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Comparative Zoology

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 1

NOTES ON THE AMERICAN SOFT-SHELL TURTLES
WITH SPECIAL REFERENCE TO AM YD A AGASSIZII

By Leoxhard Stejxeger

With Thirty Plates

CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
May, 1944

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Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 1

NOTES ON THE AMERICAN SOFT-SHELL TURTLES
WITH SPECIAL REFERENCE TO AMYDA AGASSIZII

By Leonhard Stejneger

With Thirty Plates

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

May, 1944

PREFATORY NOTE

BY

T. Barbour

As has been well known, for many, many years before his death
Doctor Stejneger was engaged in the preparation of a general treatise
on the fresh water and land Testudinata of North America. After his
death it was found that by far the greatest part of his notes were in the
fragmentary state that often holds in an early stage of preparation
for publication. The only section which might be said to be in pub-
lishable form and to reflect the mature conclusions of the author was
this part comprising his summary of the genus Amyda. For many and
obvious reasons it seems desirable to bring this to public attention and
by the same token it seems presumptuous to change it.

The Museum of Comparative Zoology has the good fortune to
present to the public what will perhaps be the last example of the work
of this distinguished veteran worker in our field. By the consent of the
authorities of the United States National Museum this institution has
permission to publish this work, a tribute of honor and respect to the
author, being published here only because the situation induced by
the state of war makes it inconvenient for the Government Printing
Office to handle the work at this time.

No. 1. — Notes on the American Soft-Shell Turtles ivith Special Reference

to Amy da agassizii1

By Leonhard Stejneger

INTRODUCTION

The number of living indigenous soft-shell turtles of the family
Trionychidae in North America is less than a half dozen, while more
than a half hundred fossil forms, the earliest dating as far back as the
Cretaceous, have been described by paleontologists from the same
region which embraces the habitable part of North America east of
the Rocky Mountains and northern Mexico.

The living American forms have been considered from time to time
by prominent herpetologists as belonging to several genera under
varying names, according to their views on the genotypes. Thus
Agassiz recognized three genera and so did Baur and Hay; Cope and
True reduced the generic subdivisions to two. Boulenger, Siebenrock,
and recent systematists have regarded them as congeneric; the name
has varied accordingly.

NOMENCLATURE

In the early stages of systematic herpetology, before the idea of the
"genotype" had become generally accepted, there was no fixed rule
which would designate the type of genera with more than one species.
A sort of "selection by the first reviser" was early practiced. Thus if
an author subsequent to the creator of a plurispecific genus, in sub-
dividing the latter, retained this name for a single species, applying
one or more new names to the rest of the species, his selection, as a
rule, was respected, and thus the idea of type selection by "elimina-
tion" became a more or less general practice. But because of the un-
willingness or ignorance of some workers and the varying and uncer-
tain application of this method, great confusion in the nomenclature of
genera resulted. This is what happened to Geoffroy's generic term
Trionyx (1809), which has been the cause of so much confusion. In
1816 Oken was the first one to subdivide the plurispecific genus, re-
serving as he did the name Trionyx for the single species T. granosa,
leaving the others under the name Amy da. In 1830 Wagler reported
Oken's monotypic selection of Trionyx granosa, but substituted As-
pidonectes for Amyda. The confusion was caused by Gray who in his

1 Published with the aid of a special gift from Mr. G. R. Agassiz.

6 bulletin: museum of comparative zoology

Synopsis Reptilium; 1831, erected the monotypic genus Emyda for
the same species. He did so in ignorance of Wagler's action and after
seeing Wagler's work called attention to it in the "Additions and Cor-
rections" in the same work, p. 78. Yet in the "Catalogue" of 1844 and
his later writings he still retained Emyda. Gray was followed by
Boulenger and many writers since 1889, in spite of Baur and Hay who
in 1898 and 1904, like Bonaparte in 18361 and 1857'2, upheld Wagler's
action.

Following these dissertations by Baur and Hay, the case of the
genotypes of the genera Amy da and Trionyx was discussed by me in
1905 (Science, new ser., vol. 25, Feb. 10, 1905, pp. 228-229), conse-
quently before the adoption (1908) of the present wording of Article
30 of the International Rules of Zoological Nomenclature. In 1905 the
argument was based on the process of elimination, which at that time
was considered the legitimate process. On that occasion, the type of
Trionyx was decided to have been selected in 1816 by Oken as first
revisor by segregating Trionyx granosus, supposed to be synonymous
with Geoffroy's T. coromandelicus, as the sole species in the restricted
genus.

However, the adoption in 1908 of the changed form of Article 30
apparently nullified previously accepted type selections unless they
were couched in the stringent language of the new version.

The new Article 30 was framed and phrased for the purpose of doing
away with the uncertainties of personal interpretation unavoidable in
the process of elimination. It was intended that the new rule should be
applied literally; hence the first article (I a) reads: "When in the origi-
nal publication of a genus, one of the species is definitely designated as
type, this species shall be accepted as type, regardless of any other con-
siderations." (Italics mine.) To emphasize this intention the last
paragraph of the article was inserted as follows: "The meaning of the
expression 'select the type' is to be rigidly construed. Mention of a
species as an illustration or example of a genus does not constitute a
selection of a type." (Italics mine.)

Applied to the case in hand the facts are these:

In the original publication of Trionyx by Geoffroy in 1809 (Ann.
Mus. Hist. Nat., Paris, 14, pp. 1-20, pis. 1-5) none of the many species
was definitely designated as type. Not in the body of the text proper,

1 Cheloniorum Tabula Analytica, p. 8: "22 AMYDA, Schweigger (Aspidonectes WAGL.
Trionyx GRAY. BELL. Gymnopus, DUM." "23. TRIONYX WAGL. {Emyda, GRAY.
BELL. Cryptopas DUM.)"

2 Contrib, Nat. Hist. United States, 1, p. 330: "Trionyx proper in contradistinction

to Aspidonectes"; see also pp. 394-395.

stejneger: American soft-shell turtles 7

but in the Explication des planches {explanation, of the plates) at the end
of the article, occurs, however, the following mention of the trionyx
of Egypt as an illustration and example of the genus, which was
omitted in the simultaneous reprint of the article in the Bulletin of the
Philomatic Society of Paris :

"Le trionyx d'Egypte, represente planche I, vue en dessous el de cote,
nous donnant une idee exacte du port et des caracteres generiques des
trionyx, nous sommes bornes, dans les planches suivantes, a faire
figurer les seules parties caracteristiques des autres especes, telles que
leurs carapaces en a, et leurs plastrons en B." l

Even if this kind of mention as an "illustration and example" may
be characterized as a "clear intention" it is certainly not a definite desig-
nation rigidly construed.

Consequently, Geoffroy having failed to definitely designate T.
aegyptiacus as the type of Trionyx, it was left to the first subsequent
author to select the type, and that was done by Fitzinger in 1843
(Syst. Rept., p. 30) who selected T. coromandelicus Geoffroy,2 "and
such designation is not subject to change."

Having now disposed of the type designation of Trionyx for a genus3
in the subfamily Cyclanorbinae, the question of the status and type of
Amyda arises.

Among the new species described by Geoffroy in the same paper of
1809, is the Trionyx javanicus (Ann., p. 15). After a brief diagnosis he
added the following svnonvm:

"Amyda javanica SCHNYEIGGER, dans un manuscrit communique
a l'Institut."

This publication of Schweigger's monotypic generic name Amyda
clearly establishes its availability for the species congeneric with his
Amyda javanica, the description of which by Geoffroy is regarded as
identical with Boddaert's Testudo cartilaginea of 1770.

Fitzinger's use (1835) of the name Amyda for a section ( = subgenus)
with Trionyx subplamis as type, and Agassiz's subsequent (1857) ap-
plication of the same name for another genus with Amyda mntica as
type are consequently both void. Dogania is the proper name for the
former; Euamyda is now available for A. mntica as a generic or sub-
generic name.

1 The Egyptian trionyx, represented on pi. 1, viewed from below and from the side, gives us an
exact idea of the aspect and the generic characters of the trionyxes; we limit ourselves in the
following plates to figure only the parts characteristic of the other species, such as their carapaces
in a and their plastrons in b.

2 Trionyx coromandelicus Geoffroy, 1809, is a synonym of Testudo granosa Schoepff, 1792

3 Synonyms: Emyda Gray, 1831 (not of Rafinesque, 1815); Lissemys Malcolm Smith, 1931.

8 bulletin: museum of comparative zoology

Genus Amyda1 Schweigger

1809. Amyda SCHWEIGGER in Geoffroy, Ann. Mus. Hist. Nat., Paris, 14,
p. 1 (monotype Amyda javanicus = T. cartilagineus).

1816. Amyda OKEN, Lehrb. Zool., 2, p. 348 (type designated by Stejneger,
1907, Trionyx euphraticus).

1830. Aspidonectes WAGLER, Nat. Syst. Amphib., p. 134 (type designated

1843 by Fitzinger, Trionyx aegyptiacus = T. triunguis).

1831. Trionyx GRAY, Synops. Rept., p. 45 (type T. ferox) (not of Oken,

1816; Wagner 1830).

1835. Gymnopus DUMERIL and BIBRON, Erpet. Gen., 2, p. 472 (substi-
tute name for Aspidonectes Wagler) (p. 484 Gymnopodus, laps,
calam.)

1835. Platypeltis FITZINGER, Ann. Wien Mus., 1, pp. 120, 127 (type desig-
nated by Fitzinger 1843, Platypeltis ferox.)

1835. Pelodiscus FITZINGER, Ann. Wien Mus., 1, pp. 120, 127 (type desig-
nated by Fitzinger 1843, P. sine?isis.)

1842. Aspedonectes HOLBROOK, North Amer. Herp. 2 Ed, 2, p. 18

(emendation).

1843. Potamochelys FITZINGER, Syst. Rept., p. 30 (monotype Aspidonectes

javanicus).

1844. Tyrse GRAY, Cat. Tort. Brit. Mus., p. 47 (type T. nilotica = T. triun-

guis).

1856. Tryonix SAGER, Peninsular Journ. Medic Coll. Sci., 3, no. 8, Feb. 1856,

p. 361 (emendation).

1857. Amyda AGASSIZ, Contr. Nat. Hist. United States, 1, p. 398 (mono-

type A. mutica) (not of Schweigger).

1864. Rafetus GRAY, Proc. Zool. Soc. London, May 1864, p. 81 (monotype
T. euphraticus).

1864. Aspilus GRAY, Proc. Zool. Soc. London, 1864 (p. 83) (type Aspilus
cariniferus).

1869. Callinia GRAY, Proc. Zool. Soc. London, 1869, p. 221 (type, T.
spiniferus).

1895. Platyrettis KIRSCH, Bull. U. S. Fish Comm. 1894, p. 333; typogr.
err.? for Platypeltis).

1900. Aspidonectus BEYER, Proc. Louisiana Soc. Nat., 1897-1899, p. 43
(emendation).

1 Name of uncertain origin, but apparently a variant of Emys, a river turtle. In the synonymy,
the numerous generic terms based on exclusively Old World species have been omitted. Refer-
ence to these may be found in Bulletin United States National Museum No. 58, 1907, p. 514.

stejneger: American soft-shell turtles 9

The generally accepted five species of the wider genus Amy da in
North America naturally fall into three groups.

1. The A. mutica group, by Agassiz considered a distinct genus and
by many accepted as a subgenus (for which the name Eliamyda
may be substituted as Amy da Agassiz is preoccupied).

2. The A. feroz group including the species A. ferox, spinifera and
emoryi.

3. The A. agassizii group containing only one species on this conti-
nent.

The external appearance of these turtles is so much alike that the
early naturalists had difficulty in diagnosing them properly, with the
result that their taxonomic history is full of misidentifications and mis-
conceptions, even to the extent that the first group, at least on one
occasion, has been suspected of being only the sexual form of one of the
species of the second group.

As in so many of the Old World soft-shelled turtles the essential
characters are recognizable only in the bony structure, so also in our
American species. The most important ones are found in the skull and
the plastral bones and they will form the main subject of the following
discussion.

Skull Characters

The relative proportions of the skulls, their component bones and
the size and shape of the various fossae and foramina vary enormously
with age, hence comparisons have to be made between specimens of
about the same size. The individual variability, which is considerable,
increases with age very often to such a degree, — for instance the
enormous expansion of the alveolar surfaces in the old specimens of
A. ferox, — as to make even group definition difficult. Measurements
of a large number of skulls have therefore to be made, and as skulls of
approximately equal size are rarely to be had, the measurements have
to be reduced to percentages of some standard dimension. As such I
have selected the basicranial length (posterior edge of occipital con-
dyle to tip of snout). The analysis of numerous measurements has
convinced me that skulls with a basicranial length between 40 and 70
mm. (average about 55 mm.) practically represent the normal propor-
tions of important cranial dimensions of our American trionychids.
The relative proportions of the various parts have at that size reached
a sufficient stability and the figures consequently are comparable inter
se. Unfortunately, series of skulls of soft-shelled turtles are not numer-

10 bulletin: museum of comparative zoology

ous in museums, especially when reduced to specimens with a basi-
cranial length between 40 and 70 millimeters, hence the tables pre-
sented below, based as they are exclusively on United States National
Museum material, total only 41 specimens; nevertheless, the figures
are believed to be fairly representative.

An inspection of table 1 will show that Amy da agassizii possesses
smaller internal choanae (ch, plate 1, fig. 2) than the other species, but
a slightly larger intermaxillary foramen (int. max. for., plate 1, fig.
2). Were it not for the latter fact the distance between the inter-
maxillary foramen and the choanae would have been greater than it
is, (viz. 12.5 against only 7.7 to 9.7) in the other species; in other
words, in A. agassizii the longitudinal diameter of the choanae equals
their distance from the intermaxillary foramen, while in the others it
is much greater, in A. ferox even averaging twice as much.

The table further shows the greater width of the alveolar surface of
A. agassizii at the intermediary age of these specimens, but older males
(over 70 mm. basicranial length) of A. ferox acquire an increasing
width of the alveolar surfaces far exceeding in proportion even that of
A. agassizii (plate 30). The width of the alveolar surface of the
mandible in A. agassizii is twice, or nearly twice, as wide as in the
others of the corresponding size.

On the underside of the skull A. agassizii, in addition to the small
size of the choanae and the greater width of the alveolar surface, is
characterized by the position of the suture between the palatines and
the basisphenoid relative to the posterior edge of the temporal fossa.
This is coincident with the different shape of the opening of the tem-
poral fossa, the posterior edge of which is much wider and nearly at a
right angle to the axis of the skull. If therefore a line is drawn across
the base of the skull at the level of this edge, the line passes nearly
through the palatine-basisphenoid suture, while in the other species it
crosses the basisphenoid at or slightly anterior to the middle (plates
1, fig. 1; 6, 30.

The skull of A. mutica is unique among the Amydas in the slender-
ness and delicacy of its bones, especially those of the snout, which is
exceedingly narrow and elongated. The distance from the tip to the
posterior rim of the orbit is much greater than from this point to the
posterior edge of the tympanic cavity, while in the other American
species it is much shorter. This difference is mainly caused by the
reduction in A. mutica of the temporal fossa, the diameter of which
is about six tenths of that of the orbit, while in the other species it
equals or averages even slightly greater than the orbit. The weakness

SI KJNEGER: AMERICAN SOFT-SHELL TURTLES

11

of the A. mutica skull is further emphasized by the narrowness of the
interorbital spaee and the practical absence of an alveolar surface on
the mandible.

The normal skulls of the remaining American species, A. ferox, A.
spinifera and A. emoryi are essentially alike and present no striking
differences from the normal Amyda skull. They agree with A. agassizii
in the greater massiveness of the bones and general proportions of the
parts. With A. mutica they agree in the longer choanae and the con-
sequent shorter distance of the latter from the intermaxillary foramen.

Table 1

Cranial measurements reduced to
basicranial length of skull

(G

c3
fcC

X

o

spinifera

s-
O

s

0)

ferox group
collectively

c3

'-3

<

<

<

<

<

<

Basicranial length in millimeters,

average

51.8

58.3

53.8

54.0

42.8

Range of b. 1. of specimens

44-70

45-70

46-69

44-69

40-48

Number of specimens measured

8

12

9

9

30

6

Tip of snout to orbit

28.6

24.9

26.0

25.2

25.4

28.71

Orbit, horizontal diameter

19.1

20.0

19.9

20.2

20.0

20.6

Orbit to tympanic cavity

31.7

36.0

27.1

31.5

31.5

20.6

Temporal fossa, longest diameter

19.3

23.7

19.3

20.7

21.2

12.6

Interorbital width

8.5

6.0

7.4

7.8

7.1

5.1

Maxillary alveolar surface, width

11.4

6.8

7.4

8.9

7.7

5.1

Internal choanae, length

12.7

15.4

16.2

15.4

15.7

14.7

Choanae to intermaxillarv foramen

12.7

7.7

9.1

8.9

8.6

9.8

Intermaxillary foramen, length

12.2

11.3

11.2

9.8

10.8

15.6

Mandibular symphysis, length

19.5

11.8

13.2

14.3

13.1

20.3

Mandibular alveolar surface, width

10.6

5.5

5.4

5.9

5.6

1 Four specimens only, due to the fact that the bones — intermaxillaries and maxillaries of
this species are so fragile or poorly ossified that they were lost or mutilated in the preparation
of the others. The omission in .4. mutica of a measurement of the alveolar width of the mandible
indicates that it is so slightly indicated that it almost may be said to be nonexisting.

12 bulletin: museum of comparative zoology

The great changes in shape, proportions, color pattern and structure
of the soft -shelled turtles according to sex, age, and individual variabil-
ity make it impracticable to construct a workable key to the species,
hence I have confined myself to specimens in their early maturity,
when the critical characters have assumed a relative and comparable
stability unaffected by the rapid changes of youth and the often exag-
gerated individual peculiarities of old age. As the specific differences
in the skull appear fully developed and most easily appreciated in
skulls between 40 and 70 mm. in basicranial length, I have selected
such specimens as norms to which other sizes may be more or less suc-
cessfully referred, pointing out the deviations, as shown in the available
material, under the headings of the various species. As the sexual
differences are comparatively slight they have been ignored in the key,
though it may be pointed out here that the adult males differ visibly
from the females in the tail extending considerably beyond the cara-
pace while in the females it about reaches the edge of the disk; in
possessing a smaller head and a greater expansion of the plastral cal-
losities.

It is finally to be emphasized that specimens may be met with
which depart so far from the norm that they defy positive identifica-
tion by the registered characters alone or in combination. Such cases
are not particularly rare among the Testudinata.

Key to young adult specimens of North American
soft-shelled turtles

a1 Neurals normally 8, separating all the pleurals; entoplastron
bent at an angle of about 100° or more; a central callosity nor-
mally on entoplastron; temporal fossa of skull small, the longest
diameter less than two thirds the long diameter of the orbit;
distance from orbit to tympanic cavity about equals diameter of
orbit; intermaxillaries (premaxillaries) narrowly touch or are
separated by the maxillaries. Nostrils rounded, septum between
them rather wide and without lateral projecting ridge; no tuber-
cles on the leathery disk of carapace or its anterior edge {mulica
group; subgenus Euamyda).

Amyda mutica, (p. 14)

a2 Neurals normally 7 (occasionally 8), last pair of pleurals in con-
tact; entoplastron bent at an angle of about 90° or less; no cal-
losity on entoplastron; temporal fossa on skull about equal to
or longer than long diameter or orbit; distance from orbit to

stejneger: American soft-shell turtles 13

tympanic cavity much greater than diameter of orbit; maxil-
laries in contact above intermaxillaries (premaxillaries). Nostrils
crescent-shaped, internal ridge projecting on each side from the
narrow septum between them; leathery disk of carapace with
tubercles at least on anterior edge.

b1 Length of inner bony choanae greater than their distance
from intermaxillary foramen; mandibular symphysis less or
equal to length of choanae {ferox group).
c1 wSculpture of bony carapace coarsely grained, usually with
numerous irregular, more or less continuous and anasto-
mosing longitudinal ridges. (Leathery carapace of young
with longitudinal rows of tubercles; coloration peculiar.

Amyda ferox, (p. 25)

c2 Sculpture of bony carapace finely grained ; (leathery disk
of carapace of young smooth ; coloration of upper surfaces
gray, more or less marked with small dusky ocellae, or
solid spots or lines; plastron uniform white).
d1 Tubercles on anterior edge of leathery carapace well
developed, normally triangular and pointed; distinct
tubercles covering anterior and posterior flaps.
e1 Bony carapace without raised bony knobs or
"warts"; (young with one marginal dusky line on
posterior part of leathery carapace).

Amyda spinifera, (p. 43)

e2 Adult with strong tubercles on the hind part of
carapace "supported there by prominent bony
warts of the bony plates" ; (in young margin of the
leathery carapace marked posteriorly by at least
two parallel dusky lines).

Amyda spinifera aspera, (p. 56)

d2 Tubercles on anterior edge of leathery carapace poorly

developed, short and bluntly rounded; tubercles on

both flaps, if present, quite minute, except in very old

specimens.

Amyda emoryi, (p. 65)

b2 Length of inner bony choanae equals their distance from in-
termaxillary foramen; mandibular symphysis longer than
length of choanae (agassizii group).

Amyda agassizii, (p. 72)

14 bulletin: museum of comparative zoology

The mutica group

The only species so far known in this group deviates in its characters
and their combination quantitatively more from any of the others in
the genus, so much in fact that it has been regarded by outstanding
zoologists such as Agassiz, Cope, True, and Baur as representing a
"good" genus. However, the differences are of a character that rather
suggest relationship with the ferox group than a separate phylogeny.
The difference in the number and relations of neurals does not seem to
have any genetic significance in the genus Amyda. The long drawn out
and slender snout with the occasional separation of the maxillaries at
the apex and the shortening of the temporal-tympanic region indicate
modifications due to some food specialization with which the absence
of the dermal ridge in the nostrils at the tip of the proboscis and the
thickening of the septum may be correlated.1 In fact, there is indica-
tion of the ridge inside, though not reaching the opening of the nostrils.
The extreme development of the plastral callosities is purely quan-
titative and is closely approached in old specimens of Amyda emoryi,
and a small callosity on the entoplastron is often observable in other
specimens of the genus. The more circular shape of the body outline
is rather a juvenile character with which the obtuse angle of the
epiplastral bones is correlated. In none of the characters does the A.
mutica show any approach to any other group in the genus, particu-
larly not to any of the Old World soft -shell turtles, so that there seems
to be no convenience in recognizing it as a separate genus.2

Amyda mutica3 (Lesueur)
Plates 2, 3, 4

1827. Trionyx muticus LESUEUR, Mem. Mus. Hist. Nat. Paris, 16, Dec.
1827, p. 263, pi. 7 (type-locality, Wabash River, New Harmony,
Indiana; type Paris Mus. No. 787; Lesueur, collector ).— LE CONTE,
Ann. Lye. Nat. Hist. New York, 3, 1830, p. 95.— GRAY, Syn. Rept.,

1 Holbrook, curiously enough, who especially called attention to the "two characters which
always exist", viz. the "total absence of spines or tubercles" and the "great difference of the
nostrils" in describing the latter speaks of them as "closely approximated" in both ferox (in
which he included spinifera) and mutica (North American Herpetology, Edit. 2, 2, p. 13 and
p. 19) while in reality thev "are widely apart" in the latter, as pointed out bv Agassiz (Contrib.
U. S. Nat. Hist., 1, p. 398).

'Should the statement made by Dr. Stockwell (Journ. Comp. Med. Surg., 9, 1888, p. 29)
be corroborated, viz., that "the marginal ossicles in A. mutica are rudimentary; in A. spinifer
altogether wanting," the question of the generic status of A. mutica might well be reopened.

3 Latin, muticus, docked, dehorned, with reference to the absence of spines on the anterior
edge of the leathery carapace.

stejxeger: American soft-shell turtles 15

pt. 1, 1831, p. 46; Cat. Tort. Brit. Mus., 1844, p. 50; Cat. Shields
Rept. Brit. Mus., p. 1, March 8, 1856, p. 69.— DUMERIL AND
BIBROX, Erp. Gen., 2, 1835, p. 482 (lapsus for Gymnopus).— HAR-
LAN, Med. Phys. Res., 1835, p. 159 (Ohio River and tributaries).—
WIED, Reise Nord-Amerika, 1, pt. 3, 1838, p. 140 (Pittsburgh, Pa.).
-HOLBROOK, North Amer. Herpet., 1 ed., 4, 1840, p. 17, pi. 2
(Mississippi and tributaries); 2 ed., 2, 1842, p. 19, pi. 2 (Mississippi
and tributary streams).— DE KAY, Zool. New York, Rept., 1842,
p. 7 (Ohio River).— TROOST, Seventh Geol. Rep. Tennessee, 1844,
p. 39 (Tennessee; milieus misprint). — DUMERIL, Cat. Meth.
Rept. Mus. Paris, pt. 1, 1851, p. 22 (types).— STRAUCH, Mem.
Acad. Sci. St, Petersbourg, ser. 7, 5, No. 7, 1862, p. 174; 8, No. 13,
1865, p. 125; 38, No. 2, 1890, p. 118.— HOY, Geology of Wisconsin,
1, 1883, (p. 423) (Wisconsin).— BOULENGER, Cat. Chel. Brit.
Mus., 1889, p. 260, fig. 68 (Mississippi, Ohio and Saint Lawrence). —
HAY, Indiana Geol. 17 Rep., 1892, p. 551; Batr. Rept. Indiana,
1893, p. 143 (Indiana: Delphi; Madison; Terre Haute. Illinois:
Mt. Carmel).— HURTER, Trans. Acad. Sci. St. Louis, 6, 1892, p.
259 (Missouri: Mississippi River near St. Louis).— SIEBENROCK,
Sitz. Ber. Akad. Wiss. Wien, Math.-Nat. KL, 111, 1902, p. 822, fig. 5
(plastron); Zool. Jahrb. Suppl., 10, pt. 3, 1909, p. 605; Ann. Naturh.
Hofmus. Wien, 27, 1913, p. 214, sep. p. 44 (plastron); Verh. Zool.
Bot. Ges. Wien, 73, Aug. 1923, p. 192.— DITMARS, Reptile Book,
1907, p. 78, pi. 27, low. fig. (St. Louis, Mo.).— OVER, South Dakota
Geol. Nat. Hist. Surv. Bull. 12, Oct. 1923, p. 18, pi. 7 (Missouri Riv.
and eastward, South Dakota).

Gymnopodus muticus DUMERIL, Arch. Mus. Hist. Nat. Paris, 7,
1855, p. 203.

Gymnopus muticus DUMERIL and BIBRON, Erpet, Gen. 9, 1854,
p. 236.— WIED, Nova Acta Leopold.-Carol., 32, pt. 1, 1865, p. 54
(Ft. Mackenzie, Missouri River, 6-8 miles below Cedar Isl., South
Dakota).

Amyda mutica AGASSIZ, Contr. Nat. Hist. United States, 1, 1857, p.
399; vol. 2, pi. 6, figs. 6-7 (Lake Erie and Ontario; Delphi, Ind.;
Burlington, Iowa; Osage River, Missouri; Alleghany Riv., Pa.). —
GRAY, Suppl. Cat. Shield Rept. Brit. Mus., p. 1, 1870, p. 95.—
COPE, Bull. U. S. Nat. Mus., No. 1, 1875, p. 41 (middle and northern
tributaries of Mississippi and the St. Lawrence Riv.). — JORDAN,
Man. Vert. North. United States, ed. 3, 1880, (p. 168); ed. 8, 1899,
p. 206 (Canada to Ohio River, and N. W.).— CRAGIN, Trans. Kan-
sas Acad. Sci., 7, 1881, p. 116 (Kansas: Manhattan; Blue and Kansas
Rivers).— SMITH, Rep. Geol. Surv. Ohio, 4, 1882, p. 668 (Ohio
River, Ohio).— TRUE, Bull. U. S. Nat. Mus., No. 24, 1883, p. 28
(Madison, Ind.; Mt. Carmel, 111.; St. Louis, Mo.; Ft. Smith, Ark.);
Fish Industr. United States, sect. 1, 1884, p. 152.— HOY, Geol.

16 bulletin: museum of comparative zoology

Wisconsin Survey 1873-1879, 1, 1883, p. 423 (tributaries to the Mis-
sissippi in Wisconsin). — DAVIS and RICE, Illinois State Lab. Nat.
Hist. Bull. No. 5, 1883, p. 52 (North of Ohio River); Bull. Chicago
Acad. Sci., 1, No. 3, 1883, p. 32 (Illinois).— BAUR, Zool. Anz., 10,
1887, p. 99 (descr. plastron) ; Amer. Natural., 22, 1888 (1122); Proc.
Amer. Philos. Soc, 31, 1893, (p. 220).— STOCKWELL, Journ.
Comp. Med. Surg., 9, No. 1, Jan. 1888, p. 29.— HIGBY, Trans.
Wisconsin Acad. Sci., 7, 1889, p. 159 (Wisconsin: western half of
state).— GARM AN, H., Bull. Illinois Lab. Nat. Hist., 3, 1892, p.
247 (throughout Illinois: Mackinaw Creek, Woodford Co.; Quincy;
Illinois Riv., Peoria; Wabash Riv., Mt. Carmel; Ohio Riv., Cairo). —
RHOADS, Proc. Acad. Nat. Sci. Philadelphia, 1895, p. 404 (Ten-
nessee).— GAGE, Trans. American Microsc. Soc, 17, 1895, (p. 185)
(embryology).— SMITH, Proc. Linn. Soc. New York, 1898-1899,
No. 11 (1899) p. 24 (not near New York City).^-McLAIN, Notes
Coll. Rept. Arkansas, 1899, p. 1 (Ft. Smith, Ark.).— ATKINSON,
Ann. Carnegie Mus. Pittsburgh, 1, 1901, p. 154 (Allegheny Co.,
Pa.).— PAULMIER, Bull. 51, New York State Mus., 1902, p. 392
(probably in northern New York State). — MORSE, Proc. Ohio
Acad. Sci., 4, pt. 3, Spec. Pap. No. 9, 1904, p. 138 ("not recorded for
Ohio")- — ?NASH, check list of Vertebrates of Ontario, Batrachians,
Reptiles, Mammals, 1906, p. 17 (Ontario, Lake Erie, rare).
SURFACE, Zool. Bull. Pennsylvania Dep. Agric, 6, nos. 4-5,
Sept. 1, 1908, p. 119 (Pennsylvania, liable to be found in Erie
County and tributaries of the Ohio). HURTER and STRECKER,
Trans. Acad. Sci. St. Louis, 18, no. 2, 1909, p. 21 (Arkansas:
Ft. Smith; Pink Bluff; Little Rock). — HURTER, Herpet. Mis-
souri, 1911, p. 249 (Mississippi, Osage, Gasconade and Meranac
Rivers).— CLARK and SOUTHALL, Rep. U. S. Comm. Fish.,
1919 (1920) App. No. 7, p. 15 (economics). — MULLER,
Amer. Midland Natural., 7, no. 6, Nov. 1921, p. 180 (Iowa:
Fairport; habits). — BLANCH ARD, Occas. Pap. Zool. Mus.
Univ. Michigan, No. 117, July 6, 1922, p. 18 (Trotter's Landing,
Benton Co., Tenn.).— WEED, Copeia, No. 116, March 15, 1923, p.
48 (Meredosia, Ills.).— PRATT, Man. Vert. United States, 1923, p.
249.— STRECKER, Baylor Univ. Bull., 27, No. 3, Sept. 1924, p. 47
(Little Rock, Ark.); Contr. Baylor Univ. Mus., No. 7, July 15, 1926,
p. 7 (Neches River, Henderson Co., Texas, probably). — MYERS,
Proc. Indiana Acad. Sci., 35, 1926, p. 292 (Indiana).— STRECKER
and FRIERSON, Contr. Baylor Univ. Mus., No. 5, May 15, 1926,
p. 10 (Caddo and De Soto Parishes, La.).— ORTENBURGER,
Proc. Oklahoma Acad. Sci.,' 6, pt. 1, 1926, p. 100 (McCurtain and
Pushmataha Cos., Oklahoma).— STRECKER and WILLIAMS,
Contr. Baylor Univ. Mus., No. 12, Dec. 27, 1927, p. 16 (Christoval,
Tom Green Co., Texas); No. 17, Oct. 20, 1928, p. 19 (Bowie Co.: "no

stejneger: American soft-shell turtles 17

doubt")-— BURT, Occas. Pap. Mus. Zool. Univ. Michigan, No. 189,
Dec. 12, 1927, p. 9 (Manhattan, Riley Co., Kansas) ; Amer. Midland
Natural., 16, No. 3, 1935, p. 321 (Barber and Reno Cos., Kansas).—
POPE and DICKINSON, Bull. Publ. Mus. Milwaukee, 8, No. 1,
April 3, 1928, p. 82, pi. 21, figs. 5-6 (Wisconsin: Mississippi River
counties; Crawford and Pepin Cos.). — JORDAN, Man. Vert.
Northeast, U. S. (13 ed.) 1929, p. 254 (Middle and Northern tribu-
taries, Miss, and St. Lawrence rivers.) — ORTENBURGER and
FREEMAN, Publ. Univ. Oklahoma Biol. Surv., 2, 1930, p. 188
(Oklahoma: Alfalfa and Comanche Cos.). — WALKER, Copeia,
1931, No. 1, p. 12 (Scioto and Brown Cos., Ohio).— BOYER and
HEINZE, Trans. Acad. Sci. St. Louis, 26, No. 4, Apr. 1, 1934, p. 199
(Missouri: Jefferson Co.: Meranac River). — RUST, Blatt. Aquar.
Terrarienk., 45, 1934, p. ; sep. p. 12.— TAYLOR, Univ. Kansas
Sci. Bull., 22, No. 11, Apr. 15, 1935, p. 218 (Arkansas: Duvall Bluff,
Prairie Co.; Lewisville, Lafayette Co.).— DELLINGER and
BLACK, Occas. Pap. Univ. Arkansas, No. 1, June 1938, p. 46 (Ar-
kansas: Garland, Jefferson, Lafayette, Prairie, Pulaski, Sebastian,
Franklin and Lawrence Cos.). — SOLA, Bull. New York Zool. Soc,
34, No. 5, Sept.-Oct., 1931, pp. 134, 142, 155, fig. 7 upper (western)
part of Pennsylvania along Lake Erie). — CAHN, Illinois Biol.
Monogr., 16, Nos. 1-2, Aug. 31, 1937, p. 176; pis. 24, 30 fig. a; map
19(Illinois).— CONANT, Amer. Midland Natural., 20, No. 1, July,
1938, p. 154, pi. 21, fig. 1 (left), fig. 2 (Ohio: Scioto, Muskingum and
Ohio Rivers; map).— PARKER, Rep. Reelfoot Lake Biol. Sta., 3,
Jan. 1939, p. 88 (Tennessee: Reelfoot Lake, nowhere abundant). —
WELTER and CARR, Copeia, 1939, No. 3, Sept. 9, p. 130 (Ken-
tucky, east.: Triplet Co.; Fox, Fleming Co.; rare). — LOGIER, Roy.
Ontario Mus. Zool. Hanb. No. 4, p. 57 (Ontario: Lake Erie, probably
misidentified).— GENTRY, Rep. Reelfoot Lake Biol. Sta., 5, Jan.
1941, p. 75 (Tennessee: Pickett Co.); Journ. Tennessee Acad. Sci.,
16, No. 3, 1941, p. 332 (Tennessee: Obey River, Pickett Co.).—
ANDERSON, Bull. Chicago Acad. Sci., 6, No. 11, 1942, p. 219
(Missouri: Jackson Co.: Fry's Lake).— PETERS, Copeia, 1942, No.
3, Oct. 8, p. 183 (Illinois: Cumberland Co.).
1864. Potamochdys ? microcephala GRAY, Proc. Zool. Soc. London, 1864
(p. 87) (type-locality, Sarawak, Borneo!!; type in British Museum).
Callinia microcephala GRAY, Proc. Zool. Soc. London, 1869 (p.
222); 1873, p. 62, fig. 11; Suppl. Cat. Shield Rept. Brit. Mus., pt. 1,
1870, p. 108; Hand-list Shield Rept. Brit. Mus., 1873, p. 83.

Types. In the Musee d'Histoire Naturelle at Paris, No. 787 is
a shell on the plastron of which is written: "Trionix mutica Lesueur.

Wabash River par moi aout 1S27", and on a paper label pasted

on the underside of the stand: "Tortue qui n'a point le bord desou

18 bulletin: museum of comparative zoology

disque spineaux & que j'ai designee sou le nom de mutica dans

precedente note accompagniee de figure que je vous ai fait passer |?]
C. A. Lesueur." It is designated as the type on the printed label. The
bony carapace is 106 mm. in length and 107 mm. in width. A pair of
fontanelles between first pair of pleurals ; 8 neurals separate the pleurals.

No. 788 is a cotype retained by Lesueur until 1844 when it was ac-
quired by the Museum. It is a larger, mounted specimen, the bony
carapace measuring 134 mm. in length, 147 mm. in width; fontanelles
obliterated; 8 neurals, but last pair of pleurals broadly in contact.
Underside of stand inscribed "Gymnopus muticus Lesueur. Wabash.
Acquis de Mr. Lesueur 1844" (Lesueur died December 12, 1846).

The so-called "Spineless" or "Brown Softshell Turtle", the smaller
of our American species, may be easily recognized by the characters
given in the key. In the adolescent and adult carapaces the absolute
absence of tubercles on the anterior edge of the leathery disk will
identify the A. mutica even when the head is missing or the proboscis
is so mutilated as to defy examination. At the stage when the very
young specimens of all the species are nearly circular in outline and the
tubercles on the anterior edge of carapace of those species normally
possessing them may in some individuals be so indistinct as to be
doubtful, the oval nostrils without the septal tubercle will positively
identify A. mutica.

The other differential characters keyed are not always to be relied
on because of individual variation. Thus while normally the number of
pleurals is 7 pairs, separated the entire length of the bony carapace by
a series of 8 neurals, there are many and significant exceptions. Thus,
as already mentioned, the mounted cotype of the species in the Paris
Museum has the last pair of pleurals broadly in contact. Similarly the
U.S.N.M. 102910 has the eighth pair of pleurals in contact for at least
half their length; No. 54734 has all the pleurals separated by the 8
neurals, but it has only 6 pleurals on the right side, against 7 on the
left; U.S.N.M. 92605, 95134 and 029261 have only 7 neurals and 7
pairs of pleurals and the last pair is broadly in contact behind the
neurals. While visiting the Museum of Comparative Zoology during
the early days of these investigations, I examined two specimens, both
unfortunately at that time without numbers and locality, one adoles-
cent with the seventh pair of pleurals in contact behind the neurals,
while the second, an adult skeleton with a bony carapace measuring
140 mm. in length and 150 mm. in width, was still more abnormal
having 8 pleurals on the right side, against 7 on the left, and with 9
neurals separating all the pleurals.

stejneger: American soft-shell turtles 19

The greater angular width of the entoplastron is quite characteristic
of this species, but it is sometimes questionable in application because
of difficulty of exact measurement. However, it is useful where other
characters are irregular or in case one has to identify a disassociated
plastron or a single bone. The great extension of the plastral callosities
in the males is also a character of value, though the callosities on old
male A. emoryi may reach similar proportions on the hyo-, hypo-, and
xiphi-plastra. A central callosity is normally present in A. mutica on
the entoplastron though exceptional in the other species. Small callosi-
ties on the epiplastra are not uncommon in A. midica, though rare in
the others.

The sutural meeting of the maxillaries on the upper side of the
snout above the premaxillary (intermaxillary) is one of the characters
of the trionychid skulls, and is probably a normal or at least original
condition in A. mutica, but as noted above the extreme tapering and
hence weakening of the snout in this species results in the frequent
loss of these parts in the preparation of the skulls. In my series of
A. mutica skulls there are only four perfect specimens and in one of
these, No. 102677, the maxillaries are plainly in contact on the upper
side, above the premaxillary, while in the others, Nos. 53521, 54733 and
029261, the maxillaries are separated by the premaxillary.

A negative character of the young A. mutica is the absence of
ocellated or solid black rounded spots on the carapace, and of a defined
angular figure on top of the snout at the base of the proboscis. In a gen-
eral way the coloration is less distinctive than in the other species. The
yellowish margin of the leathery carapace seems to be definitely wider.

The coloration and pattern in this as well as the other species of
the genus becomes gradually more obscure with age, and varies in-
dividually as well as locally according to environmental conditions.
It may therefore not be unwelcome if I include a few color descriptions
of living or freshly killed specimens which have come under my ob-
servation when comparison with Ridgway's "Nomenclature of Colors
for Naturalists" (1886) was possible.

On September 4, 1934, the National Museum received from C. R.
Rogers two live specimens taken in Medicine Lodge River, one mile
SW of Lake City, Barber Co., Kansas. Description was at once made:
U.S.N.M. No. 95185, young adult female (leathery disk about 200
mm.). Iris bright "buff", the ring nicked slightly in front and behind
by a small blackish spot. — General color above nearly uniform
"tawny" with very faint mottlings of lighter "tawny-ochraceous" and
darker "raw umber", especially on the posterior flap, on which a faint

20 bulletin: museum of comparative zoology

submarginal dusky line borders the pale margin which is lightly
suffused with "rufous"; top and sides of head like back with a sharply
defined band of "ochraceous buff" narrowly edged with dusky; the
band continues — though fainter — anteriorly through the eye on to
the canthus rostralis converging towards the base of the proboscis
without meeting that of the other side; from the side of the occiput
indication of the band — though more irregular — on the side of the
neck; underside with a fine network of red blood vessels shining
through imparting a pinkish tinge to the soft parts which fades
gradually through "lavender" and "pearl blue" into "china-blue" on
the palms, soles and digits merging on the underside of the webs into
"tawny ochraceous" exteriorly and more pinkish interiorly; throat and
chin like the soles; lower lips whitish; callosities pale "fawn-color"
tinged centrally with blueish.

The other specimen, No. 95186, is a much smaller male (leathery
carapace length about 135 mm.) (pi. 3). Iris, a pale yellowish ring, but
the black spots are somewhat larger than in the older specimen.
Colors are also essentially like the latter, but the "ochraceous buff"
postocular band has the edges even better defined and on the side of
the neck joins the pale color of the underside which is sharply set off
from that of the upper side and extends onto the upper lip; the band
is only indicated in front of the eye by a small elongated triangular
spot of pale "ochraceous".

About the same time the National Museum received two live speci-
mens from J. H. Hall, collected in Mississippi (Marion County, Colum-
bia), both females. No. 95133, the larger one (disk approximately 185
mm. long). Iris bright buff yellow with a black horizontal bar. General
color above "clay color" with irregular blotches of pale "raw sienna";
marginal dark ring on carapace broken, faint, "hair-brown"; upper
side of neck washed with "tawny ochraceous"; postocular stripe dull
"buff -yellow", edged with dusky ("hair-brown"); front legs above as
well as dorsal and lateral surface of neck sprinkled with small dusky
spots; hind feet pale "olive yellow" with slightly larger and darker
dots and marblings; webs verging on "clay color"; underside of plas-
tron "flesh color", which on the white ground of the legs changes into
"pale blue" and on the soles and front feet verges on "heliotrope
purple"; underside of webs pinkish towards the edge; claws horny
white; callosities (none on epiplastra) "vinaceous-cinnamon". No fork
figure on top of snout; no spots or definite dusky markings on soles.

The smaller specimen, No. 95134 (leathery disk approximately 155
mm.) essentially as the larger one.

stejneger: American soft-shell turtles

21

Table 2
Cranial measurements of mutica in millimeters

CD
M

03

0J

M

03

O

a

of
PE

O

I— 1

l-c

O

a

o3

O?

&

O

e*-

i

a

o3

M

+3

O

a
'3

oT

o
1— 1

O

o
"a;

aT

03
CO

a>

5

H

43

o

o

53
«

aT

CD

co

CO
0)

a
a

CD

H

co

C

CD

e

0J

ft

CO

o

CD
bfl

<M
iO

co

41.0
11.0

i— i

CO
CM
OS

cn

o

42.0
12

00

i— i

CO

43.0

co

<N

O

48.0
14.5

o

OS
(N

O

T— t

CO

>

Basicranial length

40.0
11.5

42.8

Tip of snout to orbit

12.3

Orbit, horizontal diameter

8.0

8.5

p

9.0

9.0

9.5

7.0

8.6

Orbit to tympanic cavity

8.0

8.0

8.5

9.0

10.0

9.0

8.8

Temporal fossa, longest di-

ameter

5.0

4.-r

5

6.0

5.0

6.0

5.0

5.3

Interorbital width

2.0

2.0

..0

2.0

2.0

3.0

2.0

2.1

Maxillary alveolar surface,

width

2.0

2.5

2.0

1.5

2.0

2.5

2.0

2.1

Internal choanae, length

6.0

7.0

6.5

6.0

6.0

6.5

7.0

6.7

Choanae to intermaxillary

foramen

4.0

4.0

4.0

4.0

4.5

5.0

4.0

4.2

Intermaxillary foramen,

length

6.0

6.5

6.5

8.0

6.5

6.7

Mandibular symphysis,

length

8.0

8.5

8.0

9.0

10.0

8.0

8.6

Mandibular alveolar sur-

face, width

The colors of a freshly killed male specimen (leathery disk 204 mm.
long) collected by Dr. C. E. Burt in Kansas, Reno County, 6 miles E.
of Turon (U.S.N.M. No. 95259) were as follows: Ground color "broc-
coli brown" mottled with numerous very irregular and more or less
anastomizing, ragged-edged "sepia" spots occupying as much space as

22 bulletin: museum of comparative zoology

the ground color; posterior edge of disk pale tinged with "burnt um-
ber"; no submarginal blackish line; upper soft parts same broccoli
brown with very small and faint irregular "sepia" dots; a very pale
"russet" band, raggedly edged with blackish, from canthus rostralis
through eye and over ear slanting on side of neck halfway down the
neck ; pale lines in front of eyes do not meet those on snout nor is there
trace of a connecting line forming fork or triangle at base of proboscis ;
underside whitish with a faint glaucous blue gray tinge on throat,
underside of neck and feet; no dusky markings on feet; callosities, in-
cluding the small central one on entoplastron, pale blueish "plumb-
eous".

Geographical Distribution

Mississippi River and tributaries; north to South Dakota and
Minnesota; east to western Pennsylvania; west to Kansas, Oklahoma
& Texas.

Recorded from northern localities on the Trinity, Brazos and
Colorado Rivers.

Agassiz (Contr. Nat. Hist. U. S., 1, p. 404, footnote) writes, "De-
Kay's Trionyx ocellatus is Amyda mutica," a statement which seems
to be erroneous. The reference to DeKay's Trionyx ocellatus appears to
be the following note in his Zoology of New York (pt. 3, 1842, p. 7)
under Trionyx ferox [DeKay= T. spiniferus]: "The description given
above [p. 6: "anterior margin in the adult with numerous pointed
tubercles, which may be faintly and distantly traced in the young"]
was taken several years since, from a specimen obtained in the Mohawk
River. . . . The specimen, as I then thought, varied so much from any
description of the ferox within my reach, that I considered it to be new,
and named it ocellatus." The description of the Mohawk River speci-
men is clearly that of an adult A. spin if era, and cannot be taken as the
record of A. mutica in the Mohawk River. DeKay's mention that he
suspected ferox and muticus to be identical probably caused Agassiz's
statement.

List of specimens in U. S. National Museum

4783 (2) <? a<

id., d" adol. ?

?

7646 d" adol.

Mo., St. Louis

Engelmann

7647 juv.

U tl

?

7655 juv.

Miss., Monticello

Helen Tennison

7659-60 (2)

111.

Kennicott

7746

Ark., Arkansas R. near

Ft. Smith

Lt. Whipple

stejneger: American soft-shell turtles

23

8337 9

ad.

Indiana, Madison

9615 c?

adol.

111., Mt. Carmel

Mrs. M. E. Turner

9646 d1

adol.

11 tt

Robert Ridgway

May

1878

9647 d" adol.

II tt

tt

(«

9650 <? adol.

it a

tt

a

9651 rf" adol.

t; it

tt

tt

9727 9

ad.

it ll

J. Schneck

n

11629 juv.

?

?

11630 9

adol.

?

?

12794 (2.

I juv.

<

?&

111., Mt. Carmel

J. Schneck

13549

?

?

14780 juv.

Mo., St. Louis

J. Hurter

Dec.

20, 1887

19626 9

ad.

?

?

19627 9

adol.

o

?

21418 9

?

?

22629 juv.

Tex., Sabine R., 5 mi. S.

Jordan & Gilbert

1884

Longview.

029261 ad

.

?

Dr. G. Baur

45735-8

Iowa, Fairport, Missis-

sippi R.

Snyder

June

2, 1916

52528 <? ad.

Kansas

Roy L. Moodie

52116-8

juv.

111., Olney

Robert Ridgway

53521 9

ad.

Iowa, Fairport

Bur. Fisheries

Apr.

24, 1911

54733 9

ad.

ii it

J. Snyder

54734 9

ad.

it tt

tt

Aug.

8, 1916

54742

" (Missis-

sippi R.)

a

May

8, 1916

55525

Mo., St. Louis

J. Hurter

Mar. 23, 1907

55526

it

a

May

14, 1913

55527

Ark., Jeff. Co.

tt

1899

55528

" Little Rock

tt

June

1, 1900

55600

Tex., McLennan Co.

J. Hurter

1896

59267 9

ad.

Mo., Alexandria

E. Stringham

June

5, 1916

59268-9

Minn., Homer

F. Schrader

Sept.

8, 1916

59276

111., betw. Warsaw &

Hamilton

E. Stringham

Aug.

23, 1916

59278 9

ad.

Mo., Alexandria

June

5, 1916

59281

Iowa, Keokuk

Aug.

16, 1916

59282

tt it

June

7, 1916

59283

it it

July

1, 1916

59284

it it

June

7, 1916

59982

"Central U.S."

O. P. Hay

60054-6

Iowa, Fairport

Bur. Fish.

60561

Mo. St. Louis

J. Hurter

24

bulletin: museum of comparative zoology

71547

Okla., 6 mi. E. Ingersoll

A. I. Ortenburger

92605 cf adol.

Miss., Greenville

S. F. Hildebrand

May 29, 1933

95133-4 9 adol. Miss., Columbia

J. H. Hall

Aug. 1934

95185 9 ad.

Kans., 1 mi. S.W. Lake

City

C. R. Rogers

Aug. 31, 1934

95186 d1 juv.

Kans., 1 mi. S.W. Lake

City

<(

it

95259 a*

Kans., 6 mi. E. Turon

C. E. Burt

May 25, 1934

95260 9 adol.

U U {{

it

it

95415 9 adol.

" 3 mi. S.E. Oxford

u

1934

100422 9 adol.

La., Rayville

it

Aug., 1935

100813 pull.

Kans., Wakeeney

O. P. Hay

102612-3

111., 5 mi. S. Savannah

P. Bartsch

July 29, 1907

102677 9 ad.

Tenn., Reelfoot Lake

W. M. Perrygo &

C. Lingebach

May 6, 1937

102910 9 ad.

it it a

W. M. Perrygo &

C. Lingebach

May 8, 1937

115939

[Miss.]

[B. C. Wailes]

The ferox Group

Although quite distinct, the three species included form a rather
close group chiefly characterized by the uniformity of their skulls.
Common for all three is the large opening of the inner choanae with the
concomitant shortness of the distance of the latter from the inter-
maxillary foramen, which distinguish them from the agassizii group,
while the normal proportion of the preorbital region sets them off from
the mutica group. This statement may seem strange in view of the
opinion of many early competent herpetologists who refer to the species
of the ferox group under two different generic terms, a situation caused
by the confusion of the identity of the specimens which served as basis
for the generic concept, as will be shown later on.

Externally the ferox group differs from the mutica group by the
crescentic shape of the nasal openings, a character shared by all the
other species of the genus, and by the presence of the dermal tubercles
on the carapace. From the agassizii group, however, there is no obvious
external character by which to distinguish them as a group.

Within the group, the species from which its name is taken (because
the oldest one known) is most easily identified in the adult stage from
the other American species by the coarseness of the sculpture of its
bony callosities (pis. 9, 10), and in the early juvenile stage by its
unique coloration (pi. 19). The greater extension of the plastral
flap anteriorly beyond the carapace is also a noteworthy feature.

STEJXEGER: AMERICAN SOFT-SHELL TURTLES 25

Amyda ferox1 (Schneider)

Plates 5-10

1783.— Tesludo ferox SCHNEIDER, Naturg. Schildkr., p. 330 (Savannah
River, Georgia; type in Brit. Mus.; Dr. A. Garden, collector) (based
on Pennant, Philos. Trans., 61, pt. 1, p. 266).— GMELIN, Syst.
Nat., 1, pt, 3, 1789, p. 1039.— SCHOEPFF, Naturg. Schildkr., pt.
5, 1795, p. 102; Hist. Testud., pt. 5, 1795, p. 88 (based on Pennant).-
SHAW, Gen. Zool., 3, 1802, p. 64, pi. 17.— LATREILLE, Hist. Nat.
Rept,, 1, 1801, p. 165 (based on Pennant).— DAUDIN, Hist. Nat.
Rept., 2, 1802, p. 69 (based on Pennant).
Trionyx ferox SCHWEIGGER, Konigsberg. Arch. Naturw. Math., 1,
1812, pt. 3, p. 285 (Carolina and Florida) ; pt. 4, p. 363; Prodr. Mon.
Chelon., pt. 1, 1814, p. 15.— MERREM, Tent. Syst. Amph., 1820,
p. 20. — SAY, Journ. Acad. Nat. Sci. Philadelphia, ser. 1, 4, pt. 2,
1825, p. 218 (part: Carolina; Georgia). — HARLAN, Journ. Acad.
Nat. Sci. Philadelphia, 6, Feb. 1827, p. 32 (part: many of the rivers
of the southern states, not observed to exist further south [north ?]
than South Carolina) ; Medic. Phys. Res., 1835, p. 158.— Le CONTE,
Ann. Lye. Nat. Hist., New York, 3, 1830, p. 93 (part: Rivers of
Georgia and Florida, north to Savannah). — GRAY, Synops, Rept.,
1831, p. 45 (part); Cat. Tort. Brit. Mus., 1844, p. 49 (part); Cat.
Shield Rept. Brit. Mus., pt. 1, March 1856, p. 68 (part); HOL-
BROOK, North Amer. Herpet., ed. 1, 4, 1840, p. 9, pi. 1 (part:
Savannah and rivers emptying into northern border of Gulf of
Mexico); ed. 2, 2, 1842, p. 11, pi. 1 (part); in White's Statistics of
Georgia, 1849, Fauna and Flora, p. 13 (Georgia).— STRAUCH,
Mem. Acad. Sci. St. Petersbourg, ser. 7, 5, no. 7, 1862, p. 173
(part); 8, no. 15, 1865 p. 122. — BOULENGER, Cat. Chel. Brit.
Mus., 1889, p. 259 (Georgia, Louisiana).— DITMARS, Rept. Book,
1907, p. 74, pi. 26, lower fig. (Georgia, Florida, Louisiana). — SIE-
BENROCK, Sitzungsbr. Akad. Wiss. Wien, Math. Nat. CI., 91, pt.
1, Oct. 1902, p. 829; Zool. Jahrb. Suppl., 10, pt. 3, 1909, p. 603
(Georgia, Florida west to Louisiana); Verh. Zool. Bot. Ges. Wien,
73, Aug. 1923, p. 181. — BRIMLEY, Proc. Biol. Soc. Washington,
23, 1910, p. 18 (Georgia: MimsviUe; Florida: Orlando; Belleaire;
Green Cove Springs; St. Petersburg).— DECKERT, Copeia, No.
54, Feb. 17, 1918, p. 31 (Jacksonville, Duval Co., Fla.).— FLOWER,
Proc. Zool. Soc. London, ser. A, 1937, pt. 1, Apr. 15, pp. 16, 37
(age: 25 years + ). — POPE, Turtles of the United States and
Canada, 1939, p. 303, pi. figs. 94-97 (Southeastern Atlantic and
Gulf Coastal Plain).

1 Ferocious, the specific name evidently refers to the following sentence in Dr. Garden's
original description: "As the animal is very fierce, when it is attacked or disturbed, it often
raises itself on its legs, and will leap forward to bite its disturber or enemy, which it does with
great fury and violence".

26 bulletin: museum of comparative zoology

Amyda ferox OKEN, Lehrb. Zool., 2, 1816, p. 348.— STEJNEGER
and BARBOUR, Check List North Amer. Amph. Rept., ed. 1, 1917,
p. 124 (South Carolina to Florida and Louisiana); ed. 2, 1923, p. 140;
ed. 3, 1933, p. 153; ed. 4, 1939, p. 171.— SHUFELDT, Aquat. Life,
6, 1920, p. 27 (Georgia to Florida and Louisiana; habits). — LODING,
Alabama Mus. Nat. Hist., Paper No. 5, Sept. 1922, p. 47 (Fig
Island, Mobile Co., Ala.).— PRATT, Man. Vert. United States,
1923, p. 250.— WRIGHT, Ecology, 7, Jan. 1926, p. 84, pi. 5, fig. 3
(Okefinokee region, Ga.).— CORRINGTON, Copeia, 1927, No. 165,
Dec. 23, p. 101 (Pascagoula Swamp, near Biloxi, Harrison Co.,
Miss.).— PICKENS, Copeia, 1927, No. 165, Dec. 23, p. 43 (South
Carolina).— JORDAN, Man. Vert. Northeast U.S., (13 ed.) 1929, p.
255 S.C. to Fla. & La.— CONANT, Bull. Antiven. Inst. America, 4,
No. 3, Dec. 1930, p. 63 (Florida: Seminole; 15 m. E. of Sarasota; 18
m. S. of Ft. Myers).— HALTOM, Alabama Mus. Nat. Hist., Pap.
No. 11, 1931, p. 141 (pi. 39) (Alabama part, Mobile Co., Fig IsL). —
VAN HYNING, Copeia, 1933, No. 1, Apr. 5, p. 7 (Alachua Co.,
Fla., moderately common). — DeSOLA and ABRAMS, Copeia,
1933, No. 1, Apr. 3, p. 12 (Okefinokee Swamp, Ga.).— ALLEN,
Amer. Mus. Novit., No. 542, June 20, 1932, p. 20 (Mississippi: Han-
cock Co.).— RUST, Blatt. Aquar. Terrarienk., 45, 1934, sep. p. 12.

Platypeltis ferox FITZINGER, Syst. Rept., 1843, p. 30.— AGASSIZ,
Contr. Nat. Hist. United States, 1, 1857, p. 401 (southern states,
Georgia to western Louisiana). — GRAY, Proc. Zool. Soc. London,
1869 (p. 214); 1873 (p. 58) (part).— TRUE, Fish. Industr. United
States, sect. 1, 1884, p. 152.— BAUR, Proc. Amer. Philos. Soc, 31,
1893, p. 220— LONNBERG, Proc. U. S. Nat. Mus., 17, 1894, p.
317 (southern Florida).— WRIGHT and BISHOP, Proc. Acad. Nat.
Sci. Philadelphia, vol 67, 1915, p. 119, pi. 1, figs. 1, 2, 4; pi. 2, fig. 6
(Okefinokee Swamp, Georgia).

Aspidonectes ferox COPE, Bull. U. S. Nat. Mus., No. 1, 1875, p. 51
(Georgia to western Louisiana), — TRUE, in H. S. Thompson, South
Carolina, 1883, p. 238 (South Carolina); Bull. U. S. Nat. Mus., No.
24, 1883, p. 29 (part: Palatka, Putnam Co., Fla.; Charleston, S. C).
—DAVIS and RICE, Illinois State Lab. Nat. Hist. Bull. No. 5,
1883, p. 52 (Georgia to western Louisiana).— SHUFELT, Rep. U.S.
Nat. Mus. 1892 (1893) p. 32 (cast U.S.N.M. No. 8899).— BAUR,
Amer. Natural., 22, 1888 (p. 1121).— COKER, Bull. North Carolina
Geol. Surv., 14, 1906, p. 66 (North Carolina, introduced ?).— GOFF
and GOFF, Copeia, 1935, No. 3, Oct. 15, p. 156 (Griffin, Lake Co.,
Fla.; incubation).

1788.— Testudo mollis LACEPEDE, Hist. Nat. Quadr. Ovip. Serp., 1, Synops.
Meth. Quadr. Ovip., tab. between pp. 618 and 619 (based on Pen-
nant).— BONNATERRE, Tabl. Enc. Meth. Erpet., 1789, p. 25.

stejneger: American soft-shell turtles 27

1795.— Testudo (ferox ?) verrucosa SCHOEPFF, Naturg. Schildkr., pt. 5, p. 105
(based on Bartram's Trav. North and South Carolina, 1791, p. 177)
(type locality, eastern Florida); Hist. Testud., pt. 5, 1795, p. 90.

1801.— Testudo bartrami DAUDIN, Hist. Nat. Rept., 2, p. 74 (based on Bar-
tram, Voy. Amer. Septentr., vol. 1, p. 307).— HARPER, Amer. Mid-
land Natural., 23, No. 3, May 1940, p. 717 (Halfway Pond, Putnam
Co., Florida, "restricted type locality").
Trionyx bartrami GEOFFROY-ST. HILAIRE, Ann. Mus. Hist. Nat.
Paris, 14, Aug. 1809 (p. 18).— Le CONTE, Ann. Lye. Nat. Hist.
New York, 3, 1830, p. 96 (St. John's River, Fla.).

1809 1— Trionyx carinatus GEOFFROY-ST. HILAIRE, Nouv. Bull. Soc.
Philom. Paris, 1, No. 22, July 1809, p. 365 (type locality unknown;
type in Paris Mus.); Ann. Mus. Hist. Nat. Paris, 14, Aug. 1809, p.
14, pi. 4.

1809.— Trionyx georgianus GEOFFROY-ST. HILAIRE, Nouv. Bull. Soc.
Philomat. Paris, 1, No. 22, July 1809, p. 367 (substitute name for
T. ferox "Pennant").

1809.— Trionyx georgicus GEOFFROY-ST. HILAIRE, Ann. Mus. Hist. Nat-
Paris, 14, Aug. 1809, p. 17 (variant).

1812 ?.— Trionyx brongniarti SCHWEIGGER, Konigsberg. Arch. Naturw-
Math., 1, p. 288 (substitute name of Trionyx carinatus); Prodr. Mom
Chelon., 1814, p. 18.

1835. — Gymnopus spiniferus DUMERIL and BIBRON, Erpet. Gen., 2, p. 477
(part: Rivers of Georgia and Florida).

1835.— Trionyx harlani "Bell" HARLAN, Medic. Phys. Research, p. 159
(type locality, East Florida; type in "Mus. of Bell; Lond.").—
DeKAY, Zool. New York, Rept., 1842, p. 7 (East Florida).

Types. The type of Testudo ferox is still extant in the British
Museum.1 The species was discovered, described and illustrated by an
American; his descriptions and illustrations were published by one of
the most outstanding English zoologists of the period in the foremost
scientific journal of Europe; Linnaeus himself was simultaneously
notified of its discovery. Yet, for twelve years it remained without a
systematic name, until a German bestowed on it the name by which it
is specifically known. This unusual nomenclatorial history and the
inaccessibility of the original description to most zoologists justify its
reproduction here with a brief account of the related circumstances.

Dr. Alexander Garden, of Charleston, who was an ardent student of

1 According to Shaw (Gen. Zool., 3, pt. 1, 1802), the specimen was already then in the British
Museum. In 1831 Gray (Syn. Rept., p. 46) confirms that statement and adds that Pennant
gave the type specimen to the Royal Society. In 1844 and 1856 he records it as the "Specimen
described and figured by Pennant, (restuffed)."

28 bulletin: museum of comparative zoology

the natural history of the Carolinas, became possessed of an adult
specimen of the first softshell turtle reported from America. On
December 24, 1770, he wrote to his friend Mr. John Ellis in London
as follows:1

"I have one or two things which I think will please him [Thomas Pen-
nant] ; he shall have them by one of our spring ships ; one of them is a
species of Turtle, as yet nondescript. It is amazing how Catesby
omitted this. It is found in abundance in the Savannah river, in
Altamaha, and East Florida. It is a fresh water animal, grows to a
great size, and is as delicate as the Green Turtle, having a large
leathery cover over its back, and a head very like a Mole. I intend to
send a copy of my account of this animal to Mr. Pennant for his
American Zoology, and if I can get a drawing of it copied, I will send
him that. If I can obtain another Turtle, I shall send you one stuffed.
It has a relation to the first species of Linnaeus's last edition of the
Systema Naturae."2

Early next spring he kept his promise and sent Pennant the descrip-
tion as well as a preserved specimen. Another, better specimen, he
forwarded at the same time to Mr. Ellis with a drawing made from
life. This specimen with the drawing and the letter Pennant laid be-
fore the Royal Society in London on May 2, 1771, in the Philosophical
Transactions of which, vol. 61, they were published. Dr. Garden does
not state exactly whence the specimens came. In his letter he says that
"they are not commonly got here in Charles-town, though by chance
this last summer, I had two sent me", but as he mentions Savannah
and Altamaha rivers, it may be inferred that the two turtles were sent
him from one of those rivers, or one from each of them. One of the
specimens, "the specimen described and figured by Pennant (re-
stuffed)", according to Gray is the type in British Museum.

Dr. Garden's original description follows :

[p. 268] "They are found in large quantities in Savannah and Alta-
maha rivers; and I have been told that they are very common in the
rivers in East Florida.

They grow to very large sizes, though the largest that ever I heard
of was seventy pounds.

The Turtle, which I now have by me, weighs twenty pounds; and
probably, when I first got it, it might have weighed from twenty-five
to thirty pounds, as I have observed that it has grown poorer every

1 Smith, J. E. : A Selection of the Correspondence of Linnaeus and other Naturalists. London,
1821, 1, p. 580.

2 Linnaeus, Systema Naturae, ed. 12, 1, 1766, p. 350 (Testudo coriar.en\.

stejneger: American soft-shell turtles 29

day. I have had it now near three months, and I never could observe
that it has eaten any thing that has been given it, though a variety of
things have been tried.

It is twenty inches long from one end of the shell or covering to the
other, and fourteen inches and a half bread. The colour of this shell
or covering, in general, is dark brown, with a greenish cast.

The middle part is hard, strong, and bony; but all round the sides,
especially towards the tail and hindermost part, it is cartilaginous,
soft and pliable, resembling thick tanned sole-leather, yielding very
easily to any force in any direction whatever, but thick enough and
strong enough to defend the animal from any injury. All the hind
part of the back is full of oblong smooth knobs; and the fore part,
just where it covers the head and neck, is studded full of large knobs.
The under side of this plate is very [p. 269] beautiful, of a lively whitish
colour, interspersed with innumerable very fine ramifications of blood
vessels, running from the margin of the plate into larger and larger
branches, until the sight of them is at once lost by their entering the
body of the animal.

The under, or belly plate, or rather sternum, is of a fair whitish
colour, and extended forward two or three inches more than the back
plate, so that the head rests on it very conveniently. The hind part
of this plate is hard and bony, shaped very much like a man's riding
saddle, with two pieces for the thighs to rest on. The fore part of the
plate is pliable and cartilaginous.

The head is somewhat triangular and attenuated, rather apparently
small for the animal, but growing gradually larger towards the neck,
which is thick and long, and easily extended out (neck of the present
subject was thirteen inches and a half long) to a great length, or drawn
back again under the shell or plate.

The eyes are placed in the fore and upper part of the head, near to
one another, having pretty large loose palpebrae. The pupil is small
and lively, surrounded by a lemon-coloured iris, perfectly round, and
giving much life and fire to the eyes. When danger approaches, or
when it goes to sleep, it covers its eyes, by bringing the inner and
loose part of the lower palpebrae over its eye, like a membrana nictitans.

The upper lip and under lip are both large, but especially the upper.
The mandibula are both entire, each being one entire bone all round,
of the same shape as the mouth.

[p. 270] The nostrils are the most singular part, being a cartilagi-
nous production of at least three quarters of an inch, beyond the upper
and fore angle or point of the upper lip, perforated with two apertures

30 bulletin: museum of comparative zoology

reaching back and opening into the roof of its mouth, having a smooth
septum but fimbriated upon each side. This, at first sight, in some
manner resembles the snout of the mole; but it is tender, thin and
transparent, and cannot be intended for digging in the earth or land.

The arms are thick and strong, consisting of three distinct joints,
viz. the upper, the fore arm, and hand. The hands have each five
fingers, of which the three first are shorter and stronger, and furnished
with strong nails, or rather claws. The two last fingers have more
joints, but are smaller, and, instead of being furnished with claws, are
covered with the membrane, which is extended even beyond their ex-
tremities. Towards the back or hind part, there are two spurious
fingers, which just serve to support the membrane when extended.
The upper side of these arms and hands are covered with a wrinkled
loose skin, of a dusky greenish colour. The legs consist of the same
number of joints, and have the same number of toes as there are fingers
on the fore-feet, and these are furnished with nails in the same manner,
only there is but one spurious toe. Both the fore and hind legs are
thick, strong, and muscular; and as the animal is very fierce, when it is
attacked or disturbed, it often raises itself on its legs, and will leap
forward to bite its disturber or enemy, which it does with great fury
and violence.

[p. 271] They are likewise very strong, and of a lively whitish
colour, because they are generally, if not always, covered with the
upper plate, which, as I said before, is extended a great way behind.

The tail is large and thick, and generally as long as the hind part of
the upper plate. The anus is placed about an inch from the extremity
of the tail on the inside.

The turtle, from which these characters were taken, was a female;
after she came into my possession she laid fifteen eggs, and about the
same number were taken out of the belly when she died. The eggs were
nearly an inch diameter, and perfectly spherical.

It is esteemed very good eating, and said by many to be more
delicate than the green turtle."

On June 20, 1771, Dr. Garden wrote a letter in Latin to Linnaeus
which is translated as follows :l

"I have described and have lately sent to our friends Ellis and

Pennant, a new and very rare species of river Tcstudo, known here

by the name of the Softshelled Turtle, because the covering of its

back, especially towards the sides, is of a softish, leathery, very

1 Smith, op. cit. p. 336.

stejneger: American soft-shell turtles 31

flexible substance. This animal is found in the larger fresh-water
rivers of East Florida, Georgia and South Carolina."
Dr. Garden probably was hoping that Linnaeus might have hastened
to supply the still missing nomcn triviale as he did five years earlier
with Garden's no less startling discovery of Siren lacertina.

It will be noted that Dr. Garden himself, in the letter to Linnaeus,
identified the new discovery with the softshell occurring in East
Florida, and, subsequently, naturalists applied the name ferox indis-
criminately to all specimens from North America. Therefore, when
Lesueur in 1827 described T. spinifcrus as a distinct species and his
great countrymen Cuvier and Dumeril declared it to be only the young
of T.ferox, the identity of the two names was generally accepted, even
by Holbrook (1842) and Gray (1856). * Not until 1857 when Agassiz
demonstrated the distinctness of spinifcrus did it become generally
recognized, although even Boulenger's treatment (1889) shows that
the true characters of T. ferox, as represented by the type specimen,
were not completely understood.

The type specimen in British Museum was examined by Dr. Georg
Baur on September 7, 1888, as closely as the circumstances then per-
mitted. He noted particularly its "sehr rauhe Ornament" (very rough
sculpture), and "Keine Spines, sondern Tuberkel" (no spines, but
tubercles) evidently as compared with spinifcrus; "Schadel vom Typus
des Exempl. von Lucas. Vorderer Theil stark beschadigt, stark vorn
abfallend. Unterkief. ganz von jenem Typus. Jugale mit minim, unt.
Fortsaz." (Skull of the type of Lucas' specimen. Front part much
injured, greatly inclining anteriorly. Lower jaw entirely of that type.
Jugal with minimal lower process). He sums up thus: "Resultat : Type
von Platy pcltis ferox Exemplar von Lucas."

Lucas' "exemplar" is U.S.N.M. 8899 (tintag read upside down 6688
by Baur in his notes) which Baur had been studying with F. A. Lucas,
then curator of the division of comparative anatomy in the National
Museum. No. 8899 is a fine disarticulated skeleton still in the Museum,
of approximately the same size as Dr. Garden's type, and was col-
lected by Professor S. F. Baird, April 1877, in the St. John's River,
Florida, probably not far from Jacksonville. A fine plaster cast
painted by Schindler from his color sketch of the living specimen is
on exhibition in the Museum. A photograph of it was published in
the report of the Museum for 1882, pi. 32.

1 For some unexplained reason, however, Dumeril and Bibron ignored the priority rights of
ferox and called the combination which included the South Carolina to Florida records Gymnopus
iferus. Possibly as a disguised protest against Geoffroy's inefficient substitution of Trionyx
for Schweigger's earlier Emyda.

32

bulletin: museum of comparative zoology

Thanks to the authorities of the British Museum I have been per-
mitted to examine the type and can confirm completely Dr. Baur's
result as demonstrated by the photographs side by side (pi. 5) of the
two specimens and the measurements in table 1.

Type of Trionyx harlani. In the Medical and Physical Researches
published by him in 1835 Dr. Richard Harlan includes on p. 159 the
description of a Trionyx as follows :
"Trionyx Harlani
Trionyx Harlani, Bell, Monogr. Test, pi.
Char. Body more ventricose, soft portions of the shell less exten-
sive than in the other species. In general appearance ap-
proaching more to the genus Emys.

Inhabits East Florida. Mus. of Bell, Lond."
I have not been able to locate any such plate in Bell's Monograph
of the Testudinata. Bell, in correspondence, may have indicated that
he had in his collection a specimen from East Florida which he intended
to figure in his unfinished Monograph under the above name, but there
is no such plate among the unedited ones published in 1872 under the
title: "Tortoises, Terrapins and Turtles", neither have I found any
record of what became of Bell's specimen. J. E. Gray in the introduc-
tion of his catalogue of the Tortoises in British Museum, notices that
"the specimens presented by .... Thomas Bell, Esq. [may be re-
garded] as the types of the species described in his various papers, and

in his very beautiful Monograph of the Testudinata [by] Dr.

Richard Harlan, and Messrs. Edward and Henry Doubleday, as the
types of the North-American species described by Say, Harlan, and
others," but he has no reference to any softshell from Florida, nor is
there in any of the later catalogs.

External characters. While the skull characters are only available
in dubious and critical cases, normal specimens of A. fcrox within the
group are not difficult to identify, the older ones by the coarseness of
the sculpture of the bony carapace; the very young ones by their
unique coloration.

The coarseness of the sculpture of the bony carapace of A. ferox is
not confined to the general network of vermiculations of the surface,
but it is commonly specialized into a series of more or less prominent
longitudinal welts. The difference in relative size and pattern of the
pits and ridges which constitute the character of the "sculpture" is
difficult to describe, but a comparison of the samples figured (pis. 7,
9, 10) explains it better than words.

The unique coloration of the hatchlings and the very young speci-

stejneger: American soft-shell turtles

33

mens constitutes the most obvious and characteristic feature of the
species, but unfortunately disappears with age. In all the other species
the young are of a more or less pale olive or tawny ground color, either
uniform or marked with dusky or blackish specks of varying shape,
but mostly round dots, which when larger assume the form of ocelli
with a lighter center, combined with a nearly uniform white plastron
(except for the pinkish tinge due to the fine blood vessels shining
through in the living specimens). The prevailing feature is the dark-
ness and the saturation of the pigmentation, the big dark blotches on
the carapace, and the dark slate gray underside, in combination with
the strong contrast of the light pattern on the head and the margin of
the carapace, detailed description of which will be given below.

The anterior flap of the leathery plastron is longer and less circular
than in the other species, often extending a considerable distance for-
ward beyond that of carapace, but it appears to vary individually and
is difficult of precise definition because of absence of suitably fixed
points from which to measure.

The shape of the outline of the disk, however, is somewhat different
in the }roung of A. ferox, in as much as it is less circular than in the
other species, as will appear from table 3 which presents the measure-
ments of five hatohlings of the same brood. Similar proportions are
shown by one collected by Dr. Francis Harper in the Okefinokee
Swamp (U.S.N.M. 84603), viz. length of leathery carapace 39.5 mm.
and width 32.5 mm. It is also well illustrated in the photograph of
No. 61087, pi. 17, fig. a.

Table 3

Amyda ferox, pullus

Florida: Polk Co., Auburndale

mm.

Length of leathery disk
Width of leathery disk mm.
Height of body

mm.

61083

38.0
33.5

12.5

61084

40.5
35.5
12.0

61085

40.0
34.0
12.5

61086

41.0
34.0
12.0

61087

45.0
37.0
14.0

Average

40.9
34.8
12.6

Size. Amyda ferox is apparently the largest of the North American
softshell turtles. Authentic measurements of large carapaces are few

34 bulletin: museum of comparative zoology

and those available are of dubious value, due to uncertainty as to
identification, condition of specimen when measured, and method of
measurement whether in a straight line or along the curvature of the
shell. The largest Agassiz "had ever seen or heard of" was one from
Natchez "which measured eighteen inches and a half [470 mm.] from
the front to the hind margin of the carapace" (Contrib. Nat. Hist.
U. S., vol. 1, p. 401). The largest specimen now in the National
Museum is an old skin, with the skull in (U.S.N.M. No. 38123 from
"Florida" which measures about 17 inches along the curvature (430
mm.). Its zygomatic width is 68 mm. The plaster cast of No. 8899 —
(in exhibition series) is 438 mm. long; over the curvature it measures
460 mm. (18 inches); zygomatic width 67 mm.

But larger specimens may exist, or have been living in Florida not
long ago. The National Museum has a series of 16 weathered skulls
picked up by Dr. E. A. Mearns near Kissimee about the beginning of
this century, 14 of them larger than that of the 18 inch specimen men-
tioned above (No. 8899). The basicranial length of this one is 92 mm.,
the corresponding dimension of the 14 Kissimee skulls range from 95
to 114 mm. (aver. 103) with the zygomatic width varjing between
65 and 80 mm. (aver. 73).

Abnormal alveolar surfaces

This series of 16 skulls of evidently very old specimens shows an
extraordinary development which deserves special attention.

In the normal skulls of the species in all our specimens with a basi-
cranial length below 90 mm. the lateral outline of the snout anterior
to the orbit tapers towards the end in a fairly straight line, and the
narrow maxillary alveolar surface follows almost parallel, as in the
other species of the ferox group, irrespective of sex and age.

However, in the series of 16 Mearns skulls with basicranial length
above 84 mm. we find two different styles of snout outline and alveolar
surface as recorded in table 4.

For the illustration of the extremes of the width of the alveolar sur-
face and the outline of the maxilla it is only necessary to refer to plate
6. While greatly reduced the figures convey the difference between the
two series as the figures are of the same relative size.

Evidently the abnormal development of the maxilla as represented
by plate 6, figs. 3 and 4 are due to old age since we do not find it in the
smaller and younger specimens. It at once recalls similar conditions
in some Chinese species of the genus, upon which Father Heude, in

stejneger: American soft-shell turtles

35

Table 4
Alveolar width and outline of snout in a series of very old A. ferox.

Snout outline straight

Snout outline strongly

convex

Greatest

Greatest

U.S.N.M.

alveolar

Basicranial

Basicranial

alveolar

U.S.N.M.

No.

width

length

length

width

No.

029459

7.5 mm.

114 mm.

108 mm.

21.5 mm.

029464

107 mm.

15.5 mm.

029458

106 mm.

14.0 mm.

029454

105 mm.

14.0 mm.

029470

105 mm.

16.0 mm.

029457

104 mm.

18.0 mm.

029463

104 mm.

11.5 mm.

029451

101 mm.

15.0 mm.

029450

101 mm.

11.5 mm.

029460

100 mm.

14.0 mm.

029455

029456

7.5 mm.

100 mm.

97 mm.

11.5 mm.

029453

95 mm.

9.5 mm.

029452

029475

6.8 mm.

92 mm.

84 mm.

12.0 mm.

029462

1880, 1 based the description of numerous new genera and species. I
need only refer to his pictures of Caelognathus novemcostatus, pi. 5;
Tortisternum novemcostatum, pi. G; Cinctisternum bicinctum, pi. 9; and
especially Ceramopelta latirostris, pi. 7. Notwithstanding the fact that
he did not obtain any young specimens with these characteristics
Heude regarded them as specific or generic differences. Boulenger in
cataloguing the softshell turtles in the British Museum (1889, p. 243)
was "unable to find a single young specimen with the molar-
like alveolar surfaces" but having "found in three species, viz. T.
triunguis (Africa), T. cartilagineus (E. Indies), and T. sinensis (China)

i Mem. Hist. Nat. Emp. Chinois, 1, pp. 1-38, pis. 1-9.

36 bulletin: museum of comparative zoology

examples of the two types, i.e. on the one hand sharp-edged, com-
paratively narrow jaws, and on the other hand broad crushing alveolar
surfaces nearly meeting on the median line in front of the choanae,

in specimens which, in other respects, are undistinguishable,

[he] arrived at the conclusion that we may be in presence of a case of
dimorphism caused by a difference of diet."

This idea of a "dimorphism" due to a difference of diet does not seem
convincing. That it is not the result of old age alone seems obvious
from the fact that so many old skulls of the same size and presumably
age do not show the abnormality. That it is not due to a change of
diet affecting the whole population seems also obvious since the Kissi-
mee series are all practically from the same locality. That the differ-
ence is not a local one is proved by a weathered skull almost identical
with Kissimee No. 029462 of the above table, picked up by Dr. Francis
Harper in the Okefinokee Swamp (U.S.N.M. No. 59727) which with
a basicranial length of 88 mm. has a maxillary alveolar width of 11
mm. The fact that in the series the largest skull is normal and the
smallest abnormal coupled with the other fact that all the largest
authentically sexed specimens in the collection are female and normal,
while among the preserved specimens no abnormal male has been
recorded, suggest that the difference may be due to sex. If so, does that
indicate an individual preference among the old males for a certain
kind of diet?

The normal skull. As repeatedly noted the skull of A. ferox is built
on the same plan as that of spinifera and cmoryi and differs but slightly
in the proportionate size and relation of the various bones, but the
individual variation is so great that in some cases it is even difficult
to decide to which species an isolated skull without locality record be-
longs. A. ferox, however, may generally be diagnosed as having the
narrowest interocular width, the shortest distance between choanae
and intermaxillary foramen, and, in the medium-sized skulls, the
narrowest maxillary alveolar surface. It also averages the greatest
length from orbit to tympanic cavity combined with the shortest and
weakest mandibular symphysis. The interorbital space is usually less
in width than one third of the longest diameter of the temporal fossa,
while in the others it is usually more than one third. The anterior
processes of the prefrontals as a rule are longer and slenderer and the
angle projecting into the posterior border of the nasal fossa is conse-
quently more acute. In the older and younger specimens these differ-
ences become more obscure or may be entirely obliterated.

In a series of skulls of this group a rather common feature may be

stejneger: American soft-shell turtles 37

noted in the larger specimens of A. ferox. While in the palate of the
other species the maxillaries are in contact practically the whole length
between the choanae and the intermaxillary foramen, they may be en-
tirely or partly separated by the vomer which in many skulls may be
seen as a fork enclosing the posterior end of the foramen as shown in
all the figures on pi. 6 (very large specimens).

Coloration. Old specimens of this, as well as the other species, show
but little of the characteristic normal coloration and pattern of the
species. In A. ferox more or less faint remnants of the large brownish
blotches on the carapace may be made out on the generally dingy
"Isabella" colored ground, or disappear in the blackish or brownish
variations of the latter. But as previously indicated, the very young
ones display a distinctive and peculiar color and pattern, unknown
until figured by Ditmars (1907) and described by Wright and Funk-
houser (1915). Some of the specimens received by the National
Museum were either alive or freshly killed and their descriptions with
reference to a standard color nomenclature (Ridgway's Nomenclature
of Colors for Naturalists, 1886) were made:

U.S.N.M. No. 61087. Auburndale, Polk Co., Florida, collected by
N. R. Wood, summer of 1918. Pullus. Length of leathery carapace,
45 mm. Leathery plastron extending anteriorly beyond carapace 6 mm.
Iris pale silver gray with a horizontal black bar. — Upper surface of
carapace tawny olive (Ridgway, pi. iii, fig. 17) with dusky (dark
sepia) spots and a narrow well-defined outer edge of bright ochraceous
(R. v, 7) in strong contrast; underside of carapace and plastron slate-
gray (R. ii, 5) with scattered clay-colored spots on the former and the
anterior edge of plastron, the outer edge of disk narrowly ochraceous,
as above, though less bright; upper side of head, neck and legs olive
(R. iii, 9) with clay-colored marblings; from anterior angle of eyes to
middle of proboscis an inverted Y-shaped, pale clay-colored figure, the
fork situated halfway between eye and base of proboscis; side of head
olive with a wide well-defined angular band of yellowish buff extending
from posterior corner of eye to base of lower jaw; another similar
band, but slightly broken and brighter, almost cadmium-orange
(R. vi, 2) anteriorly, originating behind the former and descending on
the neck to past the middle; a third band, paler buff, curving around
the corner of the mouth almost meeting below the corresponding band
of the other side on the posterior third of the neck; a fourth, median
band between the last ones; soft skin of feet almost "plumbeous"
(R. ii, 15) underneath.

Four other specimens of the same brood, Nos. 61083-86, are colored

38 bulletin: museum of comparative zoology

essentially as the above. The numbers, sizes, shape and arrangement
of the dark blotches on the carapace vary to a great extent and so does
consequently the light network (really the ground color of the cara-
pace) separating them, it being slightly wider in the specimen figured
(pi. 19, fig. a). There is one feature common to all and of some signifi-
cance, viz. the more or less parallel arrangement of the outer blotches
on the posterior flap, those of the outer row nearly forming a continu-
ous dark line, the next row also coalescing on the posterior half, thus
clearly indicating the blackish submarginal rings of the other species,
as shown in the illustration just quoted.

Wright and Funkliouser (Proc. Acad. Nat. Sci. Philadelphia, vol. 67,
1915, p. 122) note that "as the specimens become older, the gayly
colored markings of the carapace become less distinct and have dis-
appeared on turtles which have attained a length of 6 inches [152 mm.]."
The National Museum has a specimen (No. 56804) from Irwin Co.,
Georgia, the carapace length of which is 100 mm., which still shows the
pattern as described above, but the tubercles on the disk are all well
developed.

Their further statement that with age "the plastron grows lighter in
color and the head uniformly darker with the markings obsolete", is
borne out by the following description made in July 1919 of a live male
specimen (now U.S.N.M. No. 62217) from Georgia, Berrien County,
with a leathery disk about 235 mm. long and 195 mm. wide: Iris
brownish gray with a brassy edge against the pupil. Carapace dark
"raw umber" (R. iii, fig. 14) with large dusky, more or less confluent
irregular marblings; top of head, neck and legs "sepia" (R. iii, fig. 3)
with paler marblings ; proboscis, snout, sides of head, and lips strongly
washed with "cinnamon" (R. iii, fig. 20); a dark brown mark across
the forehead in front of eyes continued behind them to the ear as a
series of spots, and a few spots of the same color behind corner of
mouth; on underside of neck back of the skull three longitudinal white,
dusky -edged marks; underside of body white; digits and webs strongly
washed with plumbeous (R. ii, fig. 15).

Some larger specimens may still retain traces of the original pattern.
A full grown male (U.S.N.M. No. 60496, carapace length 316 mm.)
collected at Auburndale, Florida, was received alive on February 28,
1918, and its colors at once described by me as follows: Iris dark silvery
gray with a darker horizontal bar and the inner edge forming a com-
plete bright narrow silvery ring sharply defining the pupil; inside of
nostrils dark pink; inside of mouth pale flesh-color. Ground color of
carapace bistre with anastomozing, ill-defined lines (10 to 15 mm. wide)

stejneger: American soft-shell turtles 39

of tawny olive, which isolate islands of the ground color of varying
sizes but averaging perhaps 25 mm. in diameter; neck and legs above
dark olive, head anteriorly suffused with cinnamon changing to pink
on proboscis; underside pinkish white, on neck suffused with dark
purplish gray on which dusky-edged cream-colored lines here and there
with touches of orange; palms and soles dark olive.

Aberrant Color Phase. A color anomaly shown by a specimen, the
history and description of which, because of the uniqueness of the pig-
mentation deserves to be recorded, is in the National Museum.

On April 11, 1881, there was entered in the Museum register of
reptiles a consignment of living specimens collected in March by James
Bell at Gainesville, Florida. Among them was an "Aspidonectes" of
which the artist Z. Schindler on April 9 had made a water-color sketch
for use in later painting the plaster cast for the "exhibition series".
The specimen, a female (Xo. 10545, carapace 225 mm. long, 16S mm.
wide), now a skin with skull separate, the color sketch, and the painted
plaster cast (slightly broken) are still in the Museum. The water-color
sketch and the oil-painted cast agree in all essentials, except that the
cast is (now) considerably darker. The skull and other structural
characters are those of a normal Amydaferox, but the color is an ex-
treme case of erythrochroism. The ground color of the carapace in
the water-color agrees closely with what Ridgway calls "mummy
brown" (R. iii, fig. 10), while in the cast it is more like his "chestnut"
(R. iv, fig. 9); the throat and front of neck are "rufous" (R. iv, fig. 7)
in both, though lighter in the former; top of head, back of neck, and
upper side of legs raw umber (R. iii, fig. 14) in the water-color sketch
while in the cast they are dark sepia washed with rufous; the most
conspicuous feature is the band from beneath the eye on the auricular
and temporal region which is bright vermilion in both paintings-
reminding one of the corresponding band in Pseudemys elegans) and
edged with a narrow dusky line anteriorly above and below. The un-
derside, judging from the color sketch (and the specimen) was uniform
white.

Geographical distribution

Florida north to South Carolina and west along the Gulf Coast to
Louisiana.

40

bulletin: museum of comparative zoology

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bulletin: museum of comparative zoology

List of specimens in U. S. National Museum

4373

Fla.

, Palatka

T. Glover

7651

n

ii

(i

8708

Ga.

Milledgeville

T. H. Bean

July, 1876

8899 9 ad.

it

St. Johns R.

S. F. Baird

Apr. 3, 1877

9670 d1 adol.

S. C

!., Charleston

C. C. Leslie

May, 1878

10545 9 adol.

Fla.

, Gainesville

J. Bell

Mar., 1881

10704

it

tt

it

19621 adol.

Ga.

Darien

?

20189 adol.

Fla.

, Eustis

T. Holm

March, 1893

26035 juv.

n

ponds near

Welaka

W. C. Kendall

Mar. 20, 1897

29210 9 ad.

tt

E. A. Mearns

029339 adol.

II

Eustis

H. J. Webber

029448-9

It

Kissimmee

E. A. Mearns

029450-1

029452-9 c? ad.

u

n

<(

029460-2 c? ad.

11

it

a

029463 tf1 ad.

It

a

a

029464-6 cf ad.

tt

tt

tt

029467-8

It

it

a

029470 cf ad.

It

a

a

029474 9 ad.

tt

it

tt

029475 9 ad.

tt

tt

a

029619 adol.

Ga.,

Mimsville

C. S. Brimley

38123 9 ad.

Fla.

M. L. Odell

38980-1 juv.

Ga.,

Mimsville

C. S. Brimley

June 19, 1909

51184 juv.

Fla.

ponds near

Tampa

Evermann & Kendall

Nov. 3, 1896

51417-20 hf-gr.

it

St. Petersburg

C. S. Brimley

51421 juv.

tt

Orlando

it

52476-83

Fla.,

Eureka

C. S. Brimley

Aug. 12, 1915

55316 c* ad.

Fla.

Vero

I. M. Weills

56804 juv.

Ga.

Irwin Co.

J. Hurter

Oct. 15, 1902

56805

Fla.,

Orlando

a

July, 1911

56806

it

Hillsboro Co.

tt

June 15, 1910

56807 ad.

tt

Bronson

tt

1891

59318

it

Sebastian

F. Harper

Feb. 23, 1917

59727-8 (skullk

only) <? ad.

Fla.,

Lake Okeechobee F. Harper

Feb. 6, 1917

60496 c? ad.

a

[Auburndale]

N. R. Wood

Feb., 1918

60532 adol.

a

Auburndale

a

<(

60533 ad.

a

(t

it

u

60534 9 ad.

a

a

a

u

stejneger: americais

r SOFT-SHELL TURTI

,ES <i«5

60547 & ad.

Fla., Auburndale

N. R. Wood

1918

60556 juv.

« a

■ a

Mar., 1918

60S28 eggs

" Homestead

C. A. M osier

60902-4

" Eureka

C. S. Brimley

1918

61031

" Gulf port

A. G. Reynolds

61083-7 pull.

" Auburndale

N. R. Wood

July 22, 1918

61096-109

ii it

u

Aug. 24, 1918

61352 9 ad.

" Lake Miakka

C. M. Barrett

June 18, 1918

62217 & adol.

Ga., Banks Mill Pond

U.S. Bur. Fish.

63343

Fla., Auburndale

N. R. Wood

1920

70398

Ga., Mimsville

C. R. Brimley

71068-9

?

?

71156 adol.

Fla., Plant City

C. R. Aschemeier

1926

71681 !

S.C., Greenwood

Dr. Barrett

73199

Fla., Delray

J. D. Thieme

80963

Fla.

F. H. Benjamin

Apr., 1930

84079

" 15 mi. from

.

Miami

M. K. Brady

1930

84080

" Orlando

E. T. Evans

ii

84603 pull.

Ga., Chesser's Id.,

Okefinokee Swamp

F. Harper

July 21, 1931

86492 d*

Fla., 15 mi. from

Miami

M. K. Brady

Mar., 1932

86828 <? adol.

" nr. Birdon

P. Bartsch

Aug., 1932

95767 <? adol.

" Lake Iamonia

C. R. Aschemeier

Feb. 16, 1935

103736 Juv.

" Silver Lake

ti

Mar. 7, 1938

Amyda spinifera (Lesueur)

Agassiz, in 1857, while demonstrating the distinctness of Lesueur's
species described 30 years before from the Wabash River against the
contention of contemporary authors that it was only the young of
Amyda ferox, at the same time established three additional species,
Aspidonectes as per from the state of Mississippi, A. nuchalis from
Tennessee, and A. emoryi from Texas. The latter has been generally
accepted, nuchalis has been generally ignored, and asper only recently
recognized as a distinguishable race of A. spinifera. As such it will be
treated below trinominally.

The characters attributed to A. nuchalis have lost their significance
after the accumulation of additional material of A. spinifera, and the
study of available series of specimens in connection with the present
investigation have not yielded data indicating a separable group popu-
lating the upper reaches of the Tennessee and Cumberland Rivers.

'Identity questioned.

44 bulletin: museum of compakative zoology

AMYDA SPINIFERA SPINIFERA1 (LeSUeUr)

Plates 11-15

1825. — Trionyx ferox SAY, Journ. Acad. Nat. Sci., ser. 1, 4, pt. 2, p. 218 (part:
Mississippi, Ohio, and Missouri Rivers; New York, etc.) (not of
Schneider).— LeCONTE, Ann. Lync. Nat. Hist., New York, 3, 1830,
p. 93 (part: all streams which run into the Mississippi). — HARLAN,
Med. Phys. Res., 1835, p. 158 (part).— SCHLEGEL, Fauna Japon.,
Rept., 1838, p. 30, pi. 5, fig. 5 (Cumberland, Tennessee and Ohio
Rivers).— HOLBROOK, North Amer. Herpet., 1 ed., 4, 1840, p. 9
(part); 2 ed., 2, 1842, p. 11, pi. 1 (part: Mississippi; great northern
lakes; Mohawk River, New York). — DeKAY, Zool. New York, pt.
3, Rept., 1842, p. 6, fig. 11 (Mohawk River; Hudson River near
Albany, New York).— TROOST, Seventh Geol. Tennessee, 1844,
p. 39 (Tennessee).— THOMPSON, Hist. Vermont, 1853, p. 29
(Vermont: Rivers Lamoille and Winooski). — GRAY, Cat. Tort.
Brit. Mus., 1844, p. 49 (part); Cat. Shield Rept., 1, 1856, p. 68
(part).— KENNICOTT, Trans. Illinois Agric. Soc, 1, 1855, p. 591
(Illinois: Cook Co., Lake Michigan).— STRAUCH, Mem. Acad.
Sci. St. Petersbourg, ser. 7, 5, no. 7, 1862 (part); 8, no. 13, 1865, p.
122 (part).— DITMARS, Reptile Book, 1907, pi. 27, upper fig. (not
of text).— GADOW, Cambridge Nat. Hist., 8, 1901, p. 408 (part),
p. 409, fig. 92 (North America).
Amyda ferox ORTENBURGER, Copeia, no. 170, May 1929, p. 12
(Oklahoma: LeFlore Co.); p. 28 (Oklahoma: Rogers Co.).

1827 .—Trionyx spiniferus LESUEUR, Mem. Mus. Hist. Nat. Paris, 15, Dec-
1827, p. 258, pi. 6 (type-locality, Wabash River, New Harmony*
Indiana; types in Paris Mus.; Lesueur, collector). — WIED, Reise
Nord-Amerika, 1, pt. 3, 1838, pp. 140, 141 (Pittsburgh, Pa.); Voy.
Amer. Nord, 3, 1843, p. 242.— HAY, Indiana Geol. 17 Rep., 1892, p.
554; Batr. Rept. Indiana, 1893, p. 146 (Indiana: generally distri-
buted).—HURTER, Trans. Acad. Sci. St. Louis, 6, 1892, p. 260
(Missouri: Mississippi River, Merimac River, and Illinois River). —
HAHN, Proc. U. S. Nat. Mus., 35, Dec. 1908, p. 567 (Lawrence Co.,
111.).— SIEBENROCK, Zool. Jahrb. Suppl., 10, pt. 3, 1909, p. 604
(Mississippi River and tributaries; St. Lawrence River; Hudson
River) ; Verh. Zool.-Bot. Ges. Wien, 73, Aug. 1923, p. 186.— THOMP-
SON, Thirteenth Rep. Michigan Acad. Sci., 1911, pp. 106, 107, fig. 1
(Cass Co., Michigan).— ELLIS and HENDERSON, Univ. Colorado
Stud., 10, no. 2, May 1913, p. 112 (Colorado: Weld Co.: Evans,
Cache la Poudre, South Platte River, and Greeley).

1 Spine bearing, with reference to the pointed shape of the tubercles margining the anterior
flap of the leathery carapace, characteristic of older specimens.

stejneger: American soft-shell turtles 45

Gymnopus spiniferus DUMERIL and BIBRON, Erpet. Gen., 2, 1835,
p. 477 (atlas, pi. 22, fig. 1) (Wabash River).— DUMERIL, Cat.
Meth. Coll. Rept. Mus. Hist. Nat. Paris, 1851, p. 22 (Wabash River).
— SAGER, Peninsular Journ. Med. Collat. Sci., 3, no. 8, 1856, p. 361
(anatomy). — WIED, Nova Acta Acad. Leopold. -Carol., 32, pt. 1,
1865, p. 48 (Wabash River).

Gymnopodus spiniferus DUMERIL, Arch. Mus. Hist. Nat., 7, 1856,
p. 203.

Aspidonectes spiniferus RHOADS, Proc. Acad. Nat. Sci. Philadelphia,
1895, p. 386 (Tennessee: Samburg, Obion Co.).

Aspidonectes spinifer AGASSIZ, Contr. Nat. Hist. United States, 1,
1857, p. 403; vol. 2, pi. 6, figs. 1-2 (Lake Champlain; Lake Ontario
and Erie; New York, Pennsylvania, Ohio, Indiana, Illinois, Missouri,
Michigan, Wisconsin, Iowa, Ft. Union, Montana). — MILES, First
Biennial Rep. Geol. Surv. Michigan, 1861, pp. 232, 233 (Michigan:
as far W. as Genessee Co.). — ALLEN, Proc. Boston Soc. Nat. Hist.,
1874, p. 69 (Musselshell and Yellowstone Rivers, Montana). —
COPE, Bull. U. S. Nat. Mus., No. 1, 1875, p. 51.— JORDAN, Man.
Vert. North Amer., ed. 3, 1880, p. 168; ed. 5, 1888, p. 206 (Canada
to Kentucky and Minnesota); ed. 8, 1899, p. 206.— CRAGIN,
Trans. Kansas Acad. Sci., 2, 1881, p. 116 (Kansas: Franklin and
Douglas Cos.).— CONES and YARROW, Bull. U. S. Geol. Geogr.
Surv., 4, 1878, p. 261.— SMITH, Rep. Geol. Survey Ohio, 4, 1882,
p. 668 (all streams flowing into Ohio and Lake Erie). — TRUE, Bull.
U. S. Nat. Mus., No. 24, 1883, p. 29 (Webster City, Iowa; Mt.
Carmel, Illinois; Fox River, Illinois; Ft. Laramie, Nebraska); Fish.
Industr. United States, Sect. 1, 1884, p. 152.— HOY, Geol. Wiscon-
sin, Surv. 1873-1879, 1, 1883, p. 423 (Western Wisconsin).— DAVIS
and RICE, Illinois State Lab. Nat. Hist., Bull. No. 5, 1883, p. 52
(north of Ohio River); Bull. Chicago Acad. Sci., 1, no. 3, 1883, p. 32
(Illinois).— HOY, Geol. Wisconsin, 1, 1883, p. 423 (Wisconsin).-
HUGHES, Bull. Brookville Soc. Nat. Hist., No. 2, 1884, p. 41
(Franklin Co., Indiana).— STOCKWELL, Journ. Comp. Med. Surg.,
9, 1888, p. 28.— HIGLEY, Trans. Wisconsin Acad. Sci., 7, 1889, p.
159 (south and west Wisconsin). — GARMAN, H., Bull. Illinois
State Lab. Nat. Hist., 3, 1892, p. 246 (throughout Illinois: Rock
Creek, Kendall Co.; Piano; Oregon, Ogle Co.; Peoria, Peoria Co.;
Bluff Lake, Union Co.; Wabash River, Mt. Carmel).— KIRSCH,
Bull. U. S. Fish Coram. 1894 (1895), p. 81 (Eel River, Indiana).-
McLAIN, Notes Coll. Rept. Arkansas, 1899, p. 1 (Bloomington,
Monroe Co., Indiana). — SMITH, Proc. Linn. Soc. New York,
1898-99, No. 11, 1899, p. 24.— RAMSEY, Proc. Indiana Acad. Sci.,
1900 (1901), p. 224 (Lake Winona, Indiana).— ATKINSON, Ann.
Carnegie Mus. Pittsburgh, 1, 1901, p. 154 (Monongahela River,

46 bulletin: museum of comparative zoology

Pennsylvania).— MORSE, Ohio Natural., 1, no. 8, 1901, p. 127
(Columbus, Ohio); Proc. Ohio Acad. Sci., 4, pt. 3, Spec. Pap. No. 9,
1904, p. 138 (Ohio: Columbus; Sandusky; London).— PAULMIER,
New York State Mus. Bull. 51, 1902, p. 392 (Lakes Ontario and
Erie; through Erie Canal to Hudson River). — CLARK, Rep. 4
Michigan Acad. Sci., 1904, p. 193 (Eaton Co., Michigan).— JOR-
DAN, Man. Vert. Anim. U. S., 1904, p. 206.— GIBBS, NOTE-
STEIN and CLARK, Rep. 7, Michigan Acad. Sci., 1905, p.
110 (Michigan: Brookfield, Ann Arbor, Olivet, Kalamazoo, Van
Buren, Montcalm and Allegan Cos.). — STONE, Amer. Natural.,
40, 1906, p. 168 (into Delaware Valley; Cooper's Creek and Warren
Co., New Jersey; Allegheny River, western Pennsylvania). — NASH,
Check List Vertebr. Ontario, 1906, p. 17 (Canada: Ontario, western
part; one record Ottawa River). — FOWLER, Rep. New Jersey State
Mus., 1906 (1907), p. 211, pi. 57 (Cooper's Creek; Paulin's Kill,
Hainesburg, Warren Co., New Jersey, introduced). — SURFACE,
Zool. Bull. Pennsylvania Dep. Agric, 6, nos. 4-5, 1 Sept., 1908, p.
121, fig. 2 (Pennsylvania: Indiana Co.; Somerset Co.). — REED and
WRIGHT, Proc. Amer. Philos. Soc, 48, 1909, p. 408 (Cayuga Lake,
New York).
Trionyx spinifer BOULENGER, Cat. Chel. Brit. Mus., 1889, p. 259
(Foxbury, Pennsylvania; Wabash River). — SCHNEE, Zeitschr.
Naturwiss., 72, pt. 3, Dec. 24, 1899, p. 197.— SIEBENROCK, Sitz.
Ber. Akad. Wiss. Wien, Math.-Nat, KL, 111, 1902, p. 829, fig. 10
(plastron).— HENSH AW, Occas. Pap. Boston Soc. Nat. His., 7,
pi. 1, 1904, p. 4 (Lake Champlain, Vermont).— DITMARS, Rep-
tile Book, 1907, p. 77, pi. 26, upper and middle figs.; pi. 28 (Quincy,
Adams Co., Illinois).

Platypeltis spinifer BAUR, Proc. Amer. Philos. Soc, 31, 1893, p. 220
(Wabash River, Indiana).— EVERM ANN and CLARK, Proc.
Indiana Acad. Sci., 1916 (1918), p. 473 (Lake Maxinkuckee, Indiana).

Platyrettis KIRSCH, Bull. U. S. Fish Comm. 1894 (1895), p. 333
(Maumee River Basin, Ohio and Indiana).

Amyda spinifer -POTTER, Copeia, No. 82, Oct. 15, 1921, p. 76 (Thorn-
apple River, Barry Co., Michigan).— SOLA, Bull. New York Zool.
Soc, 34, no. 5, Sept.-Oct,, 1931, pp. 134, 141, 155, fig. 6 (Lake Cham-
plain).

Callinia spinifera GRAY, Proc Zool. Soc. London, 1869, p. 222
{spicifera err. typogr.); 1873, (p. 60, figs.); Suppl. Cat. Shield. Rept.
Brit, Mus., pt. 1, 1870, p. 109.

Amyda spinifera HURTER, Trans. Acad. Sci. St. Louis, 20, 1911, p.
251 (Missouri: Mississippi, Missouri, Osage, Gasconade, Meramee
and White Rivers).— STEJNEGER and BARBOUR, Check List

stejneger: American soft-shell turtles 47

North Amer. Amph. Rept., Ed. 1, Dec. 12, 1917, p. 125 (Mississippi
River and tributaries west to Colorado, north to Montana; St.
Lawrence River and tributaries; east to Vermont, western New
York and Pennsylvania); Ed. 2, 1923, p. 141; Ed. 3., 1933, p. 153.-
WRIGHT, Copeia, No. 66, Feb. 25, 1919, p. 8 (Bays of Lake On-
tario, New York).— BABCOCK, Mem. Boston Soc. Nat. Hist., 8,
no. 3, April 1919, p. 419 (east shore, Lake Champlain, Vermont). —
CLARK and SOUTHALL, Rep. U. S. Comm. Fish 1919 (1920),
App. no. 7, p. 15, pis. 7-8 (economics).— EVERMANN and CLARK,
Lake Maxinkuckee, 1920, p. 592 (habits).— ORTENBURGER,
Copeia, No. 99, Oct. 15, 1921, p. 76 (Decatur Co., Indiana); Proc.
Oklahoma Acad. Sci., 6, pt. 1, 1926, p. 100 (McCurtrin and Push-
nataha Cos., Oklahoma). — BLANCHARD, Occas. Pap. Zool. Mus.
Univ. Michigan, No. 117, July 6, 1922, p. 18 (Reelfoot Lake, West.
Tennessee); Univ. Iowa Studies Nat. Hist., 12, no. 2, 1923, p. 24
(Little Sioux Riv., Dickinson Co., Iowa); Pap. Michigan Acad. Sci.,
5, 1925, p. 386 (10 miles W of Columbus, Indiana).— WEED,
Copeia, No. 116, March 15, 1923, p. 48 (Meredosia, Illinois).—
BISHOP, Copeia, No. 125, Dec. 31, 1923, p. 120 (Albany Co., New
York).— PRATT, Man. Vert. United States, 1923, p. 249.— LO-
GIER, Canad. Field Natural., 39, May, 1925, p. 95 (Point Pel6e;
Hamilton, Dundas Marsh, Ontario, Canada); Roy. Ontario Zool.
Mus. Handb. no. 4, 1939, p. 56, pi. 7, fig. 2; pi. 8, fig. 7 (Ontario:
Hamilton Bay and Dundas Marsh, Wentworth Co.; Thames River
at Beachville, Oxford Co.; Grand River at Dunnville, Haldimand
Co.; Long Point, Norfolk Co., and Point Pelee, Essex Co.). —
SCHMIDT, Copeia, no. 154, May 20, 1926, p. 132 (Chippewa Co.,
Wise.).— MYERS, Proc. Indiana Acad. Sci., 35, 1926, p. 292 (In-
diana); vol. 36, 1927, p. 339 (Helmsburg, Indiana). — BURT, Occas.
Pap. Zool. Mus. Univ. Michigan, no. 189, Dec. 12, 1927, p. 9 (Riley
Co., Kansas).— LINSDALE, Copeia, no. 164, July-Sep., 1927, p. 81
(Doniphan Co., Kansas); Trans. Kansas Acad. Sci., 36, 1933, p. 208
(Cowley and Lane Cos., Kansas).— RUTH VEN, THOMPSON, and
GAGE, Herpet. Michigan, 1928, p. 163, pi. 19, fig. 3 (Michigan).-
MERTENS, Zool. Gart. Leipzig, new series, 1, pt. 5-6, 1928, p. 199
(distr. New York State by canals). — GLOYD, Trans. Kansas Acad.
Sci., 31, 1928, p. 135 (Franklin Co., Kansas).— POPE and DICKIN-
SON, Bull. Publ. Mus. Milwaukee, 8, no. 1, Apr. 3, 1928, p. 82, pi. 21,
figs. 7-8 (Wisconsin: Burnett, Crawford, Grand, Oneida, Polk,
Waukesha, Chippewa, Washburn, and Pepin Cos.). — POPE, Year
Book Mus. Milwaukee, 1928 (1929), pp. 180, 183 (Vernon and
LaCrosse Cos., Wisconsin). — JORDAN, Man. Vert. Northeast U.S.
(13 ed.) 1929, p. 254.— CAHN, Copeia, no. 170, May 1929, p. 8
(Lake La Belle, Waukesha Co., Wisconsin). — FORCE, Copeia,
1930, no. 2, p. 38 (Tulsa Co., Oklahoma).— DOLLEY, Amer. Mid-

48 bulletin: museum of comparative zoology

land Natural., 14, no. 3, May, 1933, p. 203 (St. Joseph Riv., Berrien
Co., Michigan).— RUST, Blatt. Aquar. Terrarienk., 45, 1934, p. — ,
sep. p. 12.— BURT and HOYLE, Trans. Kansas Acad. Sci., 37,

1934, p. 198 (Kansas).— BURT, Amer. Midland Natural., 16, no. 3,

1935, p. 321 (Kansas: Barber, Reno and Sedgwick Cos.).— BOYER
and HEINZE, Trans. Acad. Sci. St. Louis, 26, no. 4, Apr. 1, 1934, p.
199 (Missouri: Jefferson Co., common).— NETTING, Nawakwa
Fireside (N.S.), nos. 3-4, Apr., 1935, p. 49 (Penna,: 8 counties);
Proc. Pennsylvania Acad. Sci., 10, 1936, p. 27 (Pennsylvania, In-
diana Co., Plum Creek and Crooked Creek).— BRINN, Rep. Penn-
sylvania Fish Comm. for 1938-1939, p. 127 (Pennsylvania: rec. from
9 counties in Ohio and Lake Erie drainages). — SCHMIDT and
NECKER, Bull. Chicago Acad. Sci., 5, no. 4, Sept. 27, 1935, p. 76
(Illinois: Cook Co., Kankakee Co.; Indiana: Lake Co.).— TAYLOR
Univ. Kansas Sci. Bull., 22, no. 11, Apr. 15, 1935, p. 217 (Arkansas:
Devall Bluff, Prairie Co.; Lewisville, Lafayette Co.).— BABBITT,
Bull. Boston Soc. Nat. Hist., no. 78, Jan. 1936, p. 10 (Lake Cham-
plain, Vermont, rare). — HIBBARD, Trans. Kansas Acad. Sci., 39,

1936, p. 281 (Mammoth Cave Nat. Park, Kentucky).— PARKER,
Jour. Tennessee Acad. Sci., 12, no. 1, Jan., 1937, p. 85, fig. 18 (Bayou
du Chien; Reelfoot Lake, Tennessee); Rep. Reelfoot Lake Biol.
Sta., 3, Jan. 1939, p. 88 (Tennessee: Reelfoot Lake).— CAHN,
Illinois Biol. Monogr. 16, Aug. 31, 1937, p. 184, pis. 25-27, 30 fig. b,
map 20 (Illinois).— GREETE, Herpetologica, 1, no. 4, Nov. 16,

1937, p. 116 (West Virginia: Randolph Co., Tygart River near El-
kins). — ALEXANDRE, Soc. Canadienne Hist. Nat., tract no. 39,
Apr. 1937, p. 2 (Canada: Quebec: Lake Champlain and Richelieu
River, tributary to St. Lawrence R., right side).— HENNING,
Copeia, 1938, no. 2, June 30, 1938, p. 92 (Boone Co., Missouri).—
DENNING and BLACK, Occas. Pap. Univ. Arkansas Mus., No. 1,
June 1938, p. 46 (Arkansas: Lafayette, Prairie, Chicot, Clay, Gar-
land, Lawrence, and Washington Cos.). — CONANT, Amer. Mid-
land Natural., 20, no. 1, 1938, p. 157, pi. 21, fig. 1 (right), pi. 22,
figs. 1, 2 (Ohio: map); Herpetologica, 1, no. 5, Dec. 30, 1938, p. 138
(Ohio: Lucas Co.).— NECKER, Bull. Chicago Acad. Sci., 6, no. 1,
1939, p. 10 (Illinois: Cook Co.: Evanston; Kankakee Co.: Kankakee
Riv. near Altort).— WELTER and CARR, Copeia, 1939, no. 3,
Sept. 9, p. 130 (Kentucky: Triplet Co.; Fox, Fleming Co.; rare in
East Ky.).— LOGIER, Canad. Field-Nat., 39, May, 1925, p. 95
(Point Pelee; Hamilton, Dundas Marsh, Ontario, Canada); Roy.
Ontario Zool. Mus. Handb. no. 4, 1939, p. 56, pi. 7, fig. 2; pi. 8, fig. 7
(Ontario: Hamilton Bay and Dundas Marsh, Wentworth Co.;
Thames River at Beachville, Oxford Co. ; Grand River at Dunnville,
Haldimand Co.; Long Point, Norfolk Co.; and Point Pelee, Essex
Co.).— GENTRY, Rep. Reelfoot Lake Biol. Sta,, 5, Jan. 1941, p.

stejneger: American soft-shell turtles 49

75 (Tennessee: Clay Co., Overton Co.); Journ. Tennessee Acad.
Sci., 16, no. 3, 1941, p. 332 (Tennessee: Overton, Fentress, Pickett,
Jackson, and Clay Cos.).— CAGLE, Copeia, 1942, no. 3, Oct. 8,
p. 155 (Illinois: Jackson and Williamson Cos.)
Platypeltis spinifera RUTHVEN and THOMPSON, Herpet. Michi-
gan, 1912, p. 129 (Michigan).— THOMPSON, Misc. Pap. Zool.
Michigan, 1916, p. 63 (Monroe Co., Michigan).

1827.— Trionyx ocellatus LESUEUR, Mem. Mus. Hist. Nat. Paris, 15, Dec.
1827, p. 261 (type-locality, New Harmony, Indiana).— DeKAY,
Zool. New York, pt. 3, Reptiles, 1842, p. 7 (Mohawk River, New
York).

1838. — Trionyx annulifer WIED, Reise Nord-Amerika, 1, pt. 3, p. 140 (type-
locality, Ohio River at Pittsburgh, Pennsylvania); Voy. Amer.
Nord., 3, 1843, pp. 242-243.

1844.— Tyrse argus GRAY, Cat. Tort. Brit. Mus., p. 48 (type-locality, "West
Africa, Sierra Leone?"; type in Brit. Mus.; Lord Derby, collector);
Knowlsley Menag., 1846 (pi. — ).
Trionyx argus GRAY, Cat. Shield Rept. Brit. Mus., pt. 1, March 8,
1856, p. 68.

1856. Trionyx annularis "Wied", GRAY, Cat. Shield Rept. Brit. Mus., pt.

1, March 8, 1856, p. 69 (lapsus in synonymy).

1857. Aspidonectcs nuchalis AGASSIZ, Contr. Nat. Hist. United States, 1,

p. 406 (type-localities, Cumberland and Tennessee rivers; cotypes,
Mus. Comp. Zool., nos. 1623-1625, Cumberland river).— COPE,
Bull. U. S. Nat. Mus., no. 1, 1871, p. 51.— TRUE, Bull. U. S. Nat.
Mus., no. 24, 1883, p. 5; Fish. Industr. United States, sect. 1, 1884,
p. 152 (middle western states).— DAVIS and RICE, Illinois State
Lab. Nat. Hist., Bull. No. 5, 1883, p. 52 (Cumberland and Upper
Tennessee Rivers).— JORDAN, Man. Vert. Anim. North United
States, ed. 5, 1888, p. 206 (Cumberland and Upper Tennessee Rivers) ;
ed. 8, 1899, p. 206.— RHOADS, Proc. Acad. Nat. Sci. Philadelphia,
1895, p. 404 (Cumberland River, near Nashville).

