. 350). Type species (by subsequent 
designation by Lilljeborg 1865, p. 494) : Balacnoptera robusta 
Lilljeborg 1861. 
Species 

Eschrichtius robustus (Lilljeborg) 

Balacnoptera robusta Lilljeborg 1861, p. 602. 
The Graso whale. 

Subfossil in northwestern F.urope (English Channel to Baltic 
Sea). 



1952 PACIFIC GRAY WHALE 3 

Eficliric lit ills glaucus (Cope) 

Acjapfuius glaucus Cope 1868a, p. 160, 1868b, p. 225. 

Rhnchiancdcs glaucus Cope 1869, p. 15. 

The California or Pacific gray whale; devilfish. 

Recent in North Pacific Ocean (Baja California to Korea). 

REFERENCES 

Ccderlund, B. A. 

1939. A subfossil gray whale discovered in Sweden in 1859. Zjologiska 
Bidrag fraan Uppsala, 18, pp. 269 - 286, 5 pis. 
Cope, E. D. 

1868a. [Remarks on Cetacea at meeting of 23 June.] Pioc. 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, pis. 4-11. 
Dudley, Paul 

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. 
F]rxleben, J. C. P. 

1777. Systema regni animalis . . . Classis I. Lipsiae, pp. xlviii + 
636 + [66). 
Gray, J. E. 

1864. Notes on the whalebone-whales; with a synopsis of the species. 
Ann. Mag. Nat. Hist. (3) 14, no. 83 (Nov.), pp. 345-353. 

LiLLJEBORG, W. 

1861 . Hvalben, funna i jorden paa Grason i Roslagen i Sverige. Forhandl. 
Skandinav. Naturf., 8de M0de, 1860, Ki0benhavn, 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 Eschrich- 
tius robustus. Ann. Mag. Nat. Hist. (3) 15, no. 90 (June) 
pp. 492 - 495. 



BREVIORA 

MiuseTuinn of Coumparative Zoology 

Cambridge, Mass. October 21, 1952 Number 8 

A NEW SPECIES OF THE 

(TCLOSTOME GENUS PARAMYXINE 

FROM THE GULF OF MEXICO 

By Henry B. Bigelow and William C. Schroederi 

Up to the present time the myxinids, represented in the North 
Atlantic by the unibranchiate genus Myxine 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.^ 
Paramyxine, 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 interna,l 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 1st 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, 

1 Contribution No. 610, from the Woods Hole Oceanographic Institution. 

2 We owe to Mr. Stewart Springer the opportunity of studying these interesting specimens. 



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; 1st 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 
Myxine, but a somewhat broader and more paddle-shaped caudal fin 
fold. 

The gill openings of the type specimen of atami 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 atami and with the National Museum 
specimen, in body form, with rather pointed snout and a caudal fin 
fold somewhat wider than that of Myxinr. 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; 

' As scaled from Dean's (1904, pi. 1. fig. 3) ilhistration. 



1952 



NEW SPECIES OF CYCLOSTOME 



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 atarni, 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 
1st gill opening to 6th gill 

opening 2.4-5.7 2.5 2.4 

6th gill opening to tip of caudal 

fin fold 72.0-74.3 71.4 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 13.2 

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 

cloacal pocket 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 

I From 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. S. 
National Museum specimen of atami (Fig. 6), just as it is in the Gulf 
of Mexico specimens (Fig. 5) — in Mi/xine 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 1st 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, 
pi. 1, fig. 3; Matsubara 1937, pi. 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, pi. 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 

1 The anal fin fold also, of the National Museum specimen of atami 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. mitsukurii (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 mirahiUs (CoUett) 1904, or than it is to H. collei 
(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. atlaniis 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 1st gill openings, 9, 7.8, 8.9. 

1 For a synopsis of the species of myxinids, see Holly, 1933. 

' Named after Stewart Springer who discovered this interesting cyclostome. 



6 BREVIORA NO. 8 

Distance, snout to 1st gill openings, 23.5, 23.2, 22.5. 

Distance, 1st 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 1st gill openings, 38, 34, 37. 

Number of mucous pores on each side, type (590 mm.), 505 mm. 
specimen and 338 mm. specimen: 

Snout to 1st 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 Myxine; 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 1st 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 Myxine. 

2 The teeth of the type specimen cannot be counted without dissection. 



1952 NEW SPECIES OF CYCLOSTOME 7 

The six pairs of gill openings are low down on the sides, their fore-aft 
position as given in Table 1 (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 1st 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 1st 
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 1st 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 



BREVIORA 



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. atami 
as pictured by Matsubara (1937, pi. 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 
BiGELOw, Henry B., and William C. Schroeder 

1948. New genera and species of batoid fishes. Journ. Marine Research, 

vol. 7, no. 3, pp. 543-566. 
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. 
1951a. Three new skates and a new chimaerid fish from the Gulf of 
Mexico. Journ. Washington Acad. Sci., vol. 41, no. 12, pp. 383-392. 
Brito Capello, Felix de 

1868. Catalogo dos peixes de Portugal. Jorn. Sci. Math. Phys. Nat. 

Acad. Lisboa, vol. 1, no. 3, pi. 3, fig. lA. 
1868a. Descripcaao de dois peixes novos de Portugal. Jorn. Sci. Math. 
Phys. Nat. Acad. Lisboa, vol. 1, no. 4, pp. 314-317. 
Collett, R. 

1904. Diagnoses of four hitherto undescribed fishes. . . . Selsk. For., 
1904, no. 9, 7 pp. 
Dean, Bashford 

1904. Notes on Japanese myxinoids. Journ. Coll. Sci. Imp. Univ. Tokyo, 
vol. 19, art. 2, 23 pp., 1 pi. 
Holly, 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, Kiyomatsu 

1937. Studies on deep sea fishes of Japan. IIL On some remarkable 
variations found in Paramyxine atami Dean. Journ. Imper. 
Fisheries Inst. Tokyo, vol. 22, no. 1, pp. 13-15, 1 pi. 



10 



NO. 8 




^^^'^^^'-^i^n^'.^. Ai^A.iJ&d,^iS,:.».:.«^.m^S^ 




Fig. 1. Type specimen, x 34; the mucous pores somewhat exaggerated. 

Fig. 2. Ventral view of anterior part of type specimen, x }4', 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 J-^; 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. 



E V I O R A 

MeseeitTii of Coimparative Zoology 

Cambridge, Mass. October 21, 1952 Number 9 

NEW SPECIES OF EARTHWORMS FROM THE 
ARNOLD ARBORETUM, BOSTON 

By G. E. Gates 
I. 

From Allolobophora caliginosa (Savigny) 1826 there have recently 
been split off: yl. nociurna Evans 1946, and A. iowana 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 
xxxiii. Further study showed that those specimens with tumescences 
on xxxiii were referable to caliginosa as apparently restricted by Evans 
after breeding the worms in the laboratory. The worms with no 
tumescences on xxxiii 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, J/^xxvii, or xxviii to xxxiv or V2XXXV. Tubercula 



2 BREVIORA NO. 9 

pubertates of double origin, on xxxi-xxxiii. Genital tumescences, in- 
cluding a and b setae, on ix-xi, xxx, xxxii, xxxiv, and occasionally also 
on XX vi. 

Spermathecal pores, on 9/10-10/11 on r lines, setae, male poro- 
phores, calciferous sacs, calciferous glands, typhlosole and last hearts, 
etc., as in caliginosa. 

Type locality. Arnold Arl)oretum, Boston, Mass. 

Distrihution. Known at present only from Boston and vicinity. 

Remarks. A. arnoldi is distinguished from caliginosa by the constant 
absence of genital tumescences on xxxiii, occasional presence of tumes- 
cences on xxvi (instead of xxvii), slightly greater anterior extent of the 
clitellum (xxvii or xxviii instead of 3^xxviii or xxix), greater number 
of segments, and perhaps by a slightly more anterior first dorsal pore. 

From iowana, arnoldi is distinguished by the constant presence of 
genital tumescences on xxxiv, constant absence of those tumescences 
on xxxiii, by the two-part tuberculum about as in caliginosa, 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 noctnrna, 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 arnoldi in the Boston 
area will be presented, if possible, in another contribution. 

II. 

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 0. lacieum (Orley) 1881, xxx-xxxiii 
and xxxi-xxxiv; Lumbricus rnhellvs Hoffmeister 1843 and L. castaneus 
(Savigny) 1826, xxviii-xxxi and xxix-xxxii. 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 xxviii-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 fa%mo*'a-complex only by differences in tubercula 



1952 NEW SPECIES OF EARTHWORMS 3 

and tumescences as specifically distinct. 

Allolobophora molita n. 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 ?12/13. 
Clitellum on xxviii-xxxiv, possibly reaching slightly onto xxvii. 
Tubercula pubertates, longitudinally placed bands in he, extending 
across entire lengths of xxx-xxxiii; with lateral margins nearly smooth, 
but median margins (quite definitely lateral to h lines) slightly indented 
by 30/31, 31/32, 32/33. Genital tumescences, including a and h setae, 
present on ix-xi, xxxii and xxxiii (feebly developed). 

Spermathecal pores, male porophores, setae, calciferous sacs and 
glands, typhlosole, last hearts, as in ealiginosa. 

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 ealiginosa- 
complex: caligmosa, noetvrna, 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- 
xxxiii. From all of the complex, except possibly iowana, molita is also 
distinguished by the quadripartite (instead of double) origin of the 
tubercula. In iowana, 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) 13: 98-101. 
1948. On some earthworms from Iowa, including a description of a new 

species. Ann. Mag. Nat. Hist. (11) 14: 514-516. 



E V I O R A 

Musemmti of Coiraiparative Zoology 

Cambridge, Mass. December 29, 1952 Number 10 



ON THE EARTHWORMS 
OF NEW HAMPSHIRE 

By O. 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 C1917') 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 diflferent species must have been introduced many 
times, even in the deep woods." fDarlington, in lit.^ 

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. T. Darling- 
ton, Jr. collector. 

Length, 95-105 mm. Diameter, 5.5 mm. Segments, 91 (amputee), 
106 (amputee), 156, 158. First dorsal pore, ?12/13 (1), ??13/14 (2). 
Clitellum, -Ixxvii-xxxiv. Tubercula pubertates bipartite and. as in 
A. caliginosa (Savigny) 1826. Genital tumescences, on ix-xi (4), xxx, 
xxxii and xxxiv (4), xxix (1). 

Remarks. Alive, these worms appeared to be quite without pigmient 
and they looked in that respect much as Orfolasiuni ci/avpvm (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 intersegmental 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, xxxii and xxxiv (with none on xxxiii) could also be distinguished 
from calighma hy greater rlitellar length nnd by larger segment 
number. 

The presence of worms with arnoldi 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 arvnJrJi in many other states, as well 
as in Canada, may be anticipated. 

REFERENCES 

CRATES, G. E. 

1952. New species of earthworms from the Arnold .\rborptum. Boston. 
Breviora, No. 9: 1-3. 
Smith, F. 

1 01 7. North American earthworms of the family Lumbricidae in the col- 
lections of the United States National Museum. Proc. U. S. Nat. 
Mus. 52: 157-182. 
Southern, R. 

1910. A new species of enchytraeid worm from the White Mountains. 
Proo. Acad. Nat. Sci. Philadelphia, 62: 18-20. 



E V I O R A 

Mmseiuiiii of Cormparative Zoology 

Cambridge, Mass. IMarch 20, 1953 Number 11 



CHARACTERS AND SYNONYMIES AMONG 
THE GENERA OF ANTS 

Part I 
By William L. Brown, Jr. 

Museum of Comparative Zoology 
Harvard University 

Work now under way is aimed at a revision of tlie 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. 

Myrmecia Fabricius 

Myrmecia Fabricius, 1S04, Sj'st. Piez.: 423. Genotj-pe: Myrmecia gulosa 

Fabricius, by designation of Emery, 1911. 
Myrmecia (Promyrmecia) Emery, 1911, Gen. Ins., Fasc. 118: 18-19. Sub- 
genotype: Myrmecia aherrans Forel, by original designation. NEW 
SYNONYMY. 
Myrmecia Clark, 1952, Formic. Australia, Melbourne, 1: 21. 
Promyrmecia Clark, 19.52, Ibid., p. 119. 

The references of Clark should be consulted for further synonymy. 
Clark has stoutly defended the separation of Myrmecia and Pro- 
viyrmccia 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 both directions, 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 Promyrmccia. 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 Promyrmccia. 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 Promyrmccia, although he 
wrongly maintains Myrmecia nigrocincia Fr. Smith, its type, in the 
genus Myrmecia as he has defined it. 

Rhytidoponera Mayr 

Ectatomma (Rhijtidoponera) MajT, 1862, Verh. zool.-bot. Ges. Wien 12: 731. 

Genotype: Ponera araneoides Le Guillou, bj- designation of Emery, 1911. 
Rhrjtidoponera {Chalcoponera) Emery, 1897, Ann. Mus. Stor. Nat. Geneva 

38: 548. [Sub] genotype: Ponera metallica Fr. Smith, by designation of 

Emery, 1911. NEW" SYNONYMY. 
Rhytidoponera and Cfialcoponera 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 Rhylidoponcra 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 antenna! 
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. 11 

are intergradient and fully ambiguous in a number of species, among 
them R. reticulata Forel, C. lamellinodis Santschi (paratypes), C. duhia 
Crawley, and all the species of the C. impressa Ma\T 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 Rhytidojjonrra s. sir. 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. fulgcns, numcensis, atropurpurca and 
acupuncta) make an exception to the character that I have attributed 
to the subgenus concerning the structure of the antennae (Genera 
Insectorum, fasc. 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 tnrneri 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 viayrt 
Emery, araneoidcs Le Guillou, and punctata Fr. Smith, and one toward 
the type of metallica Fr. Smith and victoriae Andre. In the farfiung 



1953 ANT CHARACTERS AND SYNONYMIES 5 

regions penetrated by Rhytidoponcra 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 Rhytidoponcra 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 obser\'er, form an effective bridge joining 
these two groups. 

Before leaving Rhytidoponcra, a word is required on the females. 
It has been assumed that Chalcoponcra species always produce normal 
winged females, and that Rhytidoponcra s. sir. 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. imprcssa group {imprcssa, chalyhaea, splcndida) 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 mctaUica group, a situation of a cjuite different sort 
is apparent. Among the common and widespread members of this 
group, such as mctaUica, tasmanicnsis, and their very close allies, 
winged or obviously dealate females are very rare or unknown. The 
victoriac 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 
victoriac floundering in the damp sand at the surf edge at Seaford, on 
Port Phillip Bay, in Victoria. Of several thousand nests of victoriae, 
mctaUica, tasmairicnsis and very close allies of these species that I have 
opened, not one (with the single noted exception above) has yielded 



6 BREVIORA NO. 11 

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 lefer 
to the very widespread occuiTence 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. sir. 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. aspcra (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 



1953 ANT CHARACTERS AND SYNONYMIES 7 

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 Rhytidoponcra 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 Triodia 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 Rhytidoponcra and Camponohis. 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 Rhytidoponern 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. viayri Emery, is, with its close relatives, a 
primal ily 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 Rhyiidoponcra species suggest the 
following hypothesis, now being tested by Haskins {in Hit.). 

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 

' 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 sufficientlj' clear to convince me that his 
species are really distinct. 



8 BREVIORA NO. 11 

or highly modified ergatoids. It is possible that apparatus for the 
retention of sperm is strongly 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. IMeager 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 

Centroinynnex Mayr, 1866, Verh. Zool.-bot. Ges. Wien 16: S94. Ckniotype: 

Cenlromyrmex bohemani Mayr, monobasic. 
Typ/i'o/eros Kara wajew, 1925, Konowia 4: 128. Genotype: Tijphloteras 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 Centromyrmex hamidatus 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 de\elopment 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 
Ti/phlotcras with Ccniromyrmcx, it becomes necessary to point out 
that the subtribe Centromyrmicini of Emery is a NEW SYNONYM 
of tribe Ponerini if the spur characters wnll not hold. 

Pristomyrmex Mayr 

Pristomyrmex Mayr, 1866, Verh. Zool.-bot. Ges. Wieu 16: 903. Genotype: 

Pristomyrmex pungeiis Mayr, monobasic. 
Odoniomyrmex Andre, 1905, Rev. Ent. Caen 24: 207. 
Hylidris Weber, 1941, Ann. Ent. See. Amer. 34: 184, 190. Genotype: Hylidris 

rnyersi 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- 
viyrmex. 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. mycrsi 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 Pristomyrmex s. str. in any direct state- 
ment. Instead, Weber gives briefly his views on generic limits, w4iich 
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 7iot 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 oftered 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 synon^Tn 
Odontomyrmex, a name put forward by Andre for a species that appears 
to have the chief definitive characters of Hylidris. IVIann (1919, Bull. 
Mus. Comp. Zool. 63: 341) delivered the coup de grace to Odonto- 



10 BREVIORA NO. 11 

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 
Museiun 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 bengkalisi Menozzi, monobasic. 
Solenomijrma Karawajew, 1935, Treubia 15: 103. Genotj'pe: Solenomyrma 

acanlhina Karawajew, monobasic. NEW SYNONYMY. 
Acalama M. R. Smith, 1948, Jour. N. Y. Ent. Soc. 56: 20.5-207. Genotype: 

Acalama donisthorpei M. R. Smith, monobasic. NEW SYNONYMY. 
This genus is very doubtfully distinct from Vollcnhovia 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 donisthorpei were 
examined, and are considered specifically distinct, but not generically 
so. Specific synonymy is formalized as follows: 

Gauromyrmex acanthinus (Karawajew) new combination 
Solenomyrma acanlhina Karawajew, 193.5, 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 bengkalisi have been 
examined through the kindness of Dr. Smith. The former species has 



1953 ANT CHARACTERS AND SYNONYMIES 11 

been compared with Karawajew's description and figure, and also 
with the West Chinese series taken by myseh', and mentioned by 
Smith {loc. cit., p. 206). The comparison gives the distinct impression 
that one is deahng 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 wartimic 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 Lcj^totliorax, 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- 
onopelta punctulata Mayr, monobasic. 

Ponera Fr. Smith (partim), 1860, Journ. Proc. Linn. Soc. London, Zoo). 4 
(suppl.): 105, nee Latreille. 

Examblyopone Donisthorpe, 1949, Ann. Mag. Nat. Hist. (11) 15: 401. Geno- 
type: Exambhjopone churchilli Donisthorpe, monobasic: vide infra. 

Renea Donisthorpe, 1947, Ann. Mag. Nat. Hist. (11) 14: 183. Genotype: 
Renea testacea Donisthorpe, monobasic. {Nee 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 



12 BREVIORA NO. 11 

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 Prlonopclta majuscula follows : 

Prionopelta majuscula Emery 

Ponera similliina Fr. Smith, 1860, Journ. Proc. Linn. Soc. London, Zool. 4 

(suppl.): 105, "worker," 9, nee Fr. Smith, 1860, op. cit., p. 104. NEW 

SYNONYMY. 
IRhopalopone simillima, Emery, 1900, Term. Fiizetek 23: 311. 1911, Gen. 

Ins. 118: 35. 
Prionopelta majuscula Emery, 1897, Term. Fuzetek 20: 595-596, worker, 9. 

Brown, 1951, Bull. Brooklyn Ent. Soc. 46: 102, Examblyopone churchilli 

synonymized. 
Prionopelta poultoni Donisthorpe, 1932, Ann. Mag. Nat. Hist. (10) 10: 462, 

nam. pro Ponera simillima Fr. Smith, IL NEW SYNONYMY. 
Examblyopone churchilli Donisthorpe, 1949, Ann. Mag. Nat. Hist. (11) 15: 

401-402, 9. 
Renea testacea Donisthorpe, 1947, Ann. Mag. Nat. Hist. (11) 14: 18.3-186, 

fig., worker, 6"; pp. 590-591, 9. NEW SYNONYMY. 
In his original description of Ponera simillima II, 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 9 " 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 majuscula better than it does either of the other two 
Prionopelta species known from the Indo-Papuasian area (P. kraepelini 
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 Mafiin 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 E. churchilli. 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 E. churchUli type. The R. testacea 
workers differ as expected from authentic workers of PrionopcUa 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). 



B R E V I 

Miiseuim of Coimparative Zoology 

Cambridge, Mass. April 23, 1953 Number 12 

A CAVE FAUNULE FROM WESTERN PUERTO 

RICO WITH A DISCUSSION OF THE 

GENUS ISOWBODON 

By Thomas E. Reynolds 

Zoology Department, Massachusetts College of Optometry 

Karl F. Koopman 

Biology Department, Queens College 

Ernest E. Williams 

Harvard University 

In spite of the diligent v^^ork 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 
Isolohodon 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 C'ueva Monte 
Grande between Mayagijez 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 cuvicri, 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 
Noctilio 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. fuUginosa 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-raandibles. 

Recorded by Anthony from six localities: Cayey, Comerio, Corozal, 
Morovis, Pueblo Viejo and Trujillo Alto. 

Artibeus 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 obliquus 

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 Isolohodon and Aphadrcvs. Even a cursory inspection of 
the molar pattern, however, rules out Aphadrcvs (Miller 1929a, pi. 2). 
The specimens then seem referable to Isolohodon. The only species of 
that genus which has previously been recognized on Puerto Rico is 
Isolohodon portoriccnsis, 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 Isolohodon portoriccnsis 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 /. portoriccnsis and require special attention. All are 
much smaller in size than fully adult /. portoriccnsis. 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 /. portoriccnsis. 

The third small mandible is larger than the other two and although 
considerably smaller than adult /. portoriccnsis 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 cjuestionable Monte 
Grande mandibles as /. portoriccnsis, 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 Isolohodon with which it has been 



1953 A PUERTO RICAN CAVE KAUNULE 5 

compared. In any Isolohodon 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 Isolohodon portori- 
ceTisis 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 Isolohodon portoricensis . 
On the other hand, it may represent a rare mutant condition in a 
subadult individual of /. 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 /. 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 Isolohodon material from Puerto Rico was carefully com- 
pared with material identified as Isolohodon 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. 

/. levir was first described by Miller (1922) as a distinct genus and 
species, Ithyodontia 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 Isolohodon 
portoricensis without comment. In 1929, after studying much more 
material from the same locality, he realized that the two molars of 
''Ithyodontia" were actually upper molars of Isolohodon. In an attempt 
to save /. levir from complete synonymy, he noted that all St. Michel 
specimens were smaller than Isolohodon from other Hispaniolan 



6 BREVIORA NO. 12 

(kitchen midden) localities. The latter, in turn, were indistinguishable 
from Puerto Rican /. 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 /. portoricensis came from kitchen 
middens, whereas /. Icvir came from owl 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 Aphaetreus ruled out the suggestion that in 
the two types of Hispaniolan Isolobodon he was dealing with large- and 
small-sized samples from the same population. In later papers (Miller 
1929b, 1930) he records /. Icvir from several other Hispaniolan locali- 
ties (San Gabriel, Monte Cristi, Constanza, Trujin), in Monte Cristi 
actually in association with /. portoricensis. 

After extensive comparisons of /. portoricensis from Puerto Rico 
and surrounding islands (AMNH material) with /. levir from Hispa- 
niola (MCZ material from Fort Liberte 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 /. levir with the largest /. 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 Ithyodontia 
levir Miller a complete synonym of Isolobodon 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 7 

Acknoiiiedgmcnts. 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 e.xtinct octodont from the island of Porto Rico, West Indies. 
Ann. N. Y. Acad. Sci., vol. 27, pp. 17-22. 

Anthony, 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. 

Miller, G. S. 

1918. Mammals and reptiles collected by Theodoor de Booy in the 
Virgin Islands. Proc. U. S. Nat. Mas., vol. 5-4, 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. 



BREVIORA 



NO. 12 






PLATI'] 



Crown views of mandibles of Isolobodon. A. "Isolohodon kvir". U.S.N.M. 
No. 255874 from Monte Cristi, Dominican Republic. B. Isolobodon portori- 
censis. A.M.X.H. "0.1" Utuado, Pvierto Rico. C. Aberrant Monte Grande 
mandible. M.C.Z. Cueva Monte Grande, Puerto Rico. (These specimen.s, all 
of about the same size, have been selected to show the extremes of variabilit.>- 
of molar pattern.) 5 x natural size. 



E V I O R A 

Mii]is(eii]im of Cooipsirative Zoology 

Cambridge, Mass. Ai'kil 23, 1953 Number 13 

FOSSILS AND THE DISTRIBUTION 
OF CHELYID TURTLES 

1. ''Hydraspis'^ leithii (Carter) in the 
Eocene of India i.s a Pelomedusid 

By Ernkst Williams 

For SO yetirs a fossil turtle from the Eocene of India has heen 
referred to the Recent South American chelyid genus Ili/drdspis (not 
of Bell 1S2S, correctly called Phrj/uopft according to Stejneger 1909. 
and Lindholm, 1929). In spite of the zoogeographical interest of this 
record the evidence for this assignment lias not ])rc\i()usly heen r(>- 
examined critically. 

In view of the prevalent idea that the family reached Austi-alia from 
.Vsia, the occurrence of a chelyid in the Eocene of India would he 
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 Jiufl the 
family assignment. 

The fossil in question was brought to scientific attention just on (m- 
100 years ago. In 1852 H. J. ( "arter in a study of the geology of the is- 
land of Homljay described as Tcsiudo Irithii* 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 Tcsiudo in a 
Linnaean sense and explicitly stated that he regarded his fossil .is close 
to "Sfrrnotharrus" (= Prlusius). He published two good plates giving 
;i reconstruction of the fossil made from nine partial specimens. These 

Tt.<tiido leHhii Carter l'<.'il' !jreo(.-(-ui)ie> Ti.-iudu Intlm Guuther 
I'-gvpt. The :iv:iilable Miui i-nrreet name of the hitter then appears t 
I.ortel 18S:j 



2 BREVIORA NO. 13 

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 Hi/drasph-; 
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 scute in the Indian fossil. Lydekker (1889b, p. 170) stated: 
"The omission of a nuchal shield in the restoration of the anterior 
l)order of the carapace is probably incorrect." Perhaps, however, 
Lydekker made this statement only on the ground that if the form 
were Ili/draspi^' it should possess a nuchal scute. Absence of a nuchal 
scute would rule out all Recent genera of Chelyidae except Chrlodina 
and Emydura (in both of which the scute may be present or absent) 
and Eheya (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 ( helyi- 
dae. In most Recent South American forn s including most cf tie 
species of Hiidraspis! ( = Phriinops) the first vertebral is on the contrary 
much larger than the second. There is, it is true, an apprcach to the 
condition of the fossil in the Australian genera Enu;dnra, Ps( iidcnndvia 
and Khrya anfl in the South American Ilydraspis fiibrw.sa (specirrers 
in the British ^Museum) and perhaps in some specimens of the South 
.Vmerican genus Ilydromdusn, 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 Ijrcad inter- 



1953 FOSSILS AND DISTRIBUTION OF CHELYID TURTLES 3 

gular extends from the anterior margin to tlie humero-pectora! sulcus, 
separating the humerals in the midhne. This pattern of scutes on the 
anterior phistral lohe is different from any which is known in the 
modern ( 'helyidae. In all the genera except Chdudina, Psrudoni/dura, 
and rarely in Kini/dura (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 Cliflodina the intergular is 
very large and separates the humerals, indeed extending deep into the 
area of the pectorals, and, except in C. iutrrgularis P'ry, it does not 
reach the anterior phistral margin, the gulars meeting in front of it. 
In Pscudemydura (Siebenrock, 1907) the intergular is like that of the 
Indian fossil in its breadth but as in Chdodinn dips deeply between 
the pectorals. The gular-intergular pattern in the exceptional Kmi/dura 
siiln/lohosa 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 Ihjdrnspis and other chelyids except the forms of the 
Australian region and Batrachrmys. 

The shell, therefore, is not a gooil 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 
ha\'e been long, in strong contrast to the condition in chelyids. The 
skull, shown l)y 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-sciuamosal arch is absent, but a jugal-quadratojugid bar is 
present. In the ("helyidae and in no other turtles the skull is emargi- 
nated from the ventral margin only, and a parieto-sciuamosal con- 
nection is (except in Chclodina) always preserved. In chelyids the 
quatlratojugal 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 desc.-iber, wh > thouT;ht the fossil was closest to the African 



4 BREVIORA NO. 13 

peloniedusid genus Prlusios. 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 Palaraspis Gray, and sometimes in Pclusios). 
An intergular separating the humerals is found in Stercogciiys podocne- 
moidcs and in FJochrh/s prrftcfa as well as in occasional individuals of 
Podocncmis cxpansa. 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 l)orders, as in some Recent Podocnemis and a nimiber 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 "pecidiar also for the under- 
side of the marginal opposite the (pectoral-humeral) suture being 
rather l)roader 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 boimding 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 
woukl 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 Hosasia (Carrington da Costa 1940) 



19.53 P^OSSILS AND DISTKIBITION OF CHELYID TITRTLES 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 DacochcJys 
(Lydekker, 1889a) because there are no comparable parts, that genus 
having been founded on a mandibular symphysis, a part that is missing 
in (barter's fossil. There is also no evidence that Dacochclys is a pelome- 
dusid.* I distinguish Carter's form, with some hesitation, from 
EJochelys Nopcsa 19.31 because the gular-intergular pattern is not 
quite that of E. prrfrcta, the type of the genus, and I am not persuaded 
that the other species referred by Nopcsa to that genus {K. major) 
belongs there. The critical feature of Elochcli/s 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 Sfcrcogrni'.'i podocvrnioidc.s (Reinach, 1903), but with Schmidt 
(1940) I do not believe that podocnrmoidts belongs to the genus 
St(rco(ir)ii/.s. Podocncmoidcs and Icifhii may indeed belong to the same 
genus, but that genus is then imnamed. 

From the better defined genera of the Pelomedusidae the Indian 
form is distinguished by the following combination of characters: 



Carteremys, 



new genu^ 



Type. Trstudo Irlthii Carter 18.52. 

Diagnosis. Skull roof much emarginate from l)ehind; opisthotics 
prolonged backwards in sharp crests; mandilile 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 scars distinctive in shape 
and position. 



* .\n iiieenious 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 conybearii Owen and therefore (according to 
Lydekker and Boulencer 1K87) with Emys ddahichei Bell. No name change is necessary if this 
synonymy is correct and, since the shell of Emys conybearii shows small lateral mesoplastra 
(Lydekker and Boulengtr 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 syn physis, which I regard as quite impossible to assign to family. 
Lydekker's device has most unfortunately brought the name Dacochelys into the literature (for 
e.xample 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 E. 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 conyhenrii Owen. 



6 BREVIORA NO. 13 

Horizon. Intertrappean Eocene of Bombay. 

Comment may now he 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 Xagpur district 
as a second occurrence of ''Hi/draspis Irlthii." The new fragment was 
much larger than typical Cartrrrmi/s Irithil and differed also in the 
much narrower intergular. Lydekker passed over the differences 
casually with a mention of variability in Hjidraspin ( = Phrynops) 
hilarii. It is unlikely that he had in hand any member of the genus 
Hydraspis or Cartcrcmys Irithil. It is probable that he had some other 
pelomedusid, and he had himself previously (1887) described a Podoc- 
nrmis Indico in the Eocene of India. 

Sukheswala (1947. an abstract only) has reported a find of a shell 
of Tc.siudn (— Cnrtrrrmy.s) IrithH in the Intertrappean of Worli Fill. 
Bombay. Here, as in the case of the specimens described by Carter, 
remains of frogs (Indohafrnchus piisillvs) 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 dift'erent 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 chelyids 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 noM- in the possession of the Geological Survey of 
India. 

Two other specimens have since been foimd 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 SurNcy in the same report was a possible young 
shell of Cartcrcmys Icithll 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 



1953 FOSSILS AND DISTRIBUTION OF CHELYID TURTLES 7 

thought of hopefully, it would appear that Caiitrciin/.s Icit/iii is at 
present known with certainty only from the Intertrappean of the 
island of Bombay. 

Acknowlcdgxicnts. Dr. A. S. Romer, Dr. P. .1. Darlington, Dr. P. E. 
Vanzolini, Dr. Karl ¥. 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 discu.ssed with Dr. Rainer Zangerl. 
Mr. Arthur Loveridge. Mr. (\ M. Bogert and Dr. H. \V. 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 Carternnys 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 my insistent questions and for the gift of the photograph reproduced 
as plate 3. 

REFERENCES 

Carrington da Costa, .). 

1940. Um novo queionio 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. 

1S70. 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 Schildkroten 
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 Me.sozoic Chelonia and a tooth of 

?(Ornithopsis). Quart. Jour. Geol. Soo. London, vol. 4.5, pp. 

227-246. 
1889b. Catalogue of the fos.sil Reptilia and Amphibia in the British 

Museum (Natural History). Part III. London. 239 pp. 
1890. Notes on certain vertebrate remains from the Nagpur district. 

IL Part of a chelonian plastron from Phisdura. Rec. Geol. Surv. 

India, vol. 23, pp. 22-23. 
Lydekker, R. and G. A. Boulenger 

1887. Notes on Chelonia from the Purbeck, Wealden and London Clay. 

Geol. Mag., ser. 3, vol. 4, pp. 270-27.5. 
NopcsA, F. 

1931. Sur des nouveau.x 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. Schildkrotenreste aus dem aegyptischen Tertiar. Abhandl. 

Senckenberg. naturf. Ges., vol. 29, pp. 1-64. 
Schmidt, 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. 

SlEBENROCK, F. 

1907. Beschreibung und Abbildung von Pseiidemydurauinbrina Sicbenr. 
und fiber ihre systematische Stellung in der Familie Chelydidae. 
Sitz-Ber. Akad. Wiss. Wien. Math-naturw. Kl. vol. 116, pp. 
120.5-1211. 
Stejneger, L. 

1909. Generic names of some chelyid turtles. Proc. Biol. Soc. Wash- 
ington, vol. 22, pj). 125-127. 

SUKHESWALA, R. N. 

1947. A fossil tortoise (Testudo leithii) from the Intertrapp?an of the 
Worli Hill, Bombay. (Abstract.) Proc. Indian Sci. Congress, 
33rd Session, vol. 3, p. 97. 

WE.ST, W. D. 

1950. General report of the Geological Survey of India for the year 1948. 
Rec. Geol. Surv. India, vol. 82, pp. 1-253. 

Z ANGER L, 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 (darity from the original reconstruction (Plate XI of Carter, 1852). 




Plate 3. Cartcrenujs leithii, photogra])h of dorsal surface of shell. (Courtesy 
Dr. R. X. Sukheswala.) 



E V I O R A 

Miuseeim of Co:mpsirsitive Zoology 

Cambridge, Mass. May _'s. 1953 Xtmher 14 

RECORD (3F A 

HERAIAPHRODITIC^ HORSE^^HOE (^RAR, 

LIMriJ'S POLYPHEMUS L. 

By John P. Baptist 

Fishery Research Biologist 

U. S. Fish & Wildlife Service 

Xewburyp( )rt . Mass . 

A hermaphroditic horseshoe crab, Liinulus puh/phciiiu.s- 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 Li mill 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 
600 mature males measured this summer ranged from 8S 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- 
du "ed 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. 



2 BREVIORA NO. 14 

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 Avere 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. Ou the testis of Limulus. Trans. Linn. Soc. Lond. (Zoo).), (2) 
2: 363-366. 

LOCHHEAD, J. H. 

1950. Xiphosura polyphemus. In Selected Invertebrate Tj^pes, edited by 
F. A. Brown, Jr. pp. 360-381. John Wiley & Sons, Inc., N. Y. 



1953 



A IIERMAPHKOniTIC HORSESHOE CRAB 




PLATE 1 

Upper — Ventral view of popterijr surface of genital operculum (enlaiKcd), 
showing right genital aperture (9 ) and left genital aperture (o^). 

Lower — Photomicrograph of section through testis (X 120). S, spei'm sacs 
containing tail-less sperm. H. hepatopancreas. C, connective tissue. 

(Photographs by Alden P. Stickney) 



NO. 14 




platp: 2 

Dorsal view of Liiiiiiliis pali/phouxs 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.) 



E V I O R A 

MuseiuitM of Comparaitive Zoology 

Cambridge, Mass. May 15, 1953 Number 15 



FURTHER NOTES ON THE EARTHWORMS 
OF THE ARNOLD ARBORETUM, BOSTON 

By G. E. Gates 

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. 



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 










suhruhicunda 


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 





Figures indicate juvenile-aclitellate-clitellate or small juvenile-late juvenile-aclitellate- 
clitellate. 

t 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. 

0. 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 
xxxii (32 + 26), xxvi-xxxii (1). Epidermis of xxv (several + 8) and 
xxxii (several) slightly modified. Tubercula pubertates, on xxviii-xxx 
(35 + 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 (33 + 27), 
xvii (2 -|- 1), xviii (1), xx (3), xxi (1), xxii (-|-1), xxiii (-1-2), xxiv 
(12 -f- 12), xxv (1 + 3), xxvi (-}-l), xxvii (26 -f- 25), xxviii (2), xxxi 
(19 -f 23), xxxii (1), xxxiii (-f 1), xxxiv (-}-l),xxxv (+1). Tumescences 
on xvi may extend well into be and aa or even be united mesially. 
Those on xxxii to xxxv are especially well developed. 

Homoeosis. Male pores, on xvi. Female pores, on xiv. Clitellum, 
on xxvi-xxxii and on xxxiii-xxxiv of left side only. Tubercula pu- 
bertates, on xxviii-xxx (right side), xxix-xxxiii (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 BREVIOEA 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 (3), 115 (2), 116 (3), 117 (4), 118 (3), 
119 (4). Clitellum, on xxvii-xxxii (40). Tubercula pubertates, on 
xxviii-xxxi (5), but extending onto xxvii (40). Genital tumescences, 
including ab, on vi? (1), vii? (1), viii (2), x (6), xi (42), xii (50), xxvi 
(56), xxvii (3). 

Abnormality. No. 1. One segment near the hind end spUt, 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-xxxii, 
tubercula on xxviii-xxxi but extending onto xxvii. Genital tume- 
scences, on xxvi, and possibly also (?) on vii-xiii. 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 of 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***), 04 (*), 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, 
3, 4, 2, 1, 1, 1, 2, 2, 4, 2, 4, 4, 1, 2, 1, 3, 2, 2; viii/5, 3, 5, 1, 2, 2, 3, 0, 
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 \C 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-viii 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 Ixvi (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 ii. 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 xii, 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 
lawati 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 Hi. 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 deferentia end blindly in xxi v. 

Autotomy and Regeneraiion. 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 ofl' 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 eiseni, 



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. subntbicunda, absent in the spring, had 
appeared. One of them, D. subruhicunda, had become dominant nu- 
merically. L. 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, E. rosea, E. tetraedra and L. rubellus 
were no longer represented, but two, 0. lacteum 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. rvbelhis. 

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, 0. lacteum, D. 
subrubieunda, 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 
Cernosvitov, L. and A. C. Evans 

1947. Lumbricidae. No. 6 in Sj'nopses 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: 497-53-1. 
Smith, F. 

1917. North American eartli worms of the family Lumbricidae in the 
collections of the United States National Museum. Proc. U. S. 
Nat. Mus. 52: 157-182. 



E V I 

Miiseuim of Coimpsirsitive Zoology 

Cambridge, Mass. May 28, 1953 Number 16 

NOTES ON THE RACES OF Micrurus frontalis 
(Dumeril, Dumeril 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. /. frontalis 
(21 ex.), M. f. altirostris (21 ex.), and M. f. pyrrhocrxjptus (4 ex.).. 

I have retained altirostris as a race o{ 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 cm-ators, 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 fm-nishing 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 7-15; ventrals 216-2'42 2 

Parietals exhibit a pale area at least anteriorly; triads on body 13-17 
(13 only rarely)'; ventrals (197) 203-220 altirostris 

2. Light nuchal area between posterior end of parietal suture and first black 

ring from M to 3J^ 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 black outer rings frontalis 

Light nuchal area between posterior end of parietal suture and first black 
ring from 5^2 to S scales in width; body triads 7-9, black median ring 
of triad usually twice as broad as the black outer rings . . pijrrhocryptus 

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 locaUty as South America, 
but there is nothing to suggest this in the description or title of 
Cope's article. 

Range. Uruguay (where /ro7i/a/?;? is apparently unknown) . Outside 
of Uruguay, except where otherwise noted, it occurs alongside /ro/ito/w 
with which it may possibly hybridize. It also occurs in the Brasilian 
states of Rio Grande do Sul, Parana, Sao Paulo (probably also in 
Santa Catarina and southern IVIato Grosso). In Argentina it occurs 
in the Gobernacion 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^), Monte Carlo (C.N.H.M. 9357). Br.\sil: Parana: 
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 cfd', (206) 208-220 in 9 9 ; subcaudals 
(16) 19-25 in cfd^, (16) 18-23 in 9 9 ; also ventrals 215-220, sub- 
caudals 19-22 in two unsexed juveniles; body triads 13-17; total 
length: largest d" (M.C.Z. 20848) 918 (859 + 59) mm.; largest 9 
(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 althostris and 
frontalis, it has not reached a point where doubts are raised as to the 
assignment of any individual. For example, an altirostris 9 (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 a.ny frontalis ; ventrals 217. 

Only one frontalis, a Nueva Italia cf (M.C.Z. 47028) with 216 
ventrals has so low a count, possibly resulting from an admixture of 
altirostris "blood". Admixture with fro7italis is further suggested by a 
9 (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 d^ (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- 
cally 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 hybridization, 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 





5 


Sao Paulo 


2 


11 



4 BEEVIORA NO. 16 

Parana 5 2 

Misiones, Argentina; Villeta, Paraguay 6 1 

Rio Grande do Sul 8 2 

M. lemniscatns 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 (Dumeril, Dmneril and Bibron) 
Elaps Frontalis Dumeril, Dumeril 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 {sensu sirido) 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), Sao Paulo, Parana, 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 Gobernacion of 
Misiones. 

Material examined. Brasil: 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 Parana (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). Paraguay: Villeta: Colonia 
Nueva Italia (M.C.Z. 47028). 

> As M.C.Z. 3298 is only labelled Santa Cruz, Brasil, it ia uncertain whether it came from 
Bahia or one of the other states. 



1953 RACES OF MICRURUS FRONTALIS 5 

Ventrals 216-236 in c^ c^ , 221-242 in 9 9 ; subcaudals 20-26 in c^ d' , 
17-23 in 9 9 ; body triads 9-15; total length: largest d" (M.C.Z. 
27652) 1121 (1058 + 63) mm., largest 9 (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 oi 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 
scales in width, are about 4 or 5 scales in width in a Rio de Janeiro cf 
(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 baUocoryphus 
Cope (1859, p. 346) as a synonym oi frontalis frontalis. 

MiCRURUS FRONTALIS PYRRHOCRYPTUS (Cope) 

Flaps 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. Argentina: (A.M.N.H. 17592); Province of 
Mendoza (M.C.Z. 15911-2). Bolivia: Santa Cruz: Buenavista, 450 
meters (M.C.Z. 20622). 

Ventrals 224-232 in c^d", 228-(231) in 9 9 ; subcaudals 24-25 in 
d'd^, 24-? (tail truncated) in 9 9 ; 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 cT (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 BoHvian 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 Serie (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, Apranio do 

1944. Notas sobre a Ofiologia Neotropica e Brasilica. Pap. Avul. Dept. 
Zool., Sao Paulo, 5: 83-94. 

BOULENGER, G. A. 

1896. Catalogue of the Snakes in the British Museum. 3: xiv + 1-727, 
figs. 1-37, pis. 1-25. 

Cope, 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. 

Schmidt, K. P. 

1936. Preliminary Account of Coral Snakes of South America. Field 
Mus. Nat. Hist. Zool. Ser., 20: 189-203. 

Serie, Pedro 

1936. Nueva Enumeraci6n Sistematica y Distribuci6n Geogrdfica de 
los Ofidios Argentinos. Inst. Mus. Univ. Nac. La Plata, 1936: 
33-61, maps. 



E V I O R A 

MiiseiieTi of Coimparative Zoology 

Cambridge, Mass. May 28, 1953 Number 17 

A NEW 0RD()VI(7AX TRILOBITE 
FROM FLORIDA 

By H. B. Whittington 

bitroihirfioii. Tlie trilohite 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 Furope 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. Bridg-e. F. 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 (rreat Britain, 
is gratefully acknowleflged. 

Age. The Florida trilobite is here regarded as related to early 
representatives of the calymenid-homalonotid groups, in particular to 
species of Colpocorj/phr, but less closely to Placsiaconiia. 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, 1S49; Barrois, 1900), Portugal (Delgado, 1908), 
central Spain (\'erneuil and Barranrle, 1855), Morocco and /Vlgeria 
(Gigout, 1949; G. and H. Termier, 1950). PJarsiacomia occurs in 
younger strata, of Llandeilo age, in Bohemia and possibly Normandy 
(Prantl and Pfibyl, 1948). The Florida strata may therefore be of 
Llanvirn-Llandeilo age, i. e. in North American terms approximately 
late Canadian or early Champlainian. 



2 BREVIORA NO. 17 

Palcogcographical ImpIicatio7is. Evidence brought forward recently 
by Harrington and Kay (1951) and Wilson (1952) has served to 
strengthen the impression that the early Ordo\'ician faunas of South 
America, the Ouachita and Appalachian troughs, and northwestern 
Europe belong to the same general province. This Hkeness, 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 
Shrf'psliire 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, i.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 Flcvi- 
cah/mene and ''Brongniaridhr , 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 CALYMENTDAE IMilne Edwards, 1840. 
Genus Colpocoryphe Novak in Novak and Perner, 1918. 



H^53 A NEW ORDOVrCIAN TRILOBITE 3 

Tiipc species. Calymene arago Houuult, 1S49, from the ()r(lo\-ician 
(Llandeilo) of north-western France. 

Discussion. The outline of the ghil)ena, position of eyes, thorax of 
13 bluntly-terminated segments, and triangular pygidiimi combine 
to suggest that the P^lorida 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, PI. 3, fig. 30; see also 
Harrande, 1852, pp. 581-582, PI. 29, figs. 21, 22; 1872, p. 40, PI. 5, 
fig. 27; Prantl and Pfibyl, 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. ram is distinguished 
by the angulate outline of the pseudo-frontal glabellar lobe, the lack 
of ring furroM'S 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 Pfibyl (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 
Colpocori/phe Novak, 1918 {in Novak and Perner, 1918), and especially 
with C. inopinaia Novak (in Pompeckj, 1898, p. 249; see also Novak 
and Perner, 1918, p. 37, PI. 4, figs. 1-16; R. and E. Richter, 1927, pp. 
73-75, PI. 5, figs. 10, 1 1 ). The pygidium of Colpocoryphr 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, PI. 2, figs. 35, 39). The Florida 
trilol)ite has this type of pygidium. The cephalon, however, differs 
from that of C. arago (Barrande, 1872, PI. 2, figs. 34, 35; PI. 8, figs. 11, 
12) in that the eye is farther forwards and outwards, and the glal)ella 
lacks the deep glabellar furrows and well-defined lateral lobes. The 
cephalon of C. inopinaia 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 rata, has the distinctive 



4 BREYIORA NO. 17 

pygidium of Colpocori/phr, 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. 
PI. , figs. 1, 2. 

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 IS, R lOE., 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 distally. 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. x\nterior 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 preghil)ellar furrow and 1 .(> mm. out from the antero- 
lateral corner of the glabella. 

Thorax of 13 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 lilunt 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. Systeme Silurien du Centre de la Boheme. vol. 1. Prague and 

Parii^. 
1872. Systeme Silurien du Centre de la Boheme. Supplement au vol. 1. 
Prague and Paris. 
Barrois, C. 

1900. Bretagne, Livret-Guide, VHP Congres geolog. internat. 36 pp. 
Paris. 
Delgado, J. F. N. 

1908. Systeme Silurien du Portugal. Etude de Stratigraphie Paleonto- 
logique. Comm. Serv. Gool. Portugal, pp. 1-245, pis. 
GiGOUT, M. 

1949. Le synclinal silurien et devonien de Foucauld (Maroc occidental). 
C. R. Acad. Sci., Paris, vol. 228, no. 15, pp. 1303-1304. 
Harrington, H. J., and M. Kay 

1951. Cambrian and Ordovician Faunas of Eastern Colombia. J. 
Paleont., vol. 25, no. 5, pp. 655-668, pis. 96, 97. 
Hawle, I., and A. J. C. Corda 

1847. Prodrom einer Monographie der ixihmischen Trilohiten. Prague, 
pp. 1-176, pis. 1-7. 



6 BREVIORA NO. 17 

King, P. B. 

1950. Tectonic Framework of Southeastern United States. Hull. Am. 
Assoc. Petrol. Geol., vol. -ii, no. 4, pp. 635-671, figs. 
Nov.iK, O., and J. Per,\er 

1918. Die Trilobiten der Zone Ddi7, von Prague und Umgebung. 
Paleontogr. Bohemiae, Ceska Ak. C. Frant. Jos. pro v6dy etc., 
no. 9, pp. 1-51 (Czech and German text), pis. 1-4. 
Prantl, F., and A. Pribyl 

1948. Classification of the Bohemian Homalonotidae (Trilobitae). 
Bull. Int. de TAcad. tcheque des Sci., 46th yr., no. 9, 24 pp., 
pis. 1-2. 
Pom PECK J, J. F. 

1898. Uber Calymmene Brongniart. Neues Jahrl)., Min. Geol. Pal., 
1898 (vol.' I), pp. 187-250. 
RiCHTER, R. and E. 

1927. tJber zwei fiir das deutsche Ordovicium bedeutsame Triloliiten. 
Senckenbergiana, vol. 9, ))t. 2, pp. 64-82, ])1. 5. 

ROUAULT, M. 

1849. Memoire 1) sur la composition du test des Triloliites; 2) sur les 
changements de formes. . . . l-5ull. Soc. Geol. France, ser. 2, vol. 
6, pp. 67-89, pis. 1, 2. 
Stubblefield, C. J. 

1939. Some Aspects of the Distribution and Migration of Trilol)ites in 
the British Lower Palaeozoic Faunas. Geol. Mag., vol. 76, pp. 
49-72. 
Termier, G. and H. 

1950. Palcontologie Marocaine, vol. 2. Actual. Sci. Industr. no. 1095, 
Palis. 
Verneuil, E. de, and J. Barrande 

1855. Description des fossiles trouves dans les terrains silurien et 
dcvonien d'Almaden, d'une partie de la Sierra Morena et des 
montagnes de Tolede. Bull. Soc. Geol. France, ser. 2, vol. 12, 
pp. 964-1025, pis. 23-29. 
Wilson, J. L. 

1952. Cambrian and Ordovician Faunas from the Marathon Region. 
Guidebook, 1952 Spring Field Trip. W. Texas Geol. Soc, pp. 
62-64. 




y 



*a.*--i 



>' >^- 





#-« 



EXPI AXATIOX OF PLATE 1 

Citlpoccnjphr t.rsu' sp. nov. x 6. IT. S. Nat. Miis. Collections. 

Fig. 1. Huhher mould of holoty])('. The black spots on the left side of the 
cephalon are caused by minute holes in the mould. Fig. 2. Holotype, ex- 
ternal mould. 



E V I O R A 

Museum of Comparative Zoology 

Cambridge, Mass. September 23, 1953 Number 18 

CHARACTERS AND SYNONYMIES AMONG 
THE GENERA OF ANTS 

Part II 
By William 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 
mjTmecologists will spend a little of their time investigating and 
formally publishing new synonymy. 

'Part I: Breviora, Mus. Comp. Zcjol. 11: 1-13 (1953). 



2 BREVIORA NO. 18 

Anochetus Mayr 

Anochetus Mayr, 1861, Europ. Formicid., p. 53. Genotype: Odontomachus 
ghilianii Spinola, 1851, by designation of Bingliam, 1903. 

Myrrnapatetes Wheeler, 1929, Amer. Mus. Novit., 349: 6. Genotype: Myrrna- 
patetes filicornis Wheeler, 1929, loc. cit., fig. 3, male, monobasic. New 
synonymy. 

Wheeler described Mynnapatetes 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., 1398: 7, figs. 5-7, 

worker. 
Lepidopone Bernard, 1953 (1952), Mem. Inst. Fran(,-aise Afr. Noire, 19 (1): 

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 
{lamottei) 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 Asphinctopone. From the 
descriptions and figures of A. silvestrii 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 3 

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 lamottei, it would seem that the 
strong relationship of all tlu-ee 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. 



Hylomyrma Forel 

Pogonomyrmex {Hylomyrma) Forel, 1912, Mem. 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 

reitteri 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 spm-s 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 x\merica, 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 Emei-y 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 gocldii 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 reifferi (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 tjqjes 
concerned than is given in their mostly inadequate descriptions. 

I believe that Irogera, considered by Emery to be a subgenus of 
Rogeria, is either very close to or synonymous with Hyloimirma. The 
two Irogera species from the New World, /. procera Emery and 
7. 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. Irogera 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 thi'oughout 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 
Ochetomyrraicini. 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 GENER-A. 5 

Ochdomyrmcx Mayi- by means of the presently-employed character 
based on the form of the clypeus. Wasmannia wiUiamsoni 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 Maj-r, 1872, Ann. Mus. Civ. Stor. Nat. Genova, 2: 147. 

Genotype: Technomyrmex sh-enuus Mayr, 1872, by designation of Bingham, 

1903. 
Aphantolepis Wheeler, 1930, Psj'che, 37: 44. Genotype: Aphantolepis quadri- 

color Wheeler, 1930, loc. ciL, 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 figm-es, 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 Tech- 
nomyrmex species by its rounded propodeum and its rather smooth. 



6 BREVIORA NO. 18 

shining integument; it is ferrugineous yellow in <?olor, with blackish 
gaster, whereas T. 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: Chapmanclla negrosensis 
Wlieeler, 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 wi-ong about the last 
character; the type possesses minute spm-s, 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 Euprenolepis, 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. Euprenolepis 
negrosensis (Wheeler) is the new combination. 

Euprenolepis may as well be considered as an independent genus for 
the time being, though further investigation 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 geei Wheeler apparently does not belong to Eupreno- 



1953 CHARACTERS AND SYNONYMIES AMONG ANT GENERA 7 

Icpis, 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 gcci 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 P'orel, 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 L (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 
x\ustralian 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 lippulum and Adlerzia froggatti from Mono- 
viorium 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 MjTmicinae will 
finally be eliminated. 



E V I O R A 

Mnaseiim 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 limitefl 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 Ci/matoccras hilli (Shattuck), Ci/mafoceras 
loeblichi Miller and Harris, and Paracymatoceras tcxanum (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 Cifmatoceras and Para- 
ci/matoccras 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 Angulithcs. 
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, Cifmafoceras, Paraci/matoceras, 
Eutrcphoceras, HcminautUus, and Anguliihrs. 

The p\Tite 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. .) . 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 (pi. 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 XAUTILOIDS 3 

There are also mature and relatively large specimens of calcareous 
preservation of Cifinatoccras hilli, Poraci/nintocrras te.ranum, and Para- 
cymaioceras sp. indet. Hyatt (1894) described and figured juvenile 
specimens of Cyuiatocrras ehgans? (Paraci/mafoceras tc.vanum), C. dr- 
,'<longchampsianus, C. simplex'!, C. mdlatum. 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 Ci/maiocrras from the Gault 
of England in the British Museum of Natural History. 

Branco (1880) figured the first and second camerae of Xaufilus 
clementinus d'Orbigny ( = C!/matocera,'i) For Naidtlits cf. deslong- 
champsianus d'Orbigny { = Ciimaioceras) 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 fovirth 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 Ciinudoccras described here. 

Durham (194(), p. 432, pi. 63, figs. 3, 5, 7) described and figured a 
juvenile specimen of Honinaid'dus cthcringfoni 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 Ci/mn- 
toceras sp. figured on plate 1, figures 10-12. 

Nautiloids in the ( "retaceous of Texas are not abundant and repre- 
sentatives of only four genera, namely Ci/matoccras, Parnciimaioccras, 
Eiifrrphoceras, and Angidlthes are known. In the Washita group 
Ci/matoceras and Paraci/mafoceras have been known for a long time. 
The small immature specimens described here probably belong in 
Cymatoceras. Juvenile specimens of Paraeymatoceras of comparable 
size already have very sinuous sutures. The adorally projecting suture 
could possibly develop into the sinuated suture of Paraeymatoceras 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); (3) Grayson formation, 4.5 miles south of McGregor and 
100 yards east of the highway, McLennan County (locality 966 of 
Adkins, 1923, p. 52); 1}/^ miles south of Bosqueville on the Belmont 
place, McLennan County; and at Bosqueville, McLennan County. 

Reposifori/. B.E.G. 297 (pi. 1, figs. 7-9), 3057 (pi. I, figs. 4-6), 3002 
(pi. 1, figs. 10-12; pi. 2, figs. 1, 2), 18750 (pi. 1, fgs. 1-3), 3273 (pi. 2, 
fig. 3J. 

Cymatoceras hilli (Shattuck) 

This species has recently been described and figured l)y 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 ()5 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-l; and along east side 
of ]\Iansfield Road, halfway between Glen Garden Country Club and 
Sycamore Creek, 3 miles southeast of Fort Worth, Tarrant ( Ounty. 
B.E.G. loc. 219-T-l 7 (locality 723 of Adkins, 1918, p. 51). 

Repo.^hory. 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, anfl 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 ril)s that are slightly sigmoichil 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 l)ut is most 
distinct at the adoral part of the specimen. 

Paraci/mntoceras differs from Ci/niatocrras 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 Paracymatoccras, 
namely P. tcvanum (Shumard), has been recorded from North America. 
That species is not uncommon in the Washita group of Texas. Para- 
cymatoccras sp. indet. 1, differs from P. fi.vanuni primarily in its very 
subdued ribbing. It is undoubtedly an immature specimen which 
makes direct comparison difficult. Most of the specimens of P. tcxanum 
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 ribl)ing in the ( "ymatoceratidae can be very subdued. The suture 
of Paracymatoccras- is very similar to that of Hcrcoglossa of the late 
Cretaceous and early Cenozoic. Pscudaganidcs 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 Paracymatoccras at this time; these are: P. aspcr 
(Oppel), the type species from Tithonian formations of Europe, P. 
trichinopolitcnsis (Blanford) from the Ariyalur group (Senonian) and 
P. rola (Blanford) from the Uttattur group (Albian) of .south India, 
P. texanum (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 Hcrcoglossa. As more Cretaceous nautiloids are dis- 
covered it will become increasingly difficult to differentiate those 
essentially smooth or weakly ribbed species of Paracymatoccras 
from Hcrcoglossa. 

Occurrence. Pawpaw formation, from a pit of the Cobb brick yard, 
}4: mile east of Sycamore Creek and 3 miles southeast of Eort Worth, 
Tarrant County (locality 716 of Adkins, 1918, p. 47). 

Repository. B.E.G. 196 (pi. 2, figs. 6, 7). 



b BKEVIORA 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. 




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 Paraci/matoceras sp. indet. 2 is associated 
with P. sp. indet. 1 and P. ff.ranum, it is highly probable that P. sp. 
indet. 2 is a juvenile smooth specimen of Paraci/matoceras. 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 



1953 TEXAS LOWER CRETACEOUS NAUTILOID.S 7 

specimens for Xautilus schiwidi (pi. 1, fig. 5a, b, c), N. ODimoni (pi. 3, 
figs. 5a, b, c), N. strambcrgcnsis (pi. 5, figs. 6a, b), and X. sp. (pi. 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 Xautilus schut'idi (pi. 1, fig. 5c) and iV. ammoni (pi. 3, 
figs. 4a, b). 

Occurrence. Pawpaw formation, one-fourth mile south of the Inter- 
national and Great Northern railway bridge across Sycamore Creek, 
4}/2 miles southeast of Fort Worth, Tarrant County, B.E.G. loc. 
219-T-23. 

Repository. B.E.G. 17425 (pi. 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 
Paraci/matoceras descril)ed here. The specimen is from the pyrite 
fauna of the Pawpaw formation. The whorls measure 6 mm. in height 
and S mm. in w'dth. The venter is broadly roun led, flanks convex, 
and the umbilical shoulders are sharply rounded. The suture has a 
broad ventral saddle and a deep rounded lateral lobe followed by a small- 
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 Paraet/niaioceras 
in having a more subquadrate outline. As with P. sp. indet. 2, this 
fragmentary specimen is thought to l)e a portion of a juvenile indi- 
vidual. ^^ hether or not it is conspecific with the other two specimens 
of Paraci/matoceras 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. 

Repositorii. B.E.G. 17415 (pi. 2, figs. 4, 5). 



S bkp:vioha 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 Hving chamber. The fianks 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 lol^e 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 eross section of Angulithes sp. indet., X 1. 

The genus Angvliihis Montfort (type species Nauiilites 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. tri- 
angularis is alternately rounded and sharply angular. There are only 
five species of Cretaceous nautiloids that appear to belong in the genus 
Angulithrs; 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. tvcstphalicus (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 tp:xas lower cretackois xaitiloids 9 

to warrant a specific name hut the record of a species of the genus 
AiKjulUhcs in North America is of interest. 

Dclfoidcnaufilus — type species Xaidilus soicrrhi/i W'ethereW — was 
proposed hy Spath (1927) for Eocene nautiloids with nautihconic 
conchs and more or less sagitate cross sections with narrowly rounded 
to angular ventral zones. The sutures form narrowly rounded ventral 
saddles, rounded lateral lohes, and a small saddle near the umbilical 
wall. The siphuncle is located near the dorsum. Spath pointed out 
the similarity of whorl shape of DcltoidonauiUus to Angullfhrs; how- 
ever, he thought that Angidithcs "... with its less sinuous suture-line 
was more closely allied, via Ay^g. flvuriousianus (d'Orhigny) with the 
regular Xautiloid stock that produced Psntdocrnoceras in the Creta- 
ceous, as it had given rise before to the less specialized Paracrnoceras 
of the raUoricnsls type." 

Examination of the various species of Angidithcs from the pAiropean 
Cretaceous and the specimen from the Pawpaw formation described 
here shows no appreciable differences between DcltoidonautUus and 
Angvlit/us. At the time Spath proposed the genus Dcltoidotifntfilus it 
was known only from the Eocene. The range of DcUoidnnauiUus is 
now known to be Paleocene to Oligocene (Miller, 1951). The few 
species of AnguUlhcs range well throughout the Cretaceous. Because 
of the similarity of Di'Jtoidonautilus and AnguJifhes the genus Deltoido- 
naufilus should be suppressed and its species placed in the genus 
Angiilithis which has priority. 

Occurnncc. Pawpaw formation, at hillside on north side of east-west 
road, .3 miles southeast of Haslet, Tarrant ( ounty, B.E.G. loc. 219-T-S 
(locality 724 of Adkins, 1918, p. 51). 

Rcpositori/. B.E.G. 17421. 



10 BREVIORA NO. 19 

REFERENCES 

Adkins, W. 8. 

1918. The Weno and Pawpaw formations of the Texas Conianchean. 

Texas Univ. Bull. 1856, pp. 1-172, pis. 1-11 (1920). 
1923. Geology and mineral resources of McLennan County. Texas Univ. 
Bull. 2340, pp. 1-202, pis. 1-4 (1924). 
Blanford, H. F., and Ferd. Stoliczka 

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, etc. (Ferd. Stoliczka). 
India Geol. Survey Mem. (Palaeontologia Indica), ser. 1 and 2, 
vol. 1, pp. 1-216, pis. 1-94. 
BosE, Emil 

1927. Cretaceous ammonites from Texas and northern Mexico. Texas 
Univ. Bull. 274S, pp. 143-312, pis. 1-18 (1928). 
Branco, W. 

1880. Beitrage zur Entwickelungsgeschichte der fossilen Cephalopoden, 
Theil II, Die Goniatiten, Cl.vmenien, Nautiliden, Belemnitiden 
und Spiruliden. Palaeontographica, Bd. 27, pp. 17-81, pis. 4-11. 
Durham, J. Wyatt 

1946. Upper Aptian Xautiloids from Colombia. Jour. Paleontology, 
vol. 20, pp. 428-434, pis. 62-64. 
Foord, 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. 
Hyatt, A. 

1894. The phylogeny of an actjuired characteristic. Am. Philos. Soc. 
Proc, vol. 32, no. 143, pp. 349-647, pis. 1-14. 

KUMMEL, B. 

1948. Environmental significance of dwarfed cephalopods. Jour. Sedi- 
mentary Petrology, vol. 18, pp. 61-64. 
LOESCH, K. C. 

1914. Die Nautilen des weissen Jura, erster Teil. Palaeontographica, 
Bd. 61, pp. 57-146, pis. 10-15 (1-6). 
Miller, A. K. 

1951. Tertiary nautiloids of West-Coa.?tal Africa. Annales du Musee du 
Congo Beige Terruren (Belgique), ser. 8, Sci. Geol., vol. 8, pp. 
1-88, pis. 1-31. 



1953 TEXAS LOWER CRETACEOUS NAUTILOIDS 11 
, and R. A. Harris 



1945. North American Cymatoceratidae (Mesozoic Xautiloidea). Jour. 
Paleontology, vol. 19, pp. 1-13, pis. 1-6. 
»ScoTT, Gayle 

1924. Some gerontic ammonites of the Duck C^reek formation. Te.xas 
Christian Univ. Quart., vol. 1, no. 1, pp. 4-31, pis. 1-9. 

1940. Paleoecological factors of Cretaceous ammonoids. Jour. Pale- 
ontology, vol. 14, pp. 299-323, figs. 1-9. 
8path, 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-S4, pis. 1-7. 

Si'ENGLER, F]RICH 

1910. Untersuchungen iiber die sudindische Kreideformation. Die 
Nautiliden und Belemniten des Trichinopolydistrikts. Beitr. 
Paliiontologie Osterreich-Ungarns u. des Orients, Bd. 23, pp. 
125-157, pis. 11-14 (26-29). 

WlXTON, W. M. 

1925. The geology of Denton County. Texas Univ. Bull. 2544, pp. 1-86, 
pis. 1-27. 



EXPLANATION OF PLATE 1 

Figs. 1-12. Cymatoceras sp. indet. all figures X2. 1-3, B.E.G. 18750, from 
Orayson 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. 




i» 



■^ J 









m 



^ 10 




11 






PLATE 1 



fJXPLAXATIOX OF PLATP] 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. 3, 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, XI. 

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, XI . 





\4 





•<: 



10 



PLATE 2 



E V I O R A 

Miasenim of Comparative Zoology 

Cambridge, Mass, September 23, 1953 Number 20 

LOWER TRIASSK^ SALT RANGE 
XAUTILOIDS 

By Bernhard Kummel 

INTRODUCTION 

Triassic nautiloid.s 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 S^Tingonaiitilidae 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 Scythian, 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: 
Mctacocrras (Mojsvaroccras) subaratuni (Keyserling) 1860, from 
northern Siberia; Plrurouautilvs dicncri Krafft and Diener. 1909, 
Griipociras {(Irypoceras) hrahinanicum (Griesbach) 1880, Grypoccras 
{Grypocrraa) hi.vagonalis Diener, 1897, Grypoccras {Grypoccras) 
lilangense Krafft and Diener, 1909, from the Himalayas; Plcuronautiliis 
suhquadrancjulus Tien, 1933, from south China; Plcuronautilus kokeni 
Freeh, 1905, Tainionautilus trachyccras Freeh, 1905, Mcnuthionautilus 
kicsJiugiri Collignon, 1933, Grypoccras (Grypoccras) bidorsatoldcs n. sp., 



2 BREVIORA NO. 20 

Grypocvras {Grypociras) acmulantf n. sp. from the Salt Range, Pakistan. 
Menuthionmdilus kiislingcri was originally described from Scythian 
strata of northern Madagascar (("ollignon, 1933). The Lower Triassic 
Dinwoody and Thaynes formations of Idaho have yielded new species 
of Mojsvaroccras, Plcuronautilus, Knoploccras, Germanonaid'lus, and 
urypoccras. The descriptions of these new species are in press (Kum- 
mel, 1953). MuUer and Ferguson (1939) list Grypoccras cf. brahmani- 
cum (Griesbach) from the Candelaria formation of Nevada. 

In the Lethaea Geognostica (Freeh, 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 Tainioncmtilus trachyceras from the C'eratite Sandstone 
(zone of Flemingites flcmingianus) and PlevronaidUus kokcni from the 
Upper C'eratite Marls (zone of Konhickiics volidus). 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 (F'rech, 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 Phuronaidilus (B.M.N.H. 
C10429 and ("10426) 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. Y. 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. Xatdihis mesodicum Hauer, 1846. 

There are approximately 22 species of Upper Paleozoic nautiloids 
assigned to Domafoceras (type species, Domedoccras umbilicatum 
Hyatt, 1891). They are characterized by being flattened ventrally and 
laterally and slightly impressed dorsally. The sutures form rounded 



1958 I.OWKR TKIASSIC SALT RANGE NATTILOIDS 3 

ventral, latenil, and dorsal lobes. Tlie siphuncle is subcentral in posi- 
tion anfl orthochoanitic. Species of Dhiinitoccras show much variation 
in the degree of involution and in the position of the siphuncle. 

The Triassic genus (iriipoccrdii has long been recognized as very 
similar to late Paleozoic domatoceratids (Mojsisovics 1902, p. 227). 
Gri/poccra.s' differs from Doniafoccras 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 Doinatocrras 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. klcihegei (17%) and D. moorci (16%), are 
transitional forms to Stniopocrras. The same data on 9 species of 
Gri/poccras 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 (Jrypoccras have angular ventral 
shoulders, the Middle and Upper Triassic species generally have more 
rounded shoulders. These data suggest that Gri/poccras is a direct 
evolutionary continuation of the late Paleozoic Domaiocrms, and the 
two genera are gradational in most characters. The principal differ- 
ences are in the presence of an annular lobe anfl the greater involution 
of Gri/poccras. Under these circumstances full generic status for 
Domaioccras does not seem warranted and it is here placed as a sub- 
genus of Grypoceras since the latter genus has priority. 

Grypoccras is widely distributed geographically, being more or le.ss 
common in the Alpine and Himalayan Triassic regions. It is also 
reported in Triassic rocks from North America, the I^ssuri 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. 
mcsodicnm 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 roimded 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 l)road, 
shallow, ventral and lateral lobe. The siphuncle is in a subventral 
position, being only 4 mm. from the venter at a point where the whnrl 
height is 22 mm. 




Fig. 1. Diagrammatic ci'()i^s-se<-tion of Grj/poccras {Grupocer as) hidorsa'.oides 
II. sp., XI. 

Remarks. G. {(Jri/poccra.s) hidoi;satoidis is closely related to G. 
(Grypoceras) brahmanicum of the Otocrras beds of the Himalayas. The 
Salt Range species is more involute and more depressed in cross section. 
The siphuncle of G. (Grypoccras) 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) Ulangcnsc from the "Mceko- 
ceras" beds of the Himalayas has a very rapidly expanding and large 
conch, and no comparison can be made with G. {Grypoccras) bidorsa- 
toides. The Dinwoody formation in Idaho has yielded a new species of 
Grypoccras that is somewhat similar to this Salt Range species but 
more compressed and with a high whorl section. Except for greater 
involution, G. (Grypoccras) bidorsaioidcs is similar to several species of 
upper Paleozoic domatoceratids. 

Occurrence. Lower ('eratite Marl, zone of Prii^nolobiis roivndatns, 
Chideru, Salt Range, Pakistan. 

Repository. B.M.X.H. ('10428, holotype. 



1953 LOWER TRIASSIC SALT RANGE NAUTILOIDS 5 

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. Tlie conch measures (K) mm. in diameter; the most 
adoral part of the last whorl measures 80 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 
n. sp., XI. 

Remarks. Grypoceras {Gri/poceras) aernulans differs from G. (G.) 
hidorsatoidrs in being more compressed and in having an arched venter, 
not concave on any part of the conch. G. (6'.) 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.) hidorsatoides is similar to late 
Paleozoic domatoceratids except for the greater involution of the 
conch. 



6 BREVIORA NO. 20 

Occurrence. Lower Ceratite Marl, zone of Prionolohus rofundatus, 
Chideru, Salt Range, Pakistan. 

Repository. B.M.N.H. C47068, holotype. 

Genus MenUTHIONAUTILUS Collignon, 1933 
Type species. Nautilus {Menuihionauiilus) kieslingcri Collignon, 1933 

MenUTHIONAUTILUS kieslingeri Collignon 
Plate 1 ; figures 3, 4; plate 2, figures 1-3 

Nautilus {M etiuthionautilus) kieslingeri C-ollignon, 1933, Annales de P;ile(jnto- 
logie, tome 22, pp. 164-166, pi. 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. Diagiammutic cross-section and repre.sentation of an extein: 
suture of Afenuthionautilus kieslingeri, XI. 



195.3 LOWER TRIASSIC SALT RANGE NAUTILOIDS 7 

The whorls are much higher than wide. The hirgest specimen (pi. 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 (pi. 2, 
fig. 3), 40 mm. in diameter, the umbilicus measures (i 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 
l)road, 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. CoUignon'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. 

("ollignon (1933) established Menuthionautilus as a subgenus of 
Xautilus and he recognized the uniqueness of this Triassic nautiloid 
with its ventral siphuncle. He compared his species with Paranautiliis 
and Gri/poccras. There appears to be little actual affinity with Para- 
na iifil us. The Paranautilidae are a stock that tended toward involute, 
globular conchs and simplified septa. There are very few compressed 
species of Paranautilus, IndonauiUus, or SlhiillotKiiifihis. The Para- 
nautilidae stem from the upper Paleozoic Liroccraiidac and themselves 
gave rise to the (li/donaidilidac of the Upper Triassic with their very 
sinuous sutures. 

The inner whorls of Mrnuthionautilus are more quadratic in cross 
section, having low arched venters and sharply rounded ventral 
shoulders. The characters of the conch of Mcnuihionmiiiliis show- 
clearly affinities to Gri/poccras of the Triassic and Griiporcras (Doinato- 
ceras) of the Pennsylvanian and Permian. The Grypoceratidae include 
the following genera: Gri/poccras, Gri/poccras (Domcdoccras), Sicnopo- 
ccras, Stcaroccras, Tifanoccras, Menuthionardilus, 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 



8 BREVIORA XO. 20 

various species. In an analysis of 10 species of (i. (l)otu(iiocvras) 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 Grypuccras range from 30 to 50 per cent. As mentioned 
above under the discussion of Gri/poccras and G. (Domatoccras) these 
two forms are believed to belong to a continuous evolutionary series. 
Mrnufhionaufiius is most likely a development out of this Domaioceras- 
Griipoccras line. The similarity of the whorl section, general plan of 
the suture, and known \'ariability of the siphuncle in the parent stock 
strongly support this relationship. 

Occvrrencc. Lower ("eratite Marl, zone of PrionoJohvs rotundnfus, 
Chideru, Salt Range, Pakistan. 

Reposiiory. B.M.N.H. C104.3.3 (pi. 2, fig. 1), ('10438a (pi. 1, figs. 
3, 4), C10438b (pi. 2, fig. 3), ('10438c (pi. 2. fig. 2). 



REFERKXCE8 

CoLMGNON, Maurice 

1933. Les Cephalopodes du Tiias Inferieur. Paleontologie de Madagas- 
car, XX. Annales de Paleontologie, tome 22, pp. 151-180, pis. 
14-20 (1-7). 

Frech, Fritz 

1903-1908. Lethaea Geognostica, Teil 2, Das Mesozoicum, Band 1, 
Trias, pp. 1-623, pis. 1-68. Stuttgart. 

Mo.Jsisovirs, Edmund von 

1873, 1875, 1902. Das Gebirge um Hallstatt. — 1. Abt. Die Cephalo- 
poden der hallstatter Kalke. Abb. geol. Reichsanst. Wien. Band 6, 
Heft 1, 2; Band 1, Suppl. - Heft, pp. 1-356, pis. 1-70 and 1-23. 
(Heft 1, pp. 1-82, pis. 1-32, 1873; Heft 2, pp. 83-174, pis. 33-70, 
1875; Suppl. -H3ft, pp. 175-356, pis. 1-23, 1902.) 

Mumper, S. W. and H. G. Ferguson 

1 939. Mesozoic stratigraphy- of the Hawthorne and Tonopah Quadrangles, 
Nevada. Geol. Soc. Am. Bull., vol. 50, pp. 1573-1624. 



EXPLANATION OF PLATE I 

The specimens illustrated on this plate are from the Lower C'eratite Marl, zone 
of Prionoluhus rotiimlatus, Chideru, Salt Range, Pakistan. The>' are deposited 
in the British Museum of Natural History, London. All figures X L 

Figs. 1, 2. Gri/poceras (Gri/poccras) t>idorsatoidcs n. s])., B.M.N.H. — 
C10428, holotype. 

Figs. 3, 4. Menuthionautilus kieslingeri Collignon. Immature specimen; 
hgure 3 shows clearly the ventral position of the siphuncle. B.M.N.H. — 
CI 0438a, i)aratype. 




i^^ 





-■ '^h 




platp: 1 



EXPLANATION OF PLATE 2 

The specimens illustrated on this plate are from the Lower Ceratite Marl, 
zone of Prionolohus rotundatus, Chideru, Salt Range, Pakistan. They are 
deposited in the British Museum of Natural History, London. All figures X 1. 

Figs. 1-3. MenuthionautUus kieslingeri Collignon. I. 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. Cl043Sc. 3. Side view of 
small paratype, B.M.N.H. C10438h. 

Fig. 4. Grypoceras {(iri/poceras) aenndans n. sp. B.M.N.H. C47068, holo- 
type. 




PLATE 2 



E V I O R A 

Mesemm of Comparative Zoology 

Cambridge, Mass. September 23, 1953 Number 21 

THE ANCESTRY OF THE FAMILY 
NAUTILIDAE 

By Bernhard Kummel 

The evolutionary history of the ammonoids and nautiloids is ((uite 
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 
hut 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 anfl generally even greater radiations. The transition 
from the Permian to the Triassic witnessed wholesale extinctions 
among the ammonoids with only the Xenodiscidae 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 
."i families and 2S genera, many of which were established by him. 
On the problem of origin of the.se post-Triassic nautiloids Spath (1927. 
p. 23) discussed ''G'ri/pocrras cf. mi:so(licum'" (Hauer) recorded by 
Trechmann (191.S, p. 181) from the Upper Triassic (Carnian, bed C) 
of the Hokonui Hills, New Zealand. Spath considered this species a 



2 BREVIORA NO. 21 

Bisiphytcs and stated that "... there is more prol)ability of BIsiphytes 
being an invokite, glol)o.«e, development of the family Syringo- 
nautilidae Mojsisovics, 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 (Xatin-al History) and for 
permission to study Trechmann's specimen. The photographs were 
taken in the British Museum l)y its staff photographer. 

The uniqueness of this species (here named Bisipluitis trcchmanni 
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 Gri/poceras and goes on to mention it 
as ". . . indistinguishable from typical BIsiphi/tcs, 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 trenfls which began in the Carboniferous. 
Three distinct lines of development are represented. One includes the 
Paranautilidae, C'lydonautilidae, 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 involute, 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 Gryponaufihis, are completely unlike Bisiphytcs trcchmanni. 
Within the Syringonautilidae there are forms which are very similar 
in conch patterns to Bisiphytcs trcchmanni. Syringoccras 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 



CO 

en 
< 


d 


&i 




y 

ID 
< 


5 

z 

c 
5 

c 
.9 

c 

c 
.2 
<n 

1 

u 






1 
1 


Bisi 
tr 


1 

• 

Dhy 
ech 


t-es 
manni 


1 

8 




1 

1 


- 


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■1 



Fig. 1. Phylogenetic diagram illustrating the relationship.s of the Liassic 
Xautilidae 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 Xorian are not 
known. 



umbilical walls. The shell bears fine radial and longitudinal lines. 
These two genera differ only in the position of the siphuncle; Si/ringo- 
crras has the siphuncle very near but not at the venter, Syringonaiiiilus 
has its siphuncle in a more central position. There are about 14 
species of Si/ringonautilus, with 9 species in the Anisian, 5 in the 
Ladinian, 3 in the Carnian, and 1 indeterminate species in the Norian 
(Some of these species occur in more than one epoch.) OF Si/ringorrrafi 



4 BREVIORA NO. 21 

there are 17 species recorded, 1 from the Anisian, 3 from the Ladinian 
1? from the Carnian, and 1 from the Norian. The pattern of develop- 
ment of these two genera is markedly different in that Si/ringonautilu! 
had a large radiation soon after its appearance and then declined 
rapidly; Syritiguccras 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 ClymcnonautUus, Juvavionautilius, and Oxi/nautilus. 
Clymcnonautilus has a conch pattern like that of Si/ringonauHlus, but 
has a very sinuous suture convergent to the type of suture present in 
the C'lydonautilidae. Juvavionauiilus, in its conch shape, is a hetero- 
chronus homeomorph of Domatoccras of the Pennsylvanian and Per- 
mian. O.vytiautilus is an involute o.xycone similar in its conch pattern 
to St('7iopoccras of the Pennsylvanian and Permian. Cli/mcnonautilus 
and Oxyncndilus are monotypic and only 6 species are known of 
J uvavion(ndilu,s . 