1861. Amyda mutica MILES, First Biennial Rep. Geol. Surv. Michigan, pp.
2323, 233 (Michigan). (Not of Lesueur).— SMITH, Science News
Suppl., 1, 1879, p. 7 (Michigan).— KIRSCH, Bull. U. S. Fish Comm.
1894 (1895), p. 41 (Indiana: Eel River); p. 333 (Maumee Riv.,
Ohio).

1865. — Gymnopus olivaceus WIED, Nova Acta Acad. Leopold.-Carol., 32,
pt. 1, p. 55, pi. 5 (type-locality, New Harmony, Wabash River,
Illinois).

1869.— Callinia spicifera GRAY, Proc. Zool. Soc. London, 1869, pp. 222
(lapsus for spinifera).

50 bulletin: museum of comparative zoology

1939.— Amyda spinifera spinifera STEJNEGER and BARBOUR, Check
List N. A. Amph. Rept., ed. 4, p. 172; ed. 5, 1943, p. 213.— ANDER-
SON, Bull. Chicago Acad. Sci., 6, no. 11, July 8, 1942, p. 219 (Mis-
souri: Jackson Co.; Missouri River near Atherton). — EDGREN,
Copeia, 1942, no. 3, Oct. 8, p. 180 (Michigan: Van Buren Co., Rey-
nold Lake).— PETERS, Copeia, 1942, no. 3, Oct. 8, p. 183 (Illinois:
Cumberland Co.).

Geographical distribution

Amyda spinifera has been credited to the fauna of South Carolina
on the strength of the specimen U.S.N.M. No. 7650 recorded in U. S.
Nat. Mus. Bulletin No. 24 (1883), p. 29, as coming from "Abbeville,
S. C". It is also the record upon which is based True's record of the
occurences of Aspidonectes spinifer in H. S. Thompson's, South Caro-
lina, 1883, p. 238. The record is undoubtedly erroneous, the ascertain-
able facts being as follows :

The specimen, recently hatched, was one of the numerous neglected
turtles found in the collection when Dr. G. Brown Goode took charge
and tintagged and registered the specimens. He entered the present
one in 1872 as No. 7650, the tin tag bearing that number. In the record
book [in Brown Goode's handwriting] it is noted as an Aspidonectes
[no specific name] ; locality : "Abbeville S. C"; and no further remarks.
There is now no other indication of its origin than an old torn scrap
of paper in the bottle with "7650 Abbeville S. C." likewise in his hand-
writing. On the back of this label, however, there is — in very faded
ink and in an entirely different handwriting — the remnant of an in-
scription, beginning and end clipped off, "ville Mis". If this remnant
really is part of the original locality record, it seems probable that it
may have read "Abbeville, Miss." and inadvertently attributed to
South Carolina as the more familiar locality. The specimen is unques-
tionably an Amyda spinifera. The whole pattern of coloration is
normal of that species, which of course at once excludes A. ferox. The
single regularly defined submarginal dusky ring on the upper side of
the disc likewise excludes A. agassizii, the species one would expect
if it were collected in Abbeville, S. C. To make perfectly sure of its
identity I have had the skull extracted. It shows all the characteristic
features of A. spinifera as distinguished from A. agassizii.

Another erroneous record of Trionyx spiniferus, viz. in Pope's
Turtles (1931) pi. 45, figs. 98, 99) showing excellent figures of two soft-
shelled turtles from the region of Columbia, South Carolina, needs
correction. Both pictures are plainly of Amyda agassizii.

stejneger: American soft-shell turtles

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52 bulletin: museum of comparative zoology

Stockwell's reference to the occurrence of Aspidonectes spinifer
"North of Athabasca Lake" (Journ. Compar. Medic. Surg., 9, 1888,
p. 28) must rest on some curious lapsus.

Where Amy da spinifer meets with A. asper is as yet conjectural.
It extends at least as far south as the northern part of the State of
Mississippi, for we have undoubted specimens from De Soto County
(U.S.N.M. No. 92606) and Lake Washington, Washington County
(No. 92607). A young male specimen from Madison Co., Northern
Louisiana (No. 83985) is likewise this form with only one marginal
stripe.

The Museum of Comparative Zoology has a specimen from Colum-
bus, Ga (M.C.Z. 1606).

Mississippi River and tributaries, west to Colorado, north to Mon-
tana; St. Lawrence River and tributaries; east to Vermont, western
New York, and Pennsylvania.

A. nuchalis, judging from the slight material at hand, is not a
strongly differentiated form. The character chiefly relied on by Agas-
siz, "the most prominent specific character . . . the marked depression
on either side of the blunt median keel," does not hold in a series of
specimens. I find quite a number so characterized among typical A.
spinifer a, and while one of the specimens from near Sevier ville, Tenn.,
has a rather flat bony carapace, the other has "the blunt keel, which
extends along the median line and slopes uniformly upon the sides,"
exactly as he describes it diagnostically for A. spinifera (p. 404).
The angle of the entoplastron, in the few examples examined, is some-
what more obtuse, between 90° and 95°, than in corresponding speci-
mens of typical A. spinifera. The sharply defined ocelli on the carapace
seem to be larger than in A. spinifera of the same age (size).

I do not at all understand Agassiz's note that "this species differs
strikingly from Asp. spinifer in the much more elongated form of the
male, and in the great development of the marginal spines and of the
tubercles upon the carapace, which project very slightly in the male
Asp. spinifer." On the contrary, in our upper Tennessee specimens,
presumably typical A. nuchalis, the carapace of the males is wider
than in the females, and the spines on the anterior edge very much
smaller, exactly as in typical A. spinifera.

U.S.N.M. 86677 from Cumberland Gap, and 86682 from 2 miles
west of Sevierville, Tennesse, are typical "nuchales."

stejneger: American soft-shell turtles

53

List of specimens in the U. S. National Museum

58

Mont., Ft. Union

F. V. Hayden

7163(029528)

Tenn., Nashville

J. Varden

7165(029529)

it it

"

7166(029530)

it a

tt

7167

n it

tt

7169 juv.

tt tt

a

7648 & adol.

Wyo., Ft. Laramie

F. V. Hayden

7649

?

?

7650 pull.

Miss., Abbeville?

?

7661 juv.

111.

R. Kennicott

8359

Ind., Madison ?

?

9654

111., Mt. Carmel

R. Ridgway

May,

1878

9717

111., Mt. Carmel

Mrs. L. M. Turner

June,

1878

9928 juv.

Iowa, Webster City

C. Aldrich

1878

11625 juv.

?

?

11631 juv.

La., Prairie Mer Rouge

?

12061 juv.

111., Mt. Carmel

L. M. Turner

14535 juv.

Mont., Ft. Custer

C. Bendire

14536 9 ad.

it a

tt

16704 9

Ala., Courtland

rP. H. Kirch, E. O.

■

Jones, and

May,

1889

16705

ti a

W. M. Andrews

tt

17823 juv.

Ark., Benton

Jordan & Gilbert

19622-3 9 & adol.

?

19625

?

?

21128-9 juv.

Ohio, Cuyahoga River

A. J. Woolman

July

25, 1893

21416-7

?

?

21567-8

Ohio, Edgerton

P. H. Kirsch

July

28, 1893

21569-70

Ind., Fish Cr., near

Hamilton

tt

July

21, 1893

21571-7

Ohio, Maumee Basin

a

1893

22711

Ind., Vincennes

R. Ridgway

24536

Mont., Ft. Custer

C. Bendire

Mar.

8, 1886

26290

Ohio, Franklin Co.

R. C. Osborn & E.

C. Williamson

June,

1897

029014 9

" Columbus

O. Davie

33494

Ind., Lake Maxin-

kuckee

B. W. Evermann

July

21, 1900

33495

Yellow River nor(

h

of Burr Oak

tt

Oct.

3, 1900

33496-501

" L. Maxinkuckee

1899-

-1900

33767

W. Va., Dry Fork,

Perryville

W. P. Hay

1900

34 bulletin: museum of comparative zoology

35404-8

Ind., L. Maxinkuckee

H. W. Clark

1900

36412

111., Illinois R.

S. P. Bartlett

42583 juv.

Ind., Long Point, L.

Maxinkuckee

Evermann & Clark

Oct.

5, 1906

42584 juv.

" L. Maxinkuckee

n a

42585 eggs

u a

a it

Nov.

17, 1906

42905-6 juv.

" Burlington

B. W. Evermann

1899

50670 juv.

Twin Lakes

H. W. Clark

July

2, 1909

51213 hf-gr.

Mich., Monroe Piers

C. Rutter

Aug.

13, 1894

51214 "

Ohio, Toledo, Grassy

Point

<<

Aug.

3, 1894

51529 ad.

Kansas

R. L. Moodie

53521

Iowa, Fairport

Bur. Fisheries

Apr.

24, 1911

53522

111., Hamilton

J. McAdams

May

1, 1915

53523-6

Iowa, Fairport

E. Snyder

Oct.

1, 1914

54421 cf ad.

Mont., Crow Agency

M. A. Hanna

Aug.

5, 1916

54422-3

it a

R. Kellogg

July

8, 1916

54730 9 adol.

Iowa, Fairport

J. Snyder

June,

1916

54731 9 adol.

a tt

a

July

2, 1916

54732

a a

tt

a

54739-41

n a

a

May

8, 1916

54743-6

a a

ti

May,

1916

54747

111., Meredosia

Freeland & Williams 1908

55680

" Madison Co.

J. Hurter

55681

" Union Co.

!(

55682

Ky., Morgan Co.

It

'55683

Kans., Greenwood Co.

It

July-Aug., 1912

55684

Mo. Stone Co.

It

June 27, 1908

55085

" St. Louis Co.

It

1913

55686

" St. Louis

It

55687

it a

tt

Apr.

9, 1905

55688

" Reynolds Co.

a

July

21, 1911

55689

" FranklynCo.

a

Aug.

20, 1911

55690

" Washington Co.

tt

June

25, 1911

59046

" St. Louis

ti

May

20, 1908

59263-6 9 adol

. Miss., Homer

F. Schrader

59267 9

Mo., Alexandria

E. Stringham

June

5, 1916

59270-5

Minn., Homer

F. Schrader

Sept.

8, 1916

59277

111., between Warsaw

and Hamilton

E. Stringham

Aug.

25, 1916

59279-80

Mo., Canton

Bur. Fisheries

59285

Iowa ?

n

59736

Mont., Crow Agency

R. Kellogg

July

23, 1916

59956

Ind., Madison

O. P. Hay

59979 d" juv.

Ind. ?

ti

stejneger: American soft-shell turtles

55

60571 juv.

III., Madison Co.

J. Hurter

70397 adol.

Okla., Red River,

McCurtain Co.

A. I. Ortenburger

1924

72387

Ind., Knox

H. W. Clark

Aug.

12, 1909

73668-9 juv.

Miss., Greenwood

I. L. Towers

1925

83985 <? adol.

La., 2 mi. E. of Mounds C. E. Burt

June

30, 1931

86677 9 ad.

Tenn., 5 mi. S.E.

Cumberland Gap

a

July

22, 1932

86681-2 d" adol

.. " 2 mi. W. Sevierville

1932

87164 juv.

" 2 mi. S. Kingston "

July

27, 1932

90441-4 juv.

Kans., 1 mi. W. Winfield

Apr.

30, 1933

91022 juv.

" Winfield

L. Hoyle

May

28, 1933

92606 pull.

Miss., Lake Cormorant S. F. Hildebrand

June

9, 1933

92607 juv.

" Greenville

a

May

31, 1933

93089-94 juv.

Mo., Dardenne Cr.,

St. Charles Co.

L. Hubricht

Aug.

16, 1931

95140 pull.

Miss., 1 mi. W. Yazoo

City

C. E. Young

Aug.

13, 1934

95261 9 ad.

Kans., 2 mi. E. of

Calista

C. E. Burt

May

25, 1934

95301

" 11 mi. S.E. of

Winfield

<<

Aug.

31, 1934

95352 9 ad.

Ark., 7 mi. N.W.

Natural Dam

it

June

7, 1934

95405 & ad.

Mo., Glaize Creek,

Jefferson Co.

A. A. Heinze

July

20, 1932

99862-75

La., Red River near

Shaw

C. E. Burt

1935

100160 9 ad.

" Bayou Chene

it

June

17, 1935

100202-12

" False River near

New Roads

n

June

17, 1935

100420-1

" Cane River near

Natchitoches

it

June,

1935

100529-30 & 9

Kans., Winfield

a

Aug.

12, 1935

100580

" Lake City

it

June,

1935

100795 pull.

Ind., Irvington

O. P. Hay

101386 9 ad.

Va., Seven Mile Ford

A. Wetmore

June

1, 1935

102705 juv.

Miss., Belzoni

S. F. Hildebrand

July

10, 1936

102911 c? juv.

Tenn., Reelfoot Lake

W. M. Perrygo &

C. Lingebach

May

8, 1937

102912

n u u

W. M. Perrygo &
C. Lingebach

it

103477-9 juv.

Vt., Swanton

L. H. Babbitt

June

21, 1937

56

bulletin: museum of comparative zoology

107786

107787

109178 pull.
113228 juv.
115980 pull.

Term., Iron Creek near

Willow Grove G. Gentry

mouth of Wolf

River
La., Jonesville G. K. Payne

July 17, 1939

July 19, 1939
June, 1940

Miss., Deer Creek C. Hollingsworth June 24, 1940

"Amyda spinifera aspera1 (Agassiz)
Plates 17, 18, 19c

1854.— Trionyx ferox WAILES, Rep. Agric. Geol. Mississippi, pp. 327, 331

(Mississippi) (not of Schneider). — Plalypeltis ferox AGASSIZ, Contr.

Nat. Hist. United States, 2, 1857, pi. 6, fig. 3 (Mobile, Ala.; Mus.

Comp. Zool., no. 1608A).— Aspidonectes ferox JORDAN, Man.

Vert. North Amer., ed. 5, 1888, p. 206 (part).
1857. — Aspidonectes asper AGASSIZ, Contr. Nat. Hist. United States, 1, p.

405 (type locality, Lake Concordia, La.; cotypes, U.S.N.M. no.

012349, Prof. B. L. C. Wailes, collector, and Mus. Comp. Zool, no.

37173).— COPE, Bull. U. S. Nat. Mus., no. 1, 1875, p. 51 (Lower

Mississippi tributaries). — DAVIS and RICE, Illinois State Lab.

Nat. Hist., Bull. no. 5, 1883, p. 52.— TRUE, Fish. Industr. United

States, sect. 1, 1885, p. 152.— BAUR, Amer. Natural., 22, 1888, p.

1122; Proc. Amer. Philos. Soc, 31, July 1893, p. 217.— BEYER,

Proc. Louisiana Soc. Nat. Hist., 1897-1899 (1900), p. 43 (Louisiana).
1892. — Trionyx agassizii HAY, Indiana Geol. 17 Rep., p. 552 (part); Batr.

Rept. Indiana, 1893, p. 144 (part) (not of Bauer 1888).
1899. — Aspidonectes agassizi JORDAN, Man. Vert. North Amer., ed. 8, p. 206

(part only) (emendation) (not of Bauer).

1919. — Amyda spinifera BABCOCK, Mem. Boston Soc. Nat. Hist., 8, no. 3,
pi. 32 (not of text; not of Lesueur).— HALTOM, Alabama Mus. Nat.
Hist., Mus. Pap. no. 11, 1931, p. 142 (Alabama: Marengo Co.: near
Demopolis, Tombigbee River).

1923.— Trionyx spiniferus agassizii SIEBENROCK, Verh. Zool. Bot. Ges.
Wien, 72, Aug. 1923, p. 188 (part: West Louisiana).

1939.— Amyda spinifera aspera STEJNEGER and BARBOUR, Checklist N.
Amer. Amph. Rept., ed. 4, p. 172 (Lower Mississippi tributaries in
Louisiana and Mississippi) ; ed. 5, 1943, p. 213 (Lower Mississippi tri-
butaries in Louisiana; rivers of Mississippi and Alabama).

1 Latin: rough; probably with reference to the "prominent warts of the bony plates," which,
according to Agassiz, "exist in no other species with which I am acquainted."

stejneger: American soft-shell turtles 57

•

Types and type locality. The two specimens specifically mentioned
by Agassiz as basis for his A. as per were "an imperfect skeleton . . .
belonging to the Smithsonian Institution and prepared from a speci-
men forwarded by Professor B. L. C. Wailes of Washington, Missis-
sippi," and "a stuffed specimen belonging to the Museum of the
University of Oxford, that has been collected during the Geological
Survey of Mississippi, under the superintendence of Professor
Wailes."

In a letter dated Washington, Miss., 8: January, 1853, Professor
B. L. C. Waile wrote to Professor S. F. Baird that he had forwarded
to him a number of reptile specimens, among them "2 shells and
crania of Trionyx ferox." They were entered in the register of the
osteological collections of the Smithsonian Institution under the
generic name Trionyx only by Professor Baird on March 21, 1853 as
numbers 1084 and 1086, received from' 'B. L. C. Wailes, Washington,
Miss." Of these specimens there are now in the National Museum: 1)
a skull marked 1084; 2) a skull, with carapace and plastron in pieces
(entoplastron and epiplastra missing), marked 108G; and 3) a carapace
numbered 12349. The specimens have no original labels attached to
them, but the numbers on the skulls are written on them in ink, and
on the inside of the carapace of No. 12349 there is written with black
ink in the same "professional" hand characteristic of all the specimens
received from Wailes: "Trionyx ferox? Lake Concordia, Louisiana,
1851, B. L. C. Wailes." This is undoubtedly the "imperfect skeleton"
examined by Agassiz; it is characterized by the "prominent warts of
the bony plates" on the posterior part of the bony disk described by
Agassiz (I.e. p. 406), which "bony warts exist in no other species with
which I am acquainted."1

The second cotype, the "stuffed specimen" received from the Uni-
versity of Oxford, Mississippi, is in the Museum of Comparative Zoo-
logy (M.C.Z. no. 37173) where I was permitted to examine it. It is an
adult male, inscribed in the same bold, handsome style as Wailes'
other specimens Trionyx ferox, but no locality data. I made the fol-
lowing notes: "Bony disk 200 mm. long, 185 mm. wide; 7 neurals,
seventh small; sculpture fine, of the spinifer style, with the bony
tubercles on the seventh pleural described by Agassiz, tubercles on the
posterior and anterior flaps of leathery disk, spinous tubercles on edge
of carapace anteriorly between legs; callosities on plastron large, al-
most meeting on the mid-line; a median large triangular callosity

1 Agassiz did not know the bony disk of the true A. ferox from Florida.

58 bulletin: museum of comparative zoology

(sides about 25 mm.) on the entoplastron ; entoplastral angle slightly
less than 90°.

The specimens which Agassi z mentions as having been received
"through the kindness of Mr. Winthrop Sargent of Natchez"1 may be
regarded as paratypes. One of them, a very large male (M.C.Z. no.
1597), I have examined and made the following measurements and
notes :

Total length of leathery carapace 450 mm.

Width of leathery carapace 370

Length of bony carapace 240

Width of bony carapace (at pleural 4) 240
Height of body (at neural 1) 88

Large rounded tubercles on front edge of leathery carapace; flat
tubercles of about same size on front and hind flaps; callosities on
xiphi-plastra meeting; closest approach of hyo-hypoplastral callosities
8 mm. ; bony ridges comparable to those on bony carapace of U.S.N.M.
no. 1086; sculpture somewhat coarser (because of larger size of speci-
men) ; entoplastral angle about 90°. Distinct traces of two black mar-
ginal rings on hind flap of carapace.

A young paratype, probably of the same origin (M.C.Z. no. 1622)
with a recent label in lead pencil "No. 1622. Type Amyda asper
(Agassiz) Lake St. John, Miss.2 W. Sargent leg. et don." measures ap-
proximately 65 mm. in length and 57 mm. in width. The secondary
marginal lines on the posterior flap of the carapace characteristic of
normal individuals of the subspecies of corresponding age are repre-
sented by two series of closely set spots (pi. 16, fig 1).

Notes on synonymy. Special attention is called to the reference to
Agassiz's illustration (Contrib., 2, pi. 6, fig. 3) of a young specimen of
A. aspera under the name PlatypeHis fcrox, as it apparently has given
rise to great confusion among southern herpetologists who, because of
it, have mistakenly identified young specimens from Alabama, Missis-
sippi, and Louisiana. The young of the true A. ferox was unknown to
Agassiz, but on account of the locality (Mobile) of the specimen figured
and its similarity in color pattern to certain soft-shell turtles from
South Carolina he assumed that it represented the Florida species.
The original of the figure 3 appears to be still at the Museum of Com-
parative Zoology bearing the number 1608A. It turns out to be a

'Agassiz, in 1856, had J. Burkhart make colored illustrations from a female of the same lot
of specimens which shows plainly the marginal black rings on the posterior flap. Reduced
copies of the drawings are herewith presented (pis. 20, 21) through the kind permission of
Dr. Thomas Barbour, Director of the Museum of Comparative Zoology.

2 Possibly a slip for Louisiana.

stejneger: American soft -shell turtles 59

young of Agassiz's own Aspidonectes as per, and the figure is herewith
reproduced under its proper name (pi. 19, fig. c) for comparison with
the young of the true A. ferox (pi. 19, fig. a and b).

Babcock's figure of A. spinifera, as quoted above, does not represent
the true nominate form of that species, but is either an A. spinifera
aspera or an A. agassizii. Dr. Babcock, in a letter of December 13,
1933, kindly informs me that the picture was made, during his absence
in 1918, from a living specimen, the origin and disposition of which
he has been unable to trace. The exact identification depends some-
what on the artist's accuracy. My best guess is that the specimen was
an A. aspera, judging by the extreme length of the proboscis.

Variation. A series of four living young adults were presented in
September, 1934, by Mr. S. E. Brand, of Canton, Mississippi, all taken
in barrow pits, three on Pearl River and one on Big Black River. The
specimens unquestionably represent the same form and, as highly
instructive, some of the notes made at the time may be of interest, the
color designations in quotation marks having reference to Ridgway's
Nomenclature of Colors, 1886: —

U.S.N.M. no. 95191, 9 ad. Iris clear "primrose yellow" with black
horizontal bar not quite reaching pupil. General color above "tawny-
olive," head, neck, and legs densely speckled with dark brown, cara-
pace with obscure, irregular blotches of "raw umber," in the center
of which one or more small blackish spots form interrupted ocelli, the
two outer rows of which posteriorly assume the form of short lines
parallel with the submarginal ring; marginal edge dull "olive-buff,"
top of head like carapace with the shallow fork on the snout chiefly
indicated by the black outlines; side of head like top down to the lips;
a slightly paler postocular band tinged with yellow and narrowly but
very distinctly margined with black, the upper line continuing back-
ward the whole length of the neck ; upper side of legs slightly paler than
carapace and more tinged with olive; under side pale flesh color caused
by the fine network of blood vessels shining through the white ground
color; plastral callosities pale "vinaceous-cinnamon" more or less
tinged with bluish; under side of feet tinged with "verditer blue" with
a wash of yellow deepening towards the outer margin, inner half
spotless, outer half, including web, marked with heavy blackish anas-
tomozing lines and spots; claws yellowish white; under side of neck
faintly mottled with obscure "pinkish vinaceous" spots; throat
strongly washed with "pale blue"; lips and under side of proboscis
tinged with "gallstone-yellow." Tubercles on anterior edge of cara-
pace triangular, pointed, about 3 mm. long; anterior flap densely

GO bulletin: museum of comparative zoology

crowded with smaller and blunter tubercles of various sizes; posterior
flap with larger, oblong, blunt tubercles, fairly regularly spaced; cara-
pace posteriorly with similar but lower and longer tubercles in fairly
regular longitudinal series. Skin covering nuchal of bony carapace
with a median longitudinal series of about 4 enlarged tubercles ex-
tending on to anterior leathery flap. Fontanelles between nuchal and
first pair of pleurals covered with smooth skin upon which are a few
tubercles. Callosities covering plastral bones rather small, leaving
median fontanelles larger than xiphiplastral callosities; no trace of
callosities on entoplastron and ectoplastron ; entoplastral angle 90°.

U.S.N.M. no. 95192, also from a "barrow pit" at the nearby Big
Black River, a young adult 9 , smaller than 95191, is structurally al-
most identical with it, except for the smaller dimension and being
slightly broader. The coloration of the living specimen was darker and
more pronounced. Iris "ochraceous buff," black horizontal bar reach-
ing pupil. Carapace "raw umber" with large irregularly buffy-edged
blotches of "mummy brown," increasing towards and infringing upon
the marginal "buffy" edge and its bordering dusky ring; towards the
margin the blotches coalesce more or less so as to form three distinct
but interrupted submarginal rings; color of upper side of neck, head,
and «legs like that of the carapace but more "tawny" and thickly
sprinkled with heavy blackish spots ; forked figure on top of snout deep
"ochraceous buff," with very distinct black edges like the ochraceous
postocular stripe; under side milk white strongly suffused with flesh-
color; palms and soles strongly tinged with tawny, palms heavily
marked with coarse blackish anastomozing lines and spots, soles al-
most devoid of markings, claws white, throat tinged with pale blue;
callosities "vinaceous-cinnamon" with slight tinge of blue in center.

U.S.N.M. no. 95193, young adult d\ from Pearl River, differs
structurally from the two described females in showing no trace of
fontanelles on carapace, in much larger plastral callosities than on
xiphiplastra nearly meeting in the middle, those covering hyoplastra
and hypoplastra only 4 mm. apart; in addition there is a large triangu-
lar callosity on the entoplastron ; the row of tubercles on the edge of
anterior flap of carapace barely indicated and no tubercles on flap and
nuchal. In addition, the skin of the carapace, bony disk as well as
lateral and posterior flaps, is densely sprinkled with very minute hard
tubercles which make the skin feel like fine sandpaper. The color
notes on the living specimens are as follows : Iris pale buff, black bar
scarcely separated from pupil. Upper side nearly uniform "tawny
olive," the region of the bony disk distinctly more olive; small, round,

stejneger: American soft-shell turtles 61

dark brown spots of varying size scattered sparingly over the carapace;
the normal margin not perceptibly lighter than rest of carapace and
barely set off from it by the obscure dusky submarginal ring; no in-
dications of additional rings by dark lines or spots; top of head and
neck like back with minute black scattered dots; lips and sides of neck
strongly washed with "gallstone yellow," with normal postocular
stripe pattern indicated; fork figure on snout normal, buff colored,
black-edged; legs above and feet coarsely spotted with black; upper
side of tail like feet with lateral blackish lines converging backwards.

U.S.N.M. no. 95194, from the same locality, is also a male, and
slightly smaller. It shares the same structural characteristics, only
the tubercles on the anterior edge of carapace are slightly larger and
more distinct; the outline of the soft carapace is more oval than
rounded ovate; the xiphiplastral callosities are continuous, but the
distance between the hyohypoplastral callosities is a trifle wider, and
the entoplastral callosity somewhat smaller; the "sandpaper" effect
of the back is very much alike, but a double series of larger flat tuber-
cles on the mid-line of the 7th pleural is quite noticeable. The colora-
tion is also much the same, only the ground color of the carapace is
lighter, and the dark spots distinct, the chief difference being the inter-
rupted rows of the three marginal rings; on the other hand, the dusky
pattern elsewhere is finer; the pale edge of the leathery disk is slightly
paler than the rest of the disk.

For an easier appreciation of the more striking variations, they are
summarized in table 7. The differences are of various significance. As
certainly indication of sex is the greater development of the plastral
callosities of the males, besides the well-known difference in the
length of the tail, the "sandpaper" effect seems to be peculiar to a cer-
tain age (or size, or season?) of young males; the greater development
of the tubercles or "spines" may be correlated with sex or with age or
with both, remembering that the females are larger than the males;
the closing of the nuchal fontanelles of the carapace seems to take
place much earlier in the males than in the females; the slight differ-
ence in the outline of the leathery disk (as seen from above) is not due
to sex, and, at the stage of the specimen under discussion, not to age,
though the general rule in these turtles is that the young are more
circular than the old ones. None of the above characters seem to be
of specific or subspecific significance. On the other hand, the presence
of two or more concentric blackish rings inside the normal Amydan
dusky ring delimiting the pale rim of the leathery carapace is of diag-
nostic value in defining the subspecies, but unfortunately it has a

62

bulletin: museum of comparative zoology

tendency to be obscured or absorbed with age. Individual specimens
may occasionally be found without these additional rings among nor-
mal populations, but they are the "intermediates" which justify the
use of the trinominal nomenclature. On the other hand, it is significant
that among a very large number of specimens from northernmost
Montana to Louisiana and from Colorado to Lake Champlain, I have
not seen an authentic specimen of Amy da spinifer with two or more
submarginal rings.

Geographical distribution

Lower Mississippi tributaries in Louisiana and Mississippi and
Alabama.

Table 7
Variations in young adult A. aspera from Mississippi

Mississippi

Pearl

Black

Pearl

Pearl

River

River

River

River

95191

95192

95193

95194

Sex

9

9

&

o*

Soft carapace length mm.

286

210

180

158

width mm.

242

182

152

140

Bony carapace length mm.

173

119

111

99

width mm.

131

85

110

98

Plastral callosities

Small

Small

Large

Large

Callosity on entoplastron

0

0

+

+

Spines on edge of carapace

Large

Median

Small

Small

pointed

pointed

blunt

blunt

Tubercles on median line of front carapace

flap

+

+

0

0

Carapace skin "sandpapered"

0

0

+

+

Carapace fontanelles

+

+

0

0

Outline of carapace (from above) oval

+

+

" " " " " ovate

+

+

stejneger: American soft-shell turtles

63

Intergrades aspera-spinifera

It has been repeatedly asserted that the only differences by which
specimens of A. asper a can be recognized in the adult state are "the
very coarse and large tubercles of the front and hind part of the cara-
pace, which extend, behind, even over the bony shield, and are there
supported by prominent warts of the bony plates" (Agassiz, Contr.,
1, p. 406) and by the fact "that in younger specimens of Asp. asper
there are . . . two or three black lines separating the pale rim of the
posterior margin, whilst there is only one in Asp. spinifer." (Agassiz,
I.e.). As Agassiz himself observes, these lines fade away "pretty soon."

Table 8
Cranial measurements of A. asper a in millimeters

Basicranial length

mm.

Tip of snout to orbit
Horizontal diameter of orbit
Orbit of tympanic cavity
Longest diameter of temporal fossa
Interorbital width
Width of maxillary alveole
Length of internal choanae
Internal choanae to intermaxillary

foramen
Length of intermaxillary foramen
Length of mandibular symphysis
Width of mandibular alveole surface

C
o

G

A

to

oj

CO
00

o

63 63

16
12
19
11

5

5.5
10

7

9

a

oo

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16
10.5
18.5
12

6

5

9

6.5
6.2

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63

16
12
19
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5.5

5

9

6

6.5

9

4

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c

3

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1-5

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00
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37

10
8
9

6.5
3
3
6

6
5
6
2.5

cj

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a

CO

23 69

6

5.5

6

4

2

2

5

3
3
3
2

o3
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c3

3
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h-1

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19
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13

5

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7
11.5
5.5

m

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02
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cj
s-,

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13.8
10.1
15.3
9.9
4.4
4.4
8.1

5.9
5.5
6.4
3.5

64

bulletin: museum of comparative zoology

Specimens of intermediate ages are difficult to identify with our present
knowledge. Apparently the double or triple rings are not always a
constant character. I have before me, through the courtesy of Miss
Fannye A. Cook, a specimen collected about a mile or two southeast
of Brookhaven, Lincoln Co., Mississippi, in a small tributary of the
Bogue Chitto, Pearl River drainage. Its carapace measures only 43
mm. in length, hence it is quite young. It seems to be an aberrant
specimen of asper a, the anomaly being in the absence of the second
dark ring on the carapace margin. From its locality it ought to be
aspera, but such abnormal (or "incompleted" or "reversed") specimens
are known.

List of specimens in the U. S. National Museum

01084

Miss., Washington?

B. L. C. Wailes

01086

H 11

ii

Cotype of
Aspidenectes asper
Agassiz

7653-4

" Monticello

H. Tennison

012349

La., Lake Concordia

B. L. C. Wailes

1851 Cotype of
Aspidonectes asper
Agassiz

13250 9

" New Orleans

R. W. Shufeldt

1883

029266

u

S. W. Harvey

029310 ad. 9

" near New Orleans

U. S. Fish Comm.

66147 juv.

" Madisonville

?

May 29, 1886

68054 cf adol.

" Roberts

R. F. Shaw

79350-1 juv.

Miss., 1 mi. W. of Mel-

R. Kellogg and

vin

N. Boss

Oct. 1929

83996 9 juv.

Ala., 3 mi. S.E. of

Coatopa

C. E. Burt

July 1, 1931

95191 9 ad.

Miss., Pearl River

S. E. Brand

Aug. 1934

95192 9 adol.

" Big Black River

ii

u

95193 d" adol.

" Pearl River

ii

ii

95194 d" adol.

u ii

a

ii

100650 c? ad.

La., near Atchafalaya

C. E. Burt

June 17, 1935

100805 pull.

Miss., Enterprise

O. P. Hay

1881

115979 9 adol.

" near Guntoun

E. & W. H. Patten

Aug. 4, 1940

115981 juv.

" Chookatonkchia

Creek

H. L. Owens

June 30, 1941

stejneger: American soft-shell turtles 65

Amyda emoryi1 (Agassiz)
Plates 24-25

1849. Trionyx ferox ROEMER, Texas, p. 171, p. 459 (at New Braunfels,
Texas, in the Guadalupe and Comal Rivers).

1857. — Aspidonectes emoryi AGASSIZ, Contr. Nat. Hist. United States 1, p.
407; 2, pi. 6, figs. 4-5 (type-locality, Rio Grande River, near Browns-
ville, Texas; cotypes, U. S. Nat. Mus. No. 7855; Mus. Comp. Zool.,
Nos. 1909, 1913; Dr. Kennedy, collector; Williamson Co., Texas). —
COPE, Bull. U. S. Nat. Mus., No. 1, 1875, p. 51 (Texas); No. 17,
1880, p. 13 (Dallas; Helotes Creek, near San Antonio, Texas). —
TRUE, Bull. U. S. Nat. Mus., No. 24, 1883, p. 29 (Matamoras,
Mexico; Texas: Brownsville, Rio Grande, Rio Seco, Braunfels; Old
Fort Cobb, Oklahoma); Fish. Industr. United States, sect. 1, 1884,
p. 152.— BEYER, Proc. Louisiana Soc. Nat., 1897-1898, p. 43
(Louisiana).— STRECKER, Trans. Texas Acad. Sci., 4, pt. 2, no.
5, p. 6 (McLennan Co., Texas); Proc. Biol. Soc. Washington, 21,
March 21, 1908, p. 79 (Bazos, Bosque River, McLennan Co., Texas,
abundant); 23, 1910, p. 121 (Bullhide Creek, McLennan Co., Texas).
Trionyx emoryi BOULENGER, Cat. Chel. Brit. Mus., 1889, p. 258 —
STRAUCH, Mem. Acad. Sci. St. Petersbourg, ser. 7, 38, no. 2, 1890,
p. 117 (Texas).— DITMARS, Reptile Book, 1907, p. 78 (tributaries
of the Rio Grande in Texas and Mexico).— SIEBENROCK, Zool.
Jahrb., Suppl. 10, pt. 3, 1909, p. 603 (Colmisneil, Tyler Co., South
Bosque Riv., Texas); Verh. Zool. Bot. Ges. Wien, 73, 1923, p. 190.—
LINSDALE and GRESSITT, Copeia, 1937, no. 4, Dec. 31, pp. 222-
225, figs. 1-3 (Colorado River: Delta, Lower California; Clark Co.,
Nevada; Mohave Co., Arizona; California Lakes, Imperial Co.,
California; transpl. to Colorado Riv.?). — LINSDALE, Proc. Ameri-
can Acad. Arts Sci., 73, no. 8, May 1940, p. 255 (Nevada: Clark Co. :
Colorado River).
Amyda emoryi STEJNEGER and BARBOUR, Checklist North Amer.
Amph. Rept., ed. 1, 1917, p. 124 (Rivers of Texas, north into south-
ern Oklahoma and Arkansas); ed. 2, 1923, p. 140; ed. 3, 1933, p. 153.
-PRATT, Man. Vert. United States, 1923, p. 249.— ? HAY, Pan-
Amer. Geol., 39, March 1923, p. 119, pi. 9, figs. 2-4 (fossil, Brazos
River at Pittbridge, Texas).— SCHMIDT, Copeia, no. 131, June 30,
1924, p. 64 (Arizona; introduced).— STRECKER, Baylor Univ.
Bull., 27, no. 3, Sept. 1924, p. 47 (eastern Oklahoma); Contr. Baylor
Univ. Mus., no. 2, Jan. 15, 1926, p. 3 (Somervell Co., Texas); no. 3,
Feb. 15, 1926, p. 4 (Liberty Co., Texas); no. 6, June 15, 1926, p. 8

1 To Col. Wm. H. Emory, U.S.A., under whose command part of the type material was col-
lected, "I take great pleasure, therefore, in dedicating this species to that distinguished officer."

(Agassiz)

66 bulletin: museum of comparative zoology

(Cibolo Creek, Boerne, Tex.); no. 7, July 15, 1926, p. 7 (Cedar Creek,
Henderson Co., Tex.); no. 19, 1929, p. 15 (Trinity Riv., Ft. Worth,
Tex.); no. 23, June, 1931, p. 16 (Colorado Riv., Trevis Co., Tex.);
Baylor Bull., 38, no. 3, Aug. 1935, p. 23 (Texas: Cibolo Creek), p. 32
(Texas: Real Co.).— ORTENBURGER, Proc. Oklahoma Acad. Sci.,
6, pt. 1, 1926, p. 100 (LeFlore Co., Oklahoma).— STRECKER and
WILLIAMS, Contr. Baylor Univ. Mus., no. 12, Dec. 27, 1927, pp.

11, 15 (San Marcos and Blanco Rivers, Texas).— RUST, Blatt.
Aquar. Terrarienk., 45, 1934, p.—, sep., p. 12.— LITTLE and KEL-
LER, Copeia, 1937, no. 4, Dec. 31, pp. 216, 221 (Mesilla Valley,
Dona Ana Co., New Mexico). — GAIGE, Univ. Michigan Stud. Sci.,

12, 1937, p. 304 (Mexico, Tamaulipas, Rio Purificaci6n, N. of Ciudad
Victoria).— DELLINGER and BLACK, Occas. Pap. Univ. Arkansas
Mus., no. 1, June 1938, p. 46 (? Arkansas: Salina Riv. near Benton?
[probably Texas, U.S.N.M. no. 17823-L.S.]).— SMITH, Ann. Car-
negie Mus. Pittsburgh, 27, 1939, p. 312 (Mexico: Tamaulipas;
Nuevo Laredo).

1870. — Aspidonectes emyda "Agassiz," GRAY, Suppl. Cat. Shield Rept. Brit.

Mus., pt. 1, p. 95 (lapsus).
1870. — Aspidonectes georgii "Agassiz," GRAY, Suppl. Cat. Shield Rept. Brit.

Mus., pt. 1, p. 109 (lapsus).

1893. Platypeltis emoryii BAUER, Proc. Amer. Philos. Soc, 31, p. 220 (emen-
dation).— Amyda emoryii STRECKER, Copeia, no. 162, 1927, p. 9
(food habits); Contr. Baylor Univ. Mus., no. 15, July 10, 1928, p. 6
(Bosque Riv. near Valley Mills, Bosque Co., Texas); no. 16, Aug. 4,
1928, p. 21 (Texas: vernacular names).

Agassiz (p. 407), as character aiding in identifying this species,
calls attention to skin of the carapace being "dotted all over with
small whitish tubercles like grains of sand." This is not a specific
character, it seems to be a condition of the skin due to season or age, as
similarly "sandpaper" specimens are encountered in several of the
species.

Skull. The skulls of A. emoryi and A. svinifera are very much
alike. The snout in emoryi is slightly shorter and somewhat broader
anteriorly, the nasal cavity relatively shorter and the angle formed by
the anterior processes of the prefrontal bones more obtuse. The
alveolar surface of the maxillaries are somewhat wider. In these re-
spects the emoryi are even closer to A. fero.v.

As the differences in the skull between A. spinifera and emoryi are
very slight, many skulls can hardly be told apart. In the former the
choanae and the intermaxillary foramen average a trifle larger. The
orbit in A. spinifera is also placed slightly more backward on the

STEJNEGER: AMERICAN SOFT-SHELL TURTLES 67

average than in A. emoryi. As a consequence the snout appears a
mere trifle longer. However, it is difficult to understand how Boulen-
ger (Cat. Chel. Brit. Mus., 18S9, pp. 245-246) came to diagnose A.
emoryi as having "the snout (on the skull) obtuse, hardly as long as the
diameter of the orbit," and the other two {spinifera and/< rox) having
it "a little longer." In 26 skulls measured by me the horizontal diame-
ter of the orbits in ferox, spinifera, and emoryi averages 20.0, 19.6, and
19.1 mm respectively and the snout (as measured from the orbit)
24.9, 26.5, and 24.4 mm. In the six emoryi measured by me the snout
is 2.5, 3.0, 4.0, 5.5, 1.5, and 2.0 mm longer than the ort.it. The relative
dimensions of snout and orbit are therefore unavailable as a diagnostic
character. It should be noted, finally, that in emoryi the alveolar
surface of the maxilla of medium sized skulls is slightly wider than in
the other two, but too slightly and variably so to be of much help in
diagnosing.

Plastron. Referring to Siebenrock's remarks about the plastron of
.4. emoryi (Sitz. Ber. Akad. Wiss. Wien, Math. Nat. KL, 111, 1902, p.
830) it is well to note that the bones are essentially as in A. spinifera.
The entoplastral angle in adults seems to be more acute. The callosi-
ties appear to be more developed in a male (U.S.N.M. no. 26426);
they are almost as large as in the A. mutica figured by him (p. 823, fig.
5), but of course without trace of callosities on entoplastron or epi-
plastra. The anterior portion of the epiplastra is rather long.

Color of live specimens

Two adult males from Houston, Texas, are colored as follows. Xo.
94335 has the top of head and dorsal aspect of neck dark olive green,
becoming gradually more green on posterior half of neck. Carapace,
bony disk Van Dyke brown, bistre on the soft parts, with irregular
blackish brown anastomozing spots. Sides of neck almost citron yellow
fading into whitish on the middle line of the ventral aspect of the neck
which anteriorly changes into verditer blue on the throat, darkening
to almost indigo blue on mentum and outer half of lower jaw and
tympanic region, and extending a little below and backwards on side
of neck; the yellow of the side of the neck fades into a dull olive yel-
low with a few scattered, almost blackish spots.

Tubercles on anterior edge of soft carapace large (3-4 mm.), tri-
angular, spaced apart by the width of their bases.

Underside white, also feet, but fingers and webs pale (dull) sage
green with obscure dark marblings.

68 bulletin: museum of comparative zoology

Their measurements differ as follows:

d* ad. 94335 & ad. 94336

Soft carapace, long 342 mm. 354 mm.

286 " 288 "

Height of body 86 " 84 "

Tip of tail beyond carapace 40 32

In No. 94336 the colors are essentially as in the other except that
on head and neck the yellow is a little deeper and that there are a
large number of small blackish spots on -sides of head, even including
lower eyelid, lips, and base of proboscis, and definite blackish lines
running from eye obliquely backward to base of lower jaw which
is also outlined by similar lines; dusky obscure large (average 15
mm.) ocellar markings on under side of neck. Tubercles like 94335.

Two specimens collected by Dr. and Mrs. A. H. Wright, U. S. N. M.
nos. 94456-7 are colored as follows. In no. 94456 the neck and iris
chrome yellow more or less pale, iris with horizontal black lens.
Carapace above uniform wood brown. Marginal edge washed with
chromium green. Upper soft parts of head and legs pale chromium
green washed with wood brown on top and sides of head. Tail above
white, with posterior central part dull chromium green washed with
cadmium. Tubercles on carapace pale buff, minute, round, densely
scattered over the whole carapace, hand, and soft parts. Underside
white. Neck underneath very pale cadmium shading anteriorly into
pale verditer blue washed with pale cadmium (on throat). Dark
markings on throat strongly tinged with bluish. Fleshy lips pale
cadmium yellow. Pale cadmium yellow on fork of snout fading away
anteriorly. Triangle, base between eyes slightly angular.

No. 94457 has the same data, but a dark ring of minute blackish
specks surrounding tubercles. Underside of foot same color as upper
chromium green parts — spotted.

stejneger: American soft-shell turtles

69

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70 bulletin: museum of comparative zoology

Another male, U.S.N.M. 104240, from Pecos River, near Dryden,
Terrell Co., Texas. F. M. Setzler, coll. shows still further variation.

Anterior border of disk very obscurely tuberculated. Skin on cara-
pace shagreened like bony disk. Skin on flaps smooth, leathery: a
round smooth area over each. The fontanelle smooth. Part of flap
behind bony disk with a regular pattern of whitish "pimples" or
tubercles, those along the middorsal line in pairs. No "sandpaper"
effect. Palms and soles yellowish white, unspotted. Carapace (day
or two after death) : flap Isabella color (Ridgway, pi. Ill, fig. 23); bony
disk more tawny olive (fig. 17); neck and limbs above pale olive gray
like pickled unripe olives, ventral half of neck yellowish white with a
faint trace of a similarly colored band from lower edge of eye back-
wards along the neck. All upper soft parts with numerous black dots
more or less arranged in longitudinal series.

Geographical distribution

Rivers of Texas, north into southern Oklahoma and Arkansas, west
(introduced?) to southeastern California, the adjacent portion of
Lower California, and Clark County, Nevada; east to western Louisi-
ana; northern Mexico.

List of specimens in the U. S. National Museum

7176

Okla., Old Ft. Cobb

E. Palmer May

4, 1868

7614-20,

Mex., Matamoras,

7662-25

Tamaulipas

L. B. Couch

7628-33

Mex., Matamoras,

Tamaulipas

n

7635-6

?

?

7637-8 juv.

Tex., Rio Bravo

A. Schott

7640 juv.

" Brazos R.

G. C. Shumard

7641 juv.

" El Paso del Norte

Dr. Webb

7642 cf adol.

" Brownsville

?

7644 juv.

n a

?

7700 (029536)

ad.

" New Braunfels

?

7701 juv.

" Rio Grande del

Norte

G. Wurdemann

7747 juv.

" Rio Seco

Capt. Pope

stejneger: American soft-shell turtles

71

7854 juv.

7855 juv.

Tex.

, Brownsville

Dr. Kennedy

(Cotype of
Aspidonedes
emoryi
Agassiz)

8925

tt

tt

J. C. Merrill

May, 1877

10789 juv.

tt

San Antonio

C. W. Schuermann June 1879

17823 juv.

Ark.

, Benton

Jordan & Gilbert

19626-8

?

?

20846 9 ado!.

Tex.

, Ft. Hancock

E. A. M earns

Jan. 28, 1893

21408 juv.

?

f

26426 ad.

Tex.

, Ft. Clark

E. A. Mearns

26427 9 ad.

it

tt

a

26428-36

u

u

u

45545

. tt

Boquillas

V. Bailey

May 25, 1901

46073

It

mouth of Devil's

River

W. Lloyd

Sept, 26, 1890

46074 juv.

II

Eagle Pass

ti

Oct. 22, 1890

55601

(1

McLennan Co.

J. Hurter

1906

66147 juv.

La.,

Madisonville

f

May 29, 1886

71627-8 adol.

Ariz

., Phoenix

V. Housholder

May 1, 1926

78515-6 juv.

Tex.

, Coleto Creek

J. D. Mitchell

Oct., 1905

78517

Tex.

, Guadalupe R.

J. D. Mitchell

Aug., 1912

83690

tt

Christoval

C. E. Burt

Apr. 25, 1931

94335 d* ad.

tt

near Houston

A. C. Chandler

94336 c? ad.

tt

Houston

a

94456-7

tt

Orange

A. H. Wright

Apr. 17, 1934

95386 c5 juv.

tt

16 V2 mi. S.E. of

Caddo Lake

C. E. Burt

Apr. 1, 1934

95773 pull.

tt

Llano River, Kimble

Co.

S. Mulaik

Aug. 10, 1933

100089 cf adol.

La.,

near Napoleonsvilk

: C. E. Burt

1935

100090 juv.

a

(i

a

a

100380 d" adol.

a

Plaquemine

a

June 8, 1935

100419 <? adol.

a

Spanish Lake near

St,

Gabriel

it

June, 1935

103678 9

Tex.

, Boquillas

T. Smith

Aug. 6, 1937

104240 9 ad.

a

Pecos River near
Dryden

F. M. Setzler

72 bulletin: museum of comparative zoology

Amyda agassizii ' (Baur)
Plates 26-30

1857 .—Piatypeltis ferox AGASSIZ, Contr. Nat. Hist. United States, pt. 1, p.

401 (part) (not of Schneider).
Aspidonectes ferox COKER, North Carolina Geol. Surv., Bull. No. 14,

1906, p. 66 (South Carolina: Darlington Co., Society Hill, Peedee

River).— CORRINGTON, Copeia, No. 172, Nov. 15, 1929, p. 82

(Congaree Riv., between Columbia and New Brookland, Lexington

Co., South Carolina).
1888. Piatypeltis agassizii BAUR, Amer. Natural., 22, p. 1121 (type locality,

Georgia; type M.C.Z. no. 37172).
Pelodiscus agassizii BAUR, Proc. Amer. Philos. Soc, 31, July 1893,

p. 218.
Trionyx agassizii HAY, Indiana Geol. 17 Rep., 1892, p. 552; Batr.

Rept. Indiana, 1893, p. 144 (part, U.S.N.M. no. 8359).
1899. Aspidonectes agassizii JORDAN, Man. Vert. North Amer., ed. 8, p.

206 (part only: Ga.) (emendation).
1923. Trionyx spiniferus agassizii SIEBENROCK, Verh. Zool. Bot. Ver.

Wien, 73, Aug. 1923, p. 188 (part: Georgia).
1939.— Trionyx spiniferus POPE, Turtles U. S., Canada, pi. 45, figs. 98-99

(Broad River near Columbia, Richland Co., South Carolina) (not of

LeSueur).

The agassizii Group

In outward appearance Amyda agassizii differs very little from the
members of the spinifera group, but the skull distinguishes it at once
from all other American soft-shell turtles. In practically all the charac-
ters which differentiate it from them it agrees with the Asiatic mem-
bers of the group represented by the collective Amyda sinensis. A.
agassizii therefore may be treated as a member of a separate group,
thus more insistently emphasizing its isolated position and« insuring
the positive identification of specimens from its comparatively re-
stricted range in the United States.

Among Louis Agassiz's collection of colored drawings of turtles by
Burkhardt (mostly from life) are several painted in November and
December 1855, and inscribed as "Trionyx ferox, Ga. Dr. Daniel."
They are excellent pictures of Amyda agassizii, two of an "adult"
specimen (upper and lower surfaces) and one (upper side) of a young
specimen, but unfortunately they are without indication of size and
locality. They were sent to Agassiz by Dr. W. B. Daniel from Sa-
vannah, Ga. (Contr. Nat. Hist. U. S., 1, p. 401). The fact that they

1 Named for Louis Agassiz to indicate that the species was included by him in his account of
Piatypeltis ferox.

stejneger: American soft-shell turtles 73

came from the recorded type locality of T. ferox evidently influenced
Agassiz to regard these specimens as topo typical of ferox and hence
he recorded them as such.

The status of Baur's Platypeltis agassizii has never been fully ex-
plained. In 1888 (Amer. Natural., vol 22, p. 1121) the name appears
for the first time in the following sentence: " Platypeltis ferox of Agassiz
is not Testudo ferox Schneider, but a new species, which may be called
Platypeltis Agassizii." No further description or indication is given,
but evidently reference is intended to Agassiz's account of the species
in his Contribution to the Natural History of the United States, pt. 1 ,
pp. 401-403. A careful examination of Agassiz's text compared with
the material available to him at the time he wrote his account shows,
that his Platypeltis ferox is a mixture of two species due to the fact
that the ranges of both species overlap in Georgia and that the startling
color pattern of the very young specimens of the true Testudo ferox of
Schneider was unknown by him. Dr. Baur, in studying Agassiz's
material in the Museum of Comparative Zoology, designated the
specimen marked "Ferox Ga. No. 1" as the type of P. agassizii with
his own hand, and it must-be accepted as such.1 In 1893, after having
examined skulls and restudying the question of the generic relation-
ships of the Trionychids (Proc. Amer. Philos. Soc, vol. 31, July 1893,
p. 217) he came to the conclusion that his P. agassizii was not only
specifically but generically different from Schneider's P. ferox and re-
ferred it to Fitzinger's genus Pelodiscus with several Asiatic species.

In the meantime Dr. O. P. Hay, assuming that Agassiz's (op. cit.)
pi. 6, fig. 3, of his young Platypeltis ferox with the strongly marked two
black marginal rings represented the young P. agassizii (which has a
similar pattern), applied the name to the form which Agassiz had de-
scribed earlier as P. asper (Indiana Geol. 17 Rep., 1892, p. 552; Batr.
Rept. Indiana, 1893, Batr. Rept. Indiana, 1893, p. 144). This con-
fusion between A. agassizii and A. asper before their diagnostic fea-
tures were well understood was caused by the specimen U.S.N.M. No.
8359, a young A. agassizii, which was alluded to by Dr. Hay (op. cit.)—
though without giving the number — under A. agassizii as follows:
"This species belongs to the Southern States from South Carolina to
Texas. A single specimen has been forwarded to the National Museum
from Madison, Ind.". The specimen has the 2 black rings of the disk
characteristic of A. agassizii very plainly marked (pi. 30,) and as this
is also the normal pattern of the very young A. asper, Dr. Hay took

1 In a letter dated Feb. 13, 93 to Stejneger, he wrote: "The only Pelodiscus agassizii which I
have seen is at Cambridge Mus. (the type)."

74

bulletin: museum of comparative zoology

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stejneger: American soft-shell turtles

75

for granted that the agassizii and asper were synonymous, an error
followed by many subsequent authors.

The fact is that the locality "Madison, Ind." attributed to specimen
No. S359 in the original book of entry on June 25, 1875, is erroneous.
There is no record as to the donor or collector of the specimen and it
was entered on the book as part of an accumulation of miscellaneous
material. It is catalogued as Trionyx ferox and there is no original
label attached to the specimen, which only bears the tin-tag 8359, at
a time when the Museum had started a campaign to collect living
reptiles and amphibians to serve as models for the series of painted
casts in the exhibition series of the North American vertebrate fauna.
The tin-tagging and uncritical re-entry of many old specimens the
record of which was lost has been the source of many errors. (Note
reference to this specimen by Cahn in his Turtles of Illinois, 1937, p.
200, under Amyda ferox).

Geographical Distribution
Rivers of Georgia and South Carolina; North into southern North

Carolina.

List of specimens in U. S. National Museum

8359 juv.
8708 9 adol.
029034 9
30822 juv.
51981 (M.C.Z. 1598)

cf adol.
66859 9 adol.
71681 9 adol.

91282-3 ad. 9 ,

juv. d"
91310 9 adol.

91311-2 juv.

Ind., Madison ??
Ga., Milledgeville
Ga. ?

Ga., Baker Co.
)

Ga., Savannah
Ga., Augusta
S.C., Greenwood

T. H. Bean
R. Hessel
Brimley Bros.

W. B. Daniel
S. F. Hildebrand
Dr. Barrett,
A. L. Pickens

July, 1876
June 7, 1902

91491 juv.

91533

92521-3 J* adol.

92583 (M.C.Z.) adol. Ga.

92584 (M.C.Z. 1601)

9 Ga., Savannah

Ga., above Price Id.,

Savannah River E. H. Wood July, 193

S.C., 5 mi. W. of Plum

Branch, Savannah R. E. H. Wood
S.C., 5 mi. W. of Plum

Branch, Savannah R. " "

S.C., Batesburg L. Brodie

" Murray Lake C. E. Burt July 5, 1933

" Parks ville K. McNeill 1933

PLATES

PLATE 1

Stejnegeh — Soft-Shell Turtles

PLATE 1

Skull of Amy da (spinifera)

Fig. 1. View from above. Fig. 2. View from below. Fig. 3. View from left
side, including mandible. Fig. 4. View of mandible from above. Fig. 5. An-
terior plastral outlines of Amyda ferox U.S.N.M. 60496 (upper); Amyda
emoryi U.S.N.M. 94456 (middle); Amyda spinifera U.S.N.M. 101386 (lower).

alv. mb.

Alveolar surface of man-

pm.

premaxillary (intermaxil

dible

lary)

alv. mx.

Alveolar surface of max-

pof.

postfrontal (postorbital)

illa

prf.

prefrontal

art.

articular

pro.

prootic (otosphenoid)

boc.

basioccipital

pt.

pterygoid

bsp.

basisphenoid (para-

qj-

quadratojugal (para-

sphenoid)

quadratum)

cho.

choana

qu.

quadrate

cond.

occipital condyle

qu. art.

articulation of quadrat*

cor.

coronoid

with lower jaw

den.

dentary

s. ang.

supraangular

exoc.

exoccipital

soc.

supraoccipital

fr.

frontal

splen.

splenial

int. max.

for. intermaxillary foramen

sq.

squamosal

J"-

j«gal

sym.

symphysial

mx.

maxillary

temp, fossa

temporal fossa (inter

orb.

orbit

temporal foramen)

pa.

parietal

tymp. cav.

tympanic cavity

pal.

palatine

vom.

vomer

paoc.

paroccipital (opisthotic)

BULL. MUS COMP. ZOOL

-temp
fossa.

Stejneger. Soft-Shell Turtles. Plate 1.

pm.

~mt max for

alv. mx

temp,
fossa

Fig. 1

Fig. 2

pro
pot ju. Pil-MJ-

sopaoc

s.ar>

Fig. 4

Fig.

I

PLATE 2

Stejneger — Soft-Shell Turtles

PLATE 2
Amyda mutica 9, Fairport, Iowa. U.S.N.M. 53521.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 2.

■ « •

V <

, w
A ¥ A

<H-

3Sri

PLATE 3

Stejneger — Soft-Shell Turtles

PLATE 3

Upper view of Amyda mutica, U.S.N.M. 95186 from Medicine Lodge River,
near Lake City, Kansas. Also left side of head.

BULL MUS COMP ZOOL.

Stejneger. Soft-Shell Turtles. Plate 3.

PLATE 4

Stejneger — Soft-Shell Turtles

PLATE 4

Lower view of Amyda mutica, U.S.N.M. 95186 from Medicine Lodge River,
near Lake City, Kansas.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 4.

PLATE 5

Stejneger — Soft-Shell Turtles

PLATE 5

Skull of type of Amyda ferox in British Museum, from Georgia.

BULL MUS COMP. ZOOL.

Stejneqer. Soft-Shell Turtles. Plate 5.

PLATE 6

Stejneger — Soft-Shell Turtles

PLATE 6

Skulls of Amydaferox (U.S.N.M. 029475, 029459, 029464 and 029462 from
Kissimmee, Florida) showing extremes in the width of the alveolar surface and
in the outline of the maxilla.

BULL. MUS COMP ZOOL.

Stejneger. Sot-Shell Turtles. Plate 6.

PLATE 7

Stejneger — Soft-Shell Turtles

PLATE 7 .

Upper view of amyda ferox, U.S.N.M. 86828, Tamiami Trail near Birdon,
Florida.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 7.

PLATE 8

Stejneger — Soft-Shell Turtles

PLATE 8

Lower view of A my da- ferox, U.S.N.M. 86828, Tamiami Trail near Birdon,
Florida.

BULL. MUS. COMP. ZOOL.

Stejneger. Suet-Shell Turtles. Plate 8.

PLATE 9

Stejneger — Soft-Shell Turtles

PLATE 9

Amyda ferox, U.S.N. M. 86492 from Tamiami Trail about 15 miles from
Miami City, Florida.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 9.

\

mm

PLATE 10

Stejneger — Soft-Shell Turtles

PLATE 10

Amydaferox, U.S.N.M. 60496 from Auburndale, Florida.

BULL. MUS COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 10.

PLATE 11

Stejneger — Soft-Shell Turtles

PLATE 11

Upper view of Amyda spinifera, U.S.N.M. 101386 from Middle Fork, Hol-
ston River, Seven Mile Ford, Virginia.

BULL. MUS. COMP. ZOOL

Stejneger. Soft-Shell Turtles. Plate 11.

i

" t«

H

'*

I

I?

Bg? '

PLATE 12

Stejneger — Soft-Shell Turtles

PLATE 12

Lower view of Amyda spinifera, U.S.N.M. 101386 from Middle Fork, Hol-
ston River, Seven Mile Ford, Virginia.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 12.

PLATE 13

Stejneger — Soft-Shell Turtles

PLATE 13

Head of Amyda spinifera, U.S.N.M. 101386 from Middle Fork, Holston
River, Seven Mile Ford, Virginia.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 13.

PLATE 14

Stejneger — Soft-Shell Turtles

PLATE 14

Cotype of Amyda nuchalis, Mus. Comp. Zool. 1623 from Cumberland River,
Tennessee. Upper view.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 14

^M

zJs

PLATE 15

Stejneger — Soft-Shell Turtles

PLATE 15

Cotype of Amyda nuchalis, Mus. Comp. Zool. 1623 from Cumberland River,
Tennessee. Lower view.

BULL MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 15.

PLATE 16

Stejneqer — Soft-Shell Turtles

PLATE 16

Upper and lower views, and side of head of Amyda spinifera aspera. Cotype,
Mus. Comp. Zool. 1622 from Lake John, Florida.

BULL MUS COMP. ZOOL

Stejneger. So't-Shell Turtles. Plate 16.

PLATE 17

Stejneger — Soft-Shell Turtles

PLATE 17

Amyda spinifera aspera, U.S.N.M. 95191 from Canton, Mississippi. Upper
view.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 17.

PLATE 18

Stejnegeh — Soft-Shell Turtles

PLATE 18

Amyda spinifera aspera, U.S.N. M. 95191 from Canton, Mississippi. Lower
view.

BULL MUS COMP. ZOOL.

Stejneqer. Soft-Shell Turtles. Plate 18.

PLATE 19

Stejneger — Soft-Shell Turtles

PLATE 19

a. Amyda ferox juv., U.S.N.M. 61087 from Auburndale, Florida, b. Amy da
ferox juv., U.S.N.M. 84603 from Chesser's Island, Okefinokee Swamp, Georgia,
c. Amyda aspera, M.C.Z. 1608A from Mobile, Alabama.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 19.

I

r9H

PLATE 20

Stejneqer — Soft-Shell Turtles

PLATE. 20

Agassiz drawing of Amyda emoryi (upper view) which corresponds fairly
closely with the type (M.C.Z. 1910) collected at Brownsville, Texas by Col.
Emory.

BULL MUS. COMP. ZOOL

Stejneger. Soft-Shell Turtles. Plate 20.

^:$itik

.y i \ > -

V

V

I

/

/

\

PLATE 21

Stbjneger — Soft-Shell Turtles

PLATE 21

Agassiz drawing of Amyda emoryi (lower view) which corresponds fairly
closely with the type (M.C.Z. 1910) collected at Brownsville, Texas by Col.
Emory.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 21.

w

m

■&•>•■

PLATE 22

Stejneger — Soft-Shell Turtles

PLATE 22

Skull of Amyda emoryi, U.S.N.M. 78517, collected at Guadalupe River,
Victoria County, Texas by J. D. Mitchell.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 22.

fi

Aft

^

PLATE 23

Stejneger — Soft-Shell Turtles

PLATE 23

Upper view of Amyda emoryi, U.S.N.M. 94336 from near Houston, Texas.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 23.

PLATE 24

Stejneger — Soft-Shell Turtles

PLATE 24
Lower view of Amyda emoryi, U.S.N. M. 94336 from near Houston, Texas.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 24.

k

1

PLATE 25

Stejneger — Soft-Shell Turtles

PLATE 25
Head of Amyda emoryi, U.S.N.M. 94336 from near Houston, Texas.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 25.

^

\

JL-.)k

PLATE 26

Stejneger — Soft-Shell Turtles

PLATE 26

Amyda agassizii Juv., U.S.N.M. 8359, wrongly recorded from Madison,
Indiana. Upper view.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 26.

i'l •

XSJii-

^flt&'i °

o

o

Q 0

r

•

\

/

PLATE 27

Stejneger — Soft-Shell Turtles

PLATE 27

Upper view of Amyda agassizii, U.S.N.M. 92521 from Parksville, South
Carolina.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 27.

A

■ >

\

1 i

#

hi,

PLATE 28

Stejneqer— Soft-Shell Turtles

PLATE 28

Lower view of Amyda agassizii, U.S.N.M. 92521 from Parksville, South
Carolina.

BULL MUS. COMP ZOOL

Stejneger. Soft-Shell Turtles. Plate 28.

PLATE 29

Stejneger — Soft-Shell Turtles

PLATE 29
Head of Amyda agassizii, U.S.N.M. 92521 from Parksville, South Carolina.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 29.

PLATE 30

Stbjneger — Soft-Shell Turtles

PLATE 30

Skull of Amyda agassizii, Philadelphia Acad. Nat. Sci. 106 ( = 371).

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 30.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 2

CONTRIBUTION TO THE ORNITHOLOGY OF THE
HAWAIIAN ISLANDS

By E. H. Bryan, Jr. and J. C. Greenway, Jr.

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

May, 1944

PUBLICATIONS
OF THE .

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1
have appeared and of the Memoirs, Vol. I to LVI.

These publications are issued in numbers at irregular intervals.
Each number of the Bulletin and of the Memoirs is sold separately.
A price list of the publications of the Museum will be sent upon
application to the Director of the Museum of Comparative
Zoology, Cambridge, Massachusetts.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 2

CONTRIBUTION TO THE ORNITHOLOGY OF THE
HAWAIIAN ISLANDS

By E. H. Bryan, Jr. and J. C. Greenway, Jr.

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

May, 1944

No. 2 — Contribution to the Ornithology of the Hawaiian Islands '
By E. H. Bryan, Jr. and J. C. Greenway, Jr.

FOREWORD

For over a year prior to the entrance of the United States into the
war, Mr. E. H. Bryan, Jr., Curator of Collections at the B. P. Bishop
Museum, and Mr. J. C. Greenway, Associate Curator of Birds at the
Museum of Comparative Zoology, collaborated in the preparation of
a Check-List of Birds of the Hawaiian Islands. The finished draft of
the list was completed early in 1941, but, due to the fact that the
senior author was called to active duty as an army reserve officer,
his final comments were not received for inclusion until recently. In
the meantime the junior author was commissioned in the Navy; con-
sequently the final preparation of the Check-List for the printer
devolved upon the undersigned. No introduction had been prepared,
but I was so fortunate as to discover in the Bryan-Greenway corres-
pondence an outline of the history of Hawaiian ornithology which,
though not prepared with such a purpose in mind, nevertheless appears
suitable as such. A brief abstract has previously appeared in the
Proceedings Hawaiian Academy of Science for 1935 (Special Publica-
tion no. 26, Bernice P. Bishop Museum) p. 5-6.

Shortly before being ordered to active duty Greenway had incor-
porated his studies of the Hawaiian drepanids in a rearrangement of
the genera of this most interesting family; with the publication of this
Check-List, the opportunity to publish this account seems opportune.

Ornithologists all wish for the safe and speedy return to their chosen
fields of Capt. Edwin H. Bryan, Jr. A. U.S. and Lieut. James C.
Greenway, Jr. U.S.N.R.

James L. Peters

Cambridge, Mass.
23 June 1943

1 Published with the aid of a special gift from Mr. G. R. Agassiz.

80 bulletin: museum of comparative zoology

HAWAIIAN BIRDS1

By E. H. Bryan, Jr.

Curator of Collections, B. P. Bishop Museum2

In 1778, when the third voyage of Captain Cook made contact
between Hawaii and the so-called civilized world, there lived in Hawaii
about 100 different kinds of birds. Most of these were well known to
the Hawaiians, each kind being called by a native name. A few species
were eaten; the feathers of several kinds were used to decorate the
kahili or royal standards of the kings and high chiefs ; feathers of the
mamo, oo, iiwi, and to a less extent those of the apapane and o-u,
were tied in decorative patterns upon fine mesh network of olona
fiber to form royal uniforms and ceremonial robes; and in one way or
another many found a place in native life and lore.

Although Captain Cook did not live to see his native land again,
and although there was no trained naturalist with this third voyage,
his companions seem to have procured specimens of about sixteen
species of Hawaiian birds, and to have carried them safely back to
ornithologists in Europe. These first native Hawaiian species to be
made known to science included the iiwi, mamo, apapane, amakihi,
akepa, akialoa, o-u, oo, thrush, elepaio, rail, crow and migrant golden
plover. A specimen of the oo-aa, was also taken, but this was not
recognized as distinct from the oo, until 1855.

The iiwi has the distinction of being the first native Hawaiian
species of bird to be technically described. Barthold Lohmann, who
sailed with Cook's last expedition, brought specimens of it to Cassel,
Germany, where it was described by Georg Forster, in 1780 as Certkia
coccinea. During the next four years John Latham gave brief descrip-
tions and popular names to nine other species in his General Synopsis
of Birds. In 1788, Gmelin gave scientific names to eleven, including
these, largely on the basis of Latham's notes. Six species can be recog-
nized from the names and descriptions given by James King in his
account of Cook's voyage.

Thus did Hawaiian ornithology get away to a good start, only to
sink into quiescent repose until toward the end of the 19th century.
This came about partly through neglect, but to quite a large extent
because of a series of unfortunate circumstances.

1 Address of the retiring president, Hawaiian Academy of Science, May 18, 1935.

2 Published by permission of the Director, B. P. Bishop Museum.

BRYAN AND GREE.WVAY: HAWAIIAN' BIRDS 81

In the first place, the types of these early species, which had been
placed in the British Museum, by some mischance became lost or
destroyed. This led to considerable confusion in later technical sum-
maries, catalogs and lists, because of the lack of authentic specimens
to which to refer. In the second place, the naturalists with Vancouver,
Kotzebue, and other early expeditions which touched Hawaii, seem
to have either quite neglected the interesting bird life, or else to have
made collections and observations which resulted in no publications
for the advancement of science.

In 1825, H. M. S. 'Blonde', under command of George Anson,
seventh Lord Byron, carried back to Hawaii the bodies of Kame-
hameha II and his queen who had died in England. On board was
Chaplain Richard R. Bloxam and his brother Andrew, an enthusiastic
young naturalist. Ornithologists of the day hoped that young Andrew
Bloxam might get some of the curious Hawaiian birds and produce
an interesting publication about them. He apparently got very little
official encouragement, in spite of which he obtained specimens of nine
species of land birds on Oahu, including the now extinct thrush. He
worked hard over his specimens, and placed them, all properly labelled,
at the disposal of the Lords of the Admiralty. The scientific "Appendix
to the Voyage of the Blonde", published in 1826, is said to have been
edited bv a woman who had onlv a few of Bloxam's notes to guide her.
This, combined with some poor judgment on the part of the ornitho-
logical gentlemen of the British Museum, who identified the specimens,
made the results, as Professor Alfred Newton put it1 "a disgrace to
all concerned, since, so far from advancing the knowledge of the
subject, it introduced so much confusion as to mislead many sub-
sequent writers," especially in the absence of the specimens, which
disappeared not long after.

A few years later another good opportunity to advance Hawaiian
ornithology was missed. Dr. J. K. Townsend, the American traveller,
and the well-known naturalist, Thomas Nuttall, made a trip together
to Hawaii. Arriving in January, 1836, they spent three months col-
lecting on Oahu and Kauai. Townsend, returning at the end of the
year, found the Prussian naturalist, Herr Deppe, at Honolulu, and
with him spent a few months in the pursuit of natural history, leaving
Hawaii in March, 1837. A few of Deppe's birds were described by
Lichtenstein, and most of Townsend's specimens have been carefully
preserved, the bulk of them in the Academy of Natural Sciences,

1 Alfred Newton, Ornithology of the Sandwich Islands, Nature, 45, p. 466, 1892.

82 bulletin: museum of comparative zoology

Philadelphia. Had Townsend only published a list of his species,
and had both men but made a scientific record of their observations,
the knowledge of Hawaiian bird life would have been greatly advanced.

In the course of a six months' stay on Hawaii in 1840, the enthus-
iastic collectors, Peale and Pickering, of the United States Exploring
Expedition, obtained a large collection of birds. Most of these speci-
mens were lost in the shipwreck of the "Peacock," one of the ships of
Commodore Wilkes' squadron. Still another misfortune occurred in
1848. Peale's report on the birds of the group was just off the press,
and only a few copies had been distributed, when the entire stock was
destroyed by a fire. It is the opinion of some ornithologists that John
Cassin's new edition of this report, published ten years later, was no
improvement on Peale's original work.

In 1852 Dr. Hartlaub wrote a review of Peale's lost work, and later
he compiled the first list of Hawaiian birds, published in 1854. He
listed but 30 species of birds (of which five have not been accepted).
Of the remaining 25, only 16 are land birds, and only 14 are perching
birds.

It remained for our own Sanford B. Dole to produce an extensive
list of the Hawaiian birds. This was published first in 1869 in the
Proceedings of the Boston Society of Natural History, and later,
with additions and corrections, in the Hawaiian Annual for 1879.
Judge Dole listed 53 species, of which four were described as new.

Although in the vicinity of Hawaii from July 27 to August 19, 1875,
the scientific party of H. M. S. 'Challenger' collected only 24 bird
skins, of which only one species was new to science, the Hawaiian
duck, described in 1878 by P. L. Sclater, as Anas wyviUiana.

Revival of Interest

Considerable credit for a revival of interest in Hawaiian birds
should go to Professor Alfred Newton, of Cambridge University,
from whose interesting account of early Hawaiian ornithology some
of the foregoing has been condensed. Because of Dr. Newton's enthu-
siasm, two young Cambridge naturalists undertook collecting which
led to magnificent contributions to Hawaiian ornithology.

The first of these was Scott B. Wilson who went to Hawaii and
collected birds at Professor Newton's request. Leaving Liverpool on
February 24, 1887, he arrived in Honolulu on April 8, having made
a brief visit at Washington, D. C, with Dr. Leonhard Stejneger,
who had done considerable work on Hawaiian birds, especially those

BRYAN* AND GKKKNWAY: HAWAIIAN BIRDS 83

sent to him at the U. S. National Museum by Valdamar Knudsen of
Kauai. Wilson stayed on the islands until the close of 1888, taking
back with him to England a large and valuable collection of bird
specimens, rich in new species. This collection formed the basis for
the beautifully illustrated "Aves Hawaiienses," issued in sections
from 1890 to 1899. Wilson again visited Hawaii, in 1896, without
adding much to scientific knowledge. Besides collecting skins, he
carefully preserved whole specimens of the birds in alcohol. Upon
these specimens Dr. Hans Gadow based anatomical studies which
led to some new and startling conclusions as to the relationships of
many Hawaiian species.

Meanwhile, the Hon. Walter Rothschild, the other naturalist
interested by Professor Newton at Cambridge, sent his collector,
Henry Palmer, to the islands to procure specimens and data. Arriving
in December, 1890, Palmer collected on nearly all the main islands
of the group in company with George C. Munro, and also made a trip
to Laysan and Midway Islands, stopping at, or at least sighting,
most of the small rocky islets and reefs en route. By August, 1893,
when he left Honolulu, Palmer had collected 1832 bird specimens,
including all but seven of the species previously known, and in addi-
tion fifteen which Rothschild described as newT to science.

Rothschild produced the elaborate "Avifauna of Laysan and the
neighbouring islands with a complete history to date of the birds of
the Hawaiian possessions," published in London in three parts, which
appeared August, 1893, November 1893, and December 1900. These
contain over 300 large, well printed, folio pages, and numerous photo-
graphs and artistic colored plates drawn by Keulemans and Frohawk.
The work enumerates 116 species of birds, giving valuable notes on
food, habits, eggs, nests, and distribution, as w7ell as descriptions.

In 1898 Dr. H. Schauinsland spent three months on Laysan, and
in his fascinating little book, printed in Bremen in 1899, he lists the
birds which he found there. In 1900 he also furnished notes on a few
birds of Molokai.

In 1892 R. C. L. Perkins came to Hawaii under the joint auspices
of the Royal Society of London and the British Association for the
Advancement of Science, writh financial assistance from Bernice P.
Bishop Museum. For a decade he made extensive collections of
Hawaiian birds, as well as of insects and other land animals. His
section of the 'Fauna Hawaiiensis' on the birds was published Novem-
ber 1903. It gives an interesting and valuable summary of their dis-
tribution and relationships, especially of the native perching birds.

84 bulletin: museum of comparative zoology

In 1901, there appeared a "Key to the birds of the Hawaiian group"
by William Alanson Bryan, followed by a number of other lists and
notes on the birds of different islands of the group, written by him
and by Alvin Seale, published by Bernice P. Bishop Museum. Also,
in 1902-3, there was printed "A complete list of the birds of the
Hawaiian possessions, with notes on their habits," by H. W. Henshaw.
All of these publications have helped to complete and bring up to date
our knowledge of Hawaiian ornithology.

By 1930 the large number of birds which had been imported from
many parts of the world prompted the production of a report by
E. L. Caum on the "Exotic birds of Hawaii," published by B. P.
Bishop Museum in 1933.

Recent articles on Hawaiian birds include: — a report on sea bird
conditions on Laysan in 1911 by Homer Dill and W. A. Bryan; four
chapters in W. A. Bryan's Natural History of Hawaii; a popular
account of bird life in the Hawaiian Islands Bird Reservation by
Alexander Wetmore; four chapters in Hawaiian Nature Notes by E. H.
Bryan, Jr. ; and various articles in journals and the daily press.

Summary of Hawaiian Birds

With this brief historical background, let us consider the different
groups of birds which are found in Hawaii, and some of the interesting
facts concerning them. The list of the birds reported as having been
found in Hawaii, outside of captivity, both native and introduced,
now numbers 232 species (see table 1.) Of these, 77 species might be
considered as endemic; 18 are sea birds which nest regularly in the
archipelago, but which have a wider range; 1 species, the night heron,
is indigenous, but is found elsewhere; 8 are regular winter migrants;
34 have been recorded from time to time, and might be classed as
occasional migrants or chance arrivals; and 94 are immigrant exotic
species. Of these introduced species, 53 are probably established, and
41 are probably not established, although the exact status of several
is difficult to determine. So little field work has been done during the
past 30 years that it is not wise to state what species are really extinct.

Without reference to taxonomic classification, most of the Hawaiian
birds may be divided into three fairly well defined groups: (1) the
birds of the mountain forests; (2) the sea birds and migrants; and
(3) the introduced species. A few lowland species, such as the two
birds of prey, the black-crowned night heron, the native duck and

BRYAN AND GRKENWAY: HAWAIIAN BIRDS

85

goose, the extinct rail, the mud hens, the stilt, and the peculiar land
birds of Laysan, now all but extinct on that island, which do not fall
into these three general groups, might be given separate consideration.

Table 1. NUMERICAL SUMMARY OF HAWAIIAN BIRDS

Family

Endemic

Indigenous

& wide-
ranging sea

Regular

Migrants

< •(■(visional

& chance

arrivals

Tnt
Estab.

roduced

Not Estab

Diomedeidae

2

Procellariidae

3

5

Phaethontidae

1

1

Sulidae

3

Phalacrocoracidae

1

1

Fregatidae
Ardeidae

1
1

1

Threskiornithidae

1

Phoenicopteridae
Anatidae

3

3

10

1

1

2

Aceipitridae
Falconidae

1

2

1

Cracidae

3

Tetraonidae

1

1

Phasianidae

16?

3

Numididae

1

Meleagrididae
Turnicidae

1
1

Gruidae

1 ?

Rallidae

5

1

1

Charadriidae

5

6

Recurvirostridae

1

Phalaropodidae
Laridae

6

2
9

2

Columbidae

7

11

Psittacidae

2

5

Strigidae
Alcedinidae

1

1

Alaudidae

2

1

Corvidae

1

Paridae

1

Timaliidae

3

1

Mimidae

1 ?

Turdidae

6

2

2

Sylviidae

2

1

Museicapidae

3

2

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Prionopidae

Sturnidae

Zosteropidae

Drepanididae

45

Meliphagidae

5

Ploceidae

Icteridae

Fringillidae

77

19 8

Total Native

104

Total Introduced

*

Grand Total

232.

The Sea Birds

1 ?

1

1

2

1

1

1

5

3

34 53 41

94

Concerning the sea birds, found throughout the Hawaiian group,
little need be said here. Most of them are species widespread in the
Pacific, and although scientists argue over their scientific names they
are fairly easy to recognize. They include: the two large albatross,
the thieving frigate, three species of homely boobies, five kinds of
noisy terns, the burrowing shearwaters and petrels, and the graceful
white-and-red-tailed tropic birds. Most of them stay well away from
man's habitations. Not that they are afraid of him. Rather, man
and sea birds are rivals for a large and steady supply of fresh fish,
without which an extensive bird colony could not exist.

President Theodore Roosevelt and other thoughtful citizens have
helped to protect our sea birds by establishing for them the Hawaiian
Islands Bird Reservation on islands and reefs to the northwest of
Kauai. Here they are not only kept from molestation, but also from
time to time efforts are made to keep their rather poor environment
from becoming any worse, through occasional visits to kill off rabbits
and plant more vegetation.

Migratory Birds

Each year there come to Hawaii large numbers of tourists, concern-
ing whom no record is kept by our efficient Tourist Bureau. They
spend no money here, but they do a very considerable amount of good.
They are the migratory birds. Arriving from a more than 2,000 mile
flight, thin and hungry, they alight on our beaches, fields and pastures
and make short work of great numbers of caterpillars, grasshoppers

BRYAN AND GBEENWA1 : HAWAIIAN BIRDS 87

and other insects, much to the benefit of agriculture. They arrive and
depart with great regularity, coming in the late summer or early fall
and leaving again in the spring, to return to nest in their cold northern
home. Included among these gentle and helpful visitors, which should
be given a better welcome than the hunters' gun, are the golden
plover, the turnstone„ wandering tatler, sanderling, curlew, various
wild ducks, and several others.

The native birds of Hawaii's lowlands, marshes, ponds, valleys,
and grassy slopes are becoming very rare. The flightless rail is gone;
the duck and goose are making a last valiant stand against extermina-
tion; the mud hens, stilts, and night heron seem to be holding their
own. One still sees Pueo, the native owl, winging his low flight above
the tops of guava bushes and across grassy hillsides at sunset, in search
of rats and lizards. But Io, the native hawk, is scarce in Hawaii, due
to the mistaken idea that it was an enemy to the poultrymen.

Birds of the Mountain Forests

The birds of the mountain forests are all native perching birds and
include 50 to 55 endemic species of that order. Twenty-three of the
twenty-four genera in this group are also endemic, only Corpus being
found elsewhere in the world. These undoubtedly make up the most
interesting part of the Hawaiian bird fauna. Whence did they come?
What were their ancestral relationships? Why are there so many
genera and species, even one whole family, found nowhere else in the
world? Why are they so rapidly becoming extinct? These are ques-
tions which interest not only the ornithologist, but also the zoologist,
the biologist, the conservation expert, and the student of evolution.
I can only try to hint at the answers to some of these questions.

These native perching birds, found in the forests of the main islands
of the group, include: the crow, found only in the Kona and Kau
districts of Hawaii; six different species of thrushes, five of which
(found on five different islands) are closely related, and one, on Kauai,
somewhat different in form, food, and habits; three closely related
species of elepaio or flycatcher, on three different islands; four species
of oo, on four different islands, and one other very distinct honey-eater
of Hawaii; and between 40 and 45 different species of native "honey
creepers," belonging to a family found nowhere else in the world.
I give this range of number for the drepanids because various authori-
ties differ in their concepts of specific differences.

88 bulletin: museum of comparative zoology

Native Hawaiian Honey Creepers

The drepanids themselves are worthy of extensive study. There
are, let us say, forty species, representing eighteen genera. These
have very diverse appearance; some are black and golden, some red
and black, some green and yellow; some have shprt bills for picking up
insects, some long, slender bills for sipping nectar from the base of
the long-tubed blossoms of the native lobelias, and some have heavy,
almost finch-like or parrot-like beaks for cracking seeds. Only three
species, the iiwi, apapane, and o-u, are found on more than two islands.
Yet it is believed, that all are the descendents of a common ancestor,
or at most two ancestral stocks.

The careful anatomical studies of Dr. Hans Gadow suggest that
their nearest relatives are the Neotropical and Central American
Coerebidae. Peter P. Suskin (International Ornith. Kongress 1926,
Verhandl., VI, pp. 375-381, 1929) suggests descent from Fringillidae
of the Malaysian region. We can, perhaps, imagine that many thou-
sands of years ago there reached Hawaii from far away tropical America
or southeastern Asia a little group of these birds. A pair of them,
at least, survived, and establishing themselves, became the common
ancestors of our interesting native family. We can but suppose that
their descendents, becoming isolated on different islands of the group,
with different kinds of environment and different sources of food
supply, developed into the diverse species which we find today, or
rather, in many cases, — yesterday, for a number are very rare, even
extinct.

Origin of Hawaiian Bird Groups

The same explanation as to origin might be offered for other groups
of native perching birds. For example, one ancestral kind of immigrant
flycatcher, arriving not so many centuries ago, has given rise to the
three species of elepaio, one each on Hawaii, Oahu, and Kauai, which
look so very much alike that they might better be considered only
geographic races, rather than different species. One immigrant an-
cestral form may have been the progenitor of the four kinds of oo,
the Oahu species of which has not been seen alive since 1837; another
was the ancestor of Chaetoptila, a Hawaii honey-eater, now probably
extinct. The thrushes descended from two ancestral stocks; one
ancestor giving rise to one species each of A-Maui on Kauai, Oahu,
Molokai, Lanai, and Hawaii, and the little, weevil-eating Puaichi

BRYAN AND GREENWAY: HAWAIIAN BIRDS 89

having descended from another. Thus, all of the native Hawaiian
perching birds might have developed, as Dr. Perkins has pointed out1
"from at least six and not more than seven successfully immigrant
species."

It would make a fascinating study, by tracing out the distribution
of the nearest relatives of these various groups of birds, to try to
determine whence they came; and, by studying their degree of develop-
ment and evolution, as well as their local distribution, to attempt to
guess how long they have been in Hawaii. One interesting point
is the fact that, although the distances between the Hawaiian Islands
are not very great, and although the native forest birds have fairly
good powers of flight, their distribution is so definitely limited and
restricted.

Why Native Species are Becoming Extinct

One hears various explanations as to why the native species of
Hawaiian birds are so rapidly becoming rare or extinct. We are told
by those who have not made a study of the subject that they were
killed off by the native feather-gatherers or chased away by the minah
birds, or their eggs destroyed by the mongoose. These, in certain
instances, may have been contributing causes, but they have certainly
not been the basic reason for the changes which have come over the
native forest life. The main reason may be stated in these few words:
Nature established a balance in Hawaii, and Nature's balance has
been upset. What is this "Balanee in Nature" and what has caused
its upset?

Long ago, before the coming of man with his ships, new plants and
animals arrived from foreign parts and established themselves in
Hawaii only at long, infrequent intervals of time. This was because
of Hawaii's extreme isolation — more than two thousand miles from
the regions from which the ancestors of most of the native species
came. The kinds which managed to arrive and gain a foothold had to
adjust themselves to their new home. If they were in harmony with
their new environment they survived; if they were not in harmony,
they either became adjusted or modified until they were, or else they
perished. There have been many types of environment in HawTaii:
bare lava flows, hot dry deserts, fertile valleys, marshy lowlands,
grassy hills, wet forests, summit bogs, bare, snow-capped peaks.

1 R. C. L. Perkins, Fauna Hawaiiensis, vol. I, pt. iv, p. 369, 1903.

90 bulletin: museum of comparative zoology

Each offered a habitat for a different type of life, or a reason for
modification or adjustment. In each different environment a balance
was established, with the ironclad policy, "fit in or perish." If a
species became over-abundant it was brought back to a proper numer-
ical balance either through lack of food supply, or by the activity of
its enemies, which took advantage of its abundance to multiply them-
selves, only in turn to be starved or eaten back to normal.

Upset of Nature's Balance

Then into this nicely balanced island world came man, with his
foreign plants and animals, his thoughtless or ruthless waste, his
clearing and burning. The nice balance of Nature was upset. Cattle,
sheep, pigs, and goats ran wild through the forests, so damaging the
undergrowth that it no longer protected the roots of the growing trees.
Rain eroded away the soil; seeds could no longer germinate; and both
forest trees and undergrowth died off. Their places were quickly
taken by foreign plants which no longer furnished food or protection
to native insects and birds. Whole associated groups, interdependent
upon each other, were disassociated. Foreign birds were imported,
bringing with them diseases against which the specialized native
birds had no immunity. With food and shelter destroyed, and danger
threatening them in the lowlands, the native birds either sought new
homes in the depths of the forests, where they in turn upset nicely
adjusted Nature, or else they became extinct.

Foreign Birds

And now, because native birds are scarce, the cry is for more and
yet more foreign birds. Some kinds have been introduced to satisfy
the desire of the hunter; pheasants from the mountains of Mongolia
or Java; grouse and quail from northwestern America, China, Japan
and Australia; partridges from the slopes of the Himalayas; doves
and pigeons from Panama, India, China, Australia, and the Malay
Isles. Some, like the skylark and cardinal, have been brought because
of sentiment or to satisfy esthetic taste. Some have been introduced
for a more useful purpose, such as the horn fly-eating willie wag-tail,
the meadow lark, even the much maligned, but very useful minah.
A few accidental escapes we could well get along without.

BRYAN' AND GREENWAY: HAWAIIAN BIRDS 91

Fortunately for Hawaii, our Territory's scientific- advisers have
advocated a policy of rigid protection for our native and useful bird
life, and a strong stand against the promiscuous importation of foreign
birds and animals. This need not include birds which are to be kept
in a proper aviary, carefully safeguarded against their escape. I can
see much that is good in recent plans for extensive breeding and
exhibition of showy and interesting birds in Honolulu, so long as they
are not allowed to go wild. But I cannot approve of wholesale intro-
duction and liberation, without adequate study by experts as to the
consequences.

You may say, now that Nature's balance has been so badly upset,
and Hawaii's avian heritage is passing out of the picture, what harm
can a little more upset do ! But the birds may not be the only part of
our natural history to be effected by such introductions. Forestry,
agriculture, and all branches of animal life are so interrelated that
one cannot tell, without considerable study, what far reaching con-
sequences may result from one foolish introduction. We have effective
quarantine laws to protect Hawaii against insect pests and plant
diseases, which we all admit help tremendously to safeguard our
island paradise. Let us try to protect what bird life we have left, and
also try to avoid the risk of further upset of Nature's balance, just
for the sake of adding a flash of color or a chirp of song.

92 bulletin: museum of comparative zoology

' CHECK-LIST OF THE BIRDS
OF THE HAWAIIAN ISLANDS

By E. H. Bryan, Jr. and J. C. Greenway, Jr.

Part I. NATIVE BIRDS

Order PROCELLARIIFORMES

Family DIOMEDEIDAE— Albatrosses

Genus Diomedea — Albatrosses

Diomedea Linnaeus, Syst. Naturae, ed. 10, 1, 1758, p. 132. Type, by subse-
quent designation, D. exulans Linn.

Diomedea nigripes Audubon

Diomedea nigripes Audubon, Ornith. Biog., 5, 1839, p. 327. (Pacific Ocean,

lat. 30° 44' N., long. 146° W.)
Black-footed Albatross, Brown Gooney
Phoebastria nigripes reischekia Mathews, Bull. British Ornith. Club, 51, 1930,

p. 29. (New Zealand.)
Diomedea chinensis Temm. in Rothschild, Avifauna of Laysan, etc., p. 55

(corrected p. 292).
Diomedea albatrus (chinensis) in Wilson and Evans, Aves Hawaiiensis, p. xxv.

Range. North Pacific, breeding on islands northwest of Hawaii;
Marshall Islands and Bonin Islands.

Recorded in Hawaiian Group from Midway (breeding), Pearl and
Hermes Reef (sight), Lisianski, Laysan (breeding), French Frigate
Shoal (breeding) (sight), Necker (sight), "Bird Rock" (Nihoa) (sight),
Kaula (breeding), Lehua (sight) (Caum). Many sight records at sea.
"This is the Gooney that follows in the wrake of ships from San Fran-
cisco to Honolulu and Hilo" (Henshaw).

Nests behind the beaches on the periphery of the islands. A hybrid,
D. nigripes x D. immutabilis, from Midway is in the Bishop Museum.

Diomedea immutabilis Rothschild

Diomeda immutabilis Rothschild, Bull. British Ornith. Club, 1, 1893, p. xlviii.

(Laysan Island.)
Gooney, White Albatross, Laysan Island Albatross

Range. Central and North Pacific.

Breeds on Laysan and Midway Islands. Nests in interior of the
islands. To be seen at sea from islands off the coast Lower California
to the Bonin Islands.

BRYAN AND GREENWAY : HAWAIIAN BIRDS 93

Recorded in the Hawaiian Group from Midway (breeding), Laysan
(breeding), Lisianski (breeding), Necker Island (breeding), Pearl and
Hermes Reef (breeding), Gardner Island, "Bird Island" (Nihoa)
(breeding). "Apparently common throughout Hawaiian main group,
though it is not known to breed on any of them" (Henshaw).

Family PROCELLARIIDAE

Genus Puffinus — Shearwaters

Puffinus Brisson, Ornithologie, 1, 1760, p. 56; 6, p. 130. Type, by tautonomy,
Puffinus Brisson = ProceUaria puffinus Briinnich.

Puffinus pacificus cuneatus Salvin

Puffinus cuneatus Salvin, Ibis, 1888, p. 353. (Krusenstern Island.)

Wedge Tailed Shearwater, Uau Kane, Uwau

Puffinus pacificus laysani Mathews, Birds of Australia, vol. 2, 1912, p. 83.

(Laysan Island.)
Puffinus knudseni Stejneger, Proceed. U. S. Nat. Museum, 11, 1888, p. 93.

(Kauai.)

Range. From about 140° west latitude (two days from San Fran-
cisco) to the Bonin Islands and south to the Equator.

Recorded in the Hawaiian Group from Midway, Pearl and Hermes
Reef (sight, Galstoff), Lisianski, Laysan (breeding), French Frigate
Shoal, Xecker (breeding), "Bird Island" (Nihoa), Kaula, Lehua,
Kauai, Oahu (breeding on Mokumanu, Manana and Popoia Islets)
(Munro).

Puffinus nativitatis Streets

Puffinus (Nectris) nativitatis Streets, Bull. U. S. Nat. Museum, 1877, no. 7

p. 29. (Christmas Island, Pacific Ocean.)
Black Shearwater, Christmas Island Shearwater

Range. Pacific Ocean from the vicinity of 25° north latitude south
to the vicinity of 25° south latitude.

Recorded in the Hawaiian Group from Midway, Laysan (breeding),
French Frigate Shoal (breeding), "Bird Island" (Nihoa) (breeding?),
Lisianski (Munro ms.), Gardner (sight record by Dr. Isenbeck re-
corded by Kittlitz 1834), Oahu (Mokulua Islet) (sight by Donagho),
Mokumanu Islet (breeding) (Munro).

Puffinus newelli Henshaw
Puffinus newelli Henshaw, Auk, 17, 1900, p. 246. (Waihee Valley, "Ulani" in

error, Maui Island, see Henshaw, Birds of Hawaiian Possessions, p. 117,

footnote.)
Newell' s Shearwater, Ao

94 bulletin: museum of comparative zoology

Range. Hawaiian Islands.

Recorded in the Hawaiian Group from ?Niihau, Kauai (breeding),
Molokai, Maui. We can find no records for Lanai.

Genus Pterodroma — Petrels

Pterodroma Bonaparte, Comptes Rendus Academie Sciences, Paris, 42, 1856,
p. 768. Type, by subsequent designation, Procellaria macroptera A. Smith.

Pterodroma phaeopygia sandwichensis (Ridgway)

Oestrelala sandwichensis Ridgway, Baird, Brewer and Ridgway's Water

Birds of North America, vol. 2, 1884, p. 395. (Hawaiian Islands.)
Dark-rumped Petrel, Uau or Uwau
Oestrelala phaeopyga In Wilson and Evans, Aves Hawaiiensis.

Range. Hawaiian Islands.

Recorded in the Hawaiian Group from Hawaii (breeding on Mauna
Kea and Mauna Loa), Kauai (breeding), Molokai (breeding) (eggs in
Bishop Museum), Lanai (Munro).

Henshaw (1902) found that the mongoose had driven these birds
from their nesting grounds on Hawaii.

Pterodroma leucoptera hypoleuca (Salvin)

Oestrelata hypoleuca Salvin, Ibis, 1888, p. 359. (Krusenstern Island, North

Pacific Ocean.)
Salvin' s White Breasted Petrel, Bonin Island Petrel

Range. Northern Pacific Ocean. Breeds on the Bonin Islands ?
and Hawaiian Islands.

Recorded in the Hawaiian Group from Midway (breeding — eggs
collected by T. M. Blackman), Laysan (breeding) (Rothschild),
(W. K. Fisher 1903), ? Kaula (Caum), Lanai (Munro).

Mathews (1934) records Pterodroma leucoptera longirostris Stejneger
(1893) as the breeding bird of the Bonin Islands. This is not the case.
Birds of the Bonin Islands differ from Japanese birds and agree with
Hawaiian birds in having grayer under wing coverts and in their
large size.

Krusenstern Rock probably does not exist, reported by Captain
Lisianski in lat. 22° 15' N., long. 175° 37' W., it has been dropped
from the Pacific Islands Pilot of the British Admiralty (ed. 1931)
and though it is recorded in the Hawaiian Islands Pilot of the U. S.
Hydrographic Office (ed. 1933), there has been no actual report of a
rock in this vicinity since 1804 when breakers were reported thirty
miles south of the supposed position of the rock. The type locality
of this bird is therefore doubtful.

BRYAN AND GREENWAY : HAWAIIAN BIRDS 95

Mathews (1934) records the breeding range of this bird as follows:
"Hawaiian Group (Laysan, Pearl, Hermes [sic], Lisianski, Krusenstern
Reef, Midway and Ocean' Islands and French Frigate Shoal)." We
cannot find that this bird has ever been recorded as breeding on any
island except Laysan.

Genus Bulweria — Petrels

Bulweria Bonaparte, Nuova Annales Sci. Nat. Bologna, 8, 1842, p. 426. Type,
by monotypy, Procellaria bulweri Jardine and Selby.

Bulweria bulwerii (Jardine and Selby)

Procellaria bulwerii Jardine and Selby, Illust. Ornith. 2, 1828, pi. 65. (Ma-
deira.)

Soft Nosed Petrel, Bulwer's Petrel

Bulweria bulweri pacifica Mathews and Iredale, Ibis, 1915, p. 607. (Imojima,
Bonin Islands.)

Bulweria anjinho Heineken in Wilson and Evans, Aves Hawaiiensis, p. 211.

Range. "Breeding on islands off the coast of China; the Bonin
Islands, Vulcan Islands, the western Hawaiians and the Marquesas
Islands in the Pacific Ocean; Madeira, the Salvages, Canary and
Cape Verde Islands in the Atlantic" (Peters).

Recorded in the Hawaiian Group from Laysan, French Frigate
Shoal (breeding), Necker Island (breeding), Kauai, Hilo, Hawaii
(Henshaw), Oahu (skin in Bishop Museum, G. C. Munro coll.).

W. A. Bryan does not distinguish between sight records and col-
lected specimens. His records are from Necker and Bird Island
(Xihoa).

We quite agree with Hartert (Novitat. Zool., 33, 1926, p. 326) that
pacifica cannot be maintained.

Family HYDROBATIDAE

Genus Oceanodroma — Petrels

Oceanodroma Reichenbach, Aves Systema Naturae, 1852 (1853 fide Peters)
p. iv. Type, by original designation, Procellaria furcata Gmelin.

Oceanodroma castro cryptoleucura (Ridgway)
Cymochorea cryptoleucura Ridgway, Proceed. U. S. Nat. Museum, 4, 1882,

p. 337. (Kauai, Hawaiian Islands.)
Hawaiian Stormy Petrel, Ake-Ake (Henshaw), Oeoe (W. A. Bryan)

Range. Hawaiian Islands.

Recorded in the Hawaiian Group from Midway (Bartsch), French
Frigate Shoal, Niihau, Kauai.

"The natives report a small petrel, which they call by the above

96 bulletin: museum of comparative zoology

name (Ake-Ake), as common on the fishing grounds five or ten miles
off the coast of Hawaii, and I have little doubt that it is this species
..." (Henshaw).

OCEANODROMA MARKHAMI TRISTRAMI Salvin

Oceanodroma tristrami Salvin, Catalogue Birds British Museum, 25, 1896,

p. 354. (Sendai Bay, Japan.)
Sooty Petrel, Tristram's Petrel, Fork Tailed Petrel.
Oceanodroma fuliginosa (Gmelin) in Rothschild, Avifauna of Laysan, etc.,

p. 308; W. A. Bryan, Birds of the Hawaiian Group, p. 13 (footnote);

Henshaw, Birds of the Hawaiian Islands, p. 119; Oceanodroma tristrami

Stejneger in Bartsch, Auk, 34, 1922, p. 486; W. K. Fisher, Bull. U. S. Fish

Commission, 23, 1923, p. 795.

Range. From Japan south and west to the Hawaiian Islands.

Recorded in the Hawaiian Group from Midway (Bartsch), Laysan
(breeding), ?"Bird Island" (Nihoa) (W. K. Fisher), Lanai (skin in
Bishop Museum, G. C. Munro coll.).

In our opinion, Salvin's description of tristrami is identifiable and
Hartert's arguments (Vogel palaarktischen Fauna, p. 1416) have
no force under the Rules of Zoological Nomenclature. Mathews
records Cymochorea (markhami) owstoni Mathews and Iredale 1915,
as the breeding bird of the Hawaiian Islands in the belief that the
name tristrami is not identifiable.

Order PELECANIFORMES

Family PHAETHONTIDAE

Genus Phaethon — Tropic or Bos'n Birds

Phaethon Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 134. Type, by sub,
sequent designation, Phaethon adherens Linn. (Gray, List Genera Birds-
1840, p. 80). For synonymy see Peters, Check-List of Birds of the World,
1, p. 77.

Phaethon rubricauda rothschildi (Mathews)

Scaeophaethon rubricauda rothschildi Mathews, Birds Australia, 4, 1915, p. 303.

(Laysan and Niihau.)
Red-tailed Tropic Bird, Bos'n Bird, Koaeula
Scaeophaethon rubricauda brevirostris Mathews, Birds Australia, 4, 1915,

p. 303. (Bonin Islands.)
Phaethon rubricauda Boddaert in Rothschild, Avifauna of Laysan, etc., pp. 33,

294, 296 and other works.

BRYAN AND GKEKXWAY: HAWAIIAN BIRDS 97

Range. Pacific Ocean from the Bonin and Marianas Islands to the
Hawaiian Islands.

Recorded in the Hawaiian. Group from Midway (breeding; eggs in
Bishop Museum, T. M. Blackman coll.), Laysan (breeding), French
Frigate Shoal (sight by W. K. Fisher), Necker, Kaula, Lehua, Niihau,
Mokumanu Islet, Oahu (sight, Munro), Lanai (sight, Munro).

Wilson and Evans record this bird breeding on Kauai and Niihau,
and occurring on Hawaii.

Phaethon lepturus dorotheae Mathews

Phaethon lepturus dorotheae Mathews, Austral Avian Rec, 2, 1913, p. 7-

(Queensland.)
White Tailed Tropic Bird, Salmon Tailed Tropic Bird, Bos'n Bird, Koae,

Haakoae
Phaethon lepturus Daudin in Rothschild, Avifauna of Laysan, etc., p. 296;

Henshaw, Birds Hawaiian Islands, p. 114, and other works.
Phaethon aethereus Bloxam (nee Linn.) in Wilson and Evans, Aves Hawaiiensis,

p. 186.

Range. Australia and certain Pacific islands.

Recorded in the Hawaiian Group from Kauai, Oahu, Maui, Molokai
(specimens in Mus. Comp. Zool.), Hawaii.

Hartert (Novitat. Zoologicae, 32, 1925, p. 276) remarks that
dorotheae is probably not distinct from lepturus, which is the breeding
bird of Mauritius. This does not appear to us to be the case. The
species is in need of revision.

Family SULIDAE

Genus Sula — Boobies

Sula Brisson, Ornithologie, 1, 1760, p. 60; 6, p. 494. Type, by tautonomy,
"Sula" = Sula leucogaster Boddaert.

Sula sula rubripes Gould

Sula rubripes Gould, Synopsis Birds Australia, pt. 4, 1838, append., p. 7.

(New South Wales = Raine Island, N. Queensland, fide Mathews.)
Tied Footed Booby

Sula piscatrix Linn, in Rothschild, Avifauna of Laysan, etc., pp. 27, 297.
Sula piscator Linn, in Wilson and Evans, Aves Hawaiiensis, p. xxv; Henshaw,

Birds Hawaiian Islands, p. 113; W. A. Bryan, Key, Birds Hawaiian

Group, p. 15.

98 bulletin: museum of comparative zoology

Range. "Islands of the Indian and tropical western and central
Pacific Oceans" (Peters).

Recorded in the Hawaiian Group from Midway, Lisianski and Laysan
(breeding), French Frigate Shoal (Bryan), "Bird Island" (Nihoa)
(breeding), Lehua (Rothschild), Niihau, but see W. K. Fisher, Bull.
U. S. Fish Comm., 23, 1903, p. 797, Kaula, Kauai, Oahu (on Mokumanu
Island) (breeding), and many sight records at sea.

Sula leucogaster plotus (Forster)

Pelecanus plotus Forster, Descriptiones Animalium, etc., (ed. Lichtenstein)

1844, p. 278. (Near New Caledonia.)
Booby, Common Brown Booby, Brown Vested Booby
Sula sula Linn, in Rothschild, Avifauna of Laysan, etc., pp. 28, 297; Wilson

and Evans, Aves Hawaiiensis, p. xxv; Henshaw, Birds Hawaiian Islands,

p. 114; W. A. Bryan, Key, Birds Hawaiian Islands, p. 15; Henshaw,

Birds Hawaiian Possessions, p. 114.

Range. "Western and central tropical Pacific to northeastern
Australia" (Peters).

Recorded in the Hawaiian Group from Midway, Lisianski and Laysan
Islands, French Frigate Shoal (sight) (Fisher), Necker (breeding),
"Bird Island "(Nihoa) (breeding), Kaula (Caum), and many sight
records at sea off Niihau, Oahu (Mokumanu Islet) (Munro).

Sula dactylatra person at a Gould

Sida personata Gould, Proc. Zool. Soc. London, 1846, p. 21. (North and
northeast coasts of Australia = Raine Island, North Queensland, fide
Mathews.)

Booby, Blue Faced Booby, Masked Booby, Masked Gannet

Sula cyanops Sundevall in Rothschild, Avifauna of Laysan, etc., p. 25; Wilson
and Evans, Aves Hawaiiensis, p. xxv; Henshaw, Birds Hawaiian Islands,
p. 113; W. A. Bryan, Key, Birds Hawaiian Group, p. 15.

Range. "Breeds on islands of the central and western tropical
Pacific Ocean" (Peters).

Recorded in the Hawaiian Group from Midway (breeding), Lisianski,
Laysan (breeding), French Frigate Shoal (breeding), Necker (breed-
ing), "Bird Island" (Nihoa) (breeding), Kaula, Lehua (Caum).

Family PHALACROCORACIDAE

Genus Phalacrocorax — Cormorants

Phalacrocorax Brisson, Ornithologie, 1, 1760, p. 60. Type, by tautonomy,
Phalacrocorax = Pelecanus carbo Linnaeus.

BRYAN AND GREENWAY: HAWAIIAN BIRDS 99

Phalacrocorax pelagicus pelagic is Pallas

Phalacrocorax pelagicus Pallas, Zoographia Rosso-Asiatica, 2, 1811, p. 303.

(Eastern Kamchatka and the Aleutian Islands.)
Pelegic Cormorant

Range. Breeds on the coasts of northeastern Asia, Aleutian Islands
to south-central Alaska; south in winter to Japan and Puget Sound.

Recorded in the Hawaiian Group {as an accidental visitor) from
Laysan 1896 (Rothschild), ?Hilo, Hawaii 1900 (recorded by Henshaw
with doubt).

Family FREGATIDAE

Genus Fregata — Frigate Birds or Man o'War Birds

Fregata minor palmerstoni (Gmelin)

Pelecanns palmerstoni Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 573.

(Palmerston Island = Palmerston Atoll, Cook Group.)
Man o'War Bird, Ima
Fregata aquila Linn, in Rothschild, Avifauna of Laysan, etc., pp. 22, 297 and

other works.

Range. "Hawaiian Islands south to New Zealand" (Peters).

Recorded in the Hawaiian Group from Midway (breeding), Pearl and
Hermes Reef, Laysan (breeding), French Frigate Shoal, Necker
(breeding), "Bird Island" (Nihoa) (breeding), Kaula, Lehua, Kauai,
Oahu.

Order CICONIIFORMES

Family ARDEIDAE— Herons

Genus Nycticorax — Night Herons

Nycticorax T. Forster, Synoptic Catalog British Birds, 1817, p. 59. Type, by
tautonomy and monotypy, Nycticorax infaustus Forster = Ardea nycti-
corax Linn.

Nycticorax nycticorax hoactli (Gmelin)

Ardea hoactli Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 630. (In Novae

Hispania lacubus = Valley of Mexico.)
Black Crowned Night Heron, Auku Aukuu, Auku Kahili or Auku Kahili
Nycticorax nycticorax naevius (Boddaert) in Rothschild, Avifauna of Laysan,

etc., p. 265.
Nycticorax griseits (Boddaert) in Wilson and Evans, Aves Hawaiiensis, p. 201,

and Henshaw, Birds Hawaiian Islands, p. 102.

100 bulletin: museum of comparative zoology

Range. Northern United States and northeastern Canada south
through Mexico and Central America and east coast of South America
to eastern Argentine.

Recorded in the Hawaiian Group from Midway (skin in Bishop
Museum — D. R. Chisholm coll.), Oahu, Molokai, Hawaii — "all
islands" (Rothschild, AYilson and Evans, Henshaw).

The records of Demigretta sacra seem to be doubtful. As Bryan
(1901, p. 21) remarks, Dole's record of 1879 might just as well refer
to the Auku.

The identity of the White Heron which Dr. Finsch saw so long ago
at Kahalui is problematical.

Family THRESKIORNITHIDAE

Genus Plegadis — Glossy Ibises

Plegadis Kaup, Skizzirte Entwickelungs-Geschichte Natiirliches System, etc.,
1829, p. 82. Type, by monotypy, Tantalus falcinellus Linnaeus.

Plegadis guarauna (Linnaeus)

Scolovax gaurauna Linnaeus, Systema Naturae, ed. 12, 1, 1766, p. 242.

(Brazil.)
White Faced Glossy Ibis

Range. Northwestern North America south to Argentina.
Recorded in the Hawaiian Group (as an accidental visitor) from
Kauai, ?Maui (Henshaw), Molokai (Munro).

Order ANSERIFORMES

Family ANATIDAE

Genus Chen — Snow Geese

Chen Boie, Isis von Oken, 10, 1822, column 563. Type, by monotypy, Anser
hyperboreus Pallas.

Chen hyperborea hyperborea (Pallas)

Anser hyperboreus Pallas, Spicilegia Zoologica, etc., 1767 (1769?), fascicule 6

1769, p. 31. (Northeastern Siberia.)
Lesser Snow Goose

BRYAN AM) GKKK.WVAY: HAWAIIAN BIRDS 101

Range. Arctic North America south in winter to temperate zone.
Probably also Siberia south in winter to Japan.

Recorded in the Hawaiian Group (as an accidental visitor) from
Maui by Rothschild who received a specimen collected there by
Brother Matthias Newell; Oahu (J. E. Whitney coll.).

Genus Axser — Geese

A user Brisson, Ornithologie, 1, 1760, p. 58. Type, by tautonomy, Anser
domesticus = Anser anser Linnaeus.

Anser albifrons gambelli Hartlaub

Anser gambelli Hartlaub, Revue et Magasin Zoologie, etc., 1852, p. 7. (Texas

and southern United States.)
American White-Fronted Goose

Range. Northwestern North America south in winter to northern
California.

Recorded in the Hawaiian Group (as an accidental visitor) from
"a lake on Mr. Clark's estate at Honokaohau," Hawaii, where it was
shot December 18, 1891 (Rothschild), Molokai (G. C. Munro coll.).

Genus Philacte — Emperor Goose

Philacte Bannister, Proceedings Acad. Nat. Sci. Philadelphia, 1870, p. 131.
Type, by monotypy, Anas canagica Sewastianov.

Philacte canagica (Sewastianov)

Anas canagica Sewastianov, Nova Acta Acad. Sci. Imp. St. Petersburg, 13,

1802, p. 349, pi. 10. (Kanaga Island, Aleutian Islands.)
Emperor Goose

Range. Breeds on coasts of Siberia and Alaska, winters chiefly
in the Aleutian Islands.

Recorded in the Hawaiian Group (as an accidental visitor) from
Puna, Hawaii (Henshaw 1903).

Genus Branta — Brant or Brent Geese

Branta Scopoli, Annus I, Historico-Naturalis, 1769, p. 67. Type, by sub-
sequent designation, Anas bernicla Linnaeus (Bannister, Proceedings
Acad. Nat. Sci., Philadelphia, 1870, p. 131).

102 bulletin: museum of comparative zoology

Branta bernicla nigricans (Lawrence)

Anser nigricans Lawrence, Annals Lyceum Nat. Hist. New York, 4, 1846,

p. 171, pi. 12. (Egg Harbor, New Jersey.)
Black Brant

Range. Arctic coasts of Siberia and Arctic North America; south
in winter to Japan and North China, and to Lower California.

Recorded in the Hawaiian Group (as an accidental visitor) from Maui
(Rothschild).

Branta canadensis minima Ridgway

Branta minima Ridgway, Proceedings U. S. Nat. Museum, 8, 1885, p. 22.

("Pacific coasts of North America"; type from St. Michael's, Alaska.)
Cackling Goose
Bernicla Munroii Rothschild, Annals and Magazine Nat. Hist. (6), 10, 1892,

p. 108. (Kauai.)

Range. Breeds on the Bering Sea coast of Alaska and the Aleutian
Islands; in winter, west of the Rocky Mountains from southern
British Columbia to southern California.

Recorded in the Hawaiian Group (as an occasional or chance visitor)
from Kauai (Rothschild), ?Hawaii (Henshaw, q. v.), Molokai (G. C.
Munro coll.).

Genus Nesochen — Hawaiian Goose

Nesochen Salvadori, Catalogue Birds British Mus., 27, 1895, pp. 81, 126.
Type, by original designation and monotypy, Anser sandvicensis Vigors.

Nesocehn sandvicensis (Vigors)

Anser sandvicensis Vigors, List Animals Garden Zool. Soc, ed. 3, 1833, p. 4.
(Hawaiian Islands.)

Hawaiian Goose, Nene

Bernicla sandvicensis in Wilson and Evans, Aves Hawaiiensis, p. 187; Hen-
shaw, Birds Hawaiian Islands, p. 103.

Range. Hawaiian Islands.

Recorded in the Hawaiian Group from Hawaii. (Reports from
Maui, Kauai and Niihau have never been confirmed.) Breeds on the
Kona coast, and above 5000 feet; from the Kona District to the north-
east side of Mauna Kea, descending to 1000 feet and sea level in
winter. Nests in lava flows or open fields. Now much reduced in
numbers, though still breeding in a wild state as well as in captivity.

BRYAN AND GREENWAY: HAWAIIAN BIRDS 103

Genus Anas

Anas Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 122. Type, by subsequent
designation, Anas boschas Linnaeus = Anas platyrhynchos Linnaeus.

Anas platyrhynchos platyrhynchos Linnaeus

Anas platyrhynchos Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 125.

(Europe, type locality restricted to Sweden.)
Mallard

Range. Temperate zones of Europe, Asia and North America.
Recorded in the Hawaiian. Group (as an accidental visitor) from
Laysan (Schauinsland), Oahu, Molokai (Perkins).

Anas wyvilliana wyvilliana Sclater

Anas wyvilliana Sclater, Proc. Zool. Soc. London, 1878, p. 350. (Hawaiian

Islands.)
Hawaiian Duck, Koloa Maoli
Anas aberti Ridgway, Proc. U. S. Nat. Mus., 1, 1878, p. 250. (Mazatlan,

Mexico.)

Range. Hawaiian Islands. Recorded from Niihau, Kauai, Oahu,
Molokai, Maui and Hawaii in ponds and lakes along the coast and
in mountain streams and bogs to 8000 feet.

Recorded recently from Kauai, Mokulua Islets, N. E. coast of
Oahu (breeding) (Munro), Molokai, highlands of Hawaii (Baldwin).

The type of Anas aberti Ridgway was probably an escaped captive.
Mazatlan was formerly a port for vessels from China and possessed a
large bird market.

Anas wyvilliana laysanensis Rothschild

Anas laysanensis Rothschild, Bull. British Ornith. Club, 1, 1892, p. xvii.

(Laysan Island.)
Laysan Teal

Range. Laysan. Recorded by Kittlitz (Museum Senckenbergianum
1834, p. 124) from Lisianski but never confirmed.

Twenty birds left in 1923 (Wetmore, Nat'l Geographic Mag., 48,
no. 1, 1925, p. 103). Nine left in 1936 (W. F. Coultas, in Hit).

104 bulletin: museum of comparative zoology

Anas crecca carolinensis Gmelin

Anas carolinensis Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 533. (Carolina to

Hudson's Bay.)
Green Winged Teal

Range. Northern North America; in winter south to West Indies
and Panama.

Recorded in the Hawaiian Group (as an accidental visitor) from
Laysan (Schauinsland), Maui (G. P. Wilder coll.), Oahu (Northwood),
Molokai (G. C. Munro coll.).

Anas acuta acuta Linnaeus

Anas acuta Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 126. (Europe —

type locality restricted to Sweden.)
Pintail, Koloa Mapu

Range. Europe, Northwestern North America east to Iowa; south
in winter to West Indies, Panama and the Hawaiian Islands.

Recorded in the Hawaiian Group (as a regular migrant in winter)
from Laysan, Kauai, Oahu (Northwood), Hawaii. "Most of the
islands; common" (Perkins). "Well known to sportsmen as a winter
visitor" (Henshaw).

Genus Mareca — W7idgeons

Mareca Stephens, in Shaw, General Zoology, 12, pt. 2, 1824, p. 30. Type, by
subsequent designation, Mareca fistularis Stephens = Anas penelope
Linnaeus (Eyton, Monograph Anatidae, 1838, p. 33).

Mareca Americana (Gmelin)

Anas americana Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 526. (Louisiana

and New York.)
American Widgeon, Baldpate

Range. Northwestern to north-central North America; south in
winter to the Atlantic coast, West Indies and Central America.

Recorded in the Hawaiian Group (as an accidental visitor in winter)
from Laysan (Schauinsland), Oahu (Northwood), Maui (Perkins),
Molokai (Munro coll.).

BRYAN AND GRKKN WAY : HAWAIIAN' KIRDS 105

Genus Chaulelasmus — Gadwall Ducks

Chaulelasmus Bonaparte, Geographical and Comparative List of Birds of
Europe and North America, 1838, p. 56. Type, by monotypy, Anas
strepera Linnaeus.

Chaulelasmus streperus (Linnaeus)

Anas strepera Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 125. (Europe —

type locality restricted to Sweden.)
Gad wall

Range. Worldwide within the northern hemisphere.
Recorded in the Hawaiian Group (as an accidental visitor in winter)
from Oahu (Perkins), Molokai (Munro coll.).

Genus Spatula — Shoveller Ducks

Spatula Boie, Isis von Oken, 1822, col. 564. Type, by monotypy, Anas
clypeata Linnaeus.

Spatula clypeata (Linnaeus)

Anas clypeata Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 124. (Coasts of

Europe; type locality restricted to southern Sweden.)
Shoveller, North West Duck, Koloa Moha

Range. Worldwide within the northern hemisphere.
Recorded in the Hawaiian Group (as a regular migrant in winter)
from Laysan, Kauai, Oahu, Molokai, Lanai, Maui, Hawaii.

Genus Bucephala — Golden Eyes, Buffle Head

Bucephala Baird, Reports, Explorations and Surveys for a Railroad from the
Mississippi River to the Pacific Ocean, 9, pt. 2, pp. L, 788, 785. Type, by
original designation, Anas albeola Linnaeus.

Bucephala albeola (Linnaeus)

Anas albeola Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 124. (America =

Newfoundland ex Edwards.)
Bvffle Head

Range. Northwestern North America; in winter eastward and
southward in the United States.

Recorded in the Hawaiian Group (as an accidental visitor) from
Laysan (Rothschild), Oahu (Northwood), Maui (Perkins).

106 bulletin: museum of comparative zoology

Genus Nyroca — Diving Ducks

Nyroca Fleming, Philosophy of Zool. etc., 2, 1822, p. 260. Type, by tautonomy,
Anas nyroca Giildenstadt.

Nyroca affinis (Eyton)

Fuligula affinis Eyton, Monograph Anatidae, 1838, p. 157. (North America.)
Lesser Scaup Duck

Range. Northern North America; south in winter to Central
America.

Recorded in the Hawaiian Group (as an accidental visitor) from
Lanai (G. C. Munro coll.).

Genus Histrionicus — Harlequin Ducks

Histrionicus Lesson, Manuel d'Ornithologie, 2, 1828, p. 415. Type, by original
designation, Anas histrionica Linnaeus.

Histrionicus histrionicus pacificus W. S. Brooks

Histrionicus histrionicus pacificus W. S. Brooks, Bull. Mus. Comp. Zool.,

Cambridge, Mass., 59, 1915, p. 393. (Cape Shipunski, Kamchatka.)
Western Harlequin Duck

Range. Eastern Siberia and northwestern North America ; south in
winter to Japan and California.

Recorded in the Hawaiian Group (as an accidental visitor) from Lay-
san (P. E. H. Bompke coll. 1906).

Genus Mergus — Mergansers

Mergus Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 129. Type, by subse-
quent designation, Mergus castor Linnaeus = Mergus serrator Linnaeus.

Mergus serrator Linnaeus

Mergus serrator Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 129. (Europe.
Type locality restricted to Sweden.)

Range. Worldwide in the northern hemisphere ; south in winter to
temperate zone.

Recorded in the Hawaiian Group (as an accidental visitor) from 0§hu
(Bryan), Hawaii (Henshaw).

BRYAX AND GRKK.WVAY: HAWAIIAN BIRDS 107

Order FALCOXIFORMES
Family ACCIPITRIDAE

Genus Buteo — Soaring Hawks and Buzzards

Buteo Lacepede, Discours . . . Tableaux methodiques mammiferes . . •
Oiseaux, 1799, p. 4. Type, by tautonom}', Falco buteo Linnaeus.

Buteo solitarius Peale

Buteo solitarius Peale, U. S. Exploring Expedition, 8, 1848, p. 62. (Island of

Hawaii.)
Hawaiian Hawk, Io
Accipiter hawaii Dole, Hawaiian Almanac for 1879 (1878) p. 43.

Range. Island of Hawaii (all parts).

Genus Circus — Marsh Hawks

Circus Lacepede. Discours . . . Tableaux methodiques mammiferes oiseaux,
1799, p. 4. Type, by subsequent designation, Falco aeruginosus Linnaeus
(Lesson, Manuel d'Ornith., 1, 1828, p. 105).

Circus cyaneus hudsonius (Linnaeus)

Falco hudsonius Linnaeus, Systema Naturae, ed. 12, 1, 1766, p. 128. (Hudson
Bay ex Edwards.)

Marsh Hawk-
Range. Northern and central North America; south in winter to

Central America.

Recorded in the Hawaiian Group (as an accidental visitor) from Oahu

(Wilson).

Genus Pandion — Ospreys

Pandion Savigny, Description d'Egypt, Oiseaux, 1809, p. 69. Type, by
monotypy, Pandion fluvialis Savigny = Falco haliaetus Linnaeus.

Pandion haliaetus carolinensis (Gmelin)

Falco carolinensis Gmelin, Systema Naturae, 1, pt. 1, 1788, p. 263. (No type

locality. Carolina ex Catesby et al.)
American Osprey

108 bulletin: museum of comparative zoology

Range. Northern North America; south in winter to the West
Indies and South America.

Recorded in the Hawaiian Group (as an accidental winter visitor)
from ?Niihau (Dole), Oahu (Perkins), ?Molokai (Dole), ?Hawaii
(Dole).

Family RALLIDAE— Rails

Genus Porzanula

Porzanula Frohawk, Annals and Magazine Nat. Hist., London, (6), 9, 1892,
p. 247. Type, by monotypy, Porzanula palmeri Frohawk.

Porzanula palmeri Frohawk

Porzanula Palmeri Frohawk, Annals and Magazine Nat. Hist., London (6), 9,

1892, p. 247. (Laysan Island.)
Laysan Rail, Laysan Crake

Range. Laysan Island, where but two remained in 1923 (Wetmore
1925); Midway Island, where introduced in the latter part of the 19th
century (Rothschild, p. xiii) and where common (Denig et al).

Genus Pennula

Pennula Dole, Hawaiian Almanac and Annual for 1879 (1878), p. 54. Type,
by monotypy, Pennula millei (sic) Dole.

Pennula millsi Dole

Pennula millei [sic] Dole, Hawaiian Almanac and Annual for 1879 (1878),
p. 54. (Hawaii.)

Hawaiian Rail, Moho

Rallus ecaudotus (sic) King, in Cook's Voyage to the Pacific Ocean, 3, 1784,
p. 119. (Hawaiian Islands); Pennula ecaudata (King) in Wilson and
Evans, Aves Hawaiiensis, p. 171, and in Perkins, Fauna Hawaiiensis, 1,
pt. 4, p. 453, and older works. This name is preoccupied by Rallus ecau-
datus Miller 1783.

Range. Formerly Island of Hawaii in open grassy country or low
scrub just below the heavy rain forest, possibly in the Olaa district,
certainly on the windward side of Kilauea and perhaps about forty
miles north along the coast. ?Molokai (Perkins). Extinct, last speci-
men seen 1884 (Rothschild), ?1893 (Henshaw). Five specimens in
museums.

BRYAN AND GREENWAY: HAWAIIAN BIRDS 109

Pennula sandwichensis (Gmelin)

Rallus sandwichensis Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 717. ("in
insulis Sandwich," based on Latham's Sandwich Rail.)

Spotted Hawaiian Rail

Rallus obscurus Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 718. ("in insulis
Sandwich," based on Latham's Dusky Rail, and other authors.)

Pennula wilsoni Finsch, Notes Leyden Mus., 20, 1898, p. 77. Hawaiian
Islands.)

Range. Unknown. Now extinct. Known only from the unique
type in Leyden, Holland. Paler than millsi with central black spot on
feathers of back.

Genus Gallinula — Coots, Gallinules

Gallinula Brisson, Ornithologie, 1, 1760, p. 2. Type, by tautonomy, Gallinula
Brisson = Fulica chloropus Linnaeus.

Gallinula chloropus sandvicensis Streets

Gallinula sandvicensis Streets, Ibis, 1877, p. 25. (Oahu, Hawaiian Islands.)
Hawaiian Gallinule, Alae, Alae Ula

Range. Hawaiian Islands. Recorded from Kauai, Oahu, Molokai,
Maui, Hawaii in ponds, rice fields and taro patches.

Genus Fulica — Coots

Fulica Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 152. Type, by tau-
tonomy, Fulica atra Linnaeus.

Fulica Americana alai Peale

Fulica alai Peale, United States Exploring Expedition, 8, 1848, p. 224.

(Hawaiian Islands.)
Hawaiian Coot, Alae Keokeo

Range. Hawaiian Islands. Recorded from Niihau, Kauai, Oahu>
Molokai, Maui, Hawaii.

Order CHARADRIIFORMES

Family CHARADRIIDAE

Genus Squatarola — Plovers

Squatarola Cuvier, Regne Animal, 1, 1817 (1816), p. 467. Type, by tautonomy,
Tringa squatarola Linnaeus.

110 bulletin: museum of comparative zoology

Squatarola squatarola (Linnaeus)

Tringa squatarola Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 149. (Europe.

Type locality restricted to Sweden by Hartert, Vogel palaarktischen

Fauna, 2, p. 1553.)
Black-bellied Plover, Beetle Head

Range. Circumpolar in the arctic; south in winter to South Africa*
India, Australia, western South America.

Recorded in the Hawaiian Group (as an accidental visitor on migration)
from Oahu (Northwood et al), Hawaii (Henshaw).

Genus Pluvialis

Pluvialis Brisson, Ornitholigie, 1760, 1, p. 46; 5, p. 42. Type, by tautonomy,
Pluvialis aurea Brisson = Charadrius pluvialis Linnaeus = Charadrius
apricarius Linnaeus.

Pluvialis dominica fulva (Gmelin)

Charadrius fulvus Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 687. (Tahiti.)

Pacific Golden Plover, Kolea

Charadrius fulvus in Rothschild and other works.

Range. Siberia and Alaska; south in winter to India, Australia,
Indo-China, Pacific Islands.

Recorded in the Hawaiian Group (as a regular winter visitor) from
Midway, Laysan, Lisianski, French Frigate Shoal, Bird Island (Nihoa),
Niihau, Kauai, Oahu, Molokai, Maui, Hawaii, "all islands" (Perkins).
Arrive in August or September; depart in April or May as a rule, but
sometimes they remain during the summer, though probably they
never breed.

Genus Charadrius — Plovers

Charadrius Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 150. Type, by
tautonomy, Charadrius hiaticula Linnaeus.

Charadrius vociferus vociferus Linnaeus

Charadrius vociferus Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 150.

(North America = South Carolina ex Catesby.)
Killdeer

Range. Arctic and temperate zones and portions of the eastern
tropical; south in winter to northern South America.

Recorded in the Hawaiian Group (as an accidental visitor) from Maui
(W. A .Bryan).

BRYAN AND GREENWAY: HAWAIIAN BIRDS 111

Family SCOLOPACIDAE

Genus Numenius — Curlews

Numenius Brisson, Ornithologie, 17G0, 1, p. 48; 5, p. 311. Type, by tautonomy,
X miliums Brisson = Scolopax arquata Linnaeus.

Numenius tahitiensis (Gmelin)

Scolopax tahitiensis Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 656. (Tahiti

ex Latham.)
Bristle Thighed Curlew, Kioea

Range. Alaska; south in winter to the Pacific islands.

Recorded in the Hawaiian Group (as a regular winter visitor) from
.Midway, Laysan, Niihau, Kauai, Oahu, Molokai, Lanai (sight)
(Rothschild), Maui, Hawaii. Like the Golden Plover (Kolea), this
bird leaves in April or May and arrives in August or September, but
sometimes remains during the summer, though it probably never
breeds on the islands.

Genus Limosa — Godwits

Limosa Brisson, Ornithologie, 1760, 1, p. 48; 5, p. 261. Type, by tautonomy,
Limosa Brisson = Scolopax limosa Linnaeus.

Limosa lapponica baueri Naumann

Limosa baueri Naumann, Naturgeschichte Vogel Deutschlands, 8, 1836, p.
429. (New Holland.)

Pacific Godwit

Limosa lapponica novaezealandiae Gray in Rothschild, Avifauna of Laysan,
etc., p. 307; Henshaw, Birds Hawaiian Islands, etc., p. 93; Bryan, Key
Birds Hawaiian Group, p. 27; Perkins, Fauna Hawaiiensis (Aves), p. 45.

Range. Northeastern Asia and northwestern North America;
south in winter to islands of the Pacific and Australia.

Recorded in the Hawaiian Group (as an occasional visitor on migration)
from Laysan (Schauinsland), Kauai (W. A. Bryan), Maui (Ridgway).

Genus Heteroscelus — Tatlers

Heteroscelus Baird, Reports, Explorations and Surveys for a Railroad from the
Mississippi River to the Pacific Ocean, 9, 1858, pp. xxii, xlvii, 728, 734.
Type, by monotypy, Totanus brevipies Vieillot.

112 bulletin: museum of comparative zoology

Heteroscelus incanus (Gmelin)

Scolopax incana Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 658. (Eimeo

(= Moorea) and Palmerston Islands.)
Wandering Tattler, Ulili
Heteractitis incanus (Gmelin) in Rothschild, Avifauna of Laysan, etc., p. 255;

Henshaw, Birds Hawaiian Islands, etc., p. 92; Bryan, Key Birds Hawaiian

Group, p. 27; Perkins, Fauna Hawaiiensis (Aves), p. 450; Totanus incanus

in Wilson and Evans, Aves Hawaiiensis, p. 151.

Range. Alaska; south in winter to the west coast of America to
Ecuador and to the islands of the Pacific, sometimes to Australia.

Recorded in the Hawaiian Group (as a regular winter resident) from
Midway, Laysan, Lisianski, Necker, French Frigate Shoal, Niihau,
Kauai, Oahu, Maui, Hawaii," .... frequenting the rocky shores of all
islands" (Henshaw). Usually arrive in August and leave in May.
Though certain birds may remain all summer, there is no breeding
record for the islands.

Genus Arenaria — Turnstones

Arenaria Brisson, Omithologie, 1760, 1, p. 48; 5, p. 132. Type, by tautonomy,
Arenaria Brisson = Tringa interpret Linnaeus.

Arenaria interpres interpres (Linnaeus)

Tringa interpres Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 149. (Europe
and North America; type locality restricted to Gotland Island, Sweden,
by Hartert, Vogel Palaartischen Fauna, p. 1566.)

Turnstone, Akekeke

Tringa oahuensis Bloxam, Voyage "Blonde", 1826, p. 132.

Range. Northern Europe and northern Asia and Alaska west of
Point Barrow; south in winter to Africa, India, China and islands of
the Pacific.

Recorded in the Hawaiian Group (as a regular winter resident) from
Midway, Lisianski, Laysan, French Frigate Shoal, Necker, Niihau,
Kauai, Oahu, Molokai, Maui, Hawaii, "All the islands" (Perkins).
Usually arrive in August or September and depart in April or May.
Certain birds remain all summer, some in flocks at 6000 feet on Halea-
kala, Maui and in the crater of Kilauea, Hawaii. Subspecific identifica-
tion of winter birds is impossible. It seems more probable that most
of the birds to be seen in the Hawaiian Islands are interpres, though in-
dividuals of morinella, which is the bird that breeds in North America
east of Point Barrow, Alaska, may also reach the islands.

BRYAN AND GREENWAY: HAWAIIAN' BIRDS 113

Genus Capella — Snipe

Capella Frenzel, Beschreibung Vogel und Eyer . . . Wittenborg . . . , 1801,
p. 58. Type, by nionotypy, Scolopax coelestis Frenzel = Scolopax gallinago
Linnaeus.

( apella delicata (Ord)

Scolopax delicata Ord in Wilson, American Ornithology, 9, 1825, p. 218.

(Pennsylvania.)
Wilson's Snipe, Jack Snipe
Gallinago delicata in Rothschild, Avifauna of Laysan, etc., p. 253; Henshaw,

Birds Hawaiian Islands, p. 94; Perkins, Fauna Hawaiiensis (Aves), p. 451.

Range. Arctic and temperate North America; south in winter to
South America.

Recorded in the Hawaiian Group (as an occasional visitor on migration)
from Laysan, Oahu, Molokai, Maui, Hawaii.

Genus Crocethia — Sanderling

Crocethia Billberg, Synopsis Fauna Scandanaviae, 1, pt. 2, 1828, p. 132. Type
by nionotypy, Charadrius calidris Linnaeus = Trynga alba Pallas.

Crocethia alba (Pallas)

Trynga alba Pallas in Vroeg, Beredeneerde Catalogus, etc., Adumbratiunculae,
1764, N. 320, p. 7. ("Noordsche Zeekusten" = coast of northern Holland.)

Sanderling, Hunakai

Calidris aretiaria (Linnaeus) in Rothschild, Avifauna of Laysan, etc., p. 259;
Wilson and Evans, Aves Hawaiiensis, p. 153, 159; Henshaw, Birds
Hawaiian Islands, etc., p. 93; Perkins, Fauna Hawaiiensis (Aves), p. 451,
and other works.

Range. Arctic coasts of Europe, Asia, America ; south in winter to
temperate zone and southern hemisphere — Cape of Good Hope,
Malaya, Australia, Islands of the Pacific, southern South America.

Recorded in the Hawaiian Group (as a regular winter resident) from
Laysan, Niihau, Kauai, Maui (Ridgway), Oahu, Molokai, Hawaii,
"Most, and probably all, islands" (Perkins).

We have followed the A. O. U. Check-list and Peters (Birds of the
^Yorld) in recognizing but one form of this species. If more are recog-
nized the name of the bird which visits the islands will be Crocethia
alba tridactyla Pallas, the supposed form which breeds in northwestern
North America.

114 bulletin: museum of comparative zoology

Genus Erolia — Sandpipers

Erolia Vieillot, Analyse nouvelle ornithologie . . . 1816, p. 55. Type, by
monotypy, Erolia variegata Vieillot = Scolopax testacea Pallas.

Erolia acuminata (Horsfield)

Totanus acuminatum Horsfield, Transactions Linnaean Soc. London, 13, pt. 1,

1821, p. 192. (Java.)
Sharp-tailed Sandpiper
Heteropygia acuminata in Rothschild, Avifauna Laysan, etc., p. 255, 307;

Tringa acuminata in Henshaw, Birds Hawaiian Islands, p. 94; Perkins,

Fauna Hawaiiensis (Aves), p. 451.

Range. Eastern Asia; south in winter to south Pacific islands,
Malaya, New Guinea, Australia.

Recorded in the Hawaiian Group (as an occasional visitor on migra-
tion) from Midway, Laysan, Kauai, Oahu, Molokai (specimen in
Bishop Museum), Maui, Hawaii.

Erolia melanotos (Vieillot)

Tringa melanotos Vieillot, Nouveau Dictionnaire Hist. Nat., 34, 1819, p. 462.

(Paraguay.)
Pectoral Sandpiper
Tringa maculata in Henshaw, Birds Hawaiian Islands, etc., p. 94; Perkins,

Fauna Hawaiiensis (Aves), p. 451.

Range. Arctic zone of eastern Asia and North America; south in
winter to South America.

Recorded in the Hawaiian Group (as an accidental visitor on migration)
from Oahu (W. A. Bryan), Hawaii.

Genus Himantopus— Stilts

Himantopus Brisson, Ornithologie, 1760, 1, p. 46; 5, p. 33. Type, by tau-
tonomy, Himantopus Brisson = Charadrius himantopus Linnaeus.

Himantopus himantopus knudseni Stejneger

Himantopus knudseni Stejneger, Proceedings U. S. Nat. Mus., 10, 1887, p. 81,

pi. 6, fig. 2. (Kauai, Hawaiian Islands.)
Hawaiian Stilt, Aeo, Kukuluaeo

Range. Hawaiian Islands, where recorded from Niihau, Kauai,
Oahu, Molokai, Maui near lakes and ponds or on exposed mud flats.
"I have never seen it, however, upon Hawaii, nor have I been able to
learn of its presence there" (Henshaw).

B RY A N A ND GR E E N W A V : 1 1 A W AIIAN BIRDS 11 5

Family PHALAROPODIDAE

Genus Phalaropus — Phalaropes

Phalaropus Brisson, Ornithologie, 1760, 1, p. 50; 6, p. 12. Type, by tautonomy,
Phalaropus Brisson = Tringa fulicaria Linnaeus.

Phalaropus fulicarius (Linnaeus)

Tringa fulicaria Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 148. (Hudson

Bay ex Edwards.)
Red Phalarope
Crymophilus fulicarius in Rothschild, Avifauna Laysan, etc., p. 251; Henshaw,

Birds Hawaiian Islands, p. 95; Bryan, Key Birds Hawaiian Group, p. 25.

Range. Circumpolar in arctic; south in winter to seas off the coasts
of West Africa and Chile.

Recorded in the Hawaiian Group (as an irregular visitor on migration)
from Laysan, Kauai, Oahu, Maui, Hawaii, "evidently accompanied
the Akekeke (Arenaria interpres) to the uplands to feed" (!) (Hen-
shaw).

Phalaropus lobatus (Linnaeus)

Tringa tobata {Lobata in Emendanda, p. 824) Linnaeus, Systema Naturae, ed.

10, 1, 1758, p. 148. (Hudson Bay ex Edwards.)
Northern Phalarope

Range. Circumpolar in Arctic Zone; south in winter to oceans in
southern hemisphere.

Recorded in Hawaiian Group (as a chance visitor on migration) from
Kauai.

Family LARIDAE

Genus Larus — Gulls

Larus Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 136. Type, by subsequent
designation, Larus marinus Linnaeus (Selby, Catalogue generic and sub-
generic types . . . Aves, 1840, p. 48).

Larus delawarensis Ord

Larus delawarensis Ord in Guthrie's New Geographical . . . Grammar, etc.,
2nd American ed., 2, 1815, p. 319. (Delaware River, below Philadelphia.)
Ring-billed Gull

116 bulletin: museum of comparative zoology

Range. Northwestern and northeastern North America; south in
winter, sometimes as far as the Greater Antilles and Mexico.

Recorded in the Hawaiian Group (as an accidental visitor) from
Molokai and Maui or Hawaii (Bryan).

Larus argentatus smithsonianus Coues

Larus smithsonianus Coues, Proceedings Academy Nat. Sci., Philadelphia,

1862, p. 296. (Eastern and western coasts of North America.)
Herring Gull

Range. North America in arctic and northern temperate zones;
south in winter, sometimes to northern tropics.

Recorded in Hawaiian Group (as an accidental visitor) from Laysan.

Larus californicus Lawrence

Larus californicus Lawrence, Annals Lyceum Nat. Hist., New York, 6, 1854,

p. 79. (Near Stockton, California.)
California Gidl

Range. Northwestern North America from the Mackenzie River
east to Saskatchewan, south to west central California and north-
western Wyoming; south in winter to southern California and western
Mexico.

Recorded in the Hawaiian Group (as an accidental visitor) from Maui
or Hawaii (Bryan, 1901).

Larus pipixcan Wagler

Larus pipixcan Wagler, Isis von Oken, 1831, col. 515. (Mexico.)
Franklin's Gull

Larus franklini Swainson and Richardson in Henshaw, Birds of Hawaiian
Islands, p. 127; Bryan, Key Birds Hawaiian Group, p. 6.

Range. Interior of northwestern North America; south in winter
to coasts of the Gulf of Mexico and western coasts of South America.

Recorded in the Hawaiian Group (as an accidental visitor) from Maui
(one record).

Larus glaucescens Naumann

Lams glaucescens Naumann, Naturgeschichte Vogel Deutschlands, etc., 10,

1840, p. 351. ("Nord-Amerika".)
Glaucous-uinged Gull

BRYAX AM) GREENWAT: HAWAIIAN BIRDS 117

Range. Northeastern Asia arid northwestern North America;
south in winter to China and Japan and the Aleutian Islands to Cali-
fornia.

Recorded in the Hawaiian Group (as an accidental visitor) from Lay-
san, Hawaii, "Probably . . . other islands, especially to Oahu" (Hen-
shaw).

This gull sometimes follows ships ta the Hawaiian Islands.

Larus hyperborevs Gunnerus

Larus hyperboreus Gunnerus, in Leem . . . Beskrivelse Finmarkens Lapper,

etc., 1767, p. 226, (footnote). (Northern Norway.)
Glaucous Gull, Point Barrow Gull

Larus glaucus Brunnich, Ornithologia borealis, etc., 1764, p. 44.
Larus barrorianus Ridgway, Auk, 3, 1886, p. 330; in Henshaw, Birds Hawaiian

Islands, p. 126; Bryan, Key Birds Hawaiian Group, p. 6.

Range. Circumpolar in the Arctic; south in winter to coast of
temperate zones, straying sometimes far.

Recorded in Hawaiian Group (as an accidental visitor) from Laysan,
Kauai, Lanai, Maui.

Larus Philadelphia (Ord)

Sterna Philadelphia Ord in Guthrie's New Geographical . . . Grammar, etc.,
2nd American ed., 2, 1815, p. 319. (No locality = near Philadelphia,
Pennsylvania.)

Bonaparte s Gull

Range. Northwestern North America ; winters on the Atlantic and
Pacific coasts.

Recorded in the Hawaiian Group (as an accidental visitor) from Kauai
(Rothschild). '

Genus Rissa — Kitti wakes

Rissa Stephens, in Shaw's General Zoology, 13, pt. 1, 1826, p. 180. Type, by
monotypy, Rissa brunnichii Stephens = Larus tridactylus Linnaeus.

Rissa tridactyla pollicaris Ridgway

Rissa tridactyla pollicaris attributed to Stejneger by Ridgway in Baird,
Brewer and Ridgway, Water Birds North America, 2, 1884, p. 202.
(Kotzebue Sound, Alaska.)

Western Kittiwake

118 bulletin: museum of comparative zoology

Range. Eastern Siberia, Islands of Behring Sea, Alaska; south in
winter to Japan and California.

Recorded in the Hawaiian Group (as an accidental visitor) from
Laysan (fragments in Bishop Museum).

Genus Sterna — Terns

Sterna Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 137. Type, by tau-
tonomy, Sterna hirundo Linnaeus.

Sterna paradisaea Pontoppidan

Sterna Paradisaea Pontoppidan, Danske Atlas, 1, 1763, p. 622. (Christansoe,

Denmark ex Briinnich 1764.)
Arctic Tern

Range. Circumpolar in Arctic and parts of northern temperate
zones; south in winter to the Antarctic Ocean.

Recorded in the Hawaiian Group ( as an accidental visitor) from
Oahu, Hawaii (Henshaw).

Sterna sumatrana sumatrana Raffles

Sterna sumatrana Raffles, Transactions Linnaean Soc. London, 13, pt. 2, 1822,

p. 329. (Sumatra.)
Black naped Tern
Sterna melanauchen Temminck in W. A. Bryan, Birds Hawaiian Group, p. 8;

Perkins, Fauna Hawaiiensis (Aves), p. 464; Henshaw, Birds Hawaiian

Islands, p. 124.

Range. Western and southern Pacific and eastern Indian Oceans.
Recorded in the Hawaiian Group (as an accidental visitor) from
Kauai, Hawaii.

Sterna anaethetus lunata Peale

Sterna lunata Peale, U. S. Exploring Expedition, 8, 1848, p. 277. (Vincennes

Island = Kauehi Island, Tuamotus.)
Bridled Tern, Gray backed Tern, Pakalakala
Haliplana lunata in Rothschild, Avifauna of Laysan, etc., v, vii, ix, 37.

Range. Pacific Ocean from the Hawaiian Islands south to the
Moluccas.

Recorded in the Hawaiian Group from Laysan (breeding), ? Lisianski,
Gardner (sight, Rothschild), ? French Frigate Shoal (Fisher ? ex-
Rothschild), Necker, Niihau, Kauai, Oahu, Hawaii.

BRYAN AND GREEXWAY : HAWAIIAN BIRDS ] 1!)

Sterna fuscata oahuensis Bloxam

Sterna Oahiu nsis Bloxam, Voyage "Blonde", 1826, p. 251. (Oahu.)

Wideawake, Sooty Tern, Ewa< wa

Haliplana fuMginosa. (Gmelin) in Rothschild, Avifauna of Laysan, etc., vii,
viii, 39; Sterna fuliginosa (Gmelin) in Wilson andEvans, AvesHawaiiensis-
p. 137; Bryan, Key Birds Hawaiian Group, p. 8; Perkins, Fauna Hawai-
iensis (Aves), p. 464; Henshaw, Birds Hawaiian Islands, p. 122, et al.

Range. Pacific Ocean from the Bonin Islands and the Hawaiian
Islands southward.

Recorded in the Hawaiian Group from Midway (breeding), Pearl and
Hermes Reef, Lisianski, Laysan (breeding), ? Gardner (sight, Roths-
child), French Frigate Shoal (breeding), Necker (breeding), Bird Rock
(Nihoa), Kaula, Kauai, Oahu (Moku Manu).

Genus Procelsterna — Terns

Procelsterna Lafresnaye, Magazin Zoologie, 1842, Oiseaux, pi. 29, p. 1. Type,
by monotypy, Procelsterna tereticollis Lafresnaye = Sterna teretirostris
Lafresnaye.

Procelsterna certjlea saxatilis W. K. Fisher

Procelsterna saxatilis W. K. Fisher, Proceedings U. S. Nat. Mus., 26, 1903,

p. 559. (Necker Island.)
Necker Island Tern

Range. Marcus Island and western Hawaiian Islands.
Recorded in the Hawaiian Group from French Frigate Shoal, Necker
(breeding), Bird Island (Nihoa), Kaula.

Genus Ano us Stephens — Noddies

Anoiis Stephens, in Shaw's General Zoology, 13, pt. 1, 1826, p. 139. Type, by
subsequent designation, Anoiis niger Stephens = Sterna stolida Linnaeus
(Gray, List Genera Birds, 1840, p. 79).

Anous stolidus pileatus (Scopoli)

Sterna pileata Scopoli, Deliciae florae et faunae insubricae, etc., fascic. 2,
1786, p. 92. (No type locality — Philippine Islands ex Sonnerat, Voyage
aux Indes, etc., 1782.)

Noddy

Anous stolidus (Linnaeus) in Rothschild, Avifauna of Laysan, etc., p. 41 ; Wilson
and Evans, Aves Hawaiiensis, p. 141 ; Bryan, Key Birds Hawaiian Group,
p. 9; Perkins, Fauna Hawaiiensis (Aves), p. 464; Henshaw, Birds Ha-
waiian Islands, p. 124.

120 bulletin: museum of comparative zoology

Range. Indian and western Pacific Oceans, Seychelles and Mada-
gascar, Bonin and Hawaiian Islands south to Australia.

Recorded in the Hawaiian Group from Midway (breeding), Laysan
(breeding), French Frigate Shoal, Necker, Bird Island (Nihoa),
Kaula Lehua (breeding), Oahu.

Anous minutus melanogenys G. R. Gray

Anous melanogenys G. R. Gray, Genera Birds, 3, 1846, pi. 182. (No type
locality; figure agrees with Hawaiian bird teste Hartert.)

Hawaiian Tern, Noio

Anous hawaiiensis Rothschild in Rothschild, Avifauna of Laysan, etc., vii, xi,
43, 285; Wilson and Evans, Aves Hawaiiensis, p. 143; Perkins, Fauna
Hawaiiensis (Aves), p. 464; Henshaw, Birds Hawaiian Islands, p. 125;
Microanous hawaiiensis Bryan, Key Birds Hawaiian Group, p. 9; et al.

Range. Hawaiian Islands.

Recorded in the Hawaiian Group from Midway (breeding), Laysan
(breeding), Lisianski (breeding), Necker, Bird Rock (Nihoa), Lehua,
Niihau (sight, Rothschild), Kauai, Oahu (Mokuloa and Kaohi Kaipu)
? Molokai (W. K. Fisher), Hawaii.

Genus Gygis — Fairy Terns

Gygis Wagler, Isis von Oken, 1832, col. 1223. Type, by monotypy, Sterna
Candida Gmelin.

Gygis alba rothschildi Hartert

Gygis alba rothschildi Hartert, Novitates Zoologicae, 34, 1927, p. 18. (Laysan
Island.)

White Tern, Love Bird

Gygis alba (Sparrman) or Gygis Candida Wagler in Rothschild, Avifauna of
Laysan, etc., pp. vii, xiii, xiv, 35, 285; Wilson and Evans, Aves Ha-
waiiensis, p. 145; Perkins, Fauna Hawaiiensis, p. 145; Henshaw, Birds
Hawaiian Group, p. 126; Gygis alba kitllitzi Hartert in Bryan, Key Birds
Hawaiian Group, p. 9; et al.

Range. Islands northwest of Kauai.

Recorded in the Hawaiian Group from Midway (breeding), Laysan
(breeding), Lisianski, French Frigate Shoal, Necker (breeding),
Bird Rock (Nihoa), Kaula.

BRYAN AND GREENWAY: HAWAIIAN UIRDS 121

Order STRIGIFORMES

Family STRIGIDAE

Genus Asio Brisson

Asio Brisson, Ornithologie, 1, 1760, p. 28. Type, by tautonomy, Asio Brisson
= Strix otus Linnaeus.

Asio flammeus sandwichensis (Bloxam)

Strix sandwnchensis Bloxam, Voyage "Blonde", 1826, p. 250. (Hawaiian

Islands.)
Hawaiian Owl, Pueo
Asio acciplrinus in Wilson and Evans, Aves Hawaiiensis, p. 133; Perkins,

Fauna Hawaiiensis (Aves), p. 448.

Range. Kauai (breeding), Oahu (breeding), Molokai (? breeding),
Maui, Hawaii (breeding).

Order CORACIIFORMES

Family ALCEDINIDAE

Genus Ceryle — Kingfishers

Ceryle Boie, Isis von Oken, 1828, col. 316. Type, by subsequent designation ,
Alcedo rudis Linnaeus, Gray, List Genera Birds, 1840, p. 11.

Ceryle alcyon caurina Grinnell

Ceryle alcyon caurina Grinnell, University California Publications, Zool., 6>
1910, p. 388. (Graveyard Point, Montague Island, Prince William
Sound, Alaska.)

Western Belted Kingfisher

Ceryle alcyon (Linnaeus) in Henshaw, Birds Hawaiian Group, p. 77.

Range. Alaska south to southern California; south to northern
and central Mexico in winter.

Recorded in the Hawaiian Group (as an accidental visitor) from
Hawaii.

Order PASSERIFORMES— Perching Birds

Family CORVIDAE

Genus Corvus — Crows

Corvus Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 105. Type, by Linnaean
tantonymy, Corvus corax Linnaeus.

122 bulletin: museum of comparative zoology

Corvus hawaiiensis Peale

Corvus hawaiiensis Peale, U. S. Exploring Expedition, 8, 1848, p. 106. (Kara-
kakua [Kealakekua] Bay, Hawaii.)

Hawaiian Crow, Alala

Corvus tropicus "Linnaeua", Bloxam in Voyage "Bonde", 1826, p. 250 (Sand-
wich Islands) is a nomen nudum. Corvus tropicus Gmelin to which Bloxam
refers is not identifiable.

Corvus tropicus Gm. in Wilson and Evans, Aves Hawaiiensis, p. 1 ; Henshaw,
Birds Hawaiian Possessions, etc., p. 36.

Range. Edges of mountain forest from 1000 to 6000 feet in Kona
and Kau districts of the Island of Hawaii.

Family MUSCICAPIDAE— Flycatchers, Thrushes, Warblers
Subfamily TURDINAE

Genus Phaeornis — Hawaiian Thrushes

Phaeornis Sclater, Ibis, 1, 1859, p. 327. Type, by original designation and
monotypy, Phaeornis obscura (Gmelin).

Phaeornis obscura obscura (Gmelin)

Muscicapa obscura Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 945. "(in

insulis Sandwich.")
Ornao, Oman, Hawaii Thrush, Kamao, A-Maui.

Range. Endemic to Island of Hawaii.

Phaeornis obscura lanaiensis Wilson

Phaeornis lanaiensis Wilson, Annals and Magazine Nat. Hist. (6), 7, 1891 »

p. 460. (Lanai.)
Lanai Thrush, OXomao, Olomau

Range. Mountain forests of Lanai. Now very rare.

Phaeornis obscura rutha W. A. Bryan

Phaeornis rutha W. A. Bryan, Occasional Papers B. P. Bishop Mus., Honolulu,

4, no. 2, 1908, p. 81. (Molokai.)
Molokai Thrush, Olomao

Range. Mountain forests of Molokai.

This poorly marked form averages somewhat larger (wing averages
92 mm. and 95 mm.) and slightly darker than lanaiensis.

BRYAN AND GRKKNWAV: HAWAIIAN BIRDS 123

Phaeornis ? obscurus oahensis Wilson and Evans

Phaeornis oahensis Wilson and Evans, Aves Hawaiiensis (Introduction),

1899, p. XIII (Oahu).
Oahu Thru ah

Range. Formerly mountain forests of Oahu. Now extinct.

Specimens are supposed to have been collected by Andrew Bloxam,
naturalist on H. M. S. 'Blonde', but they have disappeared. Wilson
and Evans described the bird from Bloxam's manuscript notes.

Phaeornis obscura myadestina Stejneger

Phaeornis myadestina Stejneger, Proceed. U. S. Nat. Mus., 10, 1887 (1888),
p. 90. (Kauai.)

Phaeornis myiadestina Wilson and Evans, Aves Hawaiiensis, p. 117; Roths-
child, Avifauna Laysan, p. 63.

Kauai Thrush, Kamau, Kamao

Range. Forests of Kauai.

Phaeornis palmeri Rothschild

Phaeornis palmeri Rothschild, Avifauna Laysan Island, etc., pt. 2, 1893, p. 67.

(Halemanu, Kauai.)
Puaiohi

Range. Region of Halemanu, Kauai. Never recorded from other
parts of the island. Now very rare and possibly extinct.

This bird, because of its voice and habits, which differ from those
of other members of the genus (fide Perkins et al.), and because it
apparently breeds in the same region as myadestina, we must treat
as a distinct species.

Subfamily SYLVIINAE

Genus Acrocephalus- — Miller Birds

Acrocephalus Naumann, Land u. Wasservogel nordl. Deutschland, Nachtrage,
4, 1811, p. 199. Type, by subsequent designation, Sylvia turdoides Meyer
= Acrocephalus arundinaceus (Linnaeus).

We can see no reason for retaining the genus Conopoderas Billberg.
Some of the island forms are closer to the type of Acrocephalus than
to that of Conopoderas.

124 bulletin: museum of comparative zoology

ACROCEPHALUS FAMILIARIS (Rothschild)

Tatare familiaris Rothschild, Annals and Magazine Nat. Hist. (6), 10, 1892,

p. 109. (Laysan Island.)
Laysan Miller Bird

Range. Formerly Laysan Island, now extinct.

Acrocephalus kingi ( Wetmore)

Conopoderas kingi Wetmore, Condor, 26, 1924, p. 177. (Nihoa.)
Nihoa Miller Bird

Range. Nihoa (Bird Rock).

In employing binomials we do not mean to imply that these forms
are distinct species. The present application of specific names is
unsatisfactory. This genus needs revision.

Subfamily MUSCICAPINAE

Genus Chasiempis Cabanis

Chasiempis Cabanis, Archiv fur Naturgeschiehte, 13, 1847, Bd. 1, p. 207.
Type, by original designation, "Muscicapa sandvicensis Latham" =
Chasiempis sandwichensis (Gmelin).

Though Cabanis cited Muscicapa sandvicensis Latham, Latham
himself never used this name. Gmelin 's description of Muscicapa
sandwichensis was taken from Latham's "Sandwich Fly-catcher."

Chasiempis sandwichensis sclateri Ridgway

Chasiempis sclateri Ridgway, Proceed. U. S. Nat. Mus., 4, 1881, (1882), p. 337.

(Waimea, Kauai.)
Kauai Elepaio, Apekepeke, Amakahi, Kahuna-Ka-lai-woa
Chasiempis dolei Stejneger, Proceed. U. S. Nat. Mus., 10, 1887 (1888), p. 90.

(Kauai.)

Range. Forests of Kauai.

Both Hawaiians and scientists were long under the impression that
the white-rumped form (i\makahi or Elepaio) and the brown-rumped
form (Apekepeke) were distinct species. Perkins showed that the
latter is an immature plumage in which the birds sometimes breed.

BRYAN AND GREENWAY: HAWAIIAN BIRDS 125

Chasiempis sandwichensis gayi Wilson

Chasiempis gayi Wilson, Proceed. Zool. Soc. London, 1891, p. 165. (Oahu.)
Oahu Elepaio

Range. Forests of Oahu.

Chasiempis sandwichensis sandwichensis (Gmelin)

Muscicapa sandwichensis Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 945.

("in insulis Sandwich" = Hawaii.)
Hawaii Elepaio, Ono-Ka-Ia
Chasiempis rirfgwayi Stejneger, Proceed. U. S. Nat. Mus., 10, 1887 (1888),

p. 87, in key, p. 89. (Hawaii.)
Chasiempis ibidis Stejneger, Proceed. U. S. Nat. Mus., 10, 1887 (1888), p. 87,

in key, p. 89.

Range. Forests of Hawaii.

Stejneger's Chasiempis ibidis refers to plate 1, figure 2 in the Ibis
of 1S85. This specimen had no locality other than "Chili," as Sclater
remarks.

Subfamily DREPANIDIXAE

Genus Viridonia

Yiridonia Rothschild, Annals and Magazine Nat. Hist. (6), 10, 1892, p. 112.
Type, by monotypy, Viridonia sagittirostris Rothschild.

Viridonia sagittirostris Rothschild

Viridonia sagittirostris Rothschild, Annals and Magazine Nat. Hist. (6), 10,

1892, p. 112. (Lower Hilo slope of Mauna Kea, Hawaii.)
Green Solitaire

Range. Mountain rain forests of the windward side of Hawaii near
the Wailuku River at about 1200 to 4000 feet.

Genus Palmeria

Palmeria Rothschild, Ibis, 1893, p. 113. Type, by original designation and
monotypy, Palmeria mirabilis Rothschild = P. dolei I Wilson).

126 bulletin: museum of comparative zoology

Palmeria dolei (Wilson)

Himatione dolei Wilson, Proceed. Zool. Soc. London, 1891, p. 166. (Maui.)

Akohekohe, Crested honey eater

Palmeria mirabilis Rothschild, Ibis, 1893, p. 113.

Range. Mountain forests of Molokai and Maui. Now very rare.

Genus Himatione

Himatione Cabanis, Museum Heineanum, Th. 1, 1850, p. 99. Type, by subse-
quent designation, Himatione sanguinea (Gmelin).

Himatione sanguinea fraithii Rothschild

Himatione Fraithii Rothschild, Annals and Magazine Nat. Hist. (6), 10, 1892,

p. 109. (Laysan Island.)
Laysan Honey Creeper

Range. Formerly Laysan Island, now extinct.

Himatione sanguinea sanguinea (Gmelin)

Certhia sanguinea Gmelin, Systema Naturae, 1, pt. 1, 1788, p. 479. (Sandwich

Islands.)
Apapane, Akakani

Range. Mountain forests of ?Niihau, Kauai, Oahu, Molokai,
Maui, Hawaii.

Genus Vestiaria

Vestiaria Fleming, Philosophy of Zoology, 2, 1822, p. 246. Type, by original
designation, Certhia vestiaria = Vestiaria coccinea (Forster).

Vestiaria coccinea coccinea (Forster)

Certhia coccinea Forster, Goettingisches Magazin Wissenschaften, etc., 1,

1780, p. 347.
Iiwi, Iiwi popolo, Iiwi polena (for various phases of plumage).
Vestiaria coccinea suavis Bangs, Proceed. Biol. Soc. Wash., 24, 1911, p. 29.

(Molokai.)

Range. Mountain forests of Kauai, Oahu, Molokai, Maui, Lanai,
Hawaii.

BRYAN AND GREENWAY: HAWAIIAN BIRDS 127

Tor full synonymy see Rothschild, Avifauna Laysan Island, etc.

Vestiaria coccinea suavis Bangs was described as paler, with a larger
bill. In our opinion, the former is an individual variation and the latter
is non-existent. In our short series (28) there is an average difference
of 2 mm. in wing length, but this is not sufficient for formal recognition.

Genus Drepaxis — Hawaiian Honey Creepers

Drepanis Temminck, Manual d'Ornithologie, ed. 2, 1, 1820, p. LXXXVI .
Type, by subsequent designation (Gray, 1840), Certhia pacifica Gmelin.

Drepaxis pacifica (Gmelin)

Certhia pacifica Gm., Systema Naturae, 1, pt. 1, 1788, p. 470. ("in insulis

amicis" in error ex Latham. Hawaiian Islands.)
Mamo, Hoha or Hoho (so rendered by older writers).

Range. Formerly mountain forests of Hawaii (old records for
Kauai are almost certainly in error). Now extinct; last specimen seen
at Kaumana 1000-1500 feet above Hilo, 1898.

Drepaxis fuxerea Newton

Drepanis funerea Newton, Proceed. Zool. Soc. London, p. 690. (Molokai.)

Ma?no, Oo-nuku-umu, Hoa, Black Mamo, Perkins' Mamo

Drepanorhamphus funerea Newton in Rothschild, Avifauna Laysan Island,

etc., p. 165; Bryan, Birds Hawaiian Group, p. 42; Perkins, Fauna Ha-

waiiensis (Avesj, p. 402.

Range. Mountain forests of Molokai.

The genus Drepanorhamphus was established by Rothschild (Avi-
fauna Laysan, etc., pt. 3, 1900, p. 163. Type, by monotypy, Drepanis
funerea Newton). The characters that separate it from Drepanis are
the lack of the long, decomposed, yellow under tail coverts and the
shape of the nostril which is long and narrow, not rounded. It seems
to us that they are alike in so many ways that they form a single genus
separating them from other genera, but that they were probably dis-
tinct species.

Genus Hemigxathus — Akialoa

Hemignathus Lichtenstein, Abhandlungen Konig. Akademie Wissensch
Berlin, 1839, p. 449. Type, by subsequent designation, Hemignathus
lucidus Licht. (Gray, List Genera Birds, 1841, p. 16).

128 bulletin: museum of comparative zoology

In our opinion, the group with long lower mandibles and those with
short lower mandibles may well be considered as congeneric. If it is
desired to separate them generically then the former will require a
new generic name and the latter (heretofore known as Heterorhynchus
will have to be called Hemignathus, since the two groups, as named
heretofore, have the same type as designated by Gray and Lafresnaye.
Rothschild's arguments (Avif. Laysan Id., etc., p. 79) have no force
under the rules of zoological nomenclature (Art. 30, II, g).

Hemignathus obscurus procerus Cabanis

Hemignathus procerus Cabanis, Journal f. Ornith., 1889, p. 331. (Kauai.)
Kauai Akialoa, "Iiwi"

Range. Mountain forests of Kauai from the lowest forest zone
600-900 feet to 4000 feet.

Hemignathus obscurus ellisianus (Gray)

Drepanis {Hemignathus) ellisiana Gray, Cat. Birds Tropical Islands Pacific,

1860, p. 9. (Oahu.)
Oa.hu Akialoa, Iiwi (Jibi or Kipi)
Hemignathus lichtensteini Wilson in Aves Hawaiiensis, p. 65, and Perkins,

Fauna Hawaiiensis (Aves), p. 425.

Range. Formerly mountain forests of Oahu; now extremely rare;
perhaps confined to Waianae Mountains (Northwood).

Hemignathus obscurus lanaiensis Rothschild

Hemignathus lanaiensis Rothschild, Bull. Brit. Ornith. Club, 1, 1893, p. xxiv.

(Lanai.)
Lanai Akialoa

Range. Formerly mountain forests of Lanai, now very rare, pos-
sibly extinct.

Hemignathus obscurus obscurus (Gmelin)

Certhia obscura Gm., Systema Naturae, 1, pt, 1, 1788, p. 470. (Sandwich

Islands = Hawaii.)
Haivaii Akialoa

Range. Formerly mountain forests of Hawaii, now probably con-
fined to localized areas in the rain forests of the windward side (see
Baldwin, 1941, p. 19).

BRYAN AND (iKKF.NW AY: HAWAIIAN BIRDS 129

Hemignathus lucidus iiaxapepe Wilson

Hemignathus hanapepe Wilson, Annals Magazine Nat. Hist. (6), 4, 1889,

p. 401. (Kauai.)
Nukupu'u
Heterorhgnrhus hanapt /« in Rothschild, Avifauna Laysan, etc., p. 101 ; Perkins,

Fauna Hawaiiensis (Aves), p. 430; Henshaw, Birds Hawaiian Islands,

p. 43.

Range. Mountain forests of Kauai in Waimea district near the
Hanapepe River, 2000 to 3000 feet, very rare and local.

Hemignathus lucidus lucidus Lichtenstein

Hemignathus lucidus Lichtenstein, Abhandlungen der Konigl. Academie
Wissensch. Berlin, 1S39, p. 451, pi. 5, figs. 2, 3. (Oahu.)

Oahu Akiapolaau

HeterorhyncJms lucidus in Rothschild Avifauna Laysan, etc., p. 105; Perkins,
Fauna Hawaiiensis (Aves), p. 430; Henshaw, Birds Hawaiian Islands,
p. 41.

Akiapolaau

Range. Formerly mountain forests of Oahu ; now almost certainly
extinct.

Ten specimens were thought to exist in museums but there are not
that many, for some have been misidentified.

Hemignathus lucidus affinis Rothschild

Hemignathus affinis Rothschild, Ibis, 1893, p. 112. (Maui.)
Maui Akiapolaau

Range. Recorded from forested slopes of Haleakala Mountain,
Maui, 4000 to 4500 feet. Now local and probably very rare.

Hemignathus lucidus wilsoni (Rothschild)

Heterorhynchus wilsoni Roths., Avifauna Laysan, etc., 1893, pt. II, p. 97.

(Hawaii).
Akiapolaau
Hemignathus olivaceus Lafr. in Wilson and Evans, Aves Hawaiiensis, p. 75.

Range. Mountain forests of Hawaii from about 1500 to 6700 feet.
Still exists but not common.

130 bulletin: museum of comparative zoology

Genus Chlorodrepanis — Amakihi

Chlorodrepanis Perkins in Wilson and Evans, Aves Hawaiiensis, pt. 7, p. xxi
(introduction), June 1899. Type, by subsequent designation, Chlorodre-
panis stejriegeri Wilson (Richmond, Proceed. U. S. Nat. Mus., 24, 1902,
p. 673).

Chlorodrepanis parva (Stejneger)

Himatione parva Stejneger, Proceedings U. S. Nat. Mus., 10, 1887, p. 94.

(Kauai.)
Alawi, Anauanii, Anianau
Oreomyza parva in Rothschild, Avifauna Laysan, etc., p. 119.

Range. Forests of Kauai.

This is a very curious little bird for which Wilson and Evans pro-
posed the name Rothschildia (pre-occupied). Mathews in 1925 re-
named it Magumma. Our treatment is based on that of Perkins
(Ibis, 1895 and Fauna Hawaiiensis (Aves), p. 411).

Chlorodrepanis virens stejnegeri (Wilson)

Himatione stejnegeri Wilson, Proceedings Zool. Soc. London, 1889 (1890),

p. 446. (Kauai.)
Kauai Amakihi

Range. Forests of Kauai.

Chlorodrepanis virens wilsoni (Rothschild)

Himatione wilsoni Rothsch., Bull. Brit. Ornith. Club, 1, 1893, p. xliii. (Maui.)

Molokai Amakihi, Lanai Amakihi, Maui Amakihi

Himatione kalaana Wilson and Evans, Aves Hawaiiensis, 1896, p. 28. (Molo-
kai.)

Himatione chloridoides Wilson and Evans, Aves Hawaiiensis, 1896, p. 28.
(Lanai.)

Range. Forests of Molokai, Lanai, Maui.

This form is very close to chloris but differs in having a slightly
larger bill and (in mature specimens) it has a slightly paler back.
Differences between specimens from Molokai, Lanai and Maui alleged
to exist (an ill-defined or well-defined yellow stripe from bill to eye as
well as the shade of green on the back) fall within the range of indi-
vidual variation.

BRYAN AND GREENWAT: HAWAIIAN BIRDS 131

Chlorodrepanis virens virens (Gmelin)

Certhia virens Gm., S}'stema Naturae, 1, pt. 1, 17S8, p. 479. (in insulis Sand-
wich = Hawaii.)
Hawaii Amakihi

Range. Forests of Hawaii.

Genus Loxops — Akakane

Loxops Cabanis, Archiv fur Naturgeschichte, Berlin, 13, 1847, Bd. 1, p. 330.
Type, by original' designation, Fringilla coccinea Gm. For synonymy see
Rothschild, Avifauna Laysan, etc., p. 167.

Loxops caeruleirostris (Wilson)

Chrysomitridops caeruleirostris Wilson, Proceedings Zool. Soc. London, 1889

(1890), p. 445. (Kauai.)
Ou-holoicai, Akekee (fide Perkins)
Chrysomitridops caeruleirostris in Wilson and Evans, Aves Hawaiiensis, p. 59.

Range. Forests of Kauai.

Loxops coccinea rufa (Bloxam)

Fringilla rufa Bloxam in Voyage "Blonde", 1826, p. 250. (Oahu.)
Akepa, Akepeuie

"Loxops wolstenkolmei" Roths. — the plate so lettered in Rothschild Avifauna
Laysan, etc., p. 177.

Range. Formerly forests of Oahu, now very rare and perhaps
extinct. Last record May 20, 1S93.

Loxops coccinea ochracea Rothschild

Loxops ochracea Rothschild, Ibis, 1893, p. 112. (Maui.)
Ochraceus or Maui Akt peuie

Range. Forests of Maui.

Loxops coccinea coccinea (Gmelin)

Fringilla coccinea Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 921. (in insulis

Sandwich = Hawaii.)
Akepa, Akepeuie

Range. Forests of Hawaii. "Now rare" (Baldwin).

132 bulletin: museum of comparative zoology

Genus Paroreomyza

Paroreomyza Perkins, Ibis, 1901, p. 583. Type, by original designation,
Himatione maculata Cabanis.

In our opinion, the forms formerly listed under this name and those
listed under the name Oreomystis Stejneger 1903 are congeneric. It is
true that females of the former lack the color of the males, but the
birds are otherwise alike from a generic point of view.

Paroreomyza bairdi bairdi (Stejneger)

Oreomyza bairdi Stejneger, Proceedings U. S. Nat. Mus., 10, 1887 (1888), p. 99.

(Kauai.)
Akikihi
Oreomyza bairdi in all works on Hawaiian birds.

Range. Forests of Kauai above 1000 feet.

Paroreomyza bairdi mana (Wilson)

Himatione mana Wilson, Annals and Magazine Nat. Hist. (6), 7, 1891, p. 460.

(Hawaii.)
Hawaii Creeper, Olive-green Creeper
Oreomyza mana in all works on Hawaiian birds.

Range. Forests of Hawaii except (fide Perkins) in lower elevations
on the Kona or leeward coast. Now uncommon (fide Baldwin 1941).

In our opinion "Oreomyza perkinsi" Rothschild is in all probability
a hybrid, Chlorodrepanis virens x Paroreomyza mana. It has been listed
with other hypothetical forms and records.

Paroreomyza maculata maculata (Cabanis)

Himatione maculata Cabanis, Museum Heineanum, Th. 1, 1850, p. 100, in

footnote. (Oahu.)
Oahu Creeper
Himatione maculata Cabanis in Wilson and Evans, Aves Hawaiensis, p. 43;

Oreomyza maculata in Rothschild, Avifauna Laysan Island, etc., p. 113;

W. A. Bryan, Key Birds Hawaiian Group, p. 48; Perkins, Fauna Ha-

waiiensis, (Aves) p. 417; Henshaw, Birds Hawaiian Islands, p. 50.

Range. Mountain forests of Oahu above 1500 feet. "Scarcest of
the five native perching birds which are likely to be seen." (North-
wood 1940).

BRYAN AND OKEENWAY : HAWAIIAN BIRDS 133

Paroreomyza maculata flammea (Wilson)

Loxops flarnmea Wilson, Proceedings Zool. Soc. London, 1889 (1890), p. 445.
(Kalae, Molokai.)

Kakawahie

Loxops flammea Wilson in Wilson and Evans, Aves Hawaiiensis, p. 39; Oreomyza
flammea Wilson in Rothschild, Avifauna Laysan Island, etc., p. 121;
W. A. Bryan, Key Birds Hawaiian Group, p. 48; Perkins, Fauna Hawaii-
ensis (Aves), p. 417; Henshaw, Birds Hawaiian Islands, p. 49.

Range. Mountain forests of Molokai above 1500 feet.

Paroreomyza maculata Montana (Wilson)

Himatione montana Wilson, Proceedings Zool. Soc. London, 1889 (1890), p.
446. (Lanai.)

Alauhiio

Himatione montana Wilson in Wilson and Evans, Aves Hawaiiensis, p. 45;
Oreomyza montana in Rothschild, Avifauna Laysan Island, etc., p. 117;
W. A. Bryan, Key Birds Hawaiian Group, p. 47; Perkins, Fauna Hawaii-
ensis (Aves), p. 417; Henshaw, Birds Hawaiian Islands, p. 51.

Range. Mountain forests of Lanai.

Paroreomyza maculata newtoni (Rothschild)

Himatione newtoni Rothschild, Bull. British Ornith. Club, 1, 1893, p. xlii.

(Maui.)
Maui Creeper
Himatione newtoni Rothschild in Wilson and Evans, Aves Hawaiiensis, p. 11;

Oreomyza newtoni Rothschild in Rothschild, Avifauna Laysan Island, etc.

p. 115; W. A. Bryan, Key Birds Hawaiian Group, p. 48; Perkins, Fauna

Hawaiiensis (Aves), p. 417; Henshaw, Birds Hawaiian Islands, p. 49.

Range. Mountain forests of Maui.

Genus Ciridops

Ciridops A. Newton, Nature, 45, 1892, p. 469. Type, by monotypy, Fringilla
anna Dole.

Ciridops anna (Dole)

Fringilla anna Dole, Hawaiian Almanac, 1879, p. 49. (Hawaii.)
Waaihaiuane

134 bulletin: museum of comparative zoology

Range. Formerly mountain forests of Hawaii. Recorded from
Kona and Hilo districts in or near Loulu or Hawane palms (Prit-
chardia). Never common within the memory of man, it is now prob-
ably extinct. Last specimen taken February 1892 on the Kohala
Mountains.

Genus Pseudonestor

Pseudonestor Rothschild, Bulletin British Ornith. Club, 1, 1893, p. xxxv.
Type, by monotypy, Pseudonestor xanthophrys Rothschild.

Pseudonestor xanthophrys Rothschild

Pseudonestor xanthophrys Rothschild, Bulletin British Ornith. Club, 1, 1893,

p. 36. (Maui.)
ParroUbilled Koa Finch

Range. Maui; confined to Haleakala above 5000 feet. Rare.

Genus Psittirostra

Psittirostra Temminck, Manuel d'Ornithologie, etc.', 1, 1820, p. 70. Type, by
monotypy, Loxia psittacea Gmelin.

This name has often been emended to Psittacirostra; the original
spelling is, however, as above.

Psittirostra psittacea psittacea (Gmelin)

Loxia psittacea Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 844. (in insulis
Sandwich = Hawaii.)

Ou

Psittacirostra psittacea oppidana Bangs, Proceedings Biol. Soc. Washington,
24, 1911, p. 30. (Molokai.)

Examination of large series has convinced us that characters alleged to differen-
tiate this form from psittacea (paleness, size) fall within the range of
individual variation.

Dysmorodrepanis munroi Perkins (see list of hypothetical forms).

Range. Forests of Molokai, Lanai, Maui, Hawaii below 5000 feet.
Now very rare, possibly extinct on Lanai.

BRYAN AND GREEXWAY : HAWAIIAN' BIRDS 135

?PsiTTIR0STRA PSITTACEA DEPPEI Rothschild

PsiUirostra psittacea deppei Rothschild, Bull. Brit. Ornith. Club, 15, 1905-,
p. 45. (Oahu.)

Oahu Ou

Psittacirostra psittacea (Gmelin) (part) in Wilson and Evans, Aves Hawaiiensis,
p. 80; PsiUirostra olivacea Rothschild in Rothschild, Avifauna Laysan
Island, etc., p. 193; W. A. Bryan, Key Birds Hawaiian Group, p. 54
(note); Henshaw, Birds Hawaiian Islands, p. 66; Perkins, Fauna Ha-
waiiensis (Aves), p. 435.

Range. Formerly forests of Oahu, now extinct.

There is great individual variation in adults of the populations of this
bird on every island. Particularly, there is a tendency toward albinism
noticeable in small populations. Rothschild describes this form as
differing from others in having the middle of the breast, belly, feathers
of the tibia and under tail coverts buffy whitish, in having a shorter
wing. In view of the fact of this known tendency toward albinism and
that Rothschild's own measurement for the wing of deppei (3.75
inches = 97 mm.) falls exactly within the size range of psittaeea
(95-100 mm.), we think the validity of this subspecies is questionable.

Genus Loxioides — Palila

Loxioides Oustalet, Bulletin Soc. Philom. Paris, ser. 7, 1, 1877, p. 99. Type, by
monotypy, Loxioides bailleui Oustalet.

Loxioides bailleui Oustalet

Loxioides bailleui Oustalet, Bulletin Soc. Philom. Paris, ser. 7, 1, 1877, p. 100.

(Hawaii.)
Palila
Loxioides bailleui Oustalet in all works relating to Hawaiian birds.

Range. Upper forest zones of Hawaii 4000 to 7000 feet in Kona and
Hamakua districts. Usually in or near mamane trees (Sophora).

Genus Rhodacanthis — Koa Finch

Rhodacanthis Rothschild, Annals and Magazine Nat. Hist. (6), 10, 1892, p.
110. Type, by subsequent designation, Rhodacanthis palmeri Rothschild.

136 bulletin: museum of comparative zoology

Rhodacanthis palmeri Rothschild

Rhodacanthis Palmeri Rothschild, Annals and Magazine Nat. Hist. (6), 10,

1892, p. 111. (Kona, Hawaii.)
Orange Koa Finch

Range. Above 4000 feet in mountain forests of Kona and Kau
districts, Hawaii. Usually in Koa trees (Acacia Koa). Now very rare,
possibly extinct. Last specimens collected (1892 or 1893) by Perkins.

?Rhodacanthis flaviceps Rothschild

Rhodacanthis flaviceps Rothschild, Annals and Magazine Nat. Hist. (6), 10'

1892, p. 111. (Kona, Hawaii.)
Yellow-headed Koa Finch

Range. Presumably the same as Rhodacanthis palmeri with which
it was associated when collected by Palmer for Rothschild.

These specimens differ in having yellow, not red, heads and in being
smaller (wing 94-95 mm.). No specimens have since been found.
Perkins (Fauna Hawaiiensis (Aves), pp. 436-437) suggests that these
are not two distinct species but that only one dimorphic form is in-
volved. Since the genus is now very rare it is probable that this
problem can never be solved.

Genus Telespyza

Telespyza Wilson, Ibis, 1890, p. 341. Type, by monotypy, Telespyza cantans
Wilson.

Telespyza cantans cantans Wilson

Telespyza cantans Wilson, Ibis, 1890, p. 341. (Midway Island in error =

Laysan Island.)
Lay san Finch
Telespyza flavissima Rothschild, Annals and Magazine Nat. Hist. (6), 10,

1892, p. 110. (Laysan.)

Range. Formerly on Laysan Island, where now extinct. Midway
Island, where introduced.

Telespyza cantans ultima W. A. Bryan

Telespiza ultima W. A. Bryan, Auk, 34, 1917, p. 71. (Nihoa.)
Nihoa Finch

Range. Nihoa.

BRYAN AND GREEXWAY: HAWAIIAN BIRDS 137

Genus Chloridops

Chloridops Wilson, Proceedings Zool. Soc. London, 1888, p. 218. Type, by
monotypy, Chloridops kona Wilson.

Chloridops kona Wilson

Chloridops kona Wilson, Proceedings Zool. Soc. London, 1888, p. 218. (Kona,

Hawaii.)
Kona Finch, ? Palila

Range. Kona district of Hawaii, in mountain forests from 3500 to
5500 feet. Never abundant, now very rare, perhaps extinct.

Family MELIPHAGIDAE
Genus Moho

Moho Lesson, Traite d'Ornithologie, livr. 4, 1830, p. 302. Type, by monotypy,
Merops fasciculatus Latham = Moho nobilis Merrem.

Mohohina Mathews, Bulletin British Ornith. Club, 45, 1925, p. 93. Type, by
original designation, Acridocercus bishopi Rothschild. "Differs from
Moho Lesson in having plumes on the side of the face" (Mathews).

Pseudomoho Mathews, Bulletin British Ornith. Club, 45, 1925, p. 93. Type, by
original designation, Mohoa braccatus Cassin. "Differs from Moho Lesson
in having quite a different tail formation and in not having any orna-
mental plumes" (Mathews).

Mohornis Mathews, Systema Avium Australasianarum, pt. 2, 1930, p. 800.
Type, by original designation, Moho apicalis Gould.

Moho nobilis braccatus (Cassin)

Mohoa braccata Cassin, Proceedings Acad. Nat. Sci. Philadelphia, 7, 1855,

p. 440. (Kauai.)
O-O, A-A
Acridocercus braccatus Cassin in Wilson and Evans, Aves Hawaiiensis, p. 99;

Perkins, Fauna Hawaiiensis (Aves), p. 445.

Range. Forests of Kauai. Now rare.

Moho nobilis apicalis Gould

Moho apicalis Gould, Proceedings Zool. Soc. London, 1860, p. 380. (Owhyie

in error = Oahu.)
Oahu 0-0
Acridocercus apicalis Gould in Wilson and Evans, Aves Hawaiiensis, p. 103;

Perkins, Fauna Hawaiiensis (Aves), p. 445.

138 bulletin: museum of comparative zoology

Range. Formerly forests of Oahu, now extinct. Last specimen
collected by Herr Deppe in 1837.

Moho nobilis bishopi (Rothschild)

Acrulocercus bishopi Rothschild, Bulletin British Ornith. Club, 1, 1893, p. xli-

(Molokai.)
Bishop's 0-0
Acrulocercus bishopi Rothschild in Wilson and Evans, Aves Hawaiiensis, p. Ill ;

Perkins, Fauna Hawaiiensis (Aves), p. 445.

Range. Forests of Molokai. Now rare.

Moho nobilis nobilis (Merrem)

Gracula nobilis Merrem, Avium rar . . . Icones . . ., 1, 1786, p. 7. (Hawaii.)

(fide Davies-Sherborne)
0-0 '
Acrulocercus nobilis Merrem in Wilson and Evans, Aves Hawaiiensis, p. 105;

Perkins, Fauna Hawaiiensis (Aves), p. 445.

Range. Formerly forests of Hawaii. Now probably extinct.

Genus Chaetoptila

Chaetoptila Sclater, Ibis, 1871, p. 358. Type, by original designation, Entorniza
? angustipluma Peale.

Chaetoptila angustipluma (Peale)

Entorniza angustipluma Peale, U. S. Exploring Expedition, 8, 1848, p. 147.

(Hawaii.)
Kioea (?)

Range. Formerly mountain forests of Hawaii. Now extinct.
Recorded from "between the lower Volcano House and the crater of
Kilauea" where Mills collected his specimens (fide Rothschild who
quotes Palmer in Birds Laysan Island, etc., p. 216).

"District of Hilo near Olaa" (fide Wilson and Evans, Aves Hawaii-
ensis, p. 114).

. . . "supposed to have come from Olaa, it is very doubtful whether
the Chaetoptila was found there. It is much more probable that it
was confined to the high plateau between the mountains and the
upper edges of the forest bordering this, where it was observed by
Pickering and Peale of the U. S. Exploring Expedition" (Perkins,
Fauna Hawaiiensis (Aves), p. 445). All these speculations are open
to question.

BRYAN AND GREENWAY: HAWAIIAN' BIRDS 139

Part II. HYPOTHETICAL FORMS AND RECORDS

Family ARDEIDAE— Herons

Demigretta sacra (Gmelin)

Ardea sacra Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 640.

Range. Asiatic coasts and islands of the Pacific.
Recorded in the Hawaiian Group from Kahalui, Maui (a single sight
record by Finsch).

Family LARIDAE— Gulls

What is now thought to be a hybrid Larus hyperboreus Gunnerus x
Larus argentatus vegae Palmen is recorded as Larus nelsoni Henshaw
(Auk, 1, 1SS4, p. 250) by Oberholser (Auk, 35, 1918, p. 349) from
Hilo, Hawaii.

The specimen is said to be a young female and is in the U. S. National
Museum, Washington.

Subfamily DREPANIDINAE

Paroreomyza perkinsi (Rothschild)

Oreomyza perkinsi Rothschild, Avifauna Laysan Island, etc., pt. 3, 1900, p.

129. (Puulehua, Hawaii.)
Perkins's Creeper

Range. Known from a single specimen, the type, now in the
American Museum of Natural History, New York.

This specimen is intermediate in color and form of bill and is, in
our opinion, a hybrid Chlorodrepanis virens x Paroreomyza mana.
Although the feeding habits of these two birds differ, we do not think
that this would prevent occasional breeding, Perkins to the contrary
(Fauna Hawaiiensis (Aves), p. 417).

Genus "Sassius" Rothschild and Hartert

Sassius Rothschild and Hartert, Bulletin British Ornith. Club, 46, 1926, p. 51.
Type, by monotypy, Sassius simplex Rothschild and Hartert.

"Sassius simplex" Rothschild and Hartert

Sassius simplex Rothschild and Hartert, Bulletin British Ornith. Club, 46
1926, p. 51. (Sandwich Islands.)

140 bulletin: museum of comparative zoology

This curious little skin was discovered by Dr. Hartert in the
Naturhistorisches Museum in Vienna. He thought it to be an un-
described Drepanid at the time, but since then other ornithologists
have thought it to be more probably an artefact made of the skins
of Sun Birds (Nectariniidae), an African and Asiatic family (see
Meise, Proceedings 8th Internat. Orn. Congress, 1934, p. 123).

Genus "Dysmorodrepanis"

Dysmorodrepanis Perkins, Annals and Magazine Nat. Hist. (9), 3, 1919, p. 250.
Type, by monotypy, Dysmorodrepanis munroi Perkins.

"Dysmorodrepanis munroi" Perkins

Dysmorodrepanis munroi Perkins, Annals and Magazine Nat. Hist. (9), 3,
1919, p. 251. (Kaiholena Valley, Lanai.)

This curious bird is, in our opinion, an aberrant specimen of
Psittirostra psittacca (see Greenway, Auk, 56, 1939, p. 479). The type
and only specimen is in the B. P. Bishop Museum, Honolulu.

THE GENERIC ARRANGEMENT OF THE DREPANIDINAE

By James C. Greenway, Jr.

The inter-relationships of the genera of Drepanidinae have by
some been considered solved by the work of R. C. L. Perkins Fauna
Hawaiiensis (Aves) whose arrangement Sharpe followed In his Hand-
List. Perkins' arrangement, however, is not entirely satisfactory,
for in some cases he has used characters which form what appear to
be, even on the evidence he himself uses, rather unnatural groups.

There are no morphological characters to prove that these genera
are properly placed in a single subfamily, and Perkins' and Gadow's
works, which are classical, are at the same time not convincing, since
these birds have no single character in common. However, the ex-
tremely divergent genera are connected by intermediates, and the
hypothesis cannot be disproved. Whether the group sprang from a
"Coerebine" or a "Tanagrine" stock, or both, is not known. It is
therefore useless to speculate about the specialization of forms.

Within the subfamily we have two groups which may be called
"Coerebine" and "Tanagrine", simply because they resemble super-
ficially these groups. The first have tubular tongues with well devel-

BRYAN AND GREENWAY: HAWAIIAN BIRDS 141

oped brush-like tips. This group includes Viridonia, Palmeria,
Himatione, Vestiaria, Drepanis, Hemignathus, Heterorhynchus,
Loxops and Chlorodrepanis. At this point there is a sharp demar-
cation line, for the tongues of the following genera are not tubular,
those of Paroreomyza and Ciridops being concave and with a modified
brush (from this point of view intermediate). Through Psittirostra
there is gradation to the fleshy tongues of Loxioides, Rhodacanthis
and Telespiza, which are the markedly "Tanagrine" types. In spite
of the sharp difference in the shape of the tongue there is no other
character which indicates a break in the continuity of intermediate
forms, Chlorodrepanis and Paroreomyza being so close that, were it
not for the tongue, they would be considered to be congeneric under
present concepts of generic limits. Loxops, too, is intermediate,
having the bill and general form reminiscent of a goldfinch but with
a typically Drepanine tongue From two points of view this, is rather
an aberrant genus, for the shape of bill and tongue do not conform
as in other genera, and the bill with its "loxian twist" is of course
very curious. According to Perkins, it is a convergent parallelism of
both habit and form.

Perkins' arrangement groups Drepanis, Vestiaria, Ciridops, Palmeria
and Himatione together, and this assemblage he characterizes as
follows: "The genera of the first group are characterized by the
truncate apices of the primaries, except in the anomalous Palmeria,
and by the plumage of the young which is always partly black or of
a dull colour." This is an unnatural grouping and cannot be accepted.
A careful examination of these forms reveals the fact that only
Vestiaria and Himatione have truncated primaries. Furthermore,
adults of Telespiza ultima are melanistic and the melanism in cantons
is quite as impressive as in Vestiaria, Perkins to the contrary. Melan-
ism, in any event, is a character of doubtful value, such widely diver-
gent groups as Coereba and Charmosyna being classic examples of
melanistic mutation.

Although no linear arrangement is satisfying, it would appear to
be much more natural to place the tubular ("Coerebine") and the
more fleshy tongued ("Tanagrine") Drepanidinae at opposite ends of
the list, with the intermediate genera, Chlorodrepanis, Loxia, Parore-
omyza and Ciridops between them. Hemignathus is distinctly
"Coerebine" and should be placed close to Drepanis. Ciridops is of
course a very curious bird. The tongue is, however, intermediate,
and even though the color pattern (black wings and tail) resembles
Vestiaria slightly, as Perkins remarks, it would probably be better

142 bulletin: museum of comparative zoology

placed as an intermediate rather than with the typical Drepanidinae.
Ideally, Chlorodrepanis might head the list, leading us from the
Coerebinae into the Drepanidinae.

As in the "Coerebine" forms, so in the "Tanagrine" forms we are
gradually led from a tongue which in Pseudonestor approaches a
tubular form through Psittirostra, in which the Drepanine form is
still observable, to Loxioides, Rhodacanthis, Telespyza and Chloridops.
I have not been able to find any information about the tongues of the
last two genera, but Gadow places them (p. 246) near Rhodacanthis
which, he remarks, has the "most compact" tongue of all. There is
no doubt about the fact (which Gadow points out) that the narinal
structure follows, in effect, a parallel course.

It may well be argued that the last four genera should be lumped
in a single genus, as Rothschild suggests. Loxioides is the oldest name.
Lack of information about the tongues of these genera deters me
from this course.

No new facts are presented here, the anatomical information having
been presented by Gadow (in Wilson and Evans, Aves Hawaiiensis),
Rothschild and Perkins. I have examined tongues of Paroreomyza,
Chlorodrepanis, Himatione and Psittirostra. The interpretation of
evidence collected by Perkins, since accepted, seems to be faulty,
and a, to me, more natural linear arrangement, suggested by Gadow,
follows :

Viridonia

Palmeria

Himatione

Vestiaria

Drepanis

Hemignathus

Chlorodrepanis

Loxops

Paroreomyza

Ciridops

Pseudonestor

Psittirostra

Loxioides

Rhodacanthis

Telespyza

Chloridops

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 3

OBSERVATIONS ON CHINESE GOMPHINE
DRAGONELIES

By James G. Needham

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

June, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1, 2
have appeared and of the Memoirs, Vol. I to LVI.

These publications are issued in numbers at irregular intervals.
Each number of the Bulletin and of the Memoirs is sold separately.
A price list of the publications of the Museum will be sent upon
application to the Director of the Museum of Comparative
Zoology, Cambridge, Massachusetts.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 3

OBSERVATIONS ON CHINESE GOMPHINE
DRAGONFLIES

By James G. Needham

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

June, 1944

No. 4 — Observations on Chinese Gomphine Dragonflies
By James G. Needham

Chinese dragonflies first caught my attention in 1927, when I
received an unanticipated invitation to spend a year in China visiting
and conferring with departments of Biology in the Universities of that
country under the auspicies of the China Foundation for the Promo-
tion of Education and Culture. I looked about for aids to the study of
the local dragonflies and found there were none. There were only bare
descriptions of the adults of many species, printed in half a dozen
languages, and well scattered through the zoological literature of the
world. Nothing was known of the immature stages.

Since I was invited to lend what aid I might to the study of biology
in China, I conceived the idea that by supplying a manual for the
study of the one group of insects with which I already had some prac-
tical acquaintance at home, I might help Chinese students in the study
of their local fauna. Indeed, I was quite sure that such aid might be
more real and the results more lasting, than any that might come from
merely lecturing to them. So I began at once gathering together the
materials for a manual, going in the field to collect dragonflies as I had
opportunity, and enlisting the aid of local collectors wherever possible.

I started with no collection at all, and with almost no personal ac-
quaintance with the Oriental Odonate fauna. The literature of the
group was largely lacking in that country. Wherefore, such time as
could be taken from teaching and conferences, I devoted to collecting
and studying dragonflies.

After my arrival in September I did a little collecting of autumnal
species in and about Peiping, with the invaluable aid of Dr. Chen-fu
Wu of Yen Ching University, and Dr. Chung-lo Liu of Tsing Hua
University, and their advanced students. They guided me to the best
collecting places in some of the most beautiful aquatic situations in
North China. During the winter I was able to work on my collections
in the private laboratory of my good friend, Dr. N. Gist Gee, who was
himself a lifelong student of the Chinese fauna, and a distinguished
specialist in fresh-water sponges. In the spring he went with me to the
Yang-tze valley; to Soochow and to Hang Chow Universities where
other generous collaborators were found. I spent the month of April
in Nanking; that was my real harvest season. In Nanking I had the
generous assistance of Dr. C. Ping and a number of his research stud-
ents. There I had considerable time for collecting adult dragonflies,
and for working out partial life histories. In May I returned to Peip-

146 bulletin: museum of comparative zoology

ing, and thereafter^war conditions prevented further field work. It
was with profound regret that I had to leave China at the very opening
of the best collecting season. Then I returned home to America, bring-
ing all the collections, made and borrowed. Acknowledgment is made
in the Manual of the many sources of the borrowed material.

The Manual was written at my home in Ithaca, in such intervals
of time as I could take from teaching and departmental administra-
tion. I wrote it for the use of Chinese students in the study of their
homeland fauna. I sought merely to provide them with concise de-
scriptions, keys and tables for families, genera and species; and in all
the larger genera, where species are difficult of identification, to pro-
vide figures of the genital characters that are the ultimate criteria for
species.

In matters of classification I considered it in the interest of students
that I should keep to the older and simpler family groupings, rather
than use the many recently introduced and still untested subdivisions
of the families, concerning the validity of which the specialists are not
as yet in entire agreement. I wanted to provide something that the
college students could use; and I have had the satisfaction of knowing
that they have used the Manual successfully.

In only one group, the Gomphinae, did I add any considerable num-
ber of new species. Seventeen of these were in the large and hetero-
geneous genus Gomphus of the older authors. Among them were new
heterogeneities that I could not fit to some of the subdivisions of that
genus that had been recently proposed. I contemplated further work
upon them when more adequate collections and more time should
make that possible; but little new material has come to hand. On
completion of the Manual borrowed specimens were returned to their
owners ; but I kept duplicates of the few species that were represented
by more than one specimen; and I took occasion, while the others were
in my hands, to make photographs of the wing venation of most of
them. And now that, in retirement, I have time for more adequate
study, I have only this scanty material available. A careful restudy
of the wing venation of the Chinese Gomphines is hereinafter at-
tempted.

The venation of the basal half of the hind wing appears not to have
been thoroughly explored; for in it there are structural characters
whose taxonomic value has been quite overlooked. The late lamented
E. B. Williamson discovered some new characters in these parts, and
used them to good purpose in his well known Burmese paper of 1907.
There are yet other unused characters to which I want to direct at-

NEEDIIAM: CHINESE GOMPHINE DRAGONFLIES

147

tention here. To them I will apply convenient terms for use in the
descriptions that are to follow. The new terminology will be merely
supplemental to that used in my Manual and fully illustrated in
Plate I of that volume.

In the accompanying figure is shown a diagram based on a careful
drawing of the base of the hind wing of Gomphus campestris Ndm.,
labelled to supply names for the parts hereinafter used. The principal
longitudinal veins, Costa, Subcosta, and Radius, Media, Cubitus and

Cu, en,

Fig. 1. Base of the hind wing of Gomphus (? Burmagomphus* campestris Ndm.

Anal, are labelled in the figure at both ends of each vein, with appended
numerals designating the branches of some of them; four important
cross braces, nodus (n), subnodus (sn), arculus (ar), and anal crossing
(Ac) and three other parts, middle fork (Mf), anal loop (AL), and
membranule (to), illustrated in Plate I of the Manual, are repeated in
this diagram.

New designations here introduced are as follows: intermedian cross-
f ins (i. med.: in this wing there is but one) lie in the space between
veins M3 and M4 and beyond the arculus to the middle fork (Mf).

In the anal area of both wings the cells that lie in line alongside the
anal vein from base to gaff (g)1 are called paranals. In the fore wing
there is oftenest but a single row of them: there is always more than

1 The gaff is the fused portion of veins C'u2 and Al.

•

148 bulletin: museum of comparative zoology

one row in the hind wing and the cells of the first row are highly
differentiated and are of great systematic importance. In our diagram
they are labelled n, o, p, and AL. Cells n and o are constantly present
as large single cells in all Gomphinae known to me, with the single
exception of Anormogomphus ; cells p and AL may be divided by cross-
veins. An anal loop (AL) is said to be present only when this area is
definitely bounded by a strong vein in the rear (with that boundary
generally much stronger than shown in this figure); this loop is often
enlarged and divided into several cells.

For the cell rows running in the opposite direction, from front to
rear, (more especially for the single row alongisde vein Al extending
from the hind angle of the triangle to the wing margin and numbered
1, 2, and 3 in the diagram) I use Williamson's name post anals.

The areas into which the broad anal field of the hind wing is divided
by the three branches of the anal vein may be designated as the first
(x), second (y) and third (z) anal interspaces. Each lies behind the
branch bearing its own number (principal veins being numbered
around the wing margin from front to rear). These interspaces differ
in breadth and slant, and in number, size and arrangement of the cells
composing them. Though little noticed hitherto, they offer excellent
systematic characters. The third interspace (z) is modified into the
anal triangle of the male.

GOMPHUS s.lat.

I wish now to make a further analysis of the species that I lumped
together in the great genus Gomphus in the Manual. As evidenced by
wing venation, these species fall into natural groups as follows:

Group 1. G. abdominalis only

For it I propose the generic name GASTROGOMPHUS. Its
characters are: a very long, thick abdomen, about a third longer than
the hind wing; anal vein 3 arises generally after, and sometimes
opposite the anal crossing; no basal subcostal cross vein, and no cross
veins in any of the triangles; first and fifth antenodals thickened; a
single row of large paranal cells in the fore wing; anal triangle of the
male three celled, and four postanals cells in the hind wing (see
Manual. PI. I, fig. 4); appendages of the male of about equal length
and divergence (Manual. PI. VI, fig. 2).

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 149

It should be noted also that in the one known species there is a very
wide differentiation in size among the cells of three wing areas; very
large before the level of the areulus, a little smaller outward to a line
drawn from the stigma to the hind angle of the wing, and much smaller
thence outward to the margin.

The nymph (Manual. PI. VII, figs. 1 and la) differs from all known
related forms in having neither dorsal hooks nor lateral spines; in
having the front border of the median labial lobe doubly produced
(bilobed) and fringed at the sides of a bare median notch; and in
having the strongly incurving terminal third of the lateral lobe very
feebly denticulate on its concave inner margin.

Group 2. Xenogomphus, gen. now Type G. agricola Selys.

Characters: middle fork (Mf) unsymmetrical, askew forward; gaff
as long as or usually longer than the inner side of the triangle; inter-
median crossveins 3/1 in fore and hind wing respectively; anal triangle
of the male hind wing usually of five cells; no basal subcostal antenodal
crossvein; first and fifth antenodals thickened; no anal loop, but
usually two complete rows of cells in the first anal interspace; male
caudal appendages of equal length but the branches of the inferior
much more widely outspread (see Manual. PI. VI, figs. 6 and 6a).

Here belong also G. succumbens Ndm. (Peking Soc. N. H. Bull. 5, 3,
1930), G. citimus Ndm., G. lautus Ndm., and probably also (judging
by similarity in form of male appendages) G. svrn-hcdini Sjostedt from
Szechuan and G. ekichibui Fraser and G. mclampus Selys from Japan.
This is the only group of species in the Gomphus series of Williamson
that has the middle fork (Mf) unsymmetrical.

The nymph (see Manual PI. VII, figs. 2 and 2a) is depressed with
strictly lanceolate abdomen bearing short triangular lateral spines on
segments 7 to 9, and low dorsal hooks on segments 3 to 9, the latter
very small at the front and regularly increasing in size to rearward to
the 8th segment. The middle lobe of the labium is prominent, tri-
angularly produced with a pair of little teeth at its slightly truncate
tip, and fringes of marginal hairs at either side. The terminal fourth
of the lateral lobe beyond the base of the strong movable hook is
roundly incurved to meet the denticulate inner border, without
forming a distinct terminal hook.

The type species of the two preceding genera are both pond species,
common in central China, where I made rearings of both of them
repeatedly.

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Group 3. Eogomphus, gen. nov. Type Gomphus neglectus Ndm.

Characters : Triangles of both fore and hind wings long in the axis
of the wing, and generally four-sided by failure of the long sides to
meet at the outermost angle ; both usually traversed by a single cross
vein; bridge vein shortened distally, the distance from subnodus to
oblique vein being about a third of the distance of subnodus from the
middle fork; gaff nearly as long as the inner side of the hind wing
triangle; basal subcostal crossvein present in fore wing, absent in the
hind; vein A2 weak and angulated so as to be almost unrecognizable;

Fig. 2. The wings of male (basal part) and female Eogomphus neglectus Ndm.

for a considerable distance from the wing margin the paired long veins
inclose more than one cell row, the greatest doubling between M3 and
M4, less between Oil and Cu2, least between Rs and M2; behind
Cu2 in the fore wing are two or three cell rows traversed by ill de-
veloped accessory branch-like sectors; and middle fork (Mf) sym-
metrical.

This genus is perhaps nearest to Davidius of the Gomphus series, but
it differs in having the fore wing triangle longer and not angulated on
the outer side; the intermedian crossveins are little reduced, being 5/2
in fore and hind wings respectively. This character transgresses the
lines heretofore drawn between the Gomphine and Epigomphine
series, as does also the general aspect of the rather elongate wings.

The nymph is unknown. The nymph figured in the Manual on
Plate VII, figures 3 and 3a, referred to on page 67 as possibly belonging
to this species, is the nymph of Merogomphus.

NEEDHAM: CHINESE GOMPHINE DRAGONFLEIS 151

The three teneral specimens representing this most unusual Gom-
phine came to me just before the manuscript of the Manual left my
hands. One male was retained for the Cornell University Collection.

After a reexamination of it, I may here add another note of de-
scription.

The face and top of the head and dorsum of the first abdominal
segment are densely hairy. The dorsum of the bilobed occiput is
thinly clad with short hair that is parted in the middle and outspread
flat both ways therefrom. The spines on femora and tibiae are very
numerous, short and in uneven alignment in the outer row on the hind
femur, with the last one in the row but little longer than the others;
those of the middle femur are more than twice as long, equally num-
erous and they form an even regular row.

The superior caudal appendages are widely divergent from the base,
tapering and convergent only toward their tips. Each ends in a small
black tooth. Below the base is a large inferior branch that ends in a
blunt black tooth. The inferior appendage is little more than half as
long as the superiors and its tips have less than half their spread. It
is quadrangular, with a straight hind margin from the angles of which
arise two stout branches that project straight to rearward. Each ends
in a blunt black upturned tooth. The genitalia of the second segment
are rather prominent. The anterior hamule somewhat resembles the
cheliped of a craw-fish with both tips bluntly rounded, the anterior tip
slightly longer and inflexed around the other. The posterior hamule is
perhaps twice as long as the anterior. There is a bulbous enlargement
of its upper third, bearing on its inner side a cluster of about a dozen
small black denticles; then suddenly tapering to a claw-like incurved
tip. The peduncle ("vesicle") of the penis slopes down to rearward,
and is deeply cleft on the anterior side for the reception of the greatly
expanded penis tip. It is nearly bare except for the edges of this cleft
and the hood-like inner side of it. The second joint is clavate toward
its tip, and lacks an apical spine. The reflexed third joint bears a re-
markable enlargement at its tip; a deeply cupped expansion that
carries a suggestion of likeness to an irregular flower. Above, the rim
of the cup is deeply emarginate, and within it arise two petal-like lobes.
Far out from its center projects a bifid stigma-like process ending in
a pair of blunt, recurved, flabellate tips, below which projects a spine
of half their length.

The apical carina of the tenth abdominal segment is produced to
rearward in a low bare triangular prominence, on either side of which
is the usual line of black denticles.

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Group 4. Merogomphus Martin. Type M. paviei Martin

A more careful examination of the single known female specimen of
Gomphus torpens Ndm. of the Manual leads me to conclude that it
should have been placed in this genus and associated with Merogomphus
vandykei Ndm., for it has the following characters : vein A3 arises op-
posite the anal crossing (Ac), and not beyond it; basal subcostal cross
vein (an) present in the fore wing; intermedial! crossveins 3/1 in fore
and hind wing respectively, middle fork (Mf) symmetrical; gaff more
than half as long as the inner side of the triangle ; two rows of paranal
cells in the fore wing; first anal interspace (x) wider than the second
(y), and no anal loop. It also has a half a dozen very large spines in the
outer row on the hind femur.

A nymph of this genus was figured in the Manual without name
(Plate VII, figs. 3 and 3a) and described on pages 66 and 67. It is
probably the nymph of M. vandykei, as determined by a recent study
of the venation of its crumpled wings.1

Group 5. Mesogomphus Foerster. Type Gomphus cognatus Rambur

The Gomphus brevipennis Ndm. of the Manual belongs here, as
evidenced by the following characters: vein. A3 arises just before or
opposite the anal crossing (.4c); two rows of paranal cells in the fore,
wing; intermedian crossveins 2/1 in fore and hind wing respectively;
first and fifth antenodals thickened; middle fork (Mf) symmetrical;
arculus unusually close to the triangle of both fore and hind wings and
the front side of the subtriangle much shorter than the inner side of
the triangle; four postanal cells in the hind wing; four or five cells in
the male anal triangle; no anal loop, and the first anal interspace
wider than the second.

The nymph has been described and figured for one species of this
genus, M. balneorum by Needham and Gyger (Philippine Jour. Sci.
63, 33,P1.X, figs. 125 and 126) and for two others, M. lineatus and M.
reinwardti by Lieftinck (Tijd.v.Entomol. 77,21, 1934).

Group 6. Burmagomphus Williamson. Type B. williamsoni

Fraser

The Gomphus dolus Ndm. of the Manual belongs here. In venation
it is very close indeed to the type species as figured by Williamson

1 The method used was that described by Dr. May K. Gyger in Entomological News, 50,
p. 21, 1939.

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 153

(Proc. U. S. Nat. Mux. 33, 298, fig. 27, 1907). The combination of
venational characters by which this genus has been set apart is as
follows: In the fore wing, a single row of paranal cells; a small triangle
slightly angulated near the middle of its outer side; a long close
parallel of veins M4 and (Ail beyond it with but two intervening
rows of cells out to the level of the oblique vein, with a rather sudden
widening thereafter; and in the hind wing but three postanal cells.

The nymph of this genus has been mentioned, and given an un-
intelligible two-line description by Fraser in Fauna of British India:
Odonata 2, 212, 1934.

Another small species that would appear to belong near to Burma-
gomphus is the one a portion of whose hind wing is shown in the
accompanying figure of Gomphus campestris Ndm. of the Manual.
(See figure 1.) It is of small size (hind wing 25 mm), with slightly
angulated outer side to the hind wing triangle, and only three postanal
cells. The single-celled anal loop is quite like that of B. icilliamsoni.
There are these discordant characteristics: there is a basal subcostal
crossvein in both fore and hind wings; one or two cells of the paranal
row in the fore wing are divided, and the double row of cells beyond the
triangle does not extend outward beyond the level of the nodus.1
The venation as a whole is more sparse, there being but 11:7/8:7
nodal crossveins in fore and hind wing respectively and only three
crossveins under the bridge.

My material is inadequate for determining whether these differ-
ences are constant enough to justify generic separation. Since this
species has been taken on the campus of Yen Ching University, it
should be possible for some one there to obtain additional specimens
including also its immature stages.

Group 7. Gomphus s. str. Type Libellula vulgatissima Linn

The remaining species appearing under this generic name in the
Manual may be allowed to remain so for the present. They show a
general conformity to the type, but with numerous small divergencies
which I shall now try to indicate in so far as they appear in the vena-
tion of the wings. They all seem to have the following characters in
common: middle fork (Mf) symmetrical; vein A3 arises beyond the
anal crossing (Ac); intermedian crossveins generally 2/1; paranal
cells in the fore wing more than a single row, some cells at least being

1 This disagreement applies also to Fraser's figure of Burmagomphus pvramidalis, Faun.
British India Odonata: 2, 212, fig. 66, 1934.

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divided; postanal cells four to seven; anal triangle of the male gener-
ally three celled; and no basal subcostal cross veins.

Recalling Burniagomphus are the two small shortwinged species
G. arvalis Ndm. and G. sowerbyi Ndm. These have a well defined one
celled anal loop with the base of vein Al kinked around its outer
corner. The outer side of the nearly equilateral fore wing triangle is
slightly angulated in the middle. The arculus is between the first
and second antenodal cross veins. The gaff is about half as long as
the inner side of the hind wing triangle. Vein M4 is slightly undulate,
and there are four post anal cells. This latter character prevents plac-
ing them in Platygomphus, as does also the well developed 3-celled
anal triangle in the male. De Selys placed a question mark before his
Platygomphus occultus when he placed it in that genus. It belongs
rather with the above named pair. The three might possibly be made
the basis of a new genus; but until Burmagomphus and Platygomphus
are better defined, and until Gomphus campcstris has found its place,
and until the nymphs of all of them are made known, another name
would only add to the confusion.

Two somewhat larger species of the Manual, G. intinctus and G.
collar is, are like the three preceding in most characters, but the
arculus is nearer the second antenodal crossvein, and intinctus has
five postanal cells. In all five the gaff is about half as long as the inner
side of the triangle.

A peculiar species that is known unfortunately from only a single
female specimen is G. edax Ndm. The triangles are both elongated in
axis of the wings, the outer end of the hind wing triangle being turned
up at the end like a sled runner (see the next figure) ; there are seven
postanal cells ; the branches of the anal vein are aslant outward.

Another peculiar species, described later by me (lAngnan Sci. Jour.
10, 227, 1931) from a single female specimen taken in Hainan is
G. hoffmani. It has the first and sixth antenodals thickened, the
arculus at or beyond the second; no basal subcostal antenodal cross-
vein, the fore wing with two rows of paranal cells, no large cells in the
anal area behind the first paranal row, and the first anal interspace
much wider than the second. Added to this array of differences there
is a peculiarly elongated three-celled anal loop, around the outer end
of which vein Al makes a short sharp bend. Also the hind wing is
widest at the nodus. The gaff is as long as the inner side of the tri-
angle, which latter in the fore wing is a little longer than the front side.
Whether this wing is quite normal I cannot say.

There remain six large species (hind wing 37-40 mm.) that conform

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 155

more closely (still none too well) to the type of the genus. One,
G. cuneatus Xdm. of the Manual appears to be distinguished by having
the veins M3 and M4 not at all undulate, strictly parallel and regularly
curved and inclosing somewhat larger cells than in the other five
species. In general it has a more open venation. It also has a longer
gaff, almost as long as the inner side of the triangle, and a shorter
kink in the base of the anal vein at the outer angle of the one celled
anal loop. There is in the type specimen but one extra cell in the
otherwise single paranal row of the fore wings.

Of the five remaining species, G. endicotti appears to be separable by
reason of the shortness of the front side of the fore wing triangle — not
longer than the inner side of the same; and G. flavicomis (Peking Soc.
Nat. Hist. Bull. 1, 2, 1930) by having its anal crossing close to the
inner end of the subtriangle in the hind wing — less than half its own
length therefrom.

Finally G. amicus is separable from the remaining two by the
shortness of its gaff — less than half the length of the inner side of the
front wing triangle; and these last two may be separated by the posi-
tion of the arculus in relation to antenodal crossveins: it is midway
between the first and second in G. clathratus, and at or very close to
the second in G. septimus.

THE FRAMEWORK OF THE WING ABOUT THE

TRIANGLES

In the preceding pages I have been pointing out the best single vena-
tional characters that I have been able to find for distinguishing each
of the species listed in my Manual under GOMPHLS (all of them ex-
cept G. sornnolens, of which I now have neither wings nor photographs
of venation available). I now wish to present in the form of a table
some correlations of characters especially to show the relations of the
parts of the strong framework of the wing that are in or around the
triangles.

As a standard of comparison I take the part marked a, which forms
the one common side of triangle and subtriangle, and which is formed
in development about the posteriorly deflected portion of the main
Cubital trachea. Two additional reasons for its selection are (1) its
central location, its ends being in contact with all the other parts
compared; and (2) its relative constancy in length. A little comparison

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showed all the other parts to be more variable. Next in constancy was
the part marked e, which is formed about the main Anal trachea.

The part a was given an assigned value of 10, and the length of all
the other parts were estimated in tenths of it. That is the meaning of
the numerals in the central columns of the table. These values are

Fig. 3. The parts about the triangle (T) in the wing hind of Gomphus edax Ndm. to illustrate
the terms used in the following table.

merely estimates made under inspection with a lens, without careful
measurements: wherefore allowance (possibly up to 10%) may have
to be made for errors of judgment, and additional allowance for varia-
tion in individual specimens.

The accompanying enlarged diagram, based on a drawing of the
wing of Gomphus edax Ndm. will be useful for comparison.

In general it may be said concerning venational characters that the
conjunctions and proportions and directions of the component parts
of the strong framework of the wing, and the layout of the spaces be-
tween principal veins and their branches offer far more dependable
taxonomic characters than are to be found in the number of interven-
ing crossveins.

NEEDHAM: CHINESE GOMPHIXE DRAGOXFLU SS

157

Venational Characters in 20 Species of Gomphus

Species

Hind
wing

I n ter-
med. CVS.

f/h tv.

a

Length of

parts about

hind W. triangle

b e d e f

9

extra
paran.

lis

post
a rials A3sAc

A. loop
present

abdominalis

35

2/1

10

17 18 9

11 5

5

0

4

opp

yes

agricola

25

3/1

10

16 18 10

11 6

12

0

4

out

no

amicus

40

2/1

10

16 18 11

12 8

4

3

4

in

yes

arvalis

29

2/1

10

18 20 12

12 7

4

2

4

in

yes

campestris

clathratus

collaris

26
38

31

2/1
3/1
2/1

10
10
10

12 13 11
17 19 10
15 16 10

12 5
11 7

11 7

5
5

4

1
2

2

3 •

4
4

n
n

n

yes
yes
yes

cuneatus

38

2/1

10

15 16 10

11 7

7

1

5

in

yes

dolus

23

2/1

10

9 11 9

11 4

6

0

3

m

no

edax
endicotti

35
34

3/1

2/1

10
10

18 20 11
17 19 10

12 6
11 7

8
9

2
3

5 i
5 i

n

n

yes
yes

flavicornis

37

2/1

10

17 19 10

11 5

4

2

5 ]

n

yes

gideon
hoffmanni

36
34

4/2
2/1

10
10

16 17 10
15 16 10

11 6

11 7

5
9

3
4

4
5

n
n

yes
no

intinctus

31

2/1

10

14 15 10

11 7

4

4

5

in

yes

neglectus

36

5/2

10

19 20 10

11 5

7

0

5

out

no

occultus

30

2/1

10

14 15 9

10 5

7

1

4

m

yes

septimus
sowerbyi

• 40
29

2/1
2/1

10
10

13 15 9

14 15 9

11 6
11 6

7
4

3
2

5

4

in
in

yes
yes

torpens

30

3/1

10

15 16 9

11 4

10

2

4 i

n

no

Column 1. Length of hind wing in millimeters.

Column 2. Number of crossveins joining the sectors of the arculus between the arculus
and the middle fork in fore and hind wing.

Column 3. Relative lengths of the parts about the triangle of the hind wing in terms of
tenths of the length of the inner side of the triangle.

Column 4. Number of extra paranal cells in the front wing (more than the single row always
present).

Column 5. Number of cells in the postanal row on the proximal side of vein 1st A between
the triangle and the hind margin of the wing.

Column 6. Position of origin of vein 3d A: in, proximal to it; opp, opposite it; out, distal
to it.

Column 7. Anal loop of hind wing.

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Leptogomphus unicornis Ndm.

A study of the wings of the single known specimen of this species
shows it to have been misplaced in the Manual in the genus Davidius.
It has more in common with Leptogomphus, including (1) the form of
the wings; (2) the lack of brace vein to the stigma; (3) the trigonal
interspace regularly widening outward to the wing margin; (4) the
small triangles ; (5) the narrow fore wing subtriangle ; (6) the little ex-
panded anal area of the wing; and (7) the 3rd anal interspace (z)
longer in the axis of the wing than wide. All these I regard as primitive
characters, of relative fixity.

Fig. 4. Wings of ? Leptogomphus unicornis Ndm.

It seems to differ from Leptogomphus as represented by its type
species, L. sauteri Selys, in having but a single row of very large
paranal cells in the fore wing, in having no basal subcostal antenodal
crossvein, and in having crossveins in all the triangles and in the
supratriangular space of both the wings. I present a figure of the vena-
tion to call attention to these discrepancies. At first glance they seemed
to me to be so great as to call for generic separation; but on further
experience with the Epigomphus alliance, I think they are very un-
reliable variants, having only specific value.

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 159

TWO NEW SPECIES AND SOME NEW RECORDS

Since the publication of the Manual and of the Additions and Cor-
rections (in the Peking Soc. of Nat. Hist. 5, 1-10, 1930) only one small
collection of dragonflies has come to me from China. It was from Dr.
Ting-wei Lew. It contained two new species of Gomphines and es-
tablished a few new records.

Gomphurus gideox spec. nov.

Length 63 mm.; abdomen 45; hind wing 38.

This is a blackish species with spatulately dilated abdomen; face
black with an oblong stripe of yellow covering about half of the lab-
rum. A similar somewhat larger isolated stripe covers the top of the
frons. The remainder of the top and rear of the head is black.

The synthorax is black in front with a pair of isolated dorsal stripes
that are divergent downward, not reaching the divided cross stripe on
the collar. An antehumeral stripe of yellow is represented by a very
small spot high up near the crest, and a thin faint streak low down on
the side, its lower end. Behind the broad black humeral band the
sides are yellow with a black line on the third lateral suture, that con-
nects with the back subalar carina above, and below runs down behind
the coxa of the middle legs. A vestige of a middle stripe is present in
front of the spiracle. Legs black beyond the short bicolored basal
segments. Wings hyaline, with a faint tinge of yellow in the mem-
brane. Ante- and postnodal crossveins are 16:12 and 10/12 in fore
and hind wing respectively.

The abdomen is very moderately enlarged on the two basal segments
narrowly cylindric on segments 3 to 7, and spatulate on 7 to 10, with
widely flaring lateral expansion of the margins of 7, 8 and 9. The
dorsum of 1 is mainly yellowT and a narrow lanceolate spot appears on
the base of 2. The sides of 1 and 2 are mainly yellow, and also the base
of 3. Basal yellowish rings on 3 to 7 become narrower to rearward,
with only fine yellow intersegmental lines across the apices of 7, 8,
and 9: segment 10 and appendages wholly black. The relative length
of the last four segments middorsally is about as 11; 9: 10:5; and ap-
pendages, on the same scale as 7. Diffuse large yellow spots cover
about half of the sides of segments 8 and 9. Caudal appendages are as

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shown in figure. The posterior hamules of the male, completely hiding
the anterior ones, project strongly downward even beyond the level
of the "vesicle", and taper to claw-like sharp tips that are directed
forward.

Fig. 5. The abdomen of Gomphurus gideon sp.n., dorsal view.

The female is very similar in coloration, with the yellow areas a little
more extended, especially on the abdomen. The subgenital plate is
divided deeply into two blunt equilateral triangles that extend to
rearward across about a fifth of the 9th sternite. The supraanal plate
is shining black above, yellow beneath.

A single pair, type and paratype, collected in Chengtu, Szechuan
on June 29th, 1929, and sent me by Dr. Lew. They are now in the
Cornell University Collection.

Because of the striking dilatation of the 7th and 8th segments of the
abdomen, shown in the figure herewith, and general accord in caudal
appendages and in venation, I have placed this species in Gomphurus;
a genus hitherto known only from North America. Another Chinese
species, Gomphus kryenbergi Ris, compared by its describer with the
American Gomphus scudderi, doubtless belongs beside it. G. gideon has
however a somewhat more copious venation than the American
species, with four cells in the anal triangle in both right and left hind
wings of the male: intermedian cross veins 4/2 in both male and female,
and six postanal cells.

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 161

Davidius serenus spec. nov.

Female; length 41 mm.; abdomen 31; hind wing 27.

This is a small blackish species with yellow sides. The head is all
black except the outer sides of the mandibles and a broad transverse
stripe across the very low prominence of the frons, which are yellow.
The thorax is black in front except for a pair of opposed 7-marks that
just meet at the middle of the collar. The stalks of the 7s are slightly
tapered upward and blunt at their isolated upper ends. There is no
antehumeral yellow stripe at all. Behind the very broad black humeral
band the sides are mainly yellow with a narrow black stripe on the
third lateral suture that is connected forward with the humeral above
and below. The black of the ventral side extends upward at the middle
suture to cover the spiracle. The legs are black. The long slender hind
femora are sagged downward in the middle and beset underneath with
more than a score of slender black short subequal spines.

Wings hyaline. Ante- and postnodals 13:12/9:11 in fore and hind
wing respectively. There is an extra cubito-anal crossvein in the fore
wing, and there is a single row of cells behind the anal vein.

Abdomen mostly black with a diminishing amount of yellow on the
sides of segments 1 to 7; segment 1 mostly yellow dorsally and 2 with
a lanceolate streak of the same color. On the sides of segments 3 to 5
the yellow is broken into a row of three spots; reduced to two spots on
6, and to a single spot on 7: 8 to 10 black. Appendages yellow.

The subgenital plate of the female is oblong flat, slightly tapering to
rearward, with a deep notch at the tip, and about four fifths as long
as the venter of the 9th segment.

This species is nearly allied to D. trox Ndm., but differs in being
smaller in size, in having the labium all black, in lacking the pale spot
that is a vestige of the antehumeral stripe, in lacking the J-spot at the
rear of the side of the thorax, and in having the abdominal segments
wholly black and the appendages yellow.

There is a single specimen collected at Ruling, China in July 1933
by Dr. Ting-wei Lew, and now in the Cornell University collection.

Among the specimens sent me by Dr. Lew were five females of
Gomphus septimus Ndm. The male was described in the Manual, p. 61.
The female, heretofore unknown, is like the male in coloration, with
a basal yellow halfring on abdominal segment 7 more conspicuous than
in the male. The relative length of the last four abdominal segments
is as 15:12:10:7, with the appendages 8, on the same scale. The promi-
nent subgenital plate is scoop-shaped or shaped like the spout on a

162 bulletin: museum of comparative zoology

pitcher, triangular, black, more than half as long as the venter of the
9th segment, and directed conspicuously downward. Among the rather
stout spines on the hind femur are four to six stronger than the others,
but intermixed with the others, and none of them is as long as the
femur is thick.

Two of the specimens came from Foochow in May; one from Mt.
Poliang ding, near Ho-kiang and Ming-kiant Fukein; and two from
Tu-ching, Min-giang in Fukien.

Dr. Lew sent also a fine pair of Mcgalogomphus sommeri Selys from
Kuling; a species that has hitherto been without definite locality
assignment in China.

TWO CORRECTIONS FOR THE MANUAL AND
A CONFIRMATION

(1) Agricnemis amelia Ndm. (Manual, p. 256) is a synonym of
Ischnura delicata Hagen, which is in turn considered by some authori-
ties to be a synonym of Ischnura aurora Burmeister.

(2) Taolcstes ncctans Ndm. (Manual, p. 256) is correctly described
and illustrated, but the nymph associated with it belongs elsewhere.
The female type specimen was presented to me along with a cast
nymphal skin, supposedly reared, and with several nymphs that had
been collected at the same spot. I uncritically accepted them as be-
longing together. The nymphs were not well preserved; but in a
recent examination of one of the best of them I find enough venation
still remaining in its wing pads to show that the antenodal crossveins
are numerous. That is sufficient to show that they are not Taolcstes.

The nymph is structurally very similar to those now known belong-
ing to the genus Euphaca. Judging by its size, it should be Euphaca
opaca.

A word about the placement of Taolcstes. As explained at the be-
ginning of this article, I followed the older and easier system that
segregated the Lestinae from the others on one principal character:
middle fork (Mf) nearer the arculus than the nodus. But in doing so
I pointed out (p. 226) the nonconformity of Taolcstes with the true
Lestinae.

Two additional species have since been described from China by
Erich Schmidt (Koitowia 10, 178-183, 1931) as species of Rhipidolcstes:
R. bidens and R. truncatidens; they conform much more closely to
Taolcstes, not only in having the middle fork nearer the arculus than

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 163

to the nodus, but also in many other points of venation, and in the
male genitalia. The species T. nectans differs from both in having un-
marked hyaline wings, much more open venation, and a shorter and
thicker stigma.

Concerning the nymph of Megalestes, I noted on page 229 of the
Manual that Laidlaw had described a nymph that he referred by sup-
position to M. major (Ind. Mus. Record, 19, 185-187, 1920). I said I
was not convinced that Laidlaw's nymph belonged to Megalestes.
My remark was more than justified: this, notwithstanding Leiftinck's
oracular pronouncement (Treubia 17, 58-61, 1939) in support of Laid-
law's supposition. I obtained an almost identical, certainly congeneric
nymph of Rhinagrion philippinum from Luzon, well preserved, and
showing so complete venation in its wing pads as to leave no doubt as
to its identity. It is described and figured in the Philippine Journal of
Science, 70, 266, Plate 15, figs. 206-213 and 215-216, 1939. Laidlaw's
nymph is Rhinagrion.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 4

P3AMMOCHARIDAE

(Spider-Wasps)

Notes and Descriptions

By Nathan Banks

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

June, 1944

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These publications are issued in numbers at irregular intervals.
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Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 4

PSAMMOCHARIDAE

(Spider-Wasps)

Notes and Descriptions

By Nathan Banks

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

June, 1944

No. 4 — Psammocharidae1

(Spider- Wasps) : Notes and Descriptions

By Nathan Banks

In continuation of previous studies on the fine collection of Psam-
mocharidae in the Museum of Comparative Zoology I present descrip-
tions of some new genera and species and notes on others. Neotypes
are made for two of Say's species and descriptions of varietal forms.
Some of these varietal forms illustrate the faunal divergence of the
New England area, which I have previously noted in a Myrmeleonid,
Hesperolcon abdominalis Say. Descriptions are given of the males of
three West Indian species previously known only from the females.
Keys are made for the species of Dipogon and for the males of the
species of Ageniella known to me from the Eastern States. A compari-
son of our large black Arizona Pepsis with a series of both sexes of the
related Cuban form and the South American Pepsis grossa shows that
our form is a distinct species.

a. NEARCTIC
Batazoxus interruptus Say

Say's diagnosis says "metathorax at tip bifasciate with yellow", and
the pleura with two yellow spots. It came from Indiana. This is the
form common in the middle states, west to Kansas and Nebraska, and
in Texas, North Carolina, etc. I have selected as neotype a female
from Chicago, collected by Prof. C. T. Brues, which agrees with the
description.

There is a form in the northeastern states which I call

Batazoxus ixterruptus var. cressoni var. now

The thorax and propodeum are black, apical margin of propodeum
with a broadly interrupted yellow line, no broad yellow band above it,
no marks on pleura, the hind margin of pronotum narrowly yellow,
and the scutellum yellow. The yellow on each side of face in the typical
form is here reduced to a slender orbital line, the clypeus is usually
black. The abdomen shows no pale at bases of segments, and the base
of first segment is usually black. The wings are darker. In the male
the yellow is likewise greatly reduced. The size is generally smaller.

The holotype is a female from Holliston, Mass., 4 August. Para-
types from Holliston from 24 June to 7 September; Wood's Hole,

'Published with the aid of a special gift from Mr. R. G. Agassiz.

168 bulletin: museum of comparative zoology

Mass., (C. T. Brues); Orient, N. Y., 12 July; White Plains, N. Y.,
Sept.; Wellesley, Mass., 13 July (Bolster); Trenton, N. J.; Penna.
(Melsheimer) ; Charter Oak, Penna, 11 August, (Kirk); and Glen-
carlyn and Falls Church in northern Virginia from 14 June to 30 July.

Type M. C. Z. No. 25729.

Pompilus ichneumonides D. T. (ichneumoniformis Patton) and P.
willistoni Patton belong to Batazonus. The latter I consider as a form
of interruptus, and the former as B. navus Cress.

Arachnophroctonus Ferrugineus Say

Say says "Ferrugineous; wings violet; pleura and metathorax
black." Inhabits U. S. The type was a male.

A specimen which agrees with the above and also with the fuller
description I have selected as neotype; it is from Falls Church, Va.;
23 July.

There are three varieties which can be separated on color; there is
scarcely any difference in structure, and the genital plate of the male
has a prominent median carina in all.

A. ferrugineus var. unicolor var. nov.

Some years ago Viereck labeled a specimen "unicolor"; it was from
the far west. The body is wholly ferrugineous except a small black
mark at the base of the abdomen ; nothing dark on thorax nor on pro-
podeum, nor even a trace of dark bands on abdomen; moreover the
wings are a yellowish brown, very much paler than those of the typical
form. The structure appears to be the same except that the clypeus is
not so deeply emarginate in the middle. The holotype is from Oak
Creek Canon, Arizona, 6000 ft., July (F. H. Snow) in the M. C. Z.
no. 25730. Others are from Tucson, and southern Arizona (Bequaert).

A. ferrugineus var. annexus var. nov.

From parts of Texas come specimens intermediate in some ways
between unicolor and typical ferrugineus. There is no black on thorax
nor on propodeum, but the abdomen above shows at least traces of a
dark band at end of first and second segments. The wings are just as
dark as in typical form, which separates it from unicolor.

Holotype is from Fedor, Lee Co., Texas, June 15, 1909 (Birkmann);
paratypes from Fedor in June and September, also Dallas, Texas
(Boll); Austin, Texas, 13 October (Brues); and Davis Mts., Texas, 28
June (Englehart). Type, M. C. Z. no. 25731.