On the basis of the prominent peripheral strigations this species is 
placed in Bisiphyfcs and not in Syringocrras or Syringonaufilus. The 
latter two genera are more evolute than Bisiphyfcs, the umbilicus 
approximating 25 to 30 per cent of the diameter of the conch. The 
umbilicus of B. trcchmanni is about 19 per cent of the diameter of the 
conch. Among typical Liassic Bisiphyfcs the width of the umbilicus 
ranges from around 20 per cent to completely occluded conches as in 
B. simiU'imns Foord and ( "rick. 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 Domatoccras-Crypoccras line in the Grypo- 
ceratidae and the Mctacoccras-Mojsmroccras line in the Taino- 
ceratidae. The shape of the whorl section and suture of B. frcchnianni 
is quite similar to such species of Syringoccras as S. crcdneri Mojsiso- 
vics, S. I'vohdus Mojsisovics or species of Syringonautilus as S. lilianus 
(Mojsisovics), and N. longobardicvs (Mojsisovics). 

Early Jurassic Nautilidae are accomodated in 5 genera including 
Bisiphyfcs INIontfort, 18G8, Cenoccras Hyatt, 1883, Sphacronaidilus 
Spath, 1927, Digonioceras Hyatt, 1894, Ophionautihis Spath, 1927. 
Sphacronaidilus is monotypic, Digonioceras has 4 species, and Ophio- 
uaufilus only 5 species. The remaining species of early Jurassic 
Nautilidae belong to Bisiphytcs and Cenoccras. The Paraceno- 
ceratidae, Hercoglossidae, and Cymatoceratidae are derived from the 



1953 ANCESTRY OF THE FAMILY NAUTILIDAE 5 

Nautilidae. 

Liassic nautiloids liave received rather thorougli treatment by 
Prinz {19()()) 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 prol)ably originally roinided without angular 

shoulders (perhaps broader than high). 
2.) The primitive sculpture consisted of radial and longitudinal 
striae of ecjual strength. A smooth shell is a secondary develop- 
ment. 
'.].) 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. 
().) The umbilicus was originally open. 

Pia considered Nautilus .striafu.s as being morphologically ^'ery 
similar to the primitive original stock of Liassic nautiloids. 

Of the known I'pper Triassic nautiloid stocks, only the Syringo- 
nuufilidar 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. Bi.siphi/fi's is tlius considered a late Triassic oft'-shoot 
of the Syringonautilidae, as first suggested by Spath (1927, p. 23), 
and in the (Hrect line of ancestry of all later nautiloids. 

SYSTEMATIC ' DESCRIPTION 

Eamily NAUTILIDAE d'Orbigny, 1S4(3 

Genus BiSIPHYTES Montfort, 1S08 

BiSIPHYTES TRECHMANNI n. sp. 

Plate 1, figures 1,2,)); text figure 2 

dry pore ras cf. nicfiodiciau (Hau"r). Ti'i'cliinaiin. (^)uai1. .loui'. Gcol. Soc. 
Loiulon, vol. 73, i)p. 1S1-1S2, l'.»!s. 

The single specimen upon which this species is based is a moderately 
large phragmocone. It measures 71 mm. in diameter, 3S 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. Di;igrummatir cross section of Bisiphyles 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, ("arnian, IJed (", Otamita, Hokonui 
Hills, New Zealand. 

Repository. British Museum (Natural History) (' 21947. 



1953 ANCESTRY OF THE FAMILY NAVTILIDAE 7 

rp:ferexce8 

PlA, JULU s 

1914. rntersuchungen iiber die liassischen Xautiloidca. Bcitr. Pala- 
oiitologie Osterreich-Ungarns u. des Orients, Bd. 27, pp. 19-86, 
pis. 4-10. 

Prinz, Jrurs 

1906. Die Xautiliden in dcr uiitcreii Jura-Periode. Ann. Hist.-Xat. 
Mui^. Xat. Hung., vol. 4, pp. 201-243, pis. 3, 4. 

SPATH, L. F. 

1927. Revision of the Jura.'^.sic cephalopod fauna of Kachh (C'utch). 

India Geo). Survey Mem. (Palaeontologia Indira), n. ser., vol. 9, 

mem. 2, pp. 1-84, pis. 1-7. 
1934. Catalogue of the fossil cephalopods in the British Museum 

(Xatural History), Part 4, The Ammonoidea of the Trias (I). 

pp. 1-521, pis. 1-18. 

Trkchmaw, C. T. 

1918. The Trias of Xew Zealand, (^uart. .lour. Geol. Soe. London, vol. 
73, pp. 165-246, pis. 17-25. 




a . 

O 05 

CQ 5 




^ .2 ffi 
H O 5 

hJ .=i ^ 










E V I O R A 

MmseiiijiM of Coioipsirative Zoology 

Cambridge, Mass. October 13, 1953 Number 22 

NOTES ON THE ANT, 
LEPTOTHORAX OBLIQUICANTHUS COLE 

(Hymenoptera : Formicidae) 

By Robert E. Gregg 

Department of Biology, University of Colorado 

A new species of Lepfothorax 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 Lepfothorax. 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 Lepfothorax, it is thought advisable to publish 
it herewith. 

As Dr. Cole points out, the ant differs from the closely related 
members of the tricarinafus-fexanus complex, and particularly from 
L. t. ncomexicanus in its shorter scapes, larger epinotal spines, and 
differently shaped petiole, etc. 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 ohliquicanthus, 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) ohliquicanthus. 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. AVe 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 Leptothorax, thus: 

Leptothorax (Leptothorax) obliquicanthus Cole 

The specimens of ohliquicanthus 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 obi i qui cant hus 
raises the question of their possible function and adaptive value. 
Dr. Creighton (1930, p. 121), in discussing a similar phenomenon in 
Solcnopsis (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 

Cole, A. C. 1953. Notes on the genus Leptothorax in New Mexico and a 

description of a new species. Proc. Ent. Soc. Washington, 55: 27-30. 
Creighton, W. S. 1930. The New World species of the genus Solenopsis. 

Proc. Amer. Acad. Arts Sci., 66: 39-151. 
Creighton, W. S. and R. E. Gregg, 1953. An appeal to the International 

Commission on Zoological Nomenclature for the conservation of certain 

generic and subgeneric names of ants. MS. 
Smith, M. R. 1950. On the status of Leptothorax Mayr and some of its 

subgenera. Psyche, 57: 29-30. 



R E V I O R A 

Museiuim of Comparative Zoology 

Cambridge, Mass. October 23, 1953 Number 23 

A NEW SPECIES OF HYPOGNATHA 
FROM PANAMA 

By Arthur 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. nasiita 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. 



NO. 23 



Genus HypoGNATHA Guerin, 1839 
Hypognatha elaborata sp. nov. 
(Figures 1-10) 
Male holoiyi)c. 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 



> 






External Anatomy of Hypognatha elaborata sp. nov. 
Figures 1-5 
Fig. 1. Eyes, clypeus, and cephalic horns of male, from in front. 
Fig. 2. Median cephalic horn of male in profile. 
Fig. 3. Lateral cephalic horn and LE of male. 
Fig. 4. Left male palp, ventro-lateral view. 
Fig. 5. 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 sHghtly 
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. 

Maxillae, 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. 
2. 
3. 
4. 
Palp 

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 middleof the 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 0. 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. 



.975 


.487 


.769 


.975 


.430 


3.636 


.968 


.462 


.682 


.858 


.412 


3.382 


.682 


.252 


.396 


.418 


.308 


2.056 


.902 


.330 


.550 


.770 


.363 


2.915 


.352 


.176 


.240 




.572 


1.340 



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) . 

Maxillae. 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 stiflf 
bristles (Fig. 7). 



1953 



A NEW SPECIES OF HYPOGNATHA 




External Anatomy of Hypognatha elahorata sp. nov. 

Figures 6-10 

Fig. 6. Eyes and clypeus of female allotype, from in front. 

Fig. 7. Sternum of female allotype. 

Fig. 8. Female allotype, dorsal view. 

Fig. 9. Epigynum of allotype, posterior view. 

Fig. 10. Epigynum of disvsected paratype to show complete posterior exposur 



Tarsi Totals 



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. 

Femora Patellae Tibiae Metatarsi 

(All measurements in millimeters) 

.520 .715 .921 

. .520 .650 .814 

.380 .418 .418 

.440 .616 .780 



1. 


.975 


2 


.975 


3. 


.812 


4. 


1.100 



.440 


3.571 


.418 


3.377 


.352 


2.380 


.400 


3.336 



6 BREVIORA NO. 23 

Spines. True spines almost completely lacking. A few weak spines 
may be recorded as follows: First leg: femm- 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. 

Ahdomcyi. 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 difl^erent 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 pro|jided 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. 

Hypognatha nasuta O. p. Cambridge, 1896 

(Figures 11-15) 

H. nasutn 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 



1953 



A NEW SPECIKS OF HYPOGNATHA 



horns are relatively much broader than in H. clahorata 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. clahorata 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 H ypognatha nasuta O. P. Cambi-idj. 
Figures 11-15 



Fig. 1 1 . Eyes, clypeus, and cephalic horns of male. 

Fig. 12. Median cephalic horn of male in profile. 

Fig. 13. 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. Cambridge. 1889-1905. Arachnida-Araneida. 

Vols. I-II. In: Biologia Centrali-Americana. Dulau & Co , London. 
Simon, Eugene. 1892-1903. Histoire Naturelle des Araignees. Deuxieme 

Edition. 2 Vols. Libraire Encyclopedique 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. 



E V I 

Museiiiiin of Compsirative Zoology 

Cambridge, Mass. January 27, 1954 Number 24 



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^ 

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 x\LBATROSS 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. sinua- 
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 Anaeantho- 
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 PSEUDORAJIDAE, 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 S-pringcria Bigelow and Schroeder 
1951, and perhaps in the skate Psammobatis m?Va(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 Gannan (Proc. Boston Soc. Nat. Hist.. Vol. 19, 1S77, p. 207) described the snout of mira 
as terminating in an "acute point", which is pictured as pricklv in his classic monograph 
(Mem. Mus. Comp. Zool., Vol. 36, 1913, PI. 27, Figs. 3-.^)). 



1954 NEW FAMILY, GENUS AND SPECIES OF BATOID FISHES 3 

characters that are not as strictly alternative as seemed once to be the 
case. 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- 
hatis, Actobatus, Rhinopiera 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 Myliobafis, in Dasyatis (referred to by him as Trygon), or 
in Taejiiura. 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 Garinan, 1913, PI. 27, Fig. 1. 

- 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. 

' See Garman (1913, Pis. 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 ofr 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 elasinobranchs. 
s From base of rostral filament. 



1954 NEW FAMILY, GENUS AND SPECIES OF BATOID FISHES 5 




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, 1st 1.0, 1.1; 3rd 1.2, 1.2; 5th 0.7, 0.7; distance 
between inner ends, 1st 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 . , ^ 

m male of 262 mm. 



Teeth: ^ in type; o^ in female of 428 mm. 



28 



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 ^ as long as orbit, 
soft, narrow triangular, tapering to very slender but blunted tip. 







Fig. 2. Fseuduraja Jischcri. 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 pelvics about 
80 per cent as great as length of orbit, its length, from center of cloaca 
to tip, about 1.5 times as gi-eat as distance from center of cloaca to 
base of rostral filament; the lateral caudal folds confined to about 
posterior 3^ 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 

Hearing the level of the caudal fin. The breadth of the folds is about 
3^2 S'S great as the height of the caudal above its axis, or about % as 
great as the length of the first gill openings. 

Entire upper surface of disc 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 disc and of anterior % 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 }/i 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 sm-face of pel vies 
wholly naked. Lower surface of disc 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 cm-tains smooth edged; those of the two 
sides of head actually separated by a space about }/2 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, 

its contom- not known for adult males. Teeth ofi7t~npW9S' those of 

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 ^q 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 }4 as great as its 
length, its depth below axis about j^ 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 diagnostic shape, the anterior outline directed nearly transversely 
to the main axis of disc and tail, and so long that the tips of the pelvics 
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 



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 pel vies 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 Breviraja. 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 % 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 Yi, 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. 

&ize. 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. 

Distinctive characters. This skate resembles R. hathyphila 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 disc and tail that we took 
it for a specimen of hathyphila on our first cursory glance. But a closer 
examination showed that it differs from hathyphila in a considerably 
more obtuse anterior contour of its disc (Fig. 3A) 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 disc 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. olscni. 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 
disc, 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 Nortli 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; ORP^GON Sta. 534, northwestern part 
of Gulf of Mexico, Lat. 27°32'N; Long. 93°02'\V; 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. 



12 



NO. 24 




Fig. 4. Raja fuliginea, type specimen 306 mm. long, U. S. 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 1st 1.3; 3rd 1.3; 5th 1.0; distance between 
inner ends, 1st 12.9; 5th 7.5. 

Distance: from tip of snout to center of cloaca 39.3; from center of 
cloaca to 1st 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. 

Pelvics. anterior margin 14.4. 

Teeth: ^. 

Disc 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 % (66^) of distance rearward from 
tip of snout toward level of axils of pectorals. 

Tail with very narrow lateral folds along posterior % (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 3-4 irregular rows along anterior % 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 
% of tail, and, again, rearward from level of origin of first dorsal fin. 

Snout in front of orbits about 23^ 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 L7 
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 j^, with low triangular cusp, blunted at 

tip, arranged in quincunx in juvenile male, probably also in female. 

P'irst 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 % 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 disc 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 

BiGELOw, Henry B., and William C. Schroeder 

1948. New genera and species of batoid fishes. Sears Found., .Jour. Mar. 
Res., Vol. 7, No. 3, pp. 543-566. 

1950. New and little known cartilaginous fishes from the Atlantic. 
Bull. Mus. Comp. Zool., Vol. 103, pp. 385-408, 7 pis. 

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. 

1951a. Three new skates and a new chimaerid fish from the Gulf of 
Mexico. Jour. Washington Acad. Sci., Vol. 41, No. 12, pp. 383-392. 

Chandler, Asa C. 

1921. A new species of ray from the Texas coast . . . Proc. U. S. 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, 

pis. 1-85, A-N. 
1913. The plagiostomia. Mem. Mus. Comp. Zool., Vol. 36, xiii, 515 pp. 

Atlas, 75 pis. 

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" . . . 
' Kept. U. S. Comm. Fish. (1893), pp. 393-476, 16 pis. 

Holt, 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 pis. 

Mueller, Johannes 

1841. Fortgesetzte Untersuchungen iiber die Pseudobranchien. Arch. 
Anat. Physiol., Jahrg. 1841, pp. 263-277. 



E V I O R A 

mseimmi of Cooiiparative Zoology 



Cambridge, Mass. JAxrARY 2S, 1954 Number 25 

A NEW MIOCENE SPECIES OF PELU8I0S 
AND THE EVOLUTION OF THAT GENUS 

By Ernest Williams 

Amoiij,^ the reptilian remains from the island of Rusinga in Lake 
\'ietoria, Kenya ( olony, sent for determination to tlie British Museum 
(Xatural History) are the greater part of the carapace and a smaller 
part of the plastron of an apparently new species of Pcluiiios. 

Dr. ^^'. 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: 

Pehsios ursLXGAE, new species 
Ti/pc: Coryndon Museum Ru FodlT a partial carapace and 
plastron. 

Horizon: Miocene of iiusinga Island, Lake \'ict()ria, Kenya ( olony.' 
Diaf/no.s-i.s: A Pciusios belonging to the a(laii.soiiii-(/aboiif7i.si.s 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 x'ertebral region \ery shallowly 
excavated, quite without keel; first vertebral scvite much larger than 
\ertebral 2 and wider than long; vertebrals 2, 3, and 4 slightly longer 
than wide; mesoplastra extremely narrowed medially, barely meeting. 
The living species of Pclusios fall into two sections: 
One, which is northern and western in distribution, comprises two 
species, I', (iddiisoiili and /-". gahoticu.sis. 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 

' Por a suniiijarv of the geologv and the Mioeene fauna of Rusinga and adjacent areas .see 
Kent (1944). 



2 BREVIORA NO. 25 

into and filling up the inter\'a,l left l)y the tapered margins of the 
mesoplastra. 

The other group within the li\ing membei's of the genus is less 
restricted in distribution. One of its species — P. suhnigcr — overlaps 
most of the range of the first group and in fact extends bej'ond that 
range on the west to the (ape Verde Islands. On the east this same 
species extends to Zanzibar, the Seychelles, Mauritius and ^ladagascar. 
On the north, however, this species does not extend beyond British 
East Africa into the Sudan range of P. adansonii. 

The group typified by P. subnigcr is distinguished l)y 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. rnsingac are clearly with the first of these 
two living groups: the tapered mesoplastra clearly indicate this 
position. From P. adansonii, however, P. nisingac difters (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. gahoncnsis 
it difl'ers 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 fo.ssil species (I utilize the data 
of Loveridge, 1941, which I have, however, verified on other material). 



P. adansonii 


P. gabonensis 


P. nisinyae 


Sulcus between humerals 


Sulcus between humerals 


Unknown 


3-4 times as long as 


1 1 2-2 times as long as 




that between pectorals. 


that between pectorals. 




Mesoplastra tapered 


Mesoplastra tapered 


Mesoplastra strongly 


medially only poste- 


medially both anteriorly 


tapered medially 


riorly, thus a 


and posteriorly, thus an 


anteriorly and 


transverse hinge with 


ol)lique suture with both 


posteriorly, 


the hj'oplastra but an 


hyo- and hypoplastra. 


hardly m(>eting. 


oblique suture with the 






hypoplastra. 






A keel on the anterior 


A nodose keel in the 


No keel, the 


four vertebrals 


young, lost in the 


vertebral region 


throughout life. 


old. 


somewhat 

depressed. | 



1954 EVOHTIOX OF PEI.I SIOS 3 

Vortebrals alwut as lonj; At least vcrt('l)ials 1 V(Mtcl)ial 1 wider 

as broad in adults. to H hroadcr tlian lonjj than long, 

in adults. vcrtebrals 2-4 

longer than wide. 
Height in longtli Height in length Height in length 

about 2.(1 times. 2.;^ to 3.8 times. about 4 times. 

Shell (iistineti.\- Shell not broadened Shell distinctly 

broadened posteriorly. ])osteriorl\-. broadened 

posteriorly. 
Known maxinnun size: Known maximum size: Estimated size: 

185 mm. 2r)!) mm. 24.T mm. 

/'. rufiinqar thus contrives to combine some of the characters of both 
the two living memliers of its group. It occurs, also, outside — south 
and east — though not far outside, the present limits of its group. 
Only P. srihuujcr of the alternative group is known from Lake Victoria 
today. 

Three fossil species of Pcliisios have been previously descril)ed: 
P. rudolphi Aramhourg from the Lower Pleistocene of Omo, founded 
on a partial plastron and carapace (type in Paris Museum); P. 
df'icitzianus \. Reinach represented by fragments from the Middle 
Pliocene of Wadi Xatrun (type formerly in Munich, now destroyed); 
and P. blanckcnhorni Dacque, a skull from the Lower Miocene of 
Moghara (type in Berlin?). In addition and not previously recorded 
there are ahimdant fragments (Nairobi Museum) and a complete shell 
(British Museum No. R oTOl) of P. sinuatus (a still living species of 
the P. subnigcr group) from Bed I, Pleistocene of Oldinai. 

The fossil P. sinuafiis need not be compared with P. ru.simior. 
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 ohl specimens of Recent P. niuuatus. 
It may provisionally be accepted as ancestral to the P. sinuafiis of the 
later Pleistocene and of the Recent. 

P. drwitzianiis was originally described on quite inadequate material 
which, however, was still sufficient to place it as a member of the 
P. subnigcr group. It was redescribed from much better material by 
Dacque (1912), who at the same time discovered that a supposed 
Pliocene species of Pclomcdu.s-d {P. pUoccnica v. Reinach) was a 
synonym of this species. P. dcnihiaiuis as a member of the alternative 



4 brf:viora no. 25 

group requires no comparison with P. nisiiu/(tc. 

There remains, however, P. hlnnckriihorni, which is from a deposit 
apparently equivalent in age and Aery similar in fauna to that of the 
Lower Miocene of Rusinga Island, but 2000 miles distant. P. hlauclcm- 
horni and P. rusitigac cannot be compared, since one is l)ased on a 
skull, the other on a shell. The skull of P. hJanckcnhorni, inadequately 
described and figured only in dorsal view by Dacqur, seems similar to 
that of P. gahoncnsis. It may, therefore, belong to the same group 
within the genus as P. rusingac, and it is not impossible that the latter 
is a synonym. 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. 
hkmckrnhorni, are founded solely upon the skull. It is a remarkable 
fact that Pchisios and Pclomcdusa, 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. blanckcuhorni may be either 
specifically identical with P. nisingae, or specifically dift'erent, or it 
may belong to a dift'erent 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 l)a'-ed 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 dift'erence 
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. rusingac and P. hlanckcnhorni (if this is really a Priusiu.s), 
occurring in the Lower IVIiocene, aie 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 Prioiiicdiisidae is certainly very old; it probably stems 



1954 EVOLUTION OF PELT'SIOS 5 

ultimately from the pleurosternuls of the Upper Jurassic and the 
( "retaceous. Unfortunately the form which has l)een suggested as the 
oldest representative of the family, PJatiirhclnidcs ni/nssdc Haughton, 
is incompletely known and doubtful as to agi'. It li;is small laterally 
placed mesoplastra and is therefore not an obviously primitive form. 
Mesoplastra meeting in the midline are certainly primitive for turtles 
and Pilu.sio.s would therefore be more primitive in this respect, Kulrss 
the larger mesoplastra of P. lusios are a secondary development (see 
below). The single known character in which Platychdoides differs 
from Pelonu'dusa as ordinarily conceived — the absence of the median 
plastral fontanelle — does not in fact separate it from that genus, 
since, as I have been al)le to determine on British Museum specimens 
from Uganda and the Sudan, the median fontanelle is sometimes 
lacking in even small specimens of PcJomrdusa. The beds from which 
PJatiichcJoidcs ni/a.s-.sai' derives are Cretaceous in age, but to what part 
of the ( 'i-etaceous 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 Xorth and South America, England, Italy (del Zigno, 1887), 
Egypt, Congo and India. 

It is a curious fact that every one of these older members of the 
family that arc sufficiently known is pelomedusine in type rather 
than pelusiine, that is: the mesoplastra are small and lateral elements, 
as in Pcluiitcdusd ami Pl(tti/clirli>ld(;s, not large elements meeting in the 
center of the plastron as in Pciu^ios and the pleurosternids. Nor is it 
at all likely that this o!)servation is 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 ccmdition 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 Pdusios are fully 
congruent. The members of the genus Pclusios form a structural series 
in regard to the size of the mesoplastra, P. rus'nKjar having the most 



f) BREVIORA NO. 25 

reduced mesoplastra, P. gahouru.'n'.s the next, P. adnnsonii next, and 
the P. siihnlgrr group the most fully developed mesoplastra. A 
structural series is always ambiguous unless the time dimension can 
be added, but P. rusitigac as the oldest shell belonging to the genus 
seems to provide this time element. 

The similarity of the skulls of Prloiiitdu.sa and Prlu.s-ioff further 
suggests relationship, and the existence of a species of PclomeduMi 
(P. progahata v. Reinach) anterior in time (Lower Oligocene) to the 
earliest (Lower ]VIiocene) Pchi.sios further supports the view that 
Pehmos is a relatively late and specializefl genus directly derived 
from Pelomcdusa. 

P. ru.s-ingac 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. (". 
Arambourg for the pri^•ilege of examining the type of P. rudolphi at 
the Paris Museiuu, 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 AL Jean Guibo 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 (niggenheim Fellowship during the year 
1952-53. 

RKFEKEXCES 
Arambourg, C. 

1948. Contrihutioii a I'otudc geologic [uc ft j)iileoiitologique du bassin 
du Laf Hodolphe et de la basse vakV tie i'Omo. 2me Partie. 
Paleontologio in Mis-sion Scientific [ue de I'Omo. 1932-1933. 
Vol. 1, fasc. 3, pp. 231-559. 
Dacqce, E. 

1912. Die fo.'^silen Schildkioten Aegypteiis. Geol. Paleont. Ahhandl., 
vol. 14, i)p. 273-333. 
Hatghton, S. H. 

192S. On some reptilian remains from the dinosaur bed.** of Xvassalaiid. 
Trans. Roy. Soe. S. Africa, vol. 16, pp. 67-75. 
Kknt, p. H. 

1944. The Micjcene beds of Kaviroiido, Keii\a. (^uart. .Jour. Geol. Soe. 
London, vol. 100. ])p. S5-ll(i. 



1954 EVOLUTION OF PELUSIOS 7 

LoVERIDCK, A. 

1941. Revision of the African tcnapin of the family Pclomcdusidac. 
liull. Mils. Comp. Zoo!., vol. SS, pp. 4()7-524. 

Rkinach. a. von 

1908. Sc-hiklkroteiHcstc aus dem uegyptischen Tcrtiar. Al)han(li. 
SenckenlxMg. Xaturf. (les., vol. 29, pp. l-()4. 

ZlGNO, A. DEL 

1887a. Chelonii scopcrti nei terreni cenozoici drlle i)i('al|M' Veiieti. Mem. 

R. Inst. Veneto, vol. 23, pp. 119-129. 
lS87b. Chelonio scoperto nel calcare nummulitico de avesa pressa Verona. 

Ibid., vol. 23, pp. 135-145. 



--^■m^. 








PLATr: 1 

Pehisios rusliH/(U', dorsal view of type shell 




PLATi; -2 
I'liufiioa rumngue, ventral view of iy\)v sli 




PLATE S 
Feliisios sinuatus {B.M.y^.H. H. o7(il). dorsal vit-w of shell Irom Bed 1, 
Pleistocene of Olduvai. 




PJ.ATl'] 4 
Pelasios sinuutiis (B.M.X.H. H. oTOl), ventral view of shell 
Pleistocene of Olduvai. 



E V I O R A 

Meseem of Comparative Zoology 

Cambridge, Mass. February 3, 1954 Number 26 



A PRELIMINARY LIST OF THE 

EARTHWORMS OF NORTHERN NEW JERSEY 

WITH NOTES 

By H. Davies 

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 featiu-es 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 

I 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 

II 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), 143(1), 147(1), 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-Hke, indentation at xxxii on upper side. 
157a band-like, tripartite origin, on xxxi-xxxiii. Intersegmental 

lines distinct. 
157b L. As 157a. 

R. Band-like, slightly indented at 31/32 and 32/33. 
161a \ 

) Band-like, tripartite origin, on xxxi-xxxiii. 
162a I 

162b ) 



1954 EARTHWORMS OF NEW JERSEY 3 

In each case the lateral portion of the tuberculum is translucent 
while the median portion is opaque and appears as a hand. 

Table I 
External characteristics of Allolobophora caliginosa 







Number 






Specimen 




of 


Tubercula 


Genital 


No. 


Locality 


segments 


Clitellum pubertatis 


tumescences 



150a 


Dover 


133 


27-135 


31-33 


30 32 33 34 


150b 


Dover 


179 


27-i35 


^30-33 


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 


161a 


Pompton 












Plains 


161 


27- 34 


31-33 


30 32 33 34 


161b 


Pompton 












Plains 


1.56 


27- 34 


31-33 


33 34(R) 


162a 


Paterson 


159 


27- 34 


31-33 


30 32 33 


162b 


Paterson 


130 
(amputee) 


27- 34 


31-33 


30 32 33 34 



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, 1 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 ?'?? copula approximately 4 inches 
underground. 

This is the second record of this species in North America, Gates 
(1953, p. 518) having found it in the x\rnold 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). 



NO. 26 



Allolobophora longa Ude 1885 



Dover, sandy soil in garden, Nov. 1951, 3 elitellate specimens. 
Morristown, in garden soil (clay), May 10, 1953, 1 elitellate 

specimen. 
Dover, in garden on 2nd St., June 8, 1953, 3 elitellate specimens. 
Beatystown, damp soil in meadow, June 9, 1953, 1 elitellate 

specimen, 1 aclitellate (post sexual?) specimen. 

Genus BiMASTOS Moore 1893 
BiMASTOS PALUSTRis Moorc 1895 

Flanders, in wet moss in rocky stream near Route 31, Nov. 1951, 

4 elitellate specimens. 
Hardwick, in ditch in forest (running water), April 1952, 3 

elitellate specimens. 
Ironia, in wet moss by stream, Succasunna-Ironia Road, May 

3, 1952, 1 elitellate specimen. 
Swartswood, under moss on log by stream. May 29, 1952, 15 

elitellate specimens in association with E. tetraedra. 
Mt. Freedom, in wet moss on marshy ground by stream, April 

27, 1952, 7 elitellate and 1 juvenile specimens. 
Shongum, Raynor Road, in very wet sandy loam by stream, 
March 29, 1953, 4 elitellate specimens. 
This species seems to have an affinity for running water and is 
apparently more restricted in this respect than Eiseniella 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. 

Sperm atophores 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 elitellate 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. ruhida. 
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 ElSENIA 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 



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 liy stream. May 3, 1952, 

2 clitellate and 4 juvenile specimens. 

Shongum Lake, in wet soil by stream. May 1952, 1 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 Innnbergi 





Number 












of 




Tubercula 


Regener- 




Condition 


segments 


Clitellum 


pubertatis 


ation 


Remarks 


aclitellate 


77 


— 


26-28 


56/57 




clitellate 


62 


24-30 


26-i29 


— 


Amputee 


clitellate 


110 


P3-30* 


26-28 


— 


Amputee 


aclitellate 


129 


— 


26, 27, 28 


— 




juvenile 


76 


— 


— 


— 




juvenile 


116 


— 


— 


— 




immature 


48 


— 


■ — 


7/8 


Apparent head 
regenerate 


clitellate 


44 


§24-30 


26-28 


— 


Amputee 


clitellate 


79 


?24-30* 


26, 27, 28 


— 


Amputee 


clitellate 


51 


24-30* 


26-^29 


— 


Amputee 


clitellate 


142 


?24-30* 


26-i29 


— 




juvenile 


133 


— 


— 


101/102 




juvenile 


132 


— 


— 


47/48 




juvenile 


74 


— 


— 


— 


Amputee 


juvenile 


118 


— 


— 


— 




juvenile 


132 


— 


— 


— 




juvenile 


135 


— 


— 


— 




juvenile 


104 


— 


— 


— 




clitellate 


125 


24-30* 


26-i29 


— 








*Clitellum feebly developed 







1954 



ElARTHWORMS OF NEW JERSEY 



There are few references to this worm and Httle 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-^31 


26-31 


12(L), 13(R) 


44b 


114 


25-32 


29-131 


26-31 


11, 12(L), 12(R) 


61 


109 


25-31 


29- 30 


26-31 


11, 12 


55 


128 


26-32 


29- 31 


26-32 


12(R) 


122 


131 


26-32 


29- 31 


25-32 


None 


120 


130 


i26-32 


30- 31 


12(L), 11,12(R) 
26-32 


None 



Further description of tubercula pubertatis : 
No. 44a Tubercula bluntly elliptical from 28/29 to |31 without 
marginal incisions. Intersected by furrow 29/30 only. 



8 BREVIORA NO. 26 

44b Shape as 44a but extending to |31. 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 tuberculimi 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. maccdonica 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 ElSENIELLA 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 Hoffmeistcr 1843 

Stokes State Forest, Big Flat Brook, under leaves, May 30, 1950, 
1 clitellate specimen. 

LuMBRicus TERRESTRis Linuacus 1 758 
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 

xxii (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 | in. in diameter and ^ 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 11 

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 Idnnbergi 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 Uinicola, 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 arnoldl 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 Phcrdima 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 subriibicunda 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. Allolohophora 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 Eiseniella tctraedra 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 are recorded for the first time 
from this state, Dendrobaena viammalis being reported for the first 
time outside of Europe. Bimastos palustris and Eisenia lonnhergi are 
considered to be endemic and are widely distributed in the northern 
part of the state. Allolohophora limicola was found for the second 
time outside Europe and in three localities. Three species of Pheretima 
were present in a garden. 



REFERENCES 

Cernosvitov, 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. 

Gates, 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 pahistrls, a new oligochaete. Jour. 
Morph., 10: 473-496. 

Olson, H. W. 

1940. FJarthworms of New York State. Amer. Mu.s. Nov., No. 1090: 1-9. 

S.MITH, F. 

1917. North American earthworms of the family Lumbricidae. Proc. 
U. S. Nat. Mus., 52: 157-182. 



E V I 

MeseiLim of Comparative Zoology 

Cambridge, Mass. February 5, 1954 Number 27 



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 FAsenia foctida (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 E. foctida, 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 deferential 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" [idem, 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. foctida 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? 

' This means: segmental loration of the external openings of the deferent ducts, i. e.. the 
male pores. GEO 



2 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 



9-10 

10 

10-11 

9, 11 

L9, LIO, RIO. 
R9, RIO, LIO. 
RIO, LIO, Lll 



17 



L9, LIO, R9, Lll. 
L9, LIO, RIO, Rll 
R9, RIO, L9, Lll. 
R9, Rll, LIO 



Spiral abnormality in metamerism of the 
spermathecal region 



29 



* On the left 



ide 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 necessary 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 E. 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 tliree 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''. 



- In 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 lapcftis calami for seminal vesicles. The latter 
are four pairs, in ix-xii, in E. 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 sperraa- 
thecae in two segments were quadrithecal and, as he supposed, E. 
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 Eisenia 
lonnbergi (Michaelsen) 1894 with spermathecal pores on 8/9-10/11, 
Dendrobaena odaedra (Savigny) 1826 with the pores on 9/10-11/12. 
E. lonnbergi, 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 som-ce 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 foetida, and has only once been reported 
from the vicinity of a manure pile. D. octacdra does have the same 
red pigmentation that characterizes E. foetida, uniformly distributed 
(instead of in transverse bands) as in many individuals of E. 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 E. 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 E. foetida and two species which have not yet 
been studied in this country, Allolobophora terrestris (Savigny) 1826 
and Lumhricus 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 


5 


pages 


EL 2/3 


1 


- 


- 


- 


447 


EL 3/4 


3 


1 


- 


- 


447, 455 


EL 4/5 


- 


2 


- 


- 


448, 456 


EL 5/G 


- 


1 


1 


- 


455 


EL 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 
spermatheoae. 

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 103^ 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. 

< The number of segments in the head regenerates of E. 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 5 

SUMMARY 

Individuals of some sexthecal species were frequently used along 
with E. foctida 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. octacdra is likely to have been inadvertently taken for E. 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 

Gates, 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. 

MORGA.V, T. H. 

1S95. A study of metamerism. Quart. Jour. Micros. Sci., 37: 395-476. 



E V I 

Museium of Comparative Zoology 

Cambridge, Mass. Fklriary S. 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 Kmys turnavrnsis 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. 

Kmys fiimauensis 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 vertel)ral 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 ^'ertebrals 
less obviously but unmistakably had a similar great lateral expansion. 
Von Meyer commented: "Von alien mir bekannten Schildkroten mit 
Grenzeindriicken zeichnet sich vorliegende durch den Mangel an 
Seitenschuppen aus. . . . Der Verlauf der Grenzeindriicke sonst ist in 
dieser Schildkrote so regelmassig dass der Mangel an Seitenschuppen 
unmoglich fiir eine zufallige Erscheinung oder fiir eine Abnormitat 
gehalten werden kann: er wird der Species wirklich zugestanden haben, 
und es wird sich eigentlich nur um Entscheidung der Frage handeln, 
ob der ganzliche Mangel einer Schuppenart in einer Schildkrote 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 mochte ich um so mehr 
bis zur Kenntniss der fehlenden Theile der Schildkrote, namentlich 
(les Bauchpanzers zuriickhalten, da die hervorgehobene Abweichung 
im Hautskelet, so auffallend sie ist, mit einer Abweichung in der Zahl 
oder Form der knochernen Theile, wenigstens so weit diese verliegen, 



2 BREVIORA NO. 28 

nicht verbunden sich zeicht; die knochernen Theile sind vielmehr 
Emys entsprechend gebildet. Zur P^rrichtung jedoch einer neuen 
Species war wohl hinreiehender Grund vorhanden." 




Fig. 1. Clemmydopsis turnauensis (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 
Teitiary 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 Eviys turnaucnsis but which he referred to a new genus and 
species as Clcmmydopsis soproncnsis. The new form differed from 
Emys turnaucnsis in the shape of the neurals. In Cleminijdopsis so- 
pronensis neurals one to three were hexagonal, short-sided behind, and 
neural four quadrilateral, while the first neural was oval, the second 




Fig. 2. Clenuniidopsis sopronensis Boda. A. Type carapace, after Boda. 
B. Plastron, after Thenius. 

to fourth hexagonal, short-sided in front, in Kmi/s turnaucnsis. 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 Sarniatian Miocene 
of Steinheim in Wiirttemberg material very similar to the unique type 
of Emys fnr7iavcnsis but which he regarded as new, giving it the name 
Clcmmys stcinhcimensis. 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 turnaucnsis and Clcmmydopsis 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 turnauensis as they quite clearly did in sieinheimensis. He 
relegated both forms to the genus Clnnviys, regarding the single point 
of difference from Clemviys as usually understood — the absence of 
the first and second costals — as of specific value only. 

In the case of Clemmydojms soproncnsis, 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 steinhcimensis 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 CJrmmys 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 
Miocan und Pliocan auf, um alsbald wieder zu verschwinden. Man 
konnte versucht sein in Dacque'schem Sinne von einer IModestromung 
zu sprechen, denn ein besonderer Zweck diirfte dieser Einrichtung 
kaum zugrunde liegen. Mit der Annahme einer naheren 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 Szalal's generic assignment (pre- 
ferring, however, the synonymous name Nicoria) in reporting new 
finds of sopronensis from a new locality Brunn-Vosendorf near ^'ienna. 
Thenius' material is important in that it completes our knowledge of 
sopronensis by providing (from dissociated specimens of all the plastral 
paits) the characters of the plastron, heretofore unknown. 

Thanks to the kindness of the authorities of the Staatliche IVIuseum 
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 sieinheimensis 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 steinheimemis may best be synonymized 
with turnauensis. The two named forms are equivalent in age, not far 
distant in locahty and distinguished by a single quite trivia] character 
which may well be only an individual peculiarity of the unique type 
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", "Clemviydopsis, 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 
(ryptodira were placed in the genus Emys. 

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 soprotiensis is a quite extraordinary phenomenon apparently 
not closely approached by any recoided 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 appealed 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 



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 Clemm.i/s type neurals. 
There are indeed a number of examples of evolution in neural shape 
displayed in the fossil record. The fossil tortoise Stylcmys nebrascensis 
of the Oligocene cf North America regularly has all the neurals after 
the first hexagonal short-sided in front : the more highly evolved species 
of the same genus from the John Day beds (lowest jMiocene) regularly 
have one or more octagonal neurals. Hadrianus with primitive neurals 
evolved into later subgenera of Testudo with modified neurals. There 
is also among Recent forms considerable intraspecific variation in 
neural shape. 

In the present instance turnaunisis 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 other. 

Except for neural shape there are no known differences between 
turnauensis and soproncnsis 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 

1. gulars narrow, not overlapping 
humerals 

2. femoral scutes longer than 
pectorals 

3. no anal notch 

4. first vertebral encroaching on at 
least the first, second and third 
peripherals 

5. first neural rectangular-oval, 
neurals 2-8 hexagonal, short- 
sided in front, all moderately 
broad 

6. fifth vertebral and fourth costal 
moderate in size 



soipronensis 
gulars broad, overlapping hu- 
merals 

pectoral scutes longer than 
femorals 

a distinct though nan-ow anal 
notch 

first vertebral encroaching only 
on the first and second peri- 
pherals 

neurals 1-3 hexagonal, short - 
sided behind, neural 4 quadri- 
lateral, neurals .5-8 hexagonal, 
short-sided in front, all still 
broader than in turnauensis 
fifth vertebral and fourth costal 
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 Clcmmiidopsis 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 Clcmmys, but this is an untestable hypothesis, which it is 
useless to consider. The question is then a simple one: Is a single 

charactei 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 — Clemviiidoysis 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 Tcstudo dcnticulata and one in Chri/seinys pida 
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 Notochdys with the 
single species A. plati/nofa which differs from Clcmmys 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 Clcmmy- 
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 Clcmmy.'i, where Staesche has already placed it, and 
the Pliocene form in Gcoemyda ( = Nicoria) to which Szalai and 
Thenius have already referred it. This, however, is an inadmissible 
solution since it would make the genus Geocmyda polyphyletic. 