Typical ferrugineus also occurs in Texas.

hanks: psammocharidae 169

A. FERRUGINEUS var. NIGRESCENS var. nOV.

In the northeastern part of the country there is a much darker form;
the wings are nearly black; the abdomen, in the female, shows the
dark bands at tips of first and second segments. The thorax and pro-
podeum are wholly black above and below, and on pleura; the femora
are at least partly black, the hind femora often wholly so; in the
female the tip of tibia and of some tarsal joints also black; the antenna
is nearly wholly black, basal joint only remaining pale; in the male the
basal segment may be more black than usual; in females the face
above the antennae is more or less darkened. The clypeus is less deeply
emarginate than in typical form; otherwise I note no structural
difference.

The holotype is a female from Arlington, Va., 1 1 August; paratypes
from Fort Lee district, X. J., August; Woods Hole, Mass., (Brues);
Holliston, Mass., 27 July (Bks.), and Coldbrook, Conn., 14 July
(Wheeler). Type, M, C. Z., no. 25732.

Anoplius imbellis spec. nov.

Black throughout, wings not darker at tip. Moderately long black
hair on head, thorax, and propodeum; some stiff black bristles above
tip of abdomen, and some slender curved ones above and below; no
hair on pleura. ( 'lypeus fully two and one-half times as broad as long,
below truncate; antennae slender, third joint fully five times as long
as broad, fourth about two-thirds as long as third. Face but little
narrower above than below; a median line from antennae to the an-
terior ocellus, ocelli subequal, laterals a little nearer to each other than
to eyes; pronotum deeply angulate behind; propodeum with no dis-
tinct median groove; abdomen moderately broad at base, slender and
pointed at tip; mid and hind tibiae with short spines above; long spur
of hind tibia nearly three-fifths of basitarsus; claws with short, erect
tooth.

In fore wings the basal vein a little before the transverse, marginal
cell more than length from the wing-tip, outer side straight; second
submarginal cell a little longer than broad, receiving the first recurrent
near tip; third submarginal triangular or nearly so, outer side convex,
receiving the second recurrent (lightly curved) near middle.

Length 9 7.5 to 8.5 mm.

What is doubtless the male is similar, but little more slender; the
antennae heavy, third joint about three times as long as broad; the
third submarginal cell triangular or even pedicellate, shorter than the

170 bulletin: museum of comparative zoology

second; the fourth and fifth ventral segments are covered with a
brush of black hair, denser and longer on sides ; the genitalia show a
median plate, narrow, almost pointed at tip, and with a median,
scarcely elevated ridge.

Length c? 7 to 8 mm.

Females from Corvallis, Oregon, 30 August, 4 September, and
Hillsboro, Oregon, 25 September.

Males from Corvallis, 6, 25, 30 August, and Cornelius, Oregon,
4 August (Scullen coll.).

Type in Oregon Agricultural College, paratypes there and in
Museum of Comparative Zoology no. 25734.

Nannopompilus texanus spec. nov.

Similar in structure to AT. subviolaceus. Abdomen only faintly
bluish; fore wings scarcely smoky, but a broad dark band at tip; hind
wings hyaline, smoky at tip. The body, antennae, and basal joints of
legs sericeous, strong on lower face, clypeus, coxae, and posterior sides
of propodeum. Antennae fully as short as in Ar. subviolaceus, head,
thorax, and propodeum base as in that species. Venation similar, the
second submarginal cell nearly as high as long; the recurrent veins end
beyond middle of cells. The legs also similar, except that the spines
of the comb are longer and thickened, and some a little curved, but
none plainly spatulate. Marginal cell a little more than its length from
tip of wing.

Length 7 mm.

One female from Richmond, Fort Bend Co., 22 June, (Bequaert)
and another from Fedor, Lee Co., (Birkmann), both Texas. Type
M. C. Z. No. 25S95. Separated from A:. subviolaceus by the much
paler wings, sericeous body, stouter spines.

Gymnochares texana spec. nov.

Black, pronotum slightly sericeous near hind border, and in front
near collar; antennae dark ferruginous; wings slightly darkened
throughout, tip a little darker, but not a band. Clypeus more than
twice as broad as long, lower edge truncate; each side above clypeus
the face is faintly sericeous, also the base of mandible; face with nearly
parallel sides, except near top; a very distinct median groove from an-
terior ocellus to antennae, latter slender, third joint longer than ver-
tex-width and one and a half times longer than fourth, latter but little

banks: psammocharidae 171

if any longer than fifth. Hind ocelli a little nearer the eyes than to each
other; vertex with a few fine, long hairs; pronotum almost angulate
behind, faintly depressed near hind border, and with a median groove
reaching forward; pro and mesonotum without hair, as also the pro-
podeum, latter with a very distinct median groove above; on the sides
of the posterior slope and extending over the side are distinct but not
strong transverse ridges, fading out toward center. Abdomen slightly
compressed near tip, scarcely a trace of hair above toward tip, below
a few near tip.

Legs slender as in other species; hind femora above near tip with
three well-separated minute hair-pits, rather fainter than in allied
species; hind tibia with only a few scattered minute spines, three
minute ones on apical half of outer side; inner spur of hind tibiae not
one-half as long as basitarsus, latter with only a trace of spines below.

Fore wings with marginal cell about its length from wing-tip, about
two and a half times as long as broad, not quite as slender as in birk-
manni; second submarginal cell fully one and a half times as long as
broad, receiving the first recurrent a little before tip (nearer middle in
birkmanni); third submarginal cell only a little longer than second,
narrowed one-third above, receiving the second recurrent a little be-
yond middle; basal vein before transverse; cubitus extending out to
margin as in other species. In hind wing the anal ends much before
the fork.

Length 8 mm.

One female from Austin, Texas, May (Melander), Type, M. C. Z.
25704.

Similar in general to birkmanni, but readily separated by the groove
on propodeum.

Nannochilus gen. now

Two of our species described under Ageniella, externa and congrua,
do not have a petiolate, but a sessile abdomen; they are therefore re-
lated to Priocnemis, but differ from that genus in lacking teeth or even
bristles on the hind tibia. The claws are cleft, the inner process being
very broad and obliquely truncate, and the pulvillus is extremely
large. The venation is similar to that of Pseudagenia, but the basal
vein is interstitial with the transverse. Besides Ageniella externa Bks.
which is the genotype, there is A. congrua Cress., and a new species
described herewith.

172 bulletin: museum of comparative zoology

Nannochilus osoria spec. nov.

Body black, abdomen sometimes more brown; antennae and legs
brown, sometimes the mid and hind femora are somewhat rufous, and
the front tibiae and tarsi are usually pale yellowish, all spurs pale, but
not snow-white. Clypeus and lower face with silvery pubescence, little
on vertex, but more prominent on coxae and on each side of propodeum
behind the spiracles.

Fore wings rather evenly dusky, tip not darker; veins brown to
black; hind wings more hyaline.

Clypeus broad, short, and truncate below; lateral ocelli a little
nearer to each other than to eyes; antennae moderately long and
slender, much as in iridipennis; pronotum arcuate behind, finely
granular, a median groove on basal part; abdomen with short pale hair
on venter; basal segment not nearly twice as long as broad behind.

Fore wings with submarginal cells much broader than the marginal
cell; second submarginal about as broad as long, receiving the first
recurrent near middle; third submarginal hardly longer below, but
larger, the posterior side almost angulate in middle, so the cell is no
broader below than above, receiving the second recurrent vein near
middle; basal vein interstitial with nervellus, its lower part not
strongly curved. In hind wings the anal vein ends before the fork of
cubitus.

Length of fore wing 5 to 6.5 mm.

Holotype male from Falls Church, Virginia, 12 July; paratypes from
the same locality from 28 June to 7 September; also from Great Falls,
Virginia, 20 June, and Chain Bridge, Virginia, 23 June. Type M. C. Z.
No. 25907.

Three females agree in general; dull black, sericeous, venation as in
male, spurs pale, clypeus truncate below; third antennal joint a little
longer than fourth; pronotum broadly arcuate behind; inner spur of
hind tibiae hardly one-half of basitarsus; basal segment of abdomen
broad and sessile.

Length of fore wing of female, 6 mm.

All from Falls Church, Virginia, 13 September.

Priocnemis occldentis spec. nov.

Closely related to the Eastern P. nothus; in color the wings are uni-
formly darker and the abdomen is entirely reddish, the last few seg-
ments not darkened as in nothus. The venation the same as that of
nothus even to the bend in the lower part of the submarginal cross-

banks: psammocharidae L73

vein; the propodeum is less evenly rounded than in nothus, seen from
the side there is almost an angle at the turn, the posterior slope steeper
than in nothus.

The second plus third antenna] joints not equal to vertex-width;
ocelli subequal, in a slightly broader triangle than nothus, the laterals
much nearer to each other than to eyes; pronotum angulate behind;
no median groove on propodeum, the hue striae as in nothus; the spines
and spurs on legs as in that species.

Length 6.5 to 8 mm.

From Corvallis, Oregon, 20, 22, 23 July, 6, 23, 25, 30 August, and
4 September, also Forest Grove, 1 June; Blooming, 26 July; and Hills-
boro, 24 September, all in Oregon.

Type in Oregon State College, paratypes there and in the M. C. Z.
no. 25733.

Priocxemis nebulosus Dahlb.

Dahlbom says the wings are violaceous, mentions the emarginate
clypeus, and locality South Carolina. From Florida in the Graenicher
collection are two males and a female which agree better with the
description than the form from Virginia northward that, following
Cresson, I previously identified as nebulosus.

In the Florida female the clypeus is more plainly emarginate in the
middle, the angle each side is almost a tooth; the wings are more
evenly dark than the northern form, which will be known as P. pul-
chrina Cresson, based on the male.

In the male of nebulosus the wings are dark as in the female (in
pulchrina the male has hyaline wings with a smoky tip), and the black
median stripe on the face is broader, especially on the clypeus. The
legs are marked as in pulchrina. In both sexes the second submarginal
cell is not as long as in pulchrina.

A female from Larkins, May. males, South Miami, 26 April, and
Lake Apopka, Winter Garden, 25 April, all Florida.

Priophanes

There are several among the described species of Priocnemis which
have a distinct petiole connecting the abdomen to the propodeum,
while true Priocnemis has no petiole. For the petiolate forms I pro-
posed a new genus which is more related to Pseudagenia and Ageniella
than to Priocnemis. The venation is much like Pseudagenia, the basal
vein ends before the transverse, and in hind wings the anal vein ends
before the cubital fork. The hind tibiae have three rows of spines

174 bulletin: museum of comparative zoology

above the middle one with more or less distinct teeth; last joint of mid
and hind tarsi bare, the claws toothed; no distinct "beard" under
head; venter of female with a groove on second segment; the mesoster-
num not prominent laterally.

Type Priocnemis facetus Cress.

Also included are P. agcnoideus, arizonica, arcuatus, holonis, n. sp.,
placitus, relictus.

Priophanes holonis spec. nov.

Black; abdomen reddish all over; wings hyaline, apex broadly
dusky.

In general close to P. directa of Texas; the pronotum angulate be-
hind, the spurs black, legs and antennae entirely black; and head,
thorax, and abdomen of the same shape as P. directa. It differs in
having the teeth on hind tibiae much smaller, hardly noticeable, except
from inner side, and more especially in venation. The marginal cell is
longer, its outer side more oblique, and not quite its length from tip of
wing (in directa more than its length from tip of wing) ; the second and
third submarginal cells are proportionally longer than in directa, and
the lower outer corner of third submarginal cell is scarcely the length
of third submarginal from the outer margin of wing (in directa more
than length of third submarginal cell from outer margin of wing).
The first recurrent ends at or before middle of second submarginal,
and second recurrent before middle of third submarginal cell. The
basal vein, which is curved as in directa, ends only a little before the
transverse vein.

Length 7.5 mm. to 8.5 mm.

Holotype from Urbana, Illinois, 20 July (Bequaert); paratypes
Columbus, Ohio, August (Bequaert), also 6, 21 July (Gillaspy); and
MacCollum, Coweta Co., Georgia, 8 June (Bequaert). Type M. C. Z.
25892, paratypes there and Ohio State University.

Ageniella delicata spec. nov.

Dull black, abdomen with all of second, most of first, and part of
third segments yellowish rufous; tip of abdomen with white spot;
mandibles yellowish at tips; antennae brown, basal joint pale yellow
beneath; all coxae black, rest of front legs pale yellowish, mid legs
with tibiae and femora pale, tarsi brown, hind legs with femora pale,
tibiae and tarsi dark brown; spurs of hind legs dark brown, others
paler; wings hyaline, tip not darker, veins yellowish to brown.

banks: psammocharidae 175

( "lypeus and lower face with silvery pubescence; notum and pleura
somewhat sericeous.

Body slender; clypeus broad, somewhat rounded below; lateral
ocelli much nearer each other than to eyes; vertex convex; antennae
long and slender, third joint hardly longer than fourth; pronotum
deeply arcuate behind; propodeum finely granular, median groove
faint; basal segment of abdomen more than twice as long as broad
behind.

Fore wings rather short, the submarginal cells not as broad as the
marginal, latter nearly its length before tip of wing; second submargi-
nal about one-third longer than broad, receiving the first recurrent
vein near base; third submarginal plainly longer on lower side than
second, but narrowed nearly one-third above, receiving the second re-
current (almost angulated) a little before middle; lower part of basal
vein bulging forward, ending a little before the nervellus.

Mid and hind legs very slender and long, smooth, inner spur of hind
tibiae two-thirds of basitarsus.

Length of fore wing 4.2 mm.

One male from Falls Church, Va., 22 August. Type M. C. Z. No.
25910.

Ageniella restricta spec, now

Body dull black, more or less sericeous, most noticeable on the
coxae and pleura; clypeus and lower face with dense white pubescence;
abdomen with basal two-thirds of second segment yellowish rufous;
tip of abdomen with a white spot; legs brown to black, front legs with
tibiae and tarsi pale yellowish, hind spurs dark, others paler. Fore
wings dusky, tip rather darker, veins dark brown; hind wings almost
hyaline.

Body slender; clypeus truncate below; antennae slender, third joint
a little longer than the fourth, on the style of A. iridipcnnis; lateral
ocelli plainly nearer each other than to eyes; vertex rather strongly
convex; pronotum arcuate behind; propodeum finely granular, sloping
toward tip, median line scarcely visible; abdomen slender, basal seg-
ment more than twice as long as broad behind.

In fore wings the submarginal cells as broad as the marginal cell,
latter not quite its length from the tip of wing; second submarginal
nearly one and a half times as long as broad, receiving the first recur-
rent before basal third; third submarginal cell little longer below than
second, but broader, narrowed hardly one third above, receiving the
second recurrent (nearly evenly curved) before middle: lower part of

176 bulletin: museum of comparative zoology

basal vein only slightly bulging, and ending a little before the nervel-
lus; mid and hind legs long and smooth, inner spur of hind tibia more
than one-half but hardly two-thirds of basitarsus.

Length of fore wing 5 mm.

One male from Falls Church, Virginia, 22 August, on leaves of
tulip-tree with honey-dew. Type M. C. Z. No. 25909. Closely re-
lated to A. delicata, but differing in less reddish on abdomen, basal
joint of antennae black, shorter spurs, darker wings and venation, and
broader submarginal cells.

Ageniella neglecta spec. nov.

Body, antennae, legs, wholly black, hind spurs black, others paler;
fore wings nearly hyaline, slightly darker near tip, veins brown.
Clypeus and lower face with white pubescence, thorax and coxae
somewhat sericeous, not on abdomen.

Clypeus truncate below; antennae short and rather thick, the third
joint little more than twice as long as broad at tip ; pronotum broadly
arcuate behind; propodeum (from side) slightly rounded, no median
line; basal segment of abdomen not quite twice as long as broad be-
hind. Legs short, smooth, inner spur of hind tibiae little more than
one-half of basitarsus.

In fore wings the marginal cell is fully its length before tip of wings,
the third submarginal cell nearly three times its length from margin,
and almost as broad as the marginal cell; second submarginal cell a
little longer than broad, receiving the first recurrent at basal third;
third submarginal a little longer than the second, only a little nar-
rowed above, receiving the slightly curved second recurrent near tip;
lower part of basal vein only a little convex and ends on the nervellus ;
in hind wings the anal ends much before cubital fork.

Length of fore wing 3.5 mm.

One male from Boulder, Colorado, 26 August, 1908 (S. A. Rohwer).
Type M. C. Z. No. 25908.

Readily known by small size, deep black color, short legs, and the
large apical field beyond venation.

Ageniella accepta var. conflicta var. nov.

Differs from the typical form in having a black spot on side of scutel-
lum and on side of metanotum as well as at base of fore wing, usually
all run together in a black stripe; area between ocelli black; body more

banks: psammochajridae 177

slender than typical form; the pale area of fore wing between the first
brown band and the stigma] band is more suffused with pale brown, so
that the wing is not so plainly three-banded; the basal part of wing
is more yellowish, and the brown at apex is a paler brown than in
typical acccpta.

Holotype, a female from Falls Church, Va., 5 July; paratypes also
from Falls Church from 19 June to 13 Sept.; from Glencarlyn, Va.,
26 July, and from Riverhead, L. I., X. Y., 1 August (W. T. Davis).
Type M. C. Z. no. 25735.

Synoptic table of males of eastern species
of Ageniella at present known to me

1. Tip of abdomen with a distinct white spot; third cubital cell longer below

than above 2

Tip of abdomen wholly dark 11

2. Some reddish or yellowish on dorsum of abdomen; basal segment of abdo-

men more than twice as long as broad 3

No reddish nor yellowish on dorsum of abdomen 7

3. All spurs snow-white ; femora rufous birkmanni

At least hind spurs dark 4

4. Clypeus with a small pale spot on each side, and a small orbital spot .festina-
Clypeus wholly dark 5

5. A small yellow orbital line or spot; marginal cell much less than its length

from tip of wing; inner spur of hind tibiae scarcely more than one-half of

basitarsus; basal joint of antennae dark fratemella

Marginal cell nearly its length from tip of wing; no pale orbital line or spot;
inner spur of hind tibiae more than one-half of basitarsus 6

6. Basal joint of antennae pale yellowish below, femora yellowish or rufous,

paler than rest of legs; red of abdomen on all of second and parts of first

and third segments delicata

Basal joint of antennae black below; femora more brown, darker than rest
of legs; red on abdomen only on part of second segment restricta

7. Two pale spots or a white band on hind border of pronotum ; clypeus mostly

whitish; spurs snow-white calcarata

No pale mark on pronotum 8

8. A small yellowish spot or short stripe on orbital line a little above clypeus;
long spur of hind tibiae about one-half of basitarsus; vertex usually shin-
ing; fore wings more than five millimeters long agilis

No such spot or stripe on orbital line; fore wings not over five millimeters
long 9

178 bulletin: museum of comparative zoology

9. Femora rufous or yellowish, sometimes tibiae also; inner spur of hind

tibiae hardly more than half of basitarsus texana

Femora and tibiae black; inner spur of hind tibiae more than one-half of
basitarsus 10

10. Abdomen very slender, second and third segments with pendent side-mem-

brane, basal segment plainly more than twice as long as broad behind.

pctiolata

Abdomen broader, no side-membrane to second and third segments; basal

segment twice as long as broad behind, tip of fore wing black norata

11. Abdomen with some reddish or yellowish on basal half above 12

Abdomen wholly black above 14

12. Basal segment of abdomen not twice as long as broad behind; third sub-

marginal cell with angulate hind border perfecia

Basal segment of abdomen more than twice as long as broad; small species.

13

13. Clypeus black; second segment of abdomen dark in middle, yellowish on

sides; third submarginal cell higher than long; hind spurs dark; basal
segment of antennae black; third submarginal angulate behind.

minuscula

Clypeus mostly pale; spurs white; basal joint of antennae pale above and

below; hind border of third submarginal cell not at all angulate behind.

apicipennis

14. Clypeus with some pale, at least on lower edge; palpi pale 15

Clypeus wholly dark 17

15. Third submarginal cell with outer border angled in middle, so cell no

broader below than above; front coxae almost white clypeata

Third submarginal cell plainly longer on lower side than on upper 16

16. Hind border of pronotum slightly pale; basal joint of antennae white below.

hestia
Hind border of pronotum shows no paler band; basal segment of abdomen
not twice as long as broad behind; antennae thicker than usual.

crassicornis

17. Third submarginal cell with hind border angled in middle, so cell no

broader below than above 18

Third submarginal cell with outer side sloping, and lower border longer
than upper 19

18. Hind spurs dark; second recurrent arises near the middle of outer part of

cubitus; small species eximia

All spurs nearly white; second recurrent arises plainly beyond the middle of
outer part of cubitus ; larger species osoria

19. All spurs snow-white; basal segment of abdomen fully twice as long as

broad behind virginica

Basal segment not quite twice as long as broad behind; all spurs not white

20

banks: psammocharidae 17!)

20. Third suhmarginal cell much longer than high; a distinct median groove

on propodeum species?

Third submarginal cell scarcely as long as high, often plainly higher than
long below; no distinct groove on propodeum 21

21. Hind femora reddish (or yellowish) on outer half, base black or brown.

aludra
If hind femora pale it is also on basal part 22

22. Second recurrent arises little if any beyond middle of outer part of cubitus.

23
Second recurrent arises plainly beyond middle of outer part of cubitus . . 24

23. Front femora, tibiae, and tips of coxae pale; third submarginal cell always

longer below than above; hind legs often partly pale tenella

Front femora, tibiae, and coxae not pale; third submarginal cell often no
longer below than above; legs black, a small species eximia

24. Front tarsi yellowish; basal segment of abdomen about one and one-half

times as long as broad behind iridipennis

Front tarsi not yellowish; basal segment of abdomen not one and one-half
times as long as broad behind atrata

Nemagenia subgenus nov. of Ageniella

Pronotum above as long as mesonotum ; marginal cell its length from
tip; basal vein plainly before nervellus; inner spur of hind tibiae not
one-half of basitarsus; propodeum very long, nearly flat, from side
it is only slightly curved; otherwise like Ageniella.

Type Pompilus (Agenda) longulus Cresson.

Only the male is known; described from North Dakota, I have
specimens from Fedor, Texas (Birkmann).

Dipogon texanus spec. nov.

Head and abdomen black, clypeus and thorax rufous, legs dull
black to brown, tarsi partly paler, coxae rufous, antennae yellowish,
a narrow-black band at tip of each joint, except basal and apical; fore
wings hyaline, a narrow black band across over the basal vein, and a
very broad blackish band across over submarginals and third discoidal
cells, not occupying the tip of marginal cell.

Face and vertex with short appressed white hair ; abdomen also with
pale appressed hair, longer and erect hair near tip; venter with bands
of pale hair. Hair basket under head of white bristles. Structure in
general similar to D. brevis; very little hair on thorax or propodeum;
venation similar, but marginal cell hardly as broad, but angulate on
hind border; second submarginal about twice as long as the third; first

180 bulletin: museum of comparative zoology

recurrent ending much before middle of second submarginal, second
recurrent ending near base of third submarginal cell; medius reaches
margin; legs slender as usual; long spur of hind tibia about two-fifths
of basitarsus.

Length 6.5 mm.

From Brownsville, Texas, 11 to 16 June (Darlington). Type
M. C. Z. No. 25896.

Dipogon sericea spec. nov.

Body black; antennae with first and second joints black, beyond
yellowish, some joints narrowly dark at tips; legs black, tarsi and the
front tibiae rufous. Fore wings brown, beyond the marginal cell
snow-white, the marginal cell and below and beyond it darker than
elsewhere, stigma black; hind wings slightly infuseate.

Thorax and abdomen densely clothed with appressed gray pu-
bescence, in places somewhat yellowish, thorax with rather long, erect
white hairs, and shorter ones on the abdomen, near tip darker. Face
above antennae with bright yellowish pubescence, across face half way
up to vertex is a band of erect black bristles, and similar bristles on
vertex; the lower edge of clypeus is yellow, rest black, clothed with
white hairs, white hairs back of eyes; hair basket of fine, pale bristles.

Legs with the femora, mid and hind tibiae covered with sericeous
pile, the femora with fine, long hairs below. Hind tibiae above with a
groove and a row of fine short hairs, long spur about one-third of the
basitarsus. Antennae rather slender, third joint not much longer than
fourth, together they equal vertex-width.

Thorax shorter than in Eastern species, but fully as broad ; pronotum
broadly arcuate behind; propodeum appears to have a broad median
furrow, but all covered with the appressed hair.

In fore wings the venation is much like the Eastern D. sayi except
that the marginal cell is much shorter, extending only a trifle beyond
the third submarginal cell; the medius reaches to the margin, but in
the white area it is very fine. In hind wings venation as in other
species.

Length of fore wing 6 mm., body 6.5 mm.

One from Bull Prairie, Lake Co., Oregon, 22 July, Camas Prairie
Summit, 7,500 ft. (Frewing coll.) Type at Oregon State College.

The species of Dipogon so far described from the United States
can be separated by the following key :

1. The medius of fore wing plainly does not reach the margin; marginal cell not
strongly angled at end of second submarginal cell, the outer side curved.

Subgenus Adipogon — 2

banks: psammocharidae 181

The medius reaches to the outer margin; the marginal cell is strongly
angled at the end of the second submarginal cell, and broadest at this point.

Subgenus Dipoyon — 3

2. Forewings wholly dark, nearly black papaya

Forewings clear with a narrow dark band over the basal and transverse

veins, and a large dark spot over marginal cell and the two or more cells
behind it pulchripennis

3. Thorax yellowish to rufous 4

Thorax black 5

4. Head and abdomen black; antennae rufous, tips of joints narrowly black;

legs partly black texan us

Head rufous, abdomen black; antennae rufous, no dark bands; legs wholly
yellowish yraenicheri

5. Face, mesonotum, and abdomen with appressed grayish to yellowish pu-

bescence; hair basket pale; front legs partly pale 6

Face, thorax, and abdomen without such pubescence; fore wings clear with
two bands 7

6. Across face and on vertex are erect black bristles; fore wings brown, tip

snow-white; propodeum and pleura also sericeous sericea

No such bristles on face and vertex; extreme wing-tip faintly dark, no snow-
white; propodeum and pleura without appressed hair brevis

7. Front legs largely yellowish, also mandibles, and lower edge of clypeus.

caliptera
Front legs, clypeus, and mandibles black sayi

Adipogon, subgenus nov. has the basket of curved hairs on under
side of head as in typical Dipogon, but the median vein of fore wing
does not extend to the outer margin, and the marginal cell is not
angled below.

Type is Pompilus pulchripennis Cresson.

Pepsis pattoni spec. nov.

Our large black Arizona Pepsis has been identified by Fox and my-
self as P. obliquerugosa Lucas, a form described from Cuba. A few
years ago Salman considered both to be P. grossa Fabr. of northern
South America. With a series of both sexes of the three forms I con-
sider each a separate species.

As with the others, the wings are black with a narrow pale band at
tip, not as broad, and in the female not as white as in obliquerugosa.
On the propodeum the side tubercles are very prominent, sometimes
almost pointed, in obliquerugosa much lower and less noticeable. At

182 bulletin: museum of comparative zoology

the middle of the turn there is in both species a high ridge; in obliqueru-
gosa its top is straight or a little convex, in pattoni the top is plainly
emarginate in the middle. The oblique ridges on the propodeum so
characteristic of obliqucrugosa, are fewer, less oblique, and sometimes
none oblique. In pattoni the third joint of antennae in female is a
little shorter than in obliqucrugosa in specimens of same wing-length.
In the male the parameres of the genitalia are slender and tapering
toward tip; in grossa (see Lucas figure 24) they are heavier, and not at
all tapering; in obliqucrugosa the parameres are not as broad as in
grossa, but do not taper as much as in pattoni; in the latter the tip of
the subgenital plate is broadly convex, in the Cuban form it is plainly
emarginate in middle. The females of all three forms have long, stiff,
curved bristles under the front femora, and with our ncphclc and a few
South American species they form a natural group. Lucas placed the
male of ncphclc asformosa and says that the genitalia are the same as in
grossa, and gives no separate figure. The males of this section have
the subgenital plate slender and furnished with a broad dense crest of
long black hairs.

P. obliqucrugosa is somewhat the largest of the three species, fore
wing sometimes 45 mm. P. pattoni sometimes about 42 mm., fre-
quently 40 mm. P. grossa rarely 40 mm., often near 35 mm.

Holotype of P. pattoni 9 is from Palmerlee, Arizona, July (Bieder-
mann coll.), M. C. Z. No. 25805; allotype from southern Arizona;
paratypes are from Palmerlee, Tucson, Ola, Ft. Grant, Pinalerro
Mts., Santa Catalina Mts., all Arizona, from late June to September,
taken by Snow, Bequaert, Morse, and Wheeler. One from mountains
near Pomona, California, by H. C. Fall.

b. ANTILLEAN

Batazonus gundlachi Cress.

Described from female only.

Male. It is very similar to female, and marked the same; thorax
and abdomen brownish red, the clypeus is reddish in middle, above
antennae are two short curved dark streaks, there is some yellow each
side of scutellum (as well as the post-scutellum) ; a large yellow spot
just above base of mid coxae; propodeum broadly yellow across apical
half; abdomen black at extreme base, and faintly dark across tip of
first segment, a broad yellow band across base of third segment above
and below; legs reddish, except the paler tarsi which are dark at tips

banks: psa mmoch arid ae 183

of joints; inner spur of hind tibia about three-fifths of basitarsus. The
third submarginal cell is much longer than high, as in the female.

Length of fore wing 13 mm.

Allotype from San Bias, Trinidad Mts., Cuba, 24 April (G. E. Folk).

Batazonus hookeri Rohwer

Described from female only.

The male is largely yellowish; there is a triangular black spot below
antennae, two broad black stripes above, narrowly separated; two
yellow lines on mesonotum, the mesopleura has two slightly separated
elongate, yellow spots; the propodeum above is yellow, black at ex-
treme base, usually extended back in middle to divide or partly divide
the yellow. The dark bands on abdomen above are brown to black,
occupying hardly one-half of the segment, basal segment dark only at
tip; coxae black, with a yellow mark; femora partly dark, tip pale; tips
of tarsal joints black; inner spur of hind tibiae about four-fifths of the
basitarsus.

Length of fore wing 5 to 7 mm.

Allotype from Hatillo, Puerto Rico, January ; others from Mayaguez
in November and Cartagena in May, both Puerto Rico, all through
Mr. Ramos.

PSAMMOCHARES PARSONSI Spec. nOV.

Body, legs, and antennae wholly deep black; wings mostly black, but
paler in third discoidal cell and toward base both in front and behind;
hind wings strongly fumose.

Clypeus over three times as broad as long, nearly truncate below,
a few hairs on lower part; ocelli in a low triangle, the laterals a little
nearer to the eyes than to each other; a median line above base of
antennae, latter slender, third joint equal vertex-width, last joint only
about one-half of third joint; vertex with a few long, erect hairs,
shorter black hair on front.

Pronotum broadly arcuate behind; mesonotum with a few long hairs
each side, pleura with only scattered short hairs; propodeum short, no
distinct median groove, with rather short hair above.

Abdomen hairy toward tip below, above with stiff bristles just be-
fore tip, otherwise a few hairs on venter near hind border of segments.

Femora smooth, without hairs; front tarsi with a distinct, though
short comb, a spine at middle of the second tarsal joint; hind tibiae
quite heavily spined; two irregular rows above, some as long as the

184 bulletin: museum of comparative zoology

width of the joint; inner spur of hind tibia more than one-half of
basitarsus.

In fore wings the marginal cell is about its length before tip ; second
submarginal cell longer below than high, one-third narrowed above,
receiving the first recurrent vein near tip; third submarginal cell only
a little longer than second and likewise narrowed one-third above,
receiving the second recurrent vein (evenly curved) beyond the
middle. In hind wings the anal ends beyond the fork.

Length of fore wing 10 mm.

Females from Buenos Aires, Trinidad Mts., Cuba, 17 to 23 June
(C. T. Parsons).

In appearance like P. pcrpilosus but easily separated by absence of
hairs on femora and very much less hair on body. Type M. C. Z.
No. 25742.

NOTIOCHARES ANTILLANA spec. nov.

Male. Very much like AT. cvbensis, wholly black, with blue and
violet reflections, and agreeing closely in structure to that species.
The only difference of importance is that the male genital plate, which
is N . cubensis ends in a sharp-pointed tooth each side, in N. antiUana
ends in short, rounded lobe, alike in all three specimens.

Length of fore wing 8 to 11 mm.

From Barbados, August to December (Spencers). Type M. C. Z.
No. 25743.

Episyron cressoni Dewitz

The male is black; a white spot on each side of clypeus; a broad
orbital streak yellowish white, reaching nearly to vertex; a narrow
yellowish band across tip of pronotum, a white spot each side on basal
part of third abdominal segment, the last dorsal segment somewhat
rufous, but white at tip; the mid femora rufous except base and tip,
the hind femora rufous except basal third, and the hind tibiae rufous
except extreme tip, spurs are pale; inner one of hind tibia nearly equal
to basitarsus.

Length of fore wing 6 mm.

Allotype from Mona Island, April (Ramos).

Priocnemis ursula spec. nov.

Body black, much of it covered with a fine sericeous pile or bloom;
clypeus, mandibles, basal joint of antennae, and a few joints beyond

banks: psammocharidae IS.")

yellowish, rest of antennae brown; legs black; wings hyaline, venation
black.

Clypeus above about as broad as vertex, nearly evenly rounded
below, fully two and one-half times as broad as long, with a few fine
hairs; front with fine, pale hairs, and vertex with some longer ones;
ocelli in nearly equilateral triangle, laterals a little (but not much)
nearer to each other than to the eyes; a short median groove above
base of antennae, latter slender, second plus third joints not as long
as vertex-width; pronotum scarcely constricted behind, with a broadly
arcuate hind margin ; mesonotum not humped in middle, scarcely hairy,
scutellum and metanotal lobe with erect black hair; propodeum with
rather long, fine, pale hairs above.

Abdomen hairy only toward tip and somewhat beneath; legs
moderately slender, hind tibia above with about ten teeth with a
short spine, and a row of spines on outer part; inner spur of hind tibiae
two-fifths of basitarsus.

The fore wings have the marginal cell rather longer than space to the
tip of wing, three times as long as broad, outer side nearly straight;
second submarginal cell oblique, below nearly twice as long as high,
receiving the first recurrent vein beyond the middle; the third sub-
marginal cell plainly longer than second, but not as much wider as in
many species, outer margin bent near median vein, receiving the
second recurrent vein (almost straight) much before middle; in hind
wings the anal vein ends much before the fork.

Length of fore wing 6.5 mm.

Female from Villa Altagracia, San Domingo, July (Darlington)
Type M. C. Z. No. 25741.

In general very similar to P. salti Bks. which differs in the yellowish
tip to the abdomen.

Priocnemis arioles spec. nov.

Head and thorax black, abdomen reddish except black tip, the fifth
segment above bluish, thoracic notum bluish, legs black, femora and
tibiae somewhat bluish; antennae black; fore wings nearly evenly
infumate, slightly darker toward costal tip than behind, hind wings
slightly smoky.

Clypeus no wider than face below, with a few pale hairs, about two
and one-half times as broad as long, truncate below; vertex as broad
as face below, scarcely convex, with a few erect hairs each side, none
on front; ocelli in a rather low triangle, the laterals much nearer each

186 bulletin: museum of comparative zoology

other than to eyes ; a groove from anterior ocellus to base of antennae.

Pronotum plainly contracted behind, the margin slightly arcuate;
mesonotum almost humped in middle, with one or two erect bristles
each side and a pair on scutellum, pleura bare; propodeum with only
faint, erect, pale hair above, a whitish pile across base, and an elongate
spot each side toward hind margin, surface minutely granulate, no
median groove. Abdomen hairy on venter and apical third above.
Legs not especially slender, mid tibiae with two rows of short spines
above, hind tibiae with a row of seven or eight oblique teeth, a spine
beyond each, and an outer row of short spines ; inner spur of hind tibia
hardly two-fifths of basitarsus.

Marginal cell of fore wings about its length from wing-tip, two and
one-half times as long as broad, outer side convex; second submarginal
cell a little longer than high, but little narrowed above, receiving the
first recurrent vein at middle ; third submarginal cell much longer than
second and much wider behind, one-third narrowed above, receiving
the second recurrent vein (slightly curved) a little before middle; in
hind wings the anal vein ends much before the fork.

Length of fore wing 6 mm.

Female from Constanza, Valle Nuevo, San Domingo, 21 August,
3- to 4000 ft. (Darlington). Type M. C. Z. No. 25740.

The Antillean species with a more or less reddish abdomen can be
tabulated as follows : —

1. Fore wings hyaline with two broad dark bands pulchellus

Fore wings without bands 2

2. Thorax wholly reddish christophei

Thorax black 3

3. Third submarginal cell no longer than second, higher than long parous

Third submarginal cell much longer than second, and much longer than

high 4

4. Wings hyaline; hind legs very long, body about 4.5 mm dowi

Wings plainly infuscate, hind legs not so long, body 7 mm ariolcs

Priocnemella domingensis spec. nov.

Head, propodeum, and pleura black, thorax and abdomen above
blue. Fore wings black, violaceous, hind wings infuscated, also with
violet reflections; antennae and legs black, the femora and tibiae
above, and the front coxae iridescent bluish, also basal joint of anten-
nae above.

Clypeus large, extending laterally under the eyes, with long, black

banks: psammocharidae 187

hairs, lower margin narrowly smooth and coming to a blunt point in
middle; front and vertex with long black hair, one each side on vertex
very long; third antennal joint equal to vertex-width. Ocelli close to-
gether, the laterals more than twice as close to each other as to the
eyes; groove from anterior ocellus to the antennae. Scattered erect
black hairs on notum and scutellum, short hair on pleura; pronotum
almost angled behind. Propodeum minutely, transversely striate,
more distinctly so than in P. violaceipes, above and on sides with
sparse, erect black hair.

Abdomen hairy near tip and below, scarcely so near base. Legs
slender, spined as in P. violaceipes, the mid and hind tibiae with two
rows of erect, short black spines, no teeth, and evenly short spines on
the tarsal joints; claws with an erect, small tooth; long spur of hind
tibia about two-fifths of the basitarsus.

Venation of wings as in P. violaceipes; the marginal cell long and
pointed; second submarginal oblique, about one and one-half times as
long as high, receiving the first recurrent vein near middle; the third
submarginal cell much longer, about one-third narrowed above, re-
ceiving the second recurrent also near middle, this recurrent is more
sinuously curved than that in P. violaceipes.

Length of fore wing 10.5 to 11 mm.

Two females from near and southeast of Constanza, Valle Nuevo,
San Domingo, August, 3 to 7000 ft. (Darlington). Type M. C. Z. No.
25739.

Except for the striking difference in color of thorax and abdomen,
and slight differences in structure, it is practically the same as P. vio-
laceipes of Cuba.

Priochilus

A few Neotropical species have much the appearance of Priocnemis;
however, the last joint of hind tarsi has spines beneath, but not later-
ally; the hind tibia has no real teeth like Priocnemis, but spines in
rows, •some as long as the diameter of the joint ; the claws are cleft, the
inner part broader than the outer part, and somewhat obliquely trun-
cate; the palpi are slender as in Priocnemis.

Pompilus nobil is Fabr. is the genotype; Salius opacifrons Fox from
Jamaica goes in the genus, although smaller than most others. In
South America there are regius Fabr., diversus Smith, scrupulus Fox,
sericeifrons Fox and others. Mr. Williams in his paper "Studies in
Tropical Wasps", 192S, p. 141, calls attention to nobilis and regius as
possibly forming a new genus.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 5

SCIENTIFIC RESULTS OF A FOURTH EXPEDITION
TO FORESTED AREAS IN EAST AND CENTRAL AFRICA

VI
ITINERARY AND COMMENTS

By Arthur Loveridge

With Four Plates

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

July, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1, 2,
3 and 4 have appeared and of the Memoirs, Vol. I to LVI.

These publications are issued in numbers at irregular intervals.
Each number of the Bulletin and of the Memoirs is sold separately.
A price list of the publications of the Museum will be sent upon
application to the Director of the Museum of Comparative
Zoology, Cambridge, Massachusetts.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 5

SCIENTIFIC RESULTS OF A FOURTH EXPEDITION
TO FORESTED AREAS IN EAST AND CENTRAL AFRICA

VI

ITINERARY AND COMMENTS

By Arthur Loveridge

With Four Plates

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

July, 1944

No. 5. — Scientific Results of a Fourth Expedition to Forested Areas in

East and Central Africa

VI

Itinerary and Comments

By Arthur Loveridge

INTRODUCTION

For various reasons it would appear inopportune at the present
juncture to devote the time necessary to elaborating the ecological and
zoogeographical data accumulating from this, and previous, expedi-
tions so generously sponsored by the John Simon Guggenheim Me-
morial Foundation of New York.

On several occasions, however, correspondents in Africa have
written requesting specific information about, or the latitude and
longitude of, some of the more obscure localities mentioned in the
reports already printed, so that it would seem advisable to publish
the itinerary without further delay. A synopsis of it has, indeed, been
furnished in the caption accompanying Plate 1, and it has not been
thought necessary to enlarge on this in respect to places where only
a single night was spent, with consequently little collecting.

For the others I am now supplying the latitude, longitude, altitude,
the all-important meteorological conditions prevailing at the time of
my visit, duration of stay, and position of camp. This in turn is fol-
lowed by a brief survey of the vegetational environment, or a reference
to where such an account has been published elsewhere.

The composition of the fauna in relation to life zones receives atten-
tion, more particularly for localities in Tanganyika Territory where it
has greater significance on account of our knowledge being more com-
plete. When one contemplates the vastness of the largely trackless
forests in Uganda, one feels that the results accruing from a stay of
two weeks in areas of from 120 to ISO square miles in extent, is alto-
gether too inadequate to justify one in embarking on detailed specu-
lation as to what does, or does not, occur.

Species of exceptional interest receive mention where it is thought
that they may assist in visualizing the environment, or where, because
of their rarity, other zoologists visiting the locality would wish to
know of their occurrence. Special reference is made to forms of which
the type happens to have come from the particular locality under
discussion.

192

bulletin: museum of comparative zoology

The groups already reported upon in this Bulletin are as follows :

Speci-

New to

Volume

Pages

Class

mens

Species

M.C.Z.

89

145-214

I Mammals

812

116

40

89

215-275

II Birds

809

240

10

91

183-234

III Crustacea

389

20

4

91

235-374

IV Reptiles

1802

151

17

91

375-436

V Amphibians

1081

77

10

It has been somewhat of a disappointment that no one has been able
to undertake the identification of, or to report upon, the fairly exten-
sive collections of other invertebrate groups such as mollusks, myria-
pods, and earthworms. In addition to zoological material, we pur-
chased from Baamba and Banyaruanda tribes 1013 ethnological
items, with a gross weight of almost one ton, which were sent by
truck and train over a thousand miles to the port of shipment.

For part of the data regarding altitudes, temperature, composition
of forests, etc., I am indebted to "Uganda" by Messrs H. B. Thomas
and R. Scott, 1935, and to the "Handbook of Tanganyika Territory"
by G. F. Sayers, 1930; both mines of reliable information.

There remains only to again express my deep gratitude to the John
Simon Guggenheim Memorial Foundation, and to Dr. Thomas Bar-
bour, Director of the Museum of Comparative Zoology, for making
possible this expedition, of whose harvest of herpetological material I
am now able to take full advantage in the revisionary studies of
African lizards upon which I am engaged.

ITINERARY
UGANDA

Mabira Forest, Kyagwe (Chagwe). 0°24' N., 33°0' E. Alt. 4000 feet.

As the rains were still continuing we accepted the generous offer of
Mr. L. Jarvis to occupy the late Major Cuthbert Christy's old house
at Mubango, a rubber and coffee plantation in the southern end of
the forest, five miles north from Najembe, which is twenty miles
southeast of Jinja on the Jinja-Kampala road.

Arrived at noon on November 5, and left on the 21st.

loveridge: African itinerary and comments 193

A heavy shower amounting to .76 inches fell on the afternoon of our
arrival, .17 the following day, and .75 on the 14th. Actually a shower
or two occurred almost daily but the precipitation was too small to
be registered. The average annual rainfall at Mubango is 55 inches.
The daily temperature at 7 a.m. averaged 66°, and at noon 80°.

We had come here largely because Major Christy's labours had made
the Mabira, which covers 120 square miles, type locality for Leptopelis
n. christyi, Hylambates verrucosus, Miodon g. christyi, and Aparallactus
christyi (= A. modestus). Of these we were successful in securing only
the first and last.

Despite what appeared to me to be ideal conditions, I personally
captured only three snakes during as many days primarily devoted to
searching for them both in the forest and along its edge, where drifts
of leaves between buttress roots and rotting logs provided suitable
retreats. On four succeeding days I employed an allegedly expert
snake-catcher who got nothing during that entire period. Yet some
two-score labourers, engaged in clearing undergrowth on the planta-
tion, brought in snakes at the rate of nearly two a day, and of these
the first eleven snakes represented ten different species ! This astonish-
ing variety continued until we actually had eighteen species for a total
of twenty-seven snakes. Of these one — Dipsadoboa unicolor — was new
for Uganda, another — Bothrophthalmus I. lineatus — constituted the
most easterly record for the species, though its presence here had been
forecast by Lt. Col. Pitman.

Pitman (1934, Uganda Journal, 1, pp. 7-16) has furnished an inter-
esting account of the Mabira and its fauna and flora, rendering un-
necessary further details here. Lizards, with the exception of the
arboreal Algiroides africanus, were disappointing, being of widespread
species. A single chelonian was encountered.

The dense undergrowth rendered bird collecting extremely difficult
and only twenty skins were prepared during the fortnight. More than
a score of mammals were taken, however, including such forest forms
as Cercocebus, Cercopithecus, Hcliosciurus, Tamiscus, Protoxerus
Cephalophus and an arboreal pangolin which was described as a new
race (Phataginus tricuspis mabirae).

Budongo Forest, Bunyoro. 1°42' N., 31°24' E. Alt. 4000 feet.

Camp was pitched in the mahogany nursery at BISU beside the
Buchanan Saw Mills, so that full advantage might be taken of the
three-mile-long corridor cut into the heart of the forest.

Arrived in afternoon of November 22, and left on December 7th.

Heavy downpours occurred at intervals during our stay for the rain-

194 bulletin: museum of comparative zoology

fall here is well distributed throughout the year, the annual average
being 65 inches. During our visit the daily temperature at 7 a.m. aver-
aged 60°, and at noon 82°.

Situated on the north-north-westerly slope above the escarpment
towards the northern end of Lake Albert, Budongo is rich in mahogany
and consequently considered the most valuable of all Uganda's lovely
forests. Ironwood trees are also plentiful, in some areas forming over
50% of the canopy and being largely responsible for the maintenance
of its evergreen appearance. The forest survey reveals that the second
story stratum, attaining about 80 feet, is composed in part of such
genera as Celtis and Funtumia spp., while the third story is scanty.
Bordering the forest and at the edge of clearings, a dense, often im-
penetrable, undergrowth of bush has sprung up; but in the depths of
the forest where the canopy was contiguous one might wander at will
or chase the frogs which went leaping over the damp leaves that car-
peted the forest floor.

Our visit added four amphibia to the Uganda list, the most in-
teresting being Rana christyi, for Budongo, covering about 180 square
miles, shows very close faunal relationship with the great Ituri Forest
lying away to the west in the Belgian Congo.

This is borne out by a number of interesting herpetological records
recently resulting from the collections made here by Mr. W. J. Eggel-
ing, Conservator of Forests, whom we had the pleasure of meeting.
This West African complexion was reflected by 7 of the 9 species of
snakes, and 4 of the 7 kinds of lizards collected, though one of these —
Algircrides africanus, is more central.

Budongo is type locality for 7 valid, and 2 invalid, races of birds,
only one of which I succeeded in getting, this was the gorgeous little
forest kingfisher {Myioceyx lecontei ugandae) which obligingly alighted
on a stump in camp.

Of the 10 species of mammals secured, only 4 can be considered
definitely western, the best being a huge horseshoe bat (Hipposideros
cyclops) netted in a clearing deep in the forest, where was also the great
tree on which I shot four small chipmunk-like squirrels (Tamiscus
alexandri). Here too we trapped a strange harsh-furred mouse (Lop-
huromys aquilis subsp.) whose race proved unidentifiable; perhaps a
series might reveal it as an undescribed subspecies.

Kibale Forest, Tow. 0°24' N., 30°24' E. Alt. 4200 feet.

Camp was made near the Duta River where it crosses the new road
being cut through the forest about 20 miles southeast of Fort Portal.

Arrived late on December 8, and left on the 18th.

loveridge: African itinerary and comments 195

Thunderstorms, accompanied by rain, occurred almost every after-
noon, while on two days an exceptionally heavy downpour continued
on throughout the night. The annual average rainfall is 56.87 inches.
During our stay the daily temperature at 7 a.m. averaged 59°, and at
noon 77°.

Kibale, together with the adjacent Itwara and Muhangi Forests,
covers about 276 square miles. The northern portion, traversed by
the Kampala-Fort Portal Road, is impenetrable on account of the
dense thicket undergrowth. Acting on the advice of Mr. W. J. Eggel-
ing, therefore, we selected the southern portion as being more open,
the enormous buttressed trees sufficiently dense as to prohibit under-
growth in many areas. After penetrating the forest about a mile-and-
a-half Ave had to camp for the Dura River was still unbridged; beyond
the river the road continued for another two-and-a-half miles through
beautiful forest, and it was along this stretch that most of my collecting
was done.

Excursions along this road during and immediately after heavy rain
were productive of sundry snails, slugs and worms, the latter very
numerous and one species frequently a couple of feet in length. This
larger species, with a diameter as great as one's thumb, was remarkably
iridescent. Beyond the forest proper these worms were encountered
in muddy soil beside a stream which flowed through a patch of palm
forest much frequented by elephant. Butterflies in bewildering variety
surpassed anything which I had seen anywhere in East Africa.

It was disappointing, therefore, to find the herpetofauna extremely
scarce, not merely in species but in individuals also. All our efforts re-
sulted in obtaining only 7 species of reptiles and 7 of amphibians,
though there was some compensation in the fact that all were exclu-
sively sylvicoline species with the exception of Agama atricollis and
Bujo r. regularis which are equally at home in the savanna. The head-
man of the gang engaged in felling trees with which to bridge the
Dura, informed me that they encountered from 1 to 3 snakes only per
month. During the week the only one they brought me was a Rham-
nophis a. elgonensis. It was this paucity of poikilothermous creatures
that decided me to leave after ten days instead of remaining three
weeks as originally planned.

Monkeys were the dominant form of animal life and no fewer than 5
species, of which I collected 4, were observed feeding within a hundred
yards of my tent, the source of attraction being the huge spherical fruit
borne by a certain tree. Of a dozen species of mammals collected, a
squirrel (Funisciurus p. victoriae) was the only novelty.

196 bulletin: museum of comparative zoology

Bundibugyo, Tow. 0°41' N., 29°56' E. Alt. 3200 feet.

Stayed at the comfortable rest camp at Saza headquarters for
Bwamba District.

From the evening of December 19 to 26th.

A few showers occurred during the week. No annual rainfall record
was available as the area has only been opened up recently, but statistics
are being collected at the mission. During our stay the daily tempera-
ture at 7 a.m. averaged 55°, and at noon 62°.

This area, marked on the Uganda Survey Map of 1928 as Bwamba
Forest, is now largely covered by native gardens, including plots of
coffee, cotton, and bananas, with extensive rice cultivation in the
swampy areas. Elsewhere rank grass fifteen feet high, scattered trees,
clumps of indigenous oil palms (Elacis guinecnsis) and patches of
forest. The latter, said to amount to 140 square miles, is an extension
of the great Ituri Forest on the opposite bank of the Semliki River.

Naturally, therefore, sylvicoline snakes are plentiful and a large col-
lection could have been made had we been able to stay longer. At
first, however, the Baamba mutilated them so badly that many had to
be rejected and it was only during the last few days that they began
to arrive in good condition. At least 14 of the 16 species secured were
forest forms, of which Boiga puherulenta and Pscudohaje goldii were the
second records for Uganda and Miodon g. collaris the first for this race.

Three of the five kinds of lizards taken were widespread savanna or
eastern forest forms. No attempt was made to collect birds and the
only ones preserved were the swamp-loving rail (Sarothrura p: cen-
tralis) and warbler (Prima m. immutabilis) .

Except for a young duiker, all 42 of the mammals preserved were
rodents representing 9 species which reflected our position in the west-
ern foothills of the Ruwenzori Mountains overlooking the Semliki
River, some being montane species, others races of savanna rats.

The Baamba, who give their name to this region, are a forest tribe
of semi-pigmy stock who would undoubtedly prove helpful to a natural-
ist working in this country; unfortunately, as already indicated, de-
forestation has resulted in encroachment by many savanna forms
which would probably preponderate among specimens brought in.
The big tree cobra was obtained by Bakonjo, a primitive Bantu people
living on the slopes of Ruwenzori. Most of the cultivation was being
done by Batoro, while administration was largely in the hands of
Bahima, the tall Hamitics and former overlords. We found all of these
people most friendly and only too eager to dispose of ethnological
material, much of it superseded by changing customs.

loveridge: African itinerary and comments 197

Bugoye, Ruwenzori Mountain*. 0°17' N., 30°13' E. Alt. c. 5600 feet.

( lamped in the rest camp enclosure.

December 26 to 28th, 1938, and January 21 to 24th, 1939.

Occasional showers. We had left our thermometer hanging on a tree
in the heart of Kibale Forest so thereafter we were unable to record
temperatures.

After leaving Bundibugyo we drove over the northern end of the
Ruwenzori Range, which is about 30 miles in maximum width, through
Fort Portal, skirted the eastern flank of the 65-mile-long range, then
turned off the main road and took the side road to Bugoye where it
terminates.

Bugoye, at the southeastern foot of the range, is not only the type
locality of Chamaeleo h. ellioti, of which we obtained a good series, but
the jumping-off place for any ascent of the mountain (16,800 feet)
from this direction. This was the route followed by the British Mu-
seum Expedition of 1908, an expedition which resulted in several score
of new vertebrates. It was partly in the hope of securing topotypes,
at least of the reptiles, that we now planned to visit the two most
readily accessible sites occupied by Woosnam and his party.

We stayed at Bugoye only as long as it was necessary to get to-
gether sufficient porters for the ascent, while on the return journey I
was fully occupied with packing the specimens obtained on the moun-
tain before the lorry should arrive to pick us up.

Consequently little collecting was done in this uninteresting spot
whose surrounding slopes, eroded by constant rain, are clothed only in
sparse grass with thickets and, lower down, acacia or orchard forest.

Mubuku (Mobuku) Valley, Ruwenzori. 0°24' N., 30°0' E. Alt. 6800 feet.

Camped in deep forest between the Mubuku and Mahoma Rivers,
near foot of Nyinabitaba Ridge.

Arrived about noon on December 29, 1938, and left January 9, 1939.

Rain fell on every day except one. The worst aspect of the situation
was the absence of sunshine due to overhanging clouds which cast an
evening-like gloom and stillness over the forest for hours on end. Sun-
day was the best day with 7 hours of sunshine, on Monday about half
that amount, the rest averaged between 1 and 2 hours only. The tem-
perature at midday was probably from 55° to 66°. Under such condi-
tions collecting was difficult, birds were silent except during the brief
hours of sunshine when they were engaged in feeding on the forest
canopy well out of range. Drying of skins and other specimens became
a constant nightmare.

Despite the sodden state of the undergrowth only a toad and 3

198 bulletin: museum of comparative zoology

species of frogs were encountered, all the latter (Rana f. angolensis,
Phrynobatrachus graueri, and Hyperolius ? alticola) previously recorded
from the mountain.

Reptiles were limited to 5 species of which two were new for Ruwen-
zori, these were Cnemaspis a. elgonensis and Lygosoma g. graueri. This
little pentadactyle skink, together with the four-toed L. mcleagris, of
which we secured topotypes, were relatively plentiful. Topotypes were
also obtained of the chameleons (C. j. johnstoni and C. xenorhinus) ,
the latter a great rarity apparently dwelling in the forest canopy.
Naturally C. b. rudis, obtained by the British Museum Expedition at
10,000 feet, was not met with at the lower level but I was surprised to
see no Lacerta jacksoni, a species that has been taken at 8500 feet.

A pair of crimson-winged plantain-eaters, presumably Ruwenzoror-
nis, were seen near camp one day but out of gunshot in the tallest
trees. A mountain buzzard (Buteo oreophilus) visited our lonely camp
and fell to my gun, while topotypes of half-a-dozen passerines were also
collected.

Mammals were decidedly scarce, not a bat was seen, and, though a
net was spread across a suitable clearing, nothing was taken. Colobus
were seen only on the march to this camp and all the monkeys seen
were two solitary males of Cercopithecus m. stuhlmanni, the one col-
lected would be almost a topotype of the synonym carruthersi which
was first obtained a thousand feet higher up the mountain. During
our entire stay squirrels (Heliosciurus and Tcnniscus) were observed
four times only . Of three species of rodents preserved one was a topo-
type of the interesting mouse (Hylomyscus d. denniae). A red duiker
was seen once and heard a couple of times but attempts to collect it
failed. Not a trace of any carnivore was noted.

Mihunga Ridge, Ruwenzori. 0°21' N., 30°3' E. Alt. 6000 feet.

On the outward journey we had camped for the night beneath the
fig tree — on the upper Mihunga Ridge and collected a few specimens ;
returning we passed it to pitch our tents on the lower Mihunga Ridge
on the very site formerly occupied by the British Museum Expedition
in 1908.

Night of December 28, 1938. Then from January 9 to 19th, 1939.

We had fled from the Mubuku Valley camp on account of adverse
climatic conditions, now, though only a few miles distant as the crow
flies we seemed to be in another world. On this largely treeless spur,
bounded on the north by Weria Ravine, on the south by the Kanyon-
gorogoro Ravine, we were able to profit by the hot sunshine which
lasted for the first three days until 3 p.m. and was then followed by

loveridge: African itinerary and comments 199

showers. No rain at all occurred on the four days following while dur-
ing the last week, though the sky was often covered by fleecy clouds
through which the sun had difficulty in breaking, dark clouds formed
only in the late afternoon and then frequently dispersed without preci-
pitation in our vicinity, though at times rain might be seen falling
elsewhere.

Collecting was not confined to the 6000-foot ridge for excursions
were made to the forested heights a 1000 feet higher as well as to the
foot of the ridge where it tapered out into the swamps of the Mubuku
Valley. For descriptions of this and the last camp the reader is re-
ferred to Woosnam's account (1910, Trans. Zool. Soc. London, 19, pp.
5-24), and to some important comments by Allen and Loveridge
(1942, Bull. Mus. Comp. Zool., 89, p. 148).

Almost all of the amphibians (Bufo r. regularis, Leptopclis n. christyi
and Phrynobatrachus graueri) and most of the 9 species of reptiles
taken, came from the swamp about 500 feet lower than our camp.
Here, high in the papyrus, %were coiled the green vipers (Atheris n.
nitschei) topotypes of A. woosnami of which we had come in search.
Here also we obtained the first Uganda example of the dwarf chameleon
(Brookesia s. boulengcri).

Of 30 kinds of birds collected, 4 were topotypic, while each morning
our traps yielded some topotypic rodent until we had 10 of the species
collected at Mihunga by Woosnam and his associates; 3 others were
almost topotypic the types having been taken at higher altitudes.
This region appears to be as rich in mammals as it is poor in amphi-
bians for, in addition to the score of species collected, we found three
sleeping platforms built by chimpanzees in a tree in Weria Ravine.
The Bakonjo informed me that these platforms are made by itinerant
chimpanzees which come from the forests of the Mubuku in search
of bananas and, when benighted, construct a platform. We saw one
such individual while wTe were engaged in collecting in the swamp.

Nyakabande, Kigezi. 1°44' N., 29°45' E. Alt. 6925 feet.

Stayed at the commodious rest camp.

Arrived at noon on January 25 and left on 30th. Returning from
Mushongero on February 4, we had to wait for a lorry until the 8th.

Though there was relatively little sunshine, the weather was
generally fine except for a few heavy showers.

Nyakabande is situated in a lava-strewn plain, much of which is
industriously cultivated by the Banyaruanda who gather up the larger
blocks of lava and pile them on the periphery of each small plot, where
they serve as a windbreak. There is no water in the immediate

200 bulletin: museum of comparative zoology

vicinity, and perhaps it is just as well not to visit the foul pool from
which your water supply is likely to come. Presumably it was from
this pool that the four species of amphibia obtained here came.

Nyakabande is about six miles from Kisolo (the rendering given by
the Uganda Survey on its map A 530 of 1928, often misspelled Kisoro
or Kissolo), type locality of the toad I named Bufo r. kisoloensis,
whose validity can no longer be maintained as a result of the fresh
material obtained at Nyakabande and Mushongero.

The reptile fauna of these two localities is essentially similar as we
got only three species at the former not taken also at the latter, which
is reached by a very arduous climb over the mountains.

In the rest house grounds we found a weaver (Ploceus n. graueri)
nesting in a vociferous colony of the superficially similar P. c.feminina.
At Mushongero we obtained a topotype wagtail. (Motadlla c. wellsi)
and a pair of flycatchers (Alseonax a. ruandae) which race was de-
scribed from Bufundi on nearby Lake Bunyonyi.

At both localities the Banyaruanda apparently eat the huge mole
rats (Tachyoryctes ruandae), for they brought in as many as I would
take. The species was first collected on Mt. Muhavura which is in full
view of the rest house at Nyakabande. More valuable was an otter
(Lutra m. tenuis) which I shot in the lake near Mushongero, for it was
from this lake that Hinton described the synonym L. m. mutandae.

The main object of our stopover at Nyakabande, however, was to
purchase ethnological material from the teeming tribes inhabiting this
upland plain. Collectively known as Banyaruanda, they apparently
consist of a small admixture of Hamitic overlords, the cattle-owning
Batusi and Bahororo. The bulk of the population consisting of Bantu
agriculturists known as Bahutu, while a few semi-pigmy Batwa are
present, chiefly in the vicinity of the lakes.

Mushongero, Lake Mutanda, Kigezi. 1°46' N., 29°41' E. Alt. 5925 feet.

Stayed at the rest camp, which is situated on a little peninsula pro-
jecting from the east bank near its northern end.

Arrived late in the afternoon of January 30, and left early on
February 4th.

Frequent rainstorms swept across the lake whose mountain-girt
northern half appeared to attract and hold the heavy black clouds
which overhung it during much of our brief visit. When the sun was
hidden it was decidedly chilly.

Except for its extensive papyrus swamps, Lake Mutanda offers little
indication of its proximity to the equator. In fact with its numerous
tree-covered islets and purplish mountains its general appearance is

loveridge: African itinerary and comments 201

not unlike that of a Scottish loch. The brambles, bracken, and short,
wind-swept grass clothing the lower slopes of the mountains all suggest
that it was with good reason that these uplands have been called the
Switzerland of Central Africa.

As for the name Mushongero (misspelt Mushungero on all my
labels), I suspect that there is some connection between it and Mus-
hungwe, the Lugezi name for leach, for these loathsome creatures
swarm in the shallow waters in the vicinity of the rest camp. Crabs
(Ngara: Lugezi) were also not uncommon, and the series secured have
been described by my colleague, Dr. F. A. Chace Jr., under the name
of Potamon (Geothelphnsa) mutandensis.

The waters of the lake were teeming with Xenopus 1. bunyoniensis,
originally described from nearby Lake Bunyonyi, but search of the
papyrus both by day and night residted only in the capture of a single
example of the genus Hyperolius, and that unidentifiable! The most
perplexing aspect of the amphibian fauna was the intermediate condi-
tion of many Rana fuseigula, some, as one would expect from the ter-
rain, were typical, others showed the longer hind limb of the sylvicoline
race chapini. Three other ranids taken here also seemed to suggest
that some deforestation had taken place.

This was again the case with a couple of the dozen species of rep-
tiles collected. \Ye had gone to Mushongero primarily in search of a
blind snake obtained there by Col. Pitman; hoping that an adequate
series of the creature might settle its uncertain status. In this we were
successful, the 9 examples undoubtedly referable to Typhlops blanfordii
lestradci, originally described as a fidl species from nearby Ruhengeri
but obviously related to blanfordii of the Ethiopian highlands; another
link with the latter region is the presence at Mushongero of the little
slug-eater (Duberria I. abyssinica).

BELGIAN RUANDA

Kiraga near Kisenyi, Lake Kivu. 2°38' S., 29°1S' E. Alt. c. 5800 feet.

Camp was pitched in a plantation of eucalyptus through which the
road passes about three miles above Kisenyi, a township situated on
the shores of Lake Kivu at 4800 feet.

Arrived in afternoon of February 8, and left early on the 13th.

Some showers.

Most of our collecting at Kiraga was done along the banks of, and
in the ravine cut by, the Kisenyi River, which cascaded over falls just

202 bulletin: museum of comparative zoology

below our camp. The ravine was more or less choked by luxuriant
growths of grass and sedge, but in patches cleared for gardens we
turned over piles of vegetable debris and sought our quarry in the ex-
tensive plantations of bananas.

Only 4 species of amphibia and 6 of reptiles were taken, none of
especial interest with the possible exception of Lacerta jacksoni which,
in the absence of trees, has adapted itself to a terrestrial life. By far
the most important of half-a-dozen mammalian species were a pair of
skunk-like zorillas (Poecilogale a. doggetti).

As we were debarred from killing anything but "vermin" on ac-
count of our permit for scientific collecting not having arrived from
Stanleyville, we caught the first boat from Goma (Ngoma), type
locality for a burrowing viper (Atractaspis schoutedeni), which appears
to be based on a slightly aberrant example of the widespread A. irre-
gularis, which we got.

BELGIAN CONGO

Mamm Bay, Idjivi Island. 2°12' S., 29°0' E. Alt. 4788 feet.

Camped on the lawn of Mons. van der Berck v. Heemstede's estate.

Arrived at dusk on February 14 and left on 16th, returning March
6th.

Our arrival was greeted by torrents of rain and waves lashed by a
gale.

Wading in thigh boots among the sedges of the Bay, and aided by a
flashlight, I was able to capture topotypes of Ilyperolius kivuensis,
Icwidjwicnsis, kandti, and macrodactylus, and show that actually only
two sexually dichromatic species are present. The types had been
collected by the poet Kandt, who had made his home on Idjwi Island.

Upper Mulinga River, Idjwi Island, c. 2°8' S., 29°3' E. Alt ^500 feet.

Camped beside a footpath where it crosses a stream known as the
Upper Mulinga. This, I imagine, was about 900 to 1000 feet below
the 800 metre summit of the mountain, which dominates the island.

Arrived on February 16 and left at noon on March 6th.

The weather was very varied, we enjoyed much sunshine when it
would be quite hot; violent thunderstorms accompanied by lashing
rain were not uncommon, however, and always resulted in a consider-
able drop in temperature.

1 Not 4500 feet as printed on p. 149 of the report of Mammals, 1942, Bull. Mns. Comp. Zool.,
89, pj>. 147, 214. I am indebted to Dr. J. P. Chapin for pointing out this error.

loveridge: African itinerary and comments 203

Day after day I made excursions up the mountain to the extensive
remnants of magnificent forest, even then being destroyed by natives
contrary to regulations; policing of such remote spots being difficult.
Apart from some small patches of forest, our immediate vicinity con-
sisted of pasture land, millet fields, dense patches of sedge, and
swampy areas through which meandered small rivulets.

It was in this latter habitat that I collected most of the 8 species
of amphibia taken, but the choicest of all, a tiny, long-fingered male
Arthrolcptis xenochirus, was brought to me by a native lad. Only 2 of
the species were savanna forms.

Of the 25 species of reptiles collected, all but a fourth were of forest
association and included such rarities as Miodon g. graueri and Algi-
roides vaucreselli. The island, variously spelt Idschwi, Kwidjwi, and
Kidjwi by the Germans, is type locality for Mabuya m. kicidjwicnsis
and Lygosoma blochmanni. Over fifty of each were preserved, suffi-
cient to demonstrate that the former does not differ structurally from
true M. m. maculilabris, and that the latter is constantly three-toed.
The local chameleon appeared to differ sufficiently from its continental
congeners to be named C. d. idjwiensis.

Also new were 8 examples of a warbler (Apalis eidos), and we secured
topotypes of Barbatula kandti = Pogoniulus b. jacksoni, Colivs s.
kiwuensis and Spinus c. frontalis; in all 42 species of birds were col-
lected at this camp.

Two new races of rodents (Thamnomys v. kimiensis and Leggada b.
ablutus) were discovered, the former just behind my tent, the latter
differing from the typical race occurring on the Ruwenzori Moun-
tains. An interesting aspect of the mammalian fauna of this mountain
on Idjwi was that a third of the 21 species collected were referable to
races originally described from Ruwenzori; two others (Cercopithecus
m. schoutedeni and Lophuromys a. laticeps) were topotypes of local
forms.

TANGANYIKA TERRITORY

Ujiji, Kigoma District. 4°55' S., 29°42' E. Alt. 2800 feet.

Camped beneath the giant mangoes on the western fringe of the
town.

Arrived at noon on March 9, and left early on March 10th.

Occasional heavy showers following periods of sultry weather and
overcast skies.

As, from May 22 to 29, 1930, I had already visited this famous old

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Arab settlement on the shores of Lake Tanganyika, a description of
its features will be found in 1933, Bull. Mus. Comp. Zool., 75, p. 23.

The purpose of my visit was the same in both instances, viz. to ob-
tain examples of the rare Amphisbaena phylofiniens, known only from
the two cotypes described in 1905. On the first occasion I was unsuc-
cessful but learned from the natives that it was to be found in Ruanda,
a region of rice swamps bordering the Luiche River. Almost daily,
therefore, accompanied by two assistants, I tramped over to Ruanda,
pausing en route to inform each passer-by of our object. Not an am-
phisbaenid did we find, but as a result of our ceaseless talk 4 speci-
mens were brought in by natives. Since collecting all 7 East African
members of the family, I have come to the conclusion that the habitat
which these wormlike creatures require is one of moist sand or laterite
soil. There is no evidence which would justify the view that they are
survivals of a former forest fauna in this region.

While searching for amphisbaenids we secured a good series of a
Congolese skink (Scelotes t. hemptinnei) which had not previously been
taken in Tanganyika. Another first record for the Territory was a
"two-headed snake" {Chilorhinophis gerardi) though I had postulated
its occurrence at the southeast end of the lake in 1933. The Angola
race ornatum of Lycophidion capense was a further addition to the fauna
and, from a distributional point of view, an Aparallactus c. capensis is
interesting.

Additions to the fauna among the 9 species of amphibia taken, were
Rana m. venusta and Hypcrolius kivuensis; topotypes of H. udjijiensis
and argentovittis were also captured.

Kitaya, Southern Province. 10°40' S., 40°11' E. Alt. 300 feet.

Tents were pitched on the rest camp site on the bank of the Rovuma.

Arrived at noon on March 24th and left early on April 7th.

Torrential rains fell during our stay, necessitating the employment
of porters for the first nine miles of the return journey to Mikindani,
i.e. until past a depression of water-logged black cotton soil.

Kitaya is a village, and Liwale headquarters, on the north bank of
the Rovuma, (Rowuma; Ruvuma) River, fifteen miles inland from
Rovuma Bay. The River forming the southeastern boundary between
Tanganyika Territory and Mozambique. Livingstone safaried there
from Mikindani in 1866; the two places are now connected by a very
rough motor track of about 36 miles. The vegetation is typical of the
coastal belt; baobabs, orchard forest, and rank grass on the higher
ground, which is very sandy; rice fields and sedge (Setaria palmifolia)
fringed swamps on the lower.

loveridge: African itinerary and comments 205

Huge leeches are present in the swamps, ticks in the long grass.
Aedes, Glossina, Haematopota, Stomoxys, Tabanus, and many other
biting flies were an ever-present nuisance.

The purpose of our visit was to secure topotypical material of three
species of sedge frogs {Hyperolius eitrinus, microps, and Megalixalus
fiavomaculahis) collected by Kirk when he accompanied Livingstone.
In this we were eminently successful and able to prove that the last
named is really a Hyperolius, having a horizontal pupil in life. In
addition a series of an undescribed race of this genus (//. p. rovumae)
were collected and described.

We had to depend Aery largely on our own efforts, for the local
people — Konde and Yao — were strangely disinterested in bringing in
reptiles or mammals, in fact supplied only two of the latter. Crocodiles
as man-eaters play a prominent role in local village life. Soft-shelled
turtles (Cycloderma frenulum) were laying and in consequence may
have been more conspicuous than would normally be the case. The 13
species of snakes might almost have been selected as representative of
the most widespread African forms! Except for Ichnotropis squamuhsa
the dozen species of lizards were likewise of little interest.

Birdlife was wonderful, being particularly rich in non-passerine
species such as parrots, hornbills, wood hoopoes, cuckoos, and wood-
peckers. In all 61 species of birds and a dozen different kinds of mam-
mals were collected during the 13 days spent here. Of the latter a
molossid bat (Mops angolensis orientis) was described as new.

Mikindani, Southern Province. 10°17' S., 40°7' E. Alt. 20 feet.

Tents were pitched about two miles north of the centre of the town-
ship, which gives its name to Mikindani Bay on the southeast coast of
Tanganvika Territorv, on a little rise to the left of the mainroad to
Lindi after one passes the Government pumping station. Another
noisy pumping station engine has given the name of Mchuchu to
this area.

First landed at Mikindani on the night of March 22 and accom-
plished some collecting on the following evening before leaving for
Kitaya. Returning from the latter place on April 7 in torrential rain,
we just managed to get the tents pitched before darkness fell on the
most cheerless conditions of the whole trip. Left by dhow for Lindi,
the road being under water for miles, on April 24th.

Fully 0 inches of rain fell during our sixteen days stay, on some days
continuing from dawn till dusk. The greatest single precipitation
recorded was 1 3/16th inches on April 10, for this information and
other kindnesses I am indebted to Mr. E. A. Leakey, District Officer.
The average annual rainfall is 36 inches.

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Such conditions naturally retarded outdoor studies to some extent.
Most of our collecting was carried out in the eastern environs of the
township and relatively little in the immediate vicinity of camp.
Though the low ground surrounding the latter was largely under water,
the rainy season was already so well advanced that spawning was over
for the majority of frogs, which were already widely dispersed. Of
the 13 species collected only one (Arthrolcptis xenodactylus) has any
claim to forest associations.

Such primeval forest as may have been here long since disappeared,
but the numerous mango and other trees, to say nothing of baobab and
coconut, result in* providing conditions acceptable to some sylvicoline
species. Of the 31 species of reptiles taken, however, only a cobra
(Naja melanoleuca), is a western forest form, the 7' 6" specimen pro-
viding the most southeasterly record for the species. Topotypes were
collected of the only two lizards (Lygodactylus g. grotci and Amphis-
baena orientalis) described from Mikindani.

Of the 43 species of birds collected, topotypes were shot of Francoli-
nus h. grotci, Lagnosticta r. rcichcnowi = haematocephala, and Uraegin-
thus b. mikindaniensis = niassensis, while of 3 other races described
from here we had obtained specimens from nearby Kitaya and Nch-
ingidi.

Only 7 kinds of mammals were obtained, all typical of the coastal
zone, the bat Triaenops afer being, perhaps, the most interesting.

Mbanja, Southern Province. 9°24' S., 39°45' E. Alt. Sea level to 400
feet.

Camp was made at the edge of Mitonga (Metonge) landing field
('aerodrome'), circa 375 feet.

Arrived at noon, by truck from Lindi, on April 25, and left at 8 a.m.
on May 6, 1939.

Heavy showers occurred during the first week; the second was prac-
tically rainless, the heat tempered to some extent by the southeast
monsoon, until we were struck by a gale of wind and rain on our last
night.

Mbanja (Mbanya), about 10 miles north of Lindi, is a small village
situated on a tidal estuary and almost surrounded by mangrove
swamps. The chief and his people were most friendly and helpful.
While the village itself is on clays and coral rock, the valley at whose
mouth it lies is largely composed of black cotton soil and rich mud ex-
tensively cultivated by the industrious inhabitants. Their principal
products being ground nuts, potatoes, and mahoga, with a few paupau
trees and coconut palms about each hut.

loveridge: African itinerary and comments 207

My camp, on higher ground, was largely surrounded by orchard
forest heavily interspersed with mango trees and waist-high grass
whose barbed black seeds cause considerable discomfort by working
through clothing and even puttees. To the west lay dense scrub in-
habited by very wary squirrels and blue monkey, for the cry of the
latter was heard towards sunset on several occasions. To the east,
i.e. between camp and the coast, were native gardens and the exten-
sive Kikwetu Sisal Estate. All this country is composed of bright red
and very porous soil derived from eroded limestone.

It is in this soil that Amphisbaena ewerbecki, described from here by
Werner in 1910, occurs. Our object in coming to Mbanja was solely
to get a series and in this we were entirely successful. The soil which
furnished a congenial habitat for the amphisbaenid was favoured by
six species of fossorial snakes which we collected in addition to 9 other
kinds of ophidia, a dozen lizard forms, and 36 of birds.

Game was allegedly shot out by the Germans, who maintained a
big camp near here during 1914-1917, but wild pigs were common. At
night a solitary little antelope emerged from the scrub to feed on the
foliage of the ground nuts. Other animals seen or heard, but not col-
lected, were red elephant shrews, galagos, baboons, hares, ratel and
jackal. Man-eating lions had been causing numerous deaths quite
recently in neighbouring villages.

Rondo Forest, Southern Province. 10°8' S., 39°12' E. Alt. 2700 feet.

Camped at Nchingidi, the name given to a clearing at the forest
edge approximately three miles from the nearest scattered native huts.

Arrived just before noon on May 9, and remained until the 21st.

Each evening at varying times from sunset (6 p.m.) till 11 p.m., a
succession of mist clouds blew in from the ocean and up the face of the
escarpment (at whose edge my tent was pitched) to condense on grass
and trees. The latter literally 'rained' upon my tent with every gust
of wind. At daybreak these mist clouds hemmed us in, completely
shutting out the view of the opposite escarpment, invariably persist-
ing till 8 a.m., more usually 9 a.m., occasionally even until noon. In
addition about 4 or 5 p.m. there were sharp showers, while sudden and
unexpected storms of rain swept the plateau at uncertain intervals,
and frequently heavy downpours occurred at night. The nights were
always cool but when the sun did break through by day it was very hot.

Unfortunately an error deprived me of my quinine, and repeated
drenchings when far from camp brought on a fever which, so far as I
was concerned, halted collecting during the last week.

This open forest is situated on the waterless Rondo Plateau, about

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25 miles sothwest of Lake Rutamba, (Lutamba), itself some 25 miles
by road slightly southwest of Lindi. The curious thing about this
forest, of which mvidi is the most important constituent, is the entire
absence of standing water. Even the heaviest downpours immediately
disappeared in the porous sandy soil. To obtain water for domestic
purposes, the local natives (Mwera) made a daily journey of three
hours (there and back) to the foot of the escarpment. It is not alto-
gether surprising that with water so scarce these people should con-
sider washing superfluous. Though very friendly, these Wamwera
only began to busy themselves with bringing in specimens a day or
two before our departure.

Despite the absence of water, 5 species of amphibia, all previously
taken on the Uluguru Mountains nearly 300 miles to the north, were
collected. Of these a tiny toad (Bufo micranotis rondoensis) differed
sufficiently to warrant description.

This isolated plateau had produced 3 species of fossorial reptiles
(Amphisbaena rondoensis, Melanoseps a. rondoensis, Typhlops t. ron-
doensis) which it was necessary to describe as new. Here also we
found Chlorophis maerops, Bitis gabonica, and Brookesia brevicaudata
which occur on the Usambara Mountains 350 miles to the north ; even
more surprising was an undoubted Aparallactus jacksonii of Kiliman-
jaro. The remaining reptiles — 12 kinds of snakes and 10 of lizards
were representative of the coastal plain herpetofauna.

In the past, principally during the Great War, the forest suffered
heavily from. native incursions. Subsequently these refugees were re-
moved but evidence of their destructive occupation were to be seen
in the numerous clearings, some so eroded as to be semiarid areas with
only the scantiest covering of grass, others grass- or bush-grown and
inhabited by such non-sylvicoline, house-dwelling lizards as Hemi-
dactylus mabouia and Mabiiya striata, which I assume to have been
imported by human agency.

Bird life, though not abundant, presumably owing to the absence
of standing water, held promise of being interesting so that it was a
disappointment to go down with fever just as I was turning my atten-
tion to the avifauna. A beautiful roller {Eurystomus glaucurus) of
Madagascar was preserved, while the records of the green -headed
Oriole (0. chlorocephaius) from Mt. Chiradzulu, Nyasaland and the
Usambara Mountains are at last bridged by its occurrence on the
Rondo Plateau. Two rare flycatchers (Batis reichenoioi and Erythro-
cercus I. thomsoni), originally described from Mikindani and the
Rovuma River respectively, were also among the 25 species obtained
at Nchingidi.

loveridge: African itinerary and comments 209

Doubtless the water supply is also the reason for the scarcity of
mammals, of which only 9 species were taken, the most interesting
being Rhynchocyon p. melanurus, originally described from Lindi. I
heard, but never saw, blue monkey, bushbuck, and a small duiker
which the natives called naunde.

Lindi, Southern Province. 10°0' S., 39°14' E. Alt. 50 feet.

In Lindi Hospital (one week) and at the Beach Hotel (one week).

Arrived on May 22 and sailed on June 5th.

The rains, which average 34 inches per annum, were over, and the
clay soil already baked hard by a tropical sun.

Lindi, headquarters for the Province of the same name, is situated
in Lindi Bay between Kilwa and Mikindani. A population of nearly
4000, and the excellent hygenic conditions of the township, rendered
collecting difficult during the week that I daily awaited the arrival of
a steamer. All the same we preserved 15 species of amphibia and
reptiles, 16 of birds, and 5 of mammals.

On our very first hunt, however, we uncovered the oriental blind
snake (Typhlops braminus), the fourth and fifth examples to be taken
in the Territory. T. s. mucruso was the only other snake obtained
though bundles of thatching grass, piles of palm fronds, and heaps of
rubbish, were turned over a wide area. Amphisbaena cwcrbccki was
the only lizard of interest. An intelligent fisherman told me that four
turtles are occasionally encountered in the Bay, he described the luth,
loggerhead, hawksbill, and green turtle. ,

Siga Caves, Tonga Province. 5°6' S., 29°4' E. Alt. c. 150 feet.

Camp was pitched on the trail leading to the caves but about a mile
from the entrance.

Arrived on June 7 and remained until the 17th.

A few heavy rainstorms occurred at infrequent intervals, the rainy
season being over.

The Siga Caves, I took the name from the government signpost
erected at the turn off from the main Tanga to Mombasa road, have
nothing to do with the Sigi River a few miles to the north. They are
sometimes called the Amboni or Mkulumusi Caves for the main en-
trances are less than fifty yards from the crocodile-infested Mkulumusi
River. The numerous caves are waterworn and in past times un-
doubtedly formed an underground channel for the river. There are
dubious stories of the caves extending for a mile, but at the time of our
visit every passage was flooded to within a hundred yards of the
entrance.

The fifty acres of forest which surrounds them has suffered con-

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siderably, in fact it seemed to me that little remains but a scattering
of fine trees surrounded by secondary growth and much scrub. I
might add that the caves are held in superstitious veneration by the
natives, and propitiatory rites, of which I have given a brief account
(1940, Scientific Monthly, 51, pp. 22-35) were held there at the time
of our visit. I imagined that the presence of this "spirit" was the
reason for the absence of native squatters : later I learned that during,
and immediately following, the Great War, natives had moved in but
were turned out again by the Administration on account of their reck-
less destruction of trees.

All 11 species of amphibia, as well as 13 kinds of reptiles obtained at
Siga, were typically coastal plain. One gecko (Cncmaspis a. africanus)
was definitely sylvicoline, while five others proved of assistance in de-
fining a new coastal race named Hcmidactylus t. barbouri.

No serious attempt was made to collect birds in so well-worked a
region, and only 8 species were shot, the choicest being a pair of
migrant Malagasy egrets (Egretta g. dimorpha) and my first bat-eating
hawk (Machaerhamphus a. anderssoni).

The latter was, of course, attracted by the thousands of bats which
made the caves their home. Of these bats, 96, representing 4 species,
were collected ; they ranged in size from the huge Hipposideros g. gigas
to the modest Miniopterus minor. Galagos (G. c. lasiotis) were also
plentiful, their strange cries, mingling with the staccato bark of hyrax
and the hooting or screeching of owls, rendered our nights the noisiest
of the entire safari. The 10 species of mammals, all troglodyte or
arboreal except for a spiny mouse {Acomys w. wilsoni), were of coastal
rather than of forest affinities.

Amboni Estate, Tanga Province. 5°3' S., 39°3' E. Alt. 300 feet.

Our tents were pitched beside a small area of forest, largely second-
ary, which is being carefully preserved by the Estate Management.
As Amboni Estate covers nearly 80 square miles, the precise location
may be somewhat important, we were located about a mile above
Mabokweni Village and surrounded on three sides by sisal planta-
tions.

Arrived on June 17 and remained until the 27th.

Rain occurred in scattered showers on about four cloudy days.

It seemed strange that frogs should be assembling at the conclusion
of the rainy season, yet a visit to a rice swamp close to Mabokweni
Village on the evening of our arrival, resulted in the capture of 154
polypedatid frogs of 9 different species, the females of Hylambates
maculatus certainly were gravid and about to spawn.

loveridge: African itinerary and comments 211

Between this swamp and our camp was an extensive area that had
been under sisal since 1913. It had recently been cleared and tractors
were now engaged in spreading the vegetable debris which had been
piled in long rows and left to rot for two months. By following the
twelve-ton tractors to and fro for three days, two species of ranid
frogs, 5 of lizards, and 11 of snakes were secured. The latter were
largely of fossorial types, among them a burrowing viper (Atractaspis
bibronii) which conformed to the description of katangae of the south-
ern Belgian Congo! Only one frog (Rana o. gribinguiensis) and one
snake (Calamclaps u. warreni) could be cited as largely sylvicoline.

Nor did the forest appear to have any significance as a refuge for
surviving forest forms of reptiles, the 5 species taken within its con-
fines being coastal bush or savanna species. Hornbills, both Bycanisies
and Lophoceros, were much in evidence, but no small birds were shot
though several hours were spent in looking for them.

Jumping shrews (Petrodromus s. sultani) were not uncommon on the
outskirts, and galagos of two species (G. c. lasiotis and G. s. zanzibari-
cas) were present, the latter extremely plentiful. The place was a
refuge also for a band of colobus (C. p. palliatus) and some blue
monkey (Cercopithecus m. monoidcs) though there was no thought of
protecting the latter for whose heads a reward was offered as they
attacked the sisal shoots. Bats, a single red squirrel, and a bushbuck
complete the list of mammals seen in the forest, though other species
were collected in the surrounding plantation.

Magrotto Mountain, Tonga Province. 5°8' S., 38°45' E. Alt. 2500 feet-
Camped on a low hill half-a-mile as the crow flies west of the factory
of MagrOtto Estate.

Arrived on June 27 and remained until July 20th.
The first five days were largely overclouded, raw, damp, and chilly.
Cloud-mist like a dense fog, swept into the valley each evening about
4 p.m. remaining until 9 or 10 the following morning when dispersed
by the frequent rainstorms which swept across the hills. With the
advent of the new moon the vapour was reduced to capping the higher
forested ridges ; the hours of sunshine increased and for an entire week
there were fewer showers. During the last week, however, the weather
remained dull with frequent rainstorms. Average annual rainfall 75
inches !

Magrotto Estate, which formerly occupied about 5000 acres of hill-
tops capping the mountain, is reduced to half that size today. The
more than a million coffee trees of German times have given place
to the West African oil-palm, the only plantation of this palm in all

212 bulletin: museum of comparative zoology

East Africa. My camp among these palms was almost surrounded by
a horseshoe-shaped ridge that was largely forested, a gap towards the
south admitting the clouds of vapour which, during the southwest
monsoon, are an almost regular evening phenomenon. Just below the
western foot of the camp flows a river which rises nearby. To the
north is a swampy bottom, part of whose smothering mat of vegetation
we cleared in our search for frogs.

Prior to 1900, or thereabouts, the whole mountain was heavily
forested until much was cleared for coffee planting. During the Great
War natives moved in on the crown land and destroyed almost every
tree. The hillside directly opposite my camp had been cleared and
rows of Grevillea planted as shade trees for the coffee. When the Estate
was abandoned during the war, native trees, protected by the Grevillea,
sprang up and are now between 60 and 70 feet in height. I mention
this as representing the most amazing come-back of forest which I
have seen anywhere in East Africa. The forest-floor conditions ap-
peared indistinguishable to me from those in adjacent virgin forest.

Despite this, however, animal life, which in species closely resembles
the fauna of Amani at 3000 feet in the Usambara Mountains, and only
20 miles distant across the plain, was decidedly scarce; birds were con-
spicuously absent. In part, of course, this may be seasonal, for the
three weeks spent at Amani in 1926 were during the November rains
when amphibia were breeding, hence their predators more in evidence.
The three weeks spent on Magrotto were immediately following the
coldest months in the year when the temperature falls 57° F. (14° C.)
and a proportion of poikilothermous vertebrates may be assumed to
be quiescent.

Another factor, though only affecting the snake census, might be
found in the composition of the plantation personnel. At Amani the
'hands' were largelv of the Nvamwezi tribe, who are notoriouslv less
fearful of these reptiles. At Magrotto the labour was largely drawn
from local people, who, though helpful, and willing to bring in such
snakes as they came across, displayed no great enthusiasm.

At Magrotto 21 species of amphibia were collected of which only 3
had not been taken at Amani, 1 of these was the forest-edge form
(Rana m. venusta) of the widely distributed savanna species; signifi-
cantly enough both the others were recent invaders from the coastal
plain, i.e. Rana o. oxyrhynchus (instead of the forest-edge R. o. gribin-
guiensis taken at Amani) and Arthroleptis s. stenodactylus (instead of
the sylvicoline A. s. lonnbergi taken at Amani). Three remaining
species taken at Amani though not encountered on Magrotto, were

loveridge: African itinerary and comments 213

Xenopus and Hyperolius forms typical of the coastal plain and which
almost certainly will be found on Magrotto. A good series of topotypes
of H. substriatus (= puncticulatus) were preserved.

When we examine the composition of Magrotto's reptile fauna, of
which 20 kinds were collected, we encounter a similar situation. In-
stead of Typhlops p. gierrai, whose presence one might reasonably have
expected, we encountered only the typical form; as for Neusterophis
olivaceus, the montane uluguruensis occurred in almost equal propor-
tions to the typical savanna race. To sum up, for both snakes and
lizards, precisely 50% were sylvicoline, the rest savanna, reflecting
the fact that, though we spent more time hunting in the forest, reptile
life was more conspicuous in the plantation where immigrant forms
were alike supplanting the forest fauna in the zoological and botanical
realms.

As already stated, forest birds were scarce at the time of our visit,
only 6 or 7 being obtained, of which half were bulbuls. To these might
be added the great eagles (Stephanoaetus coronatus) seen. Of the species
collected 1 (or 2) were described from the nearby Usambara Moun-
tains, 4 from Kilimanjaro, and 1 from Mt. Elgon. A total of approxi-
mately 25% of the species being sylvicoline.

Similarly with the mammals not more than 25% of the 16 species
collected were forest forms in its restricted sense, though the propor-
tion might be raised almost to 50% by the inclusion of certain doubt-
ful creatures like Heliosciurus u. undulatus of Kilimanjaro which, like
the galagoes and monkeys, are arboreal and probably as much at
home in the coastal bush as in virgin rain forest.

Tanga, Tanga Province. 5°40' S., 39°7' E. Alt. 50 feet.

At Tanga Hotel prior to embarkation.

Arrived at noon on July 21 and left on the 23rd.

Fine and hot. Average annual rainfall 59.24 inches.

Spent my last afternoon in Tanganyika searching for Typhlops
platyrhynchus, of which Tanga is type locality, in the sandy area from
which the coconut palms had been removed in order to prepare the
site for a military landing field. All that we got were 3 species of frogs
and 2 of geckos, all typical of the coastal plain life zone.

KENYA COLONY

Likoni, Set/die Province. 4°5' S., 39°39' E. Alt. 50 feet.
On board R. M. S. Dunbar Castle.
Docked alongside Kilindini Wharf July 24 and sailed on the 26th.

214 bulletin: museum of comparative zoology

Showers and sunshine. Rainfall very variable, but the annual
average about 51 inches.

Likoni is a ferry landing on the mainland opposite Kilindini, Mom-
basa Island, the second locality in bold face type appearing on many
labels. It is a region of old coconut plantations and typical coastal
vegetation. I spent two mornings there in search of additional ma-
terial of a new gecko (Hemidactylus t. barbouri) which I had taken
previously at nearby Changamwe, and which I purposed describing.
In addition to this and other lizards we captured the first three
examples of a skink (Riopa pembanum) which I had ever collected, the
species being unknown from the continent until relatively recently.
A frog, two pigmy mice (Leggada b. vicina) of a race described from
Takaungu a few miles to the north, and a good haul of invertebrates
comprised the final spoils of a trip which had lasted nine months.

PLATES

PLATE 1

Loveridge — African Itinerary

PLATE 1

Map showing Principal Collecting Localities

1938

Landing at Mombasa (25.x), except for a stopover at Naivasha and Kinan-
gop (26-31. x), Loveridge proceeded by rail direct to Jinja (1-5. xi). Thence to
Mabira Forest (5-21.xi), Budongo Forest (22.xi-7.xii), Kibale Forest (8-19.xii),
Bundibugyo near Bwamba Forest (19-26. xii), Bugoye, foot of Ruwenzori
Mountains (26-28.xii) and Mubuku Valley at 7000 ft. (29.xii-).

1939

On leaving Mubuku (-9.i) Loveridge descended down the vallev to Mihunga,
circa 6000 ft. (9-21.i), then back to Bugoye (21-24.i), Nyakabande (25-30.i),
Mushongero (30.i-4.xi), returned to Nyakabande (4-8.U); Kisenyi (8-13. ii),
Goma (13-14.ii), Mamvu on Idjwi Island (14-16. ii), Upper Mulinga on Idjwi
(16.ii-6.iii), Uvira (7-8.iii), TJjiji (9-16. iii), Dar es Salaam (18-19.iii), Mikin-
dani (22-24.iii), Mbanja (25.iv-6.v), Lake Rutamba (6-8.v), Nchingidi
(9-21.v), Lindi (22.v-4.vi), Siga Caves (7-17.vi), Amboni Estate (17-27.vi),
Magrotto Mountain (27.vi-21.vii), Tanga (21-23.vii), Kilindini (24-26.vii).

BULL. MUS. COMP. ZOOL.

Loveridge. African Itinerary and Comments. Plate 1

UGANDA.-

'■•'BuJorrao

h> Forest i ^t
rJ*/CibaU F°rzZ~Apt&

Muill/iu Valley Bl Sf JMJ\

?' -Cb* Mabira '■

*■*«' MutaXijS—-^
Mil jCpn>V>°»S"°
Nyahabaniei -f(abale

Lake KlvJikk.%-

Costermansn-

Uvira

BELGIAN
CONGO

i

K EN YA
COLONY

TAN G A
TE R R

«0C

"aS"~vtto /t,f''WW«tJo MOMBASA
Ambon! EstaleY^f^t

Stga Cai/es'T Q

-v ( 9

x/

*

A

*

V

ZIBAR
ES SALAAM

'Mindani

Scale in Miles

0

u.

100

_l_l

200

1

PLATE 2

Loveridge— African Itinerary

PLATE 2

Fig. 1, Transport by water — Canoes on Lake Mutanda

These dugouts, waiting to take us from Mushongero to the south end of the
Lake, speak volumes for the patient toilers who, with adze or other simple
tool, gouged out the hard timber from the fallen tree. Moreover, as no trees
of sufficient height to provide sixteen or eighteen-foot canoes are to be found
within many miles of the Lake, these incredibly heavy craft had to be pushed
uphill and down dale on rollers by human muscle alone.

Fig. 2. Transport by Land — Our Lorry at Mubango

African travel today involves increasing use of modern methods of transport,
resort to head-porterage being reserved for roadless regions or where the ascent
of a mountain by native track is called for. In modernizing Africa motor-
driven vehicles will soon be claiming precedence over predators and snakes
as the major menace to life and limb. The photograph depicts our native-driven
Mercedes-Bentz, which at the time was carrying my six boys atop a load con-
sisting of a ton of camp and collecting equipment, irresolutely resting on
marshy ground where faulty driving had landed her at the very start of our
'safari!'

BULL. MUS COMP. ZOOL.

Loveridge. African Itinerary and Comments. Plate 2

PLATE 3

Loveridge — African Itinerary

PLATE 3

Fig. 1. Prospecting For a Camp Site — Forest-edge, Budongo

Budongo Forest, which covers about 180 square miles, gathers about its
fringes a dense and often impenetrable undergrowth of bush. Within, however,
where the unbroken forest canopy, two-hundred feet overhead, prevents
shrubs springing up among the mighty buttress roots, one can wander at will
over the thick layer of leaves which carpet the forest floor.

Fig. 2. Tractor and Tramway are Employed for Logging at Bisu

The absence of paths and water drove us unwillingly to camp at Bisu, in
close proximity to the Buchanan Saw Mills. There access to the forest was
assured by a track cut for the tramway depicted above. Unfortunately the
noisy tractor and its train, which made the three-mile run through the forest
every twelve hours, had driven elephant, buffalo, and other large mammals to
seek quiet elsewhere.

BULL. MUS. COMP. ZOOL.

Loveridge. African Itinerary and Comments. Plate 3

I

MHMETttBl^BUHB

PLATE 4

Loveridoe — African Itinerary

PLATE 4

Fig. 1. Native Path through Mabira Forest

Paths were few and far between, yet the undergrowth was so impenetrable
that they provided the only means for reaching the deeper forest to sample its
denizens. While monkeys and squirrels fell victims to this inquisitiveness,
parrots screeched or clambered about in the tree-tops, where, with a host of
smaller birds feeding in the forest canopy, they enjoyed complete immunity,
being out of range of gunshot.

Fig. 2. A Semi-pigmy of Baamba or Batwa Stock

The Baamba clans inhabiting the Ituri Forest region northwest of the
Ruwenzori Mountains, closely resemble their Batwa kinsmen of the Congo
forests. Attired only in a civet skin, the sturdy subject of the photograph
presents a marked contrast to the tall, cotton-clad Muganda on the forest
path in Mabira.

Fig. 3. Family Group at Bundibugyo in Toro

Yet another popular style in dress is exhibited by this Bantu family relaxing
after their early morning labours in the hot region just above the Semliki
Valley. Hundreds of natives representing many tribes, visited our camp at
Bundibugyo. If I am not mistaken those shown are Batoro, the agricultural
middle-class members of this society, superior to the hunting Baamba from
whom they obtain meat in exchange for cereals, yet themselves formerly sub-
ject to their pastoral overlords, the Hamitic Bahima.

BULL. MUS. COMP. ZOOL

Loveridge. African Itinerary and Comments. Plate 4

r-£.j

#

■&

■

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 6

NOTES ON LYCAENID BUTTERFLIES

By Harry K. Clench

MUS. COMP. ZOOL
LIBRARY

SEP 2 2 1964

HARVARD
UNIVERSITY.

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

July, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1, 2,
3, 4 and 5 have appeared and of the Memoirs, Vol. I to LVI.

These publications are issued in numbers at irregular intervals.
Each number of the Bulletin and of the Memoirs is sold separately.
A price list of the publications of the Museum will be sent upon
application to the Director of the Museum of Comparative
Zoology, Cambridge, Massachusetts.

After 1941 no more Memoirs are to be published.

SEP 2 2 1964

HARVARD
Bulletin of the Museum of Comparative Zoology rsiTY

AT HARVARD COLLEGE

Vol. XCIV, No. G

NOTES ON LYCAENID BUTTERFLIES

Bv Harry K. Clench

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

July, 1944

JUL |9 1944

L I B K A K *

No. G — Notes on Lycaenid Butterflies
By Harry K. Clench

a. THE GENUS CALLOPHRYS IN NORTH AMERICA

While working on a revision of the genus Indsalia Scudder sueh
frequent recourse was had to the various American species of Callo-
phrys that it was thought advisable to prepare a brief review of them.
Unfortunately, time was all too short to do the job properly, and
genitalic examinations were not made. These notes, however, were
gotten together, in the hope that they would facilitate further work
on the genus. Such future work should certainly include a genitalic
study of all forms involved.

The first and only real revision of the American Callophrys was that
of Barnes and Benjamin' , who placed the genus on very solid footing,
especially as compared with its previous state. It was through this
paper that our first accurate knowledge of the limits and variabilities
of the several species was obtained. There have remained, however,
several points that need clarification. Probably the most important
of these is the interrelationship of the various Californian forms. The
arrangement adopted by Barnes and Benjamin (dumetorum in the
north, with subsp. perple.va in the south) is too simple, and although
it at first appears logical, subsequent research has proved it false.
There are at least three forms, and very likely several more, in Cali-
fornia that have been going under the names dumetorum and perplexa.
What has been commonly passing as true dumetorum is something
else, while true dumetorum itself appears to be rather poorly known.
These will be discussed in greater detail under the species concerned.
Unfortunately, the problem in California has been only partly solved,
the material at hand indicating much further work to be done, but of
itself insufficient to do it.

In addition to this, there has been a new species (comstocJci) pro-
posed since the work of Barnes and Benjamin, and in the present
paper a second novelty, a new subspecies of affinis, is added.

Finally, there are a number of new and interesting records, exten-
sions of ranges, particularly of affinis and sheridanii.

The genus in North America appears to be confined exclusively to
the Rocky Mountains and the area westward to the Pacific. It ex-
tends from northern Mexico in the south to Alberta and British

U923, Contrib. Nat. Hist. Lep. North America 5, p. 61-67.

218 bulletin: museum of comparative zoology

Columbia1 in the north. The species appear to be generally mountain-
loving, although they are frequently (perplexa, for example) taken at
low elevations.

The paucity of species in Asia and Europe and their close inter-
similarity, as compared with the relatively large (6) number of species
and their diversity here in America, would suggest that the Palae-
arctic species were derived from American stock. The origin of this
American stock, however, is quite another question, and one foolish
to speculate upon, in view of our present ignorance of much of the
pertinent details.

The author wishes to thank the several individuals and institutions
who have greatly assisted in this review. The American Museum of
Natural History (AMNH) very kindly loaned some material for
study. The Carnegie Museum and the U. S. National Museum
checked the types of species in their collections. Mr. Robert G. Wind,
of Berkeley, California, was very kind in sending me for study a large
amount of exceedingly interesting material, including a number of the
complex viridis-\\ke specimens from the Californian Sierras, and other
northern Californian localities. Mr. Charles L. Remington also lent
some very interesting material which was of the utmost help in pre-
paring the paper. Mr. L. P. Grey gave the author all of his Callophrys,
which included several very interesting things. The collection of the
Museum of Comparative Zoology (M.C.Z.) has, as in times past,
always been available to me through the kindness of Mr. Banks. This
collection contains much material of interest and assistance. My own
collection has been drawn on wherever possible, being designated
(H.K.C.).

Genus Callophrys Billberg

1820, Enumeratio Insectorum, p. 80. Genotype, Papilio rubi Linn. (Palae-
arctic).

Key to species

1. Fore wing underside with green covering wing down to C112 or 2A 2

- This green restricted to a more or less broad costal and outer marginal
border, leaving a large internal area of gray or fulvous 6

'The two papers on the faunae of these provinees (K. Bowman, 1919, Annotated Cheek List
of the Macrolepidoptera of Alberta, Alberta Nat. Hist. Soc., Red Deer, Alta., 16 pp.; E. H.
Blackmore, 1927, Check-list of the Macrolepidoptera of British Columbia, C. F. Banfield,
Victoria, B. C 47 pp.) each list Callophrys dumelorum. Just, what they meant is not known.
Indeed, it coidd have been almost any species in the genus, but most logically affinis or shcri-
danii. Beferences to these papers, as well as to a number of others, were omitted due to this
and similar lack of precise information, so common in this genus.

clench: notes on lycaenid butterflies 219

2. Hind wing below immaculate, or at most with a faint indication of a white

line — usually a discal row of obscure white dashes 3

- This wing with either a prominent white line, or with a series of distinct,

pure white dashes or dots 4

3. Fulvous above in both sexes widespread, dominant; underside apple-

green a. ajfmis

- Fulvous above reduced, frequently (usually?) absent in the male; green

below purer, inclining even to bluishness affinis washingtonia

4. Underside of hind wing with a solid line of white 5

- This surface with a row of prominent white spots viridis

5. Line on underside of hind wing heavy, inwardly edged with a pronounced

band of black s. sheridanii

- This line thin; the black faint or absent sheridanii neoperplexa

6. Underside of hind wing nearly immaculate, or with a row of white spots

of varying number 7

- This area with a crooked white line 8

7. Outer margin of fore wing evenly convex apama homoperplexa

- This margin convexly angled at M2 or thereabouts, and straight, or

slightly concave thence to the inner angle 9

8. Line on underside of hind wing basally edged with black, then fulvous

a. apama

- This line with no fulvous comstocki

9. Costa of fore wing below edged with fulvous dumetorum perplexa

- Costa of this surface not edged with fulvous (or at most only very slightly)

d. dumetorum

Callophrys dumetorum Boisduval

This species ranges from northern Mexico (Baja California) north
beyond San Francisco. The northern limits of its range are still un-
defined. There are apparently two subspecies of this, one very widely
known (perplexa), inhabiting the lower part of its range, and the other
(typical dumetorum) much less perfectly known, inhabiting the upper.
It is possible that the latter as considered herein consists in reality of
several subspecies.

The species shows several characters that place it close to apama.
The principal one of these is the large fulvous or gray area on the fore
wing below. The chief constant difference between the two (apama
homoperplexa and dumetorum perplexa approach each other so closely
that aside from this basic character they are almost inseparable) lies
in the shape of the outer margin. In dumetorum it is convexly angled
in the vicinity of M2, and straight or slightly concave below it to the
inner margin, while in apama the whole margin is evenly convex.

220 bulletin: museum of comparative zoology

Callophrys dumetorum dumetorum Boisduval

Thecla dumetorum Boisduval, 1852, Ann. Soc. Ent. France (2) 10, p. 291;
Oberthur, 1913, Et. Lep. Comp. 9, p. 40, pi. 236, fig. 1926; Draudt, 1919,
in Seitz, Macrolep. World, 6, p. 763, pi. 154b; id., 1924, p. 1043. (all
partim)

Thecla dumetorum: auct. (partim)

Callophrys dumetorum: auct. (partim)

The type, in the United States National Museum, came from Cali-
fornia. No definite locality was designated. Due to a lack of material
agreeing satisfactorily with the description and with Oberthur's
figure of the type (loc. cit.) no type locality has been fixed.

Localities. California: Phelan; Snow Creek; Dana Point (all
H.K.C.); Pasadena (M.C.Z.); Calistoga; Petrified Forest (both in the
collection of R. G. Wind). These are all doubtfully typical dumetorum.

Male above uniform brownish gray. Fringe white, basally gray.
Female similar but more brownish, and with a discal fulvous suffusion.

Male below with fore wing gray or fulvous, inner margin somewhat
paler. Costa and outer margin as far down as Cui green. Hind wing
uniform green with two or three white spots, the principal ones being
on the costa and in the CU1-CU2 interspace. Fringe gray, slightly
paler outwardly and at the vein-ends on the hind wing. Female
similar, but with the gray area on the fore wing more fulvous. Both
sexes frequently have a more or less strongly developed discal line on
the fore wing, usually merely an intensification of the gray or fulvous.

Length of fore wing. Male, 12.5-13 mm.; female, 13 mm.

The above rather brief description is based on three males and a
female (two males from Phelan, one from Snow Creek, and a female
from Dana Point) which may or may not be typical dumetorum.
Several points in Boisduval's description differ from those found in
these specimens (namely, the fulvous (of Boisduval) instead of gray
color of the large area on the fore wing below; the presence of a number
of white spots on the hind wing below, lacking in the above-described
specimens). Typical dumetorum was very likely described from much
further north than these four specimens, although just where is still
unknown. The Calistoga specimens, while apparently approaching
Boisduval's description a little more closely than these, still differ
enough to be closer to the more southern specimens.

These specimens, of what is here taken to be typical dumetorum,
differ from perplexa in the absence of fulvous edging on the costa of the
fore wing below, and (in the male) by the gray, instead of fulvous,
area on this surface. There are, however, hints of fulvous on this area,

clench: notes on lycaenid butterflies 221

but this may possibly be explained by the fact that the specimens
came from a region not too remote from the home of true perplexa.

Two of the three ( lalistoga males examined have a definite fulvous
shading on the hind wing above, and one of them a patch on the fore
wing as well. More specimens are needed to tell whether or not this is
a local race of dumetorum. This character is quite unusual for dume-
torum and appears not to have been noticed previously.

Callophrys dcmetorum perplexa Barnes and Benjamin

Thecla affinis: auct. (partim)

Thecla dumetorum: auct. (partim)

Callophrys dumetorum: McDunnough, 1914, Ent. Rec. and Journ. Variation,

26, p. 196. (partim)
Callophrys dumetorum race perplexa Barnes and Benjamin, 1923, Contrib.

Nat. Hist. Lep. North America, 5, p. 65; Draudt, 1924, in Seitz, Macrolep.

World, 5, p. 1043. Comstock, 1927, Butterflies of California, p. 168,

pi. 50, figs. 17, 18, 19 (partim); Hoffman, 1940, Anales Inst. Biol. Mex., 11,

p. 708 (no. 623).

Holotype, allotype and a number of paratypes in the United States
National Museum. Described from San Diego, California.

Other localities. California : Pine Valley ; Corona ; Balboa ; El Modena
(all in coll. H.K.C.). Mexico: (northern localities, adjacent to Cali-
fornia).

Differs from typical dumetorum (as considered in this paper) in the
fulvous edging on the costa of the fore wing below, and in the in-
creased fulvous in the discal region of this same surface.

Apparently restricted to southern California and adjacent Mexico.

Length of fore wing. Male, 11-13.5 mm.; female, 11.5 mm.

Callophrys comstocki Henne
Callophrys comstocki Henne, 1940, Bull. So. Cal. Acad. Sci., 39, p. 71.

Holotype and allotype from the Providence Mts., San Bernardino
County, California, in the collection of the Los Angeles County
Museum. Paratypes from the same locality in the United States
National Museum, Canadian National Collection. Also (?) in the
collections of C. H. Ingham and C. Henne.

Topotypes have been examined from the collections of A. C.
Frederick, D. B. Stallings and the author.

Above, both sexes slate gray. Male with a small oval scent-pad.
Fringe gray, paler outwardly. Hind wing but slightly scalloped.

222 bulletin: museum of comparative zoology

Below, both sexes flat grayish green. Fore wing with inner marginal
area, centrally as far costad as M3, gray. This wing crossed by an
obsolescent post-discal line of white, basally gray-black, dashes.
Hind wing with a similar, but more distinct, line, edged basally with
black. This line outwardly displaced in M3-C111, and irregular thence
to the inner margin. Fringe of both wings basally gray, outwardly
white.

Length of fore wing. Male, 10-11 mm.

Closest to apama, but differing in several respects, principally the
gray upper surface of both sexes, and the flat green below, unrelieved
by any fulvous shading. It is a very interesting and isolated species,
possibly a local modification of apama, but certainly distinct enough
to warrant full specific distinction. So far as known it is restricted to
the Providence Mts.

Callophrys apama Edwards

This species occurs over a relatively compact range from Arizona
to New Mexico and north to Colorado, becoming differentiated in the
latter regions into a distinct subspecies. The limits of range of this
species are fairly well-known. It may, however, be turned up in
eastern California (doubtful) or southern Utah. (The Carnegie
Museum has a single male from this state; I have not seen it.)

From its near relative, C. dianetorum, apama may be distinguished
by the character given above under the general remarks for dume-
torum.

Another close ally (perhaps its closest) is the recently described
C. comstocki. This latter may be separated at once by its complete
lack of fulvous shading on the underside, by its smaller size, and by
the more distorted, less prominent white discal line on the hind wing
below.

Callophrys apama apama Edwards

Thecla apama Edwards, 1882, Papilio, 2, p. 137; Draudt, 1919, in Seitz, Macro-

lep. World, 5, p. 763; id., 1924, p. 1043.
Callophrys apama: Barnes and Benjamin, 1923, Contrib. Nat. Hist. Lep.

North America, 5, p. 67.

Type locality. Fort Grant in the Graham Mts., Arizona. One male
and two females labelled "Arizona" in the Edwards collection at the
Carnegie Museum. One of the latter made a lectotype by Holland.

Other localities. Arizona: Oak Creek Canyon (6000 ft.) (M.C.Z.);
White Mts.; Catalina Mts. (both in coll. H.K.C.); Navajo Mt. (in

clench: notes on lycaenid butterflies 223

coll. R, G. Wind); Sierra Aneha Mts. (in coll. P. S. and C. L. Reming-
ton).

Above in the male dark gray with a brownish tinge. A small,
almost circular scent-pad at the upper cell-end. Fringe of fore wing
concolorous with ground (or slightly darker), paler outwardly; of
hind wing similar, but frequently almost pure white outwardly; tips of
veins Cuu Cu2j and 2A usually white.

Female similar, but with fulvous patch on fore wing and usually
on hind wing. Fringe outwardly slightly paler.

Below, fore wing largely fulvous. Inner margin gray. A post-
discal, white irregular line crosses from costa to Cu2, composed of
white dashes, basally edged with black. Base of wing and apical area
outside of post-discal line (and extending down as far as Cux) green.
Hind wing green. A post-discal line, similar in construction to that of
the fore wing, but here more tortuous, crosses from costa to inner
margin, outwardly displaced often in the form of a crude "\Y" in the
M3-Q12 region. Basal to this line runs a closely adherent band of
fulvous. Occasionally there is a definite suggestion of a Cui-Cu2 sub-
marginal spot. Fringe of fore wing basally dull fulvous, outwardly
gray; of hind wing similar, but paler outwardly and tipped at Ciii, Cu2
and 2 A with white.

Length of fore wing. Male, 10.5-13 mm.; female, 10.5-11 mm.

The typical subspecies is distinguished from its Colorado repre-
sentative by the clarity and completeness of the markings below,
which are reduced frequently to the point of complete obscurity in
homoperplexa.

Callophrys apama homoperplexa Barnes and Benjamin

Thccla dumetorum auct. (partim)

Callophrys apama race homoperplexa Barnes and Benjamin, 1923, Contrib.

Nat. Hist. Lep. North America, 5, p. 68; Draudt, 1924, in Seitz, Macro-

lep. World, 5, p. 1043.

Holotype, allotype, and several paratypes in the United States
National Museum. Holotype and allotype from Denver, Colorado.
Paratypes from Golden, Boulder, and Denver, Colorado.

Other localities. Colorado: Durango; Husted and Starr Ranch
(El Paso County) (all M.C.Z.); Boulder Canyon; Eldora (both in
coll. P. S. and C. L. Remington). New Mexico: nr. Hot Springs, Las
Vegas (7000 ft.); Rincon (both in M.C.Z.). Specimens from Golden
have also been examined.

224 bulletin: museum of comparative zoology

On the upperside it differs apparently only in the more fulvous
shading of the female, and a greater tint of fulvous in the male.

On the underside the differences are much more pronounced.
Homopcrple.va lacks, in varying stages of completeness, the white
discal lines on each wing which form such a conspicuous part of typical
apama. The green on the fore wing appears to be more greatly ex-
tended.

Length of fore wing. As in the typical subspecies.

This subspecies is very interesting, and would seem to point to-
wards a connection with affinis. This at first implausible suggestion
becomes more likely when one notices that males of this subspecies are
frequently dull fulvous, and that occasionally specimens are found in
which the green of the underside of the fore wing covers an abnormally
large part of the wing, or shows itself faintly as a discal green suffusion.
Another point in favor of this alliance is the tendency of affinis to
produce specimens with hints (sometimes startlingly complete) of a
discal white line on the hind wing below. Yet another point is that the
ranges of the two do not apparently overlap, or if so, they only occur
sympatrically in a very restricted area. More material is needed from
the zone of transition (northern Colorado and adjacent regions) to
prove or disprove this.

Prior to the work of Barnes and Benjamin, this subspecies was com-
pletely confused with dumetorum perple.va (dumetorum auct.) of
southern California, and very understandably so. The two are strik-
ingly similar, and frequently almost the only real point of distinction
(superficial) is the difference in the fore wing outline. In general, how-
ever, males of homoperplexa have a more ruddy upper surface than do
males of perple.va. The green below of homoperplexa tends to be some-
what more brassy than that of perple.va.

Extreme homoperplexa lacks any suggestion of the discal line on
either wing below. Throughout Colorado, however, and particularly
in the southern part of the state specimens are frequently found that
tend quite definitely toward the typical subspecies. The two New
Mexico examples (localities given above) are in this category as well.
Apparently apama is restricted to Arizona.

Callophrys affinis Edwards

The exact range of this species is not yet known. It extends, from
the material and information at present available, from Utah north to
Wyoming and Washington. It may possibly occur in Colorado also.

clench: notes on lycaenid butterflies 225

The Washington specimens appear sufficiently distinct to warrant
separate racial consideration. Wyoming specimens appear to be
transitional in some respects, but are referred at present to the typical
subspecies.

This species is distinguished by its large, usually entirely unmarked
green under surface, and (with the exception of the Washington race)
large amount of fulvous above in the male. Both dumetorum (Calis-
toga) and apama homoperplexa have this suffusion in the males, but
not to the extent of typical affinis.

Callophrys affinis affinis Edwards

Theda affinis Edwards, 1862, Proc. Acad. Nat. Sri. Philadelphia, p. 223;

Draudt, 1919, in Seitz, Macrolep. World, 5, p. 763; id., 1924, p. 1043.
Callophrys affinis: Barnes and Benjamin, 1923, Contrib. Nat. Hist. Lep. North

America, 5, p. 66.

The types, one male and one female, from Utah, are in the Carnegie
Museum. Silver Lake, Utah, is here selected as the type locality,
based upon two males and two females in the M.C.Z. Barnes and
Benjamin (loc. cit.) referred to some specimens from Silver Lake as
topotypes, but did not elaborate.

Other localities. Wyoming: Teton Mts. (in coll. R. G. Wind).
Edwards' description of this species is very good, and is quoted here:
"Both sexes glossy red brown, brightest in female; the male has a
smooth oval spot on disc of primaries; costa of primaries and base of
both wings blackish brown; whole hind margin edged with same
color; fringe white.

"Under side uniform apple green, except on inner margin of pri-
maries, where it is pale, brownish grey; both wings immaculate; costal
edge of primaries grey; hind margin of secondaries without crena-
tions."

Length of fore wing: Male, 12-13 mm.; female, 12.5-13.5 mm.

The typical form is large and bright. It differs from washingtonia
in the shade of apple green below, and in the large extent of fulvous
on the male above.

t

Callophrys affinis washingtonia, new subspecies

Upperside:

Male. Both wings dark gray, slightly brownish, and occasionally
suffused with dull fulvous in the disk. Fringe white, basally dark
gray.

226 bulletin: museum of comparative zoology

Female. Similar to the male, but with the fulvous more extended.
Underside:

Male. Both wings green, slightly bluish, and very faintly brassy.
Inner margin of fore wing to 2A, and frequently to C112, gray. Hind
wing occasionally with the merest suggestion of a spot in the CU1-CU2
interspace.1 Fringe white, slightly darker at the vein-ends on the
hind wing.

Female. Similar to the male, but with the addition of a very thin,
extreme marginal line of white, running from anal angle to Mi. This
is very likely an individual variant, and will probably not be found to
hold true for other females. At the anal angle is a very small white
patch, the origin of the white line, and probably also merely an
individual variation.

Length of fore wing. Male, 11-12.5 mm.; female, 12 mm.

Holotype. Male, Alta Lake, Washington, April 25, 1935 (J. C.
Hopfinger) ex coll. P. S. and C. L. Remington.

Allotype. Female, same data as holotype.

Paratypes. Two males, Brewster, Washington, May 9, 1939 (J. C.
Hopfinger?), ex coll. L. P. Grey.

Holotype and allotype, no. 26259 in the M.C.Z. Paratypes in the
author's collection.

Remarks. Differs from typical affinis in the much more reduced
fulvous above in both sexes, and in the more bluish tone of the green
below.

A series of over 15 examples from the Teton Mts., Wyoming, seems
about intermediate between washingtonia and true affinis. There is,
however, little if any variation in the amount of fulvous on the fore
wing above in the male, while this character in washingtonia varies
from a limited amount (smaller than the smallest .of the Teton Mts.
specimens) to none at all. In several of the Wyoming specimens
there is an indication of a tendency to produce a row of white dashes
on the hind wing below; in one specimen quite strongly.

Callophrys viridis Edwards

Thecla viridis Edwards, 1862, Proc. Acad. Nat. Sci. Philadelphia, p. 223.

Thecla dumetorum: auct. {partim)

Callophrys dumetorum: Barnes and Benjamin, 1923, Contrib. Nat. Hist. Lep.
North America, 5, p. 64 {partim); Comstock, 1927, Butterflies of Cali-
fornia, p. 168, pi. 50, figs. 16, 20, 21, 25 (some of these are apparently
dumetorum) {partim).

1 The allotype has an indication of dashes in other neighboring interspaces, as well as a row
of very faint dashes on the fore wing. This latter can be seen faintly in certain lights in one
of the paratypes.

clench: notes on lycaenid butterflies 227

The type locality is here selected as San Francisco, California,
based on a female in the collection of P. S. and C. L. Remington
(May 8, 1934, Wm. Hovanitz collector) that agrees excellently with
the original description. In the absence of any type at the Carnegie
Museum this specimen is here made the neoholotype.

Other localities. "California" (M.C.Z.).

The male is above uniform gray. Fringe white, gray basally and at
the vein-ends of the hind wing. Female similar, but may be largely
suffused with fulvous, leaving a dark base, costa and outer margin on
the fore wing, and a dark base only on the hind wing.

Below in both sexes both wings are uniform pea-green. Fore wing
with inner marginal area to vein Cu2 gray, save on outer margin,
where the green extends down to 2A. Costa edged with fulvous. In
the disk is a faint row of dull white dashes, three in number in the
specimen examined, in interspaces M2-M3-Cu1-Cu2. On the hind wing
this row is continued, commencing at the outer angle and proceeding
to just basad of the anal lobe on the inner margin. The marks com-
posing this line are rather displaced and irregular in size, the one in
Cux-Cu2 being the largest. Those on the fore wing and one on the costa
of the hind wing are basally edged with dark brown or black. Fringe
of both wings white, rather dull, and basally slightly darkened.

Length of fore wing. Female, 13.5 mm. (neoholotype).

This species, heretofore considered synonymous with dumetorum,
is perfectly valid. Dumetorum, as described by Boisduval,1 has a
large fulvous area on the underside of the fore wing, lacking in viridis.
This character (the extent of the green on the underside of the fore
wing), which so far as known is very constant and subject to almost
no individual variation,2 is alone enough to raise Edwards' name from
the synonymy. Oberthur's figure of the type of dumetorum. (loc. cit.)
confirms Boisduval's description in this respect. True dumetorum has
been discussed further above.

Even with viridis and dumetorum separated, the picture is still any-
thing but clear. In California, apparently right along with viridis,
occurs a very green form with an immaculate underside. With only a
limited number of specimens available, it has been impossible to de-
termine whether this is a distinct species or merely an extreme of
viridis. Either is quite possible, although the latter is the more
probable.

1". . . et le disque des ailes superieures est beaucoup plus largement roussatre, ce qui fait
que le vert domine nioius." (reference under dumetorum).

2There is a certain amount of individual variation in this character in C. apama homopler-
plera, as has already been noted, but this is apparently an indication of transition towards
another species (affinis), and as such is excusable.

228 bulletin: museum of comparative zoology

Several specimens of Callophrys have been examined from the
collection of Mr. R. G. Wind, having been collected by him chiefly in
the Sierras. These, while apparently close to viridis differ in several
respects from it. These specimens, however, also differ considerably
inter se, so that in the absence of genitalic examination or larger series,
nothing further can be said.

Callophrys sheridanii Edwards

This species occupies a long, narrow, mountainous strip from
Cloudcroft, New Mexico north as far as Brewster, Washington. It is
differentiated into at least two subspecies, the typical occurring in
Wyoming, Colorado and probably New Mexico, while the subspecies
neoperplcxa ranges from Utah north to Washington. Specimens from
the latter locality do not seem exactly typical of neoperplcxa and
when more specimens are available they may be found to belong to a
distinct race.

The chief character whereby this species may be differentiated from
any other now known is the long, nearly straight (usually) white line
on the hind wing below, frequently basally bordered with black. It is
also one of the smaller species of the genus.

Callophrys sheridanii sheridanii Edwards

Thecla sheridanii Edwards, 1877, in Carpenter, Field and Forest, 3, p. 48

(lapsus calami).
Thecla sheridanii: Draudt, 1919 in Seitz, Macrolep. World, 5, p. 763; id., 1924,

p. 1043.
Callophrys sheridanii: Barnes and Benjamin, 1923, Contrib. Nat. Hist. Lep.

North America, 5, p. 66.

In the Carnegie Museum is one female labelled "Bighorn, Mont."
(fide Sweadner, in lift.) here selected as neoholotype. Holland's selec-
tion of a lectotype from Denver, Colorado is invalid as that is definitely
not the type locality.

Other localities. Wyoming: Teton Mts. and vie. (A.M.N.H.).
Colorado: Chimney Gulch (M.C.Z.); Ft, Collins; Denver (both
Barnes and Benjamin, p. 66); Eldora; Ceal Creek (both in coll. P. S.
and C. L. Remington). New Mexico: Cloudcroft (M.C.Z.).

Male above dark gray with a slight brownish tinge. A small oval
scent-pad at the upper cell-end. Fringes of both wings white, gray
basally. Female similar, but lacking the scent-pad.

Male below with both wings slightly brassy green, irrorated faintly,

clench: notes on lycaenid butterflies 229

especially on the hind wing, with obscure black scales. Fore wing with
inner margin gray-brown. A post-discal line crosses from costa to CU2
(occasionally with an inwardly dislocated extension in ( u2-2A), white,
and basally narrowly lined with black-brown. Hind wing with a
similar line, but heavier) and running from costa to inner margin in a
nearly straight line (frequently dislocated, but seldom if ever curved).
A small, white cell-end spot is occasionally present. Fringe as on
upper surface, but slightly greenish towards outer angle. Female as
in the male.

Length of fore wing. Male, 10-12 mm.; female, 11 mm.

The typical form differs from neoperplexa in the thicker white line
below, and the more prominent basal black edging to this line. Barnes
and Benjamin state that a specimen from Cloudcroft, New Mexico is
closer to the subspecies neoperplexa. This is decidedly queer, if so,
and is not borne out by the single specimen from that locality in the
M.C.Z., which, although not perfectly typical, is close enough to
belong here for the present.

Callophrys sheridanii neoperplexa Barnes and Benjamin

Callophrys sheridanii race neoperplexa Barnes and Benjamin, 1923, Contrib.
Nat. Hist. Lep. North America, 5, p. 67; Draudt, 1924, in Seitz, Macrolep.
World, 5, p. 1043.

Holotype and allotype from Eureka, Utah. Paratypes from Stock-
ton and Silver Lake, Utah. Holotype, allotype and 2 male, 1 female
paratypes in the United States National Museum.

Other localities. Montana : Polaris (H.K.C.). Washington : Brewster
(H.K.C.). In the M.C.Z. is a series from Silver Lake.

Its differences as compared with typical sheridanii have been
pointed out above. The Brewster, Washington, specimens do not
agree perfectly with Utah neoperplexa, and may ultimately be found
racially separable. They are very close, however, and for the present
will be left under that name. It is possible that some of the Sierran
material from California, mentioned under viridis, will prove to be
southern extensions of sheridanii stock, racially modified.

Length of fore wing. Male, 10-11 mm.; female, 11-11.5 mm.

b. THE ACASTE GROUP OF THE GENUS THECLA

The species treated here form a more-or-less closely interrelated
group isolated by Draudt (1919, in Seitz, Macrolep. World, 5, p. 762)
as the tailless section of his amyntor-group, of the genus Thccla. No

230 bulletin: museum of comparative zoology

better classification can be made with accuracy at present, since the
generic subdivision of the neotropical Theclinae is a task no one has
yet attempted with any degree of completeness1.

That the "tailless section" is here considered separately from the
"tailed section" is due principally to a very great dearth of material
of the latter. The two sections are apparently very closely allied, at
least in appearance2, and are separable only in minor characters.

The acaste group3 may be distinguished from the "tailed section"
in the following external particulars : The anal angle of the hind wing is
more produced, and the anal lobe is longer, more prominent. In
general the tail is absent (see footnote no. 3) but if it is present it is
coarser (broader) and proportionately shorter than those of the other
section. Also on the hind wing, the vein-ends are here slightly more
tufted. The green below is less uniform and usually less shining. The
general appearance of this group is more reminiscent of CaUophrys, or
even, somehow, of Incisalia, while the others seem more typically
Thecline.

The various members of the acaste group are of average Thecline
size, ranging (in the length of the fore wing) from 12 (female of remits)
to 16 mm. .(female of a. acaste). The males all have scent pads4, but
they are usually small and almost unnoticeable, occasionally being of
the same color as the ground color, a rather unusual occurrence. The
males are above (excepting marialis) some shade of blue or purple,
very metallic, with dark borders of width varying with the species.
The females on this surface are duller, the blue being considerably
less metallic, and more basally restricted. Below the sexes are similar,
with a rather similar basic pattern : ground color green ; a post-discal
line (outwardly white, basally dark) on both wings; a submarginal
row of red spots on the hind wing; a marginal row of spots or patches,
or a marginal band, on both wings; a basal quadrate patch of dark
brown on the hind wing. Any or all of these may be suppressed or

:The members of this section (perhaps even more sc than those of the "tailed section") seem
to bear affinity to the genus CaUophrys Billberg. Indeed. W. D. Field (1939, Univ. Kansas
Sci nee Bull., 26, p. 347) has placed Thecla hercdotus Fabr. (a member of the "tailed section")
in this genus, without comment however.

2Due to the briefness of time available for study it was not possible to make any genitalic
preparations. These, when examined, may show further, more real differences.

3So called in preference to the "tailless section" of Draudt, since that method of division has
proved incorrect. Thecla longula (pastor) and Thecla marialis, both tailed species, are clearly
members of the group now under consideration, while a tailless species, quite obviously belong-
ing to the other section, is being described as new in another paper.

4Godman and Salvin (1887, Biol. Centr.-Am. Lep., Rhop. 2, p. 34) erroneously characterize
agricolor as lacking a scent pad.

clench: notes on lycaenid butterflies 231

variously developed. An inner marginal hand- of tan or gray is always
present on the underside of the fore wing.

The several species together occupy a wide range from Mexico to
Bolivia and southern Brazil, and even Argentina. Our knowledge of
the distribution of the individual species is very imperfect, and will
probably remain so for some time to come.

Key to species

This key was very difficult to compose, and in parts may be in-
accurate; it should be used, therefore, with this in mind. In several
of the species but one sex is known, thus making it impossible to in-
clude characters which are definitely known to apply to both male
and female of such species.

1 . Frons brown 2

- Frons green 7

2. Outer margin of hind wing below with red-gray edging, either as a definite

band, or as a series of internervular spots 3

- This margin without red-gray spots, being green, as in the rest of the

wing 6

3. Outer margin of fore wing below with red-gray edging; a well-marked

discal line as well on this surface a. agricolor (1)

- This margin without red-gray edging; discal line obsolescent or wanting . .4

4. Tail at Cu2 on secondary longula (5)

- No tail at Cu2 5

5. Small, expanse less than 30 mm. (usually about 26 mm.) remits (2)

- Larger, over 30 mm.; up to 32 mm. or so agricolor banosensis (la)

6. Male with scent pad black; underside nearly uniform green . .longuloides (4)

- Male with scent pad colored blue as in the ground color; below with bands

of darker green pseudolongula (3)

7. Upperside of male brown; tailed at Cu2 on secondary marialis (9)

- This surface of male bright blue; no tail at C112 8

8. White discal line on underside of hind wing (may tend toward obsolescence)

9

- No white discal line on this wing legionis (6)

9. White patch in center of gray area on underside of fore wing (only in

male?) portoena (8)

- No white patch in this area, male or female 10

10. Submarginal red spots on underside of hind wing absent (almost or com-
pletely); white discal line prominent acaste catharinetisis (7a)

— Submarginal red spots on underside of hind wing usually well-developed;
discal white line obsolescent a. acaste (7)

232 bulletin: museum of comparative zoology

1. Thecla agricolor agricolor (Butler and Druce)

Strymon agricolor Butler and Druce, 1872, Cist. Ent., 1, p. 105; Butler, 1873,

Lep. Exot., p. 158, pi. 57, fig. 4.
Thecla agricolor: Hewitson, 1877, 111. Diurn. Lep. Lycaenidae, p. 201; Godman

and Salvin, 1887, Biol. Centr.-Am. Lep. Rhop., 2, p. 34, pi. 52, figs. 11,

12; Draudt, 1919, in Seitz, Macrolep. World, 5, p. 762, pi. 154a; Hoffman,

1940, An. Inst, Biol. Mex., 11, p. 707.

The type is in the British Museum, presumably. It was taken in
Cartago, Costa Rica. Draudt gives the range as "Mexico to Panama."
Godman and Salvin list Jalapa, Mexico; Duenas, Guatemala; Irazu
and Rio Sucio1, Costa Rica; Bugaba and Chiriqui, Panama. Hoffman
gives "Tierra templada de Vera Cruz. Sur de Puebla. Morelos.
Valle de Mexico (2250 M.)." Specimens in the M.C.Z. are all from
Jalapa, Mexico.

Above, in the male, metallic blue, duller than in the other species.
Both wings with a rather indefinite and heavy dark border on the outer
margin. Costa of fore wing narrowly black. Costa and inner margin
of hind wing narrowly gray. Anal lobe of hind wing rusty, black-
fringed. The scent-pad is very small, almost unnoticeable (see foot-
note, p. 230), and lies just beyond the upper cell-end.

Below, fore wing gray from inner margin to M3, darker adjacent
to the cell. Remaining area green. Outer margin from apex to inner
angle with a moderately heavy grayish border. A discal line of
reddish crosses from costa, disappearing shortly below M3. A faint
submarginal line behaves similarly. Hind wing with a broad marginal
border, outwardly gray, basally reddish. The basal limit of this border
is, in the M3-CU1 and 2A-inner margin interspaces, orange. A discal
line of dark green and reddish scales crosses the wing obscurely. Base
marked with a large, quadrate, almost black patch. Anal lobe maroon.

The female is duller above than the male, with the blue more re-
stricted, leaving very broad costal and outer marginal borders on the
fore wing, and a broad outer marginal border on the hind wing. It is
almost exactly similar on this surface to pseudolongula, but the outer
limit of the blue is more sharply defined.

Below, the female is similar to the male, but with the markings
slightly brighter.

Length of fore wing. Male, 12.5-13.5 mm.; female, 14.5 mm.

'Possibly this refers to a Rio Sucio in northern Colombia, emptying; into the Gulf of Darien
just below the south-eastern tip of Panama. The specimens on which this record is based may.
therefore, be transitional to bahosensis.

clench: notes on lycaenid butterflies 233

la. Thecla agricolor banosensis, new subspecies

Upperside:

Female. Both wings bright metallic blue. Fore wing with a costal
and outer marginal border of black-brown, covering the whole cell-end-
to-apex region, and narrowing towards the inner angle. Hind wing
with a gray costal border, becoming black-brown on the outer angle
and outer margin. Inner margin gray. Anal lobe rusty, the color
extending basad slightly, and costad as far as Cu2. Fringe of both
wings black-brown, paler between the veins.
Underside:

Female. Both wings green. Fore wing with a paler, apple-green
marginal stripe and a discal row of three obscure red-brown spots
(Mi-M2-M3-Cui). Inner margin to Cu2 gray, black basad, next the
cell. Between the gray and the green, in the Cui-Cu2 interspace,
is a band of fulvous. Hind wing with a small black basal patch, a
discal transverse line of gray, thin costad of Cui, heavier thence to
inner margin. A submarginal row of obsolescent red lunules from costa
to inner margin, replaced costally by dark green. On the outer margin
is a band of hoary maroon, basally scalloped from Mi to the anal
angle. Anal lobe maroon. Fringe of both wings white-gray, darker
at the vein-ends.

Length of fore wing. Female, 15 mm.

Holotype. Female, San Pablo, Rio Pastaza, vie. Banos, Ecuador,
2200 meters(?), (Clark-Maclntyre), ex coll. F. M. Brown, in the
American Museum of Natural History.

Remarks. This subspecies differs from typical agricolor1 in several
respects, namely : the absence of the marginal hoary band on the fore
wing below, reduction in size of the black basal patch of the hind wing
below, and the reduced size of the marginal band of the hind wing
below, and its differentiation into two bands. The blue color above is
very bright, far brighter than in any other female of this group thus
far examined (except for a single female of pseudolongula, of almost
the same intensity). This character, however, is in all likelihood of
no significance, being merely an age factor.

This subspecies, with its reduced markings below, strongly sug-
gests a transition from agricolor to rcmus, which, in turn, may connect
to pseudolongula. Many more specimens are needed, however, before

'The Jalapa, Mexico, specimens and the holotype of bafwsensis were compared with Butler's
figure in the Cist. Eat. (I.e.) and the former agreed almost perfectly. They were therefore used
in the following comparison as typical agricolor, even though not topotypical.

234 bulletin: museum of compaeative zoology

this suggestion can be proven or denied. Genitalic examination would
help considerably.

2. Thecla remus Hewitson

Thecla remus Hewitson, 186S, Descr. Lycaenidae, p. 34; 1877, 111. Diurn. Lep.

Lycaetiidae, p. 201, pi. 80, figs. 655-656; Draudt, 1919, in Seitz, Macrolep.

World, 5, p. 763, pi. 154b.
Thecla deidamia Burmeister, 1879, Atlas de la Descr. Phys. Rep. Argentine, 5,

pt. 2, p. 24.

Described from Brazil. Type (female) is presumably in the British
Museum, although Mr. Goodson (who examined the British Museum's
specimens of the present group for me) wasn't exactly clear on that
point in his notes. Draudt in Seitz gives no additional information,
and does not seem to have known the species. Burmeister records it
(as deidamia) from Las Conchas, north of Buenos Aires, Argentina.
Four specimens, all females, in the M.C.Z.: three from Blumenau,
Sta. Catharina, Brazil, and one, ex coll. J. Doll, labelled "Brazilia."
One male, in the American Museum of Natural History from "Massa-
randuba-Blumenau, Brazil."

Male above brilliant iridescent blue, violet-tinted in some lights.
Both wings edged with black on the outer margins. Hind wing gray
on costa and inner margin. Anal lobe dull maroon.

Below, the male is green on both wings. Fore wing with gray on
inner margin to Cu2, darkened basally. Hind wing with a marginal
row of hoary, reddish-gray spots, almost connected. A discal row of
obsolescent, irregular spots crosses the wing, between which and the
marginal row of spots is a suggestion of a row of red dashes, in M3-
CU1-C112. Anal lobe dark maroon.

The female above with the blue much duller and more restricted;
quite similar in appearance to the females of most of the other species
of the group. Anal lobe rusty. Below green, with the inner margin of
the fore wing gray, darker basad. Hind wing on the outer margin
with a series of internervular, almost round, reddish gray spots. This
wing crossed in the disk by a rather tortuous line of white, interrupted
frequently, and of varying intensity, basally edged with blackish.
Between this line and the marginal spots are one or two, rarely more,
red dashes, slightly crescentiform. Anal lobe dark maroon. Hewit-
son's descriptions (1868, 1877) of the female tallies quite well with what
is here regarded as remus, save for a few minor differences. The fulvous
patch at the outer angle of the hind wing below mentioned by Hewit-

clench: notes on lycaenid butterflies 235

son is very likely the result of wear. He mentions in both descriptions
a band of three or four spots on the fore wing below, near the eosta.
No indication of such spots was found on the four specimens examined.
He does not mention the subminimal scries of three or four crescenti-
form dashes on the hind wing. This character, however, is apparently
variable, being lacking in one of the four M.( \Z. specimens, and almost
lacking in another. On the strength of these apparent differences it
was decided not to select a type locality for remus until either speci-
mens are found that match more closely the description, or it is
proven that he had a slightly aberrant example.

The male agrees quite well with Hewitson's 1877 description and
figure, save for the "green tint" lie speaks of. This, however, is fre-
quently caused by chemical action, and can be overlooked. It is above
close to males of pseudolongula, but may be separated by the broader,
more apically thickened outer marginal border.

Thecla deidamia is a name that ever since its publication in 1S79
appears to have been almost overlooked. Seitz made no mention of it,
and the only published reference of it known to the author is that of
Weeks (see synonymy under portocna).

Weeks referred his name to "Ruschew.", but this is in error, as a
glance at the original description will show. Ruscheweyh collected the
specimens, noted that they were probably new, and suggested the
name deidamia (in Hit to Burmeister, apparently). Burmeister, how-
ever, wrote the description and applied the name to it, wherefore it
must go to him.

With regard to the present placing of the name, it can be said only
that a comparison of Burmeister's description with Hewitson's
descriptions and figures (1868 and 1877) and several specimens of
remus revealed an almost perfect resemblance. Burmeister's locality
(Las Conchas, nr. Buenos Aires, Argentina) is quite in accord with our
present knowledge of the distribution of remus, although extending
its range somewhat southward.

3. Thecla pseudolongula, new species

Thecla longula: Hewitson, 1877, 111. Diurn. Lep. Lycaenidae, p. 200, pi. 80,
figs. 651, 652, 653, 654; Draudt, 1919, in Seitz, Macrolep. World, 5, p. 762,
pi. 154a. {nee T. longula Hewitson, 1868 (q.v.)).

Eyes narrowly ringed with white. Frons with two parallel rows of
long, partially erect, dark brown hairs, flanked outwardly (along the
rims of the eyes) with a row of rusty scales on each side and a band of

/

236 bulletin: museum of comparative zoology

shorter hairs, with rusty ones intermingled, between them. Caudad of
the antennae is a transverse row of long rusty hairs. Collar of long
hairs, bluish and rusty on top, becoming intermingled with white on
the sides. Palpi rusty outwardly, bluish white within; terminal joint
completely rusty. Antennae black above, white annulate below; club
black, tipped with fulvous and paler below (becoming white basad).
Thorax of the male above metallic blue or green, overlaid moderately
with long, anally directed hairs, heaviest next the abdomen; female
paler, more steely blue above; male below covered with dense brown
hair, paler in the female. Abdomen of the male above metallic blue,
very bright; belbw, yellow; tip above and below gray: of the female,
duller, the blue more steely and more anteriorly restricted; below
similar. Legs (absent in most of the specimens examined) apparently
brown, tarsi black and white annulate.
Upperside :

Male. Bright metallic blue, greenish in some lights, purplish in
others. The scent pad is so small as to be almost unnoticeable, con-
sisting merely of a short row of scales along the upper disco-cellular
at the cell-end. A marginal border, about 1 mm. thick, edges both
wings. On the fore wing it is slightly thicker than on the hind wing,
and towards the apex it expands still more (2-3 mm.). The anal lobe
is prominent and rust-colored. Fringe brown on fore wing; dull white
on hind wing, basally and at the vein-ends darker.

Female. Dull gray-blue, with very broad blackish borders, some-
what narrower and basally crenulate on the hind wing. Anal lobe as in
the male. Fringe also as in the male.
Underside:

Male. Both wings bright emerald-green, with the inner margin of
the fore wing broadly gray. The hind wing is marked by two trans-
verse bands of lighter green, rather indistinct, basad to the inner of
which is a suggestion (consisting usually of one or two obscure points)
of a post-discal line. In the extreme base is a smallish dark brown area.
Anal lobe rusty and adjoining it in the CU2-2A interspace is an obscure
rusty -hoary patch.

Female. Similar, but the markings more distinct.
Length of fore wing. Male, 13.5-15 mm.; female, 14-14.5 mm.
Holotypc. Male, Mapoto, Ecuador, ex A. G. Weeks collection.
Allotype. Female, no locality, ex R. M. Gray collection.
Paratypes. On male, R. Guamlo (or Guamba- label poorly written),
Ecuador, ex A. G. Weeks collection; one male, "Colombia", Oct. 10,
1913, ex F. A. Eddy collection; two males, no locality (possibly

clench: notes on lycaenid butterflies 237

Bogota, Colombia), ex A. G. Weeks collection; one female, no Locality
("So. Am."), ex C. J. Paine collection; two males, one female, vie.
Banos, Ecuador (Clark-Maelntyre), as follows: one male, Tunguragua,
1900 meters, March, 1939; one male, Runtim, 2000-2500 meters,
Nov. 26, 1938; one female, Rio Blanco, 1700-1900 meters, Oct. 19,
1938.

Holotype, allotype, and three male paratypes, no. 26223 in the
Museum of Comparative Zoology. One male and one female paratype
in the author's collection. The last three paratypes in the collection of
the American Museum of Natural History.

Remarks. Quite different from true longula, for comparison with
which, see under that species. The closest ally of pseudolongula yet
discovered appears to be longuloides, from which it differs in the
smaller scent-pad, broader borders and larger anal lobe. It is also
allied to remits, but that species has a marginal row of hoary spots on
the hind wing below.

4. Thecla longuloides, new species

Eyes hairy, narrowly ringed with white. Frons consisting of two
parallel rows of long, erect, dark brownish hairs, thickly intermingled
with rusty ones. Just outside these rows, paralleling the white eye-
margin, is a row of rusty scales. Basad of the antennae is a transverse
row of rusty hairs, and between the antennae a few white ones.
Collar of long, rusty and brown-black hairs. Palpi outwardly covered
with mingled pale blue, rusty and gray scales, inwardly almost en-
tirely pale-blue. Antennae black above, white annulate below; club
black above, below tipped with dull fulvous, backed by white. Thorax
of the male above metallic greenish blue, covered, chiefly on the
periphery (behind the head, along the sides above the wing bases and
anterior to the abdomen), with long, grayish-rusty hairs; thorax of the
female grayer blue above, without the greenish tinge: below tufted
with brown in the male, paler in the female. Abdomen of the male
above brilliant metallic green, below yellow; tip gray above and
below; abdomen of the female with the blue more anteriorly restricted
above (the remaining area gray) ; below as in the male. Legs black
and white annulate.
Upperside:

Male. Both wings brilliant metallic blue, greenish in some lights
and purplish in others. Fore vying with an almost linear black scent-
pad lying along the upper discocellular at the cell-end. Costa very

238 bulletin: museum of comparative zoology

narrowly, outer margin slightly more broadly black-bordered. The
latter thickens apically and extends briefly basad on each vein. Hind
wing with costa and inner margin gray. Outer margin narrowly black,
also extending basad for a short distance on the veins. Anal lobe small,
rusty colored. Fringe of fore wing brown, paler outwardly; of hind
wing similar, but whitish outwardly between the veins.

Female. Both wings dark gray-brown, costa and inner margin of
hind wing pale brown. Fore wing with dull steely blue from inner
margin to upper discocellular, from base to cell-end, outwardly down
to inner margin three-quarters out. Hind wing similar, leaving only
a narrow dark border on the outer margin, hazy and indistinct basad,
that thickens slightly towards the outer angle. Anal lobe as in the
male. Fringe as in the male, but darker.
Underside:

Male. Both wings uniform green. Fore wing with the inner margin
to just over Cua gray, becoming sharply black along the lower disco-
cellular. Hind wing with two very faint and obscure small white spots,
each lined basally with a few red scales: one post-discal in the CU1-CU2
interspace; the other submarginal in the 2A-3A interspace, touching
2A. The faintest indication of a submarginal band, almost un-
noticeable, consists merely of a very slight darkening of the green.
Fringe as on upper surface. Anal lobe obscurely dark rusty colored.

Female. Similar to the male, but lacking the black along the lower
discocellular of the fore wing, and with the faint submarginal band of
the hind wing a little more prominent. The green on the outer margin
of the fore wing extends down more into the (A12-IA interspace.

Length of fore wing. Male, 14 mm.; female, 13 mm.

Holoti/pe. Male, Coroico, Bolivia, May, 1899 (Wm. J. Gerhard),
ex A. G. Weeks collection.

Allotype. Female, Chulumani, Bolivia, Dec. 12, 1898 (Wm. J.
Gerhard), ex A. G. Weeks collection.

Holotype and allotype, no. 26224 in the Museum of Comparative
Zoology.

Remarks. This species is allied to pseudolongula, but may be dis-
tinguished from it, in the male, by the much narrower outer marginal
border above, and the presence of a larger, more definite scent-pad.
Both sexes have a considerably reduced anal lobe on the hind wing
(less than half the size of that occurring on pseudolongula). Below the
green is more uniform than in pseudolongula, with the light and dark
transverse bands almost non-existent. In the female the outer margin
of the fore wing does not seem to be so convex as in that sex of pscudo-

clench: notes on lycaened butterflies _'■>'>

languid, and the blue appears to be duller, although of about equal
extent.

5. Thecla longula Hewitson

Thecla longula Hewitson, 1868, Descr. Lycaenidae, p. 34. (nee longulti, Hewit-
son, 1877, and others).

Strymon pastor Butler and Druce, 1872, Cist. Ent., 1, p. 105; Butler, 1873,
Lep. Exot., p. 157, pi. 57, fig. 5; McDunnough, 1938, Mem. S. Cal. Acad.
Sci., 1, p. 24 (no. 364).

Thecla pastor: Godman and Salvin, 1887, Biol. Centr. Am. Lep. Rhop., 2, p. 34,
pi. 52, figs. 8, 9, 10; Draudt, 1919, in Seitz, Macrolep. World, 5, p. 762,
pi. 154a; Hoffman, 1940, An. Inst, Biol. Mex., 11, p. 707.

The type apparently is not in the British Museum. Mr. F. W.
Goodson, at Tring, informs me through Dr. Riley that there is no
specimen of longula (by which he meant pseudolongula, probably
exclusively) in the British Museum from Central America. The four
specimens of ''longula" from the Hewitson collection are all referable
to pseudolongula.

Five specimens of longula have been examined and compared with
the description. They seem typical, and in the absence of a type are
made neotypes, as follows:

Xcoholotypc. Male, Orizaba, Mexico, June 1941 (Stallings and
Turner).

Neoallotype. Female, same data as above.

Neoparatypes. One female, same data as above; two females,
Presidio, Mexico, June 1941 (Stallings and Turner).

Xeoholotype and Neoallotype deposited in the Museum of Com-
parative Zoology. One neoparatype in the collection of the author.
The remaining returned to the collection of Mr. D. B. Stallings and
Dr. J. R. Turner.

This species has been subjected to a rather peculiar misidentifica-
tion, for almost the whole of its existence in the literature to date.
Hewitson's original description of longula is of a totally different
insect from that of his description and figure of 1877 (in the "Illustra-
tions"), as can be seen from the following extract from it (1868):
"Underside dull green, tinted with orange at the apex of the anterior
wing1. Posterior wing without tails:2 crossed beyond the middle by

UVohably due to wear: see under remus.

2Evidently variable in this species. One of Godman and Salvia's illustrations of pastor
showed tailless, and one of the type ineotype< serii>s of bmijula is also naturally without tails.
Possibly the species is in a state of either losing or acquiring them.

240 bulletin: museum of comparative zoology

two bands of indistinct distant red-brown spots : a series of marginal
red-brown spots, irrorated with white: the lobe red-brown." Compar-
ing this description with that of his published 1877 (pseudolongida of
this paper) shows immediately that the two are not conspecific. When
the above description was checked against pastor, the true identity
of the name was shown.

6. Thecla legionis, new species

Eyes hairy, ringed with white or pale green. Frons green. Collar
and palpi in the present specimen indiscernible. Antennae black and
white annulate, nearly all black above; club black, tipped with fulvous,
more extensive and white-backed below.- Thorax pale steely blue
above, fulvous beneath, rather pallid where visible. Abdomen gray
above (pale bluish basad), cream below.
Upperside:

Female. Both wings steely blue. Fore wing broadly black on costa,
apex and outer margin. Hing wing more narrowly so on outer margin.
Costa and inner margin gray. Anal lobe rusty. Fringe of fore wing
blackish, slightly paler outwardly; of hind wing sordid white, darker
at the vein-ends.
Underside:

Female. Both wings green. Fore wing with inner margin from Cu2
gray. Cu2 and outer margin along this gray are fulvous. Base of gray,
adjoining cell, darker. Hind wing unmarked save by a series of
obscure reddish dashes in the M2-2A interspaces slightly basad of the
submarginal area, and two almost unnoticeable white costal spots,
one each in the Sc-Rs-Mi interspaces. Outward of the reddish dashes
the green seems a little paler. Fringe of fore wing fulvous, outwardly
obscured by gray. Of hind wing fulvous, outwardly whitish, darker at
the vein-ends.

Length of fore wing. Female, 12.5 mm.

Holotypc. Female, Blumenau, Sta. Catharina, Brazil (B. Pohl),
no. 26225 in the collection of the M.C.Z.

Remarks. This species, unfortunately represented by but a single
female, appears to stand closest to acaste. It has the green frons of
acaste, but the almost unproduced anal angle of, for example, remus.
Below it looks somewhat similar to a small acaste without a discal
white line on the hind wing. Above, the blue is slightly paler than in
females of acaste, but this may be due to fading. Below legionis
differs from typical acaste (which subspecies it most closely resembles)

clench: notes on lycaenid butterflies 241

in the almost complete absence of a discal white line, the only indica-
tion being two almost costal white spots, so faint as to be hardly
discernible. The fringe is also more fulvous than in either subspecies of
acaste. The fulvous edging of the inner marginal area of the fore wing
below is also absent in both acaste and catharinensis, but this character,
like the fulvous patches mentioned by Hewitson in the descriptions of
longula and remus, may be due to Avear.

7. Thecla acaste acaste Prittwitz

Thecla acaste Prittwitz, 1865, Stett. Ent. Zeit,, 26, p. 31S; Draudt, 1919, in

Seitz, Macrolep. World, 5, p. 763, pi. 154a.
Thecla lycimna Hewitson, 1868, Descr. Lycaenidae, p. 33; 1877, 111. Diurn.

Lep. Lycaenidae, p. 203, pi. 80, figs. 663, 664, 665.

The type locality of acaste is Corcovado, near Rio de Janeiro,
Brazil. Under his lycimna Hewitson (1S77) merely gives "Brazil."1
Mr. Goodson has examined the Hewitson type for me, and it is ap-
parently quite in accordance with Prittwitz' original description, and
has the characters here used to separate true acaste from catharinensis.
Rio de Janeiro, Brazil, is here selected as the type locality of lycimna,
thus better insuring its permanent synonymy. The type of acaste,
Mr. Goodson suggests, is probably either in Munich, Berlin, or Greis-
swald, granting the collections at those localities to be still intact.

Draudt gives as records: Sao Paulo, Santa Catharina and Rio
Grande do Sul. The last two will probably refer to catharinensis.
Seven examples in the M.C.Z., Rio de Janeiro and Canto Gallo, Brazil.
The range of this, the typical subspecies, appears to be quite limited.

The male above is generally similar to longuloides, but with the
scent-pad concolorous with the ground. The ground color is slightly
duller than in longuloides, especially marginally, and the outer margin
is more broadly black. Below it is uniform green, with a gray inner
margin on the fore wing, an obsolescent white transverse line on the
hind wing running from costa to inner margin, outward of which is a
row of tiny bright red spots or dashes, the most prominent in Cui-Cu2.

Female above similar to the female of pseudolongula, but with the
outer limits of the blue less definite. Below as in the male.

Both sexes differ greatly from pseudolongula and longuloides in the
possession of a green frons.

Length of fore wing. Male, 15-15.5 mm.; female, 14-16 mm.

!The original description (1868) cited no locality whatsoever.

242 bulletin: museum of comparative zoology

7a. Thecla acaste catharinensis, new subspecies
Thecla acaste: Draudt, 1919, in Seitz, Macrolep. World, 5, p. 763 (partim.

Upperside:

Male. Both wings dully shining violet blue, deepening slightly
towards the margin. Outer margin of both wings narrowly black,
thickening slightly towards the apex on the fore wing. Costa of fore
wing also narrowly black. Costa and inner margin of hind wing gray,
the latter shaded basally with bluish. On the hind wing the anal lobe
is rusty, fringed with black.

Female. Both wings black-brown with the basal two-thirds of the
fore wing and the majority of the hind wing dull steely blue, leaving a
costal, broad apical, narrower outer marginal dark border on the fore
wing, and a gray costa and inner margin on the hind wing, with a still
narrower dark outer marginal border. Anal lobe as in the male.
Underside:

Male. Both wings bright green. Fore wing inner margin gray to
Cu2, the green encroaching only at the outer margin. Base of this
gray area dark along the lower Dc. Hind wing with a prominent,
mildly tortuous white line, basally and obscurely bordered with red.
This line commences two-thirds out on the costa and proceeds nearly
straight to three-quarters out on inner margin. Anal lobe black,
shading into deep red basad. A few minute black scales in the sub-
marginal area.

Female. Similar to the male.

Length of fore wing. Male and female, J4 mm.

Holotype. Male, Santa Catharina, Brazil, ex A. G. Weeks collection.

Allotype. Female, Blumenau, Sta. Catharina, Brazil (B. Pohl).

Paratype. One female, "Brazilia", ex J. Doll Collection.

Holotype and allotype, M.C.Z. no. 26226. Paratype in the collec-
tion of the author.

Remarks. Differs from typical acaste in the more prominent and
complete white line on the under surface of the secondary, in the
absence of the submarginal red dots or dashes on the same wing below,
and, in the male, by the reduced marginal border above.

8. Thecla portoena, new species

Thecla deidamia: Weeks, 1905, 111. Diurn. Lep. (111. Unfig'd. Lep.), 1, p. 19.
(nee deidamia Burmeister: see under remits. The identity of the insect
Weeks called deidamia (loc. cit.) is determined by three specimens so
labelled in his collection, which are now made the types of portoena.)

clench: notes on lycaenid butterflies 243

Eyes hairy, ringed with white. Frons green. Collar dark rusty
above, shading to whitish below. Palpi rusty pale gray, dorsally
dark brown. Antennae black, white annulate below, and very faintly
above; club black, tipped with dull fulvous. Thorax dull black,
covered with bluish hairs, lightly on top, heavier laterally and next
the abdomen; below grayish tan. Abdomen above blue next the
thorax, gray thence to tip; below yellow, gray at the tip. Legs black
and white annulate.
Upperside:

Male. Both wings shining lavender blue, deepening towards the
margin. Fore wing with a rather broad, dark brown marginal border,
thickening apically. Costa with a similar, but narrower border.
Hind wing with costa pale gray. Inner margin gray, overlaid basad
with bluish scales. Basal area of wing overlaid with pale scales, giving
a rather hoary appearance to this region. Outer margin narrowly
black-brown, extending slightly basad on the veins. Anal lobe rusty,
black-fringed. Fringe of fore wing dark basally, paler outwardly; of
hind wing dark, white outwardly between the veins.
Underside:

Male. Both wings uniform green. Fore icing with the inner margin
gray-tan, with a white patch on the center. The green encroaches on
this gray-tan at the outer margin. Base of this area somewhat dark-
ened. Hind wing with a nearly straight white discal line, basally
edged with a few red scales, that runs from the costa towards the
inner margin, but disappears at about M3 or Cui. A submarginal series
of thin red dashes occupies the M3-2 A interpaces, and occasionally even
further costad. Anal lobe deep red, almost maroon, extending slightly,
in the form of a compact small area of reddish irroration, into the
Cu2-2A interspace.

Length of fore wing. Male, 12-13.5 mm.

Holotype. Male, Cusilluni, Bolivia, May, 1899 (Wm. J. Gerhard),
ex A. G. Weeks collection.

Paratypes. Two males: one male, same data as holotype; one male,
Coroico, Bolivia, April 20, 1S99 (Wm. J. Gerhard), ex A. G. Weeks
collection.

Holotype and one paratype, M.C.Z. no. 26227. One paratype in
the author's collection.

Remarks. T. portocna may be distinguished from both typical
acaste and its subspecies cathariensis as follows : the marginal border of
the fore wing above (male) is broader than in either; below, the white
line on the hind wing disappears before reaching the inner margin,

244 bulletin: museum of comparative zoology

while in both acaste and a. catharinensis it proceeds all the way; the
anal lobe is here smaller than in either, and colored deep red, while in
acaste and its subspecies it is black; the inner marginal area of the fore
wing below is here gray-tan, and has a central white patch, while in
acaste and catharinensis it is gray and has no such patch (in fact
portoena is the only species at present known to the writer that
possesses such a peculiar pattern character) ; T. portoena has the red
submarginal spots as in acaste (s.s.), but they are more linear, and seem
to extend further cost ad.

It is possible that portoena may be only a subspecies of acaste. The
differences between them, however, are such that two full species seem
involved, and while at present portoena seems to be a local representa-
tive of acaste, further knowledge of the distributions of the involved
forms may prove otherwise.

9. Thecla marialis, new species

Eyes ringed with pale green. Frons green. Palpi fulvous, scaled
outwardly with green ; terminal point black. Collar above green, with
pale greenish and fulvous hairs, shading to sordid gray on the sides.
Thorax above black with dull greenish-gray hairs frontad, laterally,
and next the abdomen, all back-directed; below covered with pale
rusty long hairs. Abdomen black-brown above, yellow below, dark
gray at the tip. Legs gray, with pale annulations.
Upperside:

Male. Both wings black-brown. Fore wing with the basal area over-
laid with dull olive-green, extending marginad roughly two-thirds.
On the upper cell-end is an elongated black scent-pad, rather small.
Hind wing similar but with the olive-green shading restricted more
basad. Anal lobe fulvous, this color extending basad along inner
margin for one-third its length, and along outer margin almost to Cu2.
Tail at Cu2, very short, but definitely present. Fringe of both wings
gray with a brownish tinge.
Underside :

Male. Both wings emerald-green. Fore wing with inner margin to
Cu2 gray, darkening to almost black, next the cell, and to darker gray
on the outer margin. Hind wing with an almost non-existent indication
of a post-discal line, the only real relic being a tiny white spot in
Cui-Ci^, capped by a minute red bar. Anal lobe dark maroon, capped
thinly basad by a white line.

Length of fore wing. Male, 13 mm.

clench: notes on lycaenid butterflies 215

Holotype. Male, Victoria, Mexico, February 7, 1942 (Mrs. Mary
Alice Turner), no. 26569 in the Museum of Comparative Zoology.

Remarks. From the remaining species of the group now under con-
sideration this species differs most remarkably in the utter absence of
the brilliant morpho-blue that characterizes the males. Other dis-
tinguishing characters are: the C112 tail, present only in this species
and in longula; and the spread of fulvous from the anal lobe above.
The other differences are of less importance, but can be noted by
referring to the formal description above.

This species is a striking parallel to Thecla fusius Godman and
Salvin1, which belongs to the "tailed section" of Draudt's amyntor-
group. Fusius (of which also only the male is known) is uniformly
brown above, but below is described and figured as being exactly simi-
lar to hcrodotus Fabr.2 From Godman and Salvin's figures the follow-
ing presumably significant differences have been noted : The hind wing
is longer, more produced anally (a character of the acaste group, as
opposed to the remainder ("tailed section") of the am yutor -group);
there is a very plain anal suffusion of fulvous in marialis, of which no
indication is given either in Godman and Salvin's description, or their
figure; nor is there, in marialis any indication of the bluish suffusion in
the base, as depicted in their illustration ; below there is no white line
on the hind wing of marialis, while the figure of fusius plainly shows
one, and in which fusius also agrees with hcrodotus.

There seems little doubt, therefore, that in spite of the superficial
resemblance, we are dealing with a full species, belonging even to a
separate subgroup.

The collector of this subspecies, Mrs. Mary Alice Turner, said that
she could not recall taking the specimen itself, but remembered that
collecting at Victoria wras done by an irrigation ditch, on low weeds
in open country away from the forest.

The author wishes to thank Mr. Don B. Stallings, of Caldwell,
Kansas, in whose collection the specimen formerly rested, for his kind
donation of it to the Museum of Comparative Zoology.

1 1877, Biol. Centr. Am. Lep. Rhop., 2, p. 34, pi. 52, fig. 6, 7.

2 Godman and Salvin state that the pattern below of fusius is so similar to herodotus that,
in the absence of females, they were almost inclined to regard it as a dimorphic male of that
species.

Bulletin of the Museum of Comparative Zoology
AT HARVARD COL LEG E

Vol. XCIV, No. 7

THE SOCIAL VESPIDAE OF THE GUI AN AS, PARTICU-
LARLY OF BRITISH GUIANA

By Joseph C. Bequaert

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

August, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1, 2,
3, 4, 5 and 6 have appeared and of the Memoirs, Vol. I to LVL

These publications are issued in numbers at irregular intervals.
Each number of the Bulletin and of the Memoirs is sold separately.
A price list of the publications of the Museum will be sent upon
application to the Director of the Museum of Comparative
Zoology, Cambridge, Massachusetts.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HA R YARD CO L L E G E

Vol. XCIV, No. 7

THE SOCIAL VESPIDAE OF THE GUIANAS, PARTICU-
LARLY OF BRITISH GUIANA

By Joseph C. Bequaert

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

August, 1944

Xo. 7 — The Social Vespidae of the Guianas, "particularly
of British Guiana

(Hymcnoptera)
By Joseph C. Bequaert

The present comprehensive account of the social wasps (Vespidae)
includes keys for the identification of the South American genera and
of the species known from any of the three Guianas. This area, based
on political boundaries, may be regarded as a fairly natural unit,
although the fauna is similar to that of the adjoining forests of northern
Brazil and eastern Venezuela. Its wasp fauna is, moreover, rather uni-
form and I have no doubt that all the species included in this paper
will eventually be found in British Guiana.

For the present, only the forms definitely known from British Guiana
are fully discussed, all the recorded localities of these being listed.
Social wasps live in colonies of several or many individuals and are
therefore abundant enough to be collected casually. Their nests are
also an added attraction to the general collector. Hence they are much
better known than most other Hymenoptera, with the possible excep-
tion of the social bees. Most probably few South American species
remain to be discovered, and this is particularly true of the Guianas,
one of the parts of tropical America most thoroughly investigated by
entomologists.

The Guianas are unusually rich in social wasps and would be ideal
for the study of their fascinating habits. I list from the entire area
98 structural species (and 32 additional color varieties), belonging to
16 genera (all those known from South America, except three: Synoe-
coides, Clypearia and Protonectarina). Of these, 84 species (and 18
varieties) are definitely recorded from British Guiana, 25 of them (and
8 varieties) having not yet been taken in either Dutch or French
Guiana. The wasp fauna of the British portion thus appears to be
somewhat better known than that of the others, the differences being
entirely due to insufficient collecting, in my opinion.

The following 43 species and varieties were taken at Kartabo within
the quarter square-mile area of jungle extensively studied by the New
York Zoological Society's Department of Tropical Research.

Polistes versicolor var. vulgaris
Polistes testaceicolor
Polistes melanosoma
Polistes deceptor
Mischocyttarus carbonarius

250 bulletin: museum of comparative zoology

Mischocyttarus collarellus

Mischocyttarus duckei

Mischocyttarus flavicans

Mischocyttarus labiatus

Mischocyttarus prominulus

Mischocyttarus smithii

Mischocyttarus superus

Polybia liliacea

Polybia striata

Polybia jurinei

Polybia bifasciata

Polybia bifasciata var. quadricincta

Polybia chrysothorax

Polybia micans

Polybia rejecta

Polybia occidentalis

Polybia occidentalis var. parvula

Polybia bistriata

Polybia tinctipennis var. nebulosa

Stelopolybia cajennensis

Stelopolybia pollens

Stelopolybia obidcnsis

Stelopolybia constructrix

Stelopolybia testacea

Stelopolybia pallipes var. anceps

Metapolybia cingulata

Protopolybia minutissima

Protopolybia mimdissima var. binominata

Protopolybia pumila

Apoica pallida

Apoica pallida var. arborea

Apoica pallida var. pollens

Apoica pallida var. albimacula

Brachygastra scutellaris

Synoeca surinama

Synoeca virginea

Parachartergus smithii

Sixteen additional forms listed below were taken in nearby localities,
within a fifteen-mile radius from Bartiea :

Pseudochartergus chartergoides
Mischocyttarus foveatus
Mischocyttarus heliconius
Mischocyttarus lemoulti

bequaert: social vespidae of ghe guianas 251

Mischocyttarus oecothrix

Mischocyttarus rohindicollis

Polybia catillifex

Polybia dimidiata

Polybia gorytoides

Polybia rufitarsis

Polybia sericea

Polybia singularis

Stelopolybia paraensis

Stelopolybia angulata

Stelopolybia fulvofasciata

Prot/)polybia holoxantha

Parachartcrgus fulgidipennis var. griseus

The combined two lists of 59 forms give a fair idea of the wasps to
be found in that particular section of British Guiana. Additions may
be expected, some of the more common species of the coastal lowlands,
such as Polistes canadensis and Brachygastra lecheguama, not being
included. Nevertheless, it is fairly certain that several of the Guiana
species do not occur in the Bartica area, particularly those that have
been reported only from the hilly or mountainous areas near the
Amazon watershed (Alt. Roraima, etc.).

Acknowledgments. The present study was based upon material
derived from many sources. Foremost were the collections made over
a number of years in the Kartabo district by Dr. W. Beebe and his
associates at the Tropical Research Station of the New York Zoological
Society. Additional specimens were collected by B. E. Dahlgren,
C. Geijskes, A. Mackie, the late J. G. Myers, J. Ogilvie, O. W. Rich-
ards, Neal Weber, the late W. M. Wheeler, and F. X. Williams. I
have also included Guiana records from the collections of the American
Museum of Natural History, Carnegie Museum, Cornell University
(Dept. of Entomology), Field Museum of Natural History, Museum of
Comparative Zoology and United States National Museum.

References to the original descriptions of most of the species and
color forms, and their synonyms, mentioned in this paper will be
found in Ducke's Catalogue of the Social Wasps of Brazil (1918, Rev.
Mus. Paulista, 10, pp. 313-374).

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Table of Neotropical Genera of Social Vespidae
(Based Mainly on Females and ^Yorkers)

1. Third and fourth segments of mid and hind tarsi asymmetrical,

with inner apical lobe much longer than outer one. First ab-
dominal segment much narrowed, stalk-like. . . . Mischocyttarus
All segments of mid and hind tarsi symmetrical 2

2. Thoracoabdominal muscle inserted in a narrow, slit-like furrow

at the lower end of the propodeum. First abdominal segment

short or slightly elongate, but not stalk-like Polistes

Thoraco-abdominal muscle inserted in a broadly ovate furrow at
the lower end of the propodeum 3

3. Ocelli very large, nearly as wide as basal diameter of flagellum.

Mesepisternum completely divided into an upper and a lower
plate. Abdomen long and slender, but first segment not stalk-
like Apoica

Ocelli normal or slightly swollen, much less than basal diameter of
flagellum. In doubtful cases, mesepisternum undivided 4

4. Scutellum very prominent, its vertical hind face forming an angle

with the horizontal anterior portion, which projects beyond the
short, vertical postscutellum. First abdominal segment short

and narrowed, but not stalk-like Brachygastra

Scutellum and postscutellum placed one behind the other in one
slope or convexity; scutellum sometimes slightly more raised,
but not projecting beyond postscutellum 5

5. Hind margin of postscutellum considerably extended as a sharp

angle or broad lobe into the median concavity of the propo-
deum 6

Hind margin of postscutellum either straight or forming a slight
obtuse angle or a broad curve 7

6. Clypeus usually much longer than wide, with nearly straight api-

cal margin. Outer orbits very narrow. Postscutellum divided
into a short, horizontal basal area and a vertically abrupt

apical portion Pseudochartergus

Clypeus either wider than long or at most about as long as wide,
more or less produced medially at apex. Outer orbits of normal

width. Postscutellum forming a single oblique convexity

Protopolybia

7. Labial palpi bearing a heavy, erect, curved seta some distance

from the tip. Clypeus not longer than wide, more or less pro-
duced medially at apex 8

bequaert: social vespidae of the guianas 253

Labial palpi always of 4 segments, without erect, heavy seta
before the tip. Maxillary palpi of 6 segments 10

8. Maxillary palpi of 5 segments; labial palpi of 3 segments. Small

species, with stalk-like first abdominal segment Lcipomeles

Maxillary palpi of 6 segments. First abdominal segment sometimes
narrower than second, but rarely stalk-like 9

9. Labial palpi of 4 segments Pseudopolybia

Labial palpi of 3 segments Parachartergus

10. First abdominal tergite distinctly divided into a slender basal

stalk and a broader apical portion which is part of the remaining
swollen abdomen. Outer orbits narrow. Clypeus longer than

wide, with nearly straight apical margin Charterginus

First abdominal tergite either without basal stalk or, if stalked,
the remainder of the tergite is also much narrower than the
succeeding segments and not part of them 11

1 1 . Clypeus much longer than wide, with nearly straight apical margin.

Outer orbits very narrow 12

Clypeus either wider than long or at most as wide as long, rarely
slightly longer (in some males longer than wide) ; apical margin
more or less produced, sometimes minutely bidentate. Outer
orbits normal 13

12. Abdomen subsessile, the first segment not at all stalk-like. Dor-

sum of thorax flattened. Postscutellum oblique throughout ....

Synoecoides

Abdomen with the first segment distinctly stalk-like. Dorsum of

thorax not flattened. Postscutellum with a narrow basal area,

followed by an abruptly vertical portion Clypearia

13. Abdomen subsessile, the first segment not stalk-like but posteri-

orly part of the remaining swollen abdomen. Postscutellum
with a narrow basal area slightly tuberculate in the middle,

followed by an abruptly vertical portion Chartergvs

First abdominal segment more or less stalk-like, always much
narrower than the succeeding swollen segments. Postscutellum
not so divided 14

14. First abdominal segment very long and slender, nearly as long as

thorax, with prominent spiracular tubercles Metapolybia

First abdominal segment much shorter than thorax; spiracular
tubercles weak 15

15. First abdominal segment forming a linear stalk with subparallel

sides, flattened above; second segment abruptly widened.
Clypeus with two minute apical teeth Epipona

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First abdominal segment always much wider at apex than at base.
Clypeus ending in one tooth or obtusely rounded off at apex. . 16

16. First abdominal segment with slightly prominent spiracular

tubercles ; remaining segments abruptly widened at base, sharply

conical and somewhat compressed apically Synoeca

Abdomen elongate-oval and more or less depressed 17

17. Clypeus very wide, almost twice as wide as long in the female.

Ocelli far apart, as far from one another as from the eyes

Protonectarina

Clypeus usually as wide as long or slightly longer than wide or

moderately wider; very rarely almost twice as wide as long, in

which case the ocelli are close together 18

18. Mesepisternum undivided Polybia

Mesepisternum divided by an oblique suture into an upper and a

lower plate Stelopolybia

The following names proposed for supra -specific groups of Neotropi-
cal social wasps are not given generic status in this paper.

Agelaia Lepeletier (1836), based upon Agelaia fuscicomis Lepele-
tier (1836), is at present unrecognized. The species was described
without locality. I suggest that it may have been a Polistes.

Caba R. v. Ihering (1904) = Brachygastra.

Chartergcllus J. Bequaert (1938). Subgenus of Parachartergus.

Clypeo polybia Brethes (1923) was based on a species of Mischocyt-
tarus.

Coloboclypeus Brethes (1926) is at present unrecognized. Perhaps it
was based upon a solitary vespid.

Eupolybia Dalla Torre "(1904) = Polybia.

Gymno polybia Ducke (1914) = Stelopolybia. See the discussion under
that genus.

Hypochartergus Zavattari (1906). Subgenus of Charter ginus.

Megacanthopus Ducke (1904). Subgenus of Mischocyttarus.

Melissaia Shuckard (1841) = Brachygastra.

Monacanthocnemis Ducke (1905). Subgenus of Mischocyttarus.

Myrapetra White (1841) = Polybia.

Nectarina Swainson and Shuckard (1940) = Brachygastra.

Nectarinella J. Bequaert (1938). Subgenus of Parachartergus.

Tatua H. de Saussure (1854) = E pi pona.

Xanthocaba "Cameron" Meade Waldo (1914) = Pseudo polybia.

bequaert: social vespidae of the guianas 255

Subfamily POLISTINAE

Polistes Latreille (1802)

This genus is represented in South America by more structural
species than in any other part of the World. Those known from the
Guianas or likely to occur there may be separated by the following key.

1 . Mesopleura with not even a trace of prepectal suture 2

Mesopleura at least with traces of prepectal suture (in most cases

strongly marked) 7

2. Body thickset. Abdomen ovate-fusiform, widest before mid-

length, more or less depressed apically; first tergite strongly
convex and, in profile, abruptly sloping toward the base, seen

from above as wide at apex as long or wider 3

Body slender. Abdomen elongate-fusiform, widest about mid-
length, more or less compressed apically ; first tergite moderately
convex and, in profile, gradually sloping toward the base, seen
from above longer than wide at apex 4

3. Female: head much swollen; oculo-malar space longer than one-

third of height of eye seen in front; clypeus not or barely touch-
ing eyes. Male: clypeus pentagonal, with bluntly pointed apex,
little or scarcely separated from lower inner orbits . . P. carnifex
Female: Head moderately swollen; oculo-malar space at most as
long as one-third of height of eye, usually shorter; clypeus touch-
ing inner orbits for a distance equalling one-third to one-half of
oculo-malar space. Male: clypeus subquadrate, with straight
or weakly curved anterior margin, widely separated from inner
orbits P. major

4. Propodeum very distinctly or coarsely striate 5

Propodeum finely or obsoletely striate 6

5. Occipital carina lacking over lower third of outer orbit, where the

cheek is completely rounded off into the gula; upper part of
outer orbit with an irregular slight depression near marginal

carina P. canadensis

Occipital carina continued over lower third of outer orbit, though
much weaker than in upper part, the cheek separated from the
gula by a distinct ridge; upper part of outer orbit without de-
pression near marginal carina P. apicalis

6. Occipital carina strong, continuing along lower outer orbit to base

of mandible. Humeral collar of pronotum strongly raised

P. goeldii

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Occipital carina low, absent along lower fourth of outer orbit,
where the cheek is rounded off into the gula. Humeral collar of
pronotum moderately raised P. versicolor

7. Propodeum coarsely striate. Occipital carina absent along lower

half of outer orbit, where the cheek is rounded off into the gula.
Clypeus touching eyes over a short distance in female, widely
separated from them in male. Mesepisternum divided by an

oblique suture into an upper and a lower plate P. major

Propodeum finely or obsoletely striate. Occipital carina usually
continuing along outer orbit to base of mandible 8

8. Outer orbit much widened in upper third, where the occipital

carina is strongly raised and wing-like. Male: clypeus longer
than wide, broadly rounded off at apex; all segments of flagellum

longer than wide P. occipitalis

Occipital carina not strongly raised nor wing-like at upper third of
outer orbit 9

9. Head much swollen ; outer orbit markedly wider than eye in profile ;

occipital carina slightly sinuate about mid-height. Mesepister-
num without oblique suture dividing it into an upper and a lower

plate 10

Head not swollen; outer orbit at most as wide as eye in profile;
if wider, mesepisternum partly divided by an oblique suture. 11

10. Clypeus touching eyes over a distance equal to about one-third

of length of oculo-malar space. (Head and thorax black; abdo-
men red) P. bicolor

Clypeus touching eyes over a distance equal to a little over half the
length of oculo-malar space. (Entirely oily black) . . P. deceptor

11. Body thickset. Abdomen ovate-fusiform, widest before mid-

length, more or less depressed apically; first tergite strongly
convex and, in profile, abruptly sloping toward base, seen from
above as wide at apex as long or wider. Occipital carina usually

weak along lower outer orbit near base of mandible 12

Body slender. Abdomen elongate-fusiform, widest about mid-
length, more or less compressed apically; first tergite moder-
ately convex and, in profile, gradually sloping toward base, seen
from above longer than wide at apex. Occipital carina usually
high and ridge-like along outer orbit to near base of mandible . 13

12. Mesepisternum with at least a trace of oblique suture. Clypeus of

both sexes touching eyes over a distance equal to at most one-
third of length of oculo-malar space ; lower subocular portion in
female longer than upper interocular part. Male : clypeus ending

bequaert: social vespidae of the guianas 257

in a bluntly pointed apex; thirteenth segment of antenna slightly

curved P. ruficornis

Mesepisternum without even a trace of oblique suture. Clypeus
of both sexes touching eyes over a distance equal to one-half or
more of oculo-malar space; lower subocular portion in female
about as long as upper interocular part. Male: projecting apex
of clypeus broadly rounded off; thirteenth segment of antenna
straight P. pacificus

13. Occipital carina very faint or obsolete over lower third of outer

orbit. Face of male much lengthened; clypeus higher than wide,
touching eyes over a distance equal to about one-third of length

of oculo-malar space. Thorax densely silky P. svbsericeus

Occipital carina sharp as far down as base of mandible. Face of
male not conspicuously lengthened; clypeus about as wride as
high. Thorax not densely silky 14

14. Male: Occipital carina very high and collar-like along outer orbit;

outer orbit narrower than eye in profile ; clypeus about as wide
as long; mesepisternum with a complete but very weak oblique
suture. (Head and thorax black; abdomen red. Female not

seen) P. erythrogaster

Occipital carina low, not collar-like along outer orbit 15

15. Outer orbit wider than eye in profile. Clypeus of both sexes touch-

ing eyes over a distance less than length of oculo-malar space.
Oblique suture of mesepisternum distinct in lower half, obsolete
in upper half. (Head, thorax and two basal segments of abdo-
men mostly russet with yellow markings, remainder of abdomen

black) P. testaceicolor

Outer orbit as wide as or slightly narrower than eye in profile.
Clypeus of both sexes touching eyes over a distance about equal
to length of oculo-malar space. Oblique suture of mesepisternum

lacking or very faintly indicated. (Almost entirely black)

P. melanosoma

Polistes bicolor Lepeletier (1836) is known from French Guiana,
Dutch Guiana, Brazil (Amazon Basin), Colombia and Peru.

Polistes pacificus Fabricius (1804). No doubt some of the color
forms of this species occur in British Guiana. The typical form is
known from French Guiana, as well as from Mexico, Colombia, Trini-
dad, Brazil and Peru; var. liliaciosus H. de Saussure (1854), from
French Guiana, Brazil and Peru; var. actaeon Haliday (1836) is defin-
itely known from Brazil and Paraguay, with one rather doubtful

258 bulletin: museum of comparative zoology

record from French Guiana; and var. geminatus Fox (1898) I have
seen from French Guiana and Brazil.

Polistes apicalis H. de Saussure (1858) was described from "Guiana".
There is as yet no definite record from British Guiana. I have seen it
from Guatemala, Honduras and Ecuador.

Polistes carnifex (Fabricius, 1775). This species was discussed in a
paper published in 1936 (Rev. de Entomologia, 6, pp. 376-383) It
occurs from central Mexico to northern Argentina (Misiones) in a
number of color forms, four of which I have distinguished by name.
As it is often confused with Polistes major (Palisot de Beauvois), I
have included that species also in my key, although it does not seem
to occur in the Guianas. Typical carnifex has not been reliably reported
from the Guianas, but Polistes variegata Lepeletier (1836), described
from Cayenne, may have been this form. Var. rufipennis Latreille
(1817) is known only from Panama, French Guiana (Polistes chloro-
stoma Lepeletier, 1836, from Cayenne, is a synonym) and Dutch
Guiana; and var. ochreata Spinola (1851), from Dutch Guiana, Trini-
dad, Tobago, Colombia, and Bonacca (off the coast of Honduras).
Both these color forms no doubt occur in British Guiana.

Polistes versicolor (Olivier, 1791)

One of the most common social wasps of South America, where it
varies extraordinarily in color. In a revision published in 1934 (Rev.
de Entomologia, 4, pp. 147-157), I recognized nine varieties by name.
I have since described two more (1940, Ent. News, 51, pp. 81-82).
Four of these are known from the Guianas.

1. Typical form. Only first and second abdominal tergites marked
with yellow. I have seen it from Brazil, northern Argentina, Paraguay,
Costa Rica, and British Guiana (Demerara). It was described from
French Guiana.

2. Var. vulgaris J. Bequaert (1934). The most common form of the
species, with yellow markings on at least tergites 1 to 3, often 1 to 6.
I have seen it from British Guiana (Kartabo), Panama, Colombia,
Venezuela, Trinidad, Dutch Guiana, Brazil, Paraguay, northern Ar-
gentina, Peru, Bolivia, and Ecuador.

3. Var. myops (Fabricius, 1798). Only the second tergite is marked
with yellow spots. It was originally described from French Guiana.
I have seen it from Peru. It has been recorded also from Brazil and
(perhaps by error) from Trinidad and Paraguay.

4. Var. kaieteurensis J. Bequaert (1934). Abdomen without yellow

bequaert: social vespidae of the guianas 259

markings, but the apical segments somewhat orange. Thorax with
many yellow markings. Described from British Guiana: Kaieteur.

Polistes subsericeus H. de Saussure (1854)

I have seen a specimen from British Guiana: Mt. Roraima. The
species is also known from Dutch Guiana, Brazil and Paraguay.

Polistes erythrogaster Ducke (1905)

I have seen one specimen from British Guiana: Mt. Roraima. The
species is also known from the Amazon Basin in Brazil.

Polistes testaceicolor J. Bequaert (1937)

Synonym: Vcspa analis Fabrieius, 1798 (not of Fabricius, 1775).
"Apparently a common wasp in British Guiana: Bartica; George-
town; Warina, N. W. District; Tumatumari, Potaro River; Kaieteur;
Kartabo; Mt. Everard; Arakaka, I have also seen it from Dutch
Guiana, French Guiana, Brazil (Amazon Basin), Bolivia, Colombia,
Venezuela, Peru and Costa Rica.

This species is often heavily stylopized. One female bears seven
empty pupae of Strepsiptera, protruding from the hind margins of
second (1), third (2) and fourth (2) tergites, and of third (1) and
fourth (1) sternites. One female from Bartica is labelled as taken on
carrion of agouti.

Polistes ruficornis H. de Saussure (1853)

The typical form of this species is known only from Uruguay, Para-
guay and northern Argentina. I described the var. demeraraensis
3. Bequaert (1937) from British Guiana (Mahaica River, Demerara;
Georgetown; Blairmont), and I now also refer to this form the speci-
mens from Brazil and Bolivia which I had called var. biglumoides in
1937 (Arch. Inst. Biol. Veg., Rio de Janeiro, 3, p. 180).

The unrecognized Polistes guyanensis Cameron (1912), described
from Potaro River, British Guiana, was most probably a color form of
P. ruficornis and perhaps merely a variant of var. demeraraensis.

Polistes occipitalis Ducke (1904)

British Guiana: Mt. Roraima. Also known from French Guiana,
Dutch Guiana, Brazil, Bolivia, eastern Peru, and Colombia.

260 bulletin: museum of comparative zoology

Polistes melanosoma H. de Saussure (1853)

British Guiana: Bartica; Kartabo; west bank of Demerara River.
Also known from Brazil and Paraguay.

Polistes deceptor W. A. Schulz (1905)

British Guiana: Kartabo. Also known from Dutch Guiana, Brazil
and Peru.

Polistes canadensis (Linnaeus, 1758)

This, the most widely distributed of the American Polistes, extends
from south of the Great Lakes in the United States to northern Pata-
gonia, and occurs also in some of the Lesser Antilles, but not in the
Greater Antilles. In a recent revision (1943, in process of publication),
I recognize 19 color forms; but 2 only are known thus far from the
Guianas.

1. Typical canadensis. Fairly uniformly light russet to dark
mahogany-brown; head and thorax sometimes slightly lighter than
abdomen. No yellow markings, or sometimes a narrow apical margin
on the first tergite. Wings either uniformly purplish-black or dark
russet, or more or less russet toward the tips and darker basally. This
typical form occurs in British Guiana: Demerara; Georgetown; Mt.
Roraima. It is found also in French Guiana and Dutch Guiana and
ranges from southern Arizona to northern Argentina and Rio Grande
do Sul. It does not occur in Canada.

2. Var. infuscatus Lepeletier (1836). P. canadensis amazonicus
W. A. Schulz (1905) is a synonym. Like typical canadensis, but occiput
and outer orbits more or less extensively yellow. Wings either uni-
formly purplish-black or slightly to extensively russet apically and
darker at the base. First tergite with or without narrow apical yellow
fascia. This form is perhaps more common in British Guiana than
typical canadensis: Demerara; Onverwagt. I have also seen it from
Brazil (Amazon Basin), Dutch Guiana, French Guiana, Colombia,
Panama, Peru and Ecuador.

Polistes urceolata "Klug" Erichson (1848, in Schromburgk, Reisen
in Britisch Guiana, 3, p. 590; no sex; British Guiana) was evidently
based upon some form of P. canadensis; but the description is too brief
for recognition.

bequaert: social vespidae of the guianas 261

Subfamily POLYBIINAE

Mischocyttarus H. de Saussure (1853)

Mischocyttarus is the largest Neotropical genus of social wasps,
some 73 structural species and several color forms being described to
date (1943). This number will be more than doubled in the near
future, as both Mr. O. \V. Richards and Mr. J. F. Zikan are revising
the genus and intend to describe many new forms. Meanwhile it is
possible only to enumerate those known from British Guiana.

Mischocyttarus carbonarius H. de Saussure (1853)

Megacanthopus ruficornis Cameron (1912) is probably a synonym.
British Guiana: West bank of Demerara River; Kaieteur; Kartabo.
Also known from French Guiana, Brazil, Bolivia, Peru and Panama.

Mischocyttarus cerberus Ducke (1918)

The typical form is known from Dutch Guiana and Brazil. The var.
acheron Richards (1940) was described from British Guiana: Mazaruni
Settlement.

Mischocyttarus collarellus Richards (1940)

Common in British Guiana: Mazaruni Settlement; Cuyuni River, 3
miles from Kartabo; Potaro River, on trail between Tukeit and Kaie-
teur; Moraballi Creek, Essequibo River; Kartabo; Aremu, Bartica
District. Also known from Dutch Guiana, Brazil and Panama.

Mischocyttarus collaris Ducke (1904)

British Guiana: Berbice Savannas; Courantyne River; Demerara.
Also known from Brazil.

Mischocyttarus duckei R. du Buysson (1909)
British Guiana : Kartabo. Originally described from French Guiana.

Mischocyttarus flavicans Fabricius (1804)

Megacanthopus goeldii Ducke (1905) is a synonym.
British Guiana: Kartabo; Mazaruni River. Also known from
French Guiana, Dutch Guiana, Brazil, Bolivia, Peru and Ecuador.

262 bulletin: museum of comparative zoology

Mischocyttarus foveatus Richards (1940)

Described from British Guiana: First Falls of Essequibo River;
Issororo, Northwest District; Island near Monkey Jump; Moraballi
Creek, Essequibo River.

Mischocyttarus heliconius Richards (1941)

Described from British Guiana: Moraballi Creek, Essequibo River;
Potaro River, on trail between Tukeit and Kaieteur.

Mischocyttarus injucundus (H. de Saussure, 1854)

The typical form is known from Brazil and Colombia. The var.
bimarginatus (Cameron, 1912) was described from British Guiana,
without more definite locality. I have seen it from Demerara; Maza-
runi River; Blairmont; and Georgetown

Mischocyttarus labiatus (Fabricius, 1804)

Megacanthopus atriceps Cameron (1912) is a synonym.

British Guiana: Mt. Roraima; Rupununi River; Kaieteur; west
bank of Demerara River; Kartabo; Bartica. Also known from French
Guiana, Dutch Guiana, Trinidad, Brazil, Venezuela, Paraguay, Peru,
Colombia and Panama.

Mischocyttarus lecointei (Ducke, 1904)

British Guiana : Tukeit ; Kaieteur. Also known from French Guiana,
Dutch Guiana and Brazil.

Mischocyttarus lemoulti (R. du Buysson, 1909)

British Guiana: Moraballi Creek, Essequibo River. Originally de-
scribed from French Guiana.

Mischocyttarus metathoracicus (H. de Saussure, 1854)

British Guiana: Essequibo River; Oko River, a tributary of the
Cuyuni. Originally described from French Guiana; also known from
Brazil and Peru.

bequaert: social vespidae of the guianas 263

Mischocyttarus metoecus Richards (1940)

Described from British Guiana: Mazaruni Settlement. Also known
from Dutch Guiana and Brazil.

Mischocyttarus oecothrix Richards (1940)

Described from British Guiana: Kaieteur Savanna, Potaro Valley;
Canister Falls, Cattle Trail Survey; Moraballi Creek, Essequibo River.

Mischocyttarus prominulus Richards (1941)

Described from British Guiana: Kartabo; Mazaruni Settlement.
Also known from Bolivia.

Mischocyttarus rotundicollis (Cameron, 1912)

Originally described from British Guiana, without more definite
locality. I have seen it from Kalacoon and Penal Settlement, Bartica
District.

Mischocyttarus smithii H. de Saussure (1853)

Megacantkopus violaceipennis Cameron (1912) is a synonym.
British Guiana : Kartabo ; Essequibo River. Also known from French
Guiana and Brazil.

Mischocyttarus superus Richards (1940)

Described from British Guiana: Mazaruni Settlement; Bartica-
Potaro Road; Potaro River, on trail between Tukeit and Kaieteur;
I have also seen it from Kartabo.

Mischocyttarus surinamensis (H. de Saussure, 1854)

British Guiana: Kamakusa; Courantyne River; Trail between
Tukeit and Kaieteur; Demerara. Also known from Dutch Guiana and
Brazil.

Mischocyttarus synoecus Richards (1940)

Described from British Guiana: Amatuk, Potaro River; Mazaruni
Settlement. Also known from Brazil and Panama.

Polybia nigriceps Cameron (1912), described from British Guiana,
without more definite locality, was a species of Mischocyttarus. The

264 bulletin: museum of comparative zoology

name is preoccupied by Polybia fastidiosuscula var. nigriceps Zavattari
(1906).

Mr. O. W. Richards (in litt.) includes British Guiana in the range of
Mischocyttarus drewseni (de Saussure, 1857).

Mischoeyttarus socialis (H. de Saussure, 1854) (= Vespa atra Olivier,
1791; not of Gmelin, 1790) was taken in French Guiana, Cayenne
being the type locality of Olivier's V. atra.

Mischocyttarus alfkenii (Ducke, 1904) I have seen from Dutch
Guiana.

Pseudochartergus Ducke (1905)

I have revised this genus in a recent paper (1938, Rev. de Entomo"
logia, 9, pp. 103-105), where I recognized only two species. A study
of more extensive material in the collections of Cornell University has
led me to separate a third species, Pseudochartergus panamensis (Zavat-
tari, 1906) ( = Chartergus acutiscutis Cameron, 1907). This differs from
P. chartergoides in the relatively shorter clypeus of the female (about
as wide as long) and in the lateral ridges of the propodeum being more
rounded off. All localities from British Honduras, the Republic of Hon-
duras, Panama and Colombia, which I listed in 1938 under P. charter-
goides var. cinctellus, refer to P. panamensis.

Only one of the three species is known from the Guianas.

Pseudochartergus chartergoides (Gribodo, 1891)

I distinguish by name three color forms of this species, but only the
var. cinctellus (Fox, 1898) occurs in British Guiana: Kartabo. It is
known also from Dutch Guiana and Brazil.

Charter ginus pallidibalteatus Cameron (1912), described from
British Guiana, was most probably P. chartergoides var. cinctellus.

Charterginus Fox (1898)

This genus was revised in 1938 (Rev. de Entomologia, 9, pp. 99-
103). Only one of the four species is known from the Guianas.

Charterginus huberi Ducke (1904)

British Guiana: Source of Essequibo River; Waratuk. It was de-
scribed from the Amazon Basin of Brazil. I have seen it also from
Dutch Guiana and French Guiana (Maroni). Polybia fuhicauda

bequaert: social vespidae of the guianas 265

Cameron (1912), described from British Guiana, is a synonym of
C. huberi.

Protopolybia Ducke (1905)

This genus, containing some of the smallest South American wasps,
is well represented in the Guianas. In addition to the species included
in the key, three forms described by Cameron from British Guiana
are as yet unrecognized.

Key to Guiana Species

1. Concavity of propodeum narrow, groove-like. First abdominal

tergite slender, the basal third or more stalk-like. Clypeus about

as wide as high 2

Concavity of propodeum broad, either very shallow or deeply
bowl-shaped. First abdominal tergite thickset, not or very briefly
stalk-like at base 3

2. Thorax rather dull, with distinct, scattered, medium-sized punc-

tures; abdomen impunctate. Postscutellum nearly twice as wide
as greatest length, with triangular, bluntly pointed apical exten-
sion. (Extensively and fairly uniformly testaceous-yellow ; almost

bare) P. holoxantka

Body dull or slightly shiny, usually with a few discrete larger
punctures. Postscutellum at most one and one-half times as wide
as greatest length, with the apical extension rounded off, tongue-
like. (Black, with few or many yellowish markings)

P. minutissima

3. First abdominal tergite very broad and short, cap-shaped, passing

gradually into the base of the second, which is not constricted
from it. Outer orbit unusually widened in lower half; occipital
carina low and ending some distance from base of mandible.
Clypeus nearly twice as wide as high. Thorax thickset, about as
high as long, with many coarse punctures; humeral collar high,
shouldered at the sides; postscutellum nearly twice as wide as
long; sides of propodeum bulging, somewhat compressed, though

broadly rounded off P. emortualis

First abdominal tergite not cap-shaped, always set off from the
more swollen second. Thorax longer than high; sides of propo-
deum low, depressed, though broadly rounded off 4

4. Clypeus one and one-half times to nearly twice as wide as high,

smooth and shiny. Outer orbit about as wide as eye, not reced-

266 bulletin: museum of comparative zoology

ing; occipital carina high, extending to near base of mandible.
Concavity of propodeum deep; postscutellum about one and one-
half times as wide as long; humeral collar high, more or less
shouldered at the sides. Body very shiny, with distinct, scattered,
medium-sized to large punctures, with many long, erect hairs

P. picteti

Clypeus much less than one and one-half times as wide as high.
Outer orbit narrower than eye, receding. Postscutellum slightly
though distinctly shorter than wide. Body dull or nearly dull;

erect hairs short and sparse 5

5. Thorax about one and one-fourth times as long as high. Body dull,
fairly uniformly covered with small, discrete punctures. Clypeus
about one and one-third times as wide as high. (Mostly pale
yellow) P. amarella

Thorax nearly one and one-half times as long as high. Body not or

slightly shiny, with microscopic, alutaceous sculpture and a few

small, discrete punctures. Clypeus at most one and one-fourth

• times as wide as high. (Black, marked with yellow) . .P. pumila

Protopolybia picteti (H. de Saussure, 1854) is a widely distributed
species, of which I recognize by name 9 color forms. Only one of these
has been reported from the Guianas. P. picteti var. bella (R. von Iher-
ing, 1903) was originally described (as Polybia bella) from Dutch
Guiana. It is also known from Brazil, Peru and Bolivia. It is char-
acterized by the ground color of the body being black or slightly
brownish-black, with many yellow markings on thorax and abdomen
(usually no yellow stripes on mesonotum) and 3 pale spots on the base
of tergite 2, the median spot being transverse (wider than long).

Protopolybia holoxantha (Ducke, 1904)

This species is probably rather common in British Guiana: Kama-
kusa; Moraballi Creek, Essequibo River; Penal Settlement, Bartica
District. It is also known from French Guiana and Brazil (Oyapoc
and Amazonas).

Protopolybia minutissima (Spinola, 1851)

I recognize four color forms of this species in the Guianas.

1. Pale markings few, whitish or very pale yellow; base of tergite 2
without cross-band or lateral spots 2

bequaert: social vespidae of the guianas 267

Pale markings more extensive, pale to bright yellow; base of tergite
2 with a cross-band or a pair of spots 3

2. Thorax not or very little marked with pale yellow or whitish; no

spots on propodeum typical minutissima

Thorax more profusely marked with pale yellow or whitish; a pair
of spots on propodeum var. bitwminata

3. Body profusely marked with yellow; base of tergite 2 with a con-

tinuous and usually broad cross-band var. sedula

Body less extensively marked with yellow; base of tergite 2 with a
pair of streaks or spots var. exigua

The typical form occurs in British Guiana (Kartabo), as well as in
Peru and Brazil.

The var. binominata (YV. A. Schulz, 1906) appears to be the most
common form, being known from British Guiana (Kartabo; George-
town; Demerara), Colombia, Panama, Ecuador, Peru, and Bolivia.
Polybia minutissima H. de Saussure (1854) was this form, not Spinola's
typical minutissima.

The var. exigua (H. de Saussure, 1854) is fairly common in Brazil
and has been reported from French Guiana, Trinidad, and eastern
Peru (Iquitos).

The var. sedula (H. de Saussure, 1854) is widespread and has been
taken in Dutch Guiana, French Guiana, Brazil, Venezuela, Trinidad,
Colombia, Panama (Darien), Peru, Bolivia, and Paraguay. I regard
Polybia diligens F. Smith (1857) as a synonym of sedula.

Protopolybia amarella, new species

Fig. 1

Female. Head flattened, slightly wider than thorax; seen in front,
subcircular, scarcely wider than high; from above, rectangular with
receding hind corners, about two and one-half times as wide as long;
occipital margin with a broad and shallow inward curve. Vertex and
genae margined by a fine carina, which ends a short distance from the
base of the mandible ; gena slightly narrower than eye in profile. Oculo-
malar space lacking. Inner orbits about one and one-half times as far
apart on vertex as at clypeus. Ocelli fairly large, in a nearly equilateral
triangle ; posterior ocelli slightly farther from eyes than from each other.
Interantennal shield broad, not set off, with a longitudinal groove over
upper half; antennae somewhat farther apart than from eyes. Frons
slightly convex. Clypeus at its narrowest about one and one-third

268

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times as wide as high, irregularly heptagonal, contiguous to the eyes
over about one-half of sides; apical margins moderately produced,
ending in a blunt, rounded point. Mandible with an oblique cutting
edge of four sharp, subequal teeth. Antenna: scape long, slender,
slightly curved; second segment unusually long, only slightly shorter
than third, scarcely swollen; remainder of flagellum slightly club-

Fig. 1. Protopolybia amarella J. Bequaert. Female. A, body in profile; B,
first and second tergites from above; C, postscutellum and propodeum from
behind; D, head in front view.

shaped. Thorax rather thickset; in profile, about one and one-fourth
times as long as high; from above, nearly one and two-thirds times as
long as wide before tegulae. Pronotum nearly straightly truncate
anteriorly; humeral margin with a distinct obtuse ridge which stops at
the sides, forming slight, blunt humeral angles; dorso-lateral areas
sloping; lower anterior margin of ventro-lateral areas with a swelling
indented by a deep pronotal fovea. Mesopleuron much swollen, with-
out mesepisternal suture; deep mesepimeral suture ending abruptly

bequaert: social vespidae of the guianas 269

and far from metapleuron. Upper sclerite of metapleuron ending
nearly square below, not extended along meso-metapleural suture.
Scutellum slightly and evenly convex, with a very weak median
impressed line. Postscutellum triangular, scarcely swollen, its median
area about one and one-third times as wide as long, uniformly sloping
or nearly vertical throughout, the lower extension very long, with a
free, raised, tongue-like apex. Propodeum little swollen, nearly vertical
in profile; median concavity very broad and shallow. First abdominal
tergite short and cup-shaped, with a very short, broad stalk, which is
not wing-like at the sides; seen from above, gradually widened, slightly
shorter than wide at apex, about half as wide at apex as second seg-
ment; in profile, very gradually and moderately convex. Second
tergite from above wider than long, gradually widened at base ; in pro-
file, moderately and evenly convex. Apical margins of all segments
very slightly thickened. Mid tibiae with two spurs. Claws symmetri-
cal, unarmed. Venation: second cubital cell much higher than wide;
third cubital long, about one and one-third times as long on cubitus
as on radius ; radial cell very long, acute ; basal vein ending in subcosta
Hose to the large stigma.

Dull; fairly uniformly covered with small, discrete punctures, more
weakly on frons; clypeus and mandibles almost impunctate. Pubes-
cence sparse, short, erect, somewhat longer on propodeum and abdo-
men. Eyes bare.

Pale yellow, with ferruginous blotches on antennae, tip of mandible,
tibiae, tarsi and under side of abdomen. A few brownish-black spots
as follows : a streak behind each posterior ocellus ; a transverse row of
four spots on the prehumeral depressed slope of pronotum; narrow
sutures of mesonotum, scutellum and postscutellum; a median line
over anterior half, a line on each side over posterior half, and a pair of
elongate spots on disk of mesonotum. Disk of second tergite with a
median blackish triangle showing through the yellow integument.
Wings hyaline, with a yellowish tinge; veins and stigma pale yellowish-
testaceous.

Length (h.+th.+t. 1+2): 5.1 mm.; of fore wing, 5 mm.
British Guiana: Source of Essequibo River, female holotype (J.
Ogilvie). — Colombia: La Chorrera, Rio Igara-Parana, Intendencia
Amazonas, female paratype (J. C. Bradley). — Holotype at Mus.
Comp. Zool., Cambridge, Mass.; paratype at Dept. Ent., Cornell
University.

The almost completely yellow body makes this a striking insect.

270 bulletin: museum of comparative zoology

Protopolybia emortualis (H. de Saussure, 1855)

This species occurs in two color forms.

1. Typical emortualis is more or less extensively ferruginous, partic-
ularly on the abdomen, and has pale yellowish markings. It was de-
scribed from Brazil and Ducke reported it from Dutch Guiana.

2. The var. duckei (R. du Buysson, 1905) ( = Charter -ginus duckei
R. du Buysson) differs only in the ground color of the body, including
the abdomen, being black. It was originally described from Brazil
(Bahia) ; but Ducke reports it from eastern Peru (Iquitos) and I have
seen it from British Guiana (Upper Essequibo River). It is doubtful
whether this form is worth separating from typical emortualis.

Protopolybia pumila (H. de Saussure, 1863)

Two forms of this species are known in the.Guianas.

1. Typical pumila is extensively marked with yellow, the meso-
notum often bearing yellow stripes and the base of tergite 2 a pair of
unconnected spots. I have seen it from British Guiana: Kartabo;
source of Essequibo River. It is also known from Dutch Guiana,
Brazil, Venezuela, Colombia, Peru, and Bolivia.

2. The var. rotimdata Ducke (1910) was described (as a species)
from French Guiana and is not known from elsewhere. It has few pale
markings, no stripes on mesonotum nor spots at the base of tergite 2.

Protopolybia rufo-ornata (Cameron, 1912)

This was described, as Charter ginus rufo-ornatus, from British
Guiana, without more definite locality. It is at present unrecognized.
No doubt it was a Protopolybia, possibly a color form of P. picteti.

Protopolybia sulciscutis (Cameron, 1912)

This also was described merely from "British Guiana," as Polybia
sulciscutis, and is unrecognized. The statement in the description,
"apex of postscutellum narrowed to a broad, bluntly rounded point,"
seems to place it in Protopolybia.

Protopolybia nana (Cameron, 1912)

This is another unrecognized species described by Cameron, as
Polybia nana, from "British Guiana." The name conflicts with the

bequaert: social vespidae of the guianas 271

earlier Polybia nana H. de Saussure (1803); hut, as Cameron's wasp is
probably identical with some other described Protopolybia, it would be
premature to rename it.

Brachygastra Perty (1833)

This genus has been generally called Ncctarina Swainson and
Shuckard (1840) ; but there seems to be no reason why the earlier name
Brachygastra should not be used.

Key to Guiana Species

1. Sides of propodeum compressed but not angular, forming evenly

rounded curves in profile. Head (except clypeus), thorax and
second abdominal tergite with deep, large punctures. Dorsal
(horizontal) area of scutellum four times as wide as long in the

middle B. Scutellaria

Sides of propodeum compressed into prominent, broad, blunt angles.
Second abdominal tergite with small punctures only 2

2. Dorsal (horizontal) area of scutellum nearly flat, less *han three

times as wide as long in the middle, with the hind margin rather
sharp but slightly or scarcely curved inward. Punctures of meson-
otum fairly numerous and not or scarcely stronger than those of
the base of second tergite. Clypeus about twice as wide as

high B. lecheguana

Dorsal (horizontal) area of scutellum convex, three to four times as
wide as long in the middle, with the hind margin blunt but deeply
curved inward (scutellum forming two broad lobes). Punctures
of mesonotum decidedly stronger than those of second tergite . . 3

3. Body with many erect, short hairs. Punctures of frons and meso-

notum more numerous and fairly close. Lateral angles of pro-
podeum higher. (Usually with many yellow markings)

B. bilineolata

Body with very few erect hairs. Punctures of frons and mesonotum

far apart. Lateral angles of propodeum lower. (With few yellow

markings, the thorax and head almost entirely black)

B. augusti

Brachygastra augusti (de Saussure, 1854)

The typical form of the species has distinct yellow apical bands on
tergites 1 to 6 and sternites 2 to 6 of the abdomen. It is known from

272 bulletin: museum of comparative zoology

British Guiana (source of Essequibo River), Dutch Guiana, French
Guiana, Brazil, Paraguay, Bolivia, Peru, Colombia, and Venezuela.

Brachygastra lecheguana (Latreille, 1824)

This widely distributed and common wasp, known from southern
Arizona and southern Texas to northern Argentina, was discussed in
a paper published in Entomologica Americana in 1932 (13, pp. 92-
112). The specimens from British Guiana (Arakaka), Dutch Guiana
and French Guiana are of the var. velutina Spinola (1841), which has
the thorax densely covered with silky, golden-yellow hairs.

Brachygastra scutellaris (Fabricius, 1804)

The several color forms of this species are revised in Jr. New York
Ent. Soc, 50, (1942) 1943, pp. 306-308, where I recognize six of them
by name. The three known from the Guianas may be separated as
follows :

1. Scutellutn black; postscutellum either black or with an anterior

pale cross-band. Abdomen with whitish apical margins on most

sternites and some of the tergites var. myersi

Scutellum and postscutellum mostly or entirely yellow 2

2. Second abdominal tergite mostly orange-yellow or ferruginous.

Thorax with many yellow markings ; mesonotum often with spots

or stripes var. rufiventris

Second abdominal tergite with only an apical yellow margin. Yel-
low of pronotum either lacking or restricted to the narrow, often
incomplete humeral margin typical scutellaris

Typical scutellaris is the most common form in British Guiana:
Forest Settlement, Mazaruni River; Kartabo; junction of Mazaruni
and Essequibo Rivers. Ducke lists it from Bartica, and Bodkin from
Issororo, N. W. District. It is also known from French Guiana, Brazil,
Bolivia, eastern Peru, Colombia, and Panama. Brachygastra scutellata
Spinola (1851) is a synonym of the typical form.

The var. rufiventris (de Saussure, 1854) is reported from French
Guiana, Brazil and Colombia.

The var. myersi J. Bequaert (1943) was described from British
Guiana (Mt. Roraima) and Bolivia.

bequaert: social vespidae of the guianas 273

Brachygastra bilineolata (Spinola, 1841)

Recently (1943, Jr. New York Ent. Soc, 50, for 1942, pp. 303-306)
I recognized five color forms of this species, those occurring in the
Guianas being separated in the following key:

1. Second tergite with only the apical margin yellow. Mesonotum

with short yellow longitudinal stripes or a pair of yellow spots
posteriorly, or entirely black. Wings with a honey-yellow tinge,

most of the veins testaceous typical bilineolata

Second tergite with a broad or narrow discal yellow band in addi-
tion to the apical fascia, or mostly yellow. Mesonotum always
with two yellow stripes 2

2. Second tergite mostly yellow, except for the black base and a more

or less defined transverse black discal blotch or irregular line.

Wings with a honey-yellow tinge var. Surinam ensis

Second tergite with two separate transverse yellow stripes of about
equal width, one apical, the other discal 3

3. Wings extensively tinged with honey-yellow. Veins and stigma

mostly pale var. antillarum

Wings nearly hyaline, slightly grayish, not suffused with yellow.
Veins and stigma blackish var. smithii

Typical bilineolata is known from British Guiana (Demerara;
Georgetown), French Guiana, Dutch Guiana, Venezuela, Brazil,
Colombia, and Costa Rica. I regard Nectarinia mobiana de Saussure
(1867) as a synonym of typical B. bilineolata.

The var. antillarum (Provancher, 1883) occurs in British Guiana:
Rapununi River. I have seen it also from Dutch Guiana, Trinidad,
Brazil, eastern Peru, and the Republic of Honduras.

The var. smithii (de Saussure, 1854) has been found in French
Guiana, Dutch Guiana, Brazil, and Colombia.

The var. surinamensis J. Bequaert (1943) is known only from Dutch
Guiana.

Chartergus Lepeletier (1836)

As stated under Parachartergus, the type of this genus was desig-
nated by Emile Blanchard (1840) as Vespa nidulans Fabricius (1793),
a synonym of Vespa chartaria Olivier (1791). I revised the genus in
1938 (Rev. de Entomologia, 9, pp, 113-115) under the erroneous name
Epipona. I recognize two species, of which only one occurs in the
Guianas.

274 bulletin: museum of comparative zoology

Chartergus chartarius (Olivier, 1791)

I have seen this wasp from British Guiana: Demerara; Georgetown.
It occurs also in French Guiana, Dutch Guiana, Brazil, Paraguay,
Bolivia, and Peru (Iquitos).

Synoeca H. de Saussure (1852)

Ducke (1910) recognizes only two species of Synoeca, but I believe
three may be separated on structural characters, as du Buysson did
in his Monograph (1906). Moreover, Polistes virginea Fabricius
appears to be the valid name for the species usually called S. irina
(Spinola).

1. Larger; fore wing 16 to 18 mm. long. Eyes bare. Clypeus smooth

and shiny. Propodeum either impunctate or with a few minute
punctures. Oculo-malar space of worker and queen fully as long
as sixth antennal segment. Mostly or wholly bluish-black; wings

violaceous-black S. surinama

Smaller; fore wing 13 to 15 mm. long. Eyes with a few short hairs.
Clypeus rather dull and with scattered punctures. Propodeum
densely punctate. Oculo-malar space of worker and queen
shorter than sixth antennal segment. Either bluish-black or
partly ferruginous. Wings mostly yellowish-subhyaline 2

2. Thorax fairly uniformly punctate, the punctures of mesopleura and

mesonotum scarcely smaller than those of propodeum. Scutellum
convex, with very slight median impressed line. Swollen portion
of first abdominal segment elongate-triangular seen from above.
Black costal border sharply set off from the remainder of the

wings S. chalybea

Punctures of mesopleura and mesonotum very fine, much weaker
than those of propodeum. Scutellum gibbose, with a deep median
saddle. Swollen portion of first abdominal segment trapezoidal

seen from above. Wings fairly uniformly yellowish-russet

S. virginea

Synoeca surinama (Linnaeus, 1767)

A common wasp throughout Central and South America, from
southern Mexico to southern Brazil. The size of the head varies con-
siderably in this species and seems to be correlated with the total size
of the specimen. Perhaps these differences set off the fertile females
(queens) from the workers. Several other insects are homoeochromic

bequaert: social vespidae of the guianas 275

with S. surinama. The most interesting of these is the mantispid,
Climaciella chalybea Erichson, of which I have seen a specimen from
the Upper Rio Huallaga, Peru. The collector had evidently mistaken
it for the wasp. Possibly the mantispid lives in the nest of Synoeca.

1. Typical form. Nearly entirely bluish-black, at most with russet
blotches on the mandibles. This is Vespa .surinama Linnaeus (1767),
described from Surinam, where it is common. Vespa nigricornis
Olivier (1791), Polistcs cacrulea Fabricius (1804) and Synoeca ultra-
marina H. de Saussure (1852) are synonyms.

Common in British Guiana: Arakaka; Penal Settlement, Bartica;
Rockstone, Essequibo River; -Kartabo; Georgetown; Kamakusa;
Berbice Savannas; Blairmont. I have also seen it from Costa Rica,
Colombia, Venezuela, Trinidad, French Guiana, Dutch Guiana, Brazil,
Ecuador, Peru, and Bolivia.

2. Var. cyanea (Fabricius). The lower part of the head (particularly
the clypeus) is more or less ferruginous-red; second abdominal tergite
occasionally blotched with russet. Specimens transitional to typical
surinama are frequent. This is Vespa cyanea Fabricius (1775), Synoeca
azurea H. de Saussure (1S52) and Synoeca violacca H. de Saussure
(1852).

This form is more widely distributed than typical surinama and is
known from French Guiana. I have seen it also from Mexico (Apat-
zingan, State of Michoacan), the Republic of Honduras, Guatemala,
Costa Rica, Panama, Colombia, Brazil, Paraguay, Ecuador, and
Venezuela.

Synoeca virgixea (Fabricius, 1804)

This species is rarer and not as widely distributed as S. surinama,
being apparently restricted to the northern half of South America.
It is almost entirely ferruginous or russet with very slight or no bluish
sheen. Fabricius' description of Polistcs virginea (1804) fits this wasp
exactly. The statement "Antennae nigrae articulo secundo [ = scape]
longiore, testaceo", agrees with the Synoeca, not with Apoica pallida.
Moreover, W. A. Schulz (1912, Berlin Ent. Zeitschr., 57, p. 84) recog-
nized Synoeca irina in Fabricius' type. Polistcs irina Spinola (1851)
and Synoeca testacea H. de Saussure (1854) are synonyms.

British Guiana: Kartabo; Junction of Mazaruni and Essequibo
Rivers; Penal Settlement, Bartica District; Source of Essequibo River.
I have also seen it from Dutch Guiana, Brazil, Ecuador, and Peru.

276 bulletin: museum of comparative zoology

Synoeca chalybea H. de Saussure (1852)

1. The typical form of this species, as figured by de Saussure (1854),
is nearly entirely bluish-black, with or without violaceous reflections ;
mandibles, clypeus, oculo-malar spaces and spots on mesopleura and
legs are yellowish-russet. The specific name was originally spelled
chalibea, but later (1854) corrected to chalybea by the author himself.

British Guiana: Shudihar River. I have also seen it from Brazil and
Peru and it was originally described from French Guiana.

2. S. chalybea var. splendens R. du Buysson (1906) was originally
described as a form of S. irina; but all specimens I have seen have the
puncturation of the thorax and the shape of the first abdominal seg-
ment as in S. chalybea. It is more or less extensively ferruginous or
yellowish-russet. I have seen it from Bolivia and Peru. As Dr. Wey-
rauch collected both typical chalybea and var. splendens apparently
from the same nest at Satipo, Peru, it is doubtful whether the name
splendens deserves to be retained.

Metapolybia Ducke (1905)

I have reached the conclusion that there are two structural species
in this genus, not one as Ducke claimed. They may be separated as
follows :

1. First abdominal segment slightly widened apically seen from
above and, in profile, very gradually thickened over posterior

half. Ground color of abdomen reddish-brown M . suffusa

First abdominal segment distinctly widened apically seen from
above and, in profile, rather abruptly thickened over posterior
third. Ground color of abdomen black M. clngulata

M. suffusa (Fox, 1899) has not yet been taken in the Guianas.
According to Fox' types from Corumba, Brazil, this is the wasp with
the abdomen extensively russet which Ducke (1910) mentions from
Bolivia and calls Metapolybia pediculata var. rufopicta. I have also
seen it from Peru, Bolivia and Colombia. At Restrepo (Int. Meta),
Colombia, I found M. suffusa and clngulata nesting side by side, but in
distinct colonies. They were not in the least aggressive, although the
sting was painful.

Metapolybia cingulata (Fabricius, 1804)

Schulz (1912) saw the types of Eumenes cingulata Fabricius (1804)
at Copenhagen and recognized in them Polybia pediculata H. de

bequaert: social vespidae of the guianas 277

Saussure (1854). The extent of pale yellow markings varies, Fabricius'
cingulata being based upon specimens with only the first and second
tergites margined with yellow. This is also true of Gribodo's Tatua
decorata (1896); while Polybia pediculata var. unilineata R. v. Ihering
(1904) refers to specimens with only traces of yellow markings. There
is no name available, and none seems needed, for specimens with most
or all tergites margined.

British Guiana: Kartabo; Bartica. Also known from Venezuela,
Trinidad, Dutch Guiana, French Guiana, Brazil, Paraguay, northern
Argentina, Bolivia, Peru, Colombia, Panama, Costa Rica, Guate-
mala, and Mexico (Cordoba, Vera Cruz; Tierra Colorada, Guerrero).

Epipona Latreille (1802)

Emile Blanchard in 1840 (Hist. Nat. Ins., 3, Orth. Nevr. Hem. Hym.
Lep. Dipt., p. 394) designated as type of Epipona, Vespa morio
Fabricius (1798), a synonym of Vespa tatua Cuvier (1797). This gen-
eric name must therefore be used instead of Tatua de Saussure (1854) ;
while the genus I called Epipona in 1938 should be known as Charter-
gus Lepeletier. Epipona was revised in 1938 (Rev. de Ehtomologia, 9,
pp. 115-117), when I recognized two species, only one of which occurs
in the Guianas.

Epipona tatua (Cuvier, 1797) has not yet been taken in British
Guiana; but it is known from French Guiana, Trinidad, Venezuela,
Brazil, Peru, Colombia, Panama, and Costa Rica.

Polybia Lepeletier (1836)
Key to Guiana Species

1. Propodeum with more or less distinct transverse striation extend"

ing even over the convex sides. Occipital margin of head with a

deep inward curve. Body 14 to 16.5 mm. long 2

Propodeum without or with obsolete striation; rarely a few irregu-
lar striae in the concavity or on the sides 4

2. Head very strongly swollen behind the eyes; outer orbit slightly

wider than eye in profile. Clypeus broader than high, the apex
evenly and broadly rounded off. Pronotum on the sides with a
strong but blunt humeral crest. Striation of propodeum fine,

but regular P. ujhelyii

Head moderately swollen behind the eyes; outer orbit narrower
than eye in profile. Clypeus about as high as wide, with bluntly
pointed apex. Pronotum somewhat shouldered, but not ridged

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at the humeral angles. Striation of propodeum very irregular
or partly effaced 3

3. Concavity and sides of propodeum with coarse and irregular

wrinkles and punctures. First abdominal tergite slightly swol-
len; stalk-like base passing gradually into swollen apical por-
tion (in profile view) P- striata

Striation and punctures of propodeum weak or obsolete. First
abdominal tergite strongly swollen, abruptly raised behind the
stalk-like base (in profile view) P. liliacea

4. Median concavity of propodeum distinct and more or less circular,

never longer than wide 5

Median concavity of propodeum either almost lacking or groove-
like and longer than wide 7

5. Head distinctly swollen; outer orbit fully as wide as eye in profile.

Puncturation of mesopleura fine but distinct. Black, with
scutellum and postscutellum entirely yellow. Body 12 to 14

mm. long P • jurinei

Head not swollen; outer orbit narrower than eye in profile. Meso-
pleura nearly impunctate. Smaller; body 8 to 10 mm. long.. .6

6. Pronotum with very weak, blunt humeral collar; hind margin

forming an oval curve. First abdominal segment slender;

swollen portion longer than wide at apex P. signata

Pronotum with distinct, sharp humeral collar; hind margin nearly
semi-circular. First abdominal segment shorter; swollen portion
about as long as wide at apex P- bifasciata

7. At least sides of thorax covered with dense, silky pubescence,

often with golden sheen. Yellow markings lacking or much

reduced. Body 14 to 19 mm. long 8

Thorax nowhere covered with dense, silky pubescence 11

8. Eyes with distinct short hairs. Oculo-malar space very long (as

long as fifth antenna! segment). Pronotum without humeral
collar. Black, with thorax more or less extensively and first or
first and second abdominal segments reddish-brown. Body 15

to 17 mm. long P- sericea

Eyes bare. Oculo-malar space very short or lacking 9

9. Dorsum of thorax densely covered with long golden-silky pubes-

cence. Mesopleura finely but distinctly punctate. Pronotum
without humeral collar. Mostly brownish-red to fuscous;
abdomen without paler apical margins; head black; wings more
or less ferruginous, particularly toward costal margin. Clypeus
about as long as wide. Body 15 to 17 mm. long. . P. chrysothorax

bequaert: social vespidae of the guianas 279

Dorsum of thorax only sparsely pubescent, not golden-silky.
Color different 10

10. Only pleura of thorax distinctly silky; the abdomen not. Wings

subhyaline or slightly brownish-yellow, with ferruginous costal

margin. Body 15 to 16 mm. long P. affinis

Entire body, including abdomen, silky or velvety. Either russet
to brownish-russet with testaceous apical abdominal margins,
or (var. velutiiia) almost entirely blackish without apical mar-
gins. Wings strongly tinged with honey-yellow. Thorax very
finely punctate. Pronotum without humeral collar. Clypeus
slightly wider than long. Body 14 to 18 mm. long. . . P. micans

11. Large species, 18 to 21 mm. in total length. Head and thorax

black; abdomen red; wings nearly hyaline. Thorax shiny, with
scattered punctures and long, erect hairs. Oculo-malar space

long. Clypeus much wider than long P. dimidiata

Smaller. Thorax not shiny, or the other characters mentioned
above not all present 12

12. First tergite distinctly punctate. Mesopleura and propodeum

with large punctures. Outer orbits somewhat swollen in upper
half. Clypeus slightly wider than high. Eyes with distinct short
hairs. Body black, somewhat velvety, 12 to 14 mm. long; wings

infuscate, clearer posteriorly or apically 13

First tergite usually impunctate; if distinctly punctate, the other
characters mentioned above not all present 14

13. Oculo-malar space at its shortest over half the length of the

tenth antennal segment. Concavity of propodeum narrow,
groove-like, but very weak or barely indicated in upper third

P. tincti'pennis
Oculo-malar space at its shortest less than half the length of the
tenth antennal segment. Concavity of propodeum well-devel-
oped over entire length, broader and more shallow. . P. rufitarsis

14. Oculo-malar space distinct, at least half the length of the tenth

antennal segment 15

Oculo-malar space very short or almost lacking. Clypeus always
much narrower than vertex. Outer orbit usually narrower than
eye in profile 17

15. Black, with partly black wings. Eyes hairy. A short but distinct

carina between occiput and middle part of outer orbit (not
behind vortex). A fine, sharp humeral collar. Clypeus about as

high as wide P. ignobilis

Russet, infuscate or blackish, with many pale (yellowish) mark-

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ings, particularly on the thorax (mesonotum striped). Wings
slightly tinged with yellow. No carina between occiput and
outer orbit or vertex. No humeral collar. Eyes bare. Clypeus
much wider than high. Outer orbit slightly wider than eye in
profile 16

16. Larger species; fore wing 11 to 11.5 mm. long. Oculo-malar space

about as long as tenth antennal segment, longer than the stretch
of the inner orbit contiguous to the clypeus. Clypeus somewhat

narrower P. singularis

Smaller species; fore wing 8.5 to 9.5 mm. long. Oculo-malar space
about half the length of the tenth antennal segment, shorter
than the stretch of the inner orbit contiguous to the clypeus.
Clypeus very wide ' P. emaciata

17. Large species (15 to 18 mm. long); black with extensive pale

markings; mesonotum always with two longitudinal pale stripes.
• Upper outer orbit separated from occiput by a strong carina,

which however does not continue behind vertex. Propodeum

with at least traces of striation. (P. striata and P. liliacea:

see couplet 3)
If pale longitudinal stripes are present on mesonotum, the species

is much smaller. Propodeum without traces of striae 18

18. Thorax shiny, with many distinct punctures. A distinct but short

oculo-malar space. Eyes with scattered hairs. Longitudinal
groove of propodeum deep. Russet, with yellowish markings
and infuscate areas. Wings nearly hyaline, very slightly

yellowish P. gorytoides

Thorax practically impunctate 19

19. Humeral margin of pronotum barely curved, raised on the sides

into a sharp, low collar which projects as fairly distinct rounded
angles. Groove of propodeum weak or obsolete ; propodeum with
erect hairs. No carina between occiput and vertex. Body 12

to 13 mm. long P. rejecta

Humeral margin of pronotum distinctly curved, either not raised
or with a low collar which never forms angles at the sides .... 20

20. Sides of pronotum with a distinct though low humeral collar.

Upper outer orbits and vertex not separated by a carina from
occiput. Propodeum distinctly grooved, with erect hairs. Body

9 to 12 mm. long 21

Humeral collar barely indicated or absent. In doubtful cases, with
a fine carina between occiput and vertex 22

21. Thorax slender, flattened dorsally, particularly on posterior por-

bequaert: social vespidae of the guianas 281

tion of mesonotum and on scutellum. Scutellum not grooved.

Eyes with fine, scattered hairs P. procellosa

Thorax more thickset, not flattened dorsally; scutellum convex,

with a longitudinal groove. Eyes bare P. catillifex

22. Upper outer orbits and vertex separated from occiput by a very

fine carina, somewhat interrupted medially. Body 8 to 11 mm.

long P. occidentalis

Upper outer orbits and vertex not separated from occiput by a

carina. Body 7 to 8.5 mm. long P. bistriata

Polybia lugubris H. de Saussure (1854) was originally described as
from Guiana, but apparently by error. The species has only been taken
definitely in southern Brazil and is unlikely to occur in the Guianas.
It is not included in the foregoing key.

Polybia ujhelyii Ducke (1909) is known from the Lower Amazon
(Brazil) and French Guiana. I have seen it from Bolivia.

Polybia signata Ducke (1910) is known from French Guiana, Brazil
and Panama.

Polybia a finis R. du Buysson (1908) was described from French
Guiana. Ducke records it also from Brazil and Ecuador. I do not
know this species.

Polybia emaciata Lucas (1879). This is in my opinion the oldest
valid, recognizable name for the well-known South American wasp
which builds a nest of clay with a circular entrance. It is generally
called Polybia fasciata H. de Saussure (1854), which was, however, a
misidentification of Polybia fasciata Lepeletier (1836). Lepeletier's
fasciata was certainly not the Polybia with the clay nest, the size given
being much too large (7 French lines = 15 mm.). I regard P. fasciata
Lepeletier, Vespa fasciata Olivier (1791) and Vespa fulvo-fasciata
Degeer (1773) as one and the same species of Stelopolybia. Polybia
caementaria Ducke (1904) is a synonym of P. emaciata. The species is
known from Dutch Guiana, French Guiana, Venezuela, Colombia,
Brazil, Peru, Bolivia, Panama, Costa Rica, Guatemala and the Repub-
lic of Honduras.

Polybia gorytoides Fox (1898)

British Guiana: west bank of the Demerara River; Bartica. Also
known from Brazil (Oyapoc; Amazon Basin; Chapada), Dutch Guiana
(Kwakoegron, Saramacca River), Bolivia (Buena vista, Dept. Sta.
Cruz) and Colombia (Villavicencio, Int. del Meta).

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Polybia liliacea (Fabricius, 1804)

P. liliacea and P. striata (Fabricius) (Syn. : P. syncophanta Gribodo)
are closely allied species, with the same type of coloration, namely
black, with extensive pale markings on thorax and abdomen and par-
ticularly with two broad longitudinal pale stripes on mesonotum.
These two species are structurally distinct and, as Dr. C. Geijskes
writes me, differ in the nesting habits.

In P. liliacea the striation and puncturation of the propodeum are
weak or obsolete, the concavity showing only faint traces of striae.
The postscutellum is rather strongly swollen. The first abdominal
tergite is short and much swollen, the swelling being rather abrupt in
profile, the basal stalk-like portion well-defined and at most one-third
of the total length of the tergite. Usually the color markings are very
pale yellow or ivory-yellow, the longitudinal stripes of the mesonotum
often coalescent before the scutellum; all or most of the abdominal
segments bear apical bands. This type of coloration was correctly
described by Fabricius as follows: "Caput cum antennis nigrum.
Thorax ater limbo lineolisque duabus crassis, antice abbreviatis,
postice coeuntibus, scutello lineolisque duabus sub scutello flavis.
Abdomen nigrum segmentorum marginibus flavis. Pedes nigri."

According to Dr. Geijskes, the nest of P. liliacea is very long (1.5
m.) and sausage-shaped, truncate at the lower end. It was partly
described by de Saussure (1858, Et. Fam. Vesp., 2, pp. cx-cxi) and
H. Lucas (1867).

British Guiana: Warina; Oronoque River; Georgetown; Berbice
Savannahs; west bank of Demerara River; Kartabo; Bartica; source
of Essequibo River. It is known also from French Guiana, Dutch
Guiana, Brazil, Venezuela, Colombia, Peru, Bolivia, and Ecuador.

Polybia striata (Fabricius, 1787)

Polybia sycophania Gribodo (1891) I regard as a synonym of
Fabricius' Vespa striata. Those who, like Ducke, refuse to recognize
this wasp in Fabricius' description, will have to use Gribodo's name.

In P. striata, the puncturation of the propodeum is coarse and the
striation is distinct though irregular and particularly well developed in
the concavity. The postscutellum is moderately swollen. The first
abdominal tergite is elongate and gradually widened, being slightly
swollen in profile, with the basal stalk at least one-third of the total
length and poorly defined. Usually the color markings are darker

bequaert: social vespidae of the gutanas 283

yellow, often with an orange tinge and the stripes of the mesonotum
may be free throughout. The apical fascial' of the abdomen are either
as in liliacea, or much reduced, sometimes almost lacking. Fabricius
evidently described one of these very dark specimens: "Caput,
abdomen, pedes nigra, immaculata. Thorax niger margine antico
tenuissime, lineis duabus dorsalibus, lateribus baseos obliquis, scutello
maculisque sub scutello flavis. Alae albae costa fusca." It should be
noted that some specimens with the structural characters of striata
are colored exactly like liliacea, so that the color is not a reliable char-
acter. On the other hand, I have seen no specimens of liliacea either
with the abdomen almost black or with orange-yellow markings.

The species is as common and as widely distributed as P. liliacea.
I have seen it from British Guiana : Kartabo ;Warina. Also from Dutch
Guiana, Trinidad, Brazil, Colombia, Bolivia and Peru. It was first
described from French Guiana.

According to Dr. Geijskes, the nest of P. striata is a white spherical
structure hung up high in a tree. It is of interest that the Indians in
the interior of Dutch Guiana distinguish P. striata and P. liliacea by
name. The nest of P. striata has not yet been described.

Ducke (1910) includes P. striata and P. liliacea in his key among the
species without striation on the propodeum. As the striation is fairly
distinct, at any rate in P. striata. I have inserted these species twice in
my key.