It is entirely possible, indeed probable, that Clemmys and Gcoeiayda 
as at present defined are mere form genera, but the true relationships 
within the Clcmmys-Geocmyda section of the Emydinae are still to be 
analyzed and promise 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 afPorded by a Guggenheim fellowship 
1952-1953. 



1954 CLEMMYDOPSIS 9 

REFERENCES 

BODA, A. 

1927. Clemmt/dopsis sopronensis, n.g., n.sp. aus der uuteren pannonischen 
Stufe von Sopron in Ungarn. Zentralbl. Min. etc., Abt. B, 1927, 
pp. 375-3S3 
Cain, A. 

1952. Geography, ecology and coexistence in relation to the biological 
definition of species. Evolution, vol. 7, pp. 7«i-S3. 

COKER, R. 

1910. Diversity in the scutes of Chelonia. ,Iour. Morph., vol. 21, pp. 
1-7.5. 
Gadow, H. 

1S99. Orthogenetic variations in the shells of Chelonia. in Arthur 
Wiley's Zoological Results, part 3, pp. 207-222. 
Grant, C. 

1936. An extraordinary tortoise shell. Copeia, 193'o, p]). 231-232. 
Lynn, W G. 

1937. Variation in scutes and plates in the box turtle, Terrapene rnrolina. 
.\mer. Nat., vol. 71, pp. 421-426. 

Meyer H. von 

1847. (Letter on various fossils.) Neues .Jahrb. f. Min. etc., 1S47, pp. 

181-196. 
1858. Schildkroten und Saugetiere aus der Braunkohle von Turnau in 
Steyermark. Palaeontographica, vol. 6, pp. 50-55. 
Staesche, K. 

1931. Die Schildkroten des Steinheimer Beckens. A. Testudinidae. 
Palaeontographica Suppl., vol. 8, part 2, pp. 1-17. 
Szalai, T. 

1934. Die fossilen Schildkroten Ungarns. Folia Zool. Hydrobiol., vol. 6, 
pp. 97-142. 
Thexius, E. 

1952. Die Schildkroten (Testudinata) aus der Unterpliocan von Brunn- 
Vo.sendorf bei Wien. Neues Jahrb. f. Min. etc., 1952, pp. 318-334. 



E V I O R A 

Museeirai of Coeiparative Zoology 

Cambridge, Mass. February 8, 1954 Number 29 

ABSENC^E OF MESOPLASTRA IN A 

PELOMEDUSA 

(TESTUDIXES, PELOMEDUSIDAE) 

Hy Ernest Williams 

Mesopla.stra (a pair of l)ones intercalated between liyu- and hypo- 
plastra) are believed to be primitive features in turtles. They were 
often present in Jurassic and Cretaceous genera, and two pairs of 
inesoplastra were reported by Fraas (1913) in Profrrochcrsis, one of 
the Triassic genera. Mesoplastra have, however, been lost se\eral 
times independently: ( 1 ) in the Jurassic and Cretaceous plesiochelyids 
and thalassemydids; (2) in Wealden Chiiracrphahis\ (3) in all crypto- 
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 — Plafi/chch/s — and in some Baenida.e (within the 
genus Bai'na, cf. Bai'iut rijxirid. Hay, 19(),Sj they fail to meet in the 
center. 

To my knowledge, howe\er, 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 Pclomcdusa suhrufa from I'ganda, collected by Delme 
Radcliffe and now 1905-5-19-1 in the collections of the Reptile Section, 
British Museum (Natural History). 

In Pclumcdtisa the mesoplastra are typically small and lateral. In 
HM 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 C'helidae or the 
Cryptodira. The carapace again joins the plastron perfectly without 
aft'ording any place for a trace or rudiment of mesoplastra. 



2 BREVIORA NO. 29 

Yet this specimen is clearly referable on "habituellen ]\Ierkmalen" 
to the family Pelomedusidae, the genus FcJoiucdnsa, the species 
suhrufa (the single species of the genus). 

The carapace is quite normal except for some minor asynnnetries 
and the fusion of vertel)rals 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 scutes. There is the usual median fontanelle, though it is 
small in this instance, as seems frec^uently 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 siiown in the 
excellent photographs (Plates 1 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. P^lgon, I'ganda, otherwise \ery 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 f)f 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 Prlu,s-ios the mesoplastra are 
always small and lateral, in effect vestigial, and the disappearance by 
individual variation of a \estigial feature is not too surprising. 

It is, howe^•er, of special interest in connection with the suggestion 
recently made by me that the genus Pelusloft with well-developed 
mesoplastra (meeting medially) has been derived directly from the 
genus Priomcdus'a with reduced mesoplastra by a secondary expansion 
of these elements. If this suggestion be \alid, then, taken in connection 
with the present case, we see in Pclonialusd an interesting ambivalence 
of evolutionary potentiality, one type of \ariation reversing a previ- 
ously well-defined trend to restore (with some differences) an ancestral 
condition and, on the other hand, the o])posite type of variation 



19r)4 ABSF.XCK OF MESOPLASTKA I\ A PP:LO!MKDrSA 




iMjiurc 1. Plastni of viirious lurtlcs to show |)rcsciicc or ;il)sciicc of 
nicsoi,l;isti;i. M(>so|)l;is1 rn shaded. A. I'rlu»iv,l u^ii witliout nicsoplastra. 
H. \oYm-A\l',l(,m,'ditsa. ('. a v\n'\yu\ {II ijilmwrihim). D. f>,h,sios. E. Po- 
dornemis. {W to [• alter Houlcnii'T.^ 



4 BRKVIORA , 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 occasional men ])orn 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 
selecti\'ely valuable variant, but, as the majority of the living forms 
testify, a turtle without mesoplastra is not only perfectly A'iable 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 hafl 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 obser\'ations recorded here were made during a \isit to the 
European collections, made possible by a Fellowship from the Guggen- 
heim Founflation. 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 ]\Ir. J. C. Battersby 
and Miss A. G. G. Grandison of the Reptile Section. 

hp:ferexcp:s 

Fraas, E. 

1913. I'luteruchersis, eine pleurodire Schildkrote aut; dein Keuper. 
Jahreshefte Ver. Vaterlands Wiirtt. Xatuikunde, Stuttgart, 191.3, 
pp. 13-30. 
Hay, O. p. 

1!)0S. Fo.«sil turtles of Xorth America. Carnegie Inst. Washington Puhl. 
X'o. 75, pp. 1-.5()S. 

LoVERIDfiE, A. 

1941. Revision of the American terrapins (jf tlie family Pelomedusidae. 
Bull. Mus. Comp. Zool., vol. 8S, pp. 4(57-524. 




PLATE 1 

Pelomedum s„hn,fa H.M. X.H. Xo. IWa-o-lO-l. IiitiTiial view of jilastmn. 




PLATE 



Pelomcilusa snhnifa H.M.X.H. Xo. 19()r)-r)-l<»-l. Dorsal vi<'\v o\' skull and 
shell. 



E V I O R A 

Musceiim of Coimpsirsitive Zoology 

Cambridgp:, Mass. I"i;iii;rAKV S, 1954 XrMBp:R :>() 

AKSTIVATIOX I\ A PERAIIAX LUXCJFISH 

Hy Alfred S. Romeh 

Musi'uiii of ConipMr.-itivc Zooloj^y, Harvard riiivcr-sitj' 

and 
Everett (\ Olson 

I'liivcrsity of Chicaji-o 

It is generally recognized that the survixal to modern times of the 
three living genera of dipnoan fishes — K/jicirdtix/us i Xconrdfodii.s) of 
Australia. Profa/ifrriis of Africa and Lcpido.sirm of South America, - 
is due in great measure to adaptations which tide them oxer the 
seasonal di'oughts characteristic of the tropical regions which they 
inhabit. .\ primary adaptation lies in their pos.se.s.sion of functional 
lungs which enable tliem to utilize atmospheric oxygen, supplementing 
the refluced oxygen content in stagnant waters during drought; indeefl, 
in Proiopirrus, the hmg-hreathing fimction has attained such domi- 
nance that the fi.sh is unable to sm-\i\e if air-breathing is not possible. 

But a.ir-l)reathing in itself is not entirely satisfactory as a means of 
drought sm-\i\al - particularly if, as may often happen, the stream 
or pond dries up entirely. The Australian lungfish, under such circum- 
stances, may wallow into the mud, and have some chance of escaping 
desiccation or enemies. In the African and .South .Vmerican forms, 
an additional adaptation is present the de\elopment of a technique 
of aestivation. Meml)ers of these genera are rather elongate and eel- 
like in shape, in contrast to I^piccrafndiis and to primitive lungfishes. 
On the approach of the dry season the fish burrows in coiled form into 
the mud, which hardens atiout it to form a cocoon; an opening from 
the surface allows the entrance of air, but the amount neerled is small, 
due to a great reduction of metabolic processes. The fish passes mto 
a "summer sleep" compara!)le in great measure to the hibernation of 
various northern animals — a phase which ends with the i-eturn of the 
water and tiie resumption of normal a(|uatic 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 Kplccratodu.s', and hence 
were incapable of coiling in a burrow in the fashion of the elongate- 
bodied Lcpidosiren or Protoptervs; but it is possible that burrowing 
of some sort accompanied by phsyiological features of aestivation, 
might have preceded change in body shape. The lungfish (I'nathorhiza, 
characteristic of the Clear F'ork Permian of Te.xas, has been suspected 
by both the present writers to be related to the Lepidosirrn-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 P>onomic 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 ( "ounty, 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 alcove the Lueders limestones and hence in the 
lower part of the Arroyo Formation of the ( lear F'ork 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 o\er the surface are 
numerous circular discs, somewhat like large checkers, with diameters 
which are for the most part between 5.5 and 10 cm. The material of 
the discs 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 2S 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 AKSTIVATlOX 1\ A PEKMIA.N UNGFISH 3 

the material of the cylinder Mends laterally with the superposed 
calcareous 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 rehance 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 frecjuently 
seen in section and in a few specimens fracture has shown scales in 
surface view, the largest about 20 mm. in greatest diameter. Numerous 
discs 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 a.xis, 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 l)e made out body outlines, markefl by scale sections with 
a. surrounding layer of darker matrix. 

The only lungfishes known from the ( 'lear Fork belong to tlie genus 
Gnathorhiza, mentioned above; Sagcuodus, the common Pennsylvanian 
genus, possibly antecedent to (\'rato(lu.s' and Epiccrntodus, is not 



4 BREVIOKA NO. 30 

recorded in Texas abo\'e the ^^'ichita group. But while it is probable 
that the Reed Ranch lungfish is Guathorhiza, no tooth plates have been 
found there to make identification certain. A second fintl, however, 
lias produced a number of fragmentary teeth. These are unmistakably 
the blade-like, shearing teeth of (inafhorhiza and are referable to the 
species G. dikrloda rather than G. .si rrntfi. This locality was discovered 
by the junior author and Dr. Nicholas Hotton III in 1949 and has 
been levisited several times since then. The bed in which the cylinders 
occur is located in north central Knox ( ounty, Texas, on the Waggoner 
ranch. It crops out along a small tributary to Little Mustang ("reek 
in locality KI of the junior author (see Olson, 1951 p. 104). The age 
is middle ^'ale of the Clear P^'ork Group, Lower Permian. The site is 
approximately 700 to 800 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 ^'ale locality occur in a deep 
red shale which ^■aries 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 al)out an acre. The cylinders are distributed in rather irregular 
groups of ten to twenty in unevenly spaced areas not much more than 
a yard scjuare. The longest cylinder encountered measured 45 cm., 
hut as in the case of the Arroyo specimens no upper termination has 
l)een identified. The vertical orientation shown in the specimen figured 
is constant throughout the deposit. 

Organic remains in the cylinders from the two localities dift'er 
somewhat. The most common remains in the \'ale (ylinders are ribs, 
skull plates and scales of Giiaihurhizn 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. ]\Iore abundant, 
howevei-, are partial vertebral cohunns aiul single \ertel)rae of the 
small, worm-like amphibian, Li/.soroj/hiis. In no case have these 
columns been found in the coiled conflition characteristic of Ijisonipluis 
specimens throughout the Arroyo formation. 

There can hardly be any question that the lungfishes from the two 
localities l)elong to the same genus. Some question does exist, how- 



1954 AKSTIVATION IN A PERMIAN LUNGFISH 5 

ever, with respect to specific identity. Two species, (1. .scrrata and 
(>'. dikcludn, are known from the Vale formation, l)nt only the first 
has been fonnd in the Arroyo (Olson, 1951). Tooth plates of (1. .srrratu 
are small, lower plates ranging from 10 to 12 nnn. in length, whereas 
those of G. dikcludd are between 25 and 32 mm. long. The skulls of 
adult individuals of (1. dikcloda appear to have been between 75 and 
100 mm. long; this would suggest that the skulls of (1. terrain, unknown 
as yet, would have a range of between 25 and 35 nun. The fact that 
the Arroyo and \'ale 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 extensi\e exploration of its 
exposures. Except at the Reed Ranch locality lungfish remains ai-e 
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 
(j. dikcloda 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 \'ale, 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 sur\ived 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. P'ortunately, 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 prol)ably 
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 cylinders indicates that a certain percentage of the aesti\ating 
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 l)y sediments. This probably applies to the 



6 BREVIORA NO. 30 

remains of Li/.s-orophus as well. This amphibian, like (riiatJiorhiza, had 
both aestivating and free swimming 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 Gnaihorhiza teeth in the material from the \ ale (KI) 
locality, together with the absence of identifiable remains of any other 
lungfish type in the Clear P^ork beds makes it seemingly certain that 
the burrows were made by thi?t lungfish and that alone. This evidence 
of the adoption of aestivating hal)its by Gnathorhiza strengthens the 
suggestion, gained from tooth plates and skull pattern, that this genus 
is possibly ancestral to the modern aestivators, Lcpidosiren and 
Protoplcrus. One major difi'erence 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 liurrows 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 buri-ow 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 PLATP] 

Above, a sciich; of cxpos^ed burrows at the Reed Ranch locality; a half-dollar 
gives ail indication of size. Cciitrr, a terminal section of a burrow developed 
to show an included lunglish 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. Beloir, left, a burrow in place at the KI locality; length 45 cm. Belniv, 
right, an incomjilctc !)Uii()w from the Reed Ranch locality; length 25 cm. 



8 BREVIORA NO. 30 

The genus (inathorhiza is known not only from the Texas beds, l)ut 
also from the late Pennsylvanian of Illinois. In Texas, however, it is 
very rare in the Wichita formations of the Permian where Sagrnodus, 
common in the Pennsylvanian, is fairly abvmdant: on the other hand, 
there is no trace of Sagcnodus in the ("le.ir 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 fauna from Sagcnodufi, a presumed 
non-aestivating form, to (Inathorhiza, ))etter adapted for survival under 
drought conditions. 



LITEH.\TrRK CITED 

Oi.sox, K. C. 

HUS. A pr.'liiniiuu y report on vcrtclirates from t\v' Permian Vale 

formation of Texas. Jour. Geo!., 56: 1S6-198. 
11).") 1. Fauna of Upijer Vale and Choza. 3. Lunj; fish of the Vale 

4. The skull of Gmthnrhiza dikeloda Olson. Fieltliana, Geol., 10: 

104-124. 

Ro.MER, A. S. and H. .1. S^^TH 

1934. American Carboniferous dipnoans. .Jour. Geol., 42: 700-719. 



B R E V I 

Miaseiiei of Coimparative Zoology 

Cambridge, Mass. Makc h o. 1954 Ximber 31 

NEW FRESHWATER ClASTROPOD AIOLLUSKS 
OF THE AFRICWX GEXUS LAXISTES 

By T. Pain 

London, England 
The two new species and one subspecies flescril)ed herein form jxirT 
of the collection of the Museum of Comparative Zoology at Har\ard 
College. I am deeply indebted to Dr. J. C. Bequaert and Dr. A\'. I. 
Clench for the opportunity to examine and describe them. 

La.\i.stp;s (LAXISTES) KOBELTi, new species 
Figures 3-4 

l!»lo. Lanistes .«p.? Kobelt, Mon. Amp. in Martini and Chemnitz, Svst. 
Conch. Cab.. 1, Aht. 20, Xeue Folge, pi. 30, Hgs. 4-4a. 

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. I'mbilicus 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 1.5 11 Fig. 4 

IS 23 15 10 



2 BKEVIOKA NO. 31 

Localiiy. "Africa" (probably West Africa); holotype M.C.Z. No. 
173419; paratype.s M.C.Z. No.'l73420 and in T. Pain ("oil. 

This unusual little species was apparently figured by Kobelt (1913, 
pi. 30), but he did not name it or mention it in the te.xt. 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.) kohrUi is, as the above table shows, very constant in 
size and subject to but little variation in this respect. 

LaiMstes (MELADOMis) coNNOLLYi, new species 
P'igures 12 

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 flne 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 


Wi.lt li 


Ape 
Length 


rtu 


Width 




Holotype 


40 


36 


31 




22 


Fig. 2 


Paratype 


45 


40 


32.5 




26 


Fig. 1 


" 


43 


39 


30.5 




24.5 




" 


55 


50 


38.5 




30 




" 


45 


44 


33 




27.5 





LocaUtivs. Southern Rhodesia: Mctoria P'alls, 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 ^lajor 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 FRKSHWATEK (JASTKOPOD MOLLUSKM 3 

1938, Ann. Soutfi 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. 

Lanistks (MKi-ADoMisi (j\ I M MWERiEXisis, new subspccies 
Figures 5-7 

Shell small, fairly thick. ( 'olor 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 of !)luish- 



ite color. 


()j)crcului 


1 a 


s in typical L. (M.) ovum 








Me 


isu 


rements 


in millimeters) 








Lrlljrtli 




Width 


Apertu! 
Length 


Width 




^olotype 


30 




30 


19.5 


16 


Fig. 5 


Para type 


30 




32 


21 


15 


Fig. (i 


" 


'ifi 




25 


19 


13.5 


Fig. 7 



Localities. Belgian Congo: Lake Mweru (or Moero) at Kilwa, 
holotype, collected l>y Dr. L. van den Berghe, M.C.Z. No. 109842, 
and paratypes, M.( '.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. ^L Wanson); Lake Mweru at 
Pweto, paratypes, M.( ".Z. No. 152346 (received from Dr. ^L Wanson) ; 
Lake Mweru at Moba (Chamfubu River) near Pweto, ^LC.Z. Xo. 
185682 (received from Dr. M. Wanson). 

Only the holotype (Fig. 5) and one of the paratypes (Fig. 6), l)oth 
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. (M.) ovum Peters, possibly ccMifined 
to Lake Mweru. When adult it resembles the much larger L. (M.) 
ovum procerus v. Martens, but has always a much thicker and more 
solid shell. Immature shells are always of a unif<irm olive-green color 



4 BREVIORA NO. 31 

often with traces of malleation on the body-whorl. So far as known, 
Lanisirs oman is the only species of the genus recorded from Lake 
Mweru, where it is said to live in the swamps near the coast. 




I''ig.«. 1-2, Lanistes (Meladomus) ronnolli/i. new species: 1, paratype, X'kana; 
2, holotype, Victoria Falls. Figs. 3-4, Lanistes (Lanistes) kohelti, new species, 
Africa: 3, holotype; 4, paratype. Figs. .5-7, Lanistes {Meladomus) ovum 
mweruejtsis, new subspecies, Kihva: o, holotype; <i and 7. paratypes. All figures 
slightly over natural size. 



E V I O R A 

Mmseiiiimi of Coimparative Zoology 

Cambridge, Mass. March 12, 1954 Number 32 

FOSSILS AND THE DISTRIBUTION OF 
CHELYID TURTLES 

2. Additional Reputed Chelyid Turtles on 
Northern Continents: 

Palaeaspis conybearii (Owen) — a Pelomedusid. 

By Ernest Williams 

In the previous paper of this series a re-examination of the descrip- 
tion of " Tcstvdo" Iciikii 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 Pakuasyis was proposed by Gray for Emys 
conybearii 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 Hydromcdiisa 
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, Iruncated in front and notched behind, transversely 
concave in the middle and bent up at the ends." 

Examination of the type specimen of Etnys 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 Hydromcdusa could be 
interpreted as a nuchal succeeded by a small first vertebral. This 




Fig. 1. Emys conyhearii 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 conyhearii, although it was mentioned and discussed 
by Owen in the original description and was clearly shown by him in 
his plate XXVIIIA, which is a much restored but essentially accurate 
representation of the specimen. 

E. conyhearii Owen was synonymized by Lydekker and Boulenger 
(1887) with the earlier named Emys delahechci Bell, probably justly, 
but the latter name was based upon an imperfect shell which does not 
show the region which is critical in E. conyhearii. With the example of 
Hydromedusa in mind, it is not possible without further evidence to 
dismiss the condition shown in E. conyhearii as an individual varia- 
tion. Further, since the bony areas under the first vertebral are 



1954 



FOSSILS AND DISTRIBUTION OF CHELYID TURTLES 



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. Enujs 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 E. co7iyhcarii after Owen's description of it 
revealed the presence of small lateral mesoplastral bones as in 
Podocncmis and Piiomcdusa. 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 vertebraP occurs in a chelyid genus. The elongation of the first 
vertebral and of that region of the bony shell is rather character- 
istically chelyid (though occurring also in Pdusios). If it were not for 

' 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. 




Fig. 3. Emys delabechei Bell. Dorsal view of type 
Bell). 1/5 natural size. 



(after Owen and 



Comparison of the types of Emys conyhearii 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 howerbanki 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 hoicerhanki 
under the name Podocnemis bowerbanki. They have also synonymized, 
as alread}' mentioned, Emys conybearii with Emys delabechei, again as a 
species of Podocnemis. Only the type of Emys conybearii approaches 
completeness, but all four specimens, though differing in size, are 



1954 



FOSSILS AND DISTRIBUTION OF CHELYID TURTLES 



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 




Fig. 4. Platemyshowerhanki Owen. A. Dorsal view of type shell. B. Ventral 
view of type shell. (Both after Owen and Bell.) 1/4 natural size. 

Emys conybearii. 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. 

x\ssuming 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 bowcr- 
banki (p. 64); and Palaeaspis with the type Emys conybearii (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 howerbanki Owen 
and the correct name for the single species involved will then be 
Palaeaspis howerbanki (Owen). The genus may be rediagnosed as 
follows : 



Palaeaspis Gray 1870 

Type. Emys cony bear H Owen. 

New diagnosis. A flattened pelomedusid resembling Podocneviis 
expansa in shell characters and adult size but without a vertebral keel 
and with a very elongate first neural {ca. 2Y2 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. 

Lydekker, R., and G. A. Boulenger 

1S87. 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, 
pp. 77-79. 

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. 



BREVIORA 

Musemnti 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 William L. Brown, Jr. 

Museum of ronipaiative Zoology 
-Haivaifl University 

The following notes concern a few species belongino- to the 
imprcssa and me tallica groups of the genus Rh yi idoponera 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 subgenericall.y, from Ehytidopo)\era. The present prob- 
lem consists simply of recognizing the specific groupings in ques- 
tion by means of their external characters, chiefiy utilizing the 
workers for study objects, and making the appropriate synonymic 
reductions. The salient characters will be given briefly and 
informally with a vicAv 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 ITnited States Educational (Ful- 
bright) Foundation and of the Parker Fellowship Fund of 
Harvard University. 

Other papers on various groups of RJiytidoponera, intended 
to follow this pattern, are under preparation. 



2 BREVIORA No. 33 

GROUP IMPRESSA 

This closely interrelated series of forms comprises U. impressa 
(Mayr) and its erstwhile varieties with their synonyms. R. 
aenesce^is Emery 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 metalUca group, and generally more slender. 
The head is only slightly longer than broad, and the occipital 
lobes or "ears" are much less perfectly developed than in 
metallica, so that in direct dorsal cephalic ("full face") view, 
the occipital border appears appro.ximately 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 criterion (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 concavity 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 
hnpressa group species. 

Distribution, as known, runs from New Guinea down along 
the mountain chains paralleling the ea.stern Australian coast. 
The normal habitat is rainforest or, in the southern part of the 
range in Victoria, in moist or wet sclerophyll (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. chahjhaea at least will nest directly in the soil or 
beneath rocks. 

When the nest is breached, the workers scatter rapidly and 
hide ; some workers will feign death, but not as readily nor for 
so long a time as in the case of mefaUica. 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 occurs 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 MAVR 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 reseml)ling those of 
the usual ant nest-formation pattern throughout the respective 
dry seasons in the different parts of the range. The impressa 
group species, along with just a few other forms (in-obably 
including R. aspera and R. croe.sus), are exceptional among 
Rhytidoponero 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, which 
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, I 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" \Wth very slight geog-raphieal 
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. im- 
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. chalyhaea Emery, starting prol)ably in the region just 
north of Brisbane on the mountaintops and ranging south at 
least to Melbourne in south central Victoria. Though the char- 
acters are rather slight for the separation of purpurea from 
impressa, they appear to be discontinuous together, and T have 
little doubt that they delimit two distinct species. 

R. chalyhaea, on the other hand, seems to be distinguishable 
from impressa only on the basis of coloration, and a rather ten- 
uous and cabinet-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 Brisl)ane, 
typical blue chalyhaea 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 chalyhaea, 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. 

RnYTiDOPONERA CTiALYBAEA Emery. New status 

Rhytidoponera impressa var. cJmlybaea Emery, 1901, Ann Soc. Ent. Belg.,. 

45: 51, worker. Type locality: New South Wales. 
Ectatomma (Khytidoponera) cyrus Forel, 1910, Rev. Suisse. Zool., 18: 13, 

worker, female. Type locality: Ballina, New South Wales, (W. W. 

Froggatt). New Synonymy. 
Bliytidopancra {Clialcoponcra) cyrus, Emery, 1912, Deutsch, ent. Zeitschr., 

p. 81, discussion of worker characters and relationships. 
Jthytidoponera (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 chalybaea, 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 nsually 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 ehalybaea 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 ehalybaea 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., 
iunder 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. 

Rhytidoponera impressa (Mayr) 

Ectaiomma iiiiprc.s.siini Mayr, 1876, Jour. Mus. Godeffroy, 12: 92, worker, 
female. Type locality: Gayndah, Queensland. 



6 BREVIORA No. 33 

Ehytidoponera {Cholcoponera) 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 occasionally be present on the dorsal surfaces of the body, 
but this is negligible in the material I have seen. I can find no 
other characteristics but the general body color to distinguish 
this form from chaiyhaea {q.o.). The color and the rectangularly 
rounded 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 limit 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 
volcanic soil, much of which has been cleared to grassland for 
grazing. Malanda, a little farther north and occupied by B. 
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 7 

Rhytidoponera purpurea (Emery). New status 

Ectatomrtia impressum, var. purpureum Emery, 1887, Ann. Mus. Civ. Stor. 

Nat. Geneva, (2) 5: 444, worker, female. Type locality: Ilatam, New 

Guinea (Beccari). 
Ectatomma {Rhytidoponera) imprcssum var. apletididum Forel, 1910, Rev. 

Suisse Zool., 18: 12, worker. Type locality: Kuranda, near Cairns, 

Queensland (R. Turner). New Synonymy. 

This species differs from R. chabjbaca, to \vliicli 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, slightly 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 chalyba^a, 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. T 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 scattered rainforest 
patches following the mountains up the eastern side of Cape 
York Peninsula. 

GROUP METALLTCA 

The group of species related to R. metaUica (Fred. Smith) is 
in considerable taxonomic confusion, due chiefly to the high 
degree of geographic 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-describers con- 
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 afflicting 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 amono' the smallest Rhytido- 
ponera, and are distinguished by having an evenly convex ali- 
truncal 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 steei) 
anterior and posterior faces. The treatments of R. aspera and 
R. croesus do not reciuire further general comment. 

Rhytidoponera aspera (Roger) 

Poncra metallica var. aspera Roger, 1860, Berlin, ent. Zeitschr., 4: 308, 

worker. Type locality: Australia. 
Bhytidoponera (Citalcoponcra) aspera, Enieiy, 1912, Deutscli. ent. Zeitschr., 

p. 77, worker, in key. 
Rhytidoponera (Chnlcoportera) arnoldi Forel, 191.5, 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 brilliant 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 al)out 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 correct, 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 Zealantl. 

KiiYTiDOPONERA CROESUS Emery 

Ehytidopoiirro croesiis Emerv, 1901, Ann. Ent. Soc. Belg., 45: .30, worker. 

Type locality: New South Wales. 
Ehytidoponera (CliaJcopon,era) croesxis Emery, 1912, Deutsch. ent. Zeitschr., 

pp. 77, in key, and 81, woi-ker. 
Bhytidoponcra (Clialcoponera) fastiiosa Santschi, 1916, Bull. Soc. ent. 

France, jj. 174, all castes. Type locality: Australia (Duchaussoy), 

probably New South Wales. New Synonymy. 
CJwIcoponera victoria (sic) var. andrei Wheeler and Chapman, 1925, 

Philippine Jour. Sci., 28: GO, worker. Type locality: cited in error as 

from the mountains of Victoria; recte Dorrigo. New South Wales (W. 

Heron). New Synonymy. 
B. croesns 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 described as faiituosa, most of the head, alitrunk, petiole 
and gaster is densely purple in color, whereas intermediates like 
the types of the original croesns 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 comparLson with croesus, which species he credits 
to F'orel as author. It seems that his comparison is really made 
against cynis Forel (cJialyhaea Emery, see above), and in this 
coniKCction ^lis description makes more seiLse. The variety 
amlrei 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 eotypes. 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. "'Vicforiae" 
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: BuUi 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). R. croesus is apparently re- 
stricted to the wet subtropical forests of eastern New South Wales 
and extreme southeastern Queensland. 



R E V I 

MinseiMM of Comparative Zoology 

Cambridge, Mass. July 20, 1954 Number 34 

A REVIEW OF THE COXALIS GROUP OF 
THE ANT GENUS STICTOPONERA MAYR 

By William L. Broavn, Jk. 

Museum of Comparative Zoology 
Harvard University 

The genus Stictoponera is a chiefly Oriental and Papuasian 
assemblage of tj^pical 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 Wheeleriporie Mann and Rhopalopone Emery. 
With the new composition, titictoponcra 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 distinct tooth on the dorsum of the posterior 
coxa. These species are the "typical" members of Sticioponcra 
related to the genotype, 8. coxalis (Roger). The species formerly 
called Wheeleripo7ie alhiclava 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. 



2 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. hiroi Emery. 

B. Second postpetiolar (IV abdominal) segment not costate. 

1. Ej'es placed anterior to middle of sides of head (ac- 
cording to original descriptions) : 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. hinghamii Forel, 8. 
laevior Forel, 8. 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 (Stioioponera) laevins Forel, 1905, Mitt. Naturli. Mus., Ham- 
burg, 22: 7, worker. Type locality: Tjibodas, Java. 
Stictoponera laevior var. avia Forel, 1912, Notes Mus. Leyden, 34: 96, all 
castes. 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 stylaia Menozzi, 1925, Philippine Jour. Sci., 28: 440, pi. 1, figs. 

a, b, worker. New Synonymy. 
Stictoponera menaden,^is var. obscura Santschi, 1932, Mem. Mus. Hist. Nat. 

Belg., (hors serie) 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 Sticioponera in the East Indies and the Philippines. It 
occurs on Java and Sumatra together with 8. hicolor, 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 localities, and records from 
the mainland are scarce in the literature. From the present data, 
S. menadensis appears to be a peripherally distributed form in 
the process of being replaced by S. hicolor, 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 description mentions no characters dif- 
ferentiating his variety from normal menadensis populations. 
The type of 8. 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 Sticioponera 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. McGregor). 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. S. 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) hicolor Emery, 1889, Ann. Mus. Civ. Stor. Nat. 

Geneva, 27: 493-494, worker. 
Ectatomma (Stictoponera) bicolor var. minor Forel, 1900, Jour. Bombay 

Nat. Hist. Soc, 13: 316, worker. 
Stictoponera menadensis subsp. minor, Brown, 1948, Psyche, 54: 264, 

(teneral) worker. 
Stictoponera hicolor, 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'. mefiadensis, and has the same head shape, with prom- 
inently projecting posterior occipital lobes or "ears" and a cor- 
responding, deepl}' 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 iu 
menadensis, with some series averaging a trifle smaller, but the 
color of the alitrunk is lighter and brighter, varying shades of 
orange-ferruginous. The head varies from about the same color 
as the alitrunk to pieeous, the darker color being more common. 
The median smooth strip of the menadensis mesonotum is re- 
placed in hicolor 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 STTCTOPONERA MAYR 5 

Like S. menadensis, the present species has often been mis- 
identified. It is still possible, of course, that menadensis and 
hicolor are mere geographical color representatives of one species, 
but from the present material they can still be separated. The 
record of hicolor from the Philippines is based on a male of un- 
certain species ; no authentic hicolor records are yet known for 
these islands. 

I have reviewed material from the following localities : Indo- 
china: 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 (L. G. E. Kalshoven). 
Hong Kong: no further locality (Ris). Emery described this 
species from material taken in various Burmese localities, both 
upcountry and in Tenasserim. 

Stictoponera bingiiamii Forel 

Eotatonvma (Stictoponera) hinghamii Forel, 1900, Jour. Bombay Nat. Hiat. 

Soc., 13: 137, worker, female. Type locality: Burma [Pegu Yoma?]. 
Stwtoponera horn,eensis 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 hinghamii from horneensis, 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 case, 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 allometric 
extremes of one form. I have also seen material referable to 
the same species from Tutu River, North Borneo (Mjoberg) and 
a specimen from "Pedada-B., Lampongs, Sumatra," unknown 
collector, and I feel sure that Forel's Burmese and Emerj^'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) binghnmi subsp. crassicornis Forel, 1912, Zool. 
Jahrb. Syst., (suppl.) 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 
hmghmnii very decisivel3^ 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 case, it is very unlikely that crassicornis can be 
a race of S. binghamii, 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 
Stictoponfira spiralis Karawajew, 1925, Konowia, 4: 79-81, worker. 

This species, also described from Sumatra, reads very much 
like S. crassicornis in what seem to be the significant features, and 
it is possible that the two names are synonymous. 

Stictoponera biroi Emery 
Stictoponera biroi Emeiy, 1902, Term. Fiizetek, 25: 154, worker. 

8. 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 7 

Stictoponera posteropsis Gregg 
Stictoponera posteropsis Gregg, 1&52, Psj'che, 58: 77-8§, figs. 1, 3a, 3c, 
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. 

Geneva, 27: 494, worker. Forel, 1900, Jour. Bombay Nat. Hist. Soc, 

13: 316, 317, worker. Bingham, 1903, Fauna Brit. India, Hym., 2: 

83, worker. 
Ponera rugosa Fr. Smith, 1857, Jour. Proe. Linn. Soc. London, Zool., 2: 

6^, worker; name preoccupied by Le Guillou, 1840. New Synonymy. 
Stictoponera costoda Emery, 1901, Ann. Mus. Civ. Stor. Nat. Genova, 40: 

662, worker. 
Stictoponera costata var. unicolor Forel, 1901, Eev. Suisse Zool., 9: 335, 

worker, male. New Synonymy. 
Stictoponfira rugosa var. parva Forel, 1913, Zool. Jahrb. Syst., 36: 6, worker. 

New Synonymy. 
'i Stictoponera costata var. simalurensis Forel, 1915, Tijdschr. v. Eut., 58: 

23, worker. New Synonymy, with doubt. 
Stictoponera costata var. pinealis Wheeler, 1929, Boll. Lab. Zool. Portici, 

24: 31. New Synonymy. 
Stictoponera wallacei Donisthorpe, 1932, Ann. Mag. Nat. Hist., (10) 10: 

447, noDi. pro F oner a 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 sj-nonymy with Ponera 
rugosa of Smith. He also cites differences between costata and 
a type of coxalis he received from the Koger 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 BREvroRA No. 34 

tion of these two species, as will be seen below under S. coxalis. 
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 gastric (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 
pineaUs Wheeler, from Penang, is deep reddish with the gaster 
black. The variety imicolor 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 
color 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 likely that Forel 's variants parva 
and simahirensis 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 cost ate second gastric (second 
postpetiolar or IV abdominal) segment, is in the form of large, 
rather deep punctures, foveae or pits, with more or less distinct 
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 
discal 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 punc- 
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 supplies a fnll set of intergrades w^hich is enough 
to obscure, any j)ossible taxonomic split based on this character. It 
therefore seems that Emery was correct in maintaining cosfata 
as a single taxonomic unit despite the variation he saw. His 
reasonable suggestion that F. Smith's rngosa, from Sarawak, 
equalled costata is also accepted here, and since the name rngosa 
is preoccupied, and Donisthorpe's noineri novum wallacei 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; S. Slau ; Brooketon ; Pajan; S. 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 Eoger, 1860, Berl. ent. Zeitschr., 4: 308, worker. 
Ectatomma (Stiotoponera) coxale Forel, 1900, Jour. Bombay Nat. Hist. Soc, 

13: 316, worker. Nee Bingham, 1903, Fauna Brit. India, Hym., 2: 

84, fig. 44, worker. 
Stictoponera coxalis Emery, 1901, Ann. Mus. Civ. Stor. Nat. Geneva, 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 coxalis 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 Maj^r. 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, coxalis 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 
eostae 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 include 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 S. menadensis and S. hicolor. 
When better collections are available from southern India, it may 
be necessary to re-examine the relationship of coxalis and costata. 

Note on " Stictoponera sauteri" 

The name Stictoponera sauteri (Chapman and Capco, 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 
Ectomomyrmex 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. 



R E V I O R A 

Mnaseitim of Coimparative Zoology 

CAMHRIDdE, .AfASS. JuLY 2S, 1 (KM Xl-.MBER 35 

NEW OR REI)E8(JRIBED PELOMEDUSID 
SKULLS FROM THE TERTIARY OF 

AFRICA AND ASIA 
(TESTUDINES, PELOMEDUSIDAE) 

1. Dacquemys paleonwrphd. new oenus. new species from the 
Lower Oligocene of the Fayuni, E<iyi)t. 

By Ekxp:st Williams 

In 1912 E. Dacque descrilunl a sknll from the Lower Oligoeene 
of the Payum of Eo-ypt which he referred, with doubt, to Stereo- 
genys lihyca Andrews. The skull, which is the property of the 
Staatliches Museum fiir Naturkunde in Stuttgart, has survived 
the war. Thanks to the kindness of the authorities in Stuttgart, 
and especially of Prof. P. Berckhemer and Dr. Karl Dietrich 
Adam, I have lieen able to re-examine it. 

Re-examination confirms the excellence and accuracy of 
Dacque '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 Podocnemis 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 paleonwrpha, new species. 

The genotype is at present the only species. 



No. 35 



DaCQUEMYS PALEOMORPHA, new species 



Type. No. 12645 in the paleontologieal collection of the Staat- 
liches Museum fiir Naturkunde, Stntt^art, a skull without 
mandible. 

Type locality. Diieh, Fayum of Eiivpt. 

Horizon. Lower Oligocene. 

Specific diagnosis. With the characters of the genus. 

Description. A moderately elongate skull (estimated condylo- 
basal length 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 
groove. 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 joining 
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 mandibuli, 
and curving medially to meet the backward prong of the 
opisthotic 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- 
ing (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- 



1954 DACQUEMYS PALEOMORPIIA 3 

rock, 1!)02, siicli as is pr'eseiit in some livinu- species of Podocnemis. 
The premaxillae perhaps forming a hook ; if so the hook eroded 
in the specimen. Nares much broader than high. Two strong 
ridg(^s on tlie triturating surface of the maxilla, the ridges uniting 
aiitei-ioi'ly 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 strongly inclined backward. 
The hypertrophied "carotid channel" smaller than in many 
Podocnemis but clearly specialized in the fashion peculiar to this 
group within the pelomedusids. The basisphenoid broad, sub- 
pentagonal, a long obli((ue lateral suture jjctween quadrate and 
basiocci]iital. The occipital condyle missing in the specimen. 