Polybia jurixei H. de Saussure (1854)

A widely distributed species, which I have seen from British Guiana :
Kartabo. It is also known from French Guiana, Dutch Guiana, Brazil.
Bolivia, Peru, Ecuador and Colombia (Restrepo, Int. Meta)

Polybia bifasciata H. de Saussure (1854)

This species, which extends from Mexico to southern Brazil, is
rather variable in color. In a recent paper (1943, Jl. New York Ent.
Soc, 50, for 1942, pp. 300-303), I recognize seven forms by name.
Only two of these have been taken in the Guianas thus far. Typical
P. bifasciata I have seen from British Guiana (Kartabo; Demerara
River), as well as from eastern Peru. The var. quadricincta H. de
Saussure (1854) is the most common form of the species. It occurs in
British Guiana: Bartica; Kartabo; source of Essequibo River; Warina,
X. W. District. I have also seen it from French Guiana, Trinidad,
Venezuela, Colombia, Peru and Bolivia.

284 bulletin: museum of comparative zoology

Polybia sericea (Olivier 1791)

Olivier's typical form has the head black, the thorax and first
segment of the abdomen fulvous, and the remainder of the abdomen
blackish. This is a common wasp throughout most of Central and
South America, from Guatemala to Buenos Aires. It was originally
described from French Guiana. It occurs in British Guiana (Upper
Rupununi River) and I have seen it from Dutch Guiana (Paramaribo).
Rhopalidia rufithorax Lepeletier (1836), Apoica cubitalis H. de Saussure
(1854) and Polybia melanocephala Cameron (1906) were based upon
typical P. sericea.

Most specimens I have seen from British Guiana (Demerara; Onver-
wagt ; Georgetown ; Mt. Roraima ; Bartica District) belong to the form
of the species with both first and second abdominal segments fulvous.
This is Polistes nigripennis Fabricius (1804) and, if a name is needed for
it, may be called P. sericea var. nigripennis (Fabricius). I have also
seen it from Venezuela.

Polybia chrysothorax (Weber, 1801)

Polistes aurulenta Fabricius (1804) and Polybia aurichalcea H. de
Saussure (1854) are synonyms.

Common in British Guiana : Kartabo ; Kalacoon ; Demerara ; Bartica
District; Georgetown; Forest Settlement, Mazaruni River. I have
also seen it from Dutch Guiana, Colombia, Brazil, Venezuela and
Panama.

Polybia micans Ducke (1904)

1. Typical P. micans is russet to brownish-russet with more testa-
ceous areas, particularly at the apical margins of the abdominal seg-
ments and the base of the second tergite. The tegulae and legs are
russet-testaceous, the wings strongly tinged with russet. In British
Guiana it was taken at Kartabo and Bartica. I have also seen it from
French Guiana, Dutch Guiana, Brazil, Bolivia, and Ecuador. It
appears to be somewhat nocturnal and occasionally flies to light.
Polybia sericeibalteata Cameron (1906) I regard as a synonym of
typical micans.

2. P. micans var. velutina Ducke, was described by Ducke (1907)
as a distinct species, but I am unable to separate it by structural char-
acters from P. micans. It is fairly uniformly brownish-black, the tegu-
lae and legs being also of that color. There are a few testaceous blotches
on the head. The wings are more yellowish than russet. This form is

bequaert: social vespidae of the guianas 285

rather rare in British Guiana: Arakaka. It is known also from Brazil,
Bolivia, Peru, Ecuador and Colombia.

Polybia dimidiata (Olivier, 1791)

British Guiana: West bank of Demerara River; Oronoque River,
2°42'; Penal Settlement, Bartica District. I have also seen it from
Brazil, Dutch Guiana, French Guiana, Bolivia, Peru and Colombia.

The large syrphid fly, Ceriodes braueri Williston, is perfectly homeo-
chromic with this wasp.

Polybia rejecta (Fabricius, 1798)

I recognize three color forms of this common and widely distributed
wasp, but transitional specimens are frequent.

1. Typical P. rejecta has the head, thorax, and first abdominal seg-
ment black, with very few pale yellow markings (hind margin of
postscutellum and narrow apical margin of first tergite) ; the remainder
of the abdomen dull brick-red, with or without narrow, pale margins
(sometimes ventrally only). Sometimes the red turns fuscous or black-
ish. Common in British Guiana: Mt. Everard; Arakaka; Amatuk;
source of Essequibo River; Tumatumari, Potaro River; west bank of
Demerara River. Kartabo; Bartica; Rupununi; Penal Settlement,
Bartica District; Moraballi Creek, Essequibo River. I have also seen
it from Dutch Guiana, French Guiana, Trinidad, Venezuela, Brazil
and Peru. Polybia bicolor F. Smith (1857) and Eumenes impunctus
Provancher (1888) are synonyms.

Polybia rejecta race javaryensis Cameron (1906) is a species of
Mischocyttarus.

2. Var. belizensis Cameron (1906). Entire body black, the pale
markings of the thorax as in the typical form; abdomen dorsally with
a narrow apical margin on first segment only. This is the common
form in Central America, where I have seen it from British Honduras,
Guatemala, the Republic of Honduras, Costa Rica, and Panama; but
it occurs also in Colombia, Brazil and Peru.

3. Var. litoralis Zavattari (1905). Like var. belizensis, but all or most
of the abdominal tergites with rather broad pale apical margins.
Described from Ecuador. I have seen it from Colombia, where it is
common; but Ducke's locality "Bogota" is due to an erroneous label,
as no member of the subfamily Polybiinae occurs in the vicinity of
that city.

286 bulletin: museum of comparative zoology

Polybia occidentalis (Olivier, 1791)

The most common social wasp of tropical America, from Mexico
(northward to Nuevo Leon and Tamaulipas) to northern Argentina
and Uruguay. In the Antilles proper it is known only from St. Vincent
and Grenada. It is exceedingly variable in size, color and even the
shape of the first abdominal segment, but all of the forms intergrade.
The following seem to be sufficiently distinct to deserve names.

1. Typical P. occidentalis is black, with few or many, reduced or
more extensive pale yellow or whitish markings, fairly evenly dis-
tributed over head, thorax and abdomen; most or all of the abdominal
tergites have pale apical margins; mesonotum usually unstriped.
Vespa pygmaea Fabricius (1793), Polybia pygmaea var. minor Moebius
(1856), P. pygmaea var. major Moebius (1857), P. albo-picta F. Smith
(1857), P. bohemani Holmgren (1808) and Eumenes ductus Provancher
(1888) were based on some of the variantswhich I include undertypical
occidentalis. If desired, small specimens, with more slender first tergite
and few white markings, might be called var. pygmaea (Fabricius).

This form extends over practically the entire range of the species
and is common in the Guianas. British Guiana: Kartabo; Bartica
District; Demerara; Oko River (tributary of Cuyuni River); Upper
Rupununi.

Polybia fastidiosuscula var. nigriceps Zavattari (1906) appears to be
based upon specimens of P. occidentalis much like the typical form, but
with the head entirely black and traces of pale longitudinal stripes on
the mesonotum. I have seen such specimens occasionally from colonies
of otherwise typical occidentalis and cannot regard them as more than
individual variants.

Myraptera elegans Curtis (1844) was somewhat more extensively
bright yellow, particularly on the head, and had two yellow lines on
the mesonotum. It is doubtful whether it should be given varietal
status. It is clearly transitional to var. fiavifrons.

2. P. occidentalis var. fiavifrons F. Smith (1857) has the head almost
entirely yellow (usually with an orange tinge) and is also extensively
yellow on thorax and abdomen. In particular the mesonotum bears
two broad longitudinal stripes, often fused before the scutellum.
Polybia saussurei Holmgren (1808) is not separable, according to the
types I saw in Stockholm. This form is homeochromic with the
darker specimens of P. bistriata (Fabricius) ( = occodoma de Saussure);
but the structural characters given in the key readily separate the two
species.

bequaert: social vespidae of the guianas 287

I have seen the var. flavifrons from Ecuador (Guayaquil), Peru
(Trujillo) and Brazil (Coary, Amazonas; Lassance, Minas Geraes;
Tres Lagoas, Matto Grosso; etc.).

3. Var. parvula (Fabricius, 1S()4i was originally described as almost
wholly black, with only a narrow pale apical margin on the first
abdominal segment. Many transitions connect this form with typical
ocddentalis. I am unable to separate from it P. occidentalis var.
diguetana R. du Buysson (1905).

The var. parvula often occurs in the same districts as the typical
form, but is relatively rare in British Guiana: Demerara ; Kartabo;
Kaieteur, Arakaka. I have also seen it from Mexico, Guatemala,
Costa Rica, Panama, Colombia, Venezuela, French Guiana, Dutch
Guiana, and Brazil.

4. Var. juruana R. v. Ihering (1904) is remarkable for the extension
of the yellow markings on the abdomen, the second tergite being mostly
of that color. The mesonotum, however, has no yellow stripes. I have
seen it from Brazil (Puerto America, Rio Putumayo), Bolivia, Peru,
and Venezuela; it was described from the Jurua River, Brazil.

Specimens from Aquidauana (Matto Grosso), Brazil, in which the
apical yellow margin of tergite 2 is considerably widened on the sides,
but narrow in the middle, connect var. juruana with typical occident-
alis.

5. Var. scutellaris (White, 1S41) differs from typical occidentalis only
in the yellow being restricted to scutellum and postscutellum, which
are partly or wholly of that color.

I have seen it from Brazil (Bahia; Sao Paulo), Paraguay (Villarrica),
northern Argentina (Entre Rios) and Uruguay (Montevideo).

6. Var. spilonota Cameron (1904) has not only the scutellum and
postscutellum mostly dark yellow, often with an orange tinge, but also
a large orange-yellow spot (sometimes divided) on the propodeum, as
well as other markings on head, thorax and abdomen; often, but not
always, the mesonotum bears a prescutellar spot, divided anteriorly
into two prongs; the markings of the head are rather small.

It appears to be common in Nicaragua (San Marcos, topo-
types taken with the holotype, but probably not seen by Cameron;
Managua; Chinandega; Granada). I have also seen many specimens
from Mexico, Costa Rica and Panama which provide all transitions
between typical occidentalis and var. spilonota, sometimes apparently
in the same nest.

7. Var. ruficeps Schrottky (1902) is very distinct, the head being
entirelv or mostlv brick-red to dull red, with or without vellow mark-

288 bulletin: museum of comparative zoology

ings. The remainder of the body is much marked with yellow, but
without mesonotal stripes and with the apical margins of the tergites
complete, narrowed or interrupted medially.

It is common in northern Argentina, but occurs also in southern
Brazil (Corumba, Matto Grosso), Paraguay, and Bolivia (Buena
Vista de Sta. Cruz).

Polybia procellosa Zavattari (1906)

Typical procellosa, known from Ecuador only, is mostly ferruginous-
brown, variegated with black and yellowish markings, the legs yellow-
ish-russet, the wings gray with yellowish tinge.

The var. dubitata Ducke (1910) is black with a few whitish markings,
the legs black, the wings grayish without yellowish tinge. It was
originally described from the Amazon Basin in Brazil ; but I have seen
it from British Guiana (Kurupung) and Peru.

Polybia bistriata (Fabricius, 1804)

The species generally called Polybia oecodoma H. de Saussure (1854),
is Polistes bistriata Fabricius (1804), according to Fabricius' types. I
regard it as structurally distinct, not as a color variety of Polybia
occidentalis. The characters given in my key show that it is more
closely related to P. catillifex. Cameron's Polybia brunneiceps (1912),
described from British Guiana, without more definite locality, appears
to be a synonym of P. bistriata. The ground color varies from blackish-
brown to pale russet.

P. bistriata is common in British Guiana: Kartabo; Moraballi
Creek, Essequibo River; Rockstone, Essequibo River; Demerara;
Warina, N. W. District; Kalacoon; Mt. Roraima; Penal Settlement,
Bartica District. I have also seen it from Brazil, Dutch Guiana,
Venezuela, Ecuador, Peru, and Colombia.

Polybia catillifex Moebius (1856)

British Guiana: Turesi Falls; Kamakusa; Bartica; Pakaraimo Mts.,
head of Mazaruni River. Also known from Dutch Guiana, Brazil,
Ecuador, Colombia, and Peru.

Polybia singularis Ducke (1909)

British Guiana: Bartica District; source of Essequibo River. Known
also from the Amazon Basin in Brazil, Colombia (Mapiri) and eastern
Peru.

bequaert: social vespidae of the guianas 289

As shown in the key, this is closely allied to P. cmaciata. It builds a
similar nest of clay, but with a long, narrow, slit-like (not a circular)
entrance. Such nests were described by F. Smith (1851) and H. Lukas
(1890), but without knowledge of the builder, which was first recog-
nized by Ducke (1905; erroneously referred to P. cmaciata).

POLYBIA TINCTIPENNIS Fox (1898)

The typical color form of this species is known from southern Brazil,
eastern Peru, Honduras and Panama. In British Guiana (Kamakusa;
Kartabo; source of Essequibo River) it is replaced by the var. ncbulosa
J. Bequaert (1943), with uniformly infuscated wings. This form occurs
in Trinidad and probably also in the Amazon Basin of Brazil.

Polybia rufitarsis Ducke (1904)

I have seen the typical form of this species from British Guiana:
Moraballi Creek, Essequibo River. It is also known from Brazil.

Polybia ignobilis (Haliday, 1836)

The involved synonymy of this common South American wasp will
be discussed elsewhere. Polybia atra de Saussure (1854; not Vcspa
atra Olivier, 1791) and Polybia nigra de Saussure (1858) are synonyms.

I have seen a few specimens from British Guiana: Mt. Roraima. It
is known definitely also from Panama, Colombia, Venezuela, Brazil,
Paraguay, Bolivia, Peru and Argentina.

Apoica Lepeletier (1836)

There is only one structural species of Apoica, extremely variable in
color.

Apoica pallida (Olivier, 1791)

A. pallida is a nocturnal wasp, frequently attracted by light. I have
recently examined several thousands of these wasps. The conclusions
reached, as to color variation and distribution, will be embodied in a
paper to be published elsewhere. I am able to distinguish by name five
color forms, all of which occur in the Guianas. Although each form is
somewhat variable, there are few truly transitional specimens. The
females may be separated by means of the following key.

290 bulletin: museum of comparative zoology

1. Body testaceous, pale fulvous, or darker brown, either uniformly

so or with part to most of the abdomen paler than head and
thorax. No or very few pale yellow markings (none on head and
mesonotum). Wings more or less infuscated, often blackish. . . .

var. thoracica

Body testaceous to pale fulvous, with many yellowish markings or

extensively whitish or yellow 2

2. Abdomen (except base of tergite 1) dorsally pale yellow or nearly

white, often with silvery pubescence ("frosty"); venter either
yellow or pale yellow or fulvous. Head and thorax pale fulvous
or testaceous, usually spotted with yellow. Wings subhyaline,

brownish-russet along the costa var. pallens

Dorsum of abdomen not mostly pale yellow nor silvery 3

3. All abdominal tergites with a pale yellow, narrow or wide, apical

margin; sixth tergite entirely yellow. Head and thorax spotted
with yellow (mesonotum often with two or four yellow stripes).

Wings subhyaline, slightly pale russet along the costa

var. arborea
Abdominal tergites not all margined with pale yellow 4

4. Abdomen pale fulvous (except first tergite), with apex of first tergite

and broad base of second tergite pale yellow. Head and thorax
blackish without yellow spots (not stripes on mesonotum). Wings

strongly infuscated, nearly violaceous-black var. albi macula

Pale fulvous to darker brown. Sometimes with apex of first tergite
spotted with yellow ; sixth tergite as a rule mostly yellow. Head
and thorax with many pale yellow spots. Mesonotum with longi-
tudinal stripes (at least traces). Wings slightly russet, darker
along costa typical pallida

1. Typical A. pallida. — Syn.: Vesta pallida Olivier, 1791; Apoica
lineolata Lepeletier, 1836; Polistcs translucida Spinola, 1851; Apoica
bilineolata "Lepeletier" H. de Saussure, 1854; Apoica lineata "Lepele-
tier" R. du Buysson, 1906.

A common form in British Guiana: Rockstone, Essequibo River;
Mackenzie, Demerara River; Bartica; Kartabo; Upper Rupununi
River; Demerara River; Tumatumari, Potaro River; Torani Ranch,
Berbice River; Shudihar River; Wismar; Penal Settlement, Mazaruni
River; Kamakusa. Also known from British Honduras, Trinidad,
Dutch Guiana, Brazil, Ecuador and Peru.

2. A. pallida var. thoracica R. du Buysson, 1906.

N

bequaert: social vespidae of the guianas 291

This form is probably as common as typical pallida. British Guiana :
Mackenzie, Demerara River; Tumatumari, Potaro River; source of
Essequibo River; Kuyuwini River; Shudihar River. It is known also
from Costa Rica, Panama, Colombia, Venezuela, French Guiana,
Dutch Guiana, Brazil, Bolivia and Peru.

3. A. pallida var. pallens (Fabricius). — Syn.: Polistes pallens
Fabricius, 1804; Apoica pallida Lepeletier, 1836.

Very common in British Guiana: Kamakusa; Shudihar River;
Oronoque River, 2° 42'; Kartabo; Tumatumari, Potaro River; Kuyu-
wini River; Georgetown ; source of Essequibo River. I have seen it also
from British Honduras, Guatemala, Nicaragua, Costa Rica, Panama,
Colombia, Venezuela, Trinidad, Dutch Guiana, French Guiana, Brazil,
Paraguay, Bolivia, Peru and Ecuador. It is reported from southern
Mexico.

4. A. pallida var. arborca H. de Saussure. — Syn. : Apoica arborea
H. de Saussure, 1854.

British Guiana: Kartabo; Torani Ranch, Berbice River; Shudihar
River; Tumatumari, Potaro River; Mazaruni River. Also known from
Dutch Guiana, French Guiana, Brazil, Bolivia, eastern Peru and
Ecuador.

5. A. pallida var. albimacula (Fabricius). — Syn.: Polistes albima-
cula Fabricius, 1804; Polistes albimaculata "Fabricius" H. de Saussure,
1854.

A rare form known only from Dutch Guiana and British Guiana:
Shudihar River; Kartabo; Pakaraimo Mis. at the headwaters of the
Mazaruni River.

Stelopolybia Ducke (1910)
(Including Gymnopolybia Ducke, 1914)

Ducke (1914, Zool. Lahrb., Abt. Syst., 36, pp. 317 and 327) divided
his genus Stelopolybia into two groups, restricting the earlier name to
the species which build nests in the open, but with the combs enclosed
in a paper envelope. The species building uncovered combs inside
some natural cavity, he separated as Gymnopolybia. Although he
mentioned some structural differences between these two biological
groups, he could find none that divided the species either consistently
or naturally.

Polistes angulata Fabricius (1804) was selected as the type of Stelo-
polybia by R. Lucas (1912, Arch. f. Naturgesch., 77, Bd. 4, Heft 1,

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p. 210).1 Unfortunately this is one of the species with uncovered
combs which Ducke later placed in Gymnopolybia. O. W. Richards
(1943, Proc. Ent. Soc. London, ser. B, 12, pts. 3-4, p. 45) has now
chosen Polybia vulgaris Ducke (1904), which I regard as a synonym
of Vespa fulvofasciata Degeer (1773), as the type of Gymnopolybia.
This is also one of the species with uncovered combs. It thus appears
that the designated genotypes of Stelopolybia and Gymnopolybia are
strictly congeneric, making the two names synonyms and leaving the
group of species building combs inside a paper envelope without a
distinct name. I feel, however, that the distinction between the two
groups is purely biological. The morphological differences assigned to
them by Ducke and Richards are either inadequate, showing transi-
tions or not applying to all the species of each group, or else of very
secondary importance in the general classification of the Polybiinae.
For this reason I do not propose a new name for the species which build
combs within an envelope.

H. de Saussure (1854) divided his subgenus Polybia, s. sir. into a
number of divisions : Alpha, Iota, Phi, Mu, Kappa and Omega. Accord-
ing to F. J. Griffin (1939, Jl. Soc. Bibl. Nat. Hist., 1, pp. 211-212),
these were all published the same year (1854) and antedate de Saus-
sure's use of some of them for groups of solitary wasps. Only Phi
and Mu contained species now placed in Stelopolybia, in addition to
others. In view of the divergence of opinion regarding the use of
Greek letters spelled out as generic or subgeneric names, I select types
which will make them synonyms of older names. Phi originally con-
tained 10 species, 8 of which are now placed in Stelopolybia and one in
Mischoeyttarus, one being unrecognized though probably also a Stelo-
polybia. I herewith designate as type Vespa phthisica Fabricius (1793),
now placed in Mischoeyttarus subgenus Kappa de Saussure (1854) (see
O. W. Richards, 1941, Proc. Ent. Soc. London, B, 10, pp. 125-126).
Mu comprised 11 species, 6 now placed in Polybia, 2 in Stelopolybia
and 3 in Mischoeyttarus. I herewith designate as type Myrapetra
scutellaris White (1841), now placed in Polybia and which is, more-
over, the monotype of Myrapetra White (1841).

iR. Lucas' earlier selection of P. angulata invalidates O. W. Richards' recent (1943) choice of
Polybia infernalis de Saussure as the type of Stelopolybia.

bequaert: social vespidae of the guianas 293

Key to Guiana Species

1. Dorsal area of pronotum rounded off, at most with a trace of

humeral collar 2

Pronotum with a raised humeral collar, sometimes projecting as
lateral angles. Oculo-malar space long 5

2. Small species, 10 to 12 mm. long. Eyes with rather distinct short

hairs 3

Large species, 15 to 20 mm. long. Eyes scarcely hairy. Oculo-
malar space short, but distinct 4

3. Oculo-malar space very long (about as long as ninth antennal seg-

ment). Upper plate of mesepisternum longer than high

S. cajennensis

Oculo-malar space very short (eye almost touching mandibular

condyle). Upper plate of mesepisternum nearly as long as high

S. pollens

4. Clypeus with distinct, fairly large punctures. Anterior margin of

pronotum moderately raised near the coxae. Lower outer orbit
nearly as wide as eye in profile. Larger, wing 15 to 16 mm. long

S. paracusis

Clypeus with fine punctures. Anterior margin of pronotum strongly

raised into a translucent lamella near the coxae. Lower outer

orbit narrower than eye in profile. Smaller, wing 13 to 14 mm.

long S. obidensis

5. Sides of humeral collar strongly projecting and angular. Eyes dis-

tinctly hairy 6

Sides of humeral collar rounded off, not or scarcely projecting. Eyes
with a few sparse hairs 8

6. First abdominal segment slender, longer than the combined pro-

podeum and postscutellum. Wing 14 to 15 mm. long, slightly

yellowish S. constructrix

First abdominal segment scarcely longer than the propodeum .... 7

7. Body orange-yellow with russet areas, the hind half of abdomen

black; second tergite with yellow base and apical margin. Wings

strongly russet-yellow, 17 to 18 mm. long <S. testacea

Body black rarely with a few yellow markings. Wings subhy aline,
with a yellowish tinge, about 16 mm. long S. angulata

8. Larger; wing about 14 mm. long. First abdominal segment rather

abruptly widened and bell-shaped, slightly angular at the sides
seen from above. Second tergite rather suddenly widened behind
the base S. fuho-fasciata

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Smaller; wing 9 to 11 mm. long. First abdominal segment very
gradually widened, not bell-shaped. Second tergite gradually
behind the base S. pallipes

Stelopolybia cajennensis (Fabricius, 1798)

Poly bia lignicola Ducke (1904) is a synonym.

A common species in British Guiana: Bartica; Kamakusa, Warina,
N. W. District; Kartabo; Forest Settlement, Mazaruni River. Also
known from Dutch Guiana (Saint Barbara Plain, Surinam River;
Paramaribo), French Guiana, Trinidad, Brazil, Colombia, Peru,
Ecuador, Panama, and the Republic of Honduras.

Stelopolybia pallens (Lepeletier, 1836)

The type of Rhopalidia pallens Lepeletier is now in Spinola's collec-
tion at the Turin Museum, where it was recognized by Ducke as identi-
cal with Polybia inf emails de Saussure (1854). Lepeletier 's name is
valid; he described his R. pallens as a new species, without any refer-
ence to Polistes pallens Fabricius (1804), which is not placed now in
Stelopolybia. Other synonyms are: Polistes rufina "Illiger" Erichson
(1848), Polybia ampullaria Moebius (1856); Eumenes fiavopectus
Provancher (1888); and Polybia internalis Dalla Torre (1904).

Common in British Guiana: Bartica; Kamakusa; Kaieteur; Turesi
Falls; Kartabo; Baracara, Mazaruni River; Demerara; Moraballi
Creek, Essequibo River; Courantyne River ; source of Essequibo River;
Forest Settlement, Mazaruni River; west bank of Demerara River.
Also known from French Guiana, Trinidad, Brazil, Bolivia, Peru, and
Ecuador.

Stelopolybia paraensis (Spinola, 1851)

This species occurs in three color forms.

1. Typical S. paraensis is extensively orange-yellow or pale ferrugi-
nous ventrally, blackish-brown or black dorsally with many pale yellow
markings, forming longitudinal stripes on the mesonotum and apical
fasciae on most of the tergites; base of first and second tergites also
yellowish; antennae and legs orange-russet; tegulae yellowish; wings
strongly tinged with yellow. Not rare in British Guiana: Mt. Roraima
Kalacoon; Pakaraimo Mts., head of Mazaruni River; Shudihar River.
Also known from Brazil, Peru, and Bolivia. Homeochromic with S.
obidensis.

bequaert: social vespidae of the guianas 295

2. S. paraensis var. ruficornis Ducke (1905). Mostly black; anten-
nae russet (scape darker); spots on sides of frons, outer orbits, hind
margin of pronotum, tegulae, longitudinal streaks on propodeum,
tibiae, tarsi, apical margins of some of the tergitcs and sternites, more
or less testaceous or whitish; no stripes on mesonotum. Wings as in
typical form. Originally described from the Upper Amazon, Brazil
(Japura River; Tabatinga). I have seen it from Peru (Iquitos; Rio
Tapiche). Ducke also reports it from eastern Ecuador.

Ducke records specimens from Chiriqui, Panama, transitional
between typical paraensis and ruficornis. I have seen such a specimen
from Pozuzo, Peru. It has the legs mostly yellowish and two narrow
yellowish lines on the mesonotum; otherwise it is like ruficornis.

3. S. paraensis var. (or subsp.) obscurior, new.

Female. — Body black; only the flagellum (particularly below) and
fore tibiae and tarsi somewhat russet; lower inner orbital margins
orange or dirty yellow, filling the ocular sinuses. Tegulae black. Wings
as in typical form.

Ecuador: Jatun Yucu, Rio Napo Watershed, 700 m., holotype and
paratypes (W. Clarke-Macintyre). — Colombia: Restrepo, Int. Meta,
500 m., paratype (J. Bequaert). — British Guiaxa: Mt. Roraima,
paratype (J. G. Myers); source of Essequibo River, paratypes (J.
Ogilvie). — Dutch Guiana: Brownsberg, paratypes (D. C. Geijskes).
— Holotype and paratypes at Mus. Comp. Zool., Cambridge, Mass.;
paratypes at U. S. Nat. Mus. and Amer. Mus. Nat. Hist.

Perfectly homeochromic with S. angulata, but without the humeral
angles of that species.

Stelopolybia obidensis (Ducke, 1904)

Polybia paraensis var. luctuosa W. A. Schulz (1905) is a synonym.

British Guiana: Tumatumari; Pakaraimo Mts., head of Mazaruni
River; source of Essequibo River; Kamakusa; Tukeit; Forest Settle-
ment, Mazaruni River; Turesi Falls; Kartabo; Bartica; Moraballi
Creek, Essequibo River; Penal Settlement, Bartica District. I have
seen it also from Dutch Guiana and it is known from Brazil.

Stelopolybia constructrix (de Saussure, 1854)

British Guiana: Pakaraimo Mts., head of Mazaruni River; Kama-
kusa ; Kartabo ; source of Essequibo River. Also in French Guiana and
Brazil.

296 bulletin: museum of comparative zoology

Stelopolybia testacea (Fabricius, 1804)

Common in British Guiana: Kartabo; west bank of Demerara
River; Mackenzie, Demerara River; Bartica; Pakaraimo Mts., head
of Mazaruni River; Kuyuwini River; Oko River, a tributary of the
Cuyuni River; Mt. Roraima. Also seen from French Guiana, Dutch
Guiana, Brazil, Venezuela, Bolivia and Peru.

Stelopolybia angulata (Fabricius, 1804)

The typical form of this species, without yellow markings and with
black tarsi, occurs in British Guiana: Kaieteur; Bartica; Kamakusa;
Arakaka; west bank of Demerara River; Warina, N. W. District;
Pakaraimo Mts., head of Mazaruni River; Mt. Roraima. I have seen
it also from Venezuela, Dutch Guiana, Brazil, Ecuador, Peru and
Bolivia. In Guatemala, Costa Rica, Panama and parts of Colombia it
is replaced by a closely allied species, S. panamensis (Cameron), in
which the humeral angles are much less prominent.

S. angulata var. angulicollis (Spinola, 1851) has the tips of the
femora and the entire tibiae and tarsi yellowish ; while the var. ornata
(Ducke, 1905) has, in addition, some pale yellow spots on the thorax.
Neither of these is known from the Guianas.

Stelopolybia fulvo-fasciata (Degeer, 1773)

I regard Polistes hectica Fabricius (1804) and Polybia vulgaris Ducke
(1904) as synonyms. Vespa ochrosticta Weber (1801) may also have
been the same. Vespa fasciata Olivier (1791) appears to be merely a
new name proposed for Vespa fulvo-fasciata Degeer. Polybia fasciata
Lepeletier (1836) was also the same wasp.

Common in British Guiana: Pakaraimo Mts., head of Mazaruni
River; Kamakusa; Warina, N. W. District; New River, a tributary of
the Courantyne; Penal Settlement, Bartica District; source of Esse-
quibo River; Rockstone, Essequibo River; Mt. Roraima; Baracara,
Mazaruni River; mouth of Meamo River. Also known from French
Guiana, Dutch Guiana, Brazil, Bolivia, Peru, Ecuador and Colombia.

Stelopolybia pallipes (Olivier, 1791)

The unwarranted emendation "pallidipes" was first used by Dalla
Torre (1894). The following names are either synonyms or based upon
color forms: Polybia anceps de Saussure (1854); Polybia lutea Ducke

bequaert: social vespidae of the guianas 297

(1904); Polybia myrmecophila Ducke (1905); Polybia festae Zavattari
(1906); Polybia pallipes var. centralis Cameron (1907); and Polybia
pallipes subsp. cuzcoensis Schrottky (1911).

1. Olivier's original description reads: "Vespa pallide testacea,
capite thoracisque dorso nigro maculatis, ahdomine fusco apice pallido.
Elle a environ cinq lignes de long [5 French lines = 12.3 mm.]. Les
antennes sont noires, avec les premiers articles d'un fauve pale en
dessous. La tete est d'un fauve pale, avec la partie superieure tachee
de noir. Le corcelet est d'un fauve pale, avec trois lignes noires sur le
dos. Le petiole est fauve pale. L'abdomen est obscur, avec la base
d'un fauve pale. Les pattes sont d'un fauve pale. Les ailes sont trans-
parentes." The French description agrees well with what Ducke
(1910) called " Stelopolybai pallidipes" and de Saussure (1854, Et.
Fam. Vesp., 2, PL XXV, fig 2) figures as "Polybia pallipes". In this
typical form the abdomen has the first tergite (or petiole) pale fulvous,
the second yellowish fulvous with infuscate or blackish apical margin,
the remaining segments blackish; there are no distinct yellow apical
fasciae. P. lirfea Ducke does not seem to differ from typical pallipes.

2. 8. pallipes var. anceps (de Saussure) has most of the abdomen
strongly infuscate or blackish, with distinct yellow apical margins at
least on tergites 2 and 3, sometimes also on 4 and 5; the yellow base of
the second tergite is, as a rule, sharply defined from the fulvous discal
area.

3. S. pallipes var. festae (Zavattari) is the extreme melanistic form
of the species, with the body blackish-brown or black and few yellow-
ish markings on head and sides of thorax (none on mesonotum or
abdomen). It is known only from Eastern Ecuador (San Jose, 1800
m.). I have not seen it.

4. S. pallipes var. cuzcoensis (Schrottky) has the«black abdomen of
var. festae; head and thorax are profusely marked with yellow, as in
the typical form. It appears to be characteristic of parts of eastern
Peru: Achinamiza; La Chorrera, Putumayo; Chanchamayo. I have
also seen it from Bolivia : Huachi, Rio Beni. It is evidently the form
with black abdomen mentioned by Ducke from the Upper Amazon:
Tabatinga; Iquitos.

5. S. pallipes var. (or subsp.) fulvanceps, new.

Female. Ground color orange to fuscous-yellow, darker on the
dorsum of the abdomen, more yellowish ventrally; head and meso-
notum with the same blackish markings as in the other forms of the
species; narrow base of second tergite and broad and well-marked apical
margins of tergites 2 to 5 bright yellow; tergite 6 mostly yellow.

298 bulletin: museum of comparative zoology

Wings and legs as usual. This form combines the fulvous-orange
abdomen of var. myrmecophila with the abdominal bands of var.
anccps.

Colombia: Rio Frio, Dept. Magdalena, holotype and paratypes
(G. Salt). — Peru: Colonia Perene, paratype (J. C. Bradley); Puerto
Bermudez, Rio Pichis, paratypes (J. C. Bradley); Miriantiriani,
Camino del Pichis, paratypes (J. C. Bradley). — Holot3rpe and para-
types at Mus. Comp. Zool., Cambridge, Mass.; paratypes also at
Cornell University (Dept. of Entomology).

6. S. pallipes var. myrmecophila (Ducke) has the head and thorax
of the typical form, but the abdomen is almost unicolorous pale
yellowish-russet or orange, usually with more yellow bases of second
and third tergites. Sometimes the apical margins of some of the ter-
gites are very narrowly or faintly yellowish. P. pallipes var. centralis
Cameron is clearly the same form, as was recognized by Ducke.

Typical S. pallipes was described originally from French Guiana
(Cayenne) and I have seen it from British Guiana (Mt. Roraima),
where it appears to be uncommon. I also know it from Brazil, Para-
guay, and Peru (mouth of Rio Cotuhe)

The var. anceps is the usual form in British Guiana: Kamakusa;
Kartabo; west bank of Demerara River. I have also seen it from
Dutch Guiana, Trinidad, Brazil, Venezuela, Colombia, Panama and
Bolivia.

The var. myrmecophila also occurs in British Guiana: Source of
Essequibo River; Demerara River; Rupununi River. It is more widely
distributed than any of the other forms, as I have seen it from Guate-
mala, Honduras, Costa Rica, Panama, Colombia, Ecuador, Peru and
Brazil.

Pseudopolybia H. de Saussure (1863)

The genus was revised in 1938 (Rev. de Entomologia, 9, pp. 112-
113), but a new species is described below. Of the five species known
at present, four occur in the Guianas.

1. Mesepisternum not divided by an oblique suture. Humeral margin
of pronotum straight, bluntly ridged. First abdominal segment
longer than wide, forming a broad stalk. Small; fore wing 5.5
to 6 mm. long P. pusilla

Mesepisternum completely divided by a well-marked oblique suture
into an upper and a lower plate 2

bequaert: social vespidae of the guianas 299

2. Small; fore wing 5.5 to G.5 mm. long. First abdominal segment

longer than wide, forming a distinct stalk, gradually widened
from base to apex (seen from above). Clypeus slightly produced

medially P. langi

Larger; fore wing 11 to 12 mm. long. First abdominal segment
either cap-shaped, or, if stalk-like, not gradually widened from
base to apex. Clypeus strongly produced medially 3

3. First abdominal segment slightly longer than wide, forming a broad,

bell-shaped stalk, set off from the remainder of the abdomen

P. difficilis

First abdominal segment wider than long, cap^shaped and not set

off as a stalk from the remainder of the abdomen ... P. vespiceps

Pseudopolybia vespiceps (de Saussure, 1863) has been taken in
French Guiana and Dutch Guiana, as well as in Brazil.

Pseudopolybia pusilla (Ducke, 1904)

British Guiana: Monkey Jump, Essequibo River (Oxford Uni-
versity Exped.).

The species is also known from Brazil (Para and Oyapoc). It is
readily mistaken for a Leipomeles. It differs from L. dorsata in the
much wider clypeus and stronger humeral transverse ridge. The nest
is as yet unknown.

Pseudopolybia difficilis (Ducke, 1905)

I have seen this species from British Guiana: source of Essequibo
River. It is also known from Brazil, Peru, and Bolivia.

PsEUDOPOLYBtA langi, new species. Fig. 2

Female. Head moderately flattened, slightly wider than thorax;
seen in front, slightly wider than high; from above, rectangular with
receding hind corners, about twice as wide as long; occipital margin
nearly straight. Vertex and genae not margined by a carina behind;
gena scarcely narrower than the eye in profile. Oculo-malar space
about half the length of the fourth antennal segment. Inner orbits
about one and one-third times as far apart on vertex as at clypeus.
Ocelli rather large, in an equilateral triangle; posterior ocelli about
twice as far from eyes as from each other. Interantennal shield broad,
not set off, strongly but bluntly and evenly swollen. Antennae nearly

300

bulletin: museum of comparative zoology

twice as far apart as from eyes. Frons slightly convex. Clypeus at its
narrowest about one and two-thirds times as wide as high, irregularly
pentagonal with the upper side much the longest, contiguous to the
eyes over nearly one-half of sides; apical margins slightly produced,
ending in a very broad, obtusely rounded point. Mandible with an
oblique cutting edge of four sharp teeth, the lower three subequal, the
upper one much smaller; most of outer surface flattened or slightly
depressed. Antenna: scape short and rather thick, scarcely curved;

Fig. 2. Pseudopolybia langi J. Bequaert. Female. A, wings; B, head in
front view; C, maxillary palpus; D, labial palpus; E, body in profile.

second segment short, moderately swollen ; third less than three times
the length of second; fourth and fifth about as long as wide; sixth to
eleventh slightly wider than long; twelfth slightly longer than wide
at base, with bluntly rounded tip; flagellum about equally thick
throughout. Maxillary palpi of 6 segments. Labial palpi of 4 seg-
ments, the tip of the third with a short, heavy, erect, curved seta.
Thorax moderately elongate; in profile, about one and one-third times
as long as high ; from above, nearly twice as long as wide before tegu-
lae. Pronotum evenly rounded into a semi-circle anteriorly, the
humeral margin barely indicated by a very weak transverse, blunt

bequaert: social vespidae of the guianas 301

ridge on the sides, where it forms no angles ; dorso-lateral areas sloping;
lower anterior margin of ventrolateral areas with a vertical swelling
indented by a deep pronotal fovea. Mesopleuron moderately swollen ;
mesepisternum completely divided by an oblique suture into an upper
and a lower plate; deep mesepimeral suture ending abruptly and far
from the metapleuron. Upper sclerite of metapleuro'n ending obliquely
below, with a short extension along mesometapleural suture. Scutel-
lum slightly and evenly convex, anteriorly with a fine, raised, longi-
tudinal line which continues posteriorly as an impressed line. Post-
scutellum transversely elliptical, moderately swollen. Propodeum not
swollen, moderately slanting in profile; median concavity a broad,
shallow, longitudinal groove. First abdominal tergite narrowed,
moderately elongate, very gradually widened and thickened from base
to apex, where it is less than half as wide as second tergite, triangular
in outline from above. Second tergite from above wider than long,
gradually widened at base ; in profile, slightly and evenly convex. Mid
tibiae with two spurs. Claws symmetrical, unarmed. Venation:
second cubital cell much higher than wide; third cubital longer than
wide, rectangular, about as long on cubitus as on radius; radial cell
very long, acute; basal vein ending in subcosta close to the large
stigma.

Smooth and moderately shiny, without appreciable punctures or
other sculpture. Pubescence sparse and short. Eyes with many dis-
tinct erect hairs.

Head, thorax and legs pale yellowish, with a few fuscous areas on
frons, vertex, occiput, pronotum, mesepimeral suture, propodeum
(concavity and extreme sides), hind portion of postscutellum, and
tibiae. Antennae fuscous above, russet below. Teeth of mandibles
fuscous. Mesonotum fuscous, with a pair of broad median yellow
stripes and a shorter yellow streak on each side near the tegula.
Abdomen pale fuscous to light ferruginous, more or less yellowish at
the base of the second tergite. Wings hyaline throughout; veins pale
fuscous; stigma translucent medially.

Length (h.+th.+t.l+2): 5.5 mm.; of fore wing, 5.6 mm.

British Guiana: Kamakusa, female holotype and paratypes from
one nest (Herbert Lang). Holotype and paratypes at Mus. Comp.
Zool., Cambridge, Mass.; paratypes also at Am. Mus. Nat. Hist, and
U. S. Nat. Mus.

The characteristic thick seta of the labial palpus is shorter than in
some other species of the genus and, being of a pale color, is more
difficult to see.

302 bulletin: museum of comparative zoology

Parachartergus R. v. Ihering (1904)

This genus was revised by me in 1938 (Rev. de Entomologia, 9, pp.
105-112). Unfortunately I called it there Chartergus, in the belief that
Ashmead (1902) was the first to select a genotype for that name. The
late Miss Sandhouse informed me that Emile Blanchard in 1840
(Hist. Nat. Ins., 3, Orth. Nevr. Hem. Hym. Lep. Dipt., p. 395) had
designated Vespa nidulans Fabricius (1793), a synonym of Vespa
chartaria Olivier (1791), as the type of Chartergus Lepeletier. The
name Parachartergus must therefore be used for the genus which I had
called Chartergus.

Key to Guiana Species

1. Vertex and cheeks not separated from the occiput by a carina.

Humeral margin of pronotum erect, not overlapping nortouching

the vertex P. frontalis

Vertex and cheeks separated from the occiput by a sharp carina.

Humeral margin of pronotum slanting, touching or overlapping

. the vertex 2

2. Vertex on each side, near the upper inner orbit, with a slightly

raised, impunctate area. Smaller; fore wing 8 mm. long

P. smithii
Vertex without impunctate lateral areas 3

3. Body thickset, black. Humeral margin of pronotum low, ridge-

like, opaque. Wings black, with the apical portion either some-
what paler or decidedly whitish. Fore wing 9 to 12 mm. long. .4
Humeral margin of pronotum raised into a high, sometimes trans-
lucent lamella. Wings not black with paler or whitish tips.
Fore wing 6.5 to 8 mm. long 5

4. Head and thorax with distinct, long erect hairs P. apicalis

Head and thorax covered with a grayish bloom, the erect hairs

restricted to clypeus, postscutellum and propodeum

P. fratemus

5. Body lengthened, black or ferruginous. Thorax over one and a half

times as long as wide seen from above P. fulgidipennis

Body short, thickset, mostly testaceous or russet. Thorax about

one and one-third times as long as wide seen from above

P. eolobopterus

Parachartergus eolobopterus (Weber, 1801) was taken in French
Guiana and is known also from Colombia, Venezuela, and Trinidad.

bequaert: social vespidae of the guianas 303

Parachartergus frontalis (Fabricius, 1S04), not yet reported from the
Guianas, is known from the adjoining Oyapoc district of Brazil and
Venezuela.

Parachartergus apicalis (Fabricius, 1804), frequently recorded from
the Guianas, possibly does not occur there, as it is often confused with
P.fraternus, which I regard as a distinct species.

Paraciiartergus fraterxus (Gribodo, 1S91)

1 . The typical form of this species has the tips of the wings whitish
with pale veins, being colored exactly like P. apicalis. I have seen it
from Dutch Guiana (Paramaribo).

2. In the var. concolor Gribodo (1891), the tips of the wings are
somewhat paler but not whitish and have fuscous veins. I have seen
it from British Guiana (Hepseba, Courantyne River; Arakaka),
French Guiana, Brazil, Trinidad, Colombia, and Panama. It was
first described from Venezuela.

Parachartergus fulgidipennis (H. de Saussure, 1854)

I have recognized by name five color forms of this species, two of
which occur in the Guianas.

1. The var. griseus Fox (1898) has the thorax and abdomen black,
the abdomen rather dull with many long hairs, and the head partly
pale yellow. The wings are subhyaline, broadly black along the costal
margin, without distinct cream-colored transverse patch. Charter gus
trichiosomus Cameron (1912), described from British Guiana, is a
synonym. I have seen this form from British Guiana (Moraballi
Creek, Essequibo River), Brazil, and Peru.

2. In the var. fasdipennis Ducke (1905) the color of the body is as
in var. griseus, but the abdomen is rather shiny, with sparse and short
hairs, and the wings have a distinct cream-colored transverse patch,
which interrupts the black streak along the costal margin. It was
originally described from Brazil; but I have seen it from British
Guiana: Ite Cattle Trail.

Parachartergus smithii (H. de Saussure, 1854)

The typical form of this species was taken in British Guiana:
Kartabo. It also occurs in British Honduras, Costa Rica, Colombia,
Ecuador, Peru, and Brazil.

304 bulletin: museum of comparative zoology

Leipomeles Moebius (1856)
This genus is monotypic.

Leipomeles dorsata (Fabricius, 1804)

W. A. Schulz (1912, Berlin. Ent. Zeitschr., LVII, p. 87) examined
the type of Polistes dorsata Fabricius and recognized that it was the
wasp described by Moebius (1856) as Leipomeles lamellaria. Polybia
nana H. de Saussure (1863) and Polybia spilogastra Cameron (1912)
are other synonyms.

British Guiana: Source of Essequibo River; Monkey Jump, Esse-
quibo River. Fabricius' type came from the Essequibo River. Also
known from Dutch Guiana, Brazil, Bolivia, Peru, Ecuador, and
Panama (Barro Colorado).

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 8

THE BIRD FAUNA OF THE WEST SUMATRA ISLANDS

By S. Dillon Ripley

Division of Birds
United States National Museum

With Two Plates

CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
October,; 1944 j

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Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1, 2,
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After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE

Vol. XCIV, No. 8

THE BIRD FAUNA OF THE WEST SUMATRA ISLANDS

By S. Dillon Ripley

Division of Birds
United States National Museum

With Two Plates

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

October, 1944

Xo. 8— The Bird Fauna of the West Sumatra Islands1
By S. Dillox Ripley

CONTENTS

Page

Introduction 307

Acknowledgements 308

Ornithological History 309

Geography of the islands 311

A Faunal List of the birds of the islands 317

Summary and Conclusions 415

Bibliography 427

INTRODUCTION

The islands of the East Indies stretching from the Asiatic mainland
down to the southeast for more than two thousand m'les 'nto the
Papuan and Australian regions, form a true paradise for the student
of faunal distribution and speciation. The position of this myriad of
islands large and small, each with its varying type and degree of geo-
graphic and geologic isolating mechanisms is highly interesting. The
equable climate, with a minimum of seasonal or spasmodic changes,
adds to the effectiveness of even the most limited geographic barriers.
Ten or fifteen miles of sea is enough to preclude interchange of genes
between two sedentary populations. As an example of this kind of
speciation the bird fauna of the west Sumatra islands offers a superla-
tive opportunity for study.

Several factors have influenced my decision to study the birds of the
west Sumatra islands. During 1939 I was able to visit Nias, the largest
island of the group, where I made a small collection for the Academy
of Natural Sciences of Philadelphia. When, early in 1942 I came to the
United States National Museum from Harvard, I found that I would
have an opportunity in Washington to make an exhaustive study of
what is the largest and finest collection of birds from these islands, that
of the late Dr. W. L. Abbott. Dr. Abbott's collection has never been
reported on as a whole, but various papers by the late Dr. Charles
Richmond, Dr. H. C. Oberholser, and the late Mr. J. H. Riley have

1 In the absence of the author in war service, the proof was kindly read by Dr. Herbert
Friedmann, Curator of Birds, U. S. National Museum. Editor.

308 bulletin: museum of comparative zoology

appeared from time to time describing more than one hundred and thirty
new species and subspecies, the types of which are now in the United
States National Museum. Thus without actually returning to the
islands, now out of the question, I have been able to accumulate re-
markably complete data on the fauna of this area.

ACKNOWLEDGEMENTS x

To the late Dr. Glover M. Allen of Harvard, my councillor and
friend, I owe a great deal for what has gone into this study. Professor
A. C. Romer has been most helpful with aid and advice. Mr. James
L. Peters has been kindness itself in rendering me every encouragement
and assistance. Dr. Ernst Mayr of the American Museum of Natural
History and Mr. Rodolphe de Schauensee of the Academy of Natural
Sciences have freely loaned specimens from their collections and have
given much valuable advice. To the authorities of the United States
National Museum, Dr. Alexander Wetmore, Dr. Herbert Friedmann,
and Mr. H. G. Deignan, I owe a tremendous debt of gratitude for their
generous help and wise counsel. Certain final questions of identifica-
tion remain unsettled. For this unfortunately there is no solution
until such time as free communication, field study, and research in
general return to their proper places in this now sadly maladjusted
world. That this will happen in good time is the earnest hope and
calm conviction of all those of us who now labor in alien and exciting
fields.

S. Dillon Ripley
Washington, D. C.
November, 1942.

> Originally presented as a thesis in partial fulfillment of the requirements for the degree of
Doctor of Philosophy at Harvard University.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 309

ORNITHOLOGICAL HISTORY

The first serious scientific observing and collecting on the west
Sumatra islands was done by Baron H. C. B. von Rosenberg who
visited Nias in 1854. So far as is known the list of 56 species of birds
published by Xieuwenhuisen and himself in 1SG3 as being found on
Nias, and again the list published by him in 1878 of 60 species of
birds noted on Nias and nearby islands are not based on actual collected
specimens but only on observations. The first actual collection of
birds was made by Signor Elio Modigliani during his exploration of
Nias from April through August, 1886. This collection of 62 species,
eight of which were new, was published on by Count Salvadori in 1886
and also by Modigliani himself in 1890. By this collection the known
avifauna of Nias was raised to 102 species.

Two collections were made among these islands in 1S91. One on
Nias was prepared by M. J. Claine and reported on by Dr. Oustalet
in 1892. Signor Modigliani meanwhile had gone to Enggano and his
very interesting material consisting of 23 species, seven of which were
new, was described by Salvadori in 1892. Another collection made on
Nias was that of Mr. W. Thomas, a missionary on the island, who
sent birds to Graf von Berlepsch. Some of these birds were described
by Salvadori and some by Berlepsch in 1895. Signor Modigliani, who
was still in the islands, now visited Sipora, and Salvadori listed his
collection from there of 34 species, 3 of which were new, in 1894.

A hard-working and able young entomologist, Mr. J.Z.Kannegieter,
visited Nias and the Batu Islands during the winter of 1895-96. Mr.
J. R. H. Neervort van der Poll, for whom Kannegieter was working,
presented his bird collection to the Leyden Museum, where the Nias
collection was reported on by Dr. J. Biittikofer. Dr. Biittikofer's list
raised the known avifauna of Nias to 128 species, of which 4 were
described as new in his paper (1896). Mr. van der Poll included in-
formation on the native names of these birds as well as notes on the
color of the soft parts of the specimens. The 355 Batu specimens,
except for two species described by Finsch (1899), remained unworked
until 1940 when the Academy of Natural Sciences of Philadelphia pub-
lished two papers by de Schauensee and Ripley, and de Schauensee.

The next collections made on the islands were those of the tireless .
Dr. W. L. Abbott who voyaged about the East Indian islands for
several years on a small schooner manned by an adventuresome and
daring Malay crew. Dr. Abbott started in the autumn of 1901 at the
northern end of the chain, visiting Simalur, Babi, Lasia, and the islands

310 bulletin: museum of comparative zoology

of the Banyak group, Bangkaru and Tuangku. A year later in the
autumn and winter of 1902, Dr. Abbott along with Mr. Charles Boden
Kloss of the Federated Malay States Museum went again to Simalur.
From there they sailed south to the Pagi Islands, the Batu Islands,
Tello, Tana Massa, and Tana Bala, and lastly Nias. Two years later
in the fall of 1904, Dr. Abbott visited Enggano. His last trip to the
group was to Nias in the spring of 1905. As a result of these compre-
hensive trips more than 1300 bird skins were presented to the United
States National Museum.

Several papers have been written on these collections. Dr. Charles
W. Richmond published the first on the Simalur trip in 1903 describ-
ing 19 new forms. Much later, Dr. Harry C. Oberholser (1919) wrote
up Dr. Abbott's second collection on Simalur describing one new form.
In these two papers there are color notes on the species, and in Dr.
Richmond's paper there are some short field notes made by Dr. Abbott.
On various occasions Dr. Oberholser, notably in his well known paper,
"Descriptions of One Hundred and Four New Species and Subspecies
of Birds from the Barussan Islands and Sumatra" (1912), and Mr.
Riley have published descriptions of new forms from the Abbott
collection, but no comprehensive paper on the collection as a whole
has ever appeared. Mr. Riley at the time of his death was working
on a faunal list of the birds of these islands. It is unfortunate that he
did not live to complete it. Needless to say it has been an inestimable
source of valuable information in the preparation of this manuscript.

The next collection of birds from these islands was that of Dr. E.
Jacobson and Jonkheer W. C. van Heurn made in 1913 on Simalur and
its adjacent islands. Dr. G. C. A, Junge published on this collection
in 1936, describing 4 new races and including many extremely valuable
field notes made by the collectors. There is also a good deal of material
about the eggs of some of these forms, which were collected and
measured.

Due to difficulties with the local population, Dr. Abbott was not
given permission by the Netherlands authorities to visit Siberut and
Sipora. Later in 1924 an expedition from the Raffles Museum, partly
financed by funds given by Dr. Abbott, visited these islands and a
duplicate set of these birds was given to the United States National
Museum. This collection was reported on by Messrs. F. N. Chasen
and C. Boden Kloss in 1926. They described 11 new forms. In 1927
Mr. Riley described 3 more forms from the same collection.

So far as can be discovered the next ornithological collecting on these
islands was not until 1937 when three collectors were in the field. Miss

Fig 1 Map "i Wat Suraatni l-lnrnf-, -.|mumj» depth contours,

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 311

Barbara Lawrence made a collection of 31 species and subspecies from
Nias for the Museum of Comparative Zoology at Cambridge, Mass.
Mr. J. J. Menden made a collection on North and South Pagi, part of
which is now at the same Museum. The third collection was that of
Dr. J. K. de Jong and a worker from Buitenzorg, Saiin, who spent two
months on Enggano. The first two collections have not been worked
up, but the Enggano birds were reported on by Dr. Junge (1938), add-
ing 5 species to the list of birds found on that island. In June 1939 I
spent a few days on Nias and secured specimens representing 45 species
and subspecies. This material has been written up by de Schauensee
and myself (1940), and four new forms were described.

Altogether some two hundred twenty species and subspecies have
been described from the islands of which the present paper recognizes
one hundred fourteen forms. The total list of the bird fauna of this
small archipelago runs to two hundred eighty species and subspecies
including migrants. With the exception of three races I have been
able to see specimens of all the endemic forms occurring in this group.
These three races are: Spizaetus cirrhotics vanheurni and Eurostopodus
macrotis jacobsoni from Simalur, and Coraeina striata kannegieteri
from Nias. Present conditions indicate that collecting will be impos-
sible on these islands for some time. However, I think that except for
the recording of more migrants and the extension of range of some
known forms the present list should stand for some time to come. I
do hope that in the future there will be an opportunity for more
biological field work to be done on the birds of these islands. Un-
doubtedly they serve as a splendid natural speciation laboratory.

GEOGRAPHY OF THE ISLANDS

The west Sumatra Islands lie in a chain for a distance of approxi-
mately six hundred miles paralleling the western coast of the island of
Sumatra. This chain runs roughly from northwest to southeast ex-
tending from Lat. 3°N„ Long. 95°35'E., to Lat. 5°30'S., Long.
102°24'E. The islands have never been carefully explored by a
geologist. (See map, fig. 1).

The original appearance of these islands is presumably due to the
same phenomena which caused the rise of the Barussan Mountains in
western Sumatra. Brouwer (The Geology of the Netherlands East
Indies, 1925, p. 2) has this to remark about these mountains:

"Sumatra lies along the axis of a great geanticline. Stretching along
the western part of the island, near to the coast are the Barissan [sic]

312 bulletin: museum of comparative zoology

Mountains which give to this side of the island a rough and rugged
character. . . . The present Barissan Mountains are associated with
late Tertiary and post-Tertiary mountain-building. Considerable vul-
canism occurred during a great part of the Tertiary period and even
yet there are many active volcanoes." Later (p. 84), he says: "The row
of islands to the west of Sumatra is known for its frequent and intense
earthquakes, but volcanoes are entirely lacking."

Professor G. A. F. Molengraaff in an interesting article ("Modern
Deep-Sea Research in the East Indian Archipelago", Geog. Journ.
57, No. 2, Feb. 1921, p. 95.) has a good deal to say about the forma-
tion of the west Sumatra Islands. His discussion of folding (p. 108)
follows :

"As soon as the upper portions of the folds which develop at a cer-
tain depth approach the Earth's surface and the majority of the rocks
under diminished pressure can no more be folded without being frac-
tured (van Hise's zone of fracture), the continuity of the strata will be
broken and the culminating portions of the elevating anticlinal axes
will be fractured and show at the surface as isolated portions or blocks,
their extent and shape being greatly dependent on the geological
structure and the differences in rigidity of the composing rocks. This
may suffice to explain why an elevating submarine ridge formed by
an anticlinal axis will appear at the surface as a row of blocks, i.e.
islands separated by deep channels.

"In the great geosynclinal area between the continents of Asia and
Australia one arc of folding belonging to the Alpine system, and known
as the Malay arc, appears to originate from the Burma arc, . . . and
can be followed . . . frcm the extreme north-western end of Sumatra,
through this island and the island of Java. . . as far as the Banda sea.

"In its western section where it borders on the Indian Ocean, as in
the central portion of the island of Sumatra, this arc now consists of
two major folds, as is illustrated by an ideal cross section (Fig. 2)
from the Indian Ocean towards the stable portion of the Sunda land.
It would show the following sequence:

1. Indian Ocean.

2. Sunda trough, first geosyncline.

3. Range of coastal islands girdling the west coast of Sumatra in-
cluding the Mentawei islands, first geoanticline.

- 4. Mentawei trough and corresponding trough-shaped depths,
second geosyncline.

5. Non-volcanic and volcanic mountain ranges of Sumatra and
Java, second geoanticline.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 313

6. Tertiary terrain, folded in late Tertiary and early Pleistocene
time, now practically stable, third geosyncline.

7. Stable Sunda land including the Sunda shelf."

wsw. />/ ^^ ^ *£A LCVEL ENE

1. 2. 3.4. 5. <o. 7.

Fig. 2. Schematic representation of late Tertiary, Pleistocene, and post-
Pleistocene crustal movements in the western section of the Malay geosynclinal
area. (After Molengraaff) The numbers refer to those listed above.

The northern group of islands consists of Simalur and its outliers.
Simalur is about fifty-five miles long, varying in width from five to
fourteen miles. It lies about sixty-five miles from the coast of Sumatra.
It is a rocky well-wooded island, the main mountain of which, Sibalur,
rises to 625 metres in height. Simalur is thinly populated and there are
few gardens or clearings. Besides raising sago, coconuts and vegetables
for their own subsistance, the main industry of the people is raising
water buffalo. The principal village is Sinabang in the southeastern
part of the island.

Simalur is outside the two hundred metre line which curves out from
the Sumatran coastline and envelops the Banyak islands to the south-
east. Simalur lies on its own two hundred meter bank which encloses
several nearby islands. Northwest lie the Kokos or Sa Laut Islands,
two small flat islets which are the most northerly extensions of the
entire archipelago. These two islets are two miles by one and a half
miles, and one thousand yards in diameter respectively. They are
connected with Simalur twenty-five miles to the south east by a sub-
terranean bank less than thirty-six metres in depth. They are mainly
covered with coconut trees and the only collectors to visit them were
Jacobson and van Heurn who spent a day there, August 21, 1913.
They found two forms on Kokos, Caloenas nicobarica nicobarica and
Eudynamis scolopacea simalurensis. Presumably other birds, particu-
larly migrant shore birds, occur there from time to time.

The only other islets which have been collected on in the immediate
vicinity of Simalur are Djawi Djawi, Lugu, and Pulu Pandjang in
Sinabang harbor, and Siumat, six miles off shore in an easterly di-
rection. Siumat is about two and a half miles long with some original
forest as well as coconut palms. Both Dr. Abbott and Jacobson and
van Heurn collected there getting three common species of small island

314 bulletin: museum of comparative zoology

birds: Sterna dougalli bangsi, Chalcophaps indica indica, and Halixetus
leucogaster.

South of Simalur by about fourteen miles are two small islands
separated from each other by a strait one and one quarter miles wide.
They are sometimes called the Tapah islands. Both are small low coral
islands covered with original forest and uninhabited. Lasia, the north-
ernmost is smaller, less than two miles in length, and surrounded by
fringing reefs. Babi, the larger, is about seven miles in diameter and
nearly round. Both islands lie on a shelf separated from the neighbor-
ing islands to the north, east, and south by depths greater than two
hundred meters. The only collector to visit Babi and Lasia has been
Dr. Abbott. Together these two small islands have three endemic
races not found on any of the other islands in the group. These are :
Psittacula alexandri major, Coracina striata babiensis, and Hypothymis
azurea abbotti.

The Banyak Islands lie about thirty-eight miles off the coast of
Sumatra, and forty-five miles south-east of Simalur. Bangkaru, the
most westerly, is thirty miles east of Babi. The principal islands of
the more than fifty islets and reefs in the group are : Tuangku, seven-
teen miles long by five miles wide, Bangkaru with an area of twenty
square miles, and Ujung Batu, a small narrow island. These islands
are decidedly hilly and well forested. A mountain on Bangkaru,
Amintolan, reaches a height of three hundred three meters. There is
a hill at the south end of the island. In between the land is very low
so that at any distance the appearance presented is that of two islands.
Tuangku has a mountain of three hundred thirteen meters on the north
coast. The eastern coast is low and covered with mangroves. There is
a small population on these islands of which more than half (536 in
1911) live in Tuangku.

The only collections from the Banyak islands have been those of
Dr. Abbott who visited Tuangku and Bangkaru. Richmond (1903,
p. 485) notes that more species were seen on Tuangku than Dr. Abbott
had found on Simalur. However, Abbott noted that "no large parrots,
hornbills, or barbets were seen or heard, and no drongos or orioles
were noticed."

The largest island of the group is Nias which is seventy miles long
and from twelve to twenty-two broad. It lies twenty-eight miles south
of the Banyak islands and about fifty-six miles south west of Ujung
Singkel, a projecting point of the Sumatra coastline. Nias lies on its
own two hundred meter shelf. The nearest point of contact is with
that part of the Sumatran shelf which extends south of the Banyak

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 315

islands. The elevation of Nias reaches a height of eight, hundred eighty-
six meters. In general the central part of the island is rather high
forming an irregular plateau which extends down on all sides gradually
to the coast. The general impression is of a drowned island with an
irregular sloping outline surrounded by outlying submerged rocks and
coral reefs in all directions. Further out from the shore are many
small islands such as the Hinako group which lie on a ten fathom bank
extending out from the west coast of Nias. Presumably most of these
islands have been connected with Nias at one time or another.

Nias has the largest population of any of the islands numbering
about one hundred thousand at the last census. The main centers
are in the southern part of the island where there are many large
villages. This southern part of the island has been rather heavily
deforested with the result that original forest is now found only in a few
places on the hill tops. The northern part of the island on the other
hand is populated by a somewhat less aggressive people whose culture
apparently does not demand the construction of the giant houses found
to the south.

In this part of the island much original forest remains and there are
presumably several new records to be added to the bird list of the island
as many fewer collectors have worked here. Nias has been by far the
best collected of all the islands due to its size and accessibility. Until
1942 it was the only one of the islands which was visited regularly by
an inter-island steamer sailing from Sibolga to Goenong Sitoli every
two weeks. The main trade of the island was in pigs which were much
in demand in the Chinese-populated centers on Sumatra.

Southeast of Nias by about forty-five miles lie the Batu Islands.
These consist of three large islands, Pini, Tana Massa and Tana Bala
surrounded by more than twenty smaller ones. Besides these three
large islands, Tello and Lago are the only islands that have been visited
by collectors. Tana Massa is twenty -seven miles long by five miles
broad. Tana Bala is twenty-two miles long by seven broad. Pini is
twenty miles long and six broad. All these islands are rather low and
without distinguishing features. The population is small and the origi-
nal forest largely remains. Tello, less than two miles long, is the seat
of administration and the most populated island in the group. Lago is
a small coconut covered islet three miles northeast of Tana Massa.
Most of these islands are surrounded by reefs sometimes five or six
miles out in all directions. The whole group lies within the two hundred
meter line which curves out from the Sumatra mainland and continues
southeast in a narrow belt inclosing the islands below the Batu group.

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The name for those islands to the south of Tana Bala is Mentawi.
This includes Siberut, Sipora, North and South Pagi, and the adjacent
small islands. North and South Pagi by themselves are called the
Pagi or Pagai islands. All four of these islands are inhabited, hilly,
and evidently of volcanic formation. Although these islands lie on a
relatively shallow bank connected with Sumatra to the north, there is
a definite gap directly to the east towards the nearest part of the Su-
matran mainland. Here there is the deep Mentawi basin reaching a
depth of over sixteen hundred meters. Thus except for the one link to
the north there is no evidence to indicate that the Mentawi group has
ever been connected with Sumatra.

Siberut, thirty miles to the south of Tana Bala, is sixty miles long by
fifteen to twenty-four miles in breadth. It is heavily wooded with ex-
tensive marshes along the coast. There are about eight thousand in-
habitants at an extremely primitive cultural level. The island is of all
the group perhaps the least known and least explored. The only
collecting of birds and mammals has been that of Messrs. Kloss and
Smedley in 1924.

Sipora is a small densely overgrown island about thirty-three miles
long. The low-lying land is extremely marshy, so much so that in
former years the best communication into the interior was by canoe.
The highest point on the island is three hundred thirteen meters.
Sioban, the main village is on the eastern shore and is a small copra
shipping port.

Both North and South Pagi present a slightly different picture from
Siberut and Sipora. The Pagi islands rise more steeply from the sea,
so much so that the ten fathom curve runs very close along the shore.
There are a few villages principally along the east coast but the popu-
lation is not large. Apparently the islands are principally covered with
original forest. North Pagi lies about twelve miles south of Sipora.
There are very few inhabitants and little is known of the islands.
Except for the collection made by Dr. Abbott and Mr. Kloss in 1902
there has been only the short trip of Mr. Menden in 1937. It is unfor-
tunate that no field notes were made on these two visits, for undoubt-
edly the Pagi Islands are extremely interesting from the faunistic and
geological point of view.

Twelve miles southeast of South Pagi there is a small coconut-cov-
ered island called Sanding which apparently is little more than a wide
exposed reef. It has never been visited by a scientist. South of Sanding
the two hundred meter curve ends. Some fifty-five miles southeast
of Sipora there is a small pinnacle called Mega, two miles long and

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 317

surrounded by a fringing reef. It is low and densely wooded and rises
from the one thousand meter curve. This island too has never been
scientifically explored.

Enggano, the most isolated island of this archipelago, lies about one
hundred eighty miles south east of Sipora. It is about seventy miles
from the nearest part of the Sumatra coast. The island is twenty miles
long and approximately ten broad. A range of hills runs from north-
west to southeast through the center of the island, expressing thus the
direction of upthrust throughout the group. Enggano lies entirely on
its own shelf, the surrounding seas reaching a depth of three thousand
meters. Enggano is heavily wooded and the interior is so marshy and
difficult to penetrate that communication between the villages is said
to be by boat. It is sparsely inhabited, the 1919 census having shown
only four hundred seven persons on the island. The main trade of the
island is in copra, and the principal settlement is on Aduwe, one of the
three little islands in Enggano Bay. This group is called Pulu Dua.

FAUNAL LIST OF THE BIRDS

In the following list of the birds of the west Sumatra islands, the
arrangement of the families has been that of Wetmore (A Systematic
Classification for the Birds of the World, Smith. Misc. Coll., 99, 7,
1940, 11 pp.), while the arrangement of the species within the families
has been that of Chasen (1935). All measurements are in millimeters,
the wing pressed flat against the ruler. All weights of small birds are
in grams. Where used, wing-tail ratio refers to the percentage of the
length of the tail as compared to that of the wing. Wing tip index
refers to a similar percentage of the length of the shortest primary as
compared to that of the longest primary. Color notes follow Ridgway.

Of the total of seventy-five families found on Sumatra and generally
in the vicinity of these islands, forty-six families are represented from
this archipelago with certainty. Of the missing families some members
of the Phaethontidae, Sulidae, Rostratulidae, Indicatoridae, and Arta-
midae, may be expected to turn up on these islands in the future.
Notable, however, is the lack of representatives of the Megapodiidae
and Phasianidae, which do not occur there and presumably never have
been found on these islands.

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Family SULIDAE, Boobies, Gannets

The only record for this family is a dubious one which I have not
included in my definite list of species of the islands. There is every
reason to suppose, however, that boobies may occur around these
waters from time to time.

? Sula sula rubripes Gould

Chasen (1935) records this form from the Pagi Islands. I can find
no record of a specimen from this area except that in the British
Museum Catalogue (Vol. XXVI, p. 434, 1898), which lists a juvenal
male from Nassau Island, south of Sumatra, August (Dr. Coppinger),
Voyage of H.M.S. ''Alert". In the report on this voyage (British
Museum, 1884), Dr. Coppinger notes in the summary of the trip (p. 3)
that the boat touched at Nassau Island in the Danger Group northeast
of Samoa, after leaving Tahiti on August eighteenth. There is no
mention of a stop near south Sumatra during the later part of the trip,
although a stop was made at Singapore in November of the following
year. More evidence is needed to prove the existence of this form off
the west Sumatran islands.

Family PHALACROCORACIDAE, Cormorants

The only record for this family is from Nias. There is no indication
as to whether or not the Little Cormorant is a resident in the islands .

1. Phalacrocorax niger (Vieillot)

Listed by Biittikofer (1896) from Nias on the basisof Nieuwenhuisen
and von Rosenberg's report (1863). This species has not since been
recorded from any of the islands.

Family ARDEIDAE, Herons

2. Ardea sumatrana sumatrana Raffles

Recorded from Simalur, Babi, Tuangku, Nias, and the Pagi Islands.

The record for the Pagi Islands is by Chasen and Kloss (1935).
I can find no specimen on which it is based. A nest with two well-
grown young was found by Abbott on Simalur in November. A female
adult in the collection of the National Museum was secured on the
same island, October 23, 1902. Wing, 434.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 319

3. Ardea purpurea manilensis Meyen

Found on Xias, Pini (Batu Islands), and Enggano.

An immature specimen was taken on Enggano by Modigliani, May
19, 1891. An adult female collected by Dr. Abbott, November 5, 1904,
on the same island was labeled "iris yellow". Wing, 351.

4. Butorides striatus spodiogaster Sharpe
Synonym: Butorides striatus sipora Chasen and Kloss

Occurs on Simalur, Nias, Siberut, Sipora and North Pagi.

Three specimens from Simalur, Nias, and Sipora measure: wing
(worn), c? 182.5, 9 170.5, o 180.5; tail, d* 63, 9 63.5, o60; culmen,
J1 63, 9 60, o 60.

Oberholser (1912, p. 1) named two races, actophilus from North
Pagi Island and icastopterus from Simalur, on the basis of larger size
and darker color. The types measure :

actophilus 9 ad., icastopterxis c? ad.,

January 4, 1903. December 10, 1903.
wing 196 191

tail 70 68

culmen 66.5 65.5

Another female from North Pagi, collected December 31, is also
large (wing 190, tail 67.5, culmen 62.5). These birds presumably
represent a northern race found in Siam and China which migrates
during the winter to the southern islands. Chasen and Kloss (1926)
have described sipora as being darker on the neck than javanicus,
although similar in size. The three small specimens agree with this
description, as well as appearing darker on the back and scapulars
than a large series of javanicus. However, they are unfortunately in-
separable from spodiogaster Sharpe (Bull. Brit. Orn. CI., 3, 1894,
p. xvii) of the Andamans and Nicobars. A female from Little Nicobar
collected March 3 has a wing of 180 and agrees well in color with the
west Sumatra islands.

The names carcinophilus and carcinophonus Oberholser (Journ.
Wash. Acad. Sci., 14, 1924, p. 294) and abbotti Oberholser (U. S. Nat.
Mus. Bull. 159, 1932, p. 14) are synonyms of javanicus.

5. Butorides striatus actophilus Oberholser
Synonym: Butorides javanicus icastopterus Oberholser

Migrant from the Asiatic mainland recorded from Simalur (icastop-
terus) and the Pagi islands in December and January.

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6. Ardeola bacchus (Bonaparte)

Recorded by von Rosenberg from Nias (Biittikofer, 1896) and a
sight record (Richmond, 1903) for Simalur.

7. Egretta garzetta nigripes (Temminck)

There is a record of this species for Nias in Biittikofer 's list based
on Nieuwenhuisen and von Rosenberg (1868).

8. Egretta eulophotes (Swinhoe)

A rare migrant from China where it has been seriously reduced by
plume hunters. Recorded from Sipora by Chasen and Kloss (1926)

9. Demigretta sacra sacra (Gmelin)

Occurs on Simalur, Tuangku, Nias, Pulo Tello, Sipora, the Pagi
islands, and Enggano.

Two specimens collected by Dr. Abbott on Simalur and Pulo Mirbau
off Enggano measure: 9 ad., wing 262, tarsus 68; c? im., wing 266.5,
tarsus 69.5. The immature bird is in the mottled juvenal plumage
for the white phase discussed by Mayr and Amadon (Amer. Mus.
Novit. No. 1144, Oct. 13, 1941). Curiously enough, the wing measure-
ment of the adult female is smaller than any of the series of three
hundred seventy-eight specimens measured by them.

Of the thirty-three known adults collected on these islands, nineteen
are described as being in the grey phase, thirteen in the white phase,
and one (Salvadori, 1892) in the mottled phase.

The reef heron probably nests on the small islets off the coasts of
the larger islands. Junge (1936) records a set of eggs taken July 3
on an islet in Sinabang harbor, Simalur.

10. Mesophoyx intermedia intermedia (Wagler)

The single record for this species is for Nias, Biittikofer (1896)
founded on von Rosenberg's list.

11. Nyticorax nycticorax nycticorax (Linnaeus)

Found on Nias (Biittikofer 1896) and Enggano. Two males and
a female from the latter island measure: wing, c? 270, 287, 9 281.
One male is in an erythristic state of plumage similar to that recorded
for N. n. hoactli by van Rossem (Auk, 53, 1936, p. 322). The cheeks

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 321

and nape are tinted with buff. The upper part of the back is dark
bronzy brown shading posteriorly to "Carob brown". The wings and
tail are greyish buff. The second male approaches this but is paler and
more greenish on the back. The female is nearly normal. The occur-
rence of erythrism in this species indicates that Xycticorax caledonicus
may be a form in which this type of mutation has been established as
a dominant. It will be interesting to observe the apparently new
colony of caledonicus breeding in East Java, (Hoogerwerf, Bull. Raffles
Mus., 12, 1936, p. 120), to see if the birds hybridize with nyciticorax.
The colors of the soft parts are recorded as follows : iris straw yellow;
bill black, base and lores greenish (cT), lores slaty (cf), and base
pinkish fleshy ( 9 ) ; feet pale greenish yellow (d\ 9 ), straw yellow (cf ).

12. GORSACHIUS MELANOLOPHUS MELANOLOPHUS (Raffles)

Two females collected by Dr. Abbott on Enggano seem to be the
only record for this species in the west Sumatra islands. One specimen,
collected November 22, 1904, is in immature plumage. The soft parts
are as labelled: "iris pale greenish yellow; feet brownish green; bill
black above, pale brown beneath."

13. Ixobrychus sinensis sinensis (Gmelin)

An immature male collected by Abbott on Nias, March 28, 1903, is
the second record for that island.

14. Ixobrychus cinnamomeus (Gmelin)

A single male is recorded from Enggano June 22, 1936, by June
(1938).

15. Dupetor flavicollis flavicollis (Latham)
Recorded from Xias by von Rosenberg.

Family CICOXIIDAE, Storks

The first member of this family found on these islands proves to
belong to a widely-distributed form throughout the Malay area.

16. Dissoura episcopus stormi (Blasius)

North Pagi. An adult male and an immature female taken on North
Pagi in December by Menden are in the collection of the Museum of

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Comparative Zoology. They are the first specimens of this family to
be recorded from the west Sumatra islands. The male specimen has
the following notes on the colors of the soft parts : "iris brown, bill and
feet red" (but basal third of bill is black in the skin). The immature
has the colors noted as: "iris dark brown, feet red, bill red-brown"
(not black basally in skin).

I am indebted to Mr. James L. Peters for the identification and the
information on the labels.

Family ANATIDAE, Ducks and Geese

The Javan Tree Duck is the only species recorded on the west
Sumatra islands so far. It is a wandering species throughout the
Greater Sunda islands. Very likely Dendrocyqna arcuata occurs here
from time to time, while the absence of Anas castanea gibberifrons
from this area is definitely puzzling, (see Ripley, Auk, 59, 1942, p. 98).

17. Dendrocygna javanica javanica (Horsfield)

Biittikofer (1896) lists this as the probable race encountered by
von Rosenberg. When I was on Nias in June, 1939, I was told that
ducks were common on Nias at certain seasons.

Family ACCIPITRIDAE, Hawks, Eagles

Sixteen members of this family have been recorded from these
islands of which ten are presumably residents and six migrants. Five
of these resident forms are presumed to be identical with those found
commonly throughout the Malayan area. Of the remaining five,
Spizaetus cirrhatus vanheurni is described as being smaller than its
nearest relative. The other four forms belong to the very plastic
Spilornis cheela "rassenkreis". The two races from the northern
islands, abbotti and asturinus resemble birds from the Andaman and
Nicobar Islands. The race from the Batu Islands has been united
with the population from south Sumatra. The southernmost race,
that from Sipora and the Pagi islands has characters which markedly
resemble those found in Java, Bawean and Celebes. In all these Spi-
lor?iis forms there are marked color differences.

18. Pernis apivorus ptilorhynchus (Temminck)
Siberut (Chasen and Kloss, 1926, p. 279.).

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 323

19. HALIASTUR INDUS INTERMEDIUS Blyth

Found on Simalur, Babi, Tuangku, Nias, and the Pagi islands.

Junge (1936) notes that this bird was found nesting on Simalur in
May in a coastal grove of casuarinas at a height of 20 m. above the
ground. An immature bird was collected at the spot.

20. Accipiter badius poliopsis (Hume)

An adult female collected on November 20 is recorded from Nias
by Buttikofer (1896).

21. Accipiter soloensis (Horsfield)

Taken by Abbott on Simalur in October and January. Chasen
(1935, p. 71) lists this species from Nias on the basis of Biittikofer's
record of Accipiter poliopsis. From the description of the specimen,
however, it is impossible to be sure that he did not have a specimen of
poliopsis.

22. Accipiter trivirgatus trivirgatus (Temminck)

Recorded for Nias. A male collected March 26 is in extremely worn
plumage showing no sign of new feathers. A female collected on the
same date is also very worn, but there are a few new feathers coming
in on the sides of the neck and nape. Another female collected March
23 is renewing some of the tail feathers and the inner primaries. A
few of the nape feathers and wing coverts are new also. Soft parts:
iris, cf , "bright yellow"; 9 "orange yellow"; feet, d* "yellow", 9
"pale yellow" ; bill, 9 "black above, leaden beneath". The stomach
of the female (March 23) contained lizards.

23. Accipiter virgatus gularis (Temminck and Schlegel)

Simalur, Siumat, Lasia, and Nias. Four immature females from
Simalur and Siumat and Lasia measure: wing, 182, 185, 185.5, 188.
Two of the specimens had been eating small birds. This species has
been collected from November 25 to February 12.

24. Spizaetus nipalensis alboniger (Blyth)

Found on Pulu Asu, off Simalur, Bangkaru (sight), Nias, and North
Pagi. Chasen's record (1926) for S. cirrhatus limnaetus on the Pagi
islands presumably referred to this species. Three immature males

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from Pulu Asu, Nias, and North Pagi measure: wing, 296, 308, 310.
They were taken December 25, March 13, January 6. Soft parts:
iris, "greyish-yellow, lemon yellow"; feet, "dull lemon yellow"; claws,
"black".

25. Spizaetus cirrhatus vanheurni Junge

This race was described by Junge (1936) on the basis of five speci-
mens from Simalur. The author notes that the birds are much smaller
than S. c. limnaeetus, but failed to compare them with S. nipalensis
alboniger. In measurements and in coloration, these birds seem very
similar to alboniger.

26. Ictinaetus malayensis malayensis (Temminck)

Listed by Biittikofer from Nias on the basis of Nieuwenhuisen and
von Rosenberg (1863). This mountain forest species has never been
encountered since on the west Sumatra islands.

27. Haliaeetus leucogaster (Gmelin)

A bird of the small islands off Simalur; Asu, Djawi djawi, Babi-
ketchil, all in Sinabang bay, Siumat, and Lasia, Babi, Bangkaru,
Tuangku, Nias (vide Chasen, 1935), Sipora, and the Pagi islands.

Two males were collected on Asu and Siumat in December. A nest
was seen by Abbott on Siumat on Christmas Daj^.

28. Spilornis cheela abbotti Richmond

Simalur. This well-marked race was collected by Abbott (1901) and
van Heurn (1913) at all months of the year. The wings of nine males,
six females, and one unsexed, measure: cfcf 330-362, 9 9 319-359,
o328.

Soft parts: "iris yellow; bill horn blue, brownish at tip; cere orange
yellow; inside of mouth leaden; feet dirty yellow."

Stomach contents: snakes, small crab, centipede, lizards.

Apparently an abundant bird on Simalur, this race is very distinc-
tive in appearance. The amount of spotting on the under parts is
variable. In some specimens it extends up onto the breast; in others it
is confined to the abdomen. The marking on the breast and throat is
far more noticeable than in the paler forms, due to the contrast be-
tween the dark blackish brown and reddish brown bars.

No pale individuals have so far been taken. Meise's recent revision
of this genus (J.f.O., 87, 1, 1939, p. 65) indicates that all these popula-
tions should be considered races of cheela.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 325

29. Spilornis cheela asturinus A. B. Meyer

Nias. Three males collected by Abbott measure: wing, 291, 291.5,
299.5 All three birds are in worn plumage with a few new feathers
appearing among the greater wing coverts (one male) and secondaries
(another specimen). Soft parts: iris bright yellow, feet dirty orange
yellow, cere yellow. Stomach contents: snakes and small centipedes.

A pair collected by myself for the Academy of Natural Sciences
weighed, d* 15 oz., 9 1 lb. 4 oz.

This species is common on Nias, where it may be found perching in
a high tree on the edge of an open field or moving slowly over the
gardens or hedgerows in search of its prey.

From minimus of the Northern Nicobars, which it closely re-
sembles, this form differs by the finer barring and spotting of the
lower underparts.

30. Spilornis cheela batu deSchauensee and Ripley

Tello, Tana Massa. Meise (I.e. 1939) in his splendid revision of
the genus Spilornis lumps Malay Peninsula, Sumatra, Java, and Bali
birds all under bassus, giving for wing measurements, 348-394. I am
inclined to believe that the name bido should stand for Java and Bali
birds, which are large (384, 391), and dark, with distinctly blackish
throats. Granted the last character is allelic in these birds, still it
seems to be quite constant in the Java population.

The birds from south Sumatra, on the other hand, are uniformly
small, distinctly paler in tone than bido, although in this last character,
they do not differ from typical bassus.

When de Schauensee and I described batu (1940, p. 401.), we had
only north Sumatran material available for comparison. An adult
from Tana Massa in the National Museum's collection (sex indet.
no. 179622) does not bear out our contention that this is a pale race.
It is as dark in tone as any specimen from Sumatra or the Malay
Peninsula. However, it does agree with the Academy's two birds in
being small. Four males, three females, and one sex indet. from the
east coast province of Sumatra agree with these Batu Island birds in
size and color. They come from the Katenan River, the Siak and
Little Siak rivers, Makapan, and Pulu Padang. Their measurements
are: <?d\ wing 229, 331, 346.5, 352, tail 204, 206, 217 (molt), 220;
9 9 , wing 340, 350, 354, tail 218, 221, 221 ; o wing 346, tail 215.

As can be seen, these measurements are all below 360 for the wing
and 230 for the tail. North Sumatra birds, on the other hand, measure

326 bulletin: museum of comparative zoology

much larger. Six males and four females from Atjeh measure: wing,
c? 360, 365*, 365, 388.5, 390*, 410.5; 9 371*, 375, 380, 388; tail, <? 231
255*; 9 253*. Similarly seven Malay Peninsula birds measure wing,
cf 355 (1, worn) - 397 (374); 9 385^07 (392); & 230-259 (244.6),
9 242-264 (251.6).

Under these circumstances, I prefer to restrict the name bassus to
birds from the Malay Peninsula and Sumatra south to the province
of Oostkust. I propose also to unite birds from Oostkust south with
Batu Island birds as batu, modifying the description of that race
(I.e. 1940,) to read: Pulo Tello and South Sumatra birds "differ by . . .
smaller size".

31. Spilornis cheela sipora Chasen and Kloss

Sipora and the Pagi islands. A single female from Sipora has a
wing measurement of 310 and the soft parts are indicated as: "iris
and facial skin yellow; bill pale grey; feet yellow".

This race is distinctive in having the feathers of the crest largely
white with black tips giving a noticeably piebald effect. The breast
is solidly colored, smoky bronzy black. In its coloration, sipora shows
a close relationship to the black-cheeked strain of Spilornis which con-
tinues southward with bido of Java, baweanus of Bawean, and rufipectus
of Celebes.

Family FALCONIDAE Falcons
Two species have been recorded from the islands, both migrants.

32. Falco peregrinus calidus Latham

A migrant collected on Nias and Enggano and probably seen on
Simalur (Richmond, 1903). A male from Nias collected in March has
a wing of 306.5. A female from Enggano collected in November
measures 376. The latter specimen is molting the greater wing coverts.
The soft parts of the male are given as: "iris dark brown; eyelids
yellowish grey; feet bright yellow".

33. Falco tinnunculus subsp. ?

A winter migrant recorded from Nias in November by Biittikofer

(1896).

• U.S.N.M. birds.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 327

Family RALLIDAE, Rails, Coots, Gallinules

Four species occur on the west Sumatra islands all apparently
identical with wide-ranging Malayan forms.

34. Rallus striatus gularis Horsfield
Synonym: Hypotaenidia striata reliqua Oberholser

Simalur. Two females collected in October and December are
identical with specimens from Java and Singapore. The soft parts are
marked as: "iris pale yellow brown, red; bill pinkish red, tip horn
brown; feet dull brownish purple". Found in the rice paddies.

(Rallus jouyi Stejneger is by no means a synonym of gularis. It is
a giant pale race. The male type and a female from Shanghai measure :
wing, cf 135, 9 134.5; tail, cf 50, 9 49; culmen, d" 43.5, 9 42.5.)

35. Rallina fasciata (Raffles)

Nias, Sipora, and Enggano. A male from Enggano collected in
November measures: wing 128.5. Soft parts: "iris and eyelids red; feet
vermilion."

36. Amaurornis phoenicurus javanica (Horsfield)
Synonym: Amaurornis phoenicura cleptea Oberholser

Found on Simalur, Nias, Tello, Siberut, Sipora, the Pagi islands, and
Enggano.

A common bird of the flooded rice fields and swampy meadows near
streams. Found also in the sago swamps on those islands which have
them. These birds have a harsh rattling call.

Four males from Simalur, three females from Nias, including the
type of cleptea, and four females from Sipora are inseparable on the
basis of size.

37. Porphyrio poliocephalus indicus Horsfield
Recorded from Nias by von Rosenberg (1878).

Family CHARADRIIDAE, Plovers

Four species have been found on these islands of which three are
migrants. The resident form has been separated on the basis of color,
a character which I fail to distinguish.

328 bulletin: museum of comparative zoology

38. Pluvialis dominica fulva (Gmelin)

A common migrant recorded from Simalur, Nias, Batu islands,
Siberut, and Sipora, September through April.

39. Charadrius peronii Schlegel
Synonym: Charadrius peronii chaseni Junge

A resident species recorded from Simalur and Nias. A female from
Simalur (M.C.Z. 178151) was collected in May. Junge described the
race chaseni from Simalur as being "darker on the upper parts, less
rufous on the head especially in the 9 9 . Also the rufous colour
behind the white neckcollar and on the sides of the breast is much more
pronounced than in p. peronii. The bill in chaseni is slightly heavier."

A female from Nias collected in February (two months earlier than
Junge's Simalur series) is a good deal darker than the Simalur female.
It is in much fresher plumage. A molting female from the Philippines
collected in January has new feathers as dark as the Nias bird, while
an October male from Mindanao is considerably darker than any other
bird in our series. As far as the character of the rufous breast band
goes, the two west Sumatra island birds stand midway in a series of
seven females. I cannot find any significant size difference in the bill.
I fail to see, therefore, how chaseni can be upheld.

40. Charadrius mongolus atrifrons Wagler

A migrant recorded from Simalur and Tello; this species probably
occurs on all the small beaches. Two females from Simalur collected
in December measure: wing, 130, 134.5.

41. Charadrius leschenaultii Lesson

A winter migrant. There are records for Simalur, Nias, Pini, Tello,
and Sipora from August through December. A male and two females
collected on Simalur in December measure: wing, cf 140, 9 140.5,
142.5.

Family SCOLOPACIDAE, Snipe, Woodcock, Sandpipers
Twelve species have been found on these islands, all of them migrants.

42. Numenius phaeopus phaeopus (Linnaeus)

A migrant taken on Simalur, Sipora, and Siberut. A female taken
on Simalur in November has the rump pure white. A male from

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 329

Sipora collected in February has a few spots of grey brown among the
white rump feathers. For the time being, I think it is better to call
these birds phacopus. There seems to be a good deal of intergradation
between the two forms.

43. NUMENIUS PHAEOPUS VARIEGATUS (Scopoli)

Reported from August till June on Simalur, Nias, Pini, and Tello.
There is a sight record of one or other of the races of whimbrel for the
Pagi islands.

44. Numenius arquatus orientalis C. L. Brehm

A migrant taken on Tuangku and Nias. A female from Tuangku
(January) measures: wing, 282.

45. Limosa lapponica baueri Xaumann

The single record of this migrant for the west Sumatra islands is
Xias, October (Blasius).

46. Tringa totanus eurhinus (Oberholser)

This winter visitor has been taken on Simalur, Nias, and Tana
Massa from September until May. A female from Nias collected in
March has a wing measurement of 158.

47. Tringa nebularia Gunnerus

A migrant reported from Simalur and Nias in August, September,
and October.

48. Tringa glareola Linnaeus

This winter bird was found on Simalur from January until April.

49. Actitis hypoleucos (Linnaeus)

The common sandpiper of the islands from August until April.
Recorded with certainty only from Simalur, Babi, Nias, Tello, and the
Pagi islands.

50. Arenaria interpres interpres (Linnaeus)

A female collected on Simalur in December has a wing measurement
of 148.5.

330 bulletin: museum of comparative zoology

51. Capella stenura (Bonaparte)

During migration recorded from Simalur, Nias, Sipora, and (sight
record) the Pagi islands. Two males and three females from Simalur
and Nias measure :wing, & 135, 138; 9 134.5, 135, 135.5. These birds
were collected during December, February, and March.

The Pintail Snipe is a common bird in these islands during the
winter, being found wherever there are recently drained rice fields or
along the edges of streams, sometimes in the short grass bordering
roads built through swampy areas.

52. Erolia ruficollis (Pallas)

A casual winter visitor, found on Simalur and Nias in December
and October.

53. Erolia testacea (Pallas)
Blasius (1901) records this migrant as taken on Nias in October.

Family BURHINIDAE, Thick-knees
A single resident species occurs throughout the greater Sunda area.

54. Orthorhamphus magnirostris (Vieillot)

Simalur, Babi, Tuangku, and Nias. A male from Babi collected in
January has a wing measurement of 291.5. Weight, 2.5 pounds. Soft
parts: "iris yellow, tarsi pale yellowish slaty, thighs yellow, claws
black, leaden at base". The Stone-plover feeds on crabs and is common
along sandy beaches and on exposed reefs at all seasons.

Family GLAREOLIDAE, Pratincoles
A single species of this family has been recorded twice as a migrant.

55. Glareola maldivarum Forster
A migrant from northern Asia recorded from Nias and Sipora.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 331

Family LARIDAE, Gulls, Terns

Five species have been noted as occurring on these islands. At least
one species is known to breed here, but there has been no observable
speciation in the case of this family.

56. Chlidonias hybrida javanica (Horsfield)

There is a record for Nias in December (Blasius, 1901), and von
Rosenberg (1878) includes this in his list of species met with from time
to time among the islands.

57. Sterna dougallii bangsi Mathews
Collected on Siumat in May.

58. Sterna sumatrana sumatrana Raffles

Simalur, Pandjang, Djawi-djawi, Siumat, Nias, and Enggano. Eggs
have been taken in May on Simalur and Djawi-djawi. An immature
male was taken off Nias by Abbott, October 30.

59. Thalasseus bergii cristatus (Stephens)

Mentioned by von Rosenberg (1878) as occurring in the seas about
these islands.

60. Anous stolidus pileatus (Scopoli)

Listed by von Rosenberg as seen from time to time among the
islands.

Family COLUMBIDAE, Pigeons, Doves

Twenty-one forms are recorded from the west Sumatra islands. Of
these seven belong to wide-spread species, common throughout the
Greater Sunda area. Thirteen of the fourteen other forms are endemic.
One is considered identical with Sumatran and Malayan subspecies.
Of the thirteen endemic races eleven differ markedly from their cor-
responding relative on Sumatra by larger size and rather different
coloring. Two forms do not differ in size. One, Treron fulvicollis
melopogenys, is poorly characterized and may not be a good race. The
other, Ducula aenea consobrina, differs from D. aenea only in color.

332

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The eleven well characterized forms may be tabulated as follows:

D. aenea M . ruficeps M. phasianella

T. curvirostra

En

ggano*

zzm

Mentawi
& Paei.

Xias

Simalur

BBZ2Z
i I i i

Sumatra

1. 2. 3. 4. 5. 1. 2. 3. 1. 2. 3. 4. 5. 1. -2. 3. 1. 2. 3. 4. 5.

M H difference in size.
V/J///A difference in color.

Fig. 3

In the above chart Sumatra birds are taken as standard. Their in-
dividual variation is taken as less than the arbitrary unit 1, within
which it is considered that local island populations may vary and still
be considered inseparable. The unit 2 is used to express those varia-
tions in size or color, either one of which by itself is still too slight for
separation.

Above this number the units used indicate the relative degree of
differentiation of the two factors, size and color, of the different island
populations.

61. Treron curvirostra haliploa Oberholser

Simalur. The type, an adult male, measures: wing 145.5; tail 92.5;
culmen 16.5. The soft parts are marked: iris orange, orbital skin yel-
lowish green, feet dull purple. Junge (1936) gives the measurements
of four males as : wing 146-151 ; tail 90-94; culmen 16-17.5. This race
differs from curvirostra by larger size. Thirteen males from Peninsular
Siam and Sumatra (harterti is a synonym) measure: wing 132-139;
tail 77-83.5; culmen 16-18.

Junge (I.e.) notes that a single male from Babi is quite yellowish on
the under surface. His description sounds very much like smicra from
farther south (see also Junge, 1938, p. 340.).

A bird of original forest.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 333

62. Treron curvirostra pega Oberholser

Nias. The type, an adult male, measures : wing 144; tail 88; eulmen
16. Four females measure: wing 134.5, 142, 143.5, 143.5; tail 75, 79,
79, 82; eulmen 16.5, 17, 17.5, 18. This form is larger than curvirostra
and differs both from that form and haliploa by being noticeably paler
on the under parts and by having more grey on the flanks. The soft
parts are recorded as: "iris bright ochreous ; orbital skin apple green,
pale bluish green; feet maroon red; bill yellow; cere red." Weight:
cf, 7 oz.

The female differs as the male.

Specimens with enlarged gonads were taken by myself in June.

63. Treron curvirostra smicra Oberholser

Tello, Tana Bala, Siberut, and Sipora.

The type, an immature male taken February 8 on Tana Bala, meas-
ures: wing 138.5; tail 73; eulmen 17. Four males and a female from
Siberut and Sipora measure: wing d" 141, 141.5, 145, 147.5, 9 148.5;
tail d\ 83.5, 84, 90, 9 86; eulmen & 17.5, 17.5, 17.5, 18, 9 18. Soft
parts: "iris yellow; bill pale green; cere olive; feet maroon red, crimson
lake".

This race is very bright yellow on the under surface. The breast
particularly is a rather rich shade of lemon yellow. It is the brightest
member of the species. On the upper surface it differs from its near
relatives by having the nape more distinctly yellowish green less
greyish green.

The female differs as the male.

Evidently this bird prefers a slightly different type of habitat than
either haliploa or pega, which are found on much larger islands. If
the specimen from Babi, one hundred and sixty miles to the north,
recorded by Junge, proves to belong to this race, it would seem to
indicate that smicra not only prefers a different ecological setting but
also suits its behavior to its location. Small island populations of
pigeons seem to be more subject to sporadic wandering movements
than their large island congeners.

64. Treron curvirostra hypothapsina Oberholser

Enggano. The male type, collected November 21, measures: wing
156; tail 92; eulmen 17. The soft parts are recorded as follows: "iris
vellow; eyelids dull green; bill jade green, base dull green; feet purplish
red."

334 bulletin: museum of comparative zoology

Two other males and a female measure: wing c? 149, 150, 9 141;
tail cf 86, 90.5, 9 85.5; culmen c? 18, 18.5, 9 16.

This again is a larger bird than curvirostra and much brighter yellow
on the under surface. It differs from smicra by being slightly duller on
the under parts, lacking especially the bright suffusion of yellow on the
breast found in the latter subspecies. On the upper surface, too, the
nape is more grayish-green, less tinged with yellow. Junge (1938)
records enlarged gonads in males collected in June and July.

65. Treron fulvicollis melopogenys (Oberholser)

Nias. The type and only available specimen is a female collected by
Dr. Abbott, March 18, 1903. It measures: wing 141 ; tail 85.5; culmen

16. Soft parts: "iris pinkish mauve; bill greenish blue; cere dull
crimson, eyelid gray with yellow edge; feet maroon red."

Two females of fulvicollis from Sumatra and Sarawak measure: wing
145, 150. This is a rather variable species particularly in color. Lack-
ing a larger series, however, I suppose that this single rather small
bright-yellow-throated specimen must be recognized.

66. Treron olax (Temminck)

Recorded by von Rosenberg (1878) as being on "all the larger
islands."

67. Treron vernans miza (Oberholser)

Simalur. The type, an adult female, collected November 22, meas-
ures: wing 152.5; tail 95; culmen 15. Soft parts: "feet deep red."
Three males measure: wing 156, 156.5, 158; tail 98, 98.5, 104; culmen

17, 17.

This is purely a size race. There are no color differences compared
to griseicapilla. Junge (1936) describes the plumage of a juvenal bird
collected in July. Eggs were collected in April, June, and July.

68. Treron vernans griseicapilla Schlegel
Synonym: Dendrophassa vernans mesochloa Oberholser
Synonym: Dendrophassa vernans polioptila Oberholser

Nias, Tana Massa, Tana Bala, Siberut, Sipora, North and South
Pagi, Enggano. The type of mesochloa, an adult female taken on Nias
March 18, measures: wing 145; tail 81.5; culmen 16.5. Soft parts are
recorded as : "iris pinkish-yellow; bill gray; cere dull green; feet maroon

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 335

red." A series of males and females from Nias measure: wing c? 141
155, 9 141, 146.

The type of polioptila, an adult female collected on North Pagi
January 1, measures: wing 148; tail S9.5; culmen 14.5. A series of
males and females from the Batu Islands, Sipora, North and South
Pagi and Enggano, measure: wing cf 148-156, 9 150-152.

These birds differ a good deal in size. Nias birds are more nearly the
size of Sumatran and Malay Peninsula birds, while specimens from
the islands slightly to the south, particularly North Pagi, tend to be
larger. In view of the overlapping measurements, however, it seems
to be much wiser to group these birds all under griseicapilla. None
of the local populations seem to be entitled to subspecific recognition,
either on the basis of standard deviation tabulation or of zoogeography.
Only the Simalur birds are apparently significantly larger. It is in-
teresting to note that a large series of griseicapilla from Sumatra and
the Malay Peninsula preserve far more uniformity in their wing
measurements (cf 144-151, 9 142-150) than do the birds from the
western islands. Presumably this is due to less possibility of swamping
of the island populations.

Junge (1938) records enlarged gonads in a male from Enggano taken
in June. This is a common bird in the islands. It is fond of thick
secondary growth in swampy coastal locations. As with all pigeons,
they are very fond of the fruit of various species of Ficus.

69. Leucotreron jambu (Gmelin)

Recorded from Nias in August (Blasius) and from the same island
by Hartert.

70. Ducula aenea consobrina (Salvadori)

Synonym: Carpophaga Vandepolli Biittikofer

Synonym: Muscadivores consobrina babiensis Richmond

Synonym: Muscadivores aeneus mistus Oberholser

Synonym: Muscadivores aeneus vicinus Riley

Simalur, Lasia, Babi, Tuangku, Nias, Tello, Tana Bala, Siberut,
Sipora, North and South Pagi.

The name vandepolli was founded on a discolored Nias bird (Junge,
1935).

The type of babiensis, an adult male from Babi, measures: wing
248; tail 146.5; culmen, 23. Two pairs from Lasia and Babi measure:

336 bulletin: museum of comparative zoology

wing, cf, 245, 247 (246.6); 9 237, 237; tail, <?, 144, 146.5; culmen,
<?, 24, 26.5.

The race mistus from Simalur was described as being decidedly
smaller than Nias birds. The type measures: wing, 231; tail, 135.5;
culmen, 22.5. The wing measurements of a series of males from
Simalur, 225.5-238 (231.1), as against a series from Nias, 230.5-244
(235.8), show far too much overlap to have any significance.

When the wing measurements of the three males of babiensis are
compared with those of twenty-five males from Simalur, Tuangku,
Nias, Tana Bala, Sipora, North and South Pagi, there is only a small
amount of overlap (2.5) :

wing average

adult c? d\ Babi and Lasia 245-248 (246.6)

" other islands 226.5-247.5 (234.6)

However, the range of variation is very great and accordingly I have
tested them by the formula of t used in the comparison of the means
of two small samples, (Simpson and Roe "Quantitative Zoology",
1939, p. 210). Using this I obtained a value for / on the wing measure-
ments of 1.9. This gives a probability of between .1 and .05 showing
that the deviation is not significant. This bears out the evidence of
a male from Babi recorded by Junge (1936) which measured well
within the range of consobrina.

The race vicinus from the Batu and Mentawi islands was founded
on color: "the breast and hind neck washed with much deeper
vinaceous-lilac." The color of these birds is subject to much variation
through dirt or grease on the feathers. When absolutely clean speci-
mens of each island are compared, however, color variations tend to
disappear.

From aenea these birds differ by lacking the pronounced vinaceous
wash on the head, cheeks, and sides of the neck. As a result, there is
less contrast between the gray head and the white margin around the
bill. The under parts generally tend to be more gray, less tinged with
vinaceous, although the latter tint is noticeable in dirty specimens.
The wider tail coverts seem to average darker in consobrina. The
measuring of wings in these birds is sometimes quite difficult, due to
the wing being incompletely pulled back by the skinners. This fact
undoubtedly accounts for some of the variations in measurements.

Molting specimens were collected in November and immature birds
in October, November, and January. Eggs have been taken on Simalur
in May and June. Immature birds are distinguished by a less well de-

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 337

fined division between the gray of the nape and the bronzy green of
the back. Also the marginal under wing coverts are edged with
rufous.

Soft parts of adults are given as: "iris, deep red; bill, pale leaden,
base dark leaden; cere, dull purple; feet, livia purple." Immature
birds are given as: "iris, sepia; bill, gray; cere, faintly brown; eyelids,
gray edged with pale pinkish brown; feet, purplish brown."

These big pigeons are common and conspicuous on most of the
islands, although on Nias they seem to be confined to the higher parts
of the island where some original forest is still found. The birds are
trapped a great deal by the people either as young taken from the
nest or by the use of bird lime (Malay "gutta"). Pigeons are a staple
of the native diet.

71. Ducula aenea oexothorax (Salvadori)

Enggano. Ten males and two females measure: wing, d", 247-254;
9 , 246, 249; tail, d\ 152-156; culmen, d\ 21.5-25.5.

This race differs from aenea by larger size and by lacking the vina-
ceous wash on the head. In most specimens the tail feathers are dis-
tinctly greenish instead of bluish iridescent in color. On the under
surface, however, there is an ill-defined band of strong vinaceous wash
on the breast. The under tail coverts are dull brownish green.

From consobrina this race differs by larger size and a slightly longer
tail proportionate to the length of the wing (62% compared to 57-
59%). The same characters of vinaceous breast band and more green-
ish under tail coverts apply in this case as well as with aenea.

Soft parts: iris, deep crimson; bill, pale leaden, base dark leaden;
cere, dull purple; feet, dull purplish red.

72. Ducula bicolor bicolor (Scopoli)

Simalur, Lasia, Babi, Tuangku, Nias, Tello, Lago, Pini, Sipora, the
Pagi islands, and Enggano. Specimens from Simalur, Babi, Sipora,
and Enggano collected in October, November, December, and Jan-
uary measure: wing, d\ 224-238, 9 , 223.5-231.5. An immature bird
was taken on Sipora in October. The soft parts are recorded as: iris
dark; bill leaden, black at tip; cere greenish; feet blue.

This is a bird primarily of the very small islands and reefs. They
follow the seasons flying about in small flocks in search of trees in
fruit.

338 bulletin: museum of comparative zoology

73. Ducula bicolor badia (Raffles)

Rosenberg (1878) records encountering this bird on Nias, the
Mentawi islands, and Enggano. It has not been met with since that
time.

74. Columba Argentina Bonaparte

Recorded from Simalur, Sipora, and South Pagi. There is a sight
record for Kokos. Males and females from Simalur and South Pagi
measure : wing, d\ 239-249.5; 9 227, 240. The soft parts are given as:
iris, bright red, yellow suffused with red; bill, greenish horny, dull
purple at base, apple green; cere, dull purple; tarsi, purple; toes, pale
purplish fleshy.

This is a wandering seasonal species not common in collections. It
probably occurs from time to time on all the west Sumatra islands.

75. Macropygia ruficeps simalurensis Richmond

Simalur. Two males and two females measure: wing, cf , 152.5
(type), 149; 9 , 144.5, 146; tail, tf, 164.5 (type), 169, 9 , 155, 164.5;
culmen, d\ 14 (type), 14; 9 , 13, 14.

As well as having certain differences in color noted by Junge (1936),
this race is slightly larger in size than sumatrana.

76. Macropygia phasianella hypopercna Oberholser

Simalur. The type, an unsexed bird evidently a female, measures:
wing, 172.5; tail, 177; culmen, 17. This bird is grease stained on the
upper back and rump. With only one immature specimen, it is im-
possible to determine the validity of this race. However, its resem-
blance to the Nias race is striking.

77. Macropygia phasianella modiglianii Salvadori

Nias. A male and two females measure: wing, cf, 184.5, 9, 183,
185; tail, 9, 184, 184.5; culmen, <?, 17.5, 9, 16, 17.

This race differs from emiliana of Sumatra and Java principally by
larger size. There seems to be a slight reduction also in the amount of
the irregular barring on the breast.

78. Macropygia phasianella elassa Oberholser

Siberut, Sipora, and North and South Pagi. The type, an adult
male from North Pagi, collected November 12, measures: wing, 179.5;

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 339

tail, 171 (molt); culmen, 20. The soft parts are marked: "iris, pink
with an inner narrow blue circle."

This race is distinguishable from the available specimens of modi-
glianii. In the male the upper parts are very similar but specimens of
elassa tend to have more distinct pale tips to the feathers of the back,
scapulars, and wing coverts, giving a slightly barred effect. On the
under surface, the iridescent purple sheen of the breast tends to be
confined to a rather distinct band, whereas in modiglianii, the color is
more generally dispersed. The color of the under side of the tail
feathers seems to be paler, also, and lacking in gloss.

In the female, the color differences are more apparent. The neck
and upper back are distinctly barred black on buff, giving an effect
of more contrast. The tail seems to be slightly darker. On the under
surface there is a distinct tendency to a blackish patch on the throat.
In the male, this spot is variable and sometimes absent. In the large
series of females examined, however, it is always 'present. The rest
of the under surface seems to be distinctly darker than in modiglianii.

79. Macropygia phasianella cinnamomea Salvadori

Enggano. This race represents the farthest step in a cline running
from north to south in the islands. The Nias population differs from
emiliana primarily by size. Farther south elassa has more distinct
barring, a condition which resembles that of immature birds. Also
there is a tendency towards reduction in the amount of violet iridescent
gloss spread over the plumage and the appearance of a blackish throat
patch. In cinnamomea this condition is further exaggerated. The iri-
descent gloss has disappeared. The plumage as a whole appears faded
and washed out. The black barring on the back tends to break down.
The primaries are pale rufous on their outer edges as in immature
specimens of the other races. The black throat patch is present as a
definite adult character.

An immature female is uniformly pale rufous brown all over, except
for the upper back which is irregularly shaded with black, and the
tail and inner margins of the primaries which are blackish.

Measurements: wing, & , 205.5, 207, 9, 196, 203; tail, d\ 181.5,
198, 9 , 173, 179; culmen, d\ 22.5, 22.5, 9 , 20, 20.5.

80. Chalcophaps indica indica (Linnaeus)

A wide-spread form recorded from Simalur, Siumat, Lasia (vide
Chasen, 1935), Nias, Tello, Siberut, the Pagi islands, and Enggano.

340 bulletin: museum of comparative zoology

It will probably be found to occur on all the islands with suitable
forest.

On Nias I saw the Green-wing Dove several times sunning and dust
bathing in the middle of the small island roads.

81. Caloenas nicobarica nicobarica (Linnaeus)

Recorded from Kokos, Simalur, Lasia, Babi, Nias, Batu Ids., and
Mirabau, an islet off Enggano.

This is primarily a bird of the small coral islands off shore, but
Nieuwenhuisen and von Rosenberg record it as common in the
Lagoendi Bay region of south Nias, near the present-day Telokdalem.

Family PSITTACIDAE, Parrots

Seven psittacine forms occur on the west Sumatra islands of which
six are endemic. The species Psittacula alexandri has no representa-
tives on Sumatra. The three races found on these islands are closest
to the population of the Andaman Islands. Like that race they differ
from the bird of the mainland primarily by larger size as well as certain
color characters.

The species Psittacula longicauda found on Sumatra as well as the
Andamans and Nicobars, has a single highly specialized representative
on these islands on Enggano. It differs by much larger size and rather
juvenal color characters.

There are two forms of Psittinus cyanurus on these islands. One,
from Simalur is not only much larger than the Sumatran bird, but
has certain striking color differences. The other, from the Mentawi
and Pagi Islands differs from the Sumatra bird only in larger size.

82. Psittacula alexandri cala (Oberholser)

Simalur. The type, an adult male taken October 21, measures:
wing 185, tail 198.5. The rest of the series collected by Abbott in
October and November measure: wing, d\ 173.5 (worn)-177.5, 9,
168.5-177.5; tail, cf, 179.5-196, 9, 170-182.

This race resembles abbotti from the Andaman islands very closely.
The only tenable difference seems to be that the pinkish color of the
breast, particularly in the females, is more pure, less suffused with
lilac. Other differences can apparently be matched in a large series.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 341

From fasciata this race differs by larger size, lack of any trace of
green on the forehead, and the less intense shade of green on the back.

This species is abundant in the west Sumatra islands nocking to-
gether and nesting throughout the year.

83. Psittacula alexandri major (Richmond)

Lasia and Babi. The type, an adult male from Babi, January 14,
measures: wing, 196.5, tail, 230.5. Other birds from the two islands
measure: wing, d\ 189.5, 196.5, 201.5, 9, 186, 188.5; tail, d\ 218.5,
229, 9 , 196, 200.5.

This is a very large race differing from cala primarily in size,
although the males seem to show a reduction in the light bluish suf-
fusion on the lower abdomen and vent. One female (U.S.N.M.
179112) has a thin ring of pink extending from the sides of the neck
dorsally around the nape between the bluish-gray crown and the
green back.

Soft parts: iris, pale yellow with an inner dull green ring; upper
mandible (cf), red; cere, greenish leaden, dull greenish; feet, pale
green, pale dusty green.

84. Psittacula alexandri perionca (Oberholser)

Nias. The type, an adult male collected February 22, measures:
wing, 194.5; tail, 211.5. Another male and two females measure : wing,
tf, 182, 9 , 177, 183.5; tail, <?, 189.5, 9 , (molt), 170.

This race occupies a somewhat intermediate position as regards cala
and major in size. Like major the males have less of the heavy bluish
suffusion on the lower abdomen and vent. The color of this area is
somewhat brighter than in major.

In June these birds were nesting, using holes made high up in the
trunks of dead coconut palms.

85. Psittacula longicauda modesta (Fraser)

Enggano. The two species, alexandri and longicauda, have an extra-
ordinary distribution in the greater Sunda area. It is particularly
interesting that alexandri, a species found away from the mainland,
principally on small islands, should have reached Java. This argues
a long history for the range of the species. And yet it is a representa-
tive of longicauda which has reached Enggano, the most isolated of
the west Sumatra islands.

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The race modesta. occupies one end of a cline starting with longicauda
of the mainland, Sumatra, and Borneo. The cline consists principally
of color characters which may be summarized as follows:

Subspecies

Loral
stripe

Crown

Nape

Back vs.
Scapulars

Size
Wing d" cf

longicauda

indicated

emerald
green

pink, like
cheeks

t

strong
contrast

small

defontainei

indicated

yellowish

emerald

green

paler

strong
contrast

larger

tytleri

present

yellowish
green

lacking

weaker
contrast

larger

nicobarica

present

chromium
green

lacking

weak

contrast

larger

modesta

extending

over

nostrils

indistinctly
green

present

very

pale

weakest
contrast

largest
199.5-209

The series collected by ^Abbott measures: wing, cf, 199.5-209
(205.6), 9, 195.5-209.5 (201.1); tail, d71, 182 (worn)-233, 9, 131-
151.5

Molting specimens were collected in October and November and a
fully grown immature female November 11. The soft parts are marked
as: iris, yellow, inner circle green; upper mandible, red, lower, horn
brown, black (cf); feet, greenish leaden, slaty-green, greenish.

A male and a female are heavily stained on the under parts, pre-
sumably from eating fruit.

86. Psittinus ctanurus abbotti Richmond

Simalur and Siumat. This very distinct form lies somewhere on
the borderline between a species and a subspecies. Primarily it reflects
the speciation trend of these islands by being much larger than cya-
nurus. Also, however, there are important color characters which set
it off from the typical race.

Adult male: the crown is a more uniform blue than in cyanurus,
lacking the gray wash on the nape. Also there is a sprinklingof greenish

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 343

iridescent feathers on the forehead and the ocular area. The grayish
black back and scapulars of cyanurus are lacking, being replaced with
green. The mid portion of the back is bright blue instead of pale dull
purplish blue and the rump and upper tail coverts are green instead of
blue. On the under surface greenish yellow replaces the yellowish gray
breast, abdomen, and vent, and the blue thighs of cyanurus.

Adult female: above this bird is almost uniformly green, lacking
the dull brown head of the typical race. Below, also, abbotti is uni-
formly greenish yellow.

The type, an adult male taken in December, measures: wing, 145;
tail, 58.5; culmen (from cere) 22.5; tarsus, 13. Iris, pale yellow; cere,
dull green; feet, greenish.

A series of males and females measure: wing, c? , 139-147.5 (143.5);
cf, im., 139; 9 , 134.5-141 (137.6).

An immature male collected in October has an indistinct tinge of
blue on the forehead. An adult male (U.S.N.M. 179109) has a spot of
dull red similar to that of the humeral patch among the blue feathers
of the nape.

This is a rather tame little parrot, common locally when the wild
fig trees are fruiting.

87. Psittinus cyanurus pontius Oberholser

Siberut, Sipora, and South Pagi. This race is distinguished from
cyanurus by larger size. In a large series, any color differences are
shown to vary within the species. A male from Siberut (U.S.N.M.
279754) has a very dark blackish back. Soft parts: "iris, pale yellow;
upper mandible, red tipped with yellow, lower, olive horn; cere, dull
leaden olive; feet, olive."

The type, an adult male collected in December, measures: wing,
127; tail, 50.5; culmen, 20.75. The rest of the series measures: wing,
<? , 127-134.5 (131.5); 9 , 129-132; tail, cf , 50.5-54. Birds from Su-
matra and the Malay Peninsula measure: wing, cf , 117-125 (121.6),
9, 115-123; tail, 44-47.

88. Loriculus galgulus galgulus (Linnaeus)
Synonym: Loriculus galgulus lamprochlorus Oberholser
Synonym: Loriculus galgulus dolichopterus Oberholser

Tuangku, Nias, Pini, Tello, Siberut, Sipora, Enggano.

The type of lamprochlorus, an adult male collected on Nias, March

344 bulletin: museum of comparative zoology

14, measures: wing, 79; tail, 32.5; culmen (from cere), 10.5. This race
was described by Oberholser (1912) as being "decidedly smaller, and
the colors averaging paler; female with green color more yellowish".
A series of birds from Sumatra, Borneo, and Java (three specimens
which may well be cage birds) measure: wing, cT, 78.5-83 (80.3).
The color characters also disappear in a series.