I)lsci(ssi())i. It is very easy to show that the skull here named 
Dacqucmijs palcomori^ha cannot belong to the genus Stereogenys. 
The type of Stereogenys is S. cromcri {V . Eocene of the Fayum) 
(Andrews, 1906), and the type of that species is a skull that 
diit'ers more from Dacquemys than Dacquemys does from some 
of the species of Podocnemis sensu lata. I tabulate the major dif- 
ferences below. 

Stereogenys cromeri Bacquemys paleomorpha 

Skull very broad, nearly as broad as Skull moderately elongate, one and 

long one half times as long as wide 

Squamosal not in contact with parie- Squamosal in broad contact with 

tal parietal 

Orbits exposed dorsally Orbits not exposed dorsally 

Secondary palate greatly developed, Only an incipient secondary palate 

formed by a long median suture of formed by the median union of the 

the palatines, the maxillaries not maxillaries behind the premaxil- 

meeting medially or meeting for a laries 

short distance only 

Internal choanae therefore well be- internal choanae nearly at posterior 

hind the orbits level of orbits 

Maxillary tritui'ating surface with- JNIaxillary with two ridges 

out ridging 

Pterygoids {fide Andrews 1906) con- Basisphenoid exposed ventrally in 

ceal basisphenoid ventrally normal fashion 



4 BREVIORA No. 35 

The differences here summarized and also the oreat general dif- 
ference in habitus go far beyond the amount that may be ad- 
mitted within a genus of turtles. Dacque was indeed aware of 
these differences (cf. p. 310 his paper) but he seems to have 
believed 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. 

Quite as surely as it cannot belong to Sfereoycnifs, the skull 
here discussed cannot belong to Podocneiiiis or to any other pelo- 
medusid genus in which the skull is known. 

From Shivehoemys (Swinton, 1939) it differs in much the 
same ways that it differs from Stereogenys. From Podocnemis in 
a restricted sense (type P. expansa 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 Erymnochelys (type and only living species E. 
niadagascarensis) 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 
l)alate, by the shallowness or absence of a precolumellar fossa as 
w^ell as by the squamosal-parietal contact and probably also by 
the lateral emargination of the temporal region. 

From Pclonicdusa, Pclusios and Cartereniys (Williams, 1953), 
it is sufficiently distinguished by the covered temporal region and 
especially the squamosal-parietal contact, but from at least 
Pclusios and Pelomedusa it differs in th(> characters which it 
sliares with the Pad'tcncmis group of genera, to which it must, 
indeed, he referred — the enlarged "carotid channeT' and the 
l)asioccipital-qnadrate contact etc. From Boihrey)iys (Hay, 1908) 



1954 DACQUEMYS PALEOMOBI'lIA 5 

it differs as radically as from Sfcrctxjciii/s c.o'. in ]a('kiii<|- tiic 
extraordinary pits in the maxillae so characteristic of Bt^tlncnn/s, 
in the mnch narrower skull, in the absence of a vomer, etc. 

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 described 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 it 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: Podocncmis fajuniensis Andrews, i^tercodcnijs 
libyca Andrews, and Pchjiiudusa progaleata v. Reinach. 

If Pdomedusa progaleata (material re-examined by Dacque, 
]912) is correctly assigned and there appears to be sufficient rea- 
son to think that it is, D. paleomorpha cannot I)e synonymous 
with it, since the skulls in the two genera are so very different 
that they do not belong even in the same subsection of the 
Pelomedusidae. 

Unless the Dacquemys skidl belongs to some variety of shell 
still undiscovered, decision must then lie between Podocnemis 
fajumensis and Stereogenys lihyca. This task is difficult, requir- 
ing a rather complex and tenuous argument. 

The argument may be stated in this fashion : 

The shell of Podocnentis fajiiiiK iisis differs so little from that 
of Recent EryninocJielys itiadagasravensis that, considering the 
shell alone, the distinctness of the fossil species might be cpies- 
tioned. The skvill of the fossil form might therefore be expected 
to be very like that of the Recent species, and in fact the Dae- 
qneniys skull is more similar to the skull of Erymnoehelys 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 j^ossiliilty worth con- 
sidering that the Dacque skull may belong to "Podocnemis" 
fajurnensis. 



6 BREVIORA No. 35 

111 the alternative case of Stercofjcin/s lihijcd the sliell whieh is 
the type of this species is clearly very similar to that referred to 
S. cromeri. The argument that the shells referred to S. cromeri 
really belong to that species is plausible even though not con- 
clusive. Andrews (1906, p. 298) remarks: "Although in no case 
have the carapace and plastron be(Mi found associated with the 
skull in such a manner as to leave no doul)t 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 Fodooifiiiis. the only other Pleu- 
rodiran genus found at this horizon." Accepting Andrews' in- 
terpretation of the situation, if Sfereugcnys libyca is really 
closely related to N. cromeri, it should have a skull to some degree 
resembling that of N. cromeri. The Dac(|ue skull is so very dif- 
ferent from that of *S'. cromeri that it is very unlikely to l)e the 
skull of »S'. lihyca. 

Thus the Dacque skull is unlikely t;) lie that of S. lihiica and 
might be that of P. fajumen.sis. But to assert the truth of the 
latter suggestion confidently w^ould be a bold venture. In the next 
paper of this series T 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 distinct 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 witli 
Dacquemys paleomorpha, P. fajumensis and *Sf. lihyca an anterior 
portion of mandible which he has referred to P. fajumensis. His 
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 photogra])h of tlris mandibular 
fragment made by Peter CTreen and published ])y permission of 



I 



1954 DACQUEMYS PALEOMORPIIA 7 

the Trustees of the IJritisli .Museum. It will I)e seen that the 
ridoiuo' of this niaiiilihh' wouh] fit well with the maxillary ridging 
of Dacqueuii/s pal(<jni(>rj)li(i. liut there is no strong hook at the 
mandibular symphysis such as seems to be implied by the deep 
premaxillary form of the skull. In this ease, as in the previous 
one, only discovery of associated material can verify the reference 
to genus and sjiecies. 




Fig. 1. Dorsal view of niiiiidil)lt> lefened Ijy Andrews to Prxlooieinis 
fajiniu'H.si.s. B:M (Nil) R ."wUG, from tlie Lower Oligoceiie of tlie Fayuni, 
Egypt, xl. Photograiili reinodiiced l)y periiiissioii of tlie Trustees of tlie 
British Museum. 



The photographs of the skull of Dacqucmijs paleomorpha were 
made by the Museum fiir Naturkunde in Stuttgart and 1 am very 
grateful to the authorities there for the privilege of reproduciiig 
them. 



8 BREVIORA No. 35 

PAPERS CITED 

Andrews, C. W. 

1906. A desc-iiptive catalogue of the Tertiary vertebrata of the Fayum, 
Egypt. London. 324 pp. 

Dacque, E. 

1912. Die fossilen Schildkroten Aegyptens. Geol. Palaeont. Althandl., 
vol. 14, pp. 275-337. 

Hay, O. p. 

1908. The fossil turtles of Xortli America. Carnegie Inst. Publ. No. 75. 
568 pp. 

Siebenhock, F. 

1902. Zur Systematik der Schildkrotengattung Podocnemis Wagl. Sitz. 
Ber. Akad. Wiss. Wien, math. nat. Kl., Abt. 1, vol. Ill, 
pp. 157-170. 

SWINTON, W. E. 

1939. A new fossil fresh-water tortoise from Burma. Ree. Geol. Survey 
India, vol. 74, pp. 548-551. 

Williams, E. 

1953. Fossils and the distribution of chelyid turtles. 1. " Hydraspis 
leithii" in the Eocene of India is a pelomedusid. Breviora, No. 
13, pp. 1-8. 




J'lnti' 1. 'I'ypc skull (if l><ii-<iii< hijis jKih nnini jilni, new unins, urw spi'cies. 

Stajitlii-lu's AIusi'uiii fill- Xiituikiiuilc, Stiittt;;nt Xo. ll^(U."l. Aliuve: ventral 

vieAv; Ij^'low: dorsal view. I'lidto^raiihs ri'iiroiliu-ed \>y courtesy of the 
•Staatliches ^Museum fiir Xaturkunde. xl. 



E V I O R A 



MinseiLiitjni of Comparsitive Zoology 



CA:MBKinGK, ^[\> 



August (i, 19.")4 



Number 36 



PRESENT KNOWLEDGE OF THE SNAKE 

ELACHISTODON WESTFAUIANNI 

REINHARDT 

By Carl Gans 

Carnegie Museum, Pittsl)urgh, Pa. 

and 
Erxest E. Williams 

Museum of Compaiative Zoology, Camlnidge, Mass. 



INTRODUCTION 

In 1863 Johannes Reinharclt described a peculiar snake belong- 
ing to the new genns and species Elachisiodon ivestermanni, the 
specific name referring to the collector, William Westermann, 
who obtained the initial specimen in Rnngpore (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 liy 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 Elachi- 
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 artificialitv of this categorv. 



No. 36 



Since 1863 only four additional specimens have been reported 
and there has been little 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 Elochisfodon would be especially 




Fig. 1. The distribution of Elachistodon 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, l)een 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 Malcolm 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 ELACIIISTODON 3 

This iJciper, tlieivfore, lias two oljjects. Fii-st, it is iiit<^n(l(Ml to 
provide a recapitulation of the cxistiu-i' data on E'dcltistodon in 
the light of the findings in the ucinis Ddsiipcltis, and secondly 
it is desired to reawaken interest in Elacliistodoii in the hope of 
obtaining the additional specimens upon which the more intensive 
studies might be based. 

HISTORICAL RESUME 

The initial sjiecinien was rather fully described by Reinhardt 
(1863, p. 198), whose careful investigation of its internal anat- 
omy was luidertaken l)ecause the enlarged vertebral scales of the 
new form had left him in doubt as to its taxonomic placement. 
He prepared the tooth-bearing elements of the right side of the 
skull and one of the mandibles, and checked 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 ]-)alatine and pterygoid as well as teeth and 
minuscule fangs on the maxilla and described the shape and 
appearance of the dcMitary. in comparing Ehichishxluii with 
Ddsjiix Ills he pointed out that in both there were two tyjX's of 
penetrant liypapo])hyscs. though both types in the new form were 
less well-developed. He figured the palatal and mandibular 
elements of both genera (see Fig. 1). 

The stomach and gullet of the type specimen of E. ivestennainii 
were found to be filled with an amorphous congealed mass which 
analysis indicated might be either milk or egg 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 tlie theory that the species w^as an egg-eater. 

Reinhardt also described in detail the squamation of Elachisto- 
don, mentioning the very ])eculiar 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 E. westermanni. Reinhardt placed 
the new genus in the family Rachiodontidae {sic). 

The next to refer to Elachisfodon was Giinther (1861, p. 414) 
who mentioned it on Reinhardt 's authority only. Pie 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 
Dasypeltis. 

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. 356-357) 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 listing the snakes in the Indian Mu- 
seum recorded a specimen from Bengal which seems to have 
been that earlier 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 (1913, 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 described 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 ELACTIISTODON 5 

tional specimens obtained hy Travers at Baradiulii ^ ajipeared 
to be unknown. 

Malcolm Smith (1943, p. 4m) redesci-ibed the uvnus and 
removed it from the Elaehistodontinae to tlie Dasypeltinae. lie 
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 
Plarderian gland is enlarged in this form as in Dux fi pel tin. 

MORPlIOLOdY 

(ieneral 

The following condensed diagnosis is designed to aid identifica- 
tion of specimens. For this reason it follows the summary pre- 
sented l)y Smith (1943, p. 404) with only minor modifications. 

Head fairly distinct from neck ; eye large with vertically ellipti- 
cal pupil (round fide Reinhardt) ; a large pit in the posterior 
part of the nasal shield. Body moderately elongate, feelily com- 
pressed. Tail short, subcaudals 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 ; H or 7 supralaliials, the third and fourth 
touching the eye ; 2 pairs of sublinguals. Scales smooth, in 15 
rows, 19 on the neck, the vertebral series much enlarged, 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 light 
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. 

9 784 nun. (670 + 114 mm.). Ventrals 217. Caudals 59. Rung- 
pore, Bengal. Copenhagen Museum (Reinhai'dt ). 

1 We have been unable to find Baradijihi on any map. but the Imlijin Consuhir 
Office in New York has very kindly informed us that thtTo is a railiuad station 
by that name about 30 miles north of Calcutta. 



BREVIORA No. 36 

800 mm. (670 + 130 mm.) fide Smith, (762 mm. [635 + 127 
mm.] /;V/f Wall). Ventrals213 (210). Caudals 62 (64). Near 
Mai, Jalpaiguri District. Formerly Bombay jMuseum, now 
British Museum (Natural History). 

222 mm. (186 + 35.7 mm.), Ventrals 208. Caudals 63. Bihar, 
Purnea. Indian Museum (Blanford). 




Fig. 2. E. iref<trr))ia)inl. 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 si<iiiificant integumentary similarities between Elach- 
istodon and Dasypeltis are the structure and arrangement of the 
gular and cheek scales. This similarity Avas 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 ElacMstodon is correlated wuth similar habits. 

In both Elachistodon and Dasypeltis there are two pairs of 
sublinguals closely joined and lacking am^ evidence of the median 
groove that in most snakes provides for expansion. In Dasypeltis 



1954 ELACIIISTODON 7 

there are no true <>ulai' scales at all, while in Eluchistodou there 
is but one row of these. In both of these forms the sublinouals 
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 ease 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 ditference between Elachistodon 
and Dasypeltis is the ]n-esence 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 cei-tain species of Pythonidae has 
been determined to be that of a thermosensitive range-finder 
(Noble and Schmidt, 1987), and it is known that similar struc- 
tures of as yet uninvestigated function exist in other forms. In 
Elacliisiodon nothing is known of the function of the pits. 

Scpiamation of the Posterior liody 

The enlargement of the vertebral series of dorsal scales char- 
acteristic of Elarhistodo)! 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 conjunction with 
the extreme lateral flattening of the body it might impart a 
stiffening etfect. 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 climbing members 
of the Dipsadinae. We cite the suggestion here without passing 
judgment upon its applicability to Elachistodon, which shows 
but feeble lateral compression. 

It is also to be noted that the scales of Elachislodnu ai-e smooth 
while those of Dasypeltis are strongly keeled. This character is, 
however, of little value at the generic level or above, in view of 



8 BREVIORA No. 36 

the number of eases in snakes in whieh it shows intrageneric 
variation. 

Elachiatodon e.xhibits no traee of the size reduetion, 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 pul)lished observations are available on gum structure. 
Examination of the British Museum specimen indicated that the 
gum ridges are probably covered with a somewhat less convoluted 
mass of tissue tlian are those of Dasypeltis. 

Smith and Bellairs (1947, p. 362) have noted that the Harder- 
lan gland is notably enlarged in EJachisiodon paralleling the 
con .ition in Dasypeltis. 





Fig. 8. E. wesiermanni. A, Palatal liones ; B, anterior hvpapopliysial 
vertebrae; C, posterior hypapopliysial vertebrae. Ablireviations: e, et-to 
pterygoid; mx, maxillary; pal, palatine; pi, ptergygoid. After Malcolm 
Smith (']943). 



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 9 

US (Williams) found the fifjures accurate in all particulars includ- 
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 Dasypelfis in- 
dicates few differences beyond the presence of the relatively 
enlarged grooved teeth on the posterior aspect of the maxilla in 
Elachistodoti. 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 Ddsifijeltis. The wing- 
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 l)y 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 (1043) — botli 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 Elavil ishxhnt and Dasypeltis. The appearance 
of the element in Elachisfodoii is very strange, and since the fig- 
ure of the same element in Dasy pelt's liears veiy little resem- 
blance to actual specimens seen by us, we do not care to connnent 
further in this matter (see Fig. 4). 

\%'rte))ral Cohunn and Eso]>hagus 

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 Elachistodon. 
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 egg glides along these 
runners during ingestion, but the runners do not serve any cut- 
ting or sawing function. The egg is pierced by the elongate, 
spiniform, forward pointed hypapophyses of the posterior series, 
wliicli penetrate the esophagus in all specimens (Gans, 1952, p. 
236). After the shell has been broken, the hea\y rounded boss- 
like hypapophyses of the intermediate series transmit a crushing 
force to the egg 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 postzygapophyses being laterally displaced 
and extended by spiniform processes. This offers additional sur- 
face for the action of the dorsal musculature wliich 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 Das i/ pelt is, 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, possil)ly 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 (1943, 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 (1863, 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) 






Fis'. 4. Plate from tli.- (ni^iii; 
ventral ami side views of the In 
E. tn.sh iiiKiKiii : K, (leiitary 



I (les.-iiptioii ]iy Heiiihaidt. J, /.', (', Dorsal, 
ad of K. irrstrniKiiiin : J), jialatai Ijones of 
)f E. W(.^trniuunii ; E, palatal bones of 



Dasypeltis ticabcr; G, mandible of DiusypcUia scab 



12 BREVIORA No. 36 

has examined these vertel)rae, and the t'ollowinji' 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 inclina- 
tion of the zygapophysial surfaces is apparent. 

In some ways the anterior hypapophyses of Elachisfodon are 
reminiscent of stages in the ontogeny of the intermediate hypa- 
pophyses in Dasypeltis, though th(> mounting on a pedicel sug- 
gests that they may be specialized in a different direction. Rein- 
hardt (1863, p. 203) stated that there were nine such anterior 
hypapophyses, while Smith (1943, p. 403) counted eighteen 
all of which projected through longitiulinal slits into the esopha- 
gus. Both authors believed that these are cutting instruments, 
which hardly seems likely 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 Elachistoelon the hindmost hypapophyses are block- 
shaped spines whose major specialization lies in the fact that 
they are directed forward rather than backward. They most 
nearly resemble the hypapophyses of several of the s])ecies of 
Elaphe 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 findings 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 dift'erences. 



1954 ELACHISTODON 13 

111 spite of this, it iiuiy he stated tliat all indicalions exist that 
eg-os are opened h_v I'JhicliisttxIon in a nmiiner siniilar to that 
eni]iloy(^d by Dasi/ix His and Elaphc climacophora, i.e. by exertin<i- 
a force eoneentrated by Ihe processes of the vertebrae. 

A comment may he made on Smith's Figure C (our Fig. 8 C). 
The third figured liyi)apoi)hysial carries a hypapophysis with 
what appears to be a shovel-shaped tij). While the drawing is 
entirely accurate, its two-dimensional nature does not do the 
subject justice, as this ti]) is actually defoi-med 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 conclusion in 
Ehichisiodoii. 

BIOLOGY 

No specimen of Elaclnstodon has ever been kept in captivity 
and there exist no notes on habits or habitats in the wild. With 
the exception of Reinhardt's specimen which had egg yolk (f) 
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 "eggs but not exclusively." 
The presence of grooved teeth and the nasal pit suggest that 
the species may feed also on live and homoiothermous prey. 

The only other item of interest concerns breeding habits. The 
type was a gravid female, containing seven (-tO x 11 mm.) eggs 
(Reinhardt, 1863, 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 



No. 36 



Comparison of Elaclilsfodon and Dasypcltis 

The following tables sum up the more important similarities 
and differences — at present known — of the two genera. 

Points of Similarity 

fSul)lingual shields fused, mental groove absent ; special pro- 
vision for expansion of the skin along 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- 
ing the esophagus; anterior hypapophyses generally rounded, 
with sledge-like runners or bosses ; posterior hypapophyses de- 
veloped as foi'ward-pointing spines ; oviparous. 

Points of Dift'erence 



Dasypeltis 

Nasal pit absent 

No gulars 

Scales strongly keeled 

Vertebral scale row subequal to dor- 
sals 

Some lateral rows with scales re- 
duced in size, inclined and with 
serrate keels 

Maxillary teeth minute and equal 

Intermediate hypapophyses heavy, 

rounded, boss-like, closely applied 

to centra 
Posterior hypapopliyses bear sharply 

pointed spines and penetrate 

esophagus 
Elliptical pupil 
No pterygoid teeth 



Elacliistodon 
Nasal pit present 
One row of gulars 
Scales smooth 
Vertebral scale row enlarged 

All lateral rows with scales of equal 



1-2 enlarged grooved teeth on rear 
of maxilla 

Bosses of hypapophyses separated by 
constriction from centrum, i.e. 
mounted on pedicel 

Posterior hypapophyses much less 
strongly developed, ? no penetra- 
tion 

? Bound pupil {fide Eeinhardt) 

? Pterygoid teetli {fide Reinhardt) 



1954 ELACHISTODON 15 

Inspection of the above lists reveals that the similarities be- 
t^veen the two forms are primarily those of striietiires which in 
Dasypeltis are known to l)e connected with egg' ingestion. Many 
of the diiTerences are at a level generally associated Avith a generic 
separation, while only the nasal pit and the grooved fangs might 
indicate a higher level of difference. 

It has been suggested that Ehicli'shxhtii may be a less special- 
ized form than Dasifprlfis 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 fangs as w(dl as the additional 
modifications exhibited t)y this genus. Analysis of the relation- 
ships of the genus ElacJiistodoii may indeed yield evidence shed- 
dinu' light on the genei-al jjroblem of the origin of the opistho- 
glyphs. 

Beyond the two genera which have been compared here, analy- 
sis will haAc 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 rei)orted foi- the 
most part the observations of others and have had little ojiportun- 
ity to check these a'^ainst actual specimens. It may well be that 
careful examination of a large series of specimens would force 
a revision of our understanding of the morphology of Elachisto- 
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 EJacJiistodon. It would obviously be dangerous to 
speculate at tliis time on the larger problems offered by tlu^ Indian 
genus. 

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. 

Acknowledymenis. Our thanks are due to a consideral)le num- 



16 BREVIORA No. 36 

ber of peojile Aviih 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 EJachistodon. We are indebted 
also to the authorities of the University of Michigan Museum of 
Zoolotry for the use of library facilities while in Ann Arbor. 

BIBLIOGRAPHY 

Blan-ford, W. T. 

1875. Note on (i) Elachistodon westermanni, (ii) Platiccps semifascia- 
tus, and (iii) Ahlei^harus pusillus and Bl.epharosteres agilis. 
Jour. Asiat. Soc. Bengal, vol. 44, pt. 2, pp. 207-209. 

BOULENGEB, G. A. 

1890. Reptilia and Batrachia. /» The Fauna of British India, including 
Geylon and Burma. London, xviii t -" H pp. 

1896. Catalogue of the snakes in tlie British iMuseuui (Natural His- 
tory). Ijondon, vol. H. xii, f 7L'7 pp. 

Cope, 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 DasypcUis. Zoologica, vol. 37, pt. 4, pp. 209 244. 

Gans, C. and M. Oshima 

1952. Adaptations for egg eating in the snake Elaphe cllmacopliora 
(Boie). Amer. Mus. Novitates, No. 1571, 16 pp. 
Gij'nther, a. 

1864. The reptiles of British India. London, xxvi + 4.12 pp. 

Noble, 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. 

Peters, J. A. 

1952. The snake subfamily Dipsadinae in South and Central America. 
University of Michigan Doctoral Dissertation, 369 pp. 

Reinhardt, J. 

1863. En ny Slaegt af Slangefamilien Rachidontidae. Oversigt. K. 
Danske Vidensk. Selskab., pp. 198-210. 



1954 ELACHISTODON 17 

SCLATER, W. L. 

1891. List of the snakes in the Indian Museum. Calcutta, x -f- 79 pp. 

Shaav, G. E., E. O. Shebbeare, and P. E. Barker 

1941. The snakes of northern Bengal ami Sikkiiii. Jour. Bengal Nat. 
Hist. Soc, vol. IC), pp. .")7-li7. 

Smith, M. A. 

1943. Reptilia and Aniphil.ia. //( The Fauna of British India. London, 
vol. 3 (Serpentes), 583 pp. 

Smith, M. A. and A. d'A. Bellairs 

1947. The head glands in snakes with leniarks on the evolution of 
the parotid gland and teeth of the Opisthoglypha. Jour. Linn. 
Soc, vol. 41, pp. 351-3(58. 

Wall, F. 

1913. A rare snake, EJachistodon in stt riiiaiini from the Jalpaiguri 

District. Jour. Bombay Nat. Hist. Soc, vol. 12L', p]i. 400-401. 
1923. A handlist of the snakes of the Indian Empire. Part. 111. Jour. 
Bombay Nat. Hist. Soc, vol. 29, pp. 8*;4-87S. 



E V I O R A 

Miiaseiiiinni of Compsirative Zoology 

Cambridge, Mass. August 18, 1954 Number 37 



ON THE EVOLUTION OF AN ORIENTAL 

EARTHWORM SPECIES, PHERETIMA 

ANOMALA MICHAELSEN 1907 

By G. E. Gates 

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 v-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 hermaphroditic 
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. iypica 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 Bunna, 
as for instance in the Kamaungthwe River region of Tavoy district. 
In two collections from that area, taken two months apart, cen- 
tralis, typica, insolita and the intermediates were represented by 
0+0, 377+69, 11+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- 
graphically, Karenni quite possibly is nearer the region where 
anoynala 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- 
cally. 

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 insolita 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 spennathecae 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 insolita 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 cefitralis 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 
anarsenosomphic worms can be expected to reproduce in the same 
way as hisolita. 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 insolita and ce«- 



4 BREVIOBA 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 F2 generation if "in- 
solita" is dominant to "centralis". The Mendelian relationship, 
if such it be, is however recognizable only because the mushroom 
glands of centralis have been carried over into the insolita pheno- 
type. The centralis phenotype like-wise 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 
glands 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 insolita 
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 finst 
step, in the evolution of anoniala 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- 



6 • BBEVIORA NO. 37 

ductioii, a parthenogenetic mutation appears to be fairly common. 
Thoug-h recognized in anoniala 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. diff ring ens (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 tliird 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 athecal 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- 
iima, 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 like centralis, a mutation, 
" aprostatic ", 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 g^enus. 

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 viii 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 occa- 
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 — can 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 BEEVIORA 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. Each 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. 



E V I O R A 

Mmseimm of Coimparative Zoology 

Cambridge, Mass. November 28, 1954 Number 38 

OMNIA (TRILOBITA) FROM VENEZUELA 
By H. B. Whittington 

In 1938 Leith clescribed a Middle Ordovician fauna from the 
Venezuelan Andes which included the graptolite Dicranograptus, 
a cryptolithinid trilobite, and fragmentary brachiopods and pele- 
cypods. Recently 1 received several specimens of the trilobite 
collected from the orifjinal locality by Dr. G. R. Pierce, of the 
Creole Petroleum Corporation, and sent to me by Dr. V. D. 
AVinkler and Mr. K. 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. Waage, 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 TRINUCLEIDAB Emmrich, 1884 
Subfamily CRYPTOLITHINAE Whittington, 1941 

Genus OnNIA Bancroft, 1933 

ONNiA TERRYi (Leith, 1938) 

PL 1, figs. 1-7 

Ilolotjjpe. 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 Miicuchachi (State of Merida) " (Leith, 1938, 
p. 338). 

Additional M(iteri(d. 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 Ordovieian (Leith, 1938). Later collections have 
not included additional species that might throw further light 
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, 
convex, long median spine, the shallow occipital furrow with 
deep appendiferal pits laterally. Immediately in front of the 
occipital furrow the glabella appears to be exceptionally inflated 
where it is crossed Ijy the occiput (PL 1, figs. 2, 5), and in front 
of the occiput glabellar furrows may have been present, as in 
typical cryptolithinids. The distortion and crushing of the 
specimens (PL 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 luirt of fringe is wide and projects 
posteriorly. 

Fringe with outer surface of upper lamella sloping forward 
anteriorly, but anterolaterall.y 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 l)ackward and slightly outAvard. 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 
I2 ) immediately inside the girder, and a narrower, still lower 
ridge separates the next inmost rows (lo and L^ ; 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 E2 smaller than those of Ei, laterally and antero- 
laterally in a continuous row, but anteriorly becoming irregular, 
and about 10 pits from the midline Fi-^ bifurcates and E3 is 
formed in this anterior region (PI. 1. fig. 1). The pits of E3 
are smaller and more numerous than those in B^, and are not 
radially arranged with them. E^ and Ij are the largest pits in 
the fringe, in regular rows disturbed only at the genal angle, 
and anteriorly and laterally radially arranged with each other 
and the rows inside. I2 complete in front of the glabella, I3 
commencing at about the 4th pit of To from the median line, I4 
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 
E2 and El, Ej and Ij, I^ and I^, especially laterally and antero- 
laterally. In this same region radial ridges separate the pits 
of Ej 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 : Eo, 37 and 32; Ei, 31 and 29; I^, 30 and 29; L,, 32 and 
30. No ornament is preserved on the moulds of tlie cheek lobes 
or glabella. 

Four segments of the characteristic thorax ])oorly 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-like border and steeply-descending 
margins. First axial ring and pleural furrow distinct, remainder 
of axis not subdivided, but two or three additional ])leural fur- 
rows and inter})leural ridges may be discerned running straight 
outward and backward. 

Discussion. This species is cleai-ly a cryi)tolithinid (Whitting- 
ton, 1941, pp. 23-25) but is distinguished from Cfyptolithus 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 Eo are smaller than those of Ej, especially later- 



4 BREVIORA NO. 38 

ally, and here the upper lamella is convex upward, with a de- 
pressed reo-ion adjacent to tlie cheek lobe. These same characters 
distino'uisli the Venezuelan species from species of Crypto- 
lifhoi(](s (Whittington, 1941), and in addition the cephalon of 
dryptolithoides 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- 
(jcrolitlius (see Whittington, 1941, p. 24; Bancroft, 1949, pp. 
298-299, PL 9, figs. 4, 5, 6, 8, PI. 11, fig. 38), but has more rows 
of pits in the fringe and lacks the strong radial ridges in Ej, 
it is much more like species of Onnia (Whittington, 1940, PI. 3; 
Lamont, 1948, PL 1; Bancroft, 1949, p. 299, PL 9, figs. 9-13, 
PL 10, fig. 16). These latter have E2 of small pits, Ej and Ii 
of larger pits, and the region within Ij showing a strong radial 
arrangement, particularly in the depressed lateral region. The 
girder in Onnia is strong anteriorly, but laterally concentric 
ridges of almost equal strength may run between Ei and B2, 
and/or Ij and I2 (e.g. Whittington, 1940, PL 3, fig. 5; Lamont, 
1948, PL 1, figs. 4, 5). 0. ierryi shows a comparable development 
of ridges in the outer surface of the lower lamella in the interior 
region (PL 1, figs. 2, 4). 0. terryi differs from other species of 
Onnia in the presence of E3 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 ITpper 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 e.rsul Whittington, 1953, from early Middle Ordo- 
vician rocks of Florida, also has central European-North Afri- 
can affinity. If such a province existed, it is notable that it 
appears to lie almost entirely east and south of the belt of 
LIpper Cambrian and early Ordovician "Atlantic faunas" re- 
cently portrayed by Wilson (1954, fig. 4), and that the rocks 
are mainlv clastic. 



1954 ONNIA (tRILOBITA) from VENEZUELA 5 

REFERENCES 

Bancroft, B. B. 

1949. Upper Oidoviciiui tiilobites of zonal value in .soutlieawt Shrop- 
shire. Proc. Roy. Soe., London, Ser. B., vol. 136, pp. 291-31."), 
pis. 9-11. 

Lamont, a. 

1948. B. B. Baiu-roft's geological work. 2. Upper Ordovieian of the 
Cross Fell inlier. Quarry Manager's Jour., vol. 31, pp. 41(i-418, 
466-469, pi. 1. 

Lbith, E. 

1938. A Middle Ordovieian fauna from the Venezuelan Andes. Amer. 
Jour. Sei., 5th ser., vol. 36, pp. 337-344, pi. 

Termier, G. and H. 

1950. Paleontologie Maroeaine, 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 descril>ed by Joachim Barrande. Anier. 
Jour. Sci., vol. 238, pp. 241-259, pis. 1-4. 

1941. The Trinucleidae — with special reference to North American 
genera and species. Jour. Paleont., vol. 15, pp. 21-41, x)ls. 5, 6. 

1953. A new Ordovieian trilobite from Florida. Breviora, no. 17, 
pp. 1-6, pi. 1. 

Wilson, James L. 

1954. Late Cambrian and early Ordovieian trilobites from the Marathon 
Uplift, Texas. Jour. Paleont., vol. 28, pp. 249-285, pis. 24-27. 



EXPLANATION OF PLATE 

Onnin terri/i (Leith, 1938). Caparo-Bellavista Series, Middle Ordovician, 
near the Caparo River, State of Bariiias, Venezuela. 

Fig. 1. Rubber east from external mould of cephalon, showing upper 
lamella of fringe, X 4, MCZ 5183a. Extremities of third external row of 
pits indicated l)y " E:^. " 

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 (Ei) 
and second (E2) external row of pits, girder (g), and of first (Ii), second 
(To), and third (I:^), 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 "E:^." 

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 21/2, 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, X3, MCZ 5183f. 



E V I O R A 

Miaseuioi of Comiparsitive Zoology 

Cambridge, Mass. November 24, 1954 Number 39 

NEW OR REDESCRIBED PELOMEDUSID 

SKULLS FROM THE TERTIARY OF AFRICA 

AND ASIA 

(TESTUDINES, PELOMEDUSIDAE) 

2. A Podoeneniide 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 Podocneniis 
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 basioecipital and quadrate 
— were readily visible. Further examination left no doubt of 
the novelty of this fossil as compared with any previously known 
podocnemide ^ skull, and suggested interesting problems in regard 
to its proper correlation with a shell name and in regard to its 
l)hyletic position. 

The British Museum ]\Ioghara 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 phyletic 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 
lamina'?' of the premaxillae and maxillae to each other and to 

1 I use the term "pddociu'inide" as a convenient and informal designation for 
a subsection of the family Pelomedusidae typified hy the genus Podocncmix. 



2 A PODOCNEMIDE SKULL FROM MOGHARA 1954 

the choanal margin, the presence or absence of an anterior 
process of the palatines dividing the choanal oi)ening, the pres- 
ence or absence of a median pit in the palatal snrface 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 interpai-ietal 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 
"subocular" scute in the sense of Siebenroek (1902). 

The "carotid" channel is fully enlarged in podocnemide 
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 determinal)le 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 podocnemide skulls. Tabulating first against the living 
podocnemide species (Table 1), w^e 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 Moghai-a skull with the previously known fossil 
skulls of podocnemide type (Tal)le 2), the impressive fact is the 
close agreement, in cited characters, of the British Museum skull 
and Dacquemys pahomorpha. We need in fact to inquire what 
characters assure us that we are not dealing with Dacquemys. 



NO. 39 



fci2 



O O ij 



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A PODOCNEMIDE SKULL FROM MOGHARA 1954 



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o — 


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NO. 39 BREVIORA 5 

It is unfortunate that the unique generic character of Dac- 
qiiemys — the long squamosal-parietal suture — cannot be veri- 
fied or denied for the Moghara specimen. This region has been 
broken away. 

Tliere are, however, differences which may or may not imply 
generic distinctness. The British Museum skull is broad ; that of 
Dacqueniys is elongate. The orbits are visible dorsally in the 
]\Iiocene specimen ; they are completely concealed in Oligocene 
Dacqueniys. 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 madagascarcnsis of Boulenger) 
than to Dacqueniys. 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 Erymnochelys; 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 Dacqueniys and Erymno- 
chelys, perhaps somewhat closer to Erymnochelys. 

It 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 Erymnochelys 
and is on shell characters barely, if at all, distinguishable from 
Erymnochelys madagascarcnsis. I shall hereafter refer to Podoc- 
nemis aegyptiaca Andrews as Erymnochelys aegyptiaca (An- 
drews). 

There is only one other podocnemide shell type known from 
Moghara — the form named by Fourtau (1920) "Podocnemis" 
hiamlyi. This form differs from Erymnochelys aegyptiaca and 
from all Erymnochelys in the larger size of the intergular scale 
which separates the gulars as in the Recent South American 



G A PODOCNEMIDE SKULL FROM MOGHARA 1954 

podocnemide species (Podocncmis sensu stricto and most speci- 
mens of Peltocephaliis) . If we could assume that P. hramlyi is 
a precursor of Peltocephalus then it would not be too anomalous 
for our Moghara skull to belong to this species, rather than to 
E. aegyptiaca. In the living species Peltocephalus dumeriliana 
(in which the gular-intergular pattern is typically very similar 
to that of P. Iramlyi) the skull has definite, strong similarity 
to that of ErymnocheJys 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. hramlyi as to E. 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 
expect 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- 
liara skull — on whichever fork of the phyletic tree it belongs — 
is a structural intermediate between the Recent genera, Pelto- 
cephalus or Erynmochelys, and the Oligocene Dacquemys. 

Tills, indeed, is the principal suggestion that I wish to make : 
that there is a phyletic relationship between Peltocephalus, 
Erymnochelys and Dacquemys of the sort diagrammed below 
(Fig. 1). 



Peltocepholus - Recent of 
dumeriliono S.Americo 


Erymnochelys — Recent of 
modaqoscarensis Modaaosca 

\ 


"Podocnemis" bromiyi 
Miocene of Egypt 




1 

Erynnnocheivs - 
oegyptioco 


Miocene of 
Egypt and 
E. Africo 






Dacquemys — 
paleomorptio 


Oligocene of 
Egypt 






w 








? Docquemys 
fojumensis 





Fig. 1 Presumed phyletic position of the Egyptian Mioc-ene podoi-neniidc 



NO. 39 HREVIORA 7 

It cannot be denied that this suogested ])hyh)«i('iiy I'ests u])(in 
a rather elaborate structure of inferences, any or all of which 
may be wronj;' ; nothin<i' else, however, is possible in the current 
state of the evidence. 

It may be useful, here, to compare this su^'fiestion with the 
previous phyletic scheme for "' Podociiemis'' whieh was set up 
by Dacque 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. T differ 
with Dactpie, Zangerl and others in that T separate from the 
South American group the species dumcrllidiKi (type of Pelto- 
cephali(s) and regard the few resembhmces of the latter species 
to the other South American forms — similarity in gular ]iattern 
and in cervical articulations — as convergent only. 1 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 morjihological divergence that I recognize 
three genera within this lineage. 

Of these two respects in which I ditt'er with the Dac()ue-Zaugerl 
phylogcny, the recognition of African affinities in Pdiocephalm 
dumcr'lidiKi is the more solidly founded. I shall argue this point 
at lengtli in a future discussion of the rank, characters and varia- 
tion of the living species placed by Boulenger in Podooioiiis. 
The second conception — tliat of the Dacqueinys-Efniiiiioc/ich/s- 
Pcltocephalus series — is as yet hypothesis only, and the gi'ounds 
for this view have been stated here. 

It needs to be mentioned that Dac(iue (1912) described another 
skull from Moghara which he named Sternothacrus hlancJi-en- 
Jiorni. The descrijition is very brief and the only figure is a dorsal 
view. The skull was also (piite imperfect, lacking the temporal 
and occipital regions. 

If it belongs to iStcniollido-iis {=I'chisi()s) — an assignment 
neither contradicted nor specially supiiorted 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 ; I think, nevertheless, that the interorbital width is 
greater in the skull which has been here discussed than in the 



8 A PODOCNEMIDE SKULL FROM MOGHARA 1954 

type of hlanchenhorni. Beyond this, we must rely on Dacque's 
description of differences, and his most significant remark ap- 
pears to be his observation: " Bemerkenswert erscheint, dasz die 
Maxillaria die Choane auf der Schiidelunterseite otfenbar ganz 
iiberdecken. ' ' 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 Stereogcriys-Shweboemys series (which will be discussed 
in the concluding jjaper of this series), but in any event " Sterno- 
thaerus" hlanckenhorni would appear to have no bearing on 
the problems raised l)y 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 Clreen of the British Museum staff. 

PAPERS CITED 
Andrews, C. W. 

1906. A (k'Sfiiptive catalogue of the Tertiaiy veitebrata of the Fayum. 
London, 319 pp. 
D AC QUE, E. 

1912. Die fossilen Sehildkroten Aegyptens. Geol. Palaeont. AbhandL, 
vol. 14, pp. 27r,-837. 

FOURTAU, R. 

1920. Contribution a 1 'etude des vertelnes mioeenes de 1 'Egypt. Cairo 
(Government Press), pp. 29-34. 

SlEBENROCK, F. 

1902. Zur Systematik der Sehildkroten Gattung Podocncmis Wagl. 
Sitz. Ber. K. Akad. Wiss. Wien., Math. Nat. Kl., vol. 3, pp. 1 14. 
Williams, E. 

1954. New or redescribed pelomedusid skulls from the Tertiary of 
Africa and Asia (Testudines, Pelomedusidae). 1. Dacquemys 
palcnmor2>ha, new genus, new species, from the Lower Oligoeene 
of the P'ayum, Lgy]>t. Breviora, no. 35, ])p. IS. 
Zangerl, R. 

1948. The vertel)rate fauna of the 8elma formation of Alabama. Part 
II. The pleurodiran turtles. Fieldiana: Geol. Mem., vol. 3, 
pp. 23-o6. 




PLATE 1. 
British Museum skull from Mogliara, dorsal view. Posterior end at top of 
page. 





f . ^/ 1^ 



PLATE 2. 
British Museuiii f-kiill from Moghara, ventral view. Posterior end at top of 



E V I O R A 

Miaseium of Comparative Zoology 

Cambridge, Mass. November 29, 1954 Number 40 

SOME MOLLUSKS FROM THE 

CONTINENTAL SLOPE OF NORTHEASTERN 

NORTH AMERICA^ 

By Arthur H. Clarke, Jr. 

The continental slope area of northeastern North America 
has always been a relatively unknown and unexplored region. 
Onl.v 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 II 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- 
graphic 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. 



2 BREVIOBA 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 AlMtross 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 II, 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 
geographic 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 28 species, it will be 
seen that these new data have extended the previously recorded 
bathymetric ranges of seven species (marked *) and the geo- 
graphic ranges of six species (marked f). 

List of Mollusca Collected 

YOLDIA THBACIAEFORMIS (Storer)*t 

One large dead specimen at station 72. Bathymetric range: 
400 fms. dead (this survey) ; 29-182 fms. dead (Verrill 1884). 



1954 MOLLUSKS FROM NORTHEASTERN NORTH AMERICA 3 

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: 
Arctic 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 Miiller 

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 scaly sur- 
face sculpture is entirely lacking in many individuals and in 
some entire large lots of specimens. 

Arctica islandica (Linne) 
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) ; 
Arctic 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 
Scotia (this survey). 

Panomya arctica (Lamarck) t 
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; circumpolar (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 tubercules on the upper whorls 
and golden peach on the lower whorls. The inner nacre of the 
aperture shines with iridescent pink and green. The circular 
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 1934). Geographic range: Arctic [Ocean] (Verrill 
1884) ; Labrador to oif 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 Rocque 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 (Linne) 
One small living specimen attached to Pecten vitrevs 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 Rocque 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; 341/2-1000 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 Bruguierc 
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)*t 
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 -. 270-360 fms. alive ; 225-660 f ms. dead 
(both this survey) ; 41-86 fms. alive; 6-322 fms. dead (both, Ver- 
rill 1884). Geographic range: Circumboreal (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 2i/^ times the great- 
est width. Other specimens show a length of only li/^ times the 
width. Variation is also seen in degree of sculpture, length of 
aperture in relation to overall length, etc. The possibility of the 
existence of one or more clines will be investigated. 

Neptunea despecta tornata (Gould) *t 
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 (Gmeliu) 
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 rayige: Labrador to Norway (Johnson 
1934); Arctic 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 Ilenlopen, New 
Jersey (this survey). 

CoLus pitbescens (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, 132, 139, 144, 164, 173, 184, and 225. Bathy metric 
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 (Ball 1889) ; southeast of 
Nova Scotia to east of Chincoteague Island, Virginia (this sur- 
vey). 

Pleurotomella agassizi Verrill and Smithf 
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 piinctostriatus (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. Ball 1889). 

Rossia 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 109i-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 BREVIOBA NO. 40 

Geographic range: south of Georges Bank (this survey) ; Nova 
Scotia to West Indies (Johnson 1934). 

Mastigoteuthis agassizii Verrill* 
One living specimen at station 102. Bathymetrie 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 Verrillf 
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 
Atlantic (both, Johnson 1934). 

Bathypolypus arcttcus (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 Fmidy 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 


17 


39°55' 


71°17' 


290-330 


19 


39°57' 


71°13' 


175-180 


22 


39'53' 


70°53' 


300-330 



1954 MOLLUSKS FROM NORTHEASTERN NORTH AMERICA 



90 
95 
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 



39°52' 


70°51' 


375-420 


39°52' 


70°43' 


415-440 


40°02' 


70°24' 


105-110 


40°04' 


70°12' 


190-225 


39°57' 


69°37' 


415-560 


39° 55' 


69°26' 


165-180 


40°08' 


68°19' 


400-510 


40°10' 


68°16' 


490 


40°12' 


68°08' 


400-450 


40°17' 


67°53' 


450-500 


39°59' 


69°32' 


87-90 


39°59' 


69°35' 


82-85 


42°23' 


64°58' 


230-245 


42°46' 


63° 22' 


340-350 


42"41' 


63°33' 


300-330 


42'45' 


63°47' 


330-340 


42°35' 


64°03' 


560-600 


42°40' 


64°08' 


350-380 


42°20' 


65°03' 


305-320 


42°17' 


65°06' 


320-360 


40°46' 


66°40' 


290-300 


38°13' 


73°40' 


145-160 


38°08' 


73° 45' 


205-275 


37°38' 


74°14' 


120-130 


37°45' 


74°09' 


150 


38°33' 


73°18' 


63-65 


38°39' 


73° 05' 


330-400 


39°26' 


72°10' 


690-720 


42°43' 


63° 50' 


310-335 


42°42' 


63°47' 


360-370 


42°40' 


64°10' 


240-270 


42°33' 


64°17' 


280-320 


42°32' 


64°19' 


360-420 


42°28' 


64°31' 


280-305 


42°23' 


64°52' 


265-295 


42°15' 


64° 58' 


660-705 


42°18' 


65°05' 


220-240 


40°34' 


67°02' 


290-300 


40°46' 


66°48' 


100-105 


40°45' 


66°51' 


85-87 


40°09' 


68°24' 


250-340 


40°00' 


68°49' 


600-670 


39°53' 


70°40' 


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 pis. 

Gould, A. A. 

1870. Report on the Tnvertebrata of Massachusetts. Edited by W. G. 
Binney. Boston. Pp. i-v + 1-524, 12 pis. 

Johnson, C. W. 

1934. List of marine Mollusca of the Atlantic Coast from Labrador 
To Texas. Proc. Boston Soc. Nat. Hist., 40. no. 1 : 1-203. 

La Rocque, Aurele 

1953. Catalogue of the Recent Mollusca of Canada. Nat. Mus. Canada 
129: i-i.x + 1-406. 

NicoL, David 

1951. Recent species of the veneroid pelecypod Arctica. Jour. Wash- 
ington Acad. Scl, 4L no. 3 : 102-106. 

Thorson, Gunnar 

1951. The Godthaab expedition 1928. Meddel. om Greailand, Copen- 
hagen, 8Lno. 2: 1-117. 

Tryon, G. W., Jr. 

1879. Manual of Conehology (1), 1. Cephalopoda. Acad. Nat. Sci., 
Philadelphia, pp. 1-31G, 112 pis. 

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 pis. 

1885. Third catalogue of Mollusca recently added to the fauna of 
New England. . . . Trans. Conn. Acad., 6. no. 2 : 395-452, 3 pis. 



1954 MOLLUSKS FROM NORTHEASTERN NORTH AMERICA 



11 




- 4- 

I 



E V I O R A 

Mmseitim of Comparative Zoology 

Cambridge, Mass. December 17, 1954 Number 41 



COMMENTS ON THE CLASSIFICATION 
OF RODENTS 

By Albert E. Wood 

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- 
gie history of the order is taken into account. For example, if 
the Hystricomorpha are a natural group, how can 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 include 
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. 

Lavocat (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 cases, 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 Isehyromyoidea), 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 Avell 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 (1953a). 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 Eocene. Schaub (1953a, 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 OP RODENTS 5 

extrapolation of the post-Deseadan rate of evolution would indi- 
cate 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 
gnawing 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 earliest 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 Eocene 
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 7 

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 
gnawing 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. 



EEFERENCES 

Lavocat, E. 

1951. Revision de la faune des mammiferes oligocenes d'Auvergne et 
du Velay. Editions "Sciences et Avenir," Paris, 153 pp. 

Miller, G. S., and J. W. Gidley 

1918. SjTiopsis of the supergeneric groups of rodents. Jour. Wash. 
Acad. Sci., 8: 431-448. 

SCHAUB, S. 

1951. Review of: H. G. Stehlin et S. Schaub, "Die Trigonodontie der 
simplicidentaten Nager. " Mammalia, 15: 204-210. 

1953a. Remarks on the distribution and classification of the "Hystri- 
comorpha." Verh. Naturf. Ges. Basel, 64: 389-400. 

1953b. La Trigonodontie des Rongeurs simplicidentes. Ann. Paleont., 
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 S. Schaub 

1951. Die Trigonodontie der simplicidentaten Nager. Schweiz. Pal. 
Abh., 87: 1-385. 

Wilson, 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 

Wood, A. E. 

1937. The mammalian fauna of the White Eiver 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. 

Soc. Amer., 57: 1245 (Abstract). 
1950. Porcupines, palaeogeography, and parallelism. Evolution, 4: 

87-98. 
(In Press.) A revised classification of the rodents. Jour. Mammalogy, 



Wood, 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. 



E V I O R A 

MeseiiMM of CoaMparative Zoology 

Cambridge, Mass. February 28, 1955 Number 42 

A NEW SALAMANDER OF THE GENUS 
PARVIMOLGE FROM MEXICO 

By GrEOEGE B. Rabb 

Museum of Zoology, University of Michigan 

111 the Museum of Comparative Zoology there is a specimen of 
a Mexican plethodontid salamander superficially similar to in- 
dividuals of Pseiidoeurycea 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. 

Descr-ipiion. 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 



BREVIORA 



NO. 42 



^) 



t! 



1955 A NEW SALAMANDER OF THE GENUS PARVniOLOE 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 lip. 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 



4 . 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 liml>- slightly 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 someM'hat 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 light 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 0.3 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 con- 
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, Avhich is slightly 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 S-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-oecipital condyle distance ; f rontals 
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, 0.3 ; fronto-premaxillary fonta- 
nelle width, 0.3. 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 l)oth 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-like process ex- 



195.': 



A NEW SALAMANDER OF THE GENUS PARVIMOLGE 



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 description (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 T. pennatu- 
h(s, and since the single cleared specimen referable to T. 
dubitus that I have examined likewise has entirely cartilaginous 
ones. However, I believe that most of the discrepancies between 
the two descriptions of P. fownsendi 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 


E 


20-23 


20-26 




32-42 


45-58 


54-69 


males (4) 


M 


S£.0 


£2.5 


3.7 


39.0 


50.0 


61.5 


townsendi 


R 


19-22 


16-21 




35-41 


48-58 


55-73 


females (4) 


M 


21.0 


19.0 


4.2 


ss.o 


53.0 


64.0 


praecellens 
















(female) 




35 


30 


1 


50 


75 


87 



E, 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 townseiidi 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 Pseudoeurycea and 
Chiropterotriton. Indeed, in the latter genus, C. dimidiata shows 
seemingly paedomorphic features very like those found in town- 
sendi. 

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 PARVIMOLOE 9 

LITEEATUEE CITED 

Dui«sr, E. E. 

1922. A new salamander from Mexico. Proc. Biol. Soe. Washington, 35: 

5-6. 
1926. The salamanders of the family Plethodontidae. Smith College, 
Northampton, Mass. viii + 441 pp. 
Hilton, 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., 
34. Pt. II, 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 Eica. Univ. Kansas Sci. Bull., 33, 
Pt. I, No. 6: 279-88. 

1952. The salamanders and cjiccilians of Costa Rica. Univ. Kansas Sci. 
Bull., 34, Pt. II, No. 12 : 695-791. 



R E V I O R A 

imim of Comparative Zoology 



Cambridge, Mass. x\pril 7, 1955 Number 43 

SPEED-INDUCED SKIN FOLDS IN THE 

BOTTLE-NOSED PORPOISE, 

TVliSIOPS TRUNCATUS ' 

By Fkank S. Essapian 

^rarineljuid Kesearcli T.alioratoiy 

While engaged in a study of the Ix'havior of bottle-nosed por- 
poises, Tursiops irmicdliis (Montagu), at Marine Studios, 
Marineland, Florida, over a period of three to four years, the 
author has oliserv(nl on hundreds of occasions a remarkable 
modification of the skin contoui- of these animals whenever they 
resorted to I'apid movement about the tank (Essapian 1953, 
p. 3!)!;)). Tlie circular tank, where the i)orpoises are on exhibit, 
is seventy-five feet in diameter and about fourteen feet deep 
at the center. It liouses an average of ten animals. 

Generally lost to view due to swiftness of action, this trans- 
fornmtion of the skin contour takes place when an animal accel- 
erates its s])eed 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 I'elief on ])art of an animal, or may 
extend over its entire lengtli, 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 s(^en even in an animal only a day or two old. 

The skin folds generated b}' a sudden burst of speed are by far 
the more common of the two categories, and are usually oc- 
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 

^ Coiitriliutiou No. 7.")! from the Woods Hole Occaiiogrjiiihie Institution. 



2 BREVIORA No. 43 

by the factors which motivate tlie 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 (Pis. 1 and 2). However, in the case of 
prolonged rapid movement around the tank, the animals may 
intermittently exhibit skin folds for a jieriod of one or two or 
more seconds. Often an animal at pla}' or during the mating 
season may suddenly leap out of water, exhibiting folds at the 
peak of acceleration, just ])rior to its emergence from water, 
and again after it has reentered the water (Pis. 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 rapidl.y without exhibiting any folds 
comes to an abrupt halt. Then the folds may suddenly be dis- 
played (PI. 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 w^ould 
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. Tn a Tursiops truncatus fifteen days old and weighing 
thirty pounds, the skin, including blubber, is approximately half 



1955 BOTTLE-NOSED I'ORPOISE TURSIOl'ti TRUNCATUS 3 

an inch thick. In an adult it is nearly one and a half inches 
thick on the ventral surface. Althouoii normally not a])j)arent 
to the view or touch, the jjoi-poise'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 slin<i' is shown in Plat<' 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 Avater njion 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' ditferent tyi)es 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 
swinnning normally do not display such looseness of skin as 
is here ascribed to the gi-eat baleen whales. However, this 
similarity in the tl(^xibility of skin in such divergent species sug- 
gests that all aninuds of the order ('(^tacea 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. 

Kacovitza (190;), pi). 4o-44, pi. 3, fig. 17) mentions a group of 
four HijperoodoH (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 
rai)idly at the time or whether this deformity in the skin contour 
was api)arent even when the animal had raised itself partly out 
of water. The author furnished no further explanation beyond 
ex]iressinii' 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 tlie Woods Hole Oceanographic 
Institution. 

EEFEEENCES 

Clarke, Egbert and Johan T. Euud 

1954. International cooperation in whale marking: The voyage of the 
"Enern" to the Antarctic 1953. Norsk Hvalfangst-Tidendc, 
43(3): 128-146. 

EssAPiAN, Frank S. 

1953. The birth and growth of a porpoise. Natural History (New 
York), 62(9) : 392-399, ills. 

Racovitza, E. G. 

1903. Cetaees. Resultats du voyage du S. Y. Belgica. 1897-1899. 
Eapports Scientifiques, Zoologie. 142 pp., 15 text figs., 4 pis. 



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. 




1*1. .!. A female, approximately eighteen months old, aljout to leap ont 
of the water. Note also the folds on the head of the i)nisiung five-year- 
old male. 
Photographed at F/3..J at 1/500 of a second. 




PI. i. 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 second. 




. All adult male displays folds as he comes to an abiiipt stop in a 
vertical position (second porpoise in background shows no folds). f]n- 
larged from 16 mm. film. 
Photographed at l(i frames per second. 



t 



I 



PI. 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 r/3.5 at 1/250 of a second. 




PI. 9. Tolds in the neck-shoulder region caused by turning the head side- 
ways. This is the same animal as in Plates 4, 5, and 8. 



Photographed at F/3.5 at 1/250 of a second. 



E V I O R A 

Miiseiiam of Comparative Zoology 

Cambridge, Mass. April 8, 1955 Number 44 

A NEW MLTREX FEOM MATANZAS, CUBA 
By Wn^LiAM 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 Miirex which was obtained from shell traps set in 100 fathoms 
in Matanzas Bay, Cuba. 

MuREX (MuREx) FiNLAYi, new species 
F'igures 1-3 

Description. Shell reaching 93 mm. (3% 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 subcircular and slightly oblique. Parietal lip glazed, 
adherent to the body whorl and fairly thick. Palatal lip crenu- 
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 scale- 
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 sculp- 
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 


»5.0 


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 localit^^ 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 Mure.v heauii Fischer and 
Bernardi. From M. antillamm 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 
canal. From M. heauii it differs by having fewer and much larger 
axial costae, shorter spines and not having the extensive webbing 
which characterizes M. heauii. This present new species may also 
be related to 31. aguayoi Clench and Farfante but it differs from 
aguayoi 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 
a very complex group and of very wide distribution in nearly all 
tropical and south temperate seas. 



,^. 



•4^^ 



.y^ 







Mitrex finlayi Clencli. Figs. 1 aud 3, paratypes. Fig. 
Bay. Cuba in 100 fathoms. (All slightly enlarged.) 



liolotype. Matanzas 



E V I O R A 

Museunn of Comparative Zoology 

Cambridge, Mass. June 13, 1955 Number 45 



PALAEOTAincHA OLiaOCENKW, NEW 

GENUS AND SPECIES, AN OLKiOCENE 

SALAMANDER FROM OREGON. 

By Richard van 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 V4 ^^^il© north of Goshen, Oregon. The 
specimen was sent by Dr. Herman Clark of Willamette Univer- 
sity to the Museum of Comparative Zoology for stndy. 

The strata in which the fossil was found are designated by 
Vokes ef al. (1951) as post- Fisher and Eugene (Oligocene) 
plant-bearing tuifs. These strata are described by the authors as 
"a secpience 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. Chanej' of the University of 
California has written me that he too considers the strata to be 
younger than Eugene 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 eontains 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 black in color, apparently highly carbonized. The skull is 

1 and listed by Vokes ef al. (1951) 



2 BREVIORA NO. 45 

crushed 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 tw^o 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. Paroceipital processes, 
frontosquamosal arches, and an unpaired premaxilla are char- 
acters of this fossil as well as, in Recent salamandrids, ('ynnp>i 
Tschudi, 1838, Diemiciylus Rafinesque, "1820, Eu proof us Gene, 
1838, Hypselotriton Wolterstorff, 1934, Paehytriton Boulenger, 



1955 PALAEOTARICIIA OLIGOCENICA 3 

1878, Taricha Gray, 1845, and several species of Trittirus^ 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, Pachytrito7i, 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. Euproctus 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 {iorosa, 
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 

liis iisfd liy \\(ill.Tst(irfl- ,\nd IleiTe (li»;-(5). 
- voiiHTopalatiiif (ir 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: ralaeotaricha 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 
prepubic processes. Tarsal basale formula 1, 2, 3, 4-(-5. Maxillae 
extending caudad, united by suture Avith quadrates. 

Palaeotaricha oligocenica, new 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 lift.). 

Locality: About 3 miles (5 km.) southeast of Eugene, Lane 
County, west central Oregon; southeast Y^, sec. 14, T. 18 S, R. 
3 W ; " about 550 feet east of the Southern Pacific 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 3000 ft. 
west of long. 123 deg. ... in the Eugene quadrangle" (Dr. 
Herman Clark, in litt.). 

Diagnosis: As for the genus. 



1955 PALAEOTARICHA OLIGOCENICA 5 

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 paroc- 
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 paroccipital 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 lies wholly within the vomer. The anterior 
border of the vomer is an arc of a circle (roughly), with no sharp 
l)reaks 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 Fahieotaricha oUgocenica it is horizontal for most 
of its length, then there is a short vent rally directed slope toward 
the anterior end, and then a sudden vertical drop to the condylar 
facets (PI. 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 (PI. 1, %. 1; PI. 3, fig. 1). All but the cauclalmost 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 phy'angeal count is 1-2-3-2, which, together with the carpal 
basale ff mula (1+2, 3, 4), is the same as in Taricha. 

The pchic girdle has floated free and, with the limbs, is fairly 
well preserved as a unit. The pelvic girdle is apparently a highly 
ossified and coossified structure. Zones of weakness may perhaps 
exist anteriorly between left and right halves, and between the 

1 Some A tiie things that look like rihs in Plntf 1, figure 1 are phalanges of the 
left niaaus. 



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 
prepubic processes are rounded and robust. They project later- 
ally and apparentlj^ slightly posteriad in P. olujocenica, 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 Salamanc'ridae. 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 Kecent salamandrids, Pleurodcles Michahelles, 
1830, SaJamandrina Fitzinger, 1826, and Tylofoirifon Anderson. 
1871, the maxillae may very nearly or actually touch the quad- 
rates. In the Palaeotaricha oligoccnica 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 Vs- There is, how- 
ever, a good deal of individual variation and intergradation be- 
tween these tooth-row patterns. The available skulls of T. sierrae 
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 be somewhat deep and domed, in contrast to the 



1955 PALAEOTARICHA OLIGOCENICA 7 

more-or-less flattened and widened roof of the others. T. 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 circular 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 
(cf. Palaeotaricha) or is primitive (cf. primitive extant genera of 
salamandrids, which have the premaxilla paired, and some have 
the aperture extremely large). 

A full description of the skull of T. 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, 
jide Herre (1949) and corroborated in the specimens available 
to me. 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. 
oligocenica. The closest approach of all seems to be in the 
Miocene Ckelotriton Pomel, 1853, w^hich 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, PI. 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 like dermal plates ; and, of course, the series 
just given has only descriptive validity. 

The uncinate processes of the P. oUgocen'ca ribs are longer 
than in the Recent Taricha specimens at hand, but these struc- 
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 1+2, 3, 4, 5 in the Recent Taricha, Chioglossa 
Barbo.-'a du Bocage, 1864, Salamandra Laurenti, 1768, Mertensi- 
ella Wolterstorif, 1934, and Pleurodeles Michahelles, 1830, while 
Salaniandrina with four toes on all feet apparently has 1, 2, 3, 4, 
and all other genera have 1+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 corrol)orated 
and extended b}' the specimens available to me, the carpus has 



1955 PALAEOTARICHA OLIGOCENICA 9 

basales 1+2, 3, 4, prepollex basale,' eentrale, radiale, inter- 
medium, and iilnare, in all genera. The latter two may or mav 
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 ( 19-48) agree in general with 
those of Bolkay and me, Hilton illustrates various eases 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 light of Hilton's data, the difference be- 
tween the PalaeotaricJia 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 PalaeotaricJia 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 : I have been fortunate in being able to 
examine skulls and skeletons of every genus of Recent 8ala- 
mandridae, with the single exception of Pleurodeles, and to see 
alcoholic specimens of nearly every species of all genera. It must 
be 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 (PI. 2. A) was 

1 Bolkay follows another usa{,'e in calliiit? this element laipal hasale 1 ; and what 
he calls 2 is here called 1+2. Likewise he desi^ates the prehallu.x basale as 
tarsal basale 1. and 1+2 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 21/0 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" x 7" 
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 panatomic film, and illuminated by 
two 100- and one 200-watt bulb with reflectors, each about 2 feet 
from the subject. 

ACKNOWLEDGMENTS 

r 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 sturly, and for much help and continued advice. 



1955 PALAEOTARICHA OLIGOCENICA 11 

BIBLIOGRAPHY 

BOLKAY, StEPHAN JoSEP 

1927. Materialen zu einer vergleiehenden Morphologie der Carpal- und 
Tarsalelemente der Unterfaniilie Salamandrinae. Glasnik Zeiiial 
.iskog Miizeja u Bosiii i Hercegovini, vol. 39, pp. 59-67. 

1928. Die Schiidel der Salamandrinen, niit besonderer Riicksicht auf 
ihre systeniatische Bedeutiing. Zeitsehr. Aiiat. Entwicklungs 
gesch., vol. 86, pp. 259-319. 

GoLUFUs.s, Georg August 

1831. Beitriige ziir Kenntniss verschiedener Reptilien der Vorwelt. 
Nova Acta Acad. Leop. Carol., vol. 15, pp. 61-128. 

llKKUii:, Wolf 

J 934. Die systeinatisclie Stellung von TaricJia torosa Esehliolz. Blatter 

t'iir Aquarieii- und Terrarienkunde, vol. 45, pp. 250-252. 
1935a. Die Sehwanzlurche der initteleociinen (oberlutetisehen) Braun 

kolile des Geiseltales und die Phylogenie der Urodelen unter 

Einschluss der fossilen Formen. Zoologica, vol. 33, art. 4, pp. 

1 85. 
19351). I'her 0/i(;o.v< »u'« spinosa Navas, einen fossilen Schwanzlurch aus 

deni spanischen Tertiar. Palaeont. Zeitsehr., vol. 17, pp. 91-105. 
J 941. Falaeopleurodeles haitffi nov. gen. nov. spec, ein fossiler Schwanz 

lurch aus dem Mioziin Siiddeutschlands. Zool. Anz., vol. 134, pp. 

1-17. 
1949. Neue Tatsachen zur Stammesgeschichte der Sohwanzlun-he. 

Zool. Jahrb., Abt. Syst., vol. 78, pp. 217-236. 
1950a. Schwanzlurehe aus dem PaleocJin von Walbeck. In Neue Ergeb 

nisse und Probleme der Zoologie (Klatt-Festsehrift). Leipzig, 

Geest und Portig, pp. 286-301. 
J 9501). Der dorzeitige Stand unseres Wissens iibcr die fossilen Urodelen, 

zugleieh einige kritisehe Bemerkungen iiber Boomgardia sala- 

viaiidriformis v. Huene. Neues Jahrb. Geol. Paliiont., Monatsh., 

li»50, pp. 19-25. 

IIU.TON, WlLLIAJI A. 

194S. The ('ari)us and tarsus of salamanders. Jour. Entomol. Zool., 
vol. 40, pp. 1-13. 

l.AlliK, Gl'STAV C. 

1901. Synopsis iler Wirbelthierfauna der Ijciliin. Braunkohlforniation 
und Besehreibung neuer, oder bisher unvollstiindig bekannter 
Arten. Abhandl. deutsch. naturwiss.-med. Ver. Bohmen "Lotos," 
vol. 2, art. 4, i)p. 1-80. 



12 BREVIORA NO. 45 

MjiYER, Hermann von 

1860. Salamandrinen aus der Braunkohle am Ehein und in Bolimen. 

Palaeontographica, vol. 7, pp. 47-73. 
1803. Heliarchon fiirciUatus, ein Batrachier aus der Biaimkohle von 
Eott ini feiebengebirge. Ibid., vol. 10, pp. 292-298. 
VoKES, H. E., Parke D. Snavely, Jr., and Donald A. Myers 

1951. Geology of the southern and southwestern border areas of the 
Willamette valley, Oregon. U. S. Geol. Surv. Oil and Gas Invest. 
Map. OM 110. 

WOLTERSTORF, WiLLY and WOLF HeRRE 

1935. Die Gattungen der Wassermolche der Familie Salaniandridae. 
Arch. Xaturgesch., new ser., vol. 4, pp. 217-229. 

EXPLANATION OF PLATES 
PLATE 1 

Figs. ], 2, 4. Palaeotarictlia 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; del, dorsal opening of cavum internasale ; 
en, external naris ; /, frontal ; fsa, f rontosquamosal arch ; fv, foramen vesti- 
buli; hy, parts of hyoid apparatus; in, internal naris; Ij, 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; rci, vomerine opening of cavum internasale; v, vomer. All figures 

x2y4. 

PLATE 3 

Palaeotariclui oligocenica, holotype. Fig. 1. Eight pectoral girdle and 
limb, and ribs 3 to 7. Forearm and manus in dorsal view, other parts in right 
side view, ho l-\-2, Carpal basales 1+2; be /-/-^, carpal basalejf i^/4; /(, 
humerus; mc 1, metacarpal 1; pph, prepollex basale; r, radius; ra, radiale; 
rG, rib (i ; .s, scapula; u, ulna; ul-\-i, ulnare + intermedium. 

Fig. 2. Pelvic girdle and limbs. Pelvis and left limb in ventral view, right 
limb (left side of photograph) in dorsal view, ht 4-\-5, Tarsal basales 4+5; 
c, centrale; /, femur; /e, fibulare; fi, fibula; i, intermedium; phh, prehallux 
basale; prp, prepubic process; f, tibia; ti, tibiale ; y, ypsiloid process. Both 
figures X 2^4. 




A <9 



PLATE 1 







PLATE 





PLATE 3 



BREVIORA 

Meseiuiei of Comparative Zoology 

Cambridge, Mass. June 24, 1 955 Number 46 

CAVE-FOSSIL VERTEBRATES FROM 
CAMAGUEY, CUBA 

By Karl F. Koopman 

Department of Biology, Queens College, Flushing, New York 

and RoDOLFO Ruibal 

Division of Dife 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 
cave, 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 
cave can only be described by saying that there are millions. To 
walk in any part of this cave is to Avalk 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 3 

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 cuhanus. 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 waierhousei. 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 ciihanus. A single mandible from Cave 2. A 
single periotic bone from Cave 2 is also allocated to this species. 
AH 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 sezekorni. 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- 
scribed 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 T. muscula a subspecies of T. hrasiliensis.) Repre- 
sented by two rostra and two mandibles from Cave 2. Previously 
known from all the provinces 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 torrei. 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 knowm 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) columhianus. 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. 

Battus. Represented by many partial skulls, mandibles, and 



6 BREVIORA NO. 46 

postcranial 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 postcranial 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. 

Leiocephalus sp. A large dentary that seems to represent a 
species of Leiocephalus 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 postcranial 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 owl 
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 — Battus 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) columhianus — or extinct 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. Extinct 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. 



BREVIORA NO. 46 



GUNDLAOH, J. 

1868. Eevista 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. S. 

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. 

MOHR, 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. 



E V I O R A 

Mmseiamri of Compsirative Zoology 

Cambridge, Mass. July 12, 1955 Number 47 

A NEW SPECIES OF WHIPTAILED LIZARD 

(GENUS Cnemidophorus) FROM THE 

COLORADO PLATEAU OF ARIZONA, NEW 

MEXICO, COLORADO, AND UTAH 

By Charles H. Lowe, Jr. 

Department of Zoolog.v, 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 
stictogramnius 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 innotatus 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 redescribing 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 



BREVIOBA 



No. 47 



some respects his description fits it better than inornatus." 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 



INORNATUS 
N'25 




TOTAL, 54 



140 



150 



170 



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. sacM 
innotatus (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. Prom 
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- 
specific with sacki (= 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 occur (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 sympatric in Woodland and 
Coniferous Forest habitats in central Arizona, near the northern 
limit of the geographic distribution of C. sacki 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 tAvo species have been most 



4 BREVIORA No. 47 

commonly confused. The highly significant difference in the 
dorsal scalation 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. inornatns 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 light 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 scalation (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 browTi 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 (1931:124, 127) while at the same time 
accepting and using Burger's (1950:3) nomenclature (C. i7i- 
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[i-/cj." The specimen in question (No. 
37208) is recorded as the holotype of Cnemidophorus gularis 



6 BREVIORA No. 47 

veJox 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. gidaris velox Springer. Dr. N. E. 
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 
Cnemidophorus 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 
(cotypes) 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. gidaris 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 (iri 
litt.). 

ACKNOWLEDGEMENTS 

I Avish 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 7 

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 

Barbour, T. and A. Loveridge 

1946. First supplement to typical reptiles and amphibians. Bull. Mus. 
Comp. Zool., 96:(2): 57-214. 

BUUGER, W. L. 

1950. New, revived, and reallocated names for North American whip- 
tailed lizards, genus Cnemidoplioru^. 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. S. 
Nat. Mus., 154: 1-286. 

COWLES, R. B. 

194.3. Surfat-e-niass ratio, paleoelimate and heat sterility. Amer. Nat., 
79: 561-567. 

Fitch, H. 

1940. A biogeographical study of the ordinoides Artenkreis of garter 
snakes (genus Thamnophis). Univ. Calif. Publ. Zool., 44(1): 
1-150. 

Smith, H. M. and E. H. Taylor 

1950. An annotated checklist and key to the reptiles of Mexico exclu- 
sive of the snakes. Bull. U. S. Nat. Mus., 199: 1-253. 

Springer, S. 

1928. An annotated list of the lizards of Lee's Ferry, Arizona. Copeia, 
169: 100-104, 



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BREVIORA 

MmseiLinii of Comparative Zoology 

Cambridge, Mass. October 10, 1955 Number 48 



NOTES ON AMERICAN EARTHWOBMS OF 
THE FAMILY LUMBRICIDAE. I-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. 



EiSENiA LONNBERGi (Michaelscn) 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 stumiDs), Oct. 6, 1940, 0-1-2. 
Dr. G. E. Pickford. (Clitellate specimens were copulating 
when taken.) 

Meshoraasic 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. E. Pickford. 

Tallulah Falls, Georgia. In ravine, in sandy loam in seep of 
stream, July 27, 1931, 1-0-13. J. M. Valentine per Dr. 
G. E. 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 elitellar 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 viii-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 comers 
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 (30 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-xxviii (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 TETE 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 xxiii-xxx (1, ex copula), of xxiii (1). 
The a and h setae of xx-xxx (1), xxi-xxx (1), xxii-xxx (8), xxiii- 
xxix (10), xxiii-xxx (11), xxiv-xxix (1), xxiv-xxx (5), 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 h setae of x (3) and/or the c 
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 a^iatomy. Septa 12/13-14/15 muscular and increas- 
ingly thickened posteriorly. 

Oesophageal wall slightly 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-xviii, some special muscularity in xix in at least 
one of the worms. The typhlosole begins abruptly in region of 
xxiv-xxvii 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 for a 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 c-cxv 
(c/. 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 (33), 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 ii?). Nephridial ducts 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. 
Ovisacs .small, slightly 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 (c 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- 
able but the a and h follicles of xx-xxx were conspicuously pro- 
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 hodies. 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 Eiseniella 
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 pll of 4.5 also was mentioned by Dr. Pickford. 



1955 AMERICAN EARTHWORMS OF THE FAMILY LUMBRICIDAE I-II 5 

Life 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, like 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-xi 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 viii-xii in one worm, just as happens 
in Eisenia 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 clitellar 
region is associated with GS follicles and tumescences or with 
tubercula 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 Lum'bricus terrestris L., may not take place. 

In the evolution of E. lonnhergi 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 Miehaelsen 1910 

Helodrilus (Eisenia) carolinensis Miehaelsen 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 caroli^iensis, Cernosvitov, 1942, Proc. Zool. Soc. London, 111, (B), 
p. 222. (14 specimens, 5 immature, from Durham, N. C, in Brit. Mus.) 



1955 AMERICAN EARTHWORMS OF THE FAMILY LUMBRICIDAE MI 7 

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. B. 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 
101/2 miles on road to Whitetop Mountain, July 21, 1931, 
0-1-4. Dr. G. E. 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 (c/. 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 
concave. Prostomium epilobous (28 specimens), tongue open 
and usually quite short. Setae begin on ii (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 < iC. 
Nephropores first recognizable on iii, 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 (xxiii)xxiv- 
xxxi (3), (xxiii)xxiv-xxxi(xxxii) (1), xxiii-xxxi (3), xxiii-xxxii 
/2 (1), xxiii-xxxii (1), xxiv-xxxi (6), xxiv-xxxii/2 (1), xxiv/2- 
xxxii/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. Id7in- 
hergi, on xxvii-xxix (25), perhaps occasionally extending slightly 
onto XXX and/or xxvi. 

Genital tumescences conspicuous and transversely elliptical, 
always including both a and h setae, on ix (1), xi (3), xii (3), 
xvi (2), xvii (21), xviii (5), xx (7), xxi (16), xxii (1), xxiii (1), 
xxiv (3), xxv-xxx (27), xxxi (14), xxxii (2), xxxiii (1), c-d of 
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 12/13-14/15 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 E. 
lonnhergi Michaelsen 1894, large, moniliform, in xi-xii, sepa- 
rated by a deep constriction, the posterior gland often not reach- 
ing insertion of 12/13. Gizzard in xvii-xviii, muscularity occa- 
sionally extending slightly bej^ond insertion of 16/17 or 18/19. 
Intestinal origin in xv (18). Typhlosole begins in region of xxii- 
xxvi, abruptly or rather gradually as a flat band that shortly 
becomes vertical, reaching a height of 11/2-2 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 lonnhergi; may have a deep groove on the ventral 
face. The end normally is in region of xcvii-cv (cf. Table II). 

Subneural trunk quite small, adherent to nerve cord, some- 



1955 AMERICAN EARTHWORMS OF THE FAMILY LUMBRICIDAE I-II 9 

times quite unrecognizable. Last hearts in xi (18), the hearts 
of x-xi usually slenderer than the ones in ix, none found in v. 
Nephridial ducts pass into parietes in the b gap. 

Seminal vesicles three pairs, in ix, xi-xii (18, including the 
juvenile), those of xi sometimes (5) smaller to much smaller 
than the others. Male deferent ducts apparently slightly 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 lonnhergi) 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 lonnhergi. Glands, atrial and such as are 
present on the parietes in lonnhergi and species of Bimastos are 
lacking. 

Ahnormality 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 on xxxi-xxxiii as 
well as on ix 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 II) 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. Keproduction 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 
copulation by a July 31 worm (Linville) in which no iridescence 
was recognizable on male funnels. Funnel iridescence was bril- 
liant 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 clitellar 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 elitellum 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-xxxix and in DD on the left side of xxxv-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 lonnhergi, 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 Eisenia they still have not disappeared in some 
"formae" of a species. Presence of vesicles in x of one (or 
more?) of the types of pearsei, 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. lonnhergi 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 LUMBRICIDAE I-II 11 

Table I 

Typhlosole termination and segment number 

in Eisenia lonnhergi 



Typhlosole 


Atyphlo- 










ends in 


solate 


Number of 








segment 


segments 


segments 


Locality 


Eemarlcs 




60-61» 


1 


62 


Georgia 


Amputee 




61-64* 





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 


? 


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 


? 


f 


124 






Damaged 


110 


14 


124 


^ 






110 


17 


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 Eisenia carolinensis 



Typhlosole Atyphlo- 








ends in 


solate 


Number of 






segment 


segments 


segments 


Locality 


Bemarhs 


65* 


6 


71 




Amputee 


73 


23 


96 




Juvenile. Amputee? 


79 


21 


100 




Amputee 


80 


19 


101 




Amputee 


88 


26 


114 




Amputee 


82-89t 


11 


100 


Chapel Hill 


Amputee 


90 


32 


122 




Aniputee 


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, 
t 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. 



E V I O R A 

Mijiseiiiim of Compsirsitive Zoology 

Cambridge, Mass. November 18, 1955 Number 49 

THREE NEW SHARK RECORDS FROM 
THE aULF OF MEXICO 

By Henry B. Bigelow, 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 Eimopterus 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 E. 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 EtmOPTERUS Rafinesque 1810 

Type species Squalus spinax, Linnaeus 1758 

Whitley (1939, p. 266) has recently revived the genus Aca7i- 
thidium Lowe 1839, type species Centrina nigra Lowe 1834 
which, in 1839, Lowe renamed Acanthidium pusillum, thinking 
it separable generically from Etmopteriis 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 BREVIOBA NO. 49 

pied in Etmopterus by nigrum (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' B., 
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 
imsillus, 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. schultzi (Bigelow, 
Schroeder and Springer 1953, fig. 9G), with the stouter, more 
thorn-like denticles of E. hillianus Poey 1861 (Bigelow and 
Schroeder 1948, p. 489, fig. 92B), of E. polli (Bigelow, Schroeder 
and Springer, 1953, fig. 7 C) and of E. 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 E. 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- 

tGarman (1913, p. 223) credits the earliest use of niger in this coimectiou to 
(iiinner 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. 



1955 THREE NEW SHARK RECORDS FROM THE GULF OF MEXICO 




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4 BBEVIORA 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 
either 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 (1843, PI. 6) original 
illustration of it (as Acanthidium pusillum) ; also on the colored 
illustration by Braganza (1904, PI. 2, fig. 2, as E. 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. polli, 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 E. princeps. 

Two species of Etmopteriis, 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 {hiUianus 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 E. 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 Kafinesque 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, pi. 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, pi. 1, fig. 2, pi. 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 Muller 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. S. Nat. Mus. No. 157833 ; also skin of 
a male, 855 mm. long, eastern Atlantic, exact locality not known, 
Mus. Comp. Zool. No. 662. 