The type of dolichopterus, a female from Enggano collected Novem-
ber 6, measures: wing, 83.5; tail, 34.5; culmen (from cere), 11.5. A
series of females from Sumatra, Borneo, and Java (see anted) measure:
wing, 77-84.5 (81.4). This race was described as being larger and
darker, but these differences do not appear in comparison with a
large series of birds. Junge (1938) reached the same conclusion.

The wings of some of these specimens have not been fully pulled
back after skinning. In the case of the type of dolichopterus this may
have accounted for the impression that it was a larger bird.

Family CUCULIDAE, Cuckoos

Fifteen species have been recorded from these islands of which two
are migrants. Of the remaining species four are endemic on the islands.
Of these four races, two differ by color alone while the other two differ
only by larger size. All the races are closely related to representatives
of the species found on Sumatra.

89. Cuculus fugax fugax Horsfield

Pini, Siberut. An immature specimen from Siberut collected
October first, apparently belongs to this race. Chasen's record for
nisicolor (1935) may well be founded on this and another immature
bird taken by Kloss. The bird measures: wing, 167.5; bill (from gape),
26.5. Some of the slaty adult feathers are beginning to appear on the
forehead, cheeks, and nape. There are several white feathers also on
the nape.

90. Cuculus micropterus concretus S. Muller

Tana Bala. An adult male collected February 11, tail lacking, is
the first record of this species from the west Sumatra islands. It meas-
ures: wing, 191.5; culmen, 22.5. The soft parts are recorded as: "iris,
ochreous, rim of eyelid yellow; lower mandible, gray; gape, yellow;
feet, yellow." Compared with a male from Trang and a female frojn
east Sumatra collected in January and February, this specimen is dis-

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 345

tinctly more gray on the back, wing coverts, rump, and upper tail
coverts. The underparts seem to be identical. It is unfortunate that
no more specimens were secured.

91. CUCULUS SATURATUS SATURATUS Blyth

A migrant from southern Asia discussed by Junge (1931) in a careful
revision of the species. He lists two specimens of this race from
Simalur taken in February. The other record for the west Sumatra
islands is Nias (Biittikofer 1896, p. 171).

92. Penthoceryx sonneratii fasciolatus (S. M tiller)

Pini, Tana Massa. A male and a female collected by Kannegieter
in September and October on these two islands measure: wing, d*
(worn) 114, 9 111. These birds are indistinguishable from two males
from Trang, a locality certainly within the range of malayanus Chasen
and Kloss.

93. Cacomantis merulinus threnodes Cabanis and Heine
Synonym: Cacomantis merulinus subpallidus Oberholser

Nias, Siberut, and Enggano. The type of subpallidus, an immature
male from Nias, is a rather pale bird, but an adult male from Nias is
similar to a male from Borneo. I believe that this slight difference in
coloration is due to individual variation. Two adult males from Nias
and Siberut measure: wing, 101, 101.5.

The characteristic ascending notes of the brain-fever bird are often
heard on Nias in open areas of garden and cultivation.

94. Cacomantis variolosus sepulcralis (S. Mtiller)

Simalur, Enggano. A male molting into adult plumage and an im-
mature female were taken by Abbott on Simalur, December 14 and
November 30. The soft parts were noted as:' "iris, d* gray brown; 9
reddish brown becoming gray externally; eyelids, d1 greenish yellow;
bill black, brownish yellow at base of lower mandible; inside of mouth
orange; feet yellow."

95. Chalcites xanthorhynchus xanthorhynchus (Horsfield)
Only recorded from Simalur and Sipora.

346 bulletin: museum of comparative zoology

96. SURNICULUS LUGUBRIS DICRUROIDES (Hodgson)

A female collected on Nias March 15 evidently belongs to this large
raigatory race. It measures : wing 141.5, tail 135, culmen 22. The soft
parts are labelled: "iris dark brown, bill black, feet dark leaden."

97. Surniculus lugubris barussarum Oberholser

Tana Bala. The type of barussarum, an adult female taken on Tana
Bala in February, measures: wing 136, tail 127, culmen 22. Another
female taken in the same month measures: wing 137, tail (worn) 120,
culmen 22. Both birds have distinctly forked tails, the lateral feathers
in the type being 6mm. longer than the central ones.

This race was named (1912, p. 5.) as being smaller with the bill
relatively larger than I. lugubris. Actually two males of lugxibris from
east Java have the following measurements: wing 124, 127, culmen
20.5, 21. Robinson (Birds Malay Penin. /, 1927, p. 140.) gives meas-
urements for brachyrus of the Malay Peninsula, Borneo, and Sumatra
which are very close to these two birds from Tana Bala. A female
from Singkep Island, east Sumatra taken in May measures only:
wing 128, tail 123, culmen 20.5. If this specimen may be taken as an
example of typical brachyurus, then I presume that banissarum repre-
sents a large island form. Larger series may show, however, that these
measurements are not distinctive, in which c&sebarussarum must stand
as the name for the birds from the southern Malay Peninsula, Borrfeo,
Sumatra, and the Batu Ids.

98. EUDYNAMIS SCOLOPACEA SIMALURENSIS Junge

Kokos, Simalur, Lasia, Babi. Junge (1936) characterizes this race
as differing from malayana by, "smaller wing and in the 9 also by
the darker coloured and bigger spots on the upper side and the more
strongly red-brown washed under parts".

As to size I cannot entirely agree with this diagnosis. A male from
Babi measures: wing 211, tail 197, culmen 36, while a male from
Simalur in the collection of the Museum of Comparative Zoology has
a wing measurement of only 195 according to Mr. Peters. A series of
males of malayana measure: wing 203-212.5. However, a single pos-
sibly immature female from Simalur is distinctly more rufous above
and below than any female of malayana. In this respect it is very
close to females of mindanensis. Fortunately the bill of mindanensis

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 347

is noticeably smaller. The record for Lasia is a sight one, (Richmond,
1903).

99. Eudynamis scolopacea malayana Cabanis and Heine

Nias, Tello, Tana Massa, Pagi islands. Two males, an immature
male and three females from Tello and Tana Massa measure: wing,
c? 217.5, 218; 9 im. 213, 9 209, 212, 213.

There is a good deal of individual variation in size, bill dimensions,
and color in this species. These birds are large but not significantly
so. The soft parts are recorded as: iris red; bill gray, greenish gray;
feet gray.

The record for the Pagi islands is not founded on any specimen so
far as I know, but undoubtedly these birds could be found on almost
any of the small coral islands of the area, especially in the breeding
season.

100. Rhopodytes sumatranus rodolphi Ripley

Pini. The type, an adult male collected October-November 1896,
measures : wing 154, tail 237, culmen 37.5. The soft parts are recorded
as: iris light blue, bill yellowish green, feet gray.

Although only a single specimen has been taken on the west Sumatra
islands, its bill measurements are significantly larger than a series of
specimens from Sumatra or the Malay Peninsula and adjacent islands.
In color there seems to be no perceptible differences between this and
Sumatra or southern Malay Peninsula birds. This specimen has very
little rufous on the abdomen, but I believe that individual variation
or skinning will account for it.

101. Rhinortha chlorophaea facta Ripley

Tana Massa. The type, an adult male collected February 20, 1903,
measures: wing 123.25, tail 182, culmen 29, tarsus 28.5. A female
(A.N.S.P. 56256) collected August 24, 1896, measures: wing 124, tail
179.5, culmen 30.5, tarsus 28.5. The soft parts are given as: "iris, 9
blue gray; bill, cf dark greengage green, 9 , green; feet, c? slaty, 9
gray."

These birds are noticeably larger than any other specimens of the
species in the Museum's series. Except for Banguey I. off the coast
of North Borneo this bird has never been reported on any of the small
islands in the Greater Sunda area.

348 bulletin: museum of comparative zoology

102. Rhamphococcyx curvirostris oenicaudus
(Verreaux and Verreaux)

Siberut, Sipora, North and South Pagi. This race differs from
erythrognathus of the Malay Peninsula and Sumatra by having the
greenish iridescence on the upper surface rather darker and more
bronzy, extending up onto the nape and crown and serving to darken
the gray of those areas. The tail lacks the terminal brown patch
characteristic of the other members of the species.

Below the greenish iridescence extends up over the abdomen. The
under surface of the tail is greenish blue not brown. There seems to
be no size difference within the species.

Five males, seven females, and three sex unidentified, measure:
wing, cf , 166-181, 9 165-178, o 171-179. Soft parts are recorded as:
"iris, c? bluish white, pale blue, pale china blue, 9 dark; bill, maxilla
apple green; nostril black, pale green; mandible green, base dull
crimson, dark red; orbital skin scarlet, red; feet plumbeous."

103. Centropus sinensis bubutus Horsfield

Nias, Siberut. Four males and one unsexed from Nias collected in
February and March measure: wing, & 212, 215, 218, 221, o 220, tail
<? 272, 278, 278, o 285. The variation in these birds from purplish
to greenish iridescence about the head, neck, and breast varies with
the freshness of the plumage.

Family TYTONIDAE, Barn Owls

The only species of this family found in the area is the small Bay
Owl of the greater Sunda Islands which is not uncommon on Nias.

104. Phodilus badius badius (Horsfield)
Nias. This species has only been recorded from Nias where it is
not uncommon. For further comments and field notes see de Schau-
ensee and Ripley (1939, p. 402.).

Family STRIGIDAE, Owls

Six forms of owls are found on the west Sumatra islands, all of which
are apparently endemic. Two differ from their congeners in Sumatra
and Borneo, primarily by color differences. Three of the remaining

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 349

forms are distinctly larger than their nearest relatives on the Malay
Peninsula or Sumatra, as well as differing slightly in color. One race
is smaller than its Sumatran congener.

105. Otus scops umbra (Richmond)

Simalur. The type and unique specimen, an adult male, was col-
lected by Abbott November 29, 1901. It measures: wing 143, tail 61,
culmen 20, tarsus 24. The soft parts are recorded as: iris greenish
yellow; bill pale brown, black at tip; feet pale fleshy brown. The wing
formula is 4>3>5>2>6>7>1.

This bird is presumably in the rufescent phase. The outer scapulars
have conspicuous white spots and the feathers of the lower abdomen,
vent, and flanks are freckled and mottled with white, russet, and black
in a way that seems characteristic of the scops group.

The stomach contained insects.

106. Otus scops enganensis Riley

Enggano. A single female, the type, was collected by Abbott, No-
vember 12, 1904. Its measurements are: wing (worn), tail 73, culmen
23, tarsus 30. The tips of the primaries are completely worn off, a
condition that was not mentioned in the original description. The
wings of five birds collected by deJong (Junge 1938) measure: c? ,
160, 163; 9 , 163, 163, 166. The soft parts in the type are recorded as :
"iris greenish yellow, feet pale brownish fleshy."

The type is a bird in the rufescent phase of plumage. In his dis-
cussion of this form, Junge (I.e.) showed that the brownish or gray-
brownish individuals are very close to malayanus and sunia. The
under parts, however, are more uniformly dark rufous than any
rufescent individual of scops I have seen, except for the type of umbra,
which is the most uniformly dark of all.

107. Otus bakkamoena mentawi Chasen and Kloss

Siberut, Sipora, North Pagi. In their original description (1926),
Chasen and Kloss neglected to state that this race differed from true
lempiji by much larger size. A series of these birds measure: wing,
c? 157> 165, 9 165, 165.5, 9 im. 160; tail, tf 77.5, 82, 9 72, 80, 80,
9 im. 76. Ten specimens of lempiji from Java, Sumatra, and the
Malay Peninsula measure: wing, cf 141-150.5 (145.2), 9 140-151.5
(145.1); tail, d* 65-70 (67.2), 9 65-67.5 (66.3)..

350 bulletin: museum of comparative zoology

The soft parts of these birds are listed as: "iris, brown (3), yellow
(2), feet gray." The color differences noted in the original description
are very noticeable in this series.

108. Ketupa ketupu minor Biittikofer

Nias. Compared with ketupu from adjacent areas, three males and
a female from Nias show the following measurements:

wing tail

Nias cf 304-330.5 (316.1) 143-164 (153.5)

9 304 147

Sumatra1 cf 320-349 (338.6) 158-171 (165.4)

9 335, 338 158

Borneo c? 312-343 (330.1) 153-164.5 (158.4)

9 — —

Java & 341-357 (348.4) 157-169 (164)

9 358, 362 170, 176

From the above it will be seen that minor is a tenable race but that
there is a good deal of overlap with Sumatra and Borneo birds. Two
of the Nias males are darker on the breast than any specimen of our
series of ketupu.

Three young were taken in late March. The plumage is rather uni-
form dark buffy all over except for the thighs which are whitish. Both
upper and under parts are marked with the narrow dark brown shaft
streaks found in the adult only on the under parts.

109. Strix leptogrammica nyctiphasma Oberholser

Bangkaru. The type, an adult male collected in January, measures:
wing 299.5, tail 154. A female taken at the same time measures: wing
307, tail 167.

The type differs from a male from the Kapuas River, west Borneo
(w. 298, t. 168) by having the cheeks somewhat paler and the dark
barring on the underparts slightly heavier, the bars being closer to-
gether. In size there is no difference and the color differences are so
slight that a larger series may well show these forms to be identical.

From myrtha of Sumatra these birds differ primarily by smaller
size. This bird will probably be found one day on Babi and Simalur,
provided there is some original forest left for them.

1 have padded out my series with measurements taken in the same manner by Mayr (Bull.
Raffles Mus., No 14, 1938, p. 13) and de Schauensee and Ripley (Proc. Acad. Sci. Philadelphia.
91, 1939, p. 324.)

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 351

110. Strix leptogrammica niasensis Salvadori

Xias. A male and female (M.C.Z. 194794, 194795) collected in
July 1937 by Barbara Lawrence measure: wing, a71 273, 9 280.5;
tail, d" 151.5, 9 156.5. The male is an immature probably completing
the post juvenal molt {vide Robinson, Birds Malay Benin., 2, 1928,
p. 36). The feathers of the posterior part of the crown and nape and
a few feathers in a ring on the neck below are pale whitish buff in color
and fluffy.

The iris is recorded as brown by Biittikofer, chestnut by Salvadori
(1887). Salvadori (I.e.) lists the bill and feet as pearl gray or sky blue.

Family CAPRIMULGIDAE, Nightjars

There are records of three forms from these islands of which one
has been separated on color characters, and one, Eurostopodus tem-
minckii, shows an unconfirmed tendency towards larger size. The
endemic form, Eurostopodus viaCrotis jacobsoni has no close geograph-
ical relatives except cerviniceps from the Malay Feninsula.

111. Eurostopodus temminckii (Gould)

Nias. Two males taken by Abbott in March measure: wing 211,
213; tail 133.5, 135. Two males from Johore and Banka measure:
wing 201, 203; tail 117, 126, while three males from north Sumatra
are recorded by de Schauensee and myself (I.e., 1939) as having
wings of 197, 198.5, and 201.5 mm. However, Robinson and Kloss
(Journ. Fed. Malay States Mus., 11, 1924, p. 242) give measurements
for males up to 218.

In these two specimens the buffy whitish subterminal bars forming a
ring on the nape are narrower than in any other specimens in the
National Museum series. Also one specimen (180824) has very large
black spots on the crown.

112. Eurostopodus macrotis jacobsoni (Junge)

Simalur. According to the description (1936) a darker bird than
macrotis from the Bhilippines or cerviniceps of the Malay Beninsula.
It is most interesting to find this population so isolated from the rest
of the species. Further collecting might reveal that it occurs in the
Andamans or Nicobars.

352 bulletin: museum of comparative zoology

113. Caprimulgus affinis affinis Horsfield

Recorded from Nias by Nieuwenhuisen and von Rosenberg (1863)
under the name C. maculatus.

Family MICROPODIDAE, Swifts .

Six races of swifts are found on the west Sumatra islands of which
two are endemic. One race differs in color alone, and the second differs
in color and smaller size. Both races are closely related to a wide-
spread form found throughout the greater Sunda area.

114. Collocalia lowi lowi (Sharpe)

Simalur, Babi (sight), Nias. A male from Simalur taken May 5 is
listed by Junge with a wing measurement of 131.

115. Collocalia fuciphaga fuciphaga (Thunberg)

Nias. A single female taken in June is the only record of this species
for the west Sumatra islands.

116. Collocalia vestita vestita (Lesson)
Synonym: Collocalia fuciphaga aerophila Oberholser

Simalur, Nias, Sipora. The type of aerophila, an adult male from
Nias, collected March 16, measures: wing 115; tail, shortest rectrix
48, longest rectrix 53. A male and a female from Simalur and a female
from Sipora measure: wing, d71 112.5, 9 116.5, 116; tail, <? 46 and
50, 9 41.5 and 47, 49 (incomplete). Compared with a male from
Johore, I can see no significant differences in these specimens.

117. Collocalia esculenta cyanoptila Oberholser

Simalur. Junge (1936) records three specimens taken on Simalur
in April and July as belonging to this race. He gives their measure-
ments as follows : wing, cf 107, 109, 9 102; tail, cf 41, 41. The type
and another female of cyanoptila from Bunguran-L, Natunas, measure:
wing, 9 103.5, 103.5; tail, 41.5, 39.5 (molt). Both birds are in rather
worn plumage which accounts for the admixture of purple and greenish
gloss mentioned by Oberholser in his original description (Proc. Acad.
Nat. Sci. Phila., 58, 1906, p. 205).

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 353

118. Collocalia esculenta vanderbilti de Schauensee and Ripley

Nias. The male type and a female measure : wing, cf 99, 9 97; tail,
cf 39, 9 37. This race is smaller than oberholseri and somewhat bluer
above. These little swiftlets are common over jungle trails and roads,
especially in the early morning and evening.

119. Collocalia esculenta oberholseri Stresemann

Tana Massa, Sipora and North Pagi. The type, an adult male from
North Pagi, taken in November, measures: wing 105.5, tail 42.5.
Other birds from Sipora and North Pagi measure: wing, cf 105;
9 98, 105; o 105, 105.5; tail, cf 42.5; 9 40.5, 41.5; o 42, 42.5.

From cyanoptila this race differs by being more greenish above and
with the breast and upper abdomen paler below with more tips among
the brownish gray of that area.

Family HEMIPROCNIDAE, Tree Swifts

Two races of Tree Swifts are found on these islands of which one is
endemic. It differs from its large island and mainland relatives by
larger size.

120. Hemiprocne longipennis perlonga (Richmond)
Synonym: Hemiprocne longipennis ocyptera Oberholser
Synonym: Hemiprocne longipennis thoa Oberholser

Simalur, Nias, Pini, Tana Massa, South Pagi, Enggano. The type
of perlonga, an adult female, was collected on Simalur January 2,
1902. It measures: wing 183.5, tail 112.5. The type of ocyptera, an
adult male collected March 23, 1905 on Nias measures: wing 169.
The type of thoa, an adult male from Pini taken March 7, 1903,
measures: wing 177.5, tail 102. These last two races were described in
a lucid fashion by Oberholser (1912, pp. 7, 8) without being compared
to each other, although they came from closely neighboring islands.

A total of seventeen specimens from the west Sumatra islands have
wing measurements as follows: cf 164-180 (173.1), 9 169.5-183.5
(174.9). Of this series the birds from Nias are the smallest: cf 164-
176 (169), 9 169.5. This is probably due to the fact that Nias of all
the islands of the group seems to be the most closely connected to
Sumatra from a faunistic point of view. Whether in the case of Hemi-

354 bulletin: museum of comparative zoology

procne this might be due to swamping is certainly a possibility in the
case of this distinctly aerial species.

I have examined forty-one specimens of longipennis from the Malay
Peninsula, Rhio and Lingga islands, Sumatra, Natunas, Anambas,
Borneo, Banka, and Java. Seventeen males measure: 159-170.5
(164.3). Exclusive of molting birds, twenty-three females measure:
155-173.5 (166). From these figures it will be seen that perlonga is a
tenable subspecies. However, it seems to me that in general this
species has been split into far too many subspecies. The type of
anochroa Oberholser, an adult female from the Natunas, has a greenish
suffusion in the chest region and in general is somewhat darker than
average on the underparts. But in this it can be matched by birds
from the Malay Peninsula, Sumatra, and Borneo.

Stresemann's original description of the race harterti (Novit. Zool.,
20, 1913, p. 339) indicates that he did not have any Bornean material,
for he gives no measurements of specimens from there. Individuals
from Borneo show every variation between the two extremes of colora-
tion of the underparts of this species. Some birds are paler gray on
the throat and breast with more white on the lower abdomen. Others
are darker gray with less white on the abdomen. Northern Malay
Peninsula birds seem to be the darkest, while Java birds are the
palest. In between them is every variation of what is, after all, a
rather slight range of variation. There is no difference in size. Under
these circumstances it seems to me that it would be wiser either to
recognize several microscopic degrees of variation and name all the
small local populations, or else, and this I think preferable, to lump all
these birds as longipennis. This would leave the species arranged as
follows :

Hemiprocne longipennis coronata (Tickell)
India, Ceylon, Siam to Indo-China. A distinctive rufous-throated
race.

Hemiprocne longi pennis longipennis (Rafinesque)
Peninsular Siam to Sumatra, Borneo, Java, and Bali. Dark to
paler gray below.

Hemiprocne longipennis perlonga (Richmond)
West Sumatra islands. A large race.

Hemiprocne longipennis wallacii (Gould)
Celebes to the Sula islands. More bluish on the upperparts.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 355

121. Hemiprocne comata comata (Temminck)
Synonym: Hemiprocne comata stresemanni Neumann

Nias, Pini, Tello, Tana Massa, Tana Bala, Sibe'rut, North Pagi. A
series from these islands measures: wing, cf 126-130 (127.6), 9 123
(worn)-134 (128.5). I fail to note any of the differences mentioned
by Neumann.

Family TROGONIDAE, Trogons

Two trogons occur on the west Sumatra islands. One from Nias is
an endemic race differing slightly from its nearest relative on Sumatra
and the Malay Peninsula in color and a tendency to have a larger bill.

122. Harpactes duvaucelii (Temminck)

The only record for this species is Tana Massa (de Schauensee 1940).
A male and a female taken by Kannegieter measure: wing, c? 108,
9 105.

123. Harpactes oreskios nias de Schauensee and Ripley

Nias. In the original description (1940, p. 404.) this race was said
to differ from uniformis by lacking the orange wash on the chest of the
male and having narrower white bars on the wing coverts and second-
aries. Three males and a female from Nias, when compared with birds
from Peninsular Siam and the Langkawi islands, do not show these
characters. All three of the males have a rich orange suffusion indicat-
ing that the type may not be fully adult. However, these Nias birds
do have somewhat darker more brownish crowns than the birds with
which they have been compared. Mr. de Schauensee writes me (in
litt.) that this is also true of the type. As well as this, these birds
show a tendency to a slightly larger bill. The culmen of the four Nias
specimens measures: c? 17, 17.5, 18.5; 9 18. A series of uniformis
measures: c? 15, 15, 16, 16, 17; 9 14.5, 15, 16, 16, 17.

This trogon is found in heavy secondary jungle at low heights above
the ground, often in substage growth. It has a very distinctive bark-
ing call.

Family ALCEDINIDAE, Kingfishers

The west Sumatra islands have records of twelve species and sub-
species of kingfishers. Of these two are possibly migrants leaving ten
certain resident forms. There are five distinguishable endemic forms
three of which differ primarily by size and two of which differ in color

356 bulletin: museum of comparative zoology

from their nearest relatives. Four of these birds are most closely re-
lated to Sumatran subspecies, while one, Ceyx erithacus captus, is
almost indistinguishable from C. e. motleyi of Borneo.

Ramphalcyon capensis

This is an interesting and widespread species ranging from Siam
down through the Greater Sunda islands to some of the Lesser Sundas.
Oberholser (Proc. U. S. Nat. Mus., 35, 1909, p. 657) has revised the
genus and ended by describing several races of capensis. The addi-
tion of further specimens to the National Museum's collection has
shown, I think, that this action was unwise, as the variation in plumage
of these birds is considerable.

At least in the Malay Peninsula and island part of its range, fresh
plumage in this species is assumed in July, August, or early September.
At this time the color of the pileum may approach sepia to bistre.
The feathers of the upper back, scapulars and wing coverts are rather
bright greenish blue at this time. After eight or ten months, as the
feather tips become frayed and the color of the pileum bleached, very
noticeable changes are brought about in the color of the upper parts.
March to June or July birds have the pileum colored from whitish-
buff or pale fawn to ochraceous-buff. The color of the upper back,
scapulars and wing coverts changes also, this time not due to bleach-
ing, for there is no pigment change as tests under sodium, helium, and
ultra-violet lights have shown. Instead the frayed and sun-heated
feather seem to have become physically changed, contracting the
diffraction grating with resulting effects similar to those observed in
certain beetles. At this time the color of these parts varies from dull
cerulean to indigo to purplish blue.

Birds from the Malay Peninsula and the northern part of Sumatra
tend to be somewhat darker buff on the under surface. Perhaps they
should be kept under malaccensis. Borneo, Billiton, and Java birds
tending to be rather pale beneath, could, it seems to me, quite well
be lumped together as capensis. There is no constant difference in
size in a series of measurements of more than thirty specimens.

From these birds the west Sumatra island populations differ in
being somewhat paler and more washed out on the upper surface and
by a tendency to larger size. The blue which is so bright in capensis
and malaccensis varies in these specimens from dull brownish China-
blue in a September female from Siberut, to pale indigo in a March
male from Nias.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 357

124. Ramphalcyon capensis simalurensis (Richmond)

Simalur. The type, an adult male collected by Abbott November
29, measures: wing 144.5, tail 92.5, bill (from naris) 07. 5. Two other
males and three females measure: wing, cf 143.5 (molt), 143.5 (143.8);
9 151, 155.5; tail, 9 96, 99; bill, d1 65.5, 68.5; 9 70.5, 71.2.

Stomach contents are noted as: "small fish, prawns, and crabs."
This is a common bird of the mangroves and small creeks near the sea.

Compared to malacccnsis, this series is equally dark buff below, but
on the upper surface it is very different, lacking the bright blue or
greenish blue of the upper back, scapulars, and wing coverts.

Compared to birds from the other west Sumatra islands, these
Simalur birds appear uniformly dark buff below in the same way that
malaccensis differs from capensis. The upper parts also appear notice-
ably duller and browner than the majority of other west Sumatra
island birds, but this condition may be in part due to the fact that all
the Simalur birds were collected between October 23 and November 30.

125. Ramphalcyon capensis sodalis (Richmond)

Synonym: Ramphalcyon capensis nesoeca Oberholser
Synonym: Ramphalcyon capensis isoptera Oberholser

Babi (sight), Tuangku, Nias, Pini, Tello, Siberut,Sipora, North and
South Pagi. The type of sodalis, an adult female collected January
25, measures: wing 162, tail 100.5, bill (from naris) 73.5. Another
adult female from Tuangku measures: wing 160, tail 101.5, bill 73.2.
This race was named on the basis of size, but females later added to
the Museum's collection from other islands have equalled these
measurements.

The type of nesoeca from Nias is an adult male collected March 15.
It measures : wing 148.5, tail 93, bill (from naris) 68.5. The range of
this race was given (1909, p. 674) as Nias and the Batu islands, and
the characters were: "pileum paler, back, wings, and tail much brighter
and more bluish." In four of the eight specimens this is true, but as
these are March birds taken at a time when the pale pileum and
brighter blue is characteristic of the state of wear of the plumage, I
do not feel 50% is a good enough average difference for separation.
However, it is certainly worth noting that in the slightly brighter
color of the upper parts these four Nias specimens indicate a closer
affinity with Sumatra than is the case on the other islands.

An adult male taken December 30 in Sikakap Strait between North

358 bulletin: museum of comparative zoology

and South Pagi is the type of isoptera. It measures: wing 153.5, tail
94, bill (from naris) 68.5. This race was described (1909, p. 671) as
being larger than simalurensis with the under parts lighter, the lower
back and rump more greenish. In size these birds agree with the birds
from the other islands, as well as in having paler underparts than
simalurensis. I can find no constant difference in the greenish char-
acter of the rump.

Measurements of a series of sodalis by localities follows :

place

wing

tail

bill (from naris)

Tuangku 9

160, 162

100.5, 101.5

73.2, 73.5

Nias c?

146-152.5 (148.7)

89-93 (91)

68.2-72.5 (70.1)

9

163.5

101

68.5

Batu cf

155

89

70

O

—

92.5

75

Siberut 9

156.5

97.5

71

Sipora cf

150.5-158.5 (154.1)

95.5-101 (98)

66.5-70.5 (69)

Pagi c?

150-153.5 (151.5)

89-96.5 (93)

63-70 (67.5)

9

153, 161

91.75, 95

68, 71.5

126. Alcedo atthis bengalensis Gmelin

Simalur, Tuangku (sight), Nias, Pini, Siberut. Two pairs from Sim-
alur and Nias measure: wing, cf 70.5, 69.5; 9 70, 70. These birds
were taken in November, February, and March.

127. Alcedo meninting meninting Horsfield
Synonym: Alcedo meninting subviridis Oberholser
Synonym: Alcedo meninting callima Oberholser
Synonym : Alcedo meninting proxima Richmond

Simalur, Tuangku, Nias, Pini, Tana Massa, Tana Bala, Sipora,
North and South Pagi, Enggano. The type of subviridis is a male
from Nias collected March 23. It measures : wing 62, bill (from naris)
34. This race was described by Oberholser (1912) as smaller than
meninting with more greenish upper parts. Other birds from Nias
measure: wing, tf1 64.5, 9 63.5, 65.5; bill, cf 34, 9 31.5, 32.5. Birds
from Sumatra measure: wing, <? 61-65, 9 62-66.5; bill, c? 35-36.2,
9 31.5-33.5. Of the four specimens from Nias, only one is especially
greenish on the back and rump.

The type of callima is a male collected on Tana Bala February 8,
measuring: wing 69, bill (from naris) 36.2. It was described (1912) as
larger than meninting with upper parts slightly more greenish. A male

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 359

from Pini and a female from Tana Bala measure : wing, d" 65, 9 68.5 ;
bill, cf 36.5, 9 34.5. These measurements are large, but Junge (1936)
gives measurements for Simalur birds: wing, d* 63, 66, 69; 9 65, 67,
which are largely overlapping. Again the greenish character tends to
disappear with a large series.

The type of proxima is an adult, sex undetermined, from North
Pagi, collected January 4. It measures: wing 69.5, bill 37.5. Other
birds from North and South Pagi measure: wing, cf 67.5-69, 9 71.5;
bill, cf 35.5-37.5, 9 33.5. This race was described as being very
greenish on the back and rump. Two of the specimens are particularly
so — one slightly, and the fourth, the type, can be matched by Sumatra
birds. I think this character is entirely too variable to be a serviceable
criterion for subspecies. These birds are also large, but no more so
than Batu or Simalur birds. Recently Junge (1938) has recorded a
single male from Enggano. The wing measured 63.

From these data, I would be inclined to think that probably these
kingfishers at some time or other reach all the islands of the group.
Also these birds show a tendency to larger size, particularly on Sim-
alur, the Batus, and the Pagi islands. Finally there is a tendency which
I do not consider very significant so far, for some of these birds to have
the iridescent back and rump feathers paler and more greenish.

128. Ceyx erithacus captus Ripley

Nias. This, the only representative of the species from the west
Sumatra islands, is almost identical with motleyi of Borneo. From
this race it differs by a longer bill, slightly larger size, and by the re-
duction of the forehead spot which may be lacking in some specimens.

A not uncommon bird in thick secondary growth or substage vege-
tation usually near small streams. The specimens I saw were all
perched about six feet above the ground.

129. Ceyx rufidorsus jungei Ripley

Simalur, Bangkaru (sight), Tana Massa, Tana Bala. These birds
differ from typical ruflidorsus by larger size. A series measures : wing,
cf 62-64.5 (63.2), 9 62.5-63.5 (63); tail, <? 25-26 (25.5), 9 25.5-26.5
(26); bill, cf and 9 30.5-32 (31.3).

130. Ceyx rufidorsus rufidorsus Strickland

Siberut, Sipora. A male and female adult and an immature male,
taken October 28, are in the collection. They measure: wing, c? 61,

360 bulletin: museum of comparative zoology

9 62; tail, d* 25, 9 26; bill (from naris), tf 25, 9 26. These birds
are certainly close to jungei in size, but Chasen and Kloss' measure-
ments (1926) for Siberut and Sipora birds are so small that, without
seeing the rest of their series, I would rather leave them for the time
being as rufidorsus. The fact that one of the specimens in the National
Museum from their collection is a subadult male inclines me to be-
lieve, however, that some of their other measurements may refer to
immature specimens. There is a sight record for Ceyx from the Pagi
islands, which may refer to this species.

131. Halcyon coromanda minor (Temminck and Schlegel)
Synonym: Entomothera coromanda pagana Oberholser

Simalur, Lasia, Nias, Tana Massa, Siberut, North Pagi. The type
of pagana, an adult male collected January 4 on North Pagi measures :
wing 110, tail 62, bill (from naris) 44.5. Three females from Siberut
and North Pagi measure: wing 107.5, 109.5, 110.5. The soft parts are
listed as: "iris dark, sepia; bill and feet coral red, coral, blood red."
These birds are indistinguishable from minor as is also the type of
neophora, a bird from Tapanuli Bay, northwest Sumatra.

132. Halcyon chloris laubmanniana Grote

Synonym: Halcyon chloris cyanescens * (Oberholser)

Synonym: Sauropatis chloris chloroptera Oberholser

Synonym: Sauropatis chloris amphiryta Oberholser

Simalur, Babi, Nias, Pini, Lago, Tello, Tana Massa, Siberut. Speci-
mens from this area measure as follows (t stands for type) :

wing

bill (from naris)

Simalur o"

115, 116.5, 117.5 (t),

117.5

41, 43, 44.5 (t), 44.5 (2)

9

117

46

Nias c?

113.5 (t)

42.5 (t)

9

114.5

46

O

120

43.5

North Pagi

9

111-116.5 (113)

41-43.5 (41.8)

The type of cyanescens from east Sumatra, an unsexed bird collected
July 28, measures: wing 113, bill 43.5. Specimens from Sumatra,
Banka, and Java measure up to 115.5 (wing) and 43.5 (bill). The
original description of chloroptera from Simalur (Proc. U. S. Nat. Mus.,
55, 1919, p. 379) compared it to cyanescens, but said the birds had

1 Not Halcyon cyanescens Cabanis and Reichenow, 1877.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 361

longer wings and that the upperparts were more greenish, the pileum
was darker, the blaek nuchal hand narrower, the auriculars less green
washed, and the sides and flanks noticeably tinged with huff. In
series these color characters do not appear, nor are the size measure-
ments significant when the amount of overlap is considered.

In the original description of amphiryta from Xias (I.e., p. 382) this
bird seems to stand in an intermediate position between chloroptera
and lavbmanniana. It is smaller than the former and larger than the
latter. In color it is brighter than the former and duller than the
latter. This is a form of microdissection of characters which escapes
me. The plumage changes of these birds are now well enough known
to make comparisons of shades of greenish or bluish color untrust-
worthy. Especially is this difficult when a series of birds, for example
those from Simalur (Oct., Nov., Dec.) or Nias (Feb., March), are
taken all at about the same time. Thus they may present a condition
.of uniformity of plumage which is not really valid for the population
as a whole.

Undoubtedly these birds will eventually be shown to exist on all
the small islands and reefs of this group. They are birds of the man-
groves and beaches and usually omnipresent in these locations. Junge
(1936) lists nestlings from Simalur collected in April and eggs taken
in April and May. Nesting occurs in steep banks along streams, in
hollow trees, and even in termite nests.

133. Halcyon chloris azela (Oberholser)

Enggano and Pulu Dua. This race resembles birds from the Malay
Peninsula and outlying islands, the Andamans,and northeast Sumatra,
in size, but on the average tends to be somewhat darker on the pileum
and more dull greenish on the upper back and scapulars. In view of
the isolation of this island population, it seems strange that these birds
are so slightly distinct. Actually azela appears to be simply a small
race of cyanescens, like that race somewhat duller and darker in color
than the small Malay Peninsula birds.

The type of azela, an adult male collected November 19, measures:
wing 103, bill (from naris) 38.5. A series measures : wing, c? 102 (worn)
-104.5, 9 102-104.5; bill, c? 36-40.5, 9 38.5-41.5.

The races arvistrongi, humii, and davisoni are so far as I can see,
indistinguishable. Sharpe (Cat. Birds Brit. Mus., 17, 1892, p. 277)
describes armstrongi as having green ear coverts and the black band
around the nape obsolete. This description does not seem to be valid.

362 bulletin: museum of comparative zoology

All of the National Museum's series from Peninsular Siam, the Malay
Peninsula and islands, the Andamans, and Belawan Deli, Sumatra,
have at least partially black ear coverts and a well indicated nape
band. Robinson (Birds of the Malay Penin., 1, 1927, p. 99) felt that
the kingfishers from Lower Tenasserim and the extreme north of the
Malay Peninsula were more dull than birds from elsewhere and so
might be called armstrongi. The National Museum has both dull and
bright birds from all localities, which inclines me to believe that they
should all be listed under armstrongi, the name with page priority.

134. Halcyon pileata (Boddaert)

Simalur, Babi (sight), Pini, Siberut, Sipora, North and South Pagi.
This species has not been found breeding south of Bangkok and may
well be a migrant. Specimens taken in November, December, and
January measure: wing, cf 130-131.5, 9 126.5-134. Soft parts were
recorded as: "iris dark brown, bill red washed with brown, feet coral
red." The stomach of one bird contained a lizard.

135. Halcyon concreta concreta (Temminck)

Siberut. The single record for this species is an immature male
taken by Kloss. The wing measurement (1926) is recorded as 112,
and the soft parts: iris dark, bill yellow, culmen black, feet greenish
yellow.

Family MEROPIDAE, Bee-eaters

Two species have been recorded, one of which is only a sight record.
One species which is a migrant has been collected during the winter
months.

136. Merops leschenaulti leschenaulti Vieillot

Dr. Abbott reported seeing this species on Simalur (Richmond,
1903).

138. Merops superciliosus javanicus Horsfield

A migrant taken on Simalur, Tuangku, Nias, Pini, Tello, Tana
Massa, Tana Bala, Siberut and South Pagi. The occurence in these
islands is from September 19, the earliest date, to March 16. Some of
the fall birds are immature. Their plumage is worn, the cinnamon
throat patch is undeveloped, and the upper parts are sprinkled with
blue tipped feathers. Stomach contents, winged ants.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 363

Family CORACIIDAE, Rollers

Two forms, both migrants, occur among the islands in winter. A
third resident form has been described from Simalur. It differs from
orientalis of the Malay Peninsula and Sumatra primarily in color.

139. EURYSTOMUS ORIENTALIS ABUNDUS1 Ripley

Simalur. A male collected by Abbott in December belongs to this
race. The primaries and secondaries are washed with purplish blue
nearly to the end of the outer webs. The specimen measures: wing
194, tail 99, wing-tail ratio 51, wing tip index 36. The soft parts are
recorded as, "iris dark brown, feet red".

Records from Tello, Sipora, North and South Pagi may refer either
to this or the following race, both of which migrate into this area.

140. EURYSTOMUS ORIENTALIS DEIGNANI Ripley

Nias. A male collected by Abbott in March presumably belongs
to this race, although in worn plumage it is not so easy to distinguish
the color characters which separate it from abundus. Certainly there
is no purplish-blue wash on the primaries and secondaries and the
crown is rather blackish. Measurements: wing 1ST, tail 93 (worn),
wing-tail ratio 50, wing tip index 40.

141. EURYSTOMUS ORIENTALIS OBERHOLSERI Junge

Simalur. A single female collected in October, agrees well with
Junge's original description. It measures, wing 187.5, tail 103, wing-
tail ratio 54, wing-tip index 30. Junge (1936) notes that females were
showing nesting activities in January and March. Stomach contents :
large hard coleoptera.

Family BUCEROTIDAE, Hornbills

Two species occur in these islands, one of which is considered to be
identical with Sumatra birds although showing an incipient tendency
to larger size. The other form is endemic, differing from the Sumatra
and Borneo population by definitely larger size.

1 replaces Euryslomus o. catonyx auctorum

364 bulletin: museum of comparative zoology

142. Anthracoceros coronatus convexus (Temminck)
Synonym: Hydrocissa convexa. barussensis Oberholser

Nias, Tello, Tana Massa, Tana Bala, Siberut, Sipora, North and
South Pagi. The type of barussensis is an unsexed bird, obviously a
male with the large wing measurement of 326 and a tail measuring 289.
Birds from the other islands measure as follows :

wing

tail

Nias d" d1

295-319.5 (306.7)

269-294 (282.5)

9

275

—

Batu Ids. d1 d*

314, 321

285

O (=6)

326

289

9 subad.

279

—

Sipora cT

290

—

Pagi Ids. d* &

282-299 (289.6)

273-282 (277.6)

0(= 9)

263

246

Birds from the Batu Islands are indeed larger than average but I
feel that there is too much overlap to make this race recognizable.
Measurements of over 300 for the wings of Sumatra birds are not
uncommon, and as can be seen from the above measurements there is
a large degree of overlap between Nias and Batu birds. However,these
west Sumatra Island birds do demonstrate in an incipient form a
speciation response by increased size which seems to be so character-
istic of these island bird populations.

Soft parts; iris brown, pale vandyke brown; orbital skin bluish
white; bill ivory, patch on casque and base of bill black; feet dull
plumbeous.

Weight: & 2 lbs., 9 1 lb., 15 oz.

This is a rather shy bird and wherever lumbering or gardening tends
to reduce the original forest, they become much reduced in numbers.
In the south part of Nias where there is little original forest left, they
are now quite scarce.

It is perhaps worth noting that Hydrocissa convexa zamelaena Ober-
holser of the North Natuna Ids. based like barussensis on larger size
appears to be a synonym of convexus. The type measures: wing 312;
tail 308. This seems to be another case of a small island population
tending towards larger size than a mainland or large island population.

143. Crannorhinus leucocephalus megistus Oberholser

Tana Massa, Tana Bala. The type, an adult male collected February
1 1 , on Tana Bala measures : wing 433, tail 283. Another male collected

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 3()5

in the same month has a wing of 408, while a female from Tana Massa
measures; wing 381, tail 251. I have compared these measurements
with ten males and three females from Pulu Payong, East Sumatra,
and west and southwest Borneo. They measure: wing cT 352-405
(385.2), 9 347, 347, 354. The largest Borneo bird measures 401.5.
Comparing these measurements by use of the formula of "t" for small
samples (Simpson and Roe, Quantitative Zoology, 1939, p. 211) I
arrived at a figure of 3.07 for the measurements of the males and 8+
for the females. Thus the probability for the samples being from the
same population is about 01.3% showing that the difference is signifi-
cant. More specimens from Sumatra may show that larger birds occur
there, but for the present I feel that megistus should be recognized.

Soft parts are recorded as follows: cf feet blackish green, soles
dirty yellow; cf im. iris brownish yellow, feet deep dull green, bill pale
blood red at base shading into dull greenish yellow at tip; 9 bill gen-
erally yellow, pale toward tip, mottled at base with dull brownish
green. Skin of throat deep dull slaty blue with a few pale blotches,
orbital skin paler than throat, feet grayish olive.

Weight, c? 3.5 lbs.

I have put this race in the species leucocephalus as I consider Ma-
layan, Philippine, and Celebes birds conspecific. Malayan males differ
by having the nape black instead of rufous chestnut but in two birds
of the National Museum's series this color is not pure black. A number
of small brown feathers are interspersed along the margins of the nape
patch. The shape of the bill is exactly similar although the color is
somewhat different. As leucocephalus is the oldest name I should list
the following well marked races :

Crannorhinus leucocephalus corrugatus (Temminck)
Malay Peninsula, Sumatra, Borneo.

C. I. megistus Oberholser
Batu Ids.

C. I. waldeni Sharpe
Guimaras, Negros, Panay.

C. I. leucocephalus (Vieillot)
Camiguin I., Mindanao.

C. I. cassidix (Temminck)
Celebes.

366 bulletin: museum of comparative zoology

Family CAPITONIDAE, Barbets

Two endemic forms are confined to the west Sumatra islands.
Both differ from their closest relatives on Sumatra primarily in having
larger bills with the addition of certain color characters.

144. Chotorea mystacophanes ampala Oberholser

Tana Bala, Tana Massa. The type, an adult male collected February
11, 1903 by Dr. Abbott measures: wing 104, tail 59.5, culmen 38.5.
Two males and two females measure: wing cf 99.5, 102.5, 9 102,
105.5; tail & 52, 60, 9 55.5, 56.5; culmen & 37, 38.5, 9 37.5, 39.5.

The female with the smaller measurements, collected February 28,
is not fully adult as it lacks the blue tinted throat of maturity. Other-
wise, except for a slight reduction in size of the red patch on the
crown, the plumage is complete and must be presumed to be that
assumed after the post ju venal molt.

Soft parts : feet olive.

This race is chiefly distinguished from typical mystacophanes by the
large bill. Both males and females also seem to have a somewhat
larger red crown patch, and in the female the forehead is more tinted
with blue than in the females from Sumatra and the Malay Peninsula.
Birds from Peninsular Siam seem to show a slight decrease in size
indicating that there is a continuous cline here culminating in the large
Batu Island birds.

In spite of Chasen and Kloss' remarks (Treubia, 14, 1932, p. 14.)
I cannot see the validity of the race humei from Borneo

145. Cyanops australis gigantorhina (Oberholser)

Nias. The type, an adult male collected March 26, 1903 measures:
wing 74.5, tail 38.5, culmen 20.5. A series of four males and three
females measure: wing c? 73.5-81 (76.5), 9 74.5, 75, 80.5; tail cf
38.5-41 (40), 9 37, 38.5, 42.5; culmen & 19-20.5 (20.1), 9 20.5, 21,
21.5. Soft parts are marked as: bill black with gray base, black; skin
of throat dull black; feet olive, green.

This race is characterized principally by the size of the bill which is
noticeably larger than in Sumatra birds, four males and three females
of which measure: culmen & 17-17.5 (17.2) 9 18-18.5 (18.3). The
color of the underparts is slightly variable but tends in Nias birds to
be a purer green especially in the males.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 367

Family PICIDAE, Woodpeckers

Eleven forms occur on these islands of which nine are considered to
be endemic. All nine are related to forms found on Sumatra. Of them
six differ primarily by color characters and three by size. Two of the
size races are larger than their congeners, one is smaller. Of the two
forms considered conspecific with their Sumatran relatives, one,
Dryobates hardwickii subsp. shows a slight inclination towards larger
size measurements.

146. Picus puniceus soligae de Schauensee and Ripley

Xias. This race differs from observandus of the Malay Peninsula,
Sumatra, and Borneo by the greatly reduced amount of yellow on the
prolonged occipital crest feathers. In the type and two specimens of
the series in the National Museum, yellow is quite absent. In the five
other specimens in the National Museum's series, yellow is present,
but it is overlaid by the scarlet vermilion feathers of the crown. Three
males and four females measure: wing d71 126, 126.5, 130, 9 126, 128,
130.5, 132.5.

Soft parts : iris deep crimson, red orbital skin pale blue, slaty blue,
leaden blue; bill black above, ochraceous beneath, lower ochre yellow,
yellow; feet dirty yellow, claws horn brown. Weight 3.4 oz.

As Mayr points out (Bull. Raffles Mus. No. 14, 1938, p. 30.) birds
from Borneo average somewhat smaller than Sumatra or Malay
Peninsula birds. Two females and an immature female from Banka
are larger on the average than typical observandus. They measure:
wing 9 132.5, 136.5, 9 im. 138.

This is a bird of rather thick jungle, often found in dense scrub
growth rather close to the ground.

147. Callolophus miniaceus niasense (Buttikofer)

Nias. A male, three females and one unsexed measure : wing o" 124,
9 123, 123.5, 124.5, o 125.5. Birds from the Malay Peninsula and
Sumatra measure: wing tf1 126, 127, 130, 9 126, 126, 129, 130.

As well as being slightly smaller, this race seems to be more brightly
red on the crown and scapulars.

148. Dryobates hardwickii subspec.

Nias. A single male collected March 23, measures: wing 75, tail
33.5, culmen 17. The latter measurement is larger than a series of

368 bulletin: museum of comparative zoology

birds from Java, Sumatra and Borneo which measure: culmen cf
12.5-16 (14.6) 9 13-16.5 (15.0).

A solitary specimen like this would ordinarily have no significance
in itself, but in the case of a west Sumatra island bird, this slightly
larger bill does point I think to a potential speciation effect in this
species. Further specimens may show such to be the case. Perhaps
this is the form recorded by Nieuwenhuisen and Rosenberg as Picus
percussus (1868).

149. Meiglyptes tristis micropterus Hesse
Synonym: Meiglyptes grammithorax microterus Oberholser

Nias. A series of five males and one female from Nias measure:
wing 90.5-94.5, 9 89. Two males and three females from Sumatra
measure: wing cf 93, 98, 9 91, 92.5, 95. Oberholser 's original brief
description (1912, p. 6) of microterus was "smaller". Use of formulas
(Simpson and Roe, I.e. 1939) shows that the probability of these
specimens being drawn from the same sample is greater than .1%
Thus these small differences lack any significance. In color and mark-
ing these birds are identical.

150. Meiglyptes tukki calceuticus Oberholser

Tuangku. This race was described by Oberholser (1912, p. 6) as,
"like Meiglyptes tukki tukki, but much larger". The type and only
specimen, an adult female collected January 23, 1902, measures: wing
112.2, tail 70, culmen 25. Aside from size there are no other significant
differences.

151. Meiglyptes tukki infuscatus Salvador!
Synonym : Meiglyptes tukki hylodromus Oberholser

Nias. The type of hylodromus an adult male taken March 10, from
Nias measures: wing 97.5, culmen 22. It was described without ref-
erence to Salvadori's race. The rest of the series, five males and four
females measure: wing c? 95.5-103 (98.7), 9 91.5-100.5 (97.3),
culmen c? 22-24, 9 21-21.5. These measurements agree well with
typical tukki.

This race differs from tukki of Sumatra by having less well defined
pale bars on the back and scapulars and by having the barring on the
under parts more reduced on the average.

A male of tukki from Billiton I., southeast of Sumatra (U.S.N.M.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 369

180492) seems to be an immature bird. Like the young of Dryobates
the feathers of the crown are tipped with a reddish color, in this case
scarlet. Another mark of immaturity may be the pale color of the
abdomen which lacks the characteristic dark rusty wash of the other
specimens.

152. Meiglyptes tukki batu de Schauensee and Ripley

Pini, Tana Massa. A single male in the National Museum's col-
lection has a wing measurement of 109. It differs from the type of
calceuticus by having the barring on the throat more broad and coarse,
and by having the chest more blackish. The series measured by de
Schauensee (1940) seems to indicate that batu is intermediate in size
between tukki and calceuticus.

153. Micro,pternus brachyurus celaenephis Oberholser

Nias. The type is an adult female taken March 29, 1902 measuring:
wing 109.5, culmen 21.5. Six females of badius from Sumatra, Banka,
and Billiton measure: wing 110-122 (113.9).

As may be seen from the above figures, this race does not differ from
badius by being "somewhat larger", (1912, p. 6). It does differ, how-
ever, by being "darker", although its distinctive characters are as
follows : crown with distinct dark centers to the feathers, upper parts
with heavier darker bars; under parts, breast with brownish black
subterminal spots, stomach, flanks, vent and under tail coverts
heavily barred with dark brownish black.

154. Dryocopus javensis parvus (Richmond)

Simalur. The type, an adult male collected December 3, 1901 by
Dr. Abbott, measures: wing 185, tail 124, culmen 38.5. A series of
six males and five females measure: wing d1 172-185 (177.7), 9 172-
182 (177.2).

Soft parts; iris greenish yellow, straw yellow; bill black; feet leaden.

This race is a complete miniature of typical javensis.

155. Dryocopus javensis buttikoferi (Richmond)

Nias. The type, an adult male taken March 18, 1903 measures:
wing 232.5, tail 167, culmen 55. An adult and an immature female
both collected March 25 measure respectively; wing 241, 215, culmen
53, 49.5.

370 ' bulletin: museum of comparative zoology

Soft parts: cf iris bright yellow, upper mandible black, lower
leaden, feet leaden; im. 9 iris straw yellow, feet dusty leaden.

The only discernible difference between this race and javensis is the
lack of the blackish bars found on the thighs of the typical race.

I doubt if these birds are common on Nias at the present time. The
original forest on the island is not plentiful anywhere, although there
is more of it at the north end of the island.

156. Sasia abnormis magnirostris Hartert

Nias. Two males and a female measure: wing cf 52.5, 53, 9 54;
culmen cf 14, 14.5, 9 15. A male from east Sumatra has a wing of
54 and a culmen measuring 13.5, and eight males from Borneo and
the Malay Peyinsula measure: culmen 12-13.5 (13.1). Hartert (1901)
notes that his birds have deeper bills at the base but I cannot find this
difference. Computing the probability of these culmen measurements
by use of the formula of "t" (Simpson and Roe, I.e., 1939) I arrive at
a figure for "t" of 2.7 which is not particularly significant being more
than .02%. However, as there is no overlap, I believe a larger series
would uphold the significance of these measurements especially as the
measurements of four females from Borneo and the Malay Peninsula
are 13-13.5 (13.4), giving when computed by the "t" formula, a prob-
ability for their being identical of only a very little more than .01%.

Soft parts: iris red; ocular area purple, lilac, lilac-purple; bill, upper
mandible worn brown, lower yellow; feet orange, reddish ochre.

Weight: cf 8.5, 11 grams, 9 11 gr.

In contrast to what Robinson says (Birds Malay Penin., 2, 1928,
p. 115.) I found this bird on Nias, not in deep forest, but in secondary
scrub near cuttings and gardens. In June a male with enlarged testes
was collected.

Family EURYLAIMIDAE, Broadbills

Four forms belonging to two species have been recorded from the
west Sumatra islands. Two of the forms, one from each species, are
endemic, differing by larger size, while the remaining forms are
identical with Sumatra birds.

157. Calyptomena viridis viridis Raffles

Nias, Tana Massa. This species has been taken by Modigliani and
Kannegieter. However, it has not been met with since their time. A

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 371

single male from Tana Massa recorded by de Sehauensee (1940)
apparently does not differ from the typical form.

158. Calyptomena viridis siberu Chasen and Kloss

Siberut, North and South Pagi. A series measures as follows: wing
c? 107.5, cT im. 110, 111, 112 (2), 9 108.5, 112.5, 113. Typical viridis
measures: & 92.5-100.5, 9 98-106.

In size this race agrees with continentis from Peninsular Siam.
However, the single adult male is darker than any specimen of that
form. The females also differ from females of continentis by being
somewhat darker and duller, particularly on the upper parts.

159. Eurylaimus ochromalus mecistus Oberholser

Tuangku. The type, an adult female collected January 29, 1902,
measures: wing 88, culmen 19. Two other females measure: wing
87.5, 87.5, culmen 18.5, 19. These birds are significantly larger than
any specimens of ochromalus measured by me. Apparently they
represent an isolated population on Tuangku, an island lying not more
than twenty-five miles off the west Sumatra coast.

160. Eurylaimus ochromalus ochromalus Raffles

Pini, Tana Massa. Specimens measured by de Sehauensee (1940)
fall within the range of ochromalus rather than mecistus.

wing

culmen

Batu Ids. (f

79.5-83 (81.3)

16.5-18.

9

79.5-84.5 (81.2)

16-20.

Malay Penin., a1

76-84.5 (80.7)

16.5-20.

Sumatra, Borneo 9

74.5-81.5 (78)

16.5-18.5

There seems to be no difference in color between these specimens.

Family PITTIDAE, Pittas

Two pittas occur on these islands, both of which are presumably
migrants rather than residents.

161. Pitta brachyura cyanoptera Temminck
Synonym: Pitta moluccensis lepta Oberholser

Tuangku, Nias, Sipora. The type of lepta is a male adult collected
on Nias, March 15, 1903 .It measures: wing 121, culmen 25.5. A

372 bulletin: museum of comparative zoology

male and female from Nias and Tuangku taken in January and March
measure: wing c? 121, 9 118, culmen cf 26.5, 9 25. The race lepta
was described as being smaller than moluccensis, "especially the bill".
A series of males from Sumatra measure: wing 120.5-132.5, culmen
26.5-29.

Aside from the difference in size being insignificant, Robinson's
theory ("Fasciculi Malayensis," Zool., 3, 1906, p. 96.) that this is
a migratory form rather tends to mitigate against any attempt to
delimit isolated populations.

162. Pitta sordida cucullata Hartlaub

Nias. Another migrant which has been recorded only by
Buttikofer (1896).

Family HIRUNDINIDAE, Swallows

Two species occur on the west Sumatra islands. One is a migrant,
the other conspecific with the resident form of the greater Sunda area.

163. HlRUNDO RUSTICA GUTTURALIS Scopoli

Simalur, Tuangku, Nias, Siberut. A migrant recorded from the
middle of September to the beginning of May.

164. Hirundo tahitica javanica Sparrman
Synonym: Hypurolepis javanica hypolampra Oberholser

Simalur, Nias, Pini, Tello, Enggano. The type of hypolampra is an
adult female collected on Nias, March 22, 1903. It measures: wing
106. This measurement is not "larger" than typical javanica females
which range from 103-108. Nor does this specimen appear strikingly
pale on the lower parts.

I am inclined to agree with Junge (1936, p. 46) who lumps abbotti
with javanica, as there seem to be no constant size or color differences
between the populations in the Greater Sunda area.

This is a common bird around houses and villages. It often rests
under the eaves of European houses. Nesting takes place from early
March at least through June. This species will probably be found on
all the islands of the group.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 373

Family CAMPEPHAGIDAE, Cuckoo-shrikes

Eleven forms have been recorded from these islands. Of them,
seven are endemic, all differing from the Sumatra forms primarily by
size. In four of the forms there are minor color differences which set
them apart one from the other.

165. Coracina striata simalurensis (Richmond)

Simalur. The type, an adult male taken November 19, 1901,
measures: wing 167.5, culmen 27.5, tail 120.5. A series measures:
wing c? 168, 168.5, 169, 9 165, 165.5, 167.5, 9 im. 179; culmen d>
29.5 (2), 30, 9 29, 29.5, 30, 9 im. 28.5.

This race is slightly larger than sumatrensis. The males are slightly
paler gray above and below. The females are paler also and lack the
pronounced barring of the abdomen, vent and under tail coverts found
in sumatrensis. Only on the under tail coverts are the bars distinct.
Elsewhere they are virtually obsolete. The immature female is barred
as in the adults of sumatrensis, but the bars are less black and sharp t

166. Coracina striata babiensis (Richmond)

Babi. The female type collected January 13, 1902 measures: wing
172.5, culmen 30. Another female measures: wing 172, culmen 29.

In general these specimens are very similar to simalurensis. How-
ever, the barring on the under parts is even more reduced than in the
previous race. The barring on the under tail coverts is in a shade of
gray on white rather than black on white. In color this race approaches
somewhat the tone of Coracina papuensis, a mangrove and shoreline
species of Australasia. Presumably the habitat of babiensis in the small
low-lying Banyak Islands is rather similar to that of the widely dis-
tributed papuensis.

167. Coracina striata kannegieteri (Buttikofer)

Nias. This race is evidently similar to sumatrensis but larger. I
have seen no specimens.

168. Coracina striata sumatrensis (S. Miiller)
Synonym: Graucalus crissalis Salvadori
Synonym: Artamides sumatrensis halistephis Oberholser

Siberut, Sipora, North and South Pagi. I cannot see any difference
between these birds and a series of sumatrensis from'Pulu Tioman, the

374 bulletin: museum of comparative zoology

Rhio Archipelago, and Great Karimon Island, east Sumatra. The
males are the same shade of dark gray on the upper and lower parts.
The females have the same type of barring on the underparts. In size
there is too much overlap to provide a distinctive character for
separation.

Comparative measurements follow :

wing

culmen

Siberut d"

161.5, 165.5

26, 26.5

9

161

26

Sipora c?

169, 170.5, 172

26, 27, 27

9 (6)

162-170 (166.4)

26-28 (27.1)

North Pagi d*

—

—

9

158

28.5

South Pagi & (type

! of halistephis)

166

25

c?

161

26.5

9

162.5

26.5

Sumatra, etc. d*

160-167.5 (164.1)

27.5-29 (28.3)

9

158-168 (160.7)

25.5-27.5 (26.6)

169. Coracina striata enganensis (Salvadori)

Enggano. These birds are very close to simalurensis both in size,
which varies rather widely, and in color. The females, however,
differ by having slightly less obsolete barring on the stomach and
vent. Also the barring on the under tail coverts is slightly wider and
distinctly more blackish than in simalurensis.

Three males, four females and an immature female measure: wing
cf 166, 169, 173, 9 167, 167 (moult), 174.5, 179, 9 imm. 168; culmen
cf 28.5 (2), 30, 9 26, 27, 28, 9 im. 26.5.

170. Volvocivora fimbriata compta (Richmond)

Simalur. Siberut (?). Two specimens were collected in November
and December by Abbott. The type, a not quite adult female, measures :
wing 103, tail 83, culmen 14. Another female measures: wing 100,
tail 82.5, culmen 14.5.

Compared to females of culminata this race is larger. Three females
from Sumatra and Borneo measure, wing 92, 93, 93.5.

A male from Siberut agrees is size with these specimens rather than
with males of culminata (wing 103 compared to 92,93, 94.5,96 for of d*
from Sumatra and Borneo). This bird does not seem darker than
Sumatra and Borneo birds, but lacking males of compta I must pro-
visionally assign it to this race.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 375

171. Lalage nigra nigra (Forster)
Synonym: Lalage nigra empheris Oberholser

Nias. The type of empheris, an adult male collected March 2,
measures: wing 88, tail 6S, culmen 15. The race was originally based
on the character of a paler rump, but I cannot find that the type
supports this contention.

172. Hemipus hirundinaceus (Temminck)

Recorded from Simalur by Rosenberg (1878) and from Nias by
Xieuwenhuisen and Rosenberg (1868).

173. Pericrocotus flammeus minythomelas Oberholser

Simalur. The type, an adult male collected December 12, 1901
measures: wing 94, tail 87, culmen 14.5. A series measures: wing d1
91.5, 92, 93, 93.5, 94, 94, 9 89.5, 90, 91.

This seems to be a straight size race. In series as Junge notes (1938),
the females only show color differences, not the males; see also his
1936 paper, p. 53.

174. Pericrocotus flammeus modiglianii Salvadori

Enggano. Fourteen male adults, one male immature and four
females collected on Enggano by Abbott agree well with Junge's dis-
cussion (1938, p. 350). This race is even larger than minythomelas,
wing c?<? 93.5-97.5, cT im. 94.5, 9 9 92.5-94.5. The males are some-
what more yellowish on the under surface, not as pure vermilion as in
xanthogaster. The color of the underparts of the females is inter-
mediate between xanthogaster and minythomelas, while the upper sur-
face is more like xanthogaster. Except for the fact that the under parts
of the young male are richer, somewhat more orange-tinted, particu-
larly on the tail feathers, there is no difference between this plumage
and that of the adult female. Specimens collected in November and
December.

176. Pericrocotus igneus igneus Blyth

Nias. A single record in Buttikofer (1896) is the only one for this
form in the west Sumatran islands. The bird was a female taken on
north Nias in the fall of 1895 and is listed by Junge (I.e. 1936) among
his material of the nominate race which he compared to trophis.

376 bulletin: museum of comparative zoology

Family DICRURIDAE, Drongos

There are six resident drongos on these islands. Two are conspecific
with Sumatra birds. One differs by size from a Sumatra race, and one
differs by size and color from its nearest relative on Sumatra. The re-
maining two races are close to one found on Borneo, differing from it
in larger bill size and color.

177. Dicrurus leucophaeus leucophaeus Vieillot
Synonym: Dicrurus cineraceus celaenus Oberholser

Simalur. The type of celaenus, an adult male collected November
27, 1901 measures: wing 138.5, tail 131.5, culmen 22.5. In spite of the
original description of Oberholser (1912, p. 15), "darker, particularly
on lower surface", and Junge's supporting view (1936, p. 62.), I cannot
agree that the Simalur birds differ in color from those of Java. There
is no distinct difference in size although I have measured no Javan
specimen with a wing as long as 138. It is interesting indeed that these
birds should resemble the dark Javan race rather than the pale-lored
Sumatran forms or the paler white eye-ringed birds from the islands
farther to the south.

178. Dicrurus leucophaeus siberu Chasen & Kloss

Siberut. Four males and one female of this race measure: wing cf
132, 135, 139, 141, 9 130.5; culmen c? 22-23.5, 9 23.5. This form
has somewhat more white behind the eyes and is a trifle paler on the
back. Otherwise, except for its larger bill it is very close to stigmatops
from Borneo.

The appearance of this white eye-ringed form on the west Sumatra
islands is a good argument for including the former species lencogenis
in leucophaeus. Presumably this color pattern is carried recessively
in the uniform gray Sumatran and Javan representatives.

179. Dicrurus leucophaeus periophthalmicus (Salvadori)
Synonym: Dicrurus leucogenis diporus Oberholser

Sipora and the Pagi Islands. In the original description of diporus
(1912, p. 15) Oberholser made no mention of Salvadori's race from
Sipora with which Pagi birds are identical. The type of diporus, an
adult male from North Pagi collected November 14, 1902, measures:

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 377

wing 142, tail 127, oilmen 24. This race is paler than siberu with, in
most specimens, more white carried back from the eyes and down onto
the sides of the neck. The iris is marked as, "carmine", "reddish".
Moulting birds were taken in September and October.

ISO. DlCRURUS HOTTENTOTIUS VIRIDINITENS (Salvadori)

Siberut, Sipora, and the Pagi Islands. A series of specimens have
wing measurements as follows: cf 140 (worn)-152.5 (149.1) 9 139-
148 (142.3). The iris of adult birds is marked as, "deep red" or "crim-
son". An immature bird taken on South Pagi in September has the
iris marked "brown". Another immature was taken in September on
Siberut. See Chasen and Kloss (192G, p. 293) for a discussion of the
differences between this race and the Sumatran race.

181. DlSSEMURUS PARADISEUS PLATURUS (Vieillot)

Synonym: Dissemurus paradiseus olizurus Oberholser
Synonym: Dissemurus paradiseus pachistus Oberholser
Synonym: Dissemurus paradiseus elassopterus Oberholser

Simalur, Lasia and Babi. The type of olizurus an adult male from
Simalur taken November 19, 1901, measures: wing 140, tail 305,
culmen 29.5. The type of pachistus an adult male from Lasia collected
January 5, 1902, measures: wing 155.5, tail 367, culmen 30. The type
of elassopterus, an adult male collected January 11, 1902 on Babi,
measures: wing 154.5, tail imperfect, culmen 31.

Other specimens from these localities have the following measure-
ments (excluding the tail which is too variable) :

wing

culmen

Simalur a"

143

29

9

134.5, 141, 141.5, 144.5

26.5, 27, 27, 28

Lasia d1

153.5 (2)

27.5, 28

Babi d"

147 (worn)

30.5

9

144

29.5

I agree with Junge (1936, p. 63.) that it is impossible to separate
these birds from the Sumatran platurus, including pachistus which
Junge did not mention, presumably lacking Lasia specimens. The
worn bird was collected in January.

378 ' bulletin: museum of comparative zoology

182. Dissemurus paradiseus adelphus Oberholser

Nias. The type of this race, an adult male collected March 5, 1905,
measures: wing 157, tail 347, culmen 33. A small series measures:
wing <? 153.5, 157.5, 9 147.5, 148.5, 151; culmen d1 39.5, 31, 9 28,
29.5.

These birds are larger than any measured by Kloss (Treubia, 13,
1931, p. 359.), and so should be kept as a separate race. Oberholser's
other characters of large crest feathers, bristles, racquet feather, etc.,
are, I feel, not particularly diagnostic.

Family ORIOLIDAE, Old World Orioles

Four subspecies inhabit the west Sumatra islands, three of which
differ from the Sumatra form by larger size and minor color characters.
The remaining subspecies is based on color, showing close Bornean
affinities.

183. Oriolus chinensis mundus Richmond

Simalur. The type is an adult male collected November 19, 1901.
It measures: wing 151, tail 103.5, culmen 35. Four other males meas-
ure: wing 148-154 (150.7). Besides being larger than maculatus these
birds have a much purer, more lemon, yellow colored back.

I cannot see Chasen's race edgari from the lower Malay Peninsula
and Sumatra. A female from north west Sumatra and another from
Banka both have bill measurements (from gape) of 34, thus falling well
within the measurements of maculatus. As this is the only cited
difference (Treubia, 18, Suppl., 1941, p. 118.), I fail to see how this
race can be maintained.

There is a record for this species from Nias in Buttikofer (1896).

184. Oriolus chinensis richmondi Oberholser
Synonym: Oriolus chinensis siberu Chasen and Kloss

Siberut and the Pagi Islands. The type, an adult male collected on
North Pagi December 31, 1902, measures: wing 150, tail 97.5, culmen
34. Two other males and a female measure: wing c? 145, 152.5, 9
148. Birds from Siberut measure: wing cf 149.5, 150.5, 152, 9 141.5,
148.5.

Chasen and Kloss (1926, p. 294) described siberu as being larger
and greener than richmondi without citing any figures to support their

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 379

contention. They must have been comparing males from Siberut
which were not fully adult. A single male in the U. S. National Mu-
seum's Siberut series is fully adult and is indistinguishable from
richmondi. It is quite as bright orange yellow as Pagi birds on both
upper and lower surfaces. Females and immature birds are likewise
inseparable.

185. Oriolus chinensis sipora Chasen and Kloss

Sipora. Two males and two females measure: wing c? 153 (2), 9
148, 149. These birds are very slightly larger than richmondi but they
differ primarily by being brighter on the back with broader yellow
edgings on the wings. From mundus this race is distinguished by
having a yellow speculum patch, broader yellow margins on the
secondaries, and by being somewhat more greenish above. It is curious
that this race should occur between the neighboring islands of Siberut
and the Pagi group where richmondi occurs. The inference seems
likely that Siberut and the Pagi Islands are inhabited by actual geno-
typically separable populations which have the misfortune from the
taxonomic point of view of being phenotypically indistinguishable.

186. Oriolus xanthonotus mentawi Chasen and Kloss

Siberut and Sipora. Chasen and Kloss neglected to compare this
form in their original description (1926, p. 295) with consobrinus of
Borneo. The single adult female from Siberut in the National Mu-
seum's collection is inseparable from a female from south-east Borneo
(U.S.N.M. No. 181520.). However, the two adult males from Siberut
are somewhat more boldly streaked on the breast and abdomen than
are two Borneo birds.

I think this is a poor race which might be shown to be inseparable
from consobrinus given more material. One of the females of the latter
form from the Segah river, east Borneo, is very dark gray on the
throat and the crown has broad black centers to the feathers making
it much darker than any other specimen examined.

An immature male was collected on Sipora in October.

Colors of adult: "iris crimson, bill pinkish brown, feet grayish black".

Family CORVIDAE, CROWS

Three crows occur on these islands. One is rare throughout the
Greater Sunda area. Of the other two, one shows a slight tendency

380 bulletin: museum of comparative zoology

towards a larger bill compared with specimens from Sumatra and
Borneo, while the other form is identical with the subspecies found on
Java.

187. Corvus enca compilator Richmond

Simalur, Nias. Specimens from these two islands have somewhat
larger bills than Sumatran and Bornean specimens, but I have not
seen a large enough series of birds from the latter islands to determine
whether or not this difference is constant. Following are the measure-
ments of seven males and five females from Simalur and Nias as con-
trasted with three males and three females from Sumatra and Borneo.

wing tail culmen

Simalur, Nias d" 307-321.5(313.1) 146.5-170(157.5) 63-69.5(66.1)

9 282 (\vorn)-310.5 (301.) 142.5-151 (146) 63-65.5 (63.8)

Sumatra, Borneo <? 303-332 151-166 60.6, 62.5, 64

9 303.5-318 148-169 59, 59.5, 63

The male with the largest bill measurement is from Simalur and
the tip of the bill is broken as is that of another male from Nias with a
bill measurement of 66.5.

In color these birds appear to be slightly more blackish, less purplish
on the under surface than Borneo and Sumatra birds. The iris of these
birds is marked as dark brown. A Nias female weighed 14.9 oz.

This bird on Nias was somewhat shy and suspicious of humans. I
never saw it out on the reefs or beaches where orru sometimes goes.

188. Corvus enca enca (Horsfield)

Siberut, Sipora. A small series from these islands measures: wing
d1 276.5, 287.5; 9 273, 281, 293.5; culmen & 56, 59; 9 53, 54 (2).
The large-winged female is unusual in this respect but I note that
Meinertzhagen (Novit. Zool., 33, 1926, p. 71.) gives wing measure-
ments for this form up to 299.

A female collected in October has worn primaries and newly moulted
rectrices. A name on one of the labels may be a native name : "mengga".

189. Corvus macrorhynchos macrorhynchos Wagler

Recorded from Nias by Buttikofer (1896, p. 188). This species is
apparently uncommon in the area, as it is listed by Meinertzhagen
(Novit. Zool., 33, 1926, p. 85.) as "very rare" throughout Sumatra.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 381

Family SITTIDAE, Nuthatches

A single representative of this family has been recently found on
Simalur. It does not appear to differ from the Sumatran subspecies.

190. Sitta frontalis saturatior Hartert

Simalur. Junge records this form from Simalur (1936, p. 60.) where
it was collected for the first time by Jacobson and van Heurn. He
notes that it was locally common, sometimes occurring in small family
flocks in high open forest near clearings.

Family TIMALIIDAE, Babblers

Eleven babblers occur on these islands of which five belong to
endemic races. The characters which separate these races from their
nearest relatives on Sumatra are: size, two forms; color, one form;
size and color, two forms.

191. Malacopteron cinereum niasense (Riley)

Nias. The type, an adult male taken by Abbott, March 10, 1905,
measures : wing 80, tail 64, culmen 18. Two other males, a female and
one unsexed measure: wing cf 80.5; 81.5, 9 73, - 77.5; culmen d1
16.5, 17, o 17.

This is a distinctly larger race than that from the Malay Peninsula,
Sumatra, and Borneo. The chestnut forecrown and tail are also some-
what darker. Soft parts : "iris crimson, reddish brown ; upper mandible
black, lower bluish pink; feet bluish pink, pale bluish flesh". Males
with enlarged gonads taken in June.

As pointed out by de Schauensee and myself (1939, p. 408.) this is
the species listed by Salyadori from Nias, not magnum.

192. Malacopteron affine notatum Richmond

Bangkaru. The type, an adult male collected January 17, 1902 by
Abbott, measures: wing 80, tail 68, culmen 17. Three other males and
a female measure: wing cf 75, 77,' 78, 9 73; culmen cf 15, 16, 17, 9
15.5.

Richmond's original description (1902, p. 190.) mentions that this
form differs from affine by having a sooty black cap instead of a brown-
ish one. Actually this is not true. Both birds have a sooty black cap

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in fresh plumage, but notatum is somewhat paler on the upper surface
and partially lacks the dark chest band characteristic of the nominate
race. Also it is larger in size and has a larger bill. Soft parts: "iris
brown, feet leaden."

193. Malacopteron albogulare albogulare (Blyth)

Pini. Two specimens listed by de Schauensee (1940, p. 36.) are the
only records for the west Sumatra islands. His remarks indicate that
the specimens are intermediate between albogulare and moultoni but
more specimens are needed in order to determine the status of this
population.

194. Anuropsis malaccensis malaccensis (Hartlaub)
Synonym: Anuropsis malaccensis exsanguis Oberholser

Synonym: Anuropsis malaccensis nesitis Oberholser

Tuangku, Tana Massa. The type of exsanguis, an adult male from
Tuangku collected January 24, 1902 measures: wing 72.5, tail 35,
culmen 17.5. The type of nesitis, an adult male from Tana Massa
collected February 20, 1903, measures: wing 77, tail 36, culmen 17.5.
A male, a female, and a specimen of undetermined sex from Tana
Massa measure: wing cf 78, 9 67.5, o 69; tail c? 37; culmen d71 17,
9 15, o 15.5. Sumatra and Malay Peninsula males have wings meas-
uring up to 74, and tail measurements up to 37. The culmen measure-
ments reach 17. In color Tana Massa birds are very slightly richer
more rufescent above and more buffy on the flanks and vent. The
Tuangku specimen is exactly intermediate if such fine distinctions are
possible. I concur with Riley (unpublished MSS.) in his suppression
of exsanguis but I should like to carry this a little further and declare
both forms synonyms of the nominate race. I feel that these minor
differences in size and color are too overlapping to indicate more than
a trend in speciation away from the mainland and large island birds
rather than a positive achievement.

195. Alcippe brunneicauda, brunneicauda (Salvadori)
Synonym: Alcippe cinerea hypocneca Oberholser

Pini. The type of hypocneca is an apparently adult bird of undeter-
mined sex collected March 4, 1903. It measures: wing 73, tail 59,
culmen 13. A male also collected in March measures: wing 70.5, tail

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS •">*;!

59.5, culmen 13.5. The soft parts are marked as iris and feet grey, bill
brown. In measurements a series of Malay Peninsula and Sumatra
birds completely overlap these two specimens, so that they can hardly
be "smaller" as Oberholser (1912, p. 8) diagnoses this race. I suggest
further that the fact that the two birds are in rather worn plumage
accounts for any color differences that may be present.

196. Stachyris maculata banjakensis Richmond

Tuangku. The type is an adult male collected January 24, 1902 by
Dr. Abbott. It measures: wing 91, tail 69, culmen 21. Another male
measures wing 93, tail 72, culmen 20. These measurements are con-
siderably larger than those for the nominate form from Sumatra: wing
cf 81, 84.5, 9 82; tail c? 67, 9 64; culmen c? 17.5, 20, 9 18. Color
differences are very slight. These birds are somewhat more grayish on
the upper surface but two specimens is not a large enough series to
establish this as a character.

197. Stachyris maculata hypopyrrha Oberholser

Pini, Tana Massa. The type is an adult male collected March 6,
1903 on Pini. It measures: wing 85, tail 65.5, culmen 18.5. This small
series consisting of the type, a female, and two birds of undetermined
sex, agrees well with Oberholser's original description (1912, p. 9) in
being slightly more rusty on the upper surface and on the lower flanks
and abdomen than a series of three birds from Sumatra. There is no
size difference, and the color difference is so slight that a larger series
may show that these characters are not constant.

198. Cyanoderma erythroptera fulviventris Richmond

Tuangku. The type, an adult male collected February 1, 1902,
measures: wing 61, tail 47, culmen 17. Another male and two females
measure: wing cf 63, 9 59, 60; culmen cf 16.5, 9 15, 15.5. Two speci-
mens from Sumatra measure; wing cf (type of eripella) 60.5, 9 57;
culmen cf 14.5, 9 14.5.

Besides having larger bills, these birds do present a somewhat more
fulvous appearance on the upper surface and on the flanks and lower
abdomen. Soft parts of the type are: iris brownish red, throat skin
pale blue. The feet of the females are noted as, "pale greenish brown"
and "pale brownish fleshy".

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199. Cyanoderma erythroptera erythroptera (Blyth)
Synonym: Cyanoderma erythroptera pella Oberholser

Tana Massa, Tana Bala. The type of pella, an adult male collected
on Tana Massa February 20, 1903, measures: wing 65, tail 47, culmen
15.5. Two females measure: wing 58.5, 62; culmen 14.5, 16.

These birds are fairly large but the measurements indicate a good
deal of variability in size.

In color there is no tenable distinction between these specimens and
erythroptera with which they were not compared in the original de-
scription (1012, p. 9). The soft parts of one of the females are given
as follows : "orbital skin deep cobalt, throat pale whitish blue, feet pale
olive yellow".

200. Mixornis gularis gularis (Horsfield)

Synonym: Mixornis gularis zarhabdota Oberholser
Synonym: Mixornis gularis zaptera Oberholser

Bangkaru, Tana Massa, Tana Bala. The type of zarhabdota, an
adult male from Bangkaru measures: wing 61.5, tail 52.5, culmen 15.
It was collected by Dr. Abbott, January 19, 1902. The type of zaptera,
an adult male collected February 17, 1903 on Tana Massa measures:
wing 63, tail 55, culmen 15.5. A series of gularis from the Malay
Peninsula and Sumatra measures: cf wing 58-64, culmen 14-16.

I cannot agree with de Schauensee (1940, p. 37) that Batu birds
are more heavily streaked below than ordinary gularis. The variation
in the amount and type of streaking is considerable. The type and
unique specimen of zarhabdota is rather rufous on the upper surface
but this condition can be pretty well matched by Rhio Island birds
and must be considered to be due to individual variation. Soft parts
are indicated as: "skin of ocular area cobalt, feet yellow olive".

An immature female was taken on Tana Massa February 20, 1903.
The under parts are grayish buffy except for the throat and center
of the breast where the adult yellow feathers with central shaft streaks
are beginning to appear.

201. Macronus ptilosus ptilosus Jardine and Selby
Synonym: Macronus ptilosus batuensis Riley

Tana Bala. The type of batuensis is an adult male taken February
13, 1903 by Dr. Abbott. It measures: wing 73.5, tail 65, culmen 17.
Another specimen, unsexed, measures: wing 71.5, tail 59, culmen 17.5.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 385

A series of males from Sumatra and the Malay Peninsula have wing
measurements from 67-71.5 and culmen measurements from 16-17.5
so that I do not feel that the size of these Batu birds is particularly
significant.

Of the two specimens the type has a considerable amount of gray
on the abdomen but this can be matched by individuals in the series
of ptilosus. I am inclined to think that the amount of gray in this area
is somewhat due to the makeup of the skins.

Soft parts: "skin of throat deep cobalt verging to turquoise on neck
and eyelids".

Family PYCNONOTIDAE, Bulbuls

A total of fifteen members of this family have been collected on the
west Sumatra islands. Of these only two are endemic, although five
populations which are considered conspecific with those of Sumatra
do show a tendency towards larger size. Of the two endemic races
both show slight color differences compared to their relatives on
Sumatra, while one is also larger.

202. Aegithina viridissima viridissima (Bonaparte)
Synonym: Aegithina viridissima nesiotica Oberholser

Tana Massa, Tana Bala. The type of nesiotica is an adult male
collected on Tana Bala February 5, 1903 by Abbott. It measures:
wing 62, tail 42, culmen 15. I agree with de Schauensee (1940, p. 34.)
that this is not a tenable race.

203. Aegithina tiphia horizoptera Oberholser

Nias. The type is an adult male collected on Nias March 3, 1905.
It measures : wing 62.5, tail 48.5, culmen 16.5. The soft parts are noted
as: "iris whitish; upper mandible black, lower blue gray; feet blue
gray." As was pointed out by de Schauensee and myself (Proc. Acad.
Nat. Sci. Phila., 91, 1939, p. 346.) horizoptera must stand as the
name for birds from Peninsular Siam, northern Malay Peninsula,
Sumatra and Banka as well as Nias.

204. Chloropsis sonnerati parvirostris Hartert

Nias. A single male in the collection of the Academy of Natural
Sciences measures: wing 98, tail 69, culmen 22.5. Its weight was 45
grams and the soft parts are marked as: "iris brown, feet gray".

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This bird was in breeding condition when it was collected (June
10.). It was perched in a tall tree overlooking a cleared garden and
was singing in a rich burbling manner reminiscent to me, of the
Baltimore Oriole.

205. Chloropsis sonnerati zosterops Vigors

Pini, Tana Massa. A series in the Academy of Natural Sciences'
collection from these islands measure: wing c? 101, 106, 106.5, 110,
9 99.5, 100, 102. This is larger than the average of zosterops from
other localities, although the United States National Museum has
specimens from Trang reaching 104. Certainly, however, the Batu
Island birds represent a tendency towards larger size.

206. Irena puella criniger Sharpe
Synonym: Glauconympha cyanea megacyanea Oberholser
Synonym: Irena puella bondi de Schauensee

Tuangku, Nias, Tana Massa, Tana Bala, Siberut, Sipora, South
Pagi. Oberholser's type of megacyanea is an adult female from
Tuangku, taken January 23, 1902. This specimen measures : wing 121,
tail 84, culmen 25. Soft parts are noted as follows: "iris red, bill and
feet black."

The type of bondi is an adult male (A.N.S.P. no. 56496) taken on
Tana Massa, which measures: wing 127, tail 85, culmen 25. Fifteen
males from these islands measure, wing 121-134 (124.2) while eleven
females measure, wing 115-128 (122.5). Batu birds alone measure:
wing c? 121-134 (124.1), 9 120-128 (123.9). A series of ten males
from Borneo, Banka and Sumatra measure: wing 117-126 (120.7).

I feel, therefore, that although a size difference is indicated, it is
too small to merit naming.

Three males from Tuangku, Tana- Bala and Siberut are molting
into adult plumage (September and February). The irides of these
birds are marked: "red, deep orange red". Another specimen from
South Pagi taken in December is in the full immature plumage of the
male which is similar to that of the female. The iris of this specimen
is marked as "bright brownish yellow," while that of another immature
male from Sipora (collected in December) is indicated as brown.

In the molting specimens the black feathers of the under parts seem
to appear indiscriminately. On the upper surface in the three speci-
mens examined the metallic cobalt feathers seem to appear first on
the crown, nape and upper tail coverts.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 387

207. Microscelis charlottae crypta (Oberholser)

Tana Massa. Two specimens, a female in the Academy of Natural
Sciences' collection, and an adult, sex undetermined, taken by Abbott
in February 1903, are the only ones recorded from the west Sumatran
islands. They measure: wing 9 82, o 91, culmen o 18. The iris was
marked, "dull ochreous".

For the nomenclature of this form see Ripley (Auk, 1943).

208. Brachypodius atriceps hyperemnus Oberholser

Simalur. The type is an adult male collected November 22, 1901,
which measures: wing 80, tail 68, culmen 15. Of the series of eleven
specimens, only four show the darker lower parts and upper surface
given by Oberholser as part of his diagnosis of this race (1912, p. 10).
However, these birds are somewhat larger than typical atriceps; wing
c?1 78.5-84, 9 81.5; culmen c? 12.5-14, 9 13-14.

Junge (1936, p. 55) notes that specimens of this form were breeding
in January. These birds are found in gardens and cleared land.

209. Brachypodius atriceps atriceps (Temminck)
Synonym: Microtarsus melanocephalus chrysophorus Oberholser

Nias, Siberut, Sipora, South Pagi. The type of chrysophorus is an
adult male taken on South Pagi November 15, 1902. It measures:
wing 79.5, tail 66, culmen 14.

Oberholser's original description (1912, p. 10) of chrysophorus men-
tioned that the rump and lower parts were more golden but I have
been unable to perceive this character. Nias birds were in breeding
condition in June.

Soft parts: "iris china blue; bill black, feet dark brown."

210. Microtarsus melanoleucos Eyton
Synonym: Microtarsus melanoleucos proximus Riley

Siberut. Chasen (1935, p. 195) states that there is no difference be-
tween fresh skins of this population and fresh skins from other locali-
ties. Presumably he is right as this is a species which foxes rapidly
and Riley's series from Siberut was of fresh material compared with
which Bornean birds, taken twenty years previously, look rather
brownish.

Soft parts: "iris crimson, bill black, feet blackish brown."

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211. Tricholestes criniger sericea Robinson and Kloss

Pini, Tana Massa, Tana Bala. These islands are the only ones where
this common bulbul has been found.

212. Trachycomus zeylanicus (Gmelin)
Nias. Listed as occurring there by Buttikofer (1896, p. 197).

213. Pycnonotus plumosus porphyreus Oberholser

Tuangku, Nias, Tello, Tana Massa, Tana Bala, Siberut, Sipora,
Pagi Ids. Males from the Batu Islands have large wing measurements
(up to 92 mm.) but this is a tendency, no more. Nias birds are recorded
with weights of 36, 37 gr. and were in breeding condition in June. The
iris of a large series from these islands is recorded as : "ochreous, orange,
brownish yellow, yellow, bright ochre."

214. Pycnonotus simplex simplex Lesson

Nias, Pini. Three females from Nias agree in size with mainland
birds (their weights are given as 17, 22 gr.). A single female from Pini
measures: wing 84, tail 73, culmen 14. The wing and tail measure-
ments are larger than any other specimens measured from the Malay
Peninsula, Sumatra, etc. Perhaps more specimens may show whether
this is more than a trend towards larger size. The irides of these birds
are noted as white.

215. Pycnonotus brunneus brunneus Blyth

Tuangku, Bangkaru, Nias, Tana Massa, Tana Bala. These birds
are indistinguishable from mainland and Sumatra birds although the
wing measurements reach a slightly larger size.

Weight (Nias) 25, 28, 30, 33 gr. Testes enlarged in June. "Iris red,
orange-red, pink, orange."

216. Pycnonotus erythropthalmus erythropthalmus (Hume)
Synonym: Pycnonotus erythropthalmus cyanochrus Oberholser

Synonym: Pycnonotus erythropthalmus isus Oberholser
Synonym: Pycnonotus erythropthalmus pammicrus Oberholser

Tuangku, Nias, Tana Massa. These three races of Oberholser's
(1912, p. 10, 11) were described primarily on the basis of size although
no measurements were given. Actually there is no significant variation.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 389

The type of cyanochrus is an adult male from Rupat Strait, east
Sumatra, taken February 27, 1906. It measures: wing 75, tail 67,
culinen 12.5. The type of isus is an adult male from Tuangku, taken
January 25, 1902, measuring: wing 79, tail 64, eulmen 13. The type
of pammicrus is an adult female taken on Nias March 15, 1905, It
measures: wing 67.5, tail 5S, eulmen 12.5.

A female from Tana Massa measures : wing 71.5 while three Sumatra
females measure: wing 71, 71.5, 73. These measurements are too close
to the single Nias bird to allow the recognition of pammicrus. I can
see no valid color differences between these specimens.

Soft parts: "iris red, eyelids orange-yellow".

Family TURDIDAE, Thrushes

Eleven species and subspecies are found on these islands of which
one is a migrant. Of the ten remaining forms, eight are endemic. Of
these races two are straight size races, being larger than their Sumatran
relative, four are color races, and two embody both color differences
and larger size.

217. Enicurus leschenaulti frontalis Blyth

Nias. Recorded from Nias only by Salvadori (1887, p. 40). These
birds may be referable to the following race as no specimens have
been available for comparison.

218. Enicurus leschenaulti chaseni de Schauensee

Tana Massa. The type and unique specimen is in the Academy of
Natural Sciences' collection. It is an adult male collected on Tana
Massa, September 7, 1896 by J. F. Kannegieter. It measures: wing
104.5, tail 130.5, eulmen 22.5. This race is larger than frontalis of
Sumatra and the Malay Peninsula (wing cf 90-93), and smaller than
leschenaulti from Java (wing cf 110). In color there are no differences.

219. Copsychus saularis zacnecus Oberholser

Simalur. The type is an adult male collected December 2, 1901. It
measures: wing 101.5, tail 89, eulmen 20. Two other specimens meas-
ure: wing d" 106.5, 9 97.5, eulmen & 21, 9 19.

This is a good race, differing from musicus of Sumatra and the
Malay Peninsula in the male by the presence of more grayish-buffy

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flanks, and in the female by darker upper parts and darker throat and
breast, and by having more buffy-gray on the abdomen and flanks.
There is no size difference.

A young male moulting from immature to adult plumage was taken
in December. The nape is still sooty but the rest of the back and
crown is glossy blue black. Some of the throat and upper breast
feathers are still immature, but otherwise the rest of the breast and
the center of the throat is blue-black. The belly is nearly white.

A juvenal male collected in October has the upper parts sooty ex-
cept for a patch of blue black in the center of the back extending out
onto the scapulars. The throat and breast feathers are mouse-brown
with dull white centers. The belly is buffy white. Junge (1936, p. 56)
records a clutch of eggs taken in May and gives their measurements.
The nest was found in a pandanus, and the bird is noted as being
abundant in open country.

The soft parts of the type are marked as: "iris dark brown, bill and
feet black".

220. Copsychus saularis nesiarchus Oberholser

Nias. The type is an adult male taken March 22, 1903 and measur-
ing: wing 105, tail 91, culmen 21.5. Another male measures: wing
106, culmen 18.5. This form has more white on the third and fourth
rectrices than any qualified specimens of musicus examined. Otherwise
these birds are indistinguishable Presumably, however, the females
are darker than females of the mainland form. From zacnecus this
race differs by having less of the buffy-gray wash on the flanks and
vent. Testes enlarged in June. Weight 45 gr.

I found this bird rather shy and uncommon on Nias. The only
specimens I saw were in cocoanut groves along the shore near
Telokdalem.

221. Copsychus saularis masculus Ripley

Pini, Tello, Tana Massa. The type is an adult female collected in
September on Tana Massa (A.N.S.P. no. 56670). It measures: wing
105.5, tail 92, culmen 21. Other specimens measure: wing cf 105-
108.5, 9 102, 103; culmen c? 19.5-21, 9 20.

This race differs from musicus in the male by slightly larger size,
and in the female by large size and darker more glossy upper parts,
and darker more blackish-gray on the throat and breast. From

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 391

nesiarchus this race differs by having less white on the third and fourth
rectrices. From zacnccus this race differs as musicus, lacking the buffy
color on the underparts of the male and female.
An immature female was taken in September.

222. Copsychus saularis pagiensis Richmond

Siberut, Sipora, North Pagi. An adult male collected December 22,
1902 on North Pagi is the type of this race. It measures: wing 119.5,
tail 100, culmen 24. Two females measure: wing 109.5, 111.5, culmen
20.5, 22.

This is the largest of the races from the western islands, differing
from all others primarily in size. The type has more white on the tail
than any other specimens except those from Nias. In this respect it
differs from musicus or masculus. These specimens lack the buffy
wash of zacnecus on the underparts. The females of this race differ
from those of masculus not only by being larger but also by being
slightly darker and more glossy on the upper parts and slightly darker
on the throat and breast.

I cannot see the color differences mentioned by Richmond in his
original description (1912, p. 105).

223. KlTTACINCLA MALABARICA MELANURA (Salvadori)

Synonym: Kittacincla melanura hypoliza Oberholser

Synonym: Kittacincla melanura opisthochra Oberholser

Synonym: Kittacincla melanura pagensis Oberholser

Simalur, Lasia, Babi, Nias, Siberut, Sipora, and the Pagi Islands.
This race is characterized by lacking prominent white outer rectrices,
and by darker lower parts in the male. Females differ from those of
tricolor, the Sumatran and Javan race, by lacking the white outer
rectrices, except for small pale tips, and by having glossy blue-black
upper parts throat and breast. Thus one of the important characters
of this race is the lack of any very pronounced sexual dimorphism, a
tendency that was seen also in Copsychus. In fact, except for slightly
smaller size and shorter tails, females of K. m. melanura may be dis-
tinguished from males only by their having slightly paler chestnut
under parts.

Oberholser's three races are based on size and color differences

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which are not impressive. I agree with Riley (1929, p. 29) that they
are not worth recognizing. The type of hypoliza is an adult male col-
lected on Simalur January 3, 1902. It measures: wing 92.5, culmen
171. Two other males and two females measure: wing cf 89, 91.5, 9
86, 87; culmen cf 16, 17, 9 15, 16. The type of opisthochra is an adult
female collected on Lasia January 7, 1902. It measures: wing 88.5,
culmen 17.5. A male from Babi measures: wing 95.5, culmen 17. The
type of pagensis taken January 9, 1903 on North Pagi, is an immature
male molting into adult plumage. It measures: wing 86, culmen 16.
Two males from Sipora and an unsexed bird from South Pagi are im-
mature. The Sipora birds which are largely in adult plumage were
collected in October. The South Pagi bird is molting from nestling
into immature plumage in December.

A series of topotypes from Nias measure: wing cf 92-97 (94), 9
83.5, 85; culmen cf 15.5-17.5, 9 16,17.

224. Kittacincla malabarica opisthopela Oberholser
Synonym: Kittacincla malabarica opisthisa Oberholser

Tuangku, Bangkaru, Tello, Tana Massa, Tana Bala. Banyak
Island males are not paler on the posterior lower parts as Oberholser
claims (1912, p. 13), and if their tails are longer they are only longer
by a few millimeters which is too uncertain a character for such a small
series.

Considered genetically Banyak and Batu Island birds are probably
different just as the different island populations of melanura probably
are. However, they are phenotypically too similar to merit separation
from the taxonomic point of view. This race differs from melanura by
having white on the outer rectrices although it is less in amount than
in the case of tricolor. The females, like those of melanura are more
brightly colored, more masculine than are the females of tricolor. The
type of opisthopela is a not quite adult female collected on Tana Bala
February 5, 1903. It measures: wing 90, culmen 16.5. The type of
opisthisa is an adult male taken on Tuangku January 23, 1902. It
measures: wing 100, culmen 16.5. Birds from Tuangku and Bangkaru
measure: wing cf 97, 97, 100, culmen 16, 16, 17. Batu birds (Tana
Massa and Tana Bala) measure: wing cf 96, 102, culmen 18, 18.5.

The feet of various specimens were noted as: "pink, flesh, brown
madder".

!l have not included tail measurements as I consider them to be too variable.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 393

225. Geokiciila Sibirica (Pallas)
Nias. A migrant recorded in December by Biittikofer (189f>, p. 181).

226. Geokichla ixterpres leucolaema Salvador]

Enggano. A series of eleven adults present an interesting appear-
ance when compared with adults of i. inter pres. In color leucolaema
is dull and unfinished looking as if in slightly immature plumage. The
crown and nape are dull orange brown. The back is brown with a dull
golden orange tint. The throat and breast are black, but a wide streak
of white extends from the chin down onto the upper breast. The
flanks are strongly washed with buff. In size the two forms are similar
with the exception of the bill which in leucomelaena is considerably
larger.

Measurements: wing d* 99, 102, 103.5, 104, 105.5, 9 100.5, 101 (3)
101.5, o 101; oilmen & 18-20, 9 18-19.5.

An immature male taken in December is rusty brown on the upper
parts with paler shaft streaks on the back and scapulars. Below the
bird is white with a strong buffy-brownish wash. There are a few
black feathers on the breast.

Soft parts of adults : "iris dark brown, bill black; feet pale fleshy, dull
yellowish fleshy, straw yellow."

Dr. Abbott noted on one of the labels that this thrush was common
and not at all shy. It frequents dark jungle keeping near the ground.

227. Zoothera andromedae (Temminck)

Enggano. Salvadori (1892, p. 134) records three specimens from
the island, one of them an immature bird taken in June.

Family SYLVIIDAE, Old World Warblers

Three migrants and six resident species are found on these islands.
Three of the resident populations are endemic. Of these one race differs
in color from the Sumatra form, one race differs in color and slightly
larger measurements from Sumatra birds, and one race is closest to
Bornean birds from which it differs in color and size.

228. Locustella certhiola (Pallas)
Simalur. Junge (1936, p. 58) records two migrants taken on Simalur

in

Mav.

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229. Acrocephalus arundinaceus orientalis
(Temminck and Schlegel)

Simalur. A migrant recorded by Junge (1936, p. 58) as taken there
in February and March.

230. Orthotomus atrogularis artogularis Temminck

Tuangku (sight record), Tello, Tana Massa. The specimens ex-
amined by de Schauensee (1940, p. 39) seemed to be much the same
as a series from the Malay Peninsula. Abbott notes this form as com-
mon on Tuangku (Richmond: 1903, p. 511).

231. Orthotomus sepium ruficeps (Lesson)
Synonym: Orthotomus cineraceus baeus Oberholser

Synonym: Orthotomus cineraceus ochrommatus Oberholser

Nias and the Pagi Islands. The type of baeus, an adult male taken
March 19, 1903, measures : wing 48.5, tail 44, culmen 15.5. Four males
and a female measure: wing d71 48.5, 49 (3). 9 48; culmen cf 14, 14,
15 (2). Four males and a female from the Malay Peninsula, Banka and
Billiton measure: wing o71 48.5-50.5, 9 50.5;, culmen cf 14-15.5, 9
14.5. Using these measurements I cannot agree with Oberholser's
statement (1912, p. 13) that baeus is, "like Orthotomus cineraceus cine-
raceus but smaller".

The type of ochromatus is an adult male collected on North Pagi
November 23, 1902. It measures: wing 50.5, tail 43.5, culmen 16. I
cannot see any constant difference between the four adult males from
North and South Pagi and Nias and Malay Peninsula birds to sup-
port the name ochrommatus. Three other males have wings of 48.5,
49, 51, and culmens of 14.5-15.5, so that they are not "larger" than
baeus. The degree of paleness or darkness in these specimens is too
variable to form a strict criterion of subspeciation.

232. Orthotomus sepium concinnus Riley

Siberut, Sipora. The type is an adult male collected on Sipora
October 15, 1924, by C. B. Kloss. It measures : wing 50, tail 43, culmen
14. As Riley's description points out (1927, p. 96) this is a distinctly
paler bird than any of the other forms found in Sumatra or the western
islands. The soft parts of the type are marked as: "iris ochreous; bill,
upper mandible black, lower fleshy; feet fleshy brown".

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 395

233. Cisticola juncidis malaya Lynes

Simalur, Nias, Enggano. Apparently a common form wherever
suitable conditions for its presence exist. Found in meadows on Sima-
lur. Juvenal birds were collected in February (Simalur) and June
(Enggano). Eggs were taken in March on Simalur. For their measure-
ments see Junge (1936, p. 58). Specimens in the collection from
Simalur are molting in December.

234. Phylloscopus borealis borealis (Blasius)

Nias. A migrant taken on Nias in March. Two specimens in the
collection taken by Abbott during that month are molting.

235. Gerygone fusca muscicapa Oberholser

Enggano and Palu Dua. I cannot follow Meise (Novit. Zool., 36,
1931, p. 371) who synonymizes Enggano birds under sulphured. These
birds are not smaller as Oberholser mentions in his original descrip-
tion, (1912, p. 11) but are somewhat larger with larger bills, and the
three specimens before me are definitely brighter yellow on the under
parts than any specimens I have seen from other localities, in this
respect approaching flaieola from Celebes.

The type of muscicapa is a male collected on Pula Dua, a small
island off Enggano, November 2, 1904. It measures: wing 55, tail 35,
culmen 11.5. Two females measure: wing 50, 52.5; tail 34, 35; culmen
11, 11.5. Two males and a female from Sumatra and Banka measure:
wing c? 49.5, 53.5, 9 51; tail & 32, 34, 9 32.5; culmen cf 9, 9.5, 9 10.

236. Prinia flaviventris halistona (Oberholser)

Nias. As has been pointed out previously (de Schauensee and
Ripley, 1939, p. 409) in the case of this bird, it is not conspecific with
Burnesia dysancrita with which it was compared. The latter name is
a synonym of Prinia atrogularis albogularis (latest revision, Deignan,
Smiths. Misc. Coll., 103, no. 3, 1942).

The type of halistona is an adult male collected on Nias March 22,
1905 by Dr. Abbott. It measures: wing 48, tail 59, culmen 14. Another
male and two females measure: wing c?51.5, 9 46, 47; tail d* 62, 9
46, 51; cu'men <? 14, 9 13.5, 13.5.

This race is closest to chaseni of Borneo. Like that form it is much
paler on the under surface than rafflesi of Sumatra and the Malay
Peninsula. There is more than a hint of a buffy wash on the under

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surface, particularly on the breast and flanks. Specimens of rafflesi
tend to show some yellow on the flanks but in halistona this is virtually
lacking. On the upper surface halistona lacks much of the olive green
tint found in rafflesi. Nias birds are pure smoky gray on the head and
nape with a reduced amount of olive green on the back and scapulars.
Nias birds are somewhat larger than those from Borneo. A small
series of chaseni measure: wing c? 45, 46, 9 41, o 44.5, 47.5; culmen
cf 13, 9 14, o 12.5, 13. Specimens from south and east Borneo seem
to be somewhat more buffy less yellow on the flanks than north Borneo
birds. Two December birds, one from the Kapuas river, the other
from southeast Borneo, are quite different in this respect.

Family MUSCICAPIDAE, Old World Flycatchers

A total of twelve forms are recorded from the islands, two of which
are migrants. Of the ten other forms, six are endemic. Three of these
forms show closer affinity with birds from the Andaman and Nicobar
islands than they do to their relatives on Sumatra. The remaining
three forms differ from Sumatran races by color in two cases and by
size and color in one case.

237. Alseonax latirostris latirostris (Raffles)

Siberut. The single record for this migrant species on the west
Sumatra Islands is that of Chasen and Kloss (1926, p. 286).

Hypothymis azurea

This is a plastic species which has been split rather thoroughly in
the west Sumatra islands. The main tendency in the males is assume
a darker coloration. The white area on the underparts tends to be-
come much reduced, in one form the black crown patch and breast ring
have disappeared, and in general these forms differ from that of Su-
matra by larger size.

238. Hypothymis azurea consobrina Richmond

Simalur. In color this race seems to be indistinguishable from
tytleri of the Andamans. It does, however, appear to be slightly
smaller. The type is an adult male taken on Simalur December 24,
1901. It measures wing 69, tail 68, culmen 12. A series measures:
wing cf 69, 70, 72, 74.5, cf im. 70. Two males of tytleri measure: wing
74.5 (imperfect), 81.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 397

Compared to prophata from Sumatra, this race lacks the white
belly and vent. In consobrina this area is suffused with blue. The
bill also is somewhat larger.

239. Hypothymis azurea abbotti Richmond

Babi, Lasia. The type is an adult male taken on Babi, Januaryll,
1902 by Dr. Abbott. It measures: wing 78, tail 77, culmen 13. A
series measures: wing d" 78-81 (79.5) c? im. 76.5, 9 80; culmen
12.5-13, d" im. 13. 5 9 13.

This is an interesting and distinctive form which would be con-
sidered by many authors as a full species. Besides being larger, males
lack completely the black nape patch and breast band. Also there is a
complete lack of white on the stomach and vent. I feel, however, that
these birds only show to an extreme mutational tendencies exhibited
by the other distinctive races; consobrina and richmondi, also found on
the west Sumatran islands.

The female differs from the female of prophata by being larger, by
having a bright suffusion of blue on the upper parts, and by having
smokey brown belly and vent washed with blue. The throat is brighter
and more like the throat of the male than in prophata.

Soft parts: "iris dark brown, blackish brown; bill blue, tip black,
inside of gape yellow; feet dark leaden blue".

240. Hypothymis azurea prophata Oberholser

Synonym: Hypothymis azurea isocara Oberholser

Synonym: Hypothymis azurea amelis Oberholser

Synonym: Hypothymis azurea ponera Oberholser

Tuangku, Bangkaru, Nias, Pini, Tana Massa, Tana Bala. The three
island races were named by Oberholser (Proc. U. S. Nat. Mus., 39,
1911, pp. 588-615) on the basis of slight size variations and greater or
lesser amounts of blue wash on the abdomen. I submit that these
variations in color are individual. A table of measurements follows.

The type of isocara is an adult male from Bangkaru collected Jan-
uary 1902. It measures: wing 76, tail 71, culmen 12.5. Soft parts:
"feet dull leaden blue, bill blue, tip black, inside mouth yellowish
green."

The type of amelis is an adult male collected on Nias March 21,
1903. It measures : wing 71.5, tail 72, culmen 12. In this specimen the
blue wash extends well down onto the lower abdomen but it can be

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matched by Sumatran specimens. The type of ponera is an adult male
taken on Tana Massa February 17, 1903. It measures: wing 72, tail
69.5, culmen 11.5.

Measurements for the islands are as follows:

wing

tail

culmen

Bangkaru cf

72.5, 76

70, 71

12, 12.5

9

71.5

67

11

Nias d*

68-71.5

64-72

11.5-13

9

65,66

62,63

12, 13

Batu Ids. d1

72-73

69.5-73.5

11.5-12

241. Hypothymis azurea leucophila Oberholser
Synonym: Hypothymis azurea sipora Chasen and Kloss

Siberut, Sipora, Pagi Islands. This race differs from prophata by
having more white showing on the abdomen of the male. In this it
approaches stycmi of Thailand from which it is readily separable by
the more violaceous tone of the blue parts of the plumage. The female
differs from that of prophata by being brighter, more rufous on the
upper surface. Below there is a tinting of brown in the gray of the
chest not found so distinctly in females of prophata. These birds differ
in a similar way from the female of styani.

The type of leucophila is an adult male taken on North Pagi January
8, 1903. It measures: wing 71.5, tail 66, culmen 12. Other specimens
measure wing & 70-75 (72), 9 68, 73; tail 65-71 (68), 9 62, 64;
culmen d" 10.5-13.5, 9 10.5, 13.

242. Hypothymis azurea richmondi Oberholser

Enggano. The type is an adult male collected November 22, 1904.
It measures: wing 77.5, tail 71, culmen 13.5. In color this form is
closest to tytleri and consobrina, differing from them in the more vio-
laceous tint to the plumage of the male. Apparently the female also
differs in having the upper parts more bright and rufous rather the
same way that the female of leucophila differs from prophata. The
crown and throat are brighter blue also than is the case with most
females of this species.

Soft parts: "iris dark brown, eyelids pale blue, bill blue tip black,
feet dull blue".

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 399

243. Terpsiphone paradisi procera (Richmond)

Simalur. The type is an adult male collected December 12, 1901.
It measures: wing 87.5, oilmen 18. All the males in the* series are in
the white plumage. In color they differ from males of affi/nis as pointed
out by Junge (1936, p. 49) by having the back and scapulars more pure
white. From males of nicobarica which is very slightly larger, these
birds are indistinguishable in color. The females however, are some-
what darker, more rufous brown on the upper surface than are females
of nicobarica. They are less rufous and more dull brown than females
of affinis.

Soft parts: "iris dark brown; eyelids blue; bill blue, tip black; gape
green; feet leaden blue."

An immature male was collected in November.

244. Terpsiphoxe paradisi insularis Salvadori

Xias. Two males from Nias are bright rufous above with the re-
duced metallic crown characteristic of this form. One specimen has
long tail feathers. Their wings measure: 91, 94. \Yeight, 25.5 gr.
Soft parts: "bill and ocular area vivid prussian blue, gape yellowish
green, feet milky blue." A male with short tail feathers was in breeding
condition in June.

245. Terpsiphone paradisi incei (Gould)
Xias. A migrant recorded by Biittikofer during the winter.

246. Drymophila pyrhoptera pyrhoptera (Temminck)

Pini, Tana Massa. Inseparable from Sumatra and Borneo birds.
Four males and four females measure: wing <f 82.5-84, 9 76-80.5.
Soft parts: "iris crimson, bill black, feet gray brown".

247. Rhinomyias umbratilis umbratilis (Strickland)
Synonym: Rhinomyias umbratilis eclipis Oberholser

Tana Massa. The type of eclipis is an adult male collected in Feb-
ruary 1903 on Tana Massa. It is an unique specimen and does not
seem to be separable from typical umbratilis. It measures: wing 78,
tail 63, culmen 15. The type of richmondi from Mansalar Island in
Sibolga bay, west Sumatra measures: wing 82.5, tail 65, culmen 15.

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A male from Borneo measures: wing 78, tail 64, culmen 15. I fail to
see how the single specimen of eclipis can be considered "decidedly
smaller" (1912, p. 12).

248. Culicicapa ceylonensis ceylonensis (Swainson)

Synonym: Culicicapa ceylonensis percnocara Oberholser
Synonym: Culicicapa ceylonensis pellonota Oberholser
Synonym: Culicicapa ceylonensis amphiala Oberholser

Simalur, Nias, Siberut, North Pagi. I agree with Junge (1936, p.
48) that the proposed race percnocara is untenable. The type, an adult
male was collected on Simalur November 23, 1901. It measures: wing
66, tail 51, culmen (damaged) 12. Another male measures: wing 62.5,
tail 50, culmen (damaged) 12. The type of pellonota is an adult male
from Nias collected February 20, 1905. It measures: wing 64, tail 51,
culmen 11. A female from Nias measures: wing 58, tail 48, culmen
10.5.

The type of amphila is an adult male from North Pagi taken January
8, 1903. It measures: wing 64; tail 50, culmen 10. Males and females
from North Pagi and Siberut measure: wing cf 59, 60, 63.5, 9 59.5;
culmen c? 10, 10.5, 11, 9 10.

These measurements can be equalled by a typical series from nearly
every part of the range of this bird. The color differences mentioned
in the original description (1912, p. 12) do not appear to me to be
anything more than individual variations.

Family MOTACILLIDAE, Wagtails

Four forms have been recorded, three of which are migrants. The
fourth is inseparable from Malayan and Sumatran specimens.

249. MOTACILLA CINEREA MELANOPE Pallas

A migrant recorded in the winter from Simalur, Nias, Tana Massa
and Sipora. A male was taken on Simalur in December. "Common on
marshy ground" (Junge, 1936, p. 68.).

250. Motacilla flava simillima Hartert

Another migrant taken on Simalur, Nias, Siberut, Sipora, and
Enggano from November 10 to March 19. There are specimens in the
National Museum's collection from Simalur, Nias and Enggano.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 401

251. Dendbonanthus indicts (Gmelin)

Specimens of this migrant were taken by the Abbott expedition on
Simalur, Nias, and Tana Bala. It was seen on Bangkaru but not col-
lected. It has also been recorded from Tello, Tana Massa, Siberut and
Sipora. The earliest date is October 2, the latest March 25. Junge
(1936, p. 69) records this bird as being seen in open country, once or
twice in trees.

252. Anthus richardi malayensis Eyton

Nias. A male and a female taken in March seem to belong to this
race. The male is slightly pale below, but the degree of brownish on
the upper parts is similar to males and females from Sumatra, Malacca,
Selangor and Ceylon. The specimens measure: wing cf 84, 9 81;
culmen cf 15.5, 9 15. Soft parts: "tarsi brownish, feet pale yellowish
fleshy".

Family LANIIDAE, Shrikes

Two species are found on these islands. One is a migrant, the other
is inseparable from Sumatran specimens.

253. Lanius tigrinus Drapiez
Simalur, Nias, Tello, Tana Massa, Tana Bala, Siberut, and Sipora.
A migrant taken from November 25 through March 7.

254. Pachycephala cinerea vandepolli Finsch

Synonym: Muscitrea grisola nesiotis Oberholser

Synonym: Pachycephala cinerea butaloides Stresemann

Simalur, Nias, Tello, Siberut, South Pagi. Nine specimens from
these islands seem to be all that are available for examination. Com-
paring this series with birds in comparative states of plumage from
Java, Sumatra and the Malay Peninsula I can see no tenable reasons
for recognizing nesiotis and butaloides.

The type of nesiotis is a male taken on Simalur, October 24, 1902.
It measures: wing 86, tail 66, culmen 15.5. Above, this specimen and
a female are slightly more rufescent particularly on the outer webs of
the secondaries and tertials than any specimens except those in very
fresh plumage, not always easyto find in collections. Since Junge (1936,
p. 59) cannot see any difference between his series and Sumatra birds
I am inclined to think that these specimens are perhaps not fully adult.

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The type of vandepolli is an adult male (A.N.S.P. no 56630) col-
lected August 31, 1896 by Kannegieter on Tello. It measures: wing
87, tail 65, culmen 15. The specimen is in rather worn plumage molt-
ing the crown feathers. It is inseparable from worn plumage specimens
from Sumatra1.

Measurements for the islands follow: wing d" 87, tail 65, culmen cf
15-16, 9 15.

A female on Nias was in breeding condition in June. It weighed
21.5 gr. while a male taken at the same time weighed 19.2 gr.

Family STURNIDAE, Starlings

Seven members of this family occur on the west Sumatra islands of
which one is a migrant. Among the remaining six, five are endemic
races. All five differ from Sumatra birds primarily in larger size,
although one form, Aplonis panayen&is enganensis, has a unique im-
mature plumage.

255. Sturnia sturnina (Pallas)

Simalur, Sipora. The Daurian Starling is a common winter migrant
although there are only two records for the west Sumatra Islands
(October and December).

Gracula religiosa

This species is a relatively plastic one which has had a distinct
speciation response towards larger size in the case of small island
populations. The metallic jet plumage of these birds has allowed for
no interpretation of these populations on the basis of color. There is
but one minor morphological character (the uniting of the neck lap-
pets) and this is only a tendency.

Instead, however, of the forms found on the most isolated islands
(Simalur, Enggano) being the most distinct, it is paradoxical that they
are the ones closest in size to the mainland birds. On the contrary the
birds from the Banyak Is., Nias, the Batu Is., and the Mentawi-Pagi
group, the nearest geographically to Sumatra, have acquired the dis-
tinct characters of larger size. Comparison of these latter specimens
with birds from the South China Sea reveals that there are no tenable
characters by which they can be separated, although it would seem
obvious that genetically they are not closely related. Here again tax-
onomy has to sacrifice truth to convenience.

^his form was unfortunately overlooked in de Schauensee's "Birds of the Batu Islands"
(1940).

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 403

256. Gracula religiosa religiosa Linnaeus

Synonym: Gracula javanensis miotera Oberholser

Synonym: Gracula enganensis Salvadori
Synonym: Gracula javensis baweana Oberholser

Simalur, Enggano, Pulu Dua, and Malay Peninsula (north into
Peninsular Siam), Sumatra, Rhio Archipelago, Banka, Billiton,
Borneo, Natuna Islands, Karimata Islands, Java, Bawean Island,
Bali, Kangean Islands, Christmas Island (introduced).