BREVIORA 



NO. 49 




Oj 


02 


Oj o 


s 






f^ n. 




s 


ti 


&t 


■-C 


s 




Cj 


to" 


^ CD 


0} 




-* 


^ ^ 


TJ 




w 




1 






S 




o 
be 




cT 


a 




ft --1 


c 




O 


-si 


.2 


cc 




rt 






fe 'O 


>;; 


'S 




i^ Is 


a> 




d 




re 

o 


_o 


I 




,r: 



g -e - ^ « cc 

i! S CO - ° ® 

o <v cc 'xi ^ 

^ ^ 00 ci t:3 +s 

»- -S !>; 5 9 S 

g S « -^ ^ « 

i t£ ^o o S o 



cq 



■iliV/ 






. .„ a - 1 1 

'f- fi^ ca ti 'tS -H 
12; rt Lo 



1955 THREE NEW SHARK RECORDS PROM THE GULF OP MEXICO 7 

The most evident diagnostic features that set apart C. granu- 
losus (with C. machiquensis Maul 1955) from the other two 
species of its genus {squamosiis Bonnaterre 1788 and uyato 
Rafinesque 1810) that are known from the North Atlantic 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 
careful examination under a lens (Fig. 2, 0). In uyato, 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. 

Lahial 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 pi. 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 1st 
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 1st 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; 
pelvics 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 ^3 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 
1/4 as long as eye, its anterior edge only slightly posterior to rear 
corner of eye. Nostrils a little less than i/^ 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 i/^ as 
great as from tip of snout to mouth. Mouth very low-arched, its 
gape when closed occupying about % 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 ii^fTsj 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 func- 
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 1/2 to % 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 9 

Denticles at mid-level of side below first dorsal fin avera^fing 
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 lYo times as long as between nostrils. Exposed portion of 
first dorsal spine about II/3 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 % as long as from tip of snout to level of 
5th 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 uyato 
which Rey (1928, p. 436) 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 occur 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 vicinitv of the Faroes. 



10 BREVIOKA 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 Pacific CeniropJiorus 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 
Dal ATI AS Lie HA (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 
Atlantic 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). 



REFEEENCES 

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. Scheoeder, and Stewart Sprinqek 

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. 
Bloch, M. E., and I. G. Schneider 

1801. Systema Ichthyologiae iconibus ex iliustratum. Ix + 584 pp.; 
110 pis.; 2 vols. 
Bonnaterre, p. J. 

1788. Ichthyologie in: Tab. Enc>c. Method. Trois regnes de la Nature. 
Ivi 4- 215 pp., pis. A, B + 1-100. Paris. 
Braganza, Carlos de 

1904. Esqualos obtidos nas Costas de Portugal. Keaultadoa das Invest. 
Sci. . . . Yacht "Amelia." Ichthyologia. II; 107 pp., 2 pis. 
Lisbon. 
Cloquet, Hippoltte 

1820. Le sagre, spinas niger. Diet. Sci. Nat., vol. 1, suppl., p. 93. 
Collett, R. 

1904. Diagnoses of four hitherto undescribed fishes from the depth.*! 
south of the Faroe Islands. Videns. Selskabs Porhand. Christi 
ania, 1904, no. 9, 7 pp. 
Garman, Samuel 

1913. The Plagiostomia. Mem. Mus. Comp. Zool., vol. 36, xiii -f 515 
pp., 77 pis. 
GtfNTHER, Albert 

1870. Catalogue of fishes of the British Museum, vol. 8, xxv -J- 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, Ix -f 1240 pp. 
Linnaeus, C. 

1758. Systema naturae, 10th Ed., vol. 1, 824 pp., Holmiae. 
Lowe, E. T. 

1834. A collection of fishes made in Madeira. Proc. Zool. Soc. London 

(1833), Part 1, 1834, pp. 142-144. 
1839. A supplement to a sjTiopsis 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 pis., London. 
Maul, G. E. 

1955. Five species of rare sharks new for Madeira. . . . Notul. Natur. 
Acad. Nat. Sci. Philadelphia, No. 279, 13 pp., 3 pis. 
MuLLER, Johannes, aud F. G. J. Henle 

1837. Ueber die Gattungen der Haifische und Rochen . . . Arch. Natur- 
gesch., Jahrg. 3, vol. 2. pp. 394 401, 434. 



12 BBEVIORA NO. 49 

PlETSCHMANN, 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. 
Kl., vol. 117, pt. 1, pp. 637-710, pis. 1, 2. 

PoEY, Felipe 

1858-1861. Meinorias sobre la historia natural de la isla de Cuba, vol. 
2, 442 pp., 19 pis. 
Rafinesque, Constantine Samuel 

1810. Carratteri di alcuni nuovi generi e nuovi specie di animali c 
piante deUa Sicilia. pp. iv, 105, 20 pis. Palermo. 
Rey, Luis Lozano 

1928. Fauna Iberica. Feces. Vol. 1, 690 pp., 20 pis. Inst. Nac. 
Cienciaa, Madrid. 
Tanaka, Shigeho 

1912. Figures and descriptions of the fishes of Japan. Vol. 5, pp. 71- 
86 4- [1], pis. 21-25; vol. 6, pp. 87-108, pis. 26 30. 
Whitley, Gilbert 

1939. Studies in ichthyology no. 12. Rec. Australian Mus. Vol. 20, No. 
4, pp. 264-277. 



B R E 

Mnaseum of Compsirsitive Zoology 



Cambridge, Mass. November 30, 1955 Number 50 

NEW FEOaS OF THE GENERA 

ASTEBOPHRYS AND OREOPHBYNE FROM 

NEW GUINEA 

By Aethue Loveeidge 

During the past year the Museum of Comparative Zoology has 
received from Mr. Kenneth R. Slater some interesting microhylids 
moXudm^ Asterophrys rufescens (Macleay), valvifera (Barbour), 
turpicola (Miiller) ; Xenohatrachiis rostratus (Mehely) ; Mefo- 
postira oceUata Mehely and Cophixalus oxyrhinus (Boulenger), 
taken at Omati, near Port Moresby, Papua. 

There was also a large and handsome frog which keyed down 
to Asterophrys hoettgeri (Mehely) in the synopsis to that genus 
furnished by H. W. Parker in his "Monograph of the Microhy- 
lidae" (1934:60. Brit. Mus.). However, A. hoettgeri comes from 
Halmahera Island in the Moluccas 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. no v. 

Holotypc. Museum of Comparative Zoology No. 28205, an adult 
9 , from Omati, near Port Moresby, Papua. Collected by Kenneth 
R. Slater between January and April, 1954. 

Diagnosis. Apparently closely related to A. hoettgeri (Mehely) 
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 tubercles. 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 
glandular folds ; posteriorly an obsolete, hair-like, light vertebral 
line is barely distinguishable ; thighs heavily blotched with black, 
the elongate blotches separated by irregular white lines; on the 
tilna these light lines suggest obsolete crossbars. 

Below, pale brownish spotted w4th darker browai 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 9 . 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 3 

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 
bodv cavitv. 



Oreophryne parkeri sp. nov. 

Cophixalus geislerormn Loveridge (not of Boettger), 1948, Bull. Mus. Com]). 
Zool., 101: 423. 

Holotype. Museum of Comparative Zoology No. 13964, an adult 
9 , 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 a7ithomji-hiroi section but are not very 
closely related to either species. Oreophryne anthotiyi (Boulen- 
ger), of which we have a cotype, is a more robust frog than the 
gravid holotype of parkeri, from which it differs slightly in almost 
every character, the most conspicuous being the coloring of the 
underside and : 

Eye separated from tympanum by a distance equal to IV^ times the horizontal 
diameter of the latter. Total length of adult 45 mm anthonyi 

Eye separated from tympanum jjy a distance equal to only Ys the horizontal 
diameter of the latter. Total length of gravid 5 30 mm parkeri 

0. parkeri agrees with hirui, of which we have a series, in size 
and many other respects, but differs sharply in tympanic char- 
acters. 

Tympanum scarcely distinct, very small, % to % 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.) hiroi 

Tympanum very distinct, large, % the eye diameter; disk of third digit nearly 
% the eye diameter; ratio of tibia length to head width at commissure of 
mouth 1.37 to 1.43 (2 ex.) parJceri 

The head of parkeri, as Dr. R. G. Zweifel has pointed out to 
me, is relatively small compared with those of other Oreophryne. 



4 BBEVIOBA 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). 

Color. Above, brow^n, 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 9 . 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 
hatesi 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: 303-430), I erroneously referred 
these two frogs to Cophi.mlus geislerorum Boettger. More recent- 
ly, when Dr. R. G. Zweifel of the American Museum of Natural 
History was w^orking over our material, he redetermined them as 
Orcophryne of no known species. I then submitted them to Dr. 
11. 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 0. 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. 



E V I O R A 

Mmseitam of Coimparsitive Zoology 

Cambridge, Mass. January 27, 1956 Number 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 Oligohunis 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. 



BREVIORA 



NO. 51 



Family Mustelidae 
Subfamily Mustelinae 

MiOMUSTELA ( ? ) Sp. 

Bef erred material. M.C.Z. No. 7016 (Fig. 1), incomplete left 
mandible with Mi present and alveoli for C, Pi, P2, P3, and M2 
(incomplete). 

Horizon and locality. Arikareean Miocene. Thomas Farm, Gil- 
christ County, Florida. 




'.^o^C^E^^ 




Miomustelai'l) sp. Labial, occlusal and lingual vieAvs. M.C.Z. 7016. X 3. 



Characters. Closest to Miomustela madisonae (Douglass 1903; 
Hall 1930) among previously known forms but differing as fol- 
lows : Ml 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 



double rooted and their arrangement indicates a crowded tooth 
row. Ml 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. Mo is single rooted. The ascending ramus of the 
jaw starts just posterior to Mi, causing M2 to be tilted upward 
and forward. The masseteric fossa extends forward to the an- 
terior rim of the alveolus of M2. Mental foramina are present 
below the centers of Pi and P2. 



MEASUREMENTS (IN MILLIMETERS) OF MANDIBLE a TOOTH 


LENGTH OF TOOTH ROW AS INDICATED BY ALVEOLI 

f AMTPBinC lOlAl^mM OC D Tn DnCTCDinO KilADriM rtc vt \ 


18.0 






5.7 






6.0 






2.5 




M, , HEIGHT OF PROTOCONID FROM LABIAL MARGIN OF ALVEOLUS 


5-0 ; 


M, , " " METACONID " LINGUAL 


2.5 


M, , " " FWRACONID " " " " " 


3.5 


Mj , " " HYPOCONULID" " ' 


L7 



DISCUSSION 

Plesictis (including Miistelavus) agrees with M.C.Z. No. 7016 
in the general structure of Mi, 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 Mo (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 Ml in a large series of Maries 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 distinct 
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 Miomiistela. The figure was drawn by Miss Patricia 
Washer. 

EEFEKENCES 
Clark, J. 

1937. The stratigraphy and paleontology of the Chadron formation 
in the Big Badlands of South Dakota. Ann. Carnegie Mus., 25: 
261-350. 

Douglass, E. 

1903. New vertebrates from the Montana Tertiary. Ann. Carnegie 
Mus., 2: 145-199. 

FiLHOL, H. 

1879. Etude des mammiferes fossiles de Saint-Gerand le Puy (Allier). 
Bib. Ecole Hautes Etudes, Sect. Sci. Nat., 19: 1-252. 

Hall, E. E. 

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 Chakdin, P. 

1914. Les carnassiers des Phosphorites du Quercy. Ann. Paleont., 9: 
103-192. 



1956 A SMALL MUSTELID FROM THE THOMAS FARM MIOCENE 5 

ViRET, J. 

1929. Les faunes de mammif^res de I'Oligocene superieur de la 
Limagne Bourbonnaise. Ann. Univ. Lyon, (n.s. 1), sci. med., 
47: 5 305. 

White, 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. 



E V I O R A 

Mmseiiiiini 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 HYLA 

By Walter 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. 

Bufo 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. ameri- 
canus (1), B. w. woodhousei (2), B. w. fowleri (3), B. valliceps 
(2), B. yunctatus (1), -S. compactilus (1), 5. 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 praevius 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 3 

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 
curvature 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, 
i.e., that the ilial shaft in praevius is, for the most part, less 
curved 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 fra'gmental 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. fowleri. 

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 BRBVIORA NO. 52 

ments of the fossil species are now available. These include three 
frontoparietals (UF 9892), two temporals (UP 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 



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 
ilial prominence, and is y^vy much strengthened by the shape and 







Fig. 1. Shape of the cranial 
houstonensis (after Sanders, op. 
rius; D, Bufo w. fowJeri. 



crests in four species of Bufo. A, Bufo 
rit.); B, Biifo i. ferrefitris; C, Bufo prae- 



development of the cranial ridges. A few of the characters, such 
as the height of the urostyle crest and the arching of l^e 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 f*stablished by early Miocene. 



6 BREVIORA NO. 52 

Ran A sp. 

This genus has already been reported from the Thomas Farm 
by Tihen {op. cii.). Two additional ilia are now available (UF 
5919). These remains are too frag^llentary 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, etc. They are 
not as readily referrable to the group including heckscheri, cates- 
heiana 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 (including h. holbrooki, h. hurteri and couchi) and Spea 
(including multiplicafns, homhifrons and hammondi). 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 dorsallj^, 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 
holbrooki. • 



1956 



3II0CENE ANURANS FROM FLORIDA 



Whether or not the Miocene Scaphiopits was identical to the 
species holhrooM cannot be absolutely determined at this time. 
However, the fossil form is clearly referrable to the subgenus 
Scaphiopus. Whether or not the subgenus Spca had differenti- 
ated by this time, or whether the former is ancestral to the latter 
is, of course, unknoAvn. 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. -J,. CJoiupiii isou ot till f u>m1 iiiifi-oliylitl iliuiii fioiu tli 
locality (A) with lli.it of .1 u-i cat ^pefiiiicii uf Mii-rnlti/ld (■< 
from Gainesville, Florida. The line represents 1 iinn. 



Fan. 



MiCROHYLA sp. 

Of considerable interest is the fact that one fragmentaj 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- 
gestiAC of this genus. In addition, the acetabular cup is very 
similar to that in 3Iicro]nila, in that the edges of the cavity are 
considerably raised above the surface of tbe 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 w^eakly developed, 
Ijeing 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 Fseudacris 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 ilium, 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 ilia from the 
same locality and horizon. 

The ilia of the genera Hyla, Acris and Pseudacris are easily 
separated from those of Eana and Eleuth erodactylus by the ab- 
sence of a dorsal crest. From Microhyla, Bufo and Scaphiopvs 
they are readily distinguished by the well-developed dorsal prom- 



1956 



MIOCENE ANURANS FROM FLORIDA 



iiienee, directed dorsolaterally to laterally, and by the slender- 
ness of the shaft itself. 

The ilia of Hijla (joiui have been compared with the same ele- 
ment in the following- uenera and species: Hyla cinerea (12), H. 
squirello (6), FT. fonorali)^ (2), //. crucifer (2), H. versicolor 




E F 

Fig. 3. Interspecific variation in the ilia of Hylidae. A, 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 oimata, 7 mi. northeast of Gainesville, Florida; E, 
Hyla crucifer hartramiana, 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 nigrita (4), P. ornata 
(3), P. hrimleyi (1), Acm 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 goini. 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. hrimleyi 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, goini 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 case, 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 ANURAN8 FROM FLORIDA 11 

ditioual 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 criticism 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. Ilighton for the loan of a 
ai umber of anuran skeletons. 



LITERATURE CITED 
Brattstrom, B. 

]953. The amphibians and leptiles from Eaneho La Brca. Trans. 8an 
Diego Soc. Nat. Hist., 11 (14) :365-92. 
Cooke, C.W. 

1945. Geology of Florida. Bull. Florida Geol. Surv., 29:1-339. 
Sanders, O. 

1953. A new species of toad with a discussion of morphology of the 
bufonid skull. Herpetologiea, 9(l):25-47. 
Schmidt, K. P. 

1953. A checklist of North American amphibians and reptiles. Amcr. 
Soc. Ichthyologists and Herpetologists. Univ. Chicago Press: 
Chicago, Illinois. 

Taylor, 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. 

TlHFX, J. 

1951. Anuran remains from the Miocene of Florida, with the descrip- 
tion of a new species of Biifo. Copeia, 1951 (3 ) : 230-35. 

1954. A Kansas Pleistocene herpetofauna. Copeia, 1954 (3). -217-21. 



E V I 

Mmseiimi of Comparative Zoology 

Cambridge, Mass. April 6, 1956 Number 53 

FOOD-FINDING BY A CAPTIVE PORPOISE 
(TURSIOPS TRUNCATUSy 

By William E. Schevill and Baebara Lawrence 



Introduction 


1 


Experimental arrangements 


2 


Passive location 


4 


Active or echolocation 


4 


Echolocation to determine presence of fish 


6 


Echolocation to distinguish between alternate feeding 




places 


7 


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 Tur- 
siops truncatus was confronted with the problem of finding un- 
familiar food under varying and sometimes rather complicated 
circumstances, 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 BKEVIORA 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 cm. 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 ont" 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 forced into situations he disliked, 
the tests we devised were as simple as were consistent with get- 
ting reliable answers. Under these circumstances, it soon bv^came 
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 Tvas 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 cm. 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 cm. and often less than 45 cm. ; the very 
unusual maximum was 85 cm. on 19 September. At 11 o'clock 
on a typical sunnj^ day measurements made by diving after dark- 
adaptation gave a Secchi disk reading of 23 cm. from the diving 
mask at a depth of 2.4 m., and 61 cm. at the surafee. 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. 



L 



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 cm. 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 ''creaks" 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. Bv "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," etc., by various authors, such 
as McBride and Ilebb 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 communcative. The acoustic 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 BEEVIORA 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 2V^ 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 51/^ 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 BBEVIOBA 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 cm., 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 difi'erent occasions, and though his responses 
varied, they made a very neat pattern. The first time we tried it, 
and when w^e 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 any^vhere over a six meter stretch. Nor could vision 
have helped, because he was never nearer than 61/2 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 
2 m. 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 (cf. 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 BBEVIOEA 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 cheek 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- 
customed 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 ou 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 eye 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. 

E"'ish 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 occasions 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 
cm. 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 cm. away. The other occasion was one afternoon when his 
left eye failed; then, lie 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 like 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 Tiir slops are accustomed. 



1956 FOOD-FINDING BY POEPOISE 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. cit., p. 229) "the complete absence" of evidence for echoloca- 
tion although we cited some, unrecognized, at the bottom of page 
227.^ 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 acoustic 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 maj^ 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. 

Echolocation 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 mam' 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. E. 

1953. Acoustic orientation in the oil bird. Sieatornis. Proc. Nat. Acad. 
Sci., 39 (8) : 884-893. 



1956 FOOD-FINDING BY PORPOISE 15 

Griffin, D. E., and Galambos, R. 

1941. The sensory basis of obstacle avoidance by flying bats. Jour. 
Exper. Zool., 86: 481-506. 
Kellogg, W. N., Kohlee, 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. 
McBride, Arthur F. 

(In press.) Evidence for echolocation by cetaceans. Deep-Sea Research, 
3: (2) 
McBride, A. F., and Hebb, 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. 

Wood, F. G., Jr. 

1952. Porpoise sounds. A phonograph record of underwater sounds 
. . . Tursiops truncatus and Stenella plagiodon. Published by the 
Marineland Research Laboratory. 
Wood, F. G., Jr. 

1954. Underwater sound production and concurrent behavior of captive 
porpoises, Tursiops truncatus and Stenella plagiodon. Bull. Mar. 
Sci. Gulf and Caribbean, 3 (2) : 120-133. 



E V I O R A 

MmseiMn of Compairative Zoology 



Cambridge, Mass. June 6, 1956 Number 54 



A REVISION OF THE GENUS BBACHYMELES 

(SCINCIDAE), WITH DESCRIPTIONS OF 

NEW SPECIES AND SUBSPECIES 



By Walter C. Brown 
INTRODITCTION 



The genus Brachymeles was erected by Dumeril 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. sahadenhergi were described by Fischer (1885) ; 
and Taylor described B. elerae and B. hurksi (1917), B. suluensis 
and B. vermis (1918), B. houlcngeri (1922) and B. pathfinderi 
and B. ivrighti (1925). In 1922 Taylor correctly assigned B. 
suluensis to the synonymy of B. gracilis and also pointed out that 
B. houlengeri is closely related to the latter. It now appears that 
B. houlengeri is probably best regarded as a geographically 
isolated population which is only subspecificaliy 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. schadenhergi. A re- 
examination of the type of B. bonitae, for which assistance I am 
deeply indebted to Dr. Jean Guibe, indicates that B. hurksi is 



Menlo College and Natural History Museum, Stanford University 



2 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. honitae, 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. schadenhergi 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. 
P^rt 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 

(luently 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 

Illinois 
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 Dumeril and Bibron 

BmchymtUs Dumeril and Bibron, 1839, Erpt. Gen., 5: 776 (type species: 

Brachymclcs honitae Dumeril and Bibron, 1839, by monotypy). 
Senira Gray, 1845, Cat, Lizards Brit. Mus. : 98 (type species : Senira bicoloi 

Gray, 1945, by monotypy). 
Brac'hymelvs Agassiz, 1846, Nomeu. Zool. Index Univ.: 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 Brachymeles. 

1. Limbs absent B. vermis 

Limbs present - 

•2. Limbs pentadactyl 8 

Either fore or hind limbs or both with less than 5 digits 3 

3. Digits entirely absent or a single vestigial claw present B. honitae 



t 



4 BREVIORA XO. 54 

Two or more digits present (frequently represented only as 
clawed vestiges) 4 

4. Limbs with 2 digits B. samarensis sp. nov. 
Limbs with more than 2 digits 5 

5. Limbs with 3 digits B. tridactylus sp. nov. 

Limbs with more than 3 digits 6 

6. Fore limbs with 5 digits ; hind limbs with 4 B. pathfinderi 

Fore and hind limbs with 4 digits 7 

7. Midbody scale rows -22-; scale rows between the parietals and 

the base of the tail 83-85 B, elerae 

Midbody scale rows -28-; scale rows between the parietals and 
the base of the tail -102- 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 

of the tail 92-94 B. licolor 

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 wdth no or very few dark spotted scales 

B. schadenberfji 
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; loAvev 
lateral surfaces and frequently venter with numerous dark 
Spotted scales (except in B. qracilis houlengeri) B. gracilis 

Brachymeles vermis Taylor 

Brachymeles vermis Taylor, 1918, Philip. Journ. Sci., 13:255 — Bubuan Id., 
Sulu Archipelago. 
Material examined. Jolo Id. 5 (C.A.S. 60720-22, 60857-58) ; 
Bubuan Td., Sulu Arch. 1 (C.A.S. 62489). 



1956 REVISION OP" THE GENUS BRACHYMELES 5 

Diagnosis. Habitus very slender; suout to vent length 64-76 
nun. for 3 mature specimens; limbs absent; no ear opening; no 
postnasal; 22-24 scale 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 Dumeril and Bibron 

Brachymeles bonitae Dumeril and Bibron, 1839, Erpet. Gen., 5:777 — Manila, 

Luzon Island. 
Braclnjmeles burTcsi Taylor, ]917, Pliilii). .Tourn. Sci., 12:275 — Sumagui, 
Mindoro Island. 

Material examined. Holotype (examined by Dr. Jean Gnibe) ; 
Mindoro Id. 3 (C.A.S. 62064, C.N.H.M. 22525, S.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.S. 62278-79, 62575). 

The single specimen (No. 1151, now C.A.S. 62578) referred to 
B. bonitae by Taylor (1917, 1922), Avith which he compared B. 
hurksi 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 betAveen 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 roAvs is actually an indiAndual aberration or char- 
acterizes a more or less isolated population of B. bonitae cannot 
l)e determined at this time. 

Diagnosis. Habitus A-ery slender; snout to vent length 57-82.5 
mm. for 6 mature specimens ; limbs reduced to vestiges without 
CA-idence of toes or Avith a single claAv, 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 roAA^s around the middle 
of the body for 10 specimens ; 100-106 scale rows along the mid- 
dorsal line betAveen 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 mid dorsal 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 lighter. 

Relationship. It would appear to be more closely related to 
B. elerae than to B. honitae 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 OP 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. honitae 
is 100-106. 

Range. Known only from the type locality. 

Bracuymeles 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. 




Fig. 1. Dorsal view of head of Brnchymeles tridactylus. 

Paratijpes. 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 Brachyineles, with short limbs, possessing 
three reduced, clawed digits on both the fore and hind limbs; 
scale row^s 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 w^ith 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 ; j^arietals meeting posterior to the interparietal ; a 
distinct pair of nuchals in only one specimen ; two f renals, 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 reduced, 
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 holofype. Total length 106 mm. ; snout to 
vent 65 mm. ; axilla to groin 40 mm. ; snout to forelimb 13 mm. ; 
length of hind limli 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. bo'nitae from which it differs primarily in less reduced 
limbs and the presence of three stump-like, clawed digits on each 
limb. B. honitae has lost all the digits or retained a vestige of 
only one at the tips of the minute limbs. 

Range. Known o\\\y from the type locality. 

Brachymeles elerae Taylor 

Brachymeles clcme Taylor, 1917, Philip. Journ. Sci.. 12:273 — Philippine 
Islands. 

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 gi^eatly 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 Province, 
Luzon Island. 



1956 REVISION OP THE GENUS BRACHYMELES 9 

Brachymeles wrighti Taylor 

Brachymelcs wrighti Taylor, 1925, Philip. Journ. Sei., 26:106 — Trinidad, 
northern Luzon Island. 

Material examined. Holotype (M.C.Z. 26589 : examined by Mr. 
Arthur Loveridge). 

Diagnosis. Habitus slender; snout-vent length 130 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 pathfinderi 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. 

Ra7igc. Known only from the type locality. 

Brachymeles bi color (Gray) 

Senira blcolor (part), Gray, 1845, Cat. Lizards Brit. Mus.: 98 — Philippine 
Islands. 

Material examined. Holotype 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. Early records are given as from the Philippine 
Islands. More definite information as to the distribntion of this 
species can not he stated at this time. 

Brack YMELES gracilis (Fischer) 

Several populations of this species exhibit rather conspicuous 
and constant differences in color pattern and in certain instances 
in scalation 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. hoidengeri 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 bon- 
lengcri 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). 

Braciiymeles gracilis gracilis (Fischer^ 

Eumeces (Kiopa) gracilis Fischer, 1885, Jahrli. Aviss. Aiist. Hamburg, 2: 85 — 

Mindanao Island. 
Braciiymeles sulnensis Taylor, 1918, Philip. Journ. Sei., 13:254 — Bubuan 

Island, Sulu Archipelago. 



1956 REVISION OF THE GENUS BRACK YMELES 11 

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 scale. 

Range. Known definitely from the 8ulu Archipelago and 
Mindanao. 

Brachymeles gracilis boulengeri Taylor 

Brnchymeles boulengeri (part) Taylor, 1922, Philip. Bur. Sci. 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, pis. 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 



NO. 54 



Table 1 
Comparison of subspecies of Brachymeles gracilis 







Dorsal scale rows 






No. of midbody 


between parietals 


Dorsolateral 




scale rows 


and base of tail 


stripe present 


B. g. gracilis 


R = 24-26 


R = 64-69 


Yes 


(Mindanao and 


M = 25.6 ±0.197 


M = 66.4 ± 0.201 




Basilan) 


N=36 


N=36 




B. g. boulengeri 


R = 24-26 


R = 64-66 


Yes 


(Polillo) 


M = 25.7 ± 0.305 


M = 65.2 ± 0.367 






N = 6 


X = 6 




B. g. boulengeri 


R = 26,26 


R = 63,65 


Yes 


(Luzon) 


N = 2 


N=2 




B. g. boulengeri ( '? 


) R = 24-28 


R = 63-66 


Yes 


(Bohon 


M = 26.2 ±0.158 


M = 64.0 ± 0.199 






N = 21 


N =21 




B. g. taylori 


R = 24-28 


R = 64-68 


Xo 


(Negros) 


M = 26.5 ± 0.097 


M = 65.6 ± 0.150 






N=49 


X— 49 




B. g. taylori 


R =26 


R =66 


No 


(Mindoro) 


N = l 


X = l 





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, pi. 7), not uniform but each scale 
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 row's posteriorly) is somewhat variable 
usually near Monkey skin or Army brown (Maerz and Paul, pis. 
5 and 6). Five to 7 lateral rows exhibit prominent, dark brown 
spots; the venter is Rose tan (Maerz and Paul, pi. 12), occasion- 
ally with small broAvnish 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. 

lirachymeles Ixnilengeri (i)art) Taylor, 1922, Philip. Bur. Sci. Pub. No. 
17:246. 

Holotype. N.H.M.S.U., No. 18615, a male, collected by W. C. 
Brown and D. S. 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 taylori. 

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, 
V2 km. S.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, 
llaya 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, pi. 12), frequently with scat- 
tered to numerous small brown spots. 

Bange. 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 scale 
rows between the parietals and the base of the tail, and the 
presence of a more or less distinct, 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. 

if larger samples do indeed prove the Luzon, Negros and Jolo 
populations to be the same from a taxonomic standpoint, and 
that of Palawan, Basilan, Mindanao. Ijeyte 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 
polytypic 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- 
eall}^, which have maintained disjunct distributions within this 
archipelago. The first interpretation is accepted with some reser- 
vations in this paper. 

Brachymeles schadenbekgi schadenbergi (Fischer) 

Kaineces (Biopa) schadenhergi Fischer, 1885, Jahrli. wiss. Aiist. Hamburg, 2: 
87 — • Miudanao 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, 61968-64) ; Leyte Id. 3 (N.II.M.S.U. 18701; C.N.H.M. 42779, 
42792); Bohol Id. 5 (N.II.M.S.V. 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 d= 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, pis. 2 and 3). The venter is 
whitish, lightly tinted with the above shades. 

Ratif/f. Known from the Sulu Archipelago, Mindanao, Leyto 
and Bohol Islands (probably also Palawan Island). 



16 



NO. 54 



Table 2 
Comparison of subspecies of Brachynieles schadenbergi 



Dorsal scale rows 
No. of midbody between parietals Dorso-lateral 
scale rows and base of tail stripe present 



B. s. schadenbergi 


E = 26-28 


E = 67-71 


Xo 


(Mindanao and 


M = 26.5 ±0.161 


M = 68.8 ± 0.250 




Basilan) 


N = 25 


X=25 




B. s. schadenbergi 


E =28 


E = 68,68,70 


No 


(Leyte) 


X = 3 


X=3 




B. s. schadenbergi 


E =28 


H. = 70,71 (4) 


No 


(Bohol) 


N=5 


X=5 




B. s. schadenbergi ( 


-') E =28 


E = 70 


No 


(Palawan) 


X = l 


X = l 




B. s. talinis 


E =28-32 


E = 67-70 


Yes 


(Negros) 


M = 30.4 ± 0.276 


^r = 68.2 ± 0.193 






X =11 


X = 12 




B. s. talhnis (?) 


E = 29,30,32 


E 72,72,73 


Yes 


(Jolo) 


X =3 


X=S 




B. s. talinis ( ='; 


R = 26,28 


E = 69,70 


Yes 


(Luzon) 


X ^ 2 


X = 2 





E = range, M = mean, X = number of specimens 

Brachymeles schadenbergi talinis subsp. nov. 

Holohjpe. 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, 
Dec. 1953, March, 1955: 18360-62, ridge "south of the Maite 



1956 REVISION OF THE GENUS BRACHYMELES 17 

Kiver, east slope of Cuernos de Negros, about 6-7 km. west of 
Luzuriaga, Negros Oriental, Nov., 1954, March, 1955; M.C.-Z. 
54257, 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 Brachynielcs 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 
scale rows at the shoulder), dorso-lateral light stripe, reddish in 
life, sometimes fading in older specimens; second pair of chin 
shields generally wdder, 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 BracJujinelc.s scliadnibergi talinlf;. 

Color (freshly preserved material). Dorsum is dark reddish 
brown, the lateral margins of the scales tending to be dull red- 
dish producing an indistinct, lined 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 Avhite or whitish tan in 
preservative.) These stripes begin just posterior to the eye and 
extend to the region of the hind limlis or the base of the tail. 



18 BREVIORA NO. 54 

They are most unifonu and conspicuous in juveniles, tending to 
become 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. Knowai 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 Guibe, Museum 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 honitae, Bracliymeles tvrighti, Brachymeles pathfinderi and Brachymele.s 
bicolor; to Dr. Eobert F. Inger, Chicago Natural History Museum and Dr. 
Joseph R. Slevin, California Academy of Sciences for permitting me to 
e.xamine material of this genus in the collections of their respective institu- 
tions. I also wish to tliank Dr. Inger for his kindness in suggesting that I 
describe at this time a new species from Samar Island which is represented 
Ijy a unique specimen in the collections of the Chicago Natural History 
Museum. 

This study avus 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 

BouLEXGER, George Albert 

1887. Catalogue of the lizards in the British Museum. Ed. 2. London, 
vol. 3, i-xii -f 575 pp., pis. 1-40. 

Duml'ril, Andre Marie Const.\nt, and Gabriel Bibron 

1839. Erpetologie general. Paris, vol. 5, i-viii -f 855 pp., pis. -IS-SS. 



1956 REVISION OF THE GENUS BEACHYMELES 19 

Fischer, Johann Gustav 

1885. Ichthyologische uud herpetolog'ische Bemerkungen. Part IV: 
Uber eine Kollection von Amphibien und Eeptilien von Mindanao. 
Jalirb. 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. 

Maerz, A. and M. Eea Paul 

1930. A dictionary of color. New York, McGraw Hill Book Co., i-vii + 
207 pp., pis. 1-56. 

Taylor, Edward Harrison 

1917. Brachymeles, a genus of Philippine lizards. Phil. Journ. Sci., 
vol. 12, pp. 267-277, pi. 1. 

1918. Reptiles of the Sulu Archipelago. Phil. Journ. Sci., vol. 13, pp. 
233-267, pis. 1-3. 

1922. The lizards of the Philippine Islands. Publ. 17, Phil. Bur. Sci. 

Manila, pp. 1-269, pis. 1-23. 
1925. Additions to the herpetological fauna of the Philippines, IV. 

Phil. Journ. Sci., vol. 26, pp. 97-111. 



E V I O R A 

Miuseiim of Cooiparative Zoology 

Cambridge, Mass. June 6, 1956 Number 55 



REDISCOVERY OF HYLA DOESALTS 

AND LECHBIODUS PAPUANUS 

IN NEW GUINEA 

By Arthitr IjOveridge 



Last year I had occasion to describe a new microhylid discov- 
ered by Mr. Kenneth R. Slater at Oniati, Papua. More recently 
we have received part of a collection made by Mr. Slater durino- 
a trip up the Aramia River. This river lies somewhat to the 
north of Katow (i.e. Binturi River, Papua), a locality from 
which Macleay (1877, Proc. Linn. Soc. New South Wales, 2. pp. 
135-138) described a number of amphibians, among them Litoria 
dor ml u Macleay, now known as: 



Hyla dorsalis (Macleay) 

This hylid, so far as I know, has not been taken again during 
the period of almost eight}' years that has since elapsed. Among 
the Slater material is a 20 mm. gravid 9 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 lines," 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 line from snout to anus. On either eyelid are two tiny 
tubercles that, if constant in the species, may well have been 
overlooked by Macleay. 



NO. 55 



Lechriodus papuanus (Roux) 



The same collection contained half a dozen frogs (M.C.Z. 
28382-7) bearing- a distinct resemblance to Cornufer, bnt 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 
papua7ius (Roux) which was described in 1927 from a single 
juvenile ? , 23 mm. in length, 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 / — \ -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 ? 9 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 eanthus 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 $ 5 from 
New South Wales). Range: New South Wales; Queensland; 
and (/id!e 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 LECTIRIODUS PAPUANUS 3 

point above elbow a broad or narrow blackish brown line 
wliich may break up into a series of black specks; lower lip 
white; an indistinct interorbital skin fold (4 S $, 2 9 9 
from Aramia Eiver). Range: British and Dutch New Guinea .... 

/. pap nanus 

Our knowledge of /. papuanus, hitherto derived from a single 
immature 9 , is naturally extended by the Aramia River series 
of adult $ S and 9 9 ; 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 v^_^'.shaped 
interorbital fold ; from the eye a supratympanic 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, 
witliout 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, 
21/2, their tips slightly 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 bellv smooth. 



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 $ $ , 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 5 , 18 mm. ; in 9 , 20 mm. ; 
length of hind limb from anus to tip of longest toe in $ , 11 
mm. ; in 9 , 86 mm. In size, therefore, papuanus differs little 
from fletcheri. 

Stomach contents. That of the S was found by my colleague 
Dr. W. L. Brown to consist of a ehrysomelid beetle, a termite's 
head, and remains of ants referable to the genus Odontomachus. 



E V I 

Musemiim of Compsirative 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 : hal)itus ; color pattern ; dorsal, 
ventral and midbody scale 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 occiduiis 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 he seriously regarded. 



2 BREVIOKA NO. 56 

Key to the Jamaican and Cayman, forms of Celestus: 

1. Adpressed limbs meet or fail to meet bj' less thau 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 3 

L'. 19 or fewer subdigital lamellae on fourth toe of hind foot. 

Throat mottled hewardii Gray 1845 

Range : Central and western Jamaica and Portland Point. 
21 or more subdigital lamellae on fourth toe of hind foot. 
Throat not mottled duquesneyi Grant 1940a 

Range: Portland Point only. 

3. 89-116 dorsal scales, occipital to point above vent 4 

125-139 doreal scales, occipital to point above vent 

harhouri Grant 1940li 

Range: 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 

cnisculus crusculus Garman 1187 

Range : all of Jamaica, at least in the lowlands. 

Tail dorsally lighter than back and without pattern 

cnt^cnlus 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 
grouping, 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 misiclentified as hewardii but proved easily separable 
from a hewardii with the same locality data despite the loss in 
45181 of the blue tail so much stressed by Grant as diagnostic 
for dnquesneyi. 

In the second, short-legged group the series of C. harbouri 
studied included the type and 13 specimens collected by Bar- 
l)our,' all from Mandeville, plus a single specimen from Fern 
(lulley. One character, dorsal scale count, readily separates C. 
harhouri from C. crusculiis? This difference is very marked and 
shows no overlap. Crusculus has the lowest dorsal scale count of 
any Jamaican Cclcstus (mean 101.52, range 89-116) ; harbouri 
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 harbouri consists of a 
series of dark brown chevrons with the Vs 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 harbouri 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." 

r. crusculus is a variable species also in dorsal, ventral, and 
midbody scale 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 ])e distinguished from that at Kingston, the 

1 These were labeled by him as impressiis Cope. The status of this name is very 
much iu doubt and its disposition will depend on a new evaluation of the types 
in the Philadelphia Academy. 

2 Ventral scale 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 harhouri bears rnther larjre pearly spots as compared with 
smaller punctations in cruHmthm. 



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 Ecclesdown 
(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 (CD.) 
were calculated for Ecclesdown-Boston Bay and Balaclava com- 
pared to both Mandeville and the species as a whole : 

Boston Bay-Ecclesdown Balaclava 

Mandeville CD. = 1.57 CD. = 2.32 

entire species CD. = .676 CD. =: 1.01 

As is to be expected, as compared to the entire species, the CD. 
is not significant. The Ecclesdown-Boston Bay and Balaclava 
populations are sufficiently different from the Mandeville popula- 
tion as determined by the CD. 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 ynaculaius 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 eon- 



1956 JAMAICAN AND CAYMAN CELESTUS 5 

specific with Jamaican populations but because of the tail color 
difference it may be recoo:nized 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 hewardii group 
(long-legged group) made up of C. hewardii and C. duquesneyi. 
and the crusculus group (short-legged group) comprising C. 
crusculus and C. harhouri. 

3. C. hewardii, C. crusculus and C. harhouri are sympatric at 
Mandeville. 

4. C. hewardii and C duquesneyi (sympatric at Portland 
Ridge) can be distinguished by habitus, coloration, lamellar connt 
and number of chin shields. 

5. C. harhouri and C. crusculus can be distinguished by dorsal 
scale 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. 

1 wish to express my thanks to Dr. Ernst Mayr and Dr. E. E. 
Williams for their generous and valuable advice, to Mr. Benja- 
min Slireve who made the scale counts and measurements on the 
specimen of Celestus harhouri from Fern Gulley, and to Mr. 
Garth Underwood for useful criticism of the manuscript. 

LITEEATURE CITED 

CopK, E. D. 

18G8. An examination of the Eeptilia and Batrachia obtained by the 
Orton Expedition to Ecuador and the Upper Amazon with notes 
on other species. Proc. Acad. Nat. Sci. Phila. : 96-140. 



6 BREVIORA NO. 56 

Gabman, S. 

1887. On West Indian Geckonidae and Anguidae. Bull. Essex Inst., 19: 
17-24. 

1888. Eeptiles and batrachians from the Caymans and from the 
Bahamas. Bull. Essex Inst., 20: 103-116. 

Grant, C. 

1940a. Notes on the reptiles and amphibians of Jamaica, with diagnoses 

of new species and subspecies. Jamaica Today: 151-157. 
1940b. The Reptiles in Lynn and Grant: The herpetology of Jamaica. 

Bull. Inst. Jamaica, Sci. Ser., 1: 1-65. 
1940c. The herpetology of the Cayman Islands. Bull. Inst. Jamaica, Sci. 

Ser., 2: 1-65. 

Gray, J. E. 

1845. Catalogue of the specimens of lizards m the collection of the 
British Museum. London, 289 pp. 



E V I 

MmseiLim of Comparative Zoology 

Cambridge, Mass. JuxVe 29, 1956 Number 57 



THREE NEW SPECIES OF MIMETIDAE 
(ARANEAE) FROM PANAMA 

By Arthur M. Chickering 

Allnon College, Albion, Michigan 

In 1947 I published a brief paper attempting to bring onr 
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. 



NO. 57 



Genus GelAXOE 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 vievirs, 
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 
centers. 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. 

Maxillae. Convergent ; moderately slender ; almost contiguoas 
distal to lip ; palp inserted into basal third. 

Lip. Clearly united to sternum with no sternal suture visible ; 
only slightly longer than wide near middle. 

Sternum. Scutiform in 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. 1243. 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 measuieineuts iu millimeters) 





Fiuura 


Patellae 


Tibiae 


Metatarsi 


Tarsi 


Totals 


1. 


3.770 


1.235 


3.510 


3.640 


2.015 


14.170 


2_ 


3.510 


1.100 


2.600 


2.510 


1.430 


11.150 


3. 


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 ; patella ; 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 tubereulate; posterior surface with a pair 
of chitinized areas separated by a shallow slit; ventral surface 



1956 ne\v species op mimetidae 5 

with a pair of moderately deep depressions separated hy a septum 
(Fig. 1). 

Color in alcohol. Le<>s : 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 Mtmetus Hentz, 1832 
Mimetus saetosus sp. nov. 

(Figs. 2-3) 

Female holotype. Total length 8.71 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 slightly from PLE to opposite 
interval betAveen 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 wdth 
each lateral row with five, the posterior one on each side being 
offset laterally. 

Eyes. Eight in two rows; heterogeneous; LE on prominent 
tubercles ; AME at base of a tubercle 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 
length 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 jiosition 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. 

Maxillae. 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. Scutiform ; 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. 1243. 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 



1. 


2.860 


.780 


2.730 


2.275 


1.250 


9.895 


2. 


2.340 


.650 


1.950 


1.820 


1.105 


7.865 


3. 


1.300 


.455 


1.040 


.855 


.840 


4.490 


4. 


1.820 


.528 


1.690 


1.235 


.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 
foi- 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 0-lp-O-O, prolateral 
0-1-0-0, retrolateral 0-1-0, ventral 0-lp-O-lp-O ; 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 
0-lr-O-l-O, prolateral 0-1-0-0, retrolateral 0-1-0 together with sev- 
eral other slender spines irregularly placed, ventral 0-lp-O-l-O 
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. 

Ahdomen. 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 BREVIOBA NO. 57 

spinnerets and with a fairly well developed chitinous lip. Spin- 
nerets as usual in the genus. 

Epigynum. With a moderately well ehitinized 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 lirownish spots. 

Type locality. Female holotype from Barro Colorado Island, 
C. Z., July 18, 1954. No paratypes. 

MiMETUS VARIEGATUS Sp. UOV. 

(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. Pvatio of eyes AME : ALE : PME : 



1956 



NEW SPECIES OP MIMETIDAE 



PLE = 5.5 : 4 : 4.5 : 4.5. Height of clypeus eqnal to about 
14/11 of the diameter of AMB. 




External Anatomy of Mimetus 
Figs. 4-5. Mimetus variegatus sp. nov. ; tibia and tarsus of male palp; 
two views. 



C'helicerae. 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 

Maxillae. Slender, somewhat convergent; reach considerably 
beyond lip ; with palp inserted into base ; with serrnla only at 
onter distal corner. 

Lip. About nine tenths as long as wide ; with a well developed 
sternal suture; reaches well beyond middle of maxillae. 

Sternum. Broad, scutif orm ; 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 measuieinents in millimeters) 
Femora Patellae Tibiae .Metatarsi Tarsi Totals 



1. 


2.990 


.(5S2 


3.055 


3.510 


1.365 


11.602 


2 


2.405 


.520 


2.112 


2.275 


1.000 


8.312 


3. 


1.625 


.300 


.975 


1.034 


.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 0-1-1-1-1- 
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 11 

of first femur, retrolateral apparently with only one near distal 
end, ventral ; 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 w^here 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 lp-lp-0; metatarsus 
dorsal 0-1-0-0, 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; til)ia and patella short; tarsus 
moderately complicated ; with structural features best shown in 
Figures 4-5. Base of cymbium with a lol)ule which suggests that 
of M. verecundus Chickering but is, nevertheless, quite distinct 
from that species. 

Ahdomen. Of moderate height; about 9/11 as wide as long 
at level of "shoulder humps"; 1.43 mm. long; widest at level of 
"shoulder humps" w'hich 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. Each 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 slightly 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 cons^Dicuous than in 
the male. Otherwise essentially as in that sex. 

Maxillae, 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 OE MIMETIDAE 13 

(All measurements in millimeters) 



Femora 


Patellae 


Tibiae 


Metatarsi 


Tarsi 


Totals 


2.860 


.725 


2.925 


2.730 


1.300 


10.540 


2.405 


.645 


2.145 


1.885 


1.040 


8.120 


1.495 


.390 


1.105 


1.040 


.715 


4.745 


2.080 


.455 


1.495 


1.300 


.780 


6.110 



Spines. Small spiues between the robust spines on the first 
and second legs are mueh 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 l(weak)-l; tibia dorsal 0-1-1-1-0, 
prolateral with 0-1-1-1-1-1-1-1-0 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 ; metatarsus dorsal a row of 12-15 long slender spinules, 
hardly spines, prolateral 1-1-1-1-1-0, retrolateral 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 ; 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 except 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-lp- 
0-lp-O; 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 31. 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 except 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. 

BIBLIOGEAPHY 

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. Arachnida-Araneida. Vols. III. In: Biologia Centrali-Ameri- 
cana. Dulau & Co., London. 



E V I O R A 

Miaseiiim of Cooiparative Zoology 

Cambridge, Mass. June 29, 1956 Number 58 



SEQUENCE OF PASSERINE 
FAMILIES (AVES) 

By E. Mayr and 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 
encl depending on the particular sequence adopted by the 
i-espective 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. Juuge, Chairman 

Eijksmuseum Nat. Hist., Leiden 
Prof. J. Berlioz 

Museum d'Hist. Nat., Paris 
IMof. 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 



2 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 svibmit 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 ' Checklist would be willing to adopt whatever sequence 
the committee would vote for. 

PvEPORT OF THE EDITOPvS OP PETEPtS' 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 because 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 basic 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 sho^vn 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, Pycnonotidae, Sylviidae. Timaliidae, Turdidae, Muscicapi- 



1956 SEQUENCE OK PASSERINE FAMILIES (aVES) 3 

dae, etc.), (2) New World Insect-eaters and finches, (3) crows, 
birds of paradise, and associated families. All the more peculiar 
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 (1!>04), Sharpens 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 
(1952). 

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, Sclater used it in 1930, and Stresemann 
again in the Handbuch der Zoologie (1934). 

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 ])arenthetically 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 
impossil)le. The system of songbirds abounds in artificial aggre- 
gates 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 
aggregate. 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 Pycnonotidae 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 
Estrildidae 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, Aveighty 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 oeeurred auiono^ 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 famili(»s 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, ."]. 

ACTIONS OF THE COMMITTEE 

The editors of Peters' C-hecklist requested that the members 
of the Committee express their preference between the sequences 
] , 2, 3 and 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 se(juence 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 
guide the editors of Peters' Checklist, may be stated. 

(A) There are a number of natural groups among the Oscines 
such as the thrush-flycatcher group, the "New World finches," 
and others delineated in Mayr and Amadou, 1951. In a linear 
listing unrelated groups of families 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 PachycephaJinae next to the Paridae, the 
Meliphagidae next to the Emberizinae, etc. 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 Hyposifta, 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 listed 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- 
cates that the larks may not be closely related to &jvy 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. GralUnidae and Artamidae. Should these families be trans- 
ferred from a place near the Laniidae to the neighborhood of the 
Callaeidae ? 

Answer: Yes. It is ad"^'isable for tAvo 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 rocinn and that thev 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. Ptilogonafidae and Dididae. Should not these two groups 
be retained as families? 

Answer: They are better placed as subfamilies for the reasons 
stated above under B. 

5. Pityriasis. Does the inclusion of this genus in the Prionopi- 
dae reflect true relationship? 

Anstver: 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 traditionalh^ 

6. Esirildidae. 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, Sylviidae, Muscicapidae, Fringillidae, Ploceidae, 
Emherizidae. 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 Emherizidae. 



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 Osoine Families 

Alaudidae 

Hirundinidae 

Motacillidae 

Campephagidae 

Pycnonotidae 

Irenidae 

Laniidae 

Prionopidae 

Vangidae 

Borabycillidae 

Bombycillinae 

Ptilogonatinae 

Duliivae 
Cinclidae 
Troglodytidae 
Mimidae 
Prunellidae 
Muscicapidae 

Turdinae (incl. Zeledauia) 

Tiinaliinae (incl. Chamaea) 

Paradoxornithinae 

Polioptilinae (incl. Rliamphocaenus and Microbates) 

S.vlviinae (incl. Regnlus, Leptopoeeile, Lophobasileus) 

Malurinae 

Muscicapinae 

Monarchinae 

Paehycephalinae • 

Paridae 
Sittidae 

Sittinae 

Hyposittinae 

Neosittinae 
Certhiidae 
Dicaeidae 



1956 SEQUENCE OF PASSERINE FAMILIES (AVES) 9 

Neetariuiidae 
Zosteropidae 
Melipha^dae 
Emberizidae 

Eniberizinae 

C'ardinalinae = Kichniondeninac 

Tanagrinao = Thraupinac 

Tersininae 

Coerebinae 
Parulidae 
Drepaniidae 

Vireonidae (im-l. Viieolaiiiiis + Oyclarhis) 
Icteridae 
Fringillidae 

Fringillinae 

Carduelinae 
Estrildidae 
Ploceidae 
Sturnidae 
Oriolidae 
Dicruridae 
Callaeidae 
Grallinidae 
Artamidae 
Craeticidae 
Ptilinorliynchidao 
Paradisaeidae 
Corvidae 

LITERATURE CITED 

A.O.U. (Amkkicax Ornithologists' Union) 

1931. Check-list of North American birds. 4th ed., 5L'(3 pp.. Lancaster, 
Pa. 

liERLIOZ, J. 

1950. In Grasse (ed.), Traite de Zoologie, vol. lo, Oiseaux, 1164 pp., 
Paris (Systematics pp. 845-1055). 

I'.ISWAS, 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. X. 

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. 

Gadow, H. 

1893. In H. G. Bronn's Klassen und Ordnungen des Thierreichs etc. 

Vol. 6, 4 Abth., II, Systematiseher, 301 pp. (pp. 279, 301). 
1898. A classification of Vertebrata, etc. 82 pp., London. 

Garbod, a. H. 

1876-1877. On some anatomical peculiarities which bear upon the major 
divisions of the passerine birds. Part 1, Proc. Zool. Soc. 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. 

Mathevs^s, E. M. 

1927-1930. Systema avimn 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 cerebralisation chez les oiseaux. Alauda, 14, pp. 
1-20; 15, pp. 1-15 (pt. 2), i.e., pp. 161-171 (pt. 3). 

li.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, E. B. 

1885. Catalogue of birds of the British Museum. 10, p. 1. 



1956 SEQUENCE OF PASSERINE FAMILIES (AVES) 1.1 

Sharpk, R. B. 

1899-1909. A luuul-list of the genera niul species of birds. 5 vols., London. 

Shufeldt, R. W. 

1904. An arrangement of families and the higher groups of birds. 
Amer. Nat., 38. pp. 833-857. 

Steiner, H. 

1955. Das Brutverhalten der Prachtfiiiken, Spermestidae, als Au.s 
druck ihres selbstandigen Familicncharaeters. Act. XI Cong. 
Int. Orn. (Basel), pp. 350-3.55. 

8TEJNEGER, L. H. 

1885. Birds. In Standard Natural History (J. S. Kingsley ed.). 
Boston. 

Stresemann, E. 

1934. Aves, Ilandlnieh der Zoologie. Kiikenthal and Krumbaeli ed., 7 
(2), 898 pp., Berlin and Leipzig. 

SUNDEVALL, C. J. 

1872-1873. Methodi naturalis avium disponendarum tentamen. 2 pts., 4°, 
Stockholm. 

SUSCHKIX, P. P. 

1925. The Evening Grosbeak (Hesperiphona), the only American 
genus of a Palaearctic group. Auk, 42. pp. 256-261. 

TORDOPF, H. 

1954. A systematic study of the avian family Fringillidae based on 
structure of the skull. Misc. Publ. Mus. Zool., Univ. Michigan, 
no. 81, 42 pp. 

Wallace, A. E. 

1874. On the arrangement of the families constituting the order 
Passeres. Ibis, pp. 406-416. 

Wetmore, a. and W. W. Miller 

1926. The revised classification for the fourth edition of the A.O.U. 
check list. Auk, 43. pp. 337-346. 

1951. A revised classification for birds of the world. Smithsonian 
Misc. Coll., 117, no. 4, pp. 1-22. 



E 



Museiiim of Comparative Zoology 



{'amukidge, Mass. September 12, 1956 Number 5!) 



A NEW SUBGENUS OF VHAMAELIW FROM 

RHODESIA AND NEW RACE OF MABVYA 

FROM KENYA COLONY 

By Arthui^ Loveridge 

Fift}- years ago when Boiilenger (1906, Ann. Mag. Nat. Hist., 
(7), 18, p. 346, fig.) described RhampJioleon marshalli from 
Mashonaland, he remarked on the only occurrence of this genus 
south of tlie Zambezi as being "of very great interest." In as- 
signing marshalli to Rliampholeon, 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, in whicli 
i-espect he found it agreed with other continental species of 
Ehampholeon, a group I have since suggested should be regarded 
as only a subgenus of Brookcsia. 

Dr. V. FitzSimons (1943, The Lizards of South Africa, p. 
172, pi. xxi, fig. 5) also treated marshalli as the only South Afri- 
can representative of Ehampholeon. 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 tlie 
Museum of Comparative Zoology and were examined by me when 
making a synopsis of the continental African Brookesia (sub- 
genus Ehampholeon). At that time (1951, Bull. Mus. Comp. 
Zoo]., 106, p. 182, footnote) I rejected marshalli as a Ehampho- 
leon, despite its bicuspid claws, referring it to Chamaeleo on 
account of its prehensile tail. 

Actually the bicuspid claws (present or absent in Eliam.pho- 
leon) constitute the only character in which marshalli dift'ers 
from the forms assigned to Chamaeleo ; thus it bividges the ga]) 



2 BRHVIOKA NO. 59 

between Chamaelco and the subgenus Bhamplwleon 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. 

BiCUSPIS subgenus new 

Type. Rhampholeon marshalli Boulenger; known onl}- from 
Southern Rhodesia. 

Diagnosis. General appearance and soft nasal protuberance 
resembling that of Chamaelco rather than Rhampholeon. Scales 
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, Avith an average 
of 1.86 times for our entire series of 31 marshalli. The largest 
of the series, a topotypic gravid 9 (M.C.Z. 44445), measures 
103 (68 + 35) mm. 

The only Brook esia (subgenus Rhampholeon) with a tail any- 
thing like as long proportionately as that of marshalli, is B. k. 
kersteni (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. rohecchii (Bou- 
lenger). 

Mabuya bayonh keniensis subsp. nov. 

When the Museum of Comparative Zoology received a pair of 
typical Mahuya hayonii Bocage from Chitau, Bihe District, 
Angola, in 1936, it was immediately apparent that its East 
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 Mahuya 
could be undertaken, or a decision reached as to whether hayonii 
itself should be treated as a race of gravenhorstii Dumeril and 
Bibron of Madagascar. As time for any such thorough investiga- 
tion is lacking, I propose the name Mahuya hayonii keniensis 
subsp. nov. 

Holotype. Museum of Comparative Zoology No. 29662, an 
adult $ , from the northern Uaso (Guaso) Nyiro, Sotik, Kenya 

1 As Angt'l (1942, Mem. Acad. Malgache, 36, pp. 154, 178) designated no type 
for his genus Erohiticauda, I suggest that Brookesia nams Boulenger be so 

rcgjinlfd. 



1956 LIZARDS FROM KENYA AXO RHODESIA 8 

Colony. Collected by the Smithsonian-African Expedition, 190!). 

Paratypes. A specimen from Mount Kenya (U.S.N.M. 40710), 
another from Wambiigu (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 hayonii by me (1923; 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 Mahuia hayoni (sic) by Tornier, 1896 
(Die Kreiehthiere 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. Ngurumani). 

For the loan of the paratype material in the United States 
National Museum, as well as an Angolan b. hayonii (U.S.N.M. 
26389) for comi)arative purposes. 1 am indebted to Dr. Doris 
M. Cochran. 

Diagnosis. Dorsals tricarinate, whereas in h. hayonii they are 
strongly quinquecarinate. Certain other characters may prove 
to have an average difference, but this cannot be demonstrated 
until more material of typical hayonii 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 slightly 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) supraocular (though usually not in con- 



4 HKKVIORA NO. 59 

tact with the first) ; frontoparietal single, larger than the inter- 
parietal, which separates the parietals completely ; nuchals 
multicarinate. 

Midbody scale-rows 36 (34-36), dorsals strongly tricarinate; 
preanals not or but slightly enlarged; scales on soles not or but 
slightly 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) fiecks (or ocelli) ; a 
(n-eam (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 o' (M.C.Z. 21)662), 148 (62 + 86) 
mm.; of paratype 9 (M.C.Z. 2!)663), 160 i- (82 + 78+) mm., 
tail-tip missing. 



E V I O R A 

Museianti of Comparative Zoology 

Cambridge, Mass. September 12, 195() Number 60 



A NEW SPECIES OF AGRIOGNATIIA FROM 
JAMAICA, B. W. I. 

By Arthfr M. Chickerixg 

III connection with my study of tlic genns TetragnatJia 
Latreille, 1804 from Panama I have also had the opportunity to 
study several species believed to belong to the rather poorly- 
known genus Agriognatha 0. 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 1958, Miss E. B. Bryant 
of the Museum of Comparative Zoology at Harvard College had 
l)een 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 and it is hoped that considerable time can be 
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 XO. 60 

The liolotype and allotype Avill be deposited in the collection of 
the Museum of Comparative Zoology at Harvard Collepre. 

Genus AgeIOGNATHA 0. V. Cambridge, 1806 

The genus Agriognailia 0. 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 
congeneric with A. bella (0. P. Cambridge) from Costa Rica. He 
considered it probable that Cyrtognatha Keyserling was the same 
as Agriognatha 0. 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 (0. 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 
glol)Osa 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 
described in this paper. 

Agriognatha beyantae 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. Eight in two rows as usual in the genus; LE extend 
slightly beyond border of head. Viewed from above, anterior 



1956 AGRIOONATPIA FROM JAMAICA 3 

row strongly i-pcurved, posterior row moderately recurved ; 
viewed from in front, anterior row moderately recurved, posterioi- 
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 nearlj^ 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 pi'ominent tubercle. Height of clypeus equal 
to diameter of AME. 

Chelicerae. Very divergent, nearly horizontal; in i)lace 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-bifurcate tooth near 
the base of the fang on the promargin together with three small 
teeth; the retromargin has four small teeth as shown in Figure ]. 

Maxillae. Essentially parallel but with distal third slightly 
curved outward and considerably widened; slightly 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 betw'een 
bases of fourth coxae which are separated by slightly more than 
one third of their width. 

Legs. 1243. \Yidth of first patella at "knee" .220 mm., tibial 
index of first leg 5. AYiclth of fourth patella at ''knee" .165 mm., 
tibial index of fourth leg 7. 



4 






BREVIORA 






NO. 60 




Femora 


Patellae 
(All measu 


Tibiae 
lements in 


Metatarsi 
millimeters) 


Tarsi 


Totals 


1. 


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 


7.475 


Palp 


.726 


.185 


.198 




.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 leg: : 
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 broAvn with 
many irregular darker areas especially on dorsal surfaces. 
Carapace : with a narrow lirown 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, 1.43 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 
14 of the diameter of ALE. Height of clypeus equal to slightly 
more than % of the diameter of AME. Other features essentially 
as in male. 

Chelicerae. Robust ; only slightly 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. 
4) ; some variation in placement and relative sizes of the teeth 
has been noted and must be expected among paratypes. 

Maxillae, Lip, and ^iernvm. Essentially as recorded for the 
male. 

Legs. 1243. 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) 



3.380 


.845 


3.140 


3.412 


1.365 


12.142 


2.600 


.780 


2.190 


2.632 


1.040 


9.242 


1.495 


.435 


.845 


.975 


.715 


4.465 


2.210 


.487 


1.495 


1.820 


.780 


6.792 



H BREVIORA NO. 60 

All legs with stout spines and hairs but the spines seem not to 
be specialized as in the male. The triehobothria 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. 1., June 27, 1954. Several paratypes 
of both sexes from the following localities: Blue Mountains, 
August, 1934 (Darlington) ; St. Andrews, Clydesdale, July, 1950 
(Bengry) ; Hardwar Gap, June, 1954'; Hanover, Askenish, 
trail to Dolphin's ITejid. .luue 24, 1954. 

BIBLIOGRAPHY 

Bryant, Elizabeth B. 

1940. Cuban spiders in Ihc Miiscuiii of ( 'oiuimiative Z()olo<;y. Bull. 

Mus. Comp. Zool., vol. 8(), pp. 2-19-531!, pis. 1-22. 
1945. The Argiopidae of Hispaiiiola. Bull. Mus. Ooni]). Zool., vol. 9."), 

no. 4, pp. 359-418, pis. 1-4. 

Cambridge, O. P. and F. P. Cambridge 

1889- Arachnida-Aiaueida. Vols. I and II in: Biologia Centrali- 
1905. Americana. Dulau & Co., Loudon. 

Keyserling, Gr.\f Eugex vox 

1881. Neue Spinneu aus .Xmerika. \'eiliaudl. zool. Lot. (ies. Wit^n, vol. 
31, p. 270. 

Petrunkevitch, Alexander 

1911. A synonjTiiic index-catalogue of spiders of North, Central, and 

South America, etc. Bull, .\iner. I\[us. Xat. Tlist., New York, 

vol. 29, pp. 1-809. 
1925. Arachnida from Panama. Conn. Acad. Aits and Sciences, vol. 

27, pp. 51-248, 157 figs. 

iSiuce 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 kimlly lonned these specimens for examina- 
tion. 



1956 



AGRIOGNATHA FROM JAxMAICA 



IvOEWEli, C. F. 

1942. Katalos tier Araneae. Vol. 1. Bremen. 

Simon, E. 

1897. On the spiders of the Iskmd of St. Vincent. Part III. Proc. Zool. 
Soc. London, pp. 8G0-890. 





A 




External Anatomy of Agriognatim bryantae sp. nov. 

Fig. 1. Male chelieerae, 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. 



E V I O R A 

Mnasemiim of Comparative Zoology 

Cambridge, Mass. September 14, 1956 Number 61 



OX KEGENERATKJN BY EARTHWORMS OF A 

SPECIES OF THE LUMBRICID GENUS 

DENDEOBAENA EISEN 1874. 

By G. E. Gates 

Regeneration, either cephalic or caudal, after natural or ex- 
perimental amputation, for any species of Dendrohaena has not 
been recorded hitherto, though several of the head regenerates 
attributed by Morgan to Eisenia foctida (cf. 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 tenuis 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. ruhida (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 sculp- 
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 



2 BREVIORA NO. 61 

all levels to 13/14 and equimery at least to 5/6 can be expected 
in D. ruhida when conditions are more favorable. 

The level at which the indeterminate monstrosity was devel- 
oped is close to, if not actually in, a region where regenerative 
capacity in E. foetida (cf. Gates, 1949-1950) is bipotential. 
Monstrosities such as the one under consideration often are pro- 
duced in morphogenetie regions of dual capacity. However, the 
data now available as to the results of posterior amputation in 
D. ruhida scarcely warrant anticipation of heteromorphic 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. 
137). 

Table I 
Head regeneration in Dendrohaena ruhida 



•-> 

11 


o 2 
■sSs 


! 


2 


» 1 


a; 




1 


2/3 




1 


2 


19 




o 


3/4 




1 


3 


19 


Metamerism still indistinct in regen- 
erated left halves of iv-vi. 


3 


5/6 




3 


4 


32 


Eegenerate 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 nonnal. 


6 


6/7 




4 


-3 


19 


Metamerism probably was develop- 
ing abnormally. 


' 


7/8 




- 


4 


37 


Metamerism not quite normal. Ee- 
generated ventral half of viii ab- 
normal. 


8 


7/8 




3 


3 


19 





1956 REGENERATION IN DENDROBAENA 



n 


— 1- G 

fi £.2 




1° 


a 

1 


Davs 

of 

regener 

tion 


9 


7/8 


3 + 


3 




19 ] 


10 


8/9 


4 


3 




19 1 


11 


8/9 


4 


4 




32 1 


12 


8/9 


4 


4 




32 !; 


13 


8/9 


4 


-3 




19 I 


14 


11/12 


5? 


. 4 




32 I 


15 


13/14 


•')? 


-r, 




32 I 



Proximal portion of regenerate too 
short for a normal segment. 

Metamerism apparently developing 
abnormally in regenerated riglit 
halves of ix-x. 

Regenerate segments ii-iv not quite 
nomial. 

Segment iv too large. Excised dor- 
sal half of ix had been regenerated. 

Metamerism apparently normal. 

Regenerate metamerism abnormal. 

Metamerism in regenerated left 
halves of xiv-xvi probably develop- 
ing abnormally. 
Ifi 13/14 o-O ' -3 3(1 Metamerism rather indistinct in re- 

generated portions of xiv-xv, ven- 
tral half of xiv and a small portion 
of XV. 

17 17/18 4-0 i -3 32 Regenerate terminating distally in 

rounded knob like a rudimentary 
prostomium but towards the ven- 
tral rather than the dorsal side. 

Except as indicated otherwise above, excisions appear to have been transverse 

and along intersegmental furrows. 

Stages. 1. Prostomium aJid buccal invagination as yet unrecognizable. 

2. Prostomium and mouth developed. 

3. Intersegmental furrows demarcate regenerate into segments. 

-3. Furrows still indistinct. 

4. Pigment present but obviously different from that of substrate. 

5. Setae and nephropores recognizable. 

(i. Pigment now like that of substrate, external stigmata of regen- 
eration, except for typical metamcric anonuilies, uni-ecogniz- 
able. 



NO. 61 



Posterior regeneration in D. rubida 



All 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 
(cf. Table II) 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 intersegmental furrow and 
again there are no other external indications of reorganization. 

A terminal portion of No. 1, presumablj^ comprising only the 
last segment, at time of preservation was being reorganized. 
Completion of the process apparently under way probabh^ would 
have resulted in development anteriorly of eight new setal fol- 
licles with setae and appearance of an intersegmental furrow 
demarcating 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. 

Homomorphic 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 DKNDROBAENA 

Table IT 
Kesults of posterior amputation in D. ruhida 



m2 



1 42/43 .' Teiniiiial sul)strate segment has lost nuuh 

pigment, setae and nephropores. 

2 54/55 ? An intersegmental furrow delimits small 

white an;il region from last substrate 
segment. 

3 65/66 12 Same as in No. 2. 

4 80/81 6 Anal region not delimited from last siilv 

stiate segment which still has setae and 
nephropores. 

5 85/86 7 Anal region not delLmited from last sub- 

strate segment where some of the setjie 
still are present. 

6 87/88 10 Anal region not delimited. 

Time allowed for regeneration, 19 days. 

Uiseussioii 

llomomorphic tail regeneration does not necessarily follow 
immediately after amputation and may not begin until nine 
months later. The process gets under waj' 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 foctida, in the author's earlier studies 
(Gates, 1049-1950). after r(Miioval of the posterioi- 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 B. ruhida 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. 

EEFERENOES 

Gates, G. E. 

1948. On segment formation in normal and regenerative growth of 
earthworms. Growth, 12: 166-180. 

1949-1950. Regeneration in an eartliworm, Eisenia fuetida (Savigny) 1826. 
I-III. Biol. Bull., 96:129-139; 98:36-45; 99:425-438. 

1953. On regenerative capacity of earthworms of the family Lumbii- 
cidae. Am. Midland Nat., 50: 414-419. 

1954. Anterior regeneration in a sexthecal species of lunibricid earth- 
worm. Breviora, 27:1-5. 



E V I O R A 

Muiseitami of Comparative Zoology 

Cambridge, Mass. October 3, 19o() Number 62 



A THIRD LEAF-N08ED 8PECIES OF THE LIZARD 
GENUS AXOLIS PROM SOUTH AMERICA 

By Jameh a. Peters 

J>i(i\vii T'liivcisity, Providence, Bliode IsL-iiul 
and 
(tustavo Orces-V. 

Escnehi J'olyteeiiica Xacioiinl, Qiiitd, Kcunaor 

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. 



l'"i<'lil wurk in Kcuailor d\ii-iiis' l!l-">4 liy tlic senior author was suppo 
rant from the PtMirose Fund of the Anieriean Pliilosoidiical Society. 



2 HREVIORA NO. 62 

Akolis proboscis, new species 

Type. Museum of Comparative Zoology Xo. 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 Piohincha Province, Ecuador, collected by An- 
tonio Proano, 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, flesh}" appendage, which is about as long as the head, on 
the snout. The only other species with a similar appendage is 
A7iolis pliyllorhinus 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 crest, as wtII 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 along 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 occupied 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 ANGLE FROM SOUTH AMERICA 3 

three or four scales in center of sni)raorbital 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 scales 
between canthus rostralis and upper labials anteriorly, four or 
five posteriorly, no distinguishal)le horizontal rows of loreal 
scales. Last scale in canthus rostralis followed by a single super- 
ciliary, wliich 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- 
eiliaries. Single row of three or four enlarged subocular scales 
from loreals to granular postocular region and between the 
granules 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 man}- 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 sul)- 
ocular enters labial row, followed by granular scales. First labial 
on both sides wedged between rostral and second labial, lowei- 
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 sul)labials on each side, anteriormost in contact with lower 
lal)ials, })osterior 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. 

Body very slightly compressed, vertebral angle comparatively 
sharp. Dorsal crest of raised denticulations runs down spine 
from head onto tail, Avith 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. Scales 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 fiat 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. 
Scales on throat quite granular except on moderately developed 
gnlar 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. 

Scales on all parts of limbs and feet subequal, pavimentose or 
slightly granular. Scales on dorsum of digits expanded, nmch 
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. Scales 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 i/. 
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 

Fuie Foot Hind Foot 

Left 5-11-16-17-10 4-11-19-29-18 

Right 5-11-17-17-10 4-12-14-30-18 

Scnle foinui'.ne for digits in liolotyiK". 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 phalan.x, which 
arises dorsally from the lamellar portion of the toe, are not included. 

Dorsal ground color (in alcohol) a dull gun-metal blue, with 



1 !);■)() l.EAF-NOSKD ANOLE FROM SOUTH AMEHK'A 5 

irregular, ])arallel, horizontal black streaks along vertebral mar- 
gin. These streaks vaguely line up and form a pair of lateral 
bands between the limbs. Dark brownish-black spot at shoulder. 
Jjimbs and lateral surfaces spotted with light yellowish-white ; 
limbs vaguely barred with darker blue ; skin in interstices be- 
tween scales of dorsum of digits light, giving appearance of 
banding on fingers and toes. Dorsum of head unicolor, as back ; 
temporal region and lips lighter, with faint reddish or purplish 
tinge and a marked light spot over the ear opening. Ventral 
surfaces generally lighter, M'ith 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 
described 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 Ecuador, 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 Blgr., 
eulaemus Blgr., festae Peracca, gracilipes Blgr., granuliceps 
Blgr., lotifrons Berthold, maculiventris Blgr., and peraccae 
l>lgr. ) have been previously recorded from Ecuadorian localities. 
While in the Choco area of Ecuador in 1954, the senior author 
collected four specimens of A. breviceps Blgr., three individuals 
of A. peraccae Blgr., 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 Ecuador. 
Seven additional species have been recorded in the literature as 
taken from Ecuadorian 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 Ecuador with no added informa- 
tion, and no additional specimens have been taken to establish 
a range for the species. 

There are sixteen species, including alhi Barbour, bitectus 
Cope, hreviceps Blgr., fasciatus Blgr. (with its synonyms elegans 
Blgr. and irregularis Werner), festae Peracca, fraseri Giinther 
(with its synonym devillei Blgr.), gracilipes Blgr., granuliceps 
Blgr., latifrons Berthold (with its synonym princeps Blgr.), 
lemniscatus Blgr., lemurmus Cope, macrolepis Blgr., maculi- 
ventris Blgr., notophoUs Blgr., palmeri Blgr., and peraccae 
Blgr., for which both males and females are known, and for 
which available specimens or descriptions provide many and 
obvious diiferences from A. proboscis. Females only are known of 
the species antonii Blgr. and ventrimacnlatus Blgr., but the 
descriptions given for the types shows that these species are 
quite different from A. proboscis. Only males of aequatorialis 
Werner, chloris Blgr., eulaemus Blgr., rosenbergi Blgr., 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 
'Shaughnessy, and lionotus Cope, but they are sufficiently well 
described to eliminate them as available names for A. proboscis. 

Myers and Carvalho (1945, p. 7) felt that the relationships of 
their new species were wdth 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 '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-NOSKD ANGLE FROM SOUTH AMERICA 7 

opportunity to compare data for tliis species witli tlie 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 slightly more 
prominent than that of A. transfasciatus and presumably also 
of punctatiis. In addition to the presence of keels on the ventrals, 
A. houlengeri 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 houlengeri 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 houlengeri. Two rows of 
scales, slightly 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 p^'o- 
boscis or phyllorhinus, and apparently not true of transfasciatus. 
It appears, then, that A. houlengeri is quite distinct 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, 1). 1(J5) described t<cytomycterus 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, w^hile 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 tlio Lizards in tlic British MiiscHUii. Vol. II:i-xiii, 
1-497, -24 pis. 



8 BREVIOKA SO. 62 

Cope, E. D. 

1876. Eeport on the reptiles brought by Professor James Orton from 
the Middle and Upper Amazon, and Western Peru. Jouni. 
Acad. Nat. Sci. Philadelphia, (2) 8:159-188. 

Myers, G. S. and A. L. de Carvalho 

1945. A strange new leaf-nose lizard of the genus Aiioli.'^ from Ania- 
zonia. Bol. Mus. Nac. Brasil, Zoo!., No. 43:1-14, 9 figs. 



Anoli.s proho.scis Holotype i (M. C. Z. 54800). 



E V I O R A 

Mmseiuim of Comparative Zoology 

Cambridge, Mass. October 11, 1956 Number 63 



NEW BATHYAL ISOPODA FROM THE CARIBBEAN 
WITH OBSERVATIONS ON THEIR NUTRITION 

By Robert J. Menzies 

Marine Zoologist 
Lamont Geological Observatory! 
(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 
Epibenthic Trawl (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- 
scrijit 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 

1 Lamont Geological Observatory, Contribution No. 219, Biology Program No. 7. 



2 HBEVIORA NO. 63 

the parapodial setae of some species of polyehaete 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 n. sp. 
Figure 1 
Synonyms. None. 

Diagnosis. Front concave. 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. 
Tyj^e. 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 Eurycope 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 bifurcating 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 1<^K0M THE CARIBBEAN 




Figure 1 
Figure 1. Acantkocupe .p.nosiss..na n. sp., bolotype female length 8 mm 
..,, toto; 5, maxilliped; C, first peraeopod ; B, first antenna; E, left manda 
ble ; F, seventh peraeopod. 



BREVIORA 



NO. 63 




Figure 2. Dendrotion hanseni, n. sp., holotype male, length 3.5 tiim. A. 
toto; B, first pleopod ; C, maxilliped; D, pleotelson ; E, gnathopod ; F, left 
mandible. 



1956 NEW ISOPODA FROM THE CARIBBEAN 5 

DeNDROTION HANSENI, 11. sp. 

Fifjure 2 
Synonyms. None. 

Diagnosis. Lateral ])r()eesses .spiuulate. 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 
liad been known only from Norway. 

Genus NeOANTHUKA. n. g. 

Synonyms. None. 

Diagnosis. Anthuridae with piercing and sucking mouth parts. 
Eyes 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. Neoanthura 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. 

Xeoantiiura coeca, n. sji. 

Figure -3 
Synonyms. None. 

Diagnosis. Mandibular palp triai'ticulate, lack setae. Apex of 



NO. 63 




Figure 3 
Figure 3. Neoanthura coeca, n.g., n. sp., holotype female. A-A', toto; B, 
uaxilliped ; C, first antenna ; ]), secon'l antenna ; E, uropod and telson. 



1956 



NEW ISOPODA FROM THE CARIBBEAN 




Figure 4 

Figure 4. An-tarcturm annaoidcs, n. sp., holotype male. 
B, penis; C, gnathopod; D, seventh peraeopod; E, inner 
uropod. 



A, lateral view; 
view of apex of 



S BREVIORA NO. 63 

pleotelsoii rounded with only three setae. Margins of uropodal 
i-arai 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 peraeonal somites. 

Measurements. Holotype female, length 3.75 mm.; width 0.25 
mm. 

Tijpe. 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 length 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 la Plata. It differs 
from that species in lacking a marginally serrated pleotelson 
and in lacking armed coxal plates. 

Iaxikella caribbic'a, n. 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 \yith 7 spines on dorsum, third with 9 ; fourth with 9 ; 
fifth to seventh with 2. Maxilliped with two coupling hooks. First 
antenna with 13 articles. 



1956 



NEW I80P0DA FROM THE CARIBBEAN 




Figure 5 
Figure 5. lanirella caribbica, n. sp. A, type toto; B, dactyl first peraeo- 
Ijod; C, maxilliped ; D, dactyl seventh peraeopod; E, first antenna. 



10 BREVIORA NO. 63 

Measurements. Holotyi)e 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' E. 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. 
17, 175 pp., 25 pis. 

Bonnier, J. 

1896. Edriophthalones, //( Res. Sci. Canipagne du Caudan dans le 
Golfe de Gaseogne. Ann. Univ. Lyon, vol. 26, pp. 527-689, pi. 33. 

Menzies, R. J. 

1956. New abyssal tropical Atlantic isopods with observations on their 
biology. In press. Am. Mus. Nat. History. 

Saks, G. O. 

1897- An account of the Crustacea of Norway. II, Isopoda. pp. 1-270. 
1899. 100 pis. Bergen. 



E V I O R A 

Museum of Compsirative Zoology 

Cambridge, Mass. October 11, 1956 Number 64 



RARE SPECIES OF COPEPODA, CALANOIDEA, 
TAKEN FROM THE IZU REGION 

By Otohito 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. S. 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. 




NO. 6-i 



Figure 1. Byooalanus 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; /, second antenna; g, first leg and proximal 
outer margin of endopodite; h, 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 Pseudocalamis 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 as in Pseudocalamis. Type species, Ryocalanus infelix n. sp. 

Ryocalanus INFELIX sp. nov. 

Description. Male : Length, 2.18 mm. ; eephalothorax, 1.93 mm. ; 
abdomen, 0.25 mm. ; head separate from first thoracic segment ; 
last two thoracic segments separate ; eephalothorax 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