Oberholser (1912, p. 16) described miotera as being smaller with a
more slender bill than javensis (= religiosa). The type of miotera is an
adult male taken on Simalur November 24, 1901. It measures: wing
183, tail 86, culmen 29.5. Another male and four females measure:
wing & 175, 9 170-178 (176), culmen c? 28.5, 9 29, 29.5, 30 (2).
These measurements are within the range of religiosa.

Salvadori (1892, p. 137) described enganensis as being smaller with
differently arranged wattles. This again depends on the method of
making up the skins. Seven specimens measure : wing cf 182, 9 173-
175.5 (174.3), tail & 91, 9 89.5-93; culmen cf 29.5, 9 27-31.5.

The type of baweana is a female from Bawean Island in the Java
Sea, collected November 23, 1907. It measures: wing 175, tail 82,
culmen 26.5. A male measures: wring 179, tail 78, culmen 30. The size
of the lappets mentioned in the description (Proc. U. S. Nat. Mus. 52,
1917, p. 195) is well within the range of other carefully prepared
specimens.

Eighteen other specimens of religiosa from Java, Borneo, Sumatra,
etc. measure: wing d1 177-186.5 (181.5), 9 174-185 (180.2); tail 70-
86 (79), 9 73-87 (81.5); culmen c? 28-32.5, 9 26.5-32.5.

Soft parts: "iris dark brown; bill red, tip yellow; wattles yellow,
bright yellow; feet yellow".

These birds are common to abundant in original forest. Malay
name, "beo".

257. Gracula religiosa robusta Salvadori
Synonym: Gracula javanensis ophellochlora Oberholser

Babi, Tuangku, Bangkaru (seen), Nias. The type of ophellochlora
an adult male collected on Tuangku, January 23, 1902. It measures:
wing 201, tail 88.5, culmen 34. Other birds from Tuangku measure:
wing cT 202-204 (203), 9 192; tail & 93-96 (95), 9 93; culmen &
33-34, 9 33. Birds from Babi and Nias measure: wing c? 201-210

404 bulletin: museum of comparative zoology

(205.4), 9 198-201.5; tail c?93-100 (96.5), 9 92-94; culmen d" 34-38
(36.4), 9 34, 36.

Oberholser's race (1912, p. 17) was named as being smaller with
more greenish sides to the head. It is not significantly smaller as the
measurements show nor are the sides of the heads of these birds more
greenish.

This mynah, the largest race of the species, was found in breeding
condition on Nias in June. A male weighed 425 gr., a female 420. Soft
parts: "iris dark brown; bill red, orange red at base, yellow at tip;
feet and wattles yellow". This is a bird of the high land away from the
sea, found usually in small flocks in the jungle.

258. Gracula religiosa batuensis Finsch
Synonym: Gracula javana prasiocara Oberholser

Tello, Siberut, Sipora, Pagi Islands, and Tioman Island, Anamba
and Tambelan Islands. I have examined thirty-three specimens from
all these localities and I can find no constant differences to separate
them. Finsch (Notes Leyden Mus., 21, 1899, p. 14) named this race
on the basis of greater amount of white on the wing, (a variable char-
acter) and certain differences in the size and structure of the neck
wattles. The Batu Island birds, four from Tello collected by Kan-
negieter, have all been skinned in a careful way to show the wattles.
They have been spread out over the back of the neck and dried so
that they show up exceptionally well. One of the females has the two
wattles joined together well out beyond the point covered by the crest
feathers. In two other specimens this is not the case. The crest
feathers cover the point where the two wattles join. Out of five
specimens from Siberut, only one has the two wattles joined together
beyond the crest feathers. No specimen from Sipora or the Pagi
Islands have the wattles joined. The Pagi specimens have been pre-
pared for the most part without reference to the preservation of the
neck wattles which, as a result are so shrunken that this very condition
might serve as a taxonomic character were it not known to be due to
chance.

Two females from Bunoa, Tambelan Islands have the two wattles
joined together beyond the crest feathers.

I have seen one female of religiosa from the northern Malay Penin-
sula with joined wattles.

Altogether the wattle structure seems to be an uncertain and
dubious character. I feel that on the basis of this evidence it should
not be used as a criterion of the race batuensis. It is noteworthy, how-

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 405

ever, that eleven out of seventeen specimens of robusta have the
wattles prominently connected out beyond the crest feathers.

The only real character that I can determine for this race is one of
intermediate size between robusta and religiosa. It is for this reason
that I have included prasiocara under batuensis. The measurements
are so largely overlapping that there would be no chance of deter-
mining whether unlabeled specimens had come from the west Sumatra
Islands or the south China Sea.

The type of prasiocara is an adult male from Piling in the 6\namba
group, collected August 17, 1899. It measures: wing 196, tail 89.5,
culmen 30. A small series measures: wing cf 189.5-196 (192.9), 9
174-192 (184); tail <? 85-89.5 (87.7) 9 82-89 (85.3); culmen a*
30-32, 9 29-32.

Birds from Tello, Siberut, Sipora, and the Pagi Islands measure:
wing & 173-192.5 (186), 9 170-193.5 (182.3); tail cf 78.5-87 (82.2),
9 75-87 (82.1); culmen cf 31-34 (32.2) 9 30-34 (32.1).

Soft parts of these specimens: "iris dark brown, brownish gray;
bill orange red, tip black; feet yellow" (immature specimens were
collected in August and November).

259. Aplonis panayensis altirostris (Salvadori)
Synonym: Lamprocorax chalybeus rhadinorhamphus Oberholser
Synonym: Lamprocorax panayensis nesodramus Oberholser

Simalur, Babi, Nias. I agree with Riley (1929, p. 33) that these
birds should be combined under one name. The type of rhadinor-
hamphus is an adult male collected on Simalur December 12, 1901. It
measures: wing 103.5, tail 62.5, culmen 18. The type of nesodramus
is an adult male taken January 13, 1902, on Babi. It measures; wing
106, tail 64.5, culmen 19. Three females from Simalur and Babi
measure: wing 102.5, 103 (2); tail 61.5, 62.5, 63; culmen 17, 17.5, 18.5.

A series from Nias measures : wing cf 100.5-105.5, 9 97-104(101.5),
tail tf 61 (3), 9 57-63 (60.2) ; culmen d* 19, 19.5 (2), 9 17-18.

Two immature birds were taken in February and March. The iris
of these birds is noted as: "bright crimson, red," of the immatures:
"orange red, brick red."

260. Aplonis panayensis pachistorhinus (Oberholser)
Synonym: Aplonis panayensis leptorhynchus Stresemann

Pini, Tello, Tana Massa, Tana Bala, Siberut, Sipora, South Pagi.
As de Schauensee has pointed out (1940, p. 41) there is no reason to

wing

tail

culmen

110-112

66-69

19-20

107 (2)

62,63

19,20

107.5, 112

67,68

20,21

104, 106

63.5 (2)

19-21

105-109

64-67

19-20

100-104

60-64

18.5, 19 (2)

406 bulletin: museum of compaeative zoology

recognize Stresemann's race as the measurements of the two series
overlap.

The type of pachistorhinus is an adult male taken on South Pagi
November 19, 1902. It measures: wing 109, tail 68, culmen 19.5.

Adults from the different islands measure as follows :

Batu Ids. <?

9
Siberut and Sipora c?
9
South Pagi c?

9

Immature birds were taken from September through December and
in February (Tana Bala).

261. Aplonis panayensis enganensis (Salvadori)

Enggano. Seven adults measure wing d1 112.5-116, 9 HO, 112,
116; tail c? 71-75, 9 67, 67, 75; culmen c? 18.5-19.5, 9 18.5-19.
The most distinctive character of this form is not so much the fact
that it is a large bird, the largest of these races, but rather the unique
immature plumage. Seven immature birds taken in November and
December are similar to immature examples of the rest of the species
on the upper parts, but the lower parts are very dark brown almost
black with only pale edges to the feathers of the throat and abdomen.
This last is somewhat variable, but in any case all the specimens
present a far more uniform dark and sooty appearance on the under
parts than any other form.

Another interesting character of enganensis is the bill which is
smaller than the other western island forms, being virtually identical
with that of strigatus.

Soft parts: "iris red, deep' red"; of immatures, "gray brown, green-
ish white" (?).

Family NECTARINIIDAE, Sunbirds

Many subspecies have been named from the west Sumatra islands
but I can recognize only ten forms as occurring there. Of these, two
are certainly, one possibly, endemic. One of these forms is a straight size
race, one is part larger size, part color, and the third is possibly ten-
able based on color differences. All are closely related to Sumatran
forms.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 407

262. Chalcostetha calcostetha calcostetha (Jardine)

Synonym: Chalcostetha calcostetha heliomarpta Oberholser

Synonym: Chalcostetha calcostetha siberu Chasen and Kloss

Synonym: Chalcostetha calcostetha pagicola Oberholser

Simalur, Nias, Pini, Tana Massa, Siberut, Sipora, Pagi Islands. A
total of three races have been named on the basis of small color and
size differences. I have been unable to see the color differences men-
tioned. Females all through the islands are identical with Sumatra and
Malay Peninsula specimens.

On the question of size I should say that these birds agreed with
specimens from Sumatra in being large, but within the range of
measurements for this form as a whole. Of all the females measured,
the type of heliomarpta is the largest, but so close to the other speci-
mens, that I feel sure that more females from Simalur would invalidate
this single large bird. In fact Junge's discussion (1936, p. 69) indicates
that this is the case.

The type of heliomarpta is an adult female collected December 1,
1901. It measures: wing 61.5, tail 44.5, culmen 19.5. Two males, also
from Simalur, measure: wing 62 (2), tail 52, 54; culmen 19, 19.5.

The type of pagicola is an adult male collected on North Pagi,
January 2, 1903. It measures: wing 62, tail 52, culmen 19.5.

Birds from Sumatra, Malay Peninsula, and Banka, from which local-
ity this form is not recorded in Chasen (1935, p. 273) measure: wing
c? 60-65, 9 54-60; tail d" 50-54; 9 40-43.5; culmen c? 18-19,
9 17-19.5.

Junge (1936, p. 70) writes that this form is abundant in coastal
mangroves. Nesting apparently goes on throughout the year.

263. Aethopyga siparaja siparaja (Raffles)

Synonym: Aethopyga siparaja tinoptila Oberholser

Synonym: Aethopyga siparaja melanetra Oberholser

Synonym: Aethopyga siparaja heliophiletica Oberholser

Synonym: Aethopyga siparaja niasensis Hartert

Synonym: Aethopyga siparaja photina Oberholser

Synonym: Aethopyga siparaja siberu Chasen and Kloss

Simalur, Siumat, Lasia, Babi, Bangkaru, Nias, Siberut, Sipora, the
Pagi Islands. So many races have been described of this species that
I feel somewhat at a loss as to where to begin in analyzing the various
specimens. A series of twenty-two skins from the above localities

408 bulletin: museum of comparative zoology

convinces me, however, that it is useless to attempt to separate races
from these islands. To begin with, large series are necessary for proper
identification of valid characters. Some of these forms (viz niasensis)
have been described on the basis of a single specimen.

The type of tinoptila is an adult male collected on Siumat off Simalur
in December, 1901. It measures: wing 52.5; tail 42, culmen 15. In
color it is indistinguishable from Sumatran birds.

The type of mclanetra from Lasia is an adult male taken January 5,
1902. It measures: wing 51, tail 40, culmen 14.5. The type of helio-
philetica is an adult male from Bangkaru taken January 18, 1902. It
measures : wing 51.5, tail 43, culmen 15.5. Both specimens are insepar-
able from typical siparaja in size and color.

Of seven males from Nias, three are indistinguishable from typical
siparaja. Of the remaining four specimens, two collected in February
and March are not fully adult, therefore have a slight greenish tinge
to the red of the back, and two, taken in March are in very worn
plumage making them appear slightly pale on the upper surface.

The type of photina is an adult male taken on North Pagi December
22, 1902. It measures: wing 51.5, tail 41, culmen 15. Specimens of
both this form and so-called siberu have the rump irregularly washed
with orange red, but contrary to Chasen and Kloss' statement (1926,
p. 298) this occurs occasionally in Sumatran birds at least.

This is a bird of cleared gardens, found in blossoming trees.

264. ClNNYRIS BRASILIANA MECYNORHYNCHA Oberholser

Simalur. The type of mecynorhyncha is an adult male taken on
Simalur November 19, 1901. It measures: wing 51.5, tail 30, culmen
16. Another male has a wing of 51 and a culmen measuring 16. These
measurements are larger for the culmen than any others I have meas-
ured although I hardly think that Oberholser's description (1912, p.
19) "bill very much larger", is pertinent here. A male from Nias, the
type of oenopa with which presumably the Simalur birds were com-
pared, measures: culmen 15.5!

265. ClNNYRIS BRASILIANA BRASILIANA (Gmelin)

Synonym: Cinnyris brasiliana oenopa Oberholser
Synonym: Cinnyris brasiliana hypolampis Oberholser

Nias, Tana Massa, Siberut, Sipora, and the Pagi Islands. The type
of oenopa is an adult male taken on Nias, March 14, 1905. It measures :

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 409

wing 48, tail 27.5, culmen 15.5. The type of hypolampis is an adult
male taken December 11, 1902 on South Pagi. It measures: wing 50,
tail 29, culmen 14.5. Males from Nias have culmens measuring
14.5-15.5. Males from Tana Massa down to South Pagi have culmen
measurements ranging from 14-15. Six males from Java and the
Malay Peninsula have eulmen measurements ranging from 13-15. At
this rate it seems impossible to separate these birds although it may
be said that west Sumatra island birds do have a tendency towards
slightly larger size.

Breeding birds were taken on Nias in June. They weighed : cf 6, 9
gr. 9 6 gr. Soft parts: "iris brown, feet dark brown".

266. Cinnyris jugularis polyclysta Oberholser

Enggano. This is a darker, more olive bird that pectoralis. The
culmen measurements also are larger. The type of polyclysta is an
adult male taken on Enggano November 24, 1904. It measures:
wing 56.5, tail 33, culmen 20. Two other adult males measure: whig
55.5, 56; culmen 19, 20.

267. Anthreptes simplex simplex (S. Miiller)

Nias. This species has been taken by Rosenberg, Modigliani, Claine,
and Thomas.

268. Anthreptes malacensis malacensis (Scopoli)
Synonym: Axthreptes malacensis pelloptilus Oberholser
Synonym: Axthreptes malacensis pollostus Oberholser
Synonym: Anthreptes malacensis nesaeus Oberholser

Simalur, Nias, Tello, Tana Massa, Siberut, Sipora, and the Pagi
Islands. In series from any one of these islands the measurements of
the sunbird differ. However the difference is very small, and if birds
of similar months are compared there is seen to be no difference in
color.

These measurements indicate that there is a trend towards larger
size on the west Sumatran Islands, but only the Batu series from Tello
and Tana Massa are consistently larger. However, the difference in
size is far too small to merit separation.

410 bulletin: museum of comparative zoology

Weight (Nias); d" 11-14 gr. c? im. 11 gr. Soft parts: iris brown.

wing

tail

culmen

Simalur d" (type of pelloptilus 68.5

44

19 Nov. 22, 1901

&

68.5, 69.5

44,47

18, 18.5

9

66

42

17

Nias d* (type of pollostus)

66.5

42.5

17.5 Feb. 27, 1905

&

64-68.5

41-44

16.5-17.5

9

60, 61.5

36.5, 41

15.5, 16.5

Batu Ids. c?

70.5-71

44-48

18-19

9

68-70

45

17.5-19

Siberut, Sipora &

66-68.5

42-45

17.5-19

9

64

41

16.5

Pagi Ids. d1 (type of nesaeus)

69.5

44

18.5 Nov.-Dec. 1902

&

69.5 (2)

45(2)

17.5, 18

9

65

42

17

Java, Sumatra cf

65-68.5

42-46

17-19

9

58,60

37,40

16, 17

As so often is the case on the west Sumatra islands, if there is a
tendency towards larger size in a species, the larger birds come from
Simalur, the Batu Islands or the Pagi Islands. Those populations
nearest the Sumatra-Java form come from Nias, Simalur and Siberut.

269. Chalcoparia singalensis panopsia Oberholser

Tuangku, Nias, Tana Massa. When Kloss described the race
sumatrana (Journ. Fed. Malay States Mus., 10, 1921, p. 209) he had no
specimens of the west Sumatra island population to compare his
Sumatran specimens with. Compared with a single old, faded Suma-
tran female in the collection of the Academy of. Natural Sciences,
I see no difference that is tenable between the Sumatra specimen and
the island specimens. One Nias female is brighter but other less ma-
ture specimens are less bright. More Sumatran specimens may well
show that sumairana is a synonym of panopsia.

The type of panopsia is an adult female taken on Tuangku January

25, 1902. It measures: wing 53.5, tail 37.5, culmen 13.5. Males and

females from Nias and Tana Massa measure: wing c? 51, 53, cf im.

50.5, 55, 9 48, 54.5, 9 im. 51; tail cf 39, 42, 9 37; culmen <?<? and

9 9 13-13.5.

Soft parts: "iris brown; feet gray, yellowish gray, greenish yellow".
Weight: cf 7.2, 9 9.6 gr. Immature specimens were taken (marked
cf) is in full female plumage.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS

411

270. Arachnothera longirostra longirostra (Latham)

Synonym: Arachnothera longirostra zarhina Oberholser

Synonym: Arachnothera longirostra niasensis van Oort

Synonym: Arachnothera longirostra hypochra Oberholser

Synonym: Arachnothera longirostra exochra Oberholser

Tuangku, Bangkaru, Nias, Tana Massa, Siberut, Sipora, and the
Pagi Islands.

A series from these islands reveals that this is a very plastic species
as far as length of bill is concerned. There is good deal of variation in
color among these birds depending on season and molt. Two specimens
from Nias are pale on the under parts but both were taken in March.
They agree fairly well with my specimens from Singapore.

The measurements of these specimens are as follows :

wing

culmen

Tuangku, Bangkaru

cf (type of zarhina)

68.5

43.5 Jan. 18, 1902

&

70

broken

9

62.5

38

Nias d1

69

broken

9

68.5

41

Tana Massa 9

60.5, 64

37,38

Siberut, Sipora <?

66.5-70

34-40

9

60-62.5

34

North Pagi o71 (type

of hypochra)

67.5

38 Nov. 24, 1902

&

68, 70

36.5

9

60

33

South Pagi d" (type of exochra)

69

36 Nov. 15, 1902

&

69-71

36-39

9

61-65

33-34

Soft parts: "iris brown, bill black, feet bluish-gray". Weight
(Nias), 14 gr.

271. Arachnothera chrysogenys chrysogenys (Temminck)

Synonym: Arachnothera chrysogenys pleoxantha Oberholser

Synonym: Arachnothera chrysogenys isopega Oberholser

Nias, Sipora, and the Pagi Islands. The type of pleoxantha is an
adult female taken on Nias, February 27, 1905. It measures: wing 78,
tail 36, culmen 38. A male measures: wing 83 (worn), culmen 35.5.
The type of isopega is an adult, six undetermined taken on the Pagi
Islands, December 26, 1902. It measures : wing 93, tail 40, culmen 35.5.
A female from South Pagi measures: wing 81.5, culmen 36.5.

412 bulletin: museum of comparative zoology

These specimens compare favorably with a series of birds from
Sumatra including the type of copha. Soft parts : "bill dark horn brown,
yellow along middle of culmen; feet brownish fleshy."

Family DICAEIDAE, Flower-peckers

Five forms are recorded from the west Sumatra islands four of
which are endemic. All four races differ in color from their relatives
on Sumatra, but three of the races show a tendency towards larger
size particularly in the bill measurements.

272. Dicaeum cruentatum niasense de Schauensee and Ripley

Nias. A single male measures: wing 48, culmen 11. This specimen
agrees well with the original description (1939, p. 410.). Junge's
record (1936, p. 74.) for Simalur may refer to this race.

273. Dicaeum cruentatum batuense Richmond

Pini, Tello, Sipora, South Pagi. The type is an adult male taken
March 3, 1903 on Pini. It measures: wing 48, tail 25.5, culmen 10.25.
It agrees well with Richmond's original description (1912, p. 104).
Unfortunately I have had no Sumatra specimens for comparison.
Two males from Sipora have wing measurements of 45.5. A female
from Pini measures: wing 44.5.

274. Dicaeum trigonostigmum antioproctum Oberholser

Synonym: Dicaeum trigonostigma melanthe Oberholser

Synonym,: Dicaeum trigonostigma lyprum Oberholser

Synonym: Dicaeum trigonostigmum tanamassae de Schauensee

and Ripley
Synonym: Dicaeum trigonostigmum pagense Oberholser

Simalur, Lasia, Nias, Pini, Tana Massa, Siberut, Sipora, South Pagi.
This form shows a tendency towards slightly larger size than t. trigo-
nostigmum from the Malay Peninsula and Sumatra, but the color
differences indicated in the various descriptions tend to disappear in a
series of male specimens. Female specimens from all these islands
agree, however, in differing with females of the nominate race by being
brighter on the rump and definitely brighter, more orange yellow on
the abdomen.

»

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS

413

Measurements are as follows :

wing tail

Simalur cf (type of antioproctum) 49.5 22.5

d* 54 21

9 49 22

Lasia c? (type of melanthe) 53 24.5

Nias d1 (type of I y pram) 49 20

cf 49, 50 21.5, 23

9 48.5, 50.5 20, 22

Pini c? 49.5 23.5

Siberut, Sipora & 49.5-50.5 22-23

9 47, 50 22.5, 23

South Pagi & (type of pagense) 49.5 23

cf 50 24

9 48.5 22

Sumatra, Malay Peninsula cf 45-50.5 20-23

9 46.5 20.5

culmen

11.5 Nov. 25, 1901

11.5

11

11.5 Jan. 7, 1902

11 March 21, 1903

11 (2)

10(2)

10.5

10-11.5

11 (2)

11

11

10.5

10-11

11

Immature birds were collected from September through December.
Males in breeding condition on Nias in June. Weight c? 7, 8.5 gr.,
9 7 gr. Feet dull slate color.

275. Anaimos percussus regulus de Schauensee

Tana Massa. The type and two females taken by Kannegieter in
1896, are in the collection of the Academy of Natural Sciences. They
measure: wing c? 56, 9 51.5, 52; tail cf 23, 9 24, 24.5. These birds
differ from ignicapillus by being duller and paler and more washed out
in their entire coloration. In size they are the same.

276. Anaimos maculatus maculatus (Temminck)
Synonym: Anaimos maculatus opistatus Oberholser

Nias. The type of opistatus is an adult male taken on Nias March 3,
1905. It measures: wing 51.5, tail 22.5, culmen 10.5. Another male
and a female measure: wing c? 51, 9 51, culmen cf 10, 9 10. Although
the type specimen is very slightly darker, the other male is completely
inseparable in color from a male from the Sink river, east Sumatra.
There is no difference in wing size as the Sumatra bird measures 51,
but the culmen is a trace shorter measuring 9 mm.

A female weighed 6 gr. and had the following colors of the soft
parts: "iris brownish red; base of lower mandible gray, remainder
black; feet grayish."

414 bulletin: museum of comparative zoology

Family ZOSTEROPIDAE, White-eyes

A single subspecies from Enggano is the only member of this family
on these islands. It is much larger than the Sumatran species and is
referred tentatively to the large mountain species of the Malay
Peninsula.

277. Zosterops aureiventer salvadorii Meyer and Wiglesworth

Enggano, Pulu Dua. In coloration this form is almost exactly
intermediate between Z. palpcbrosa sumatrana and Z. aureiventer
aureiventer from the Malay Peninsula. However, in size it is much
nearer aureiventer than palpebrosa, and so I am inclined to agree with
Junge (1938, p. 353) that it should be put into that species.

Five males and two females measure: wing d1 58-60, 9 56.5, 59;
tail & 38-39.5, 9 36, 37; culmen rf1 13-14, 9 12.5, 13. Soft parts:
"iris reddish brown, feet leaden".

A single male from the island of Banka has a wing of 54 mm. In
size, therefore, it must be assigned temporarily to aureiventer aurei-
venter from the Malay Peninsula although the under tail coverts are
rather more orange-yellow than in that form. This is a new record for
Banka and it may well be that a larger series will show that there is a
distinct form on this island.

Family PLOCEIDAE, Weaver-finches

Three common forms occur on the islands, all of which are pre-
sumably identical with the populations found on Sumatra.

278. Lonchura maja maja (Linnaeus)
Synonym: Munia maja simalurensis Oberholser

Simalur, Nias, Pini. The type of simalurensis is an adult male col-
lected by Dr. Abbot on Simalur, November 22, 1901. It measures:
wing 54.5, tail 31.5, culmen 13. This race was based on the males
having more white on the anterior lower parts, but this is a completely
variable character appearing irregularly in birds from the Malay
Peninsula to Java.

Another male and a female from Simalur measure: wing cf 54.5, 9
53.5; culmen d71 13, 9 12. Males and females from Nias measure:

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 415

wing of 52.5-54.5, 9 53-55 ; culmen & 11.5-12, 9 12-13.5. A female
from Pini measures: wing 54, culmen 12.

An immature bird, pale brown above, grayish buffy white below,
was collected in February on Nias. A female molting into adult plum-
age was taken on the same island in March. Soft parts, bill pale blue
gray; feet dull slaty, dark leaden blue. Weight c/1 (testes enlarged
in June) 11 gr. Eggs were taken on Simalur in June. They are de-
scribed in Junge (1936, p. 68).

This is a common form in the rice fields and in open gardens.

279. LONCHURA PUNCTULATA FRETENSIS KloSS

Nias. Recorded from this island by Salvadori (1886, p. 552) and
Oustalet (1892, p. 115).

280. Ploceus philippinus infortunatus Hartert

Nias. Salvadori, Blasius (1901, p. 71) and Oustalet have all listed
this species as occurring on Nias.

SUMMARY

A study of the faunal list of the birds of the west Sumatra islands
reveals very clearly that factors are present which have resulted in the
presence or absence of certain birds and the speciation of some of these
bird forms. These factors are difficult to analyze and evaluate. It
may be said at once, however, that there are virtually no important
ecological barriers except those noted below. So far as is known at
present, all the islands from Simalur to Enggano have the same gross
conditions of forest, swamp, and open country. Presumably food
conditions are identical. As far as climatology is concerned there is
no evidence to indicate that conditions are relatively different on any
of the islands. Thus we are reduced in this discussion to two factors
which come under the heading of isolating mechanisms. One of these
is the area of the different islands in square miles. The other is the
relative degree of isolation of the different islands one from the other,
and from Sumatra.

The area of islands within certain gross limits is obviously an im-
portant factor as far as the presence of a bird fauna is concerned.
Very small islands in this region are usually of coral, support only
limited types of vegetation, and in consequence are populated by

416 bulletin: museum of comparative zoology

birds of a reduced number of species and often of a sporadically wan-
dering type (viz., Ducula bicolor, Halcyon chloris, etc.). Larger islands
obviously tend to have different types of soil (volcanic upthrusts,
limestone outeroppings, schists), support a much larger variety of
plant and insect life, and consequently provide for a much greater
bird fauna. With increasing area comes an increasing variety of
habitats. There is, also, an increasing opportunity both through the
isolation of the habitats themselves as well as the factor of the size of
the population, which recent genetic studies of Drosophila in California
(Dobzhansky et al.) tend to show is highly important in speciation,
for small populations to be built up of the new bird arrivals.

Isolation of the islands as a factor is inescapable but it is difficult
to determine just how important it is. After all, isolation of too small
a population must certainly retard speciation just as surely as con-
stant swamping of a population must retard it. But it remains for
the following paragraphs to indicate how great or how small these
two mechanisms are in effect.

The area of these islands may be expressed roughly as follows :

Simalur (including Lasia and Babi)

500 sq.

mi.

Banyak Is.

120 "

a

Nias

1200 "

u

Batu Is.

420 "

(I

Mentawi Is. (Siberut and Sipora)

1500 "

It

Pagi Is.

660 "

11

Enggano

200 "

a

Total

4600 sq.

mi.

To understand the isolation of these islands it is necessary to take
into account the depths of the surrounding seas. Simalur is about
sixty miles from the nearest point of Sumatra. Nias is approximately
fifty miles from Sumatra, and the other large islands follow this dis-
tance closely; Siberut sixty miles, the Pagi Islands forty-five miles,
Enggano sixty miles.

These differences can not be in themselves significant. The islands
are all about the same distance away from Sumatra. However, as
there seems to be no geological evidence to show that the west Sumatra
islands have ever been directly attached to the coast of Sumatra, we
must examine the position of the depth contours to see if any other
factors than purely geologic ones can be shown to exist. At once one
fact stands out. The 200 meter curve which most authorities (Brouwer

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 417

et al.) consider as defining the limits of the drowned Sunda land in
this region, extends out in two places to embrace two groups of the
west Sumatra islands, the Banyak Is. and the Batu Is. Southeast of
the Batus there is a tenuous connection with Siberut, Sipora, and the
two Pagi Is. which, however, are separated to the east from Sumatra
by the Mentawi trough. The other islands are all separated from the
200-meter curve, although in the case of Nias the separation is very
small. Examination of the map (Fig. 1.) will make this situation
plain. It is my contention then that during past epochs the Banyak
and the Batu Is. may have acted as stepping stones or funnels from
which the rest of the islands have been primarily colonized.

In order to make this plain I should like to proceed to a more
specific examination of the bird population of these islands with ref-
erence to the particular kinds and numbers of birds found on each
island. A tabulation of the families of birds in the area shows that
forty-six have been recorded so far from the islands. Of these, how-
ever, the following are purely migrants and so may be eliminated
from this discussion : Scolopacidae, Glareolidac, Pittidae.

Several other families are resident, but are of a type which may
be considered wide-ranging or wandering within the areas they in-
habit. I feel that it is best to discount members of these families from
the discussion as they tend to obscure the primary issue concerned
here which is one of local distribution and speciation. In this con-
nection reference is made to the short discussion preceding each
family in the Faunal List. These familes are : Phalacrocoracidae (only
one uncertain record for the islands), Ardeidae (a wide ranging family
subject to uncertain and erratic local migration), Ciconiidae, Anatidae
(uncommon among the islands), Rallidae (a widely spread family with
little speciation in the Malayan area), Charadriidae, Burhinidae,
Laridae (three more widely distributed families), Tytonidae (only one
member of which has so far been recorded from the islands), Meropidae
(a migrant family except for one uncertain record), Hirundinidae (a
widely ranging form), Motacillidae (another migrant family in the
area except for one widely-distributed form), Laniidae, and Ploceidae
(all records for which are presumably of common Greater Sunda
Island forms).

The remaining families, twenty-nine in number, have been listed
below with crosses indicating their presence on various island groups.
I have included Babi and Lasia in with Simalur, as they have no
families which are not found on the larger island. I have included the
Pagi Islands with the Mentawi Islands for the same reason. The sole

418

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exception in the Mentawi, Pagi distribution, Dissoura episcopus stormi
of the Ciconiidae, will undoubtedly be found on other islands in the
future. All migrant species of these families have been eliminated in
the following list.

Resident

Simalur

Banyak

Nias

Batu Is.

Mentawi

Enggano

Family

group

Is.

Pagi group

Accipitridae

X

X

X

X

X

Columbidae

X

X

X

X

X

X

Psittacidae

X

X

X

X

X

X

Cuculidae

X

X

X

X

X

Strigidae

X

X

X

X

X

Caprimidgidae

X

X

Micropodidae

X

X

X

X

Hemiprocnidae

X

X

X

X

X

Trogonidae

X

X

Alcedinidae

X

X

X

X

X

X

Coraciidae

X

Bucerotidae

X

X

X

Capitonidae

X

X

Picidae

X

X

X

X

Eurylaimidae

X

X

X

X

Cam pephagidae

X

X

X

X

Dicruridae

X

X

X

Oriolidae

X

X

X

Corvidae

X

X

X

SUtidae

X

Timaliidae

X

X

X

Pycnonotidae

X

X

X

X

X

Turdidae

X

X

X

X

X

x .

Sylviidae

X

X

X

X

X

X

Muscicapidae

X

X

X

X

X

X

Sturnidae

X

X

X

X

X

X

Nectariniidae

X

X

X

X

X

X

Dicaeidae

X

X

X

X

Zosteropidae

X

Total

23 = 79%

14 = 48%

26 = 89%

20=68%

21=72%

13=44%

A preliminary inspection of these percentages compared to the land
areas of the islands shows, as might be expected, that Nias, the

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 419

largest single island, has the largest bird population. However, the
next largest group is the Mentawi Is., Siberut and Sipora. Siberut is a
much larger island than Simalur and yet has a smaller bird fauna.
Again Enggano, which is larger than the Banyak group, has a smaller
bird fauna. Clearly the other factor of relative isolation of the islands
must be figured in to this consideration.

A further rough inspection of these figures on the bird families
reveals other interesting facts. If the Banyak and Batu Ids. have
acted as funnels for the bird fauna it is obvious that Xias must have
profited most greatly, as it is enveloped by the two groups. Let us
reexamine the distribution of families chart with this in mind.

The Accipitridae are common to all the islands except Enggano. The
Columbidae and Psittacidae are common to all. The Cuculidae may
turn up on the Banyak Is. when more collecting is done. However,
only three out of the thirteen forms from the islands occur on Simalur,
showing that the predominant distribution of this family has been
through the Batu group.

The Strigidae occur on all of the islands except the Batu group. I
think the absence of this family from this group is due to the vagaries
of collecting rather than to any real condition. The Caprimulgidae
occur only on Simalur and Nias. This would indicate that these birds
are confined to relatively large islands and that their distribution came
through the Banyak group. The Micropodidae and the Hemiprocnidae
occur on Nias and the islands to the south, including the Batus. How-
ever, they are found only on Simalur, not the Banyak Is., which might
indicate that Simalur was colonized from Nias. The Trogonidae occur
only on the Batu Is. and Nias. Here is a case where a family has reached
the islands certainly only by the shortest route. The Alcedinidae occur
on all the islands. The Coraciidae are found as residents only on
Simalur. The nearest relative of this subspecies on any of the adjacent
islands is the race gigas from the Andamans.

The distribution of the Bucerotidae is centered around the Batu Is.
funnel, having spread only to Nias on one side and the Mentawi group
on the other. Somewhat the same is true of the Capitonidae, confined
to the Batus and Nias. The Picidae have spread from the Banyak and
Batu Is. only to Simalur and Nias. The Eurylaimidae have a similar
but more southerly distribution, not having reached Simalur, but being
found in the Mentawi Is.

In the case of the Campcphagidae, no members of this family have
been found on the two small island groups. Presumably this is due to
the small area of the islands as there are no other apparent factors

420 bulletin: museum of comparative zoology

involved. Similarly, the Dicruridae are found only on the large islands,
as are the Oriolidae and Corvidae. However, these last three families
have not reached Enggano. The Sittidae have been recorded only once
from the islands and that record is quite recent (Junge 1936).

The Timaliidae and the following six Passerine families : Pycnonot-
idae, Turdidae, Sylviidae, Muscicajndae, Sturnidae, and Nectariniidae
all show indications by their distribution of having spread out from
the central islands. The Dicacidae are absent from the Banyak group,
but may perhaps be found there later.

The last family, the Zosieropidac, is represented by only a single
form on Enggano, which gives small clue as to its origin.

Thus twenty of the families represented by resident and local forms
on the islands seem to be in support of the contention that bird distri-
bution tends to derive from the islands on the 200-meter shelf. Of
the remaining nine, five are large island forms which may well have
come from the smaller islands as stepping s-tones, but which were not
able to maintain themselves there. Certainly 68% is a reasonable
majority with which to support the contention that the Banyak and
Batu islands on the 200-meter shelf have been the main source of
bird distribution.

I have made a diagram of the relationship of the size of the islands
compared with the percentage of families found on each group. In
order to check the percentage of families represented, I have also
depicted the percentage of the species resident on each group as com-
pared to the total number of species found on the islands. This total
number, one hundred ninety-eight, is taken only from the twenty-nine
families of truly static residents. In the following chart the island
groups are listed in terms of their isolation, Enggano being the most
isolated, the Banyak group the least isolated.

In the diagram below the two solid lines represent the percent-
ages of families (heavy line) and species (lighter line) found on
each group. It will be noted that the only appreciable difference
in the % of species as contrasted with the % of families is that be-
tween the Mentawi and Pagi islands. Although both have the same
number of resident families (21 out of 29), the Pagi islands have only
forty-two resident species compared to fifty-six on the Mentawi
islands. The broken line represents the areas of the groups in square
miles. As the Mentawi and Pagi Islands are composed of several
islands rather than a single land mass, I have reduced their total areas
by one quarter, in order to attempt to rationalize the reduced effect
of a land area of discrete units. The unit figures have been chosen

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 421

arbitrarily. The two percentage series have been plotted against these
figures in terms of 0 to 10 (=100), standing for percent. The area
series has been plotted against 0 to 12 (=1200), standing for square
miles.

Enggano

SlHALUR

Pagi

MENTAWI

NlAS

Batu
Banyak

Fig. 4

2 3 + 567

= % OF FAMILIES

= % OF 5PECIE5

= % LAND AREA (RATIONALIZED)

3

In contrast to these figures, it is important to list the occurrence of
endemic races on the west Sumatra islands. In the following list I
have shown the number of resident races on the island or island group,
the number of endemic forms found there, and the resulting percentage
of endemisms computed against residents. Whenever a west Sumatra
island endemic race occurs on more than one of the island groups I
have credited it to each of the islands where it occurs.

Islands

Resident forms

Endemic forms

%

Enggano

25

16

64

Simalur

61

34

55

Mentawi and Pagi

59

24

41

Nias

82

35

43

Batu

62

27

44

Banvak

28

13

46

422

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I have attempted to compute these percentages of endemic forms
against a series representing the degree of isolation of the islands.
As these islands have been shown to be little affected by the distance
in a straight line from the Sumatra coast, I have used the factors of
position with relation to the 200-meter shelf and relative distance
from island to island.

Enggano

SlMALUR.

Mentawi and
Pagi

NlA5

Batu
Banyak

Fig. 5

12 3 4 5 6

— -.«• = % OF ENDEMISMS

-% C06FFICIENT OF ISOLATION

10

In this diagram the solid line represents the percentage of
endemic races on each island. The broken line represents the co-
efficient of isolation of the islands. The isolation series has been ob-
tained arbitrarily in the following way. The Banyak islands average
about twenty-five miles from Sumatra. This distance then is picked
as a unit. The Banyak islands are on the 200-meter shelf, therefore
they stand as 1. The Batu islands are also on the shelf, but a little
farther from shore on the average. I have listed them as 1 + . Nias is
more than twenty-five miles from the Banyak or Batu islands, but less
than fifty. It, therefore, stands as 1+ on each of these counts. It
is also off the 200-meter shelf, which I have given the arbitrary value

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 423

of 2. Thus Nias is 3+ in isolation on two counts, but as the island
is enveloped by these two funnels, so that there is twice as much chance
for birds to have arrived there, I have divided the two figures into
each other, giving Nias an isolation value of 1 + . The Mentawi and
Pagi islands are all less than twenty-five miles from each other or from
the Batu islands. They are also all on the 200-meter shelf. Thus these
islands must have an isolation value of 1. Simalur is off the shelf
(=2), and is more than twenty-five miles from the Banyak Is. Its
isolation value is thus 4. Enggano, approximately one hundred fifty
miles south-east of South Pagi, is off the shelf (three units of fifty
miles = 6), and this added to the Mentawi-Pagi figure, makes 7.

Clearly the two charts considered above indicate a striking paral-
lelism between the grographic factors considered and the distribution
and speciation of the bird forms involved.

A few words about the endemic forms from the islands are perhaps
of value here. The primary relationships of the one hundred eleven
forms from the islands are with Sumatra. Ninety-three of the forms
are closest to forms found on Sumatra. Of the remaining races, the
affinities are as follows: Nicobar and Andaman Is. 9, Malay Peninsula
4, Java 1, Borneo 4. Two additional races are conspecific with races
found on the Andamans and Nicobars, 1; and on Java, 1.

Of the characters used in separating these endemic races there are
only two which are really important. These are size and color. So far,
no single other technique has been derived for estimating speciation
effects in birds than such simple gross morphological characters as
these. It remains for some later refinement of comparative anatomy
or genetics to be discovered which can be used to analyze speciation
in birds.

Using these two simple criteria thus we have the following figures:

Systematic factors on the west Sumatra Islands

Size, larger than nearest relatives 33

Size, smaller than nearest relatives 7

Color differences 32

Size and color differences together 39

Total 111

These figures indicate very clearly that the predominant character
of the endemic races of the west Sumatra islands is size, and that by
far the majority of the size races are larger than their congeners. It is
my impression that a dominant factor in the speciation of birds

424 bulletin: museum of comparative zoology

among the small islands of this region is that of larger size, but this
remains to be shown by a more general survey.

Of the two hundred eighty forms listed from the west Sumatra
islands, only forty-six or about 16% are known migrants. These
forms belong to the Ardeidae, Accipitridae, Chafadriidae, Scolopacidae,
Glareolidae, Cuculidac, Alcedinidae, Meropidae, Coraciidae, Pittidae,
Hirundinidae, Turdidae, Sylviidae, Musdcapidae, Motacillidae, Lani-
idae, and Sturnidae. Most of these migrant birds have no close rela-
tives on the islands and presumably have had no effect in the current
bird distribution. This is an important point to bring out, for to
anyone unfamiliar with the East Indian region it probably seems in-
credible that the resident species of birds are so sedentary that specia-
tion has actually been able to take place on these small islands.

Of all the migrant species there are only four that have races
resident in the area. These are :

1. Butorides striatus actophilus, a large migrant from Asia into the
islands during the winter. B. s. spodiogaster is the resident form of the
islands, distinguishable from actophilus primarily only by smaller size.

2. Surnicidus lugubris dicuroides, a large migrant from Asia during
the winter months. S. I. barussarum is the resident form, which differs
primarily from dicruroides by smaller size.

3 . Eurystomus orientalis ab undus and Eurystomus orientalis deignani,
two migratory races during the winter months from India or China,
and northern Thailand. E. o. oberholseri from Simalur differs from
these races by color characters.

4. Tcrpsiphone paradisi incei, a migrant from China recorded only
once from the islands. Differs primarily by color from the resident
forms, T. p. proccra and T. p. insular is.

In all four of the above cases there are resident forms on the neigh-
boring islands from which these west Sumatra island endemisms are
more likely to have been derived than from the Asiatic migrants. These
are: Butorides s. spodiogaster from the Nicobars or B. s. javanicus from
Sumatra; Surniculus I. brachyurus of Sumatra; Eurystomus o. orientalis
of Sumatra and the Malay Peninsula ; Terpsiphone p. nicobarica from
the Nicobars or T. p. affinis or madzoedi from Sumatra.

CONCLUSION

The islands off the west coast of Sumatra comprise a small archi-
pelago which represents a parallel upthrust to the late Tertiary
mountain-building movements along the western side of Sumatra.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 425

Presumably all the islands have always been separated i'rom Sumatra
with the possible exception of the Banyak and Batu island groups
which lie on the 200-meter Sunda shelf. These islands, the nearest to
Sumatra, have served as stepping stones or funnels by which the
majority of the fauna has reached the rest of the islands. In the case
of the birds found on these islands, G8% of the resident forms indicate
that this has been the distributional route followed.

The birds of the west Sumatra islands are considered to belong to
two hundred eighty species and subspecies, forty-six of which are
migrants into the area, principally from the Asiatic mainland. Of the
resident forms, one hundred eleven or 47% are considered to be en-
demic to the west Sumatra islands.

After a study of the geographical and geologic factors in the en-
vironment of these islands, it is presumed that there are two main
factors which influence the distribution and speciation of the birds
found here. One factor which seems to affect the distribution of the
resident birds is the actual area of the islands. Upon tabulating the
area of the different islands and arranging them on a scale an effort
was made to find out whether or not the number of resident species
showed a similar curve. The resident bird population was determined
in two ways, first by the percentage of the total of resident families
found on each island or group, second by the percentage of the total
number of resident species. Upon comparison with the area curve a
surprisingly reasonable agreement was noted.

The other factor, the relative isolation of the islands, was measured.
By determining the distance of the Banyak Islands from the shore of
Sumatra and calling this arbitrarily "one unit" a distance scale was
set up. Then by arbitrarily attributing "two units" to any equivalent
distance off the 200-meter shelf, a relative isolation scale was set up.
The resulting figure for each island, the coefficient of isolation, was
plotted on a simple scale.

This scale was compared with another one obtained by plotting the
percentage of endemic forms as compared with the total number of
resident forms for each island. These two scales were shown also to
agree very well one with the other.

Of the forms considered by the author to be endemic on the islands
84% are shown to be derived definitely from the fauna of Sumatra.
Among the remaining forms, the Andaman and Nicobar Islands seem
to claim the largest proportion of affinities.

Among the endemic forms considered there is a small majority in
favor of larger size as the single most important systematic character

426 bulletin: museum of comparative zoology

of the islands. Of the forms which embody two characters, size and
color, larger size is important in 80% (31 out of 39) of the cases,
making a total value for this character of 58%.

Thus the birds of the west Sumatra islands demonstrate that in the
process of speciation there has been a distinct correlation between the
relative degree of isolation of the islands inhabited by them. Further-
more, there seems to be a definite trend in the speciation of these
forms. All through the different families a dominating and recurring
character is that of larger size. Lastly, the distribution of the birds of
these islands shows a correlation between the number of species found
on any island and the size of the island, as well as indicating that the
main route for the ingress of birds has been via the Banyak and
Batu Islands.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 427

BIBLIOGRAPHY

1863. NlEWENHUISEN J. T. EN ROSENBERG H. C. B. VON

Verslag omtrent het eiland Nias en deszelfs bewoners. Verhandalingen
van het Bataviaasch Genootschap van Kunsten en Wetenschappen,
Vol. 30.
1878. Rosenberg, H. C. B. von'

Der Malayische Archipel. Leipzig, 1878.

1887. Salvadori, Tommaso

Catalogo delle collezioni ornitologische fatte presso Siboga in Sumatra
e nelT isola Nias del Sig. Elio Modigliani. Ann. Mus. Civ. Storia Nat.
Genova, ser. 2, vol. 4, 1886 (1887), pp. 514-563, pis. 8-9.

1890. Modigliani, E.

Un. Viaggio a Nias. Milano: 1890, roy. 8 vol., pp.i-xvi, 1-724, pis. 1-15.

1892. Salvadori, Tommaso

Uccelli di Engano, raccolti dal. Dott. E. Modigliani. Ann. Mus. Civ.
Storia Nat. Genova, ser. 2, vol. 12, 1892, pp. 123-142.

1892. Oustalet, E. .

Catalogue des Oiseaux rapporte de l'lle Nias, par M. J. Claine en 1891.
Bull. Soc. Philomatique, Paris, ser. 8, vol. 4, 1892, pp. 107-122.

1894. Salvadori, Tommaso

Viaggio del Dr. Elio Modigliani nelle Isole Mentawei — Catalogo di
una Collezione di Uccelli di Si-pora. Ann. Mus. Civ. Storia Nat.
Genova, ser. 2, vol. 14, 1894, pp. 588-601.

1895. Berlepsch, Hans Graf von

Description of two new Species of the Genera Phoenicophaes and
Spilornis, with a note on Oriolus consobrinus Rams. Nov. Zool., vol. 2,
1895, pp. 70-75.

1896. Buttikofer, Dr. J.

On a collection of birds from Nias. Notes from the Leyden Museum,
vol. 18, 1896, pp. 161-198.

1898. Hartert, Ernst

Einiges iiber Vogel von der Insel Nias. Ornith. Monatsber., vol. 6,
1898, pp. 89-94.

1899. Flnsch, Dr. O.

On three apparently new species of birds from the Islands of Batu,
Sumbawa, and Alor. Notes from the Leyden Museum, vol. 20, 1899,
pp. 224-226.

1899. Finsch, Dr. O.

Das Genus Gracula Linn, und seine Arten nebst Beschreibung einer
neuen Art. Notes from the Leyden Museum, vol. 21, 1899, pp. 1-22,
2 pis.

428 bulletin: museum or comparative zoology

1901. Blasius, W.

Bemerkungen iiber neue Sendungen malayischer Vogel II. Vogel von
der Insel Nias, gesammelt von Hugo Raap. Journ. fur Ornith., vol.
49, 1901, pp. 60-73.

1901. Hartert, E.

Some notes on Java birds. Nov. Zool., vol. 8, February 1901,
p. 51.

1902. Richmond, Charles W.

Descriptions of eight new birds from islands off the west coast of
Sumatra. Proc. Biol. Soc. Wash., vol. 15, 1902, pp. 187-190.

1903. Richmond, Charles W.

Birds collected by W. L. Abbott on the coast and islands of northwest
Sumatra. Proc. U. S. Nat. Mus., vol. 26, 1903, pp. 485-524.
1909. Oberholser, Harry C.

A revision of the Kingfisher Genus Ramphalcyon (Pelargopsis). Proc.
U. S. Nat. Mus., vol. 35, 1909, pp. 657-680.

1911. Oberholser, Harry C.

A monograph of the Flycatcher genera Hypothymis and Cyanonympha.
Proc. U. S. Nat. Mus., vol. 39, 1911, pp. 585-615.

1912. Richmond, Charles W.

Descriptions of five new birds from the west coast of Sumatra. Proc.
Biol. Soc. Wash., vol. 25, 1912, pp. 103-106.

1912. Oberholser, Harry C.

A revision of the forms of the edible-nest swiftlet, Collocalia fuciphaga
(Thunberg). Proc. U. S. Nat. Mus., vol. 42, 1912, pp. 11-20.

1912. Oberholser, Harry C.

Descriptions of one hundred and four new species and subspecies of
birds from the Barussan Islands and Sumatra. Smithsonian Misc.
Coll., vol. 60, 1912, no. 7, pp. 1-22.

1915. Oberholser, Harry C.

A review of the subspecies of the ruddy kingfisher, Entomothera coro-
manda (Linnaeus). Proc. U. S. Nat. Mus., vol. 48, 1915, pp. 639-657.

1919. Oberholser, Harry C.

A revision of the subspecies of the white-collared kingfisher, Sauropalis
Moris (Boddaert). Proc. U. S. Nat. Mus., vol. 55, 1919, pp. 351-395

1919. Oberholser, Harry C.

Notes on Dr. W. L. Abbott's second collection of birds from Simalur
Island, western Sumatra. Proc. U. S. Nat. Mus., vol. 55, 1919, pp.
473-498.

1923. Oberholser, Harry C.

Descriptions of new East Indian birds of the families Turdidae, Syl-
vlidae, Pycnonotidae, and Muscicapidae. Smithsonian Misc. Coll.,
vol. 76, 1923, no. 6, pp. 1-9.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 429

1923. Oberholser, Harry C.

Descriptions of New East Indian Nectariniidae. Journ. Wash. Acad.
Sci., vol. 13, 1923, pp. 226-232.

1924. Oberholser, Harry C.

Descriptions of new Treronidae and other non-passerine birds from the
East Indies. Journ. Wash. Acad. Sci., vol. 14, no. 13, 1924, p. 294.

1926. Chasen, F. N. and Kloss, C. Boden

Spolia Mentawiensis — Birds. Ibis, 1926, pp. 269-306, pi. 3.

1926. Oberholser, Harry C.

Descriptions of nineteen new East Indian passerine birds. Journ.
Wash. Acad. Sci., vol. 19, 1926, pp. 515-522.

1927. Riley, J. H.

Description of a new owl from Engano Island. Proc. Biol. Soc. Wash.,
vol. 40, 1927, pp. 93-94.

1927. Riley, J. H.

Spolia Mentawiensia — Three new birds from the Mentawi Islands.
Proc. Biol. Soc. Wash., vol. 40, 1927, pp. 95-96.

1929. Riley, J. H.

A review of the birds of the islands of Siberut and Sipora, Mentawi
Group (Spolia Mentawiensia). Proc. U. S. Nat. Mus., vol. 75, art. 4,
1929, pp. 1-45.

1929. DE SCHAUENSEE, R. M.

A new barbet from the Batu Islands. Proc. Acad. Nat. Sci. Phila.,
vol. 81, 1929, p. 521.

1935. Chasen, F. N.

A Handlist of Malaysian Birds. Bull. Ramies Mus. No. 11, 1935, pp.
i-xx+389.

1935. Junqe, G. C. A.

The mysterious Carpophaga vandepolli Biittikofer. Zoologische Mede-
deelingen, 18, 1935, pp. 161-2.

1936. Junge, G. C. A.

Fauna Simalurensis — Aves. Temminckia, 1, 1936, pp. 1-76, pis. 3.

1937. Junge, G. C. A.

Further Notes on the Birds of Simalur. Temminckia, 2, 1937, pp.
197-202.

1937. Neumann, O.

Description of four new races from Sumatra and the Mentawi Archi-
pelago. Bull. Brit. Orn. Club., 57, 1937, p. 151.

1938. Junge, G. C. A.

On a Collection of Birds from Enggano. Treubia, Deel 16, An. 3,
August, 1937, pp. 339-356.

430 bulletin: museum of comparative zoology

1939. Junge, G. C. A.

Description of a New Bird from Simalur. Zoologische Mededeelingen,
22, 1939, p. 120.

1940. DE SCHAUENSEE, R. M. AND RlPLEY, S. DlLLON

Birds from Nias Island. Zoological Results of the George Vanderbilt
Sumatran Expedition, 1936-1939. Part III, Proc. Acad. Nat. Sci.
Phila., 91, 1940, pp. 399-413, pis. 2.

1940. DE SCHAUENSEE, R. M.

The Birds of the Batu Islands. Proc. Acad. Nat. Sci. Phila., 92, pp.
23-42, map 1.

1941. Ripley, S. Dillon

A New Race of Ceyx erithacus. Proc. New England Zool. Club., 19,
1941, p. 15.

1942. Ripley, S. Dillon

Ceyx erithacus and rufidorsus. Zoologica, 27, pt. 2, July 20, 1942,
pp. 55-59.

1942. Ripley, S. Dillon

Notes on Malaysian Cuckoos. Auk, 59, Oct. 1942, pp. 595-596.

1943. Ripley, S. Dillon

The name of the Sumatran crested olive bulbul. Auk, 60, April 1943,
pp. 268269.

1943. Ripley S. Dillon

Description of a new Copsychus from the Batu Islands. Notulae
Naturae, No. 114, 1943, (Jan. 28).

PLATES

PLATE 1

Riplej — Sumatran Birds

PLATE 1

Fig. 1 . Stone-paved road on Nias. Secondary growth and native plantings
of coconuts line the majority of these trails.

Fig. 2. Stream near Hilisimelano, Nias Is., June, showing the typical
condition of the stream beds during summer. The suspension bridge is a recent
innovation. On the hill in the background there are traces of the old original
forest.

BULL MUS. COMP. ZOOL.

Ripley. Sumatran Birds. Plate 1

PLATE 2

Ripley — Sumatran Birds

PLATE 2

Fig. 1. Sibarau River, Sipora Is. This is the main river of the island and
one of the important routes of communication. The banks are well planted
with coconuts and bananas near the villages.

Fig. 2. Nest of Hirundo tahitica javanica with two fledglings ready to fly
(June, 1939). This nest was built on the front porch of a government rest
house at Hilisimetano, Nias Is.

BULL. MUS. COMP. ZOOL.

Ripley. Sumatran Birds. Plate 2

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 9

FOSSIL CETACEANS FROM THE FLORIDA TERTIARY

By Remington Kellogg

With Six Plates

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

November, 1944

Zoology \

No. 9 — Fossil Cetaceans from the Florida Tertiary1

By Remington Kellogg

A number of new and strange types of extinct marine mammals
have been brought to light during the past 25 years by the commer-
cial development of the Florida phosphate beds. Several new kinds of
fossil cetaceans from these deposits were described by the late Dr.
Glover M. Allen, and others, including the small collection here de-
scribed, were awaiting his attention. Through the kindness of Dr.
Thomas Barbour, who made the necessary arrangements for my visit
to Cambridge, I was accorded the privilege of studying these specimens.
If the general composition of the Miocene marine faunas of Europe
be accepted as a valid basis for correlation, then some of the cetaceans
that have been reported to have been dug out of the Bone Valley pebble
phosphates are clearly older than the Pliocene and not younger than
the upper Miocene. Cooke and Mossom (1929, p. 164) seem to have
been the first to suggest that the extinct marine mammals found in
these pebble phosphates have been reworked from older formations,
particularly the Hawthorn formation, and this conclusion may be
applicable in part at least to some of the cetaceans, inasmuch as the
long beaked porpoises found in Polk County, Florida, are restricted
to the Miocene in European deposits. Although the remains of three
long beaked porpoises (Schizodelphis depressus, Schizodelphis bobengi,
and Pomatodelphis inaequalis), that have been collected in Polk
County, are limited to sections of rostra and mandibles, the structural
details of these fragments are so unlike those of Pliocene porpoises
there is slight possibility of mistaken identification. No complete skull
or associated skeletal parts of cetaceans have ever been reported from
the Bone Valley pebble phosphates. The geologic age of the river
porpoise (Goniodelphis hudsoni) can not be determined with certainty,
since this type of odontocete modification occurs in both the upper
Miocene and the lower Pliocene. The small sperm whale (Kogiopsis
floridana) and the balaenopterid hereinafter described seem to be
representatives of the Pliocene fauna.

Since most of the recorded species are based on portions of the rostra
and of mandibles, it may be assumed that either (1) the fossilized
skeletal elements were broken up in the course of commercial dredg-
ing and hydraulic mining, or (2) they represent reworked material
from an older formation, or (3) they were dislodged from the laminated
blue clays underlying the phosphate deposits. A more plausible explan-

1 Published with the permission of the Secretary of the Smithsonian Institution.

434 bulletin: museum of comparative zoology

ation for this mixed association of types of cetaceans, that hitherto
have been known to occur only in geologic stages of different age, may
be found when precise field studies are made of the actual occurrence
of these bones in the commercial pits. Officials at the plant of the
American Agricultural Chemical Company informed Dr. White (1942,
p. 87) that the light brown, dark brown and black specimens came
from the pebble phosphate and that the pure white specimens came
from the underlying laminated blue clays. Most of the pure white
specimens that have been examined represent odontocetes that are
considered to belong to the Miocene fauna. One notable exception is
found in the material referred to Goniodelphis hudsoni, which consists
of the grayish white type skull, the light brown ankylosed mandibular
rami, and the almost white section of the right mandibular ramus.
Some of the specimens belonging to the long beaked porpoises are
likewise grayish white. One explanation that may be offered is that
some discoloration of reworked specimens subsequently incorporated
in more recent deposits may be expected in these shallow formations.

INIIDAE

Goniodelphis hudsoni G. M. Allen

Plate 1; pi. 2, fig. 1

Type. A portion of a cranium, no. 3920, Vertebrate Paleontology
Catalogue, Museum of Comparative Zoology. Collector, H. L. Hudson.

Referred specimens. (1) A short portion of right mandibular ramus,
no. 17879, Vertebrate Paleontology Catalogue, Museum of Compara-
tive Zoology. (2) The major portion of the ankylosed mandibular
rami, no. 17881, Vertebrate Paleontology Catalogue, Museum of
Comparative Zoology. Collector, George C. Elmore, 1941.

Horizon and locality. The type and the ankylosed mandibular rami
presumably were derived from the pebble phosphate deposits, which
belong to the lower Pliocene Bone Valley formation; the short portion
of the right mandibular ramus is thought to have been removed from
the laminated blue clays, immediately below the pebble phosphate,
which are tentatively referred to the middle Miocene Hawthorn forma-
tion. All three of these specimens were found in pits of the American
Agricultural Chemical Company at Pierce, Polk County, Florida.

Description. Symphyseal portion of mandibular rami (no. 17881,
M.C.Z.), measuring 520 mm. in length, broken transversely at seven
places; external surface of right ramus (pi. 1, fig. 2) weathered behind

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERT1 \KY 435

anterior 200-210 mm.; distal 165 mm. of symphysis curves distinctly
upward toward anterior broken extremity; symphysis decreases in
diameter gradually toward anterior end, both transversely and ver-
tically; angle formed by opposite rami behind symphysis approxi-
mately 45 degrees; more than 30 alveoli in each ramus; teeth near
anterior end of symphysis implanted in pairs (although opposite
alveoli are not separated by equivalent intervals from preceding and
succeeding alveoli, the general effect is that of paired teeth); 29th and
30th alveoli (counting forward) in left ramus more or less pandurate
in outline; corresponding alveoli in right ramus more nearly elliptic al
in outline; antero-posterior expansion and side to side compression of
roots of mandibular teeth most conspicuous on seven anterior pairs of
teeth; roots of anterior mandibular teeth measure about 15 mm.
anteroposterior^' (measurements can be taken only near middle of
length of root and it is quite possible that distal end of root is some-
what more expanded) ; behind 24th pair of teeth (counting forward)
roots at alveolar level progressively appear less flattened from side to
side; teeth less regularly spaced and tend to alternate behind 20th
pair of alveoli (counting forward); roots of corresponding teeth more
noticeably swollen internally and not so conspicuously expanded
anteroposteriorly; alveolar walls broken down for a distance of 60
mm. in front of hinder end of symphysis (boundaries of individual
alveoli are so indistinctly defined that it is impossible to describe or.
measure each individually); opposite alveoli separated by a distance
of approximately 5 mm. in portion 100 to 200 mm. behind anterior
extremity of symphysis ; interval between opposite tooth rows increases
imperceptibly toward hinder end of symphysis and measures about 17
or 18 mm. in portion immediately in front of fork of rami ; no indication
of dorsoventral constriction of rami immediately behind posterior end
of symphysis corresponding to condition shown by type mandible of
somewhat larger Saurocetes argcntiniis (Burmeister, 1871, pi. 1, fig. 1);
five or six minute nutrient foramina located on external face of right
ramus below 28th and 29th alveoli (counting forward); several scat-
tered foramina on ventral surface of symphysis; approximately 5 mm.
above ventral margin of external face of symphyseal portion of left
ramus is a narrow, seemingly discontinuous groove, from which grooves
of similar width spaced apart at intervals varying from 10 to 20 mm.
extend obliquely forward and upward toward alveolar margin of
ramus; anteriormost groove curves upward to about level of center of
28th alveolus (counting forward); second groove ends near anterior
end of alveolus of 27th tooth (counting forward) ; third groove ends

436

bulletin: museum of comparative zoology

indistinctly near alveolus of 26th tooth (counting forward); fourth
groove ends abruptly about 15 mm. below alveolar level of 23rd tooth
(counting forward); fifth groove ends abruptly about 15 mm. below
alveolar level of 21st tooth (counting forward); remainder of external
surface of left ramus weathered to such an extent that original posi-
tion and direction of grooves can not be determined; on external face
of symphyseal portion of right ramus, hindermost visible groove, about
95 mm. in length, extends upward and forward from near ventral
margin of ramus to near hinder edge of alveolus of 26th tooth (counting
forward); about 15 mm. below above mentioned groove, another
similarly directed shorter groove extends toward level of anterior
margin of alveolus of 26th tooth (counting forward); anteriormost
lateral groove terminates near hinder edge of alveolus of 28th tooth
(counting forward).

Section of right mandibular ramus (no. 17879, M.C.Z.), measuring
252 mm. in length; ramus (pi. 2, fig. 1) apparently bends upward

Fig. 1. Goniodelphis hudsoni, cross section near hinder end of symphysis,
left ramus restored, no. 17879, M.C.Z.

behind level of posterior end of symphysis and seemingly increases in
dorso-ventral diameter toward coronoid process; external face of
symphyseal portion of right ramus somewhat convex; outer surfaces
of opposite rami form more or less V-shaped ridge along ventral line
of ankylosis anteriorly; ramus not distinctly constricted behind

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 437

symphysis and no indication of pits for reception of apices of teeth in
upper jaw like on ramus of Saurocetes argentinvs (Burmeister, 1871,
pi. 1, fig. 1); longitudinal groove on external face narrow, approxi-
mately 10 mm. above ventral margin (somewhat similar to groove
shown on mandible figured by Burmeister, 1871, pi. 1, fig. 1); two
small foramina on lower external face of right ramus, approximately
39 mm. apart, below 2nd and 5th teeth (counting forward); not more
than 2 teeth in right ramus wholly behind level of posterior end of
symphysis; teeth (fig 1) with simple conoidal crown, slightly curved
backward and inward toward apex; crown not noticeably laterally
compressed; yellowish brown enamel on crown unevenly wrinkled by
fine striae; neck of root below crown short, not markedly constricted;
root swollen below neck, more noticeably internally than externally,
and expanded antero-posteriorly; distal end of root conspicuously
expanded antero-posteriorly and markedly flattened frcm side to side.

Measurements (in millimeters) :

Length of ankylosed mandibular rami (no. 17881, M.C.Z.), as pre-
served 520

Right mandibular ramus, 27 anterior alveoli (4th to 30th alveolus

counting forward from hindermost) in an interval of 473

Right mandibular ramus, 6 anterior alveoli (24th to 29th alveolus

counting forward) in an interval of 119

Right mandibular ramus, 5 posterior alveoli (4th to 8th alveolus

counting forward) in an interval of 65

Dorso-ventral diameter of right ramus about 30 mm. in front of
posterior end of symphysis 39 +

Left mandibular ramus, 27th tooth (counting forward) : Right Left

antero-posterior diameter of tooth near middle of length of

root 16.8 .... 15.0

transverse diameter of tooth near middle of length of root 7.8 .... 8.5

Left mandibular ramus, 25th tooth (counting forward) :
antero-posterior diameter of tooth near middle of length

of root 15.T. . . . 17.4

transverse diameter of tooth near middle of length of root 5.5 .... 7.2

Dorso-ventral diameter of right ramus between 23rd and 24th alveoli

(counting forward) 33.2

Greatest transverse diameter of same 24.7

Length of a portion of right ramus (no. 17879, M.C.Z.) 252

1 1 teeth in an interval of 145

5 posterior alveoli (4th to 8th alveolus counting forward from hinder-
most) in an interval of 65

438 bulletin: museum of comparative zoology

Dorso-ventral diameter of right ramus at posterior end of symphysis 42.5

Dorso-ventral diameter of right ramus at level of 11th tooth (count-
ing forward) 40

Right ramus, 6th tooth (counting forward) :

antero-posterior diameter of crown at base 6.5

transverse diameter of crown at base 6.3

height of crown, inside 7.2

antero-posterior diameter of root at alveolar level 11.0

transverse diameter of root at alveolar level 9.5

Right ramus, isolated tooth (probably 8th counting forward) :

greatest length of tooth 26.3

antero-posterior diameter of root near extremity 15.2

antero-posterior diameter of expanded portion of root below crown 12.0
transverse diameter of expanded portion of root below crown .... 9.4

antero-posterior diameter of crown at base 6.5

transverse diameter of crown at base 6.2

height of crown, outside 7.8

Type skull (no. 3920, M.C.Z.), apex of supraoccipital to point of di-
vergence of opposite premaxillaries (corresponding in cross section
to level of posterior margin of hindermost alveolus in right maxil-
lary) 250

Apex of supraoccipital to level of assumed antorbital notch on right
maxillary (Allen, 1941, pi. 1) 230

16 alveoli in left maxillary in an interval of 167

8 alveoli (1st to 8th counting forward) in right maxillary in an inter-
val of ' 93

Distance between inner margins of hindermost alveoli in right and

left maxillaries 71.8

Distance between inner margins of 6th teeth (counting forward) in
right and left maxillaries 25

Distance between inner margins of 9th teeth (counting forward) in
right and left maxillaries 11

Right maxillary, 7th tooth (counting forward) :

antero-posterior diameter of crown at base 7.8

transverse diameter of crown at base 6.7

antero-posterior diameter of root at alveolar level 10.0

transverse diameter of root at alveolar level 9.5

Length of right palatal groove ( anterior wall of narial passage to an-
terior end of groove) 98

Palatal surface, anterior wall of left narial passage to anterior end of
palatal exposure of vomer 241

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 439

Remarks. Some of the extinct porpoises, which have been compared
with Goniodclphis hudsoni by Allen (1941, pp. 7-8) are considered
by the writer to have somewhat different relationships.

The extinct porpoise Saurocetes argentinus (Burmeister, 1871, p. 51)
was based on two fragments of mandibles from an unknown locality,
although the matrix indicated that these specimens had been found in
the early Pliocene deposits on the shores of the Parana River. The
larger fragment (Burmeister, 1871, pi. 1, fig. 1) comprising the hinder
portion of the symphysis, both rami having been broken off a short
distance behind the latter, is 15 inches (381 mm.) long and 2 3^2 inches
(63.5 mm.) dorso-ventrally in front of hinder end of symphysis, but
only 134 inches (44.45 mm.) at the distal end. The symphyseal por-
tion is 11 inches (279.4 mm.) long, l^i inches (38 mm.) wide at distal
end, and 2}/§ inches (55 mm.) wide at posterior end. Burmeister
estimated the length of the entire mandible to be 30 to 32 inches.
The smaller fragment (Burmeister, 1871, pi. 1, fig. 4) represents a
short piece of the right ramus from the region immediately behind the
symphysis. On the outer face of the left ramus and somewhat above
the ventral margin is a channel or furrow, beginning at about the
posterior end of the symphysis and extending forward, from which
numerous wrinkles rather evenly spaced extend obliquely forward and
upward to below the alveolar margin. In cross section (Burmeister,
1871, pi. 1, fig. 2) the symphysis is triangular, with rounded contours.
The median region between the alveoli is somewhat elevated above the
sides. There are 12 alveoli (6 teeth) in the left ramus and 7 alveoli (3
teeth) in the right ramus. The teeth are not closely approximated
anteriorly, and behind and external to each is a small circular cavity,
apparently for lodging the apex of the corresponding upper tooth
when the jaws were shut. The hindermost tooth in the left ramus
alone is situated behind the posterior end of the symphysis. The teeth
are large, having conical crowns which are slightly compressed from
side to side, somewhat curved backward, and covered irregularly with
wrinkled enamel. Between the base of the crown and the gibbous
portion of the root is a well marked neck or constriction. The extrem-
ity of the root is compressed from side to side and irregularly divided
into two or three rootlets. The detached tooth figured by Burmeister
(1871, p. 54, pi. 1, fig. 3) is 2 inches (50.8 mm.) long, of which the height
of the crown is 8 lines (18.86 mm.), the neck \x/i lines (3.18 mm.),
and the length of the root 15 lines (31.8 mm.).

Inasmuch as Saurocetes Burmeister was considered to be preoccupied
by Saurocetus Agassiz, and since he had ascertained that the teeth of

440 bulletin: museum of comparative zoology

the Argentine odontocete were quite different from those of the animal
previously described by Agassiz, Burmeister (1891a; 1891b, p. 162)
withdrew the name Saurocetes argentinus and replaced it with Sauro-
delphis argentinus. In August of the same year, Ameghino (1891b, p.
255) proposed Pontoplanodes as a substitute for the generic name
Saurocetes Burmeister, and specifically designated Saurocetes argenti7ius
as the genotype. Consequently, as pointed out by Cabrera (1926, p.
397), Saurocetes argentinus Burmeister (January, 1871, )Saurodelphis
argentinus Burmeister (June, 1891), and Pontoplanodes argentinus
Ameghino (August, 1891) are absolute synonyms. Both Rovereto
(1915, p. 143) and Cabrera prefer to employ Saurodelphis in place of
Saurocetes, notwithstanding the provisions of the International Rules
of Zoological Nomenclature (see art. 36, recommendations; "names
which differ from generic names already in use only in termination or
in a slight variation in spelling . . . are not to be rejected on this
account," as for example Polyodonta, Polyodontas, Polyodontus.).

In 1892, Burmeister described and figured an imperfect skull from
the cliffs at La Curtiembre on the shore of the Parana River. In
restoring the skull, Burmeister (1892, pi. 8, figs. 1 and 5) used the skull
of the Recent Stenodelphis as a model, but neglected to show the three
teeth preserved in the right maxillary and added the terminal portion
of the rostrum (1892, pi. 8, fig. 2) of another odontocete. Burmeister
(1892, p. 456, pi. 8, fig. 6) reconstructed the type mandible of Sauro-
cetes argentinus by adding the anterior end of the symphysis of another
individual, in which the teeth are smaller and have a very large,
laterally compressed and antero-posteriorly expanded root of irregular
form, and a high laterally compressed crown. The terminal portion of
the symphysis diminishes in height rapidly near the tip. This recon-
structed skull and mandible were referred by Burmeister to Sauro-
delphis argentinus, and much of the confusion regarding the structural
peculiarities of this extinct porpoise may be traced to this restoration.

Abel (1909, pp. 257, 271), having obtained photographs of the
above-mentioned skull and the terminal portion of the rostrum, con-
curred with the opinion written by F. Ameghino that they belonged to
two different porpoises, and stated that the cranium undoubtedly
represented a member of the Iniidae. Abel, however, adopted an ill-
advised procedure to make names available for these two porpoises.
The name Saurodelphis argentinus was restricted by Abel to this skull.
Abel (1909, pp. 258-259), furthermore, concluded that the mandible
described by Burmeister in 1871, the terminal portion of the rostrum
figured by Burmeister in 1892, as well as the mandibular fragment

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 441

described by Ameghino (1891a, p. 163, fig. 71) under the name of
Saurocetes obliquus should be designated as Pontoplanodes argentinus.

The critical analysis published by Cabrera (1926, pp. 396-403)
shows rather conclusively that the type mandible (Burmeister, 1871,
pi. 1, fig. 1; Rovereto, 1915, pi. 2, figs. 1-2) and the terminal portion
of the rostrum (Burmeister, 1892, pi. 8, fig. 2; Abel, 1909, pi. 1, fig. 3)
are referable to Saurocetes argentinus. Cabrera (1926, p. 401) has also
shown that Saurocetes obliquus (Ameghino, 1891a, p. 163, fig. 71) does
not represent the anterior part of the mandibular symphysis as stated
by Ameghino, but agrees absolutely with the terminal portion of the
rostrum of Saurocetes argentinus.

According to Cabrera (1926, pp. 402-403), the skull of Saurocetes
argentinus is distinguished from that of Inia geoffrensis by the follow-
ing details: the elevated vertex (formed by union of nasals, frontals
and supraoccipital) is higher and more inclined backward; the anterior
border of the nasal passages, which is constituted by close approxima-
tion of the premaxillaries, forms an open inverted "V"; the lateral
occipital crests are more closely approximated, the minimum distance
between the crests being 55 mm.; the supraoccipital is deeply concave
dorsally; and the upturned lateral borders of the ascending plates of
the maxillaries and the underlying lateral supratemporal extensions of
the frontals form a narrower and deeper depression than in Inia. The
mandible is constricted dorso-ventrally behind the level of the posterior
end of the symphysis. The teeth (Rovereto, 1915, p. 146) are relatively
large, the antero-posterior diameter at base of the crowns of the man-
dibular teeth varying from 19 to 22 mm.; only one tooth is situated
behind the posterior end of the symphysis; the five posterior alveoli
(1st to 5th counting forward) in the left ramus occupy an interval of
94 mm., and the 7 anterior ones (6th to 12th) 188 mm.; the hinder-
most as well as the penultimate mandibular alveoli are rounded, but
thence forward the alveoli tend to assume an elliptical form; the
alveoli on the terminal portion of the rostrum and mandible are con-
stricted medially, the outline being pandurate; the antero-posterior
diameter of the root of the 5th tooth (type rostral fragment of Sauro-
cetes obliquus Ameghino, 1891, p. 163) at level of alveolar margin is
20 mm. and the antero-posterior diameter of the same tooth at base
of the crown is 16 mm. ; the five maxillary teeth occupy an interval of
90 mm. Notwithstanding the lack of harmony in the measurements of
the alveoli when computed in accordance with the scale of reduction
indicated for published illustrations (Burmeister, 1892, pi. 8, fig. 2;
Ameghino, 1898, p. 221, figs, 86a-d) of the terminal rostral fragment,

442 bulletin: museum of compakative zoology

it would appear that the antero-posterior diameter of these alveoli is
not less than 20 mm. and probably not more than 28 mm., since the
largest alveolus on the rostral fragment described by Cabrera (1926,
p. 400) measures 23 mm. antero-posteriorly and 9 mm. transversely.

It remained for Cabrera (1926, p. 403) to discover that the incom-
plete skull, which had been identified as Saurodelphis argentinus by
Burmeister (1892, pi. 8, figs. 1 and 5; Abel, 1909, pi. 1, figs. 1-2), as
well as another skull belonging to the Museo de La Plata should be
referred to Ischyrorhynchus vanbenedeni and to suggest that it was not
impossible that the type mandible of Anisodelphis brevirostratus
(Rovereto, 1915, p. 149, pi. 4, figs. 1-2) belonged to the same porpoise.
Cabrera seems to have been the first to notice that the type of Ischyro-
rhynchus vanbenedeni (Ameghino, 1891a, p. 163, fig. 72) did not repre-
sent a portion of the mandibular symphysis as stated by Ameghino,
but that it was actually a rostral fragment. That this allocation is
probably correct is shown not only by the form of the teeth in situ,
but also by the similarity in the dimensions of corresponding parts,
the transverse diameter of the type rostral fragment of Ischyrorhynchus
vanbenedeni being 31 mm. and that of the middle portion of the ros-
trum of Burmeister's skull 30 mm.

The imperfect skull of Ischyrorhynchus vanbenedeni (Burmeister,
1892, pi. 8, figs, 1, 5; Abel, 1909, pi. 1, figs. 1, 2), which lacks the
terminal portion of the rostrum as well as the occipital region, has a
length of about 637 mm. and was at least 303 mm. in breadth across
the bases of the zygomatic processes when complete. Abel (1909, pp.
269-271 )points out that the skull of Ischyrorhynchus is characterized
as follows : the posterior ends of the ascending plates of the maxillaries
project farther backward than in Inia; the vertex (constituted by the
frontals) is pushed farther back than in Inia; the distance between the
upturned borders of the ascending plates of the maxillaries is much
less than in Inia ; the posterior wall of the nasal passages is much less
inclined forward than in Inia and consequently more of the flat nasal
bones can be seen, when viewed from above. With reference to the
anterior border of the squamosal, when seen from above, the nasal
passages as well as the vertex and the posterior ends of the maxillaries
are pushed considerably farther backward than in Inia, but the last
two distinctions mentioned by Abel are based on somewhat dubious
assumptions. In addition, the skull of Ischyrorhyiichus can be dis-
tinguished readily from that of Inia by its size, by the length of the
palatal grooves, by the relations of the palatine bones, and especially
by the shape and dimensions of the teeth. Although the entire basi-

KELLOGG: FOSSIL CETACEAN'S FROM THE FLORIDA TERTIARY 443

cranial region is destroyed, the relations of the bones in the preserved
portion of the type skull indicate that Goniodelphis was somewhat
similar to Ischyrorkynchus in these above-mentioned details. Cabrera
(1926, p. 403) concludes that the skull of Ischyrorkynchus vanbenedeni
is distinguished from that of Saurocctcs argentinus by the greater eleva-
tion and large size of the knob-like vertex, and by the open inverted
"U" contour of the anterior border of the nasal passages.

Rovereto (1915, p. 151) has published the following measurements
for the maxillary teeth (probably the 14th or 15th counting forward)
of Ischyrorhynrhus vanbenedeni: antero-posterior diameter of crown at
base, 17 mm.; transverse diameter of crown at base, 10 mm.; height
of crown, 5 mm.; interval between opposite tooth rows, 6 mm. As
regards the teeth located near the middle of the rostrum, Ameghino
(1891, p. 165) states that the average diameter of their roots at the
level of the alveolus is 13 mm.; the crown of a detached tooth is said
to measure 9 mm. antero-posteriorly and transversely at the base.
Should the allocation of Anisodelphis brevirostratus (Rovereto, 1915,
p. 150) to Isehyrorhynchus vanbenedeni be confirmed, the mandible is
characterized as follows: the anterior alveoli are separated by inter-
spaces of 18 to 24 mm.; the 7 hindermost alveoli are separated by
interspaces of 5 mm. ; at the posterior ends of the tooth rows the teeth
are opposite one another and anteriorly they are not opposite but
alternated; one tooth is situated behind the posterior end of the
symphysis.

Presumably the identical specific names lead Allen (1941, p. 7) to
confuse the skull of the extinct ziphioid whale D iochotichus vanbenedeni
(True, 1910) with that of Ischyrorhynehus vanbenedeni.

Doubtless a certain relationship exists between Goniodelphis and
Proinia (True, 1909), but not a close one. In the last mentioned
genus, the posterior wall of the nasal passages is not at all inclined
backward and the ascending plates of the maxillaries, though narrow,
are directed forward and downward, rather than outward and upward.
While both genera have been assigned to the Iniidae, they do not
exhibit any close similarities.

Our knowledge of Hcsperoeetus californicus (True, 1912) is limited
to two pieces of the ankylosed rami, comprising the anterior portion
of the symphysis, which were found in the upper Miocene upper San
Pablo formation near Rodeo, California. Contrary to Allen's assertion,
Hcsperoeetus could hardly have been larger than Goniodelphis, since the
dimensions of the corresponding portions of the symphysis are approxi-
mately the same. Nevertheless, Hesperocetus does not seem to be

444 bulletin: museum of comparative zoology

related closely either to Goyiiodelphis or to any of the described Tertiary
Iniidae. The tips of the teeth in the upper series are fitted into elon-
gated depressions in the interspaces between the alveoli in the corre-
sponding lower series. This peculiar alternation in the symphyseal
region of transverse pairs of teeth and of transverse pairs of elongated
depressions in the interspaces is not duplicated in any of the known
extinct iniids. The teeth are relatively large, the crowns at their bases
measuring from 8 to 10 mm. antero-posteriorly and from 8 to 8.5 trans-
versely. The alveoli in the longest symphyseal fragment measure 9
mm. transversely and from 14 to 20 mm. antero-posteriorly; the inter-
spaces average about 20 mm. antero-posteriorly. There are three
alveoli in the left ramus in an interval of 91 mm. The presence of
relatively large foramina, which open into grooves not more than 10
mm. in length and which are spaced at intervals of 40 to 60 mm. on
the lateral symphyseal surface of the ramus, characterize the mandibles
of H esperocetus and Inia. No trace of a longitudinal lateral furrow,
such as is present on the symphyseal portions of the mandibles of
Goniodelphis and Saurocetes, can be detected on the mandibular sym-
physis of H esperocetus.

Goniodclphis is a small-toothed iniid, whose skull and mandibles may
be distinguished readily from those of previously described river por-
poises by the following combination of characters :

The vertex is narrow, transversely widened (32 mm.), and much less
elevated than on the crania of either Inia, Saurocetes or Ischyro-
rhynchus; the posterior wall of the nasal passages is not so steep as in
Inia but is inclined backward at an angle of about 30° with the palate;
a shallow depression, about 25 mm. long and deeper ventrally, located
at the top of the posterior wall of the nasal passages, marks the position
of the flattened squarish nasal bones; the longitudinal depression
formed by upturned lateral borders of ascending plates of maxillaries
and underlying lateral supra temporal extensions of frontals is not so
deep as in Inia; the anterior border of the narial passages is constituted
by close approximation of the premaxillaries, forming a wide open in-
verted "U" like in Ischyrorhynchus; the rostrum is noticeably expanded
near the base and narrowed toward the extremity; posterior maxillary
and mandibular alveoli are rounded; anterior maxillary and mandibu-
lar alveoli are elongated, the anteriormost mandibular alveoli being
somewhat pandurate in outline and thus resembling Saurocetes; the
teeth are much smaller than those of either Saurocetes or Ischyro-
rhynchus, the antero-posterior diameter of anterior mandibular alveoli
varying from 15 to 18 mm., and the antero-posterior diameter at base

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 445

of the crowns of mandibular teeth from 5.5 to 7.8 mm.; not more than
two teeth are situated behind the posterior end of the symphysis; the
5 anterior teeth (12th to 16th counting forward) on the type skull
occupy an interval of 61 mm., which corresponds very closely with the
60 mm. interval for the same teeth on an Inia skull (no. 239667,
U S.N.M.).

DELPHINIDAE

Megalodelphis, new genus

Genotype. Megalodelphis magnidens new species.

Diagnosis. Rostrum and symphyseal portion of mandibular rami
elongated and compressed dorso-ventrally; 7 teeth in ramus behind
level of posterior end of symphysis; crowns of mandibular teeth more
or less flattened from side to side and curved inward toward apex;
enamel on crowns lightly striated, with a distinct vertical carina on
anterior and posterior cutting edges.

This odontocete is further characterized by the unusually large di-
mensions of the upper and lower jaws, in fact it is the largest known
long-beaked porpoise, either extinct or living.

Megalodelphis magnidens, new species
Plate 2, fig. 2; pi. 3.

Type. A posterior symphyseal section of the ankylosed mandibular
rami, no. 17883, Vertebrate Paleontology Catalogue, Museum of Com-
parative Zoology. Collector, George C. Elmore, 1941.

Referred specimen. A short portion of the rostrum, no. 17880, Verte-
brate Paleontology Catalogue, Museum of Comparative Zoology, Col-
lector, George C. Elmore, 1941.

Horizon and locality. Laminated blue clays, immediately below the
pebble phosphate, which are tentatively referred to the Hawthorn
formation, in pits of the American Agricultural Chemical Company at
Pierce, Polk County, Florida. Middle Miocene (Simpson, 1932, p.
425; White, 1942, p.' 87).

Description. Short section of left mandibular ramus (pi. 3, fig. 2),
including hinder portion of symphysis, crushed; hinder portion of left
ramus split lengthwise, thus separating opposite walls of alveoli;
symphyseal region dorso-ventrally flattened, judging from conforma-
tion of hinder end ; seven alveoli in left mandibular ramus behind level

446 bulletin: museum of comparative zoology

of symphysis, with teeth in situ in three alveoli; alveoli relatively large
in comparison to size of roots of teeth; two cavities (fifth and sixth
counting forward from hindermost alveolus) interpreted as represent-
ing alveoli occupied by corresponding teeth in milk dentition ; fourth
permanent tooth (counting forward) in left ramus erupting normally
and protruding far enough for apex of crown to become worn; fifth
permanent tooth (counting forward) erupting in interspace between
fifth and sixth alveoli for milk teeth, with apex of crown worn; no
remnant of sixth permanent tooth, except for dubious cavity in spon-
giosa anterior to root of fifth permanent tooth; seventh permanent
tooth (counting forward) not fully erupted and apex of crown com-
plete; eighth permanent tooth (counting forward) fully erupted, with
apex of crown worn off; depression (length, 10 mm.; width, 7 mm.)
antero-external to eighth alveolus and in interspace between eighth
and ninth alveoli for apex of corresponding tooth in upper jaw ; ninth
and tenth alveoli represented by external rims; crowns of teeth, (pi. 2,
fig. 2) flattened from side to side, curved inward toward apex; generally
blackish enamel on crown with concentric bands of lighter color and
lightly ornamented with more or less vertical anastomosing striae;
distinct vertical carina on anterior and posterior cutting edges.

Fig. 2. Mcgalodelphis magnidms, cross section of rostral fragment, 70 mm.
in front of posterior end, no. 17880, M.C.Z.

Section of rostrum (no. 17880, M.C.Z.) 336 mm. in length; whole or
portions of seven alveoli present in left maxillary, six being sufficiently
complete for measurement; eight alveoli in right maxillary less satis-
factorily preserved, the four anterior to hindermost alveolus being

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 447

nearly complete; interval between opposite rows of alveoli varying
from 35 to 40 mm.; roots of maxillary teeth attenuated and bent up-
ward and backward, judging from direction and shape of alveoli;
ankylosis of parallel premaxillaries complete, line of union marked by
irregular furrow or groove; width of ankylosed premaxillaries anteri-
orly, 43.5 mm.; lateral rostral groove broad (10+ mm. in width), pre-
sumably marking line of ankylosis of premaxillaries and maxillaries;
mesorostral gutter (fig. 2) completely enclosed by premaxillaries and
maxillaries, and wider than high; opposite maxillaries ankylosed along
mid-line; apices of teeth in mandible seemingly in contact with upper
jaw in interspaces between maxillary alveoli, the depressions (pi. 3,
fig. 1) being located external to external margin of maxillary alveoli.

Measurements (in millimeters) :

Length, as preserved, of left mandibular ramus (no. 17883, M.C.-Z.) 330
Transverse diameter of rami at level of fork at hinder end of sym-
physis, estimated 145 *

9 hindermost alveoli in an interval of 213

Anterior margin of alveolus of third tooth to anterior margin of

root of eighth tooth (counting forward from hindermost alveolus) 117.6
Left mandibular ramus:

Fourth tooth, counting forward from hindermost alveolus :

antero-posterior diameter of crown at base 13.0

transverse diameter of crown at base 9.7

height of crown inside (apex worn) 14.0

antero-posterior diameter of root below crown 17.8

Tooth in interspace between fifth and sixth alveoli for milk teeth:

antero-posterior diameter of crown at base 13.0

transverse diameter of crown at base 10.0

height of crown inside (apex worn) 14.4

antero-posterior diameter of root below crown 14.5

Seventh tooth, counting forward from hindermost alveolus:

antero-posterior diameter of crown at base 14.7

transverse diameter of crown at base 9.7

height of crown inside (apex complete) 15.0

Eighth tooth, counting forward from hindermost alveolus:

antero-posterior diameter of crown at base 15.0

transverse diameter of crown at base 12.7

height of crown inside (apex worn) 21.3

antero-posterior diameter of root below crown 18.5

Length of rostral fragment (no. 17880, M.C.Z.) 336

Transverse diameter 70 mm. anterior to hinder end of rostral frag-
ment 70.5

448 bulletin: museum of comparative zoology

Vertical diameter 70 mm. anterior to hinder end of rostral fragment 47.5

4 alveoli in right maxillary in an interval of 136

6 alveoli in left maxillary in an interval of 244

Right maxillary, counting forward from hindermost alveolus:

Second alveolus, antero-posterior diameter 25

Second alveolus, transverse diameter 18

Interspace between second and third alveoli 12

Third alveolus, antero-posterior diameter 25.4

Third alveolus, transverse diameter 17.3

Interspace between third and fourth alveoli 12.3

Fourth alveolus, antero-posterior diameter 23.0

Fourth alveolus, transverse diameter 17.7

Interspace between fourth and fifth alveoli 12.3

Fifth alveolus, antero-posterior diameter 25.5

Fifth alveolus, transverse diameter 19.4

Remarks. There are only two extinct odontocetes that approach the
Florida porpoise in size. In view of the confusion that still seems to
persist regarding the valid name for one of these porpoises, it is neces-
sary to review briefly the history of the "dauphin a longue symphyse
de la machoire inferieure" of Cuvier (1825, ed. 3, vol. 5, p. 312, pi. 23,
figs. 4-5, 9-11), to which two names, Champsodelphis macrogenius
(Fischer) and Champsodelphis bordae (Holl), were applied in 1829.
Both of the above mentioned names were based on the incomplete
mandible and the rostral fragment, which Cuvier had described and
figured. These two specimens came from the Helvetian shell marl at
Sort; 8 kilometers from Dax, DepartementLandes, France, and must be
considered as co-types. Cuvier saw in the museum at Dax the incom-
plete mandible, measuring 16 French inches [ = 433 mm.] in length, on
which there are 12 alveoli (4 teeth) in the right ramus and 18 alveoli
(10 teeth) in the left ramus. The short rostral fragment, measuring
178± mm. in length, on which there are 4 alveoli (2 teeth) in the right
maxillary and 3 alveoli (1 tooth) in the left maxillary was presented in
1803 by Borda to the Museum National d'Histoire Naturelle, Paris.

Valenciennes (1862, pp. 789-790) seems to have been the first to
observe that the extinct porpoise to which the rostral fragment
(Cuvier, 1825, pi. 23, figs. 9-11) belonged was quite distinct from the
one represented by the mandibles (Cuvier, 1825, pi. 23, figs. 4-5). In
support of his contention that the name Champsodelphis macrogenius
had been founded on specimens representing specifically distinct por-
poises, Valenciennes in describing the rostral fragment directs atten-
tion to the teeth, which are characterized by their dimenisons, by the

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 449

absence of any trace of a small heel or blunt tubercle at the base of
the. crown posteriorly (notwithstanding the statement of Cuvier), and
by the different appearance of the enamel on the crown. Valenciennes,
however, did not restrict the name Champsodelphis macrogenius to
either of the above mentioned specimens.

Brandt (1873, p. 264) decided to ignore C hampisodclphis macrogenius,
inasmuch as this name was based on parts of two distinct porpoises.
For the rostral fragment (Cuvier, 1825, p. 313, pi. 23, figs. 9-11),
Brandt (1873, p. 266) proposed the new name Champsodelphis valen-
ciennesii. Consequently, Brandt actually restricted the name Champ-
sodelphis macrogenius to the mandible (Cuvier, 1825, p. 312, pi. 23,
figs. 4-5), notwithstanding the fact that he (Brandt, 1873, p. 263) pro-
posed the new name Champsodelphis macrognathus for this specimen.
Whether or not Champsodelphis macrogenius (Fischer) has priority
over Champsodelphis bordae(Ho\\) has not been determined, since both
were published in 1829, but both of these names are many years older
than Champsodelphis macrognathus (Brandt). Since this method of
elimination is not specifically covered by the International Rules of
Zoological Nomenclature, in order to obviate any possible confusion
the mandible is herewith designated as the lectotype of both C. macro-
genius and C. bordae.

The alveoli on the rostral fragment allocated to Megalodelphis mag-
nidens are much larger than those on the type rostral fragment of
Champsodelphis valenciennesii, the measurements of the alveoli of the
former varying from 23 to 25.5 mm. antero-posteriorly, and from 17.3
to 19.4 mm. transversely, whereas an alveolus of the latter measures

18.4 mm. antero-posteriorly and 14 mm. transversely. Furthermore,
the rostral fragment of C. valenciennesii has somewhat different pro-
portions, being higher than wide (dorso-ventral diameter, 55 mm.;
transverse diameter, 53 mm.), whereas the rostral fragment referred to
M. magnidens is noticeably wider than high (dorso-ventral diameter-

47.5 mm.; transverse diameter, 70.5 mm.).

It seems almost certain that the mandible found at Sort represents
a porpoise quite different from Megalodelphis magnidens, since the
teeth have a small heel or blunt tubercle at the base of the crown poste-
riorly and a somewhat smaller crown. The dimensions of one tooth
are stated by Cuvier to be as follows: height, 15 mm., and diameter at
base approximately, 11 mm.; the interspace between alveoli is stated
to be 20 mm. A more important difference is indicated by the presence
of ten or more teeth on the ramus posterior to the level of the hinder
end of the symphysis of Champsodelphis macrogenius, whereas only

450 bulletin: museum of comparative zoology

seven teeth are present on the corresponding portion of the mandible
of Megalodelphis magnidens. Moreover, the symphyseal portion of the
mandibles of C. macrogenius is much narrower, the transverse diameter
at the hinder end of the symphysis (50 mm.) being about one-third of
that of M. magnidens (145=*= mm.), and is less markedly compressed in
a dorso-ventral direction.

Distinction between Megalodelphis magnidens and the Miocene
M acrodelphinus kelloggi (Wilson, 1935, pp. 28-58, figs. 4-9), several
specimens of which have been collected in the Pyramid Hills sand,
about 5 miles southwest of Woody, Kern County, California, is not
restricted to minor details. It is obvious at first glance that the dorso-
ventrally compressed mandibular symphysis and rostrum of M. mag-
nidens is wholly unlike that of this Miocene porpoise from California.
One may infer that the elongated mandibular symphysis and rostrum
of M . magnidens suggested a long-beaked skull similar in general shape
to the corresponding portions of Pomatodelphis, Schizodelphis, and
Zarhachis. Moreover, behind the level of the symphysis there are 14
alveoli in an interval of 203 mm. on the right ramus of M. kelloggi in
contrast to the 7 alveoli in an interval of 162.5 mm. on the correspond-
ing portion of the mandible of M . magnidens. From the dimensions, it
will be seen that M . magnidens surpassed in size the largest of all pre-
viously described fossil porpoises.

If we examine in detail the type skull and other specimens allocated
to M acrodelphinus kelloggi, we find a number of well marked peculiari-
ties that suggest rather strongly some sort of a relationship with at
least one of the less completely known porpoises from the Helvetian
shell marl near Dax, France. The dimensions and conformation of the
cross section of the rostrum of M. kelloggi (Wilson, 1935, fig. 4a) are
surprisingly close to the corresponding rostral section of C. valencien-
nesii (see C hampsodelphis macrogenius Gervais, 1859, pi. 41, fig. 6b).
It is possible that the skull of C 'hampsodelphis valenciennesii may have
had a rostrum somewhat similar to that of M. kelloggi. Although simi-
lar in general conformation, the teeth of the French form may average
slightly larger than those of the Californian porpoise, as is indicated
by the following measurements :

Champsodelphis Macrodelphinus
valenciennesii kelloggi

In left maxillary, there are 3 alveoli in an

interval of 69.5 51.0

A right maxillary tooth:

Antero-posterior diameter of crown at

base 12.2 8.0

Transverse diameter of crown at base . . 9.5 9.0

Vertical height of enamel crown 15.0 13.4

Qz-vmo ollrviiran/-**} Virvnrotro-n mncf no mono T/-»r inrn \nHii Q 1 ciTZf* variation

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 451

A careful examination of the rostral fragment, which constitutes the
type of C ham psodd phis valenciennesii , failed to convince the writer
that Abel (1899, p. 841) was correct in suggesting that this species
might belong in the genus Tursiops. It is quite obvious that valencien-
nesii does not belong in the genus Champsodelphis. Since the type
rostral fragment seems not to differ appreciably either as regards size
or conformation from the corresponding section of the rostrum of
Macrodclphinus kelloggi, valenciennesii may be assigned tentatively
to the genus Macrodclphinus.

PHYSETERIDAE

?Hoplocetus, species indet.

Referred specimens. Two teeth, no. 17886, Vertebrate Paleontology
Catalogue, Museum of Comparative Zoology. Collector, George C.
Elmore, 1941.

Fig. 3. ? Hoplocetus, species indet., lateral view of tooth. Fig. 3a.
view of tooth, no. 17886, M.C.Z.

-anterior

Horizon and locality. Presumably from the pebble phosphate de-
posits, which are referred to the Bone Valley Formation, in pits of the
American Agricultural Chemical Company at Pierce, Polk County,
Florida. Lower Pliocene (Simpson, 1930, pp. 177-185; 1932, pp. 445-
446, 469).

452

bulletin: museum of comparative zoology

Description. The crowns of these two teeth are conical, 24 to 26
mm. in height, and slightly curved backward. The shortest tooth (fig.
3, 3a) has the enamel on the crown ornamented with coarse anastomo-
sing striae. Below the base of the enameled crown the neck of the root
is deeply worn on one side. The root is gibbous near the middle and
tapers to the extremity. The pulp cavity seems to be completely closed.

The other tooth (fig. 4) is considerably longer and exhibits a more
regular conformation. The enamel on the crown is less noticeably

Fig. 4. ? Hoplocetus, species indet., lateral view of tooth, no. 17886, M.C.Z.

wrinkled, although anastomosing striae directed more or less dorso-
ventrally are present. The long axis of the root is weakly curved from
end to end. The neck of the root is slightly constricted below the base
of the crown. At the extremity of the root is an orifice for the 112 mm.
long pulp cavity which extends toward the crown; the transverse
diameter of the pulp cavity is 10 mm.

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 453

Measurements (in millimeters) :

Greatest length of tooth 190.5 138.2

Antero-posterior diameter of crown at base 20.2 20.2

Transverse diameter of crown at base 21.5 20

Height of crown inside, apex worn 24 26

Greatest antero-posterior diameter of root 41.5 45.2

Greatest transverse diameter of root 36.6 39

Remarks. Teeth of this same general appearance have been referred
to several genera. Some of these genera seem to have served as "catch-
alls" for species that, although based solely on teeth and other inade-
quate skeletal remains, are derived from formations ranging in age
from lower Miocene (Langhian) to lower Pliocene (Plaisancian).
Nevertheless, Abel (1905, p. 52) placed eight of these genera in the
synonymy of Scaldicetvs. It is difficult to justify this procedure, espe-
cially in view of the similar appearance of the teeth of a number of
valid generic types of Miocene physeteroid whales, for which readily
recognizable skulls are available, as for instance Aidophyseter, Dia-
phorocetus, Idiophyseter, Idiorophus, Orycterocetus, Physetcrula, Scaldi-
cetus, and Thalassocetus. The mandibular teeth of the Recent sperm
whale Physeter catodon are not uniform in either size or conformation,
and there is no valid basis for an assumption that less individual
variation will be exhibited by the teeth of one of these extinct physete-
roids. The 45 teeth constituting the type of Scaldicetus caretti vary in
length from 200 to 240 mm. Regardless of variance of opinions as to
the precise generic allocation of species based wholly on teeth, it seems
desirable, nevertheless, in the absence of satisfactory information in
regard to the skull to refer these tw o teeth from Florida to a described
genus. Inasmuch as these teeth exhibit a general conformation some-
what similar to those of Hoplocetus crassidens, they are referred tenta-
tively to Hoplocetus. The genotype, Hoplocetus crassidens (Beneden
and Gervais, 1880, p. 340, pi. 20, figs. 26-27), was based on two teeth
from the middle Miocene (Helvetian) shell marl at Romans, Departe-
ment Drome, France. The enamel on the crown of the smallest tooth
is more distinctly wrinkled than that on the other tooth, and the apices
of the crowds of both teeth are worn. The crown of the largest tooth,
as preserved, measures 11 mm. in height and that of the other tooth
17 mm. These two teeth, one of which has a distinctly swollen root,
have the top of the root distinctly constricted below the base of the
crown, forming a short neck. The lengths of the roots of these two
teeth are respectively 110 and 94 mm.

454 bulletin: museum of comparative zoology

Portions of skulls of three odontocetes were allocated to Diaphoro-
cetus mediatlanticus by Allen (1921, p. 154). One of these, comprising
a section of the rostrum which measures 288 mm. in length as well as
the corresponding portion of the ankylosed mandibular rami (Allen,
1921, p. 155, pi. 12, fig. 13; no. 10922, Div. Vert. Paleont., U. S. Nat.
Mus.), seems to be allied to if not identical with M egalodelphis magni-
dens. The second, a section of the ankylosed mandibular rami which
measures about 150 mm. in length (Allen, 1921, p. 155, pi. 12, fig. 14),
in all probability is not referable to "Diaphorocetus" mediatlanticus
(Kellogg, 1925, p. 13, pis. 4-5), although it does belong to a sperm
whale. The portion of the base of the skull, comprising the occipital
condyles (Allen, 1921, p. 156, pi. 9, fig. 6) may represent the same ex-
tinct sperm whale.

No record seems to have been made of the stratigraphic position of
the above-mentioned specimens in the phosphate deposits of Polk
County, and consequently the two isolated teeth as well as the symphy-
seal fragment and the portion of the base of the skull may have been
removed either from the lower Pliocene pebble phosphate deposits or
from the underlying middle Miocene laminated blue clays. The trans-
verse diameter of the ankylosed mandibular rami just in advance of
the hinder end of the symphysis is about 57 mm. and the antero-
posterior diameters of the alveoli vary from 18 to 22.5 mm. It is obvi-
ous that the roots of the two isolated teeth are too large to be lodged in
a mandible of approximately the same dimensions as this symphyseal
fragment. Even though one concedes that the discrepancy in size be-
tween the two isolated teeth (no. 17886, M.C.Z.) and the two teeth
retained in the alveoli of the symphyseal fragment (no. 15751, M.C.Z.)
may not exceed the limits of sex and age variation, no accurate com-
parisons can be made since the crowns and the necks of the roots of
the two symphyseal teeth are destroyed.

CETOTHERIIDAE

? Mesocetus, species indet.
Plate 4

Referred specimens. The right and left bullae, no. 17885, Vertebrate
Paleontology Catalogue, Museum of Comparative Zoology. Collector,
George C. Elmore, 1941.

Horizon and locality. Laminated blue clays immediately below the

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 455

pebble phosphate, which are tentatively referred to the Hawthorn
formation, in pits of the American Agricultural Chemical Company at
Pierce, Polk County, Florida.

Description. The right and left bullae (no. 17885, M.C.Z.) are fairly
complete, but are not associated with other cranial elements. On both
of these bullae the very thin anterior pedicle and the accessory ossicle
borne by it, as well as most of the posterior pedicle are missing.

These tympanic bullae have a broad furrow, which commences on
the posterior face at the base of the posterior pedicle (pi. 4, fig. 3) and
curves downward and then forward on the ventral face. This ventral
furrow, or longitudinal depression, creates the broad indentation on
the posterior border, as seen from below, and is narrowed anteriorly by
the development of a prominent oblique keel or crest (pi. 4, figs. 2, 5),
which originates at the anterior end and extends backward to about
the level of the sigmoid process. The longitudinal furrow which divides
the ventral face of the bulla into two lobes, an external (lateral) and
an internal (mesial) one, has been considered by some to constitute a
diagnostic feature of Recent Odontoceti. It is nevertheless true that
this modification is more accentuated in the Odontoceti. The external
(lateral) face, as seen from below, curves convexly from end to end,
whereas the nearly straight internal (mesial) face is slightly indented
near the middle of its length; the postero-external and postero-internal
angles (pi. 4, fig. 2) are rounded.

The greatly thickened and inwardly reflected inner (mesial) lip, or
involucrum (pi. 4, fig. 4), rather abruptly decreases in width anterior
to the level of the sigmoid process. The dorsal surface of the involu-
crum exhibits an undulating curvature, interrupted posteriorly by
rugosities and anteriorly by transverse creases. At the interior (eusta-
chian) end of the bulla, the dorsal face of the involcrum merges im-
perceptibly with the curved outer lip, but is not depressed to form a
cleft for the passage of the eustachian tube. The anterior end of the
bulla of all known odontocetes is deeply scooped out, forming a spout-
like cleft for the passage of the eustachian tube. Lillie (1910, pp. 779-
7S0) and Hanke (1914, pp. 507, 509) found no trace of this cleft in the
bullae of mysticetes. By dissection they have shown that the eustachian
tube opens into the floor of the air sinus enclosed in the pterygoid
fossa, which in turn communicates with the anterior end of the tym-
panic cavity of the bulla. This arrangement characterizes all the
mysticetes, fossil or Recent, that have been studied.

There is a deep groove on the outer (lateral) lip of the bulla between
the sigmoid process and the posterior conical apophysis. The posterior

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process (pi. 4, fig. 4) is a thin plate of bone of irregular curvature at
the base, bi-concave in front and concave behind, which projects from
the involucrum. The sigmoid process is located on the outer (lateral)
lip near the middle of the length of the bulla and the rounded distal
end of this process is twisted at right angles to the long axis of the
bulla. The elongated scar on the basal internal (mesial) border of the
sigmoid process and the presence of a small fractured area on the outer
(lateral) lip between the sigmoid process and the anterior process in-
dicates that the slender anterior process of the malleus was as rigidly
fixed as in Recent mysticetes.

Measurements of the Bullae (in millimeters)

No. 17884,
M.C.Z.

No. 17885,
M.C.Z.

Right

Left

Right

Left

Greatest antero-posterior diameter

74
41

48
73.7

73.4
41.5

47

73.5
22

57.5
37

40.5

35.5
12.5
57.5

15.4

59.5

Greatest transverse diameter of bulla (but not
including the sigmoid process)

39

Greatest transverse diameter of bulla, inner face
(opposite basioccipital) to external swelling
above sigmoid process

41.3

Greatest dorso-ventral diameter on external
face, ventral face to tip of sigmoid process
(with one arm of calipers resting on hinder
end of involucrum and end of sigmoid process
and the other arm on the ventral face of the
bulla)

34.5

Transverse diameter of sigmoid process

Greatest length of involucrum

12.8
59

Greatest distance between outer lip of bulla and
opposite (inside) face of involucrum

16

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 457

Remarks. The general conformation of these bullae suggested com-
parison with Mesocetus hungaricus (Kadic, 1907, p. 33, fig. 3), although
the antero-posterior diameter of the bulla of the latter is 70 mm.
Unfortunately, only the left bulla of M . hungaricus was found and
most of the outer (lateral) lip is destroyed. This left bulla has a longi-
tudinal furrow on the ventral face and an involucrum of approximately
the same shape and appearance; the postero-internal (mesial) angle
seems to protrude less conspicuously. The contour of the bulla of the
genotype Mesocetus longirostris (Beneden, 1886, pi. 35, figs. 2-12), as
seen from below, does not match that of these Florida bullae very
closely; the former does have a similar longitudinal furrow on the
ventral face, but on the other hand the postero-internal (mesial) angle
does not protrude to the same extent and the posterior border is
rounded and not indented medially. There is a strong possibility that
these Florida bullae may belong to one of the previously described
Miocene cetotheres, but no definite allocation will be made until more
adequate material is available for study.

? Isocetus, species indet.
Plate 5

Referred specimens. The right and left bullae, no. 17884, Vertebrate
Paleontology Catalogue, Museum of Comparative Zoology. Collector,
George C. Elmore, 1941.

Horizon and locality. Presumably from the laminated blue clays im-
mediately below the pebble phosphate, which are tentatively referred
to the Hawthorn formation, in pits of the American Agricultural
Chemical Company at Pierce, Polk County, Florida. Middle Miocene.

Description. The right and left bullae (no. 17884, M.C.Z.) are in-
complete and fractured in several places. On both of these bullae, the
sigmoid process and adjoining portion of the overarching outer (lateral)
lip, as well as the thin anterior process and the posterior process are
destroyed. The posterior conical apophysis also is destroyed on the
right bulla.

The posterior face of the left bulla (pi. 5, fig. 6) is characterized by
the deep and more or less vertical groove which terminates in the notch
between the posterior conical apophysis and the base of the thin pos-
terior process. This groove, if present, was reduced in length on the

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right bulla. The posterior face is strongly convex, the apex of this
curvature merging imperceptibly into the low longitudinal crest on the
ventral surface. This longitudinal crest divides the ventral surface of
the bulla into two sloping surfaces, an external (lateral) and an internal
(mesial) one. When viewed from below (pi. 5, fig. 5), the external
(lateral) face, lateral to the longitudinal ventral crest, exhibits an
almost three-sided contour, whereas the internal (mesial) face is some-
what depressed near the middle of its length. From a ventral view,
there are no postero-external and postero-internal angles; the posterior
end is strongly convex and the anterior end is obliquely truncated.

The greatly thickened and inwardly reflected internal (mesial) lip,
or involucrum (pi. 5, fig. 1), rather abruptly decreases in width at
about the level of the hinder edge of the anterior process. The dorsal
surface of the involucrum exhibits a sub-concave curvature from end
to end, interrupted by transverse creases. At the anterior (eustachian)
end of the bulla, the dorsal face of the involucrum merges imperceptibly
with the curvature of the thickened outer (lateral) lip, and although
slightly depressed it does not form a cleft for the passage of the eusta-
chian tube.

Judging from the curvature of the outer (lateral) lip of the left bulla
(pi. 5, fig. 4), there seems to have been a deep groove between the
greatly elongated posterior conical apophysis and the sigmoid process
which is destroyed. The laterally flattened and triangular posterior
conical apophysis projects at least 8 mm. beyond the level of the base
of the posterior process. The posterior process (pi. 5, fig. 1) is a thin
plate of bone, almost straight at the base, which projects from the in-
volucrum. The anterior process seems not to have been very broad
antero-posteriorly.

Measurements: (see table p. 456).

Remarks. Compared with the bullae of other described North
American cetotheres, the elongation of the triangular posterior conical
apophysis is most unusual. It is assumed that the right and left bullae
belonged to an immature individual since the ventral surface is but
slightly roughened and the median longitudinal crest is not fully de-
veloped. This assumption is corroborated to some extent by compari-
son with a right and a left bulla (no. 5499, U.S.N.M.), unquestionably
belonging to different individuals, which were found in the phosphate
deposits at Tigerbay, Polk County, Florida. Most of the outer (lateral)
lip and a portion of the anterior end of the right bulla are broken off,
but in the present condition it measures 81.8 mm. in length; the dis-
tance from the ventral face of the bulla to the dorsal face of the involu-

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 459

crum is 42.5 mm. [The corresponding measurement of the right bulla
from Pierce is 38.3 mm.] This right bulla has a strongly roughened
ventral surface, a high median longitudinal crest on the ventral face
which continues upward on the center of the posterior face almost to
the interval between the posterior conical apophysis and the base of
the posterior process, and deep transverse creases on the dorsal face
of the involucrum. On the internal (mesial) side, the surface of the
involucrum tends to fold over the ventral surface of the bulla. There is
a rather close agreement between the general conformation of this
right bulla and that of Isoectus depavwii (Beneden, 1886, pi. 71, figs.
3-8) from the upper Miocene Anversian stage of the Antwerp basin,
Belgium. Notwithstanding the close similarity in most details to the
Tigerbay bulla, it is well to call attention to the presence on the pos-
terior face of the left bulla of Isocetus depauicii (Beneden, 1886, pi. 71,
fig. 8) of two well developed ridges which converge near the base of
the posterior process. These ridges are not so well developed on the
right bulla from Tigerbay. The left bulla from Tigerbay is incomplete,
but is relatively smooth and approximately the same size and shape
as the bullae from Pierce. The differences observed are sufficient to
establish specific distinctness, but in the absence of corroboratory
cranial material, all four of these bullae are tentatively referred to
Isocetus.

BALAENOPTERIDAE

Balaenoptera flortdana, new species

Plate 6

Type. An essentially complete right mandible, somewhat crushed
in median region, no. 17882, Vertebrate Paleontology Catalogue,
Museum of Comparative Zoology. Collector, George C. Elmore, 1941.

Horizon and locality. Pebble phosphate, referred to the Bone Valley
formation, in pits of the American Agricultural Chemical Company at
Pierce, Polk County, Florida. Lower Pliocene.

Description. Horizontal ramus of right mandible bowed outward;
internal surface of horizontal ramus distinctly flattened, external sur-
face strongly convex; relatively short symphyseal region similar in
most respects to corresponding portions of mandibles of finbacks and
sei whales, and not roughened or pitted for insertion of connecting

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■

ligaments ; a well defined internal ledge above inferior groove extending
backward for some 400 mm. on distal end of mandible; mandible con-
stricted dorso-ventrally behind distal expansion and also in front of
coronoid process, the maximum dorso-ventral diameter of ramus being
near middle of its length (ramus of both finback and sei whale tapers

Fig. 5. Balaenoptera floridana, cross section right mandible, 100 mm. behind
anterior end, no. 17882, M.C.Z.

gradually in depth from in front of coronoid process to tip) ; cross sec-
tion of mandible 100 mm. behind anterior end (fig. 5) distinctly flat-
tened from side to side, whereas dorsal half is twice width of ventral
half in same section of an immature finback mandible (no. 16039,
U.S.N.M.); cross section 300 mm. behind anterior end (fig. 6) is ovate-
pyriform in outline in contrast to ovate outline of cross section taken
at same distance from tip of immature finback mandible ; cross section
1600 mm. behind anterior end (fig. 8) rounded ovate in contrast to
marked internal flattening of section taken in corresponding portion
of immature finback mandible; internal series of small foramina,
located in narrow longitudinal groove on alveolar edge of mandible,
for most part obliterated by erosion (internal row of foramina quite

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 461

Fig. 6. Balaenoptera floridana, cross section of right mandible, 300 mm. be-
hind anterior end, no. 17882, M.C.Z.

Fig. 7. Balaenoptera floridana, cross section of right mandible, 1000 mm.
behind anterior endjprobably some distortion from crushing, no. 17882, M.C.Z.

462 bulletin: museum of comparative zoology

conspicuous on immature and adult finback mandibles) ; mental fora-
mina on external surface concealed by reconstruction of crushed areas
(external mental foramina relatively large and located at varying in-
tervals on anterior three fourths of finback and sei whale mandibles) ;
small triangular coronoid process bent outward toward apex (coronoid
process of finback mandible distinctly elongated) ; a small subsidiary

Fig. 8. Balaenoptera floridana, cross section of right mandible, 1600 mm.
behind anterior end, no. 17882, M.C.Z.

process behind and below base of coronoid process, likewise bent out-
ward, representing anterior termination of elongated protuberance on
dorso-internal side of ramus, a distinctive characteristic of balae-
nopterine whales; maximum dorso-ventral diameter of elongated
dorso-internal protuberance 42 mm. and distance from anterior rim of
large internal dental foramen to apex of subsidiary process 115 mm.;
area in front and above large internal dental foramen more like condi-
tion exhibited by mandibles of immature finbacks than of sei whales
(no subsidiary process at anterior termination of dorso-ventrally elon-
gated protuberance on either finback or sei whale mandibles); hori-
zontal distance from hinder face of condyle to apex of coronoid process
slightly more than 18 percent of greatest length of mandibles of B.

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 463

floridana and immature finback, hut distance from anterior rim of
large internal dental foramen to apex of coronoid process relatively
greater in B. floridana; condyle (fig. 10) abruptly narrowed below level
of deep furrow on external side (posterior aspect of condyle of B. flori-

Fig. 9. Balaenoptera floridana, cross section through coronoid process, 1860
mm. behind anterior end, no. 17882, M.C.Z.

dana more like that of a young 26 foot sei whale (no. 239307, U.S.N.M.),
except that the latter possesses a well defined groove on internal face
and only a slight notch on external face) ; dorsal half of condyle trun-
cated obliquely in dorso-ventral direction and convexedly curved from
external to internal margin ; ventral half depressed medially in oblique
direction and truncated more or less at right angles to long axis of
ramus ; condyle also strongly compressed from side to side in contrast
to pronounced side to side widening of dorsal half of condyle on man-

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dibles of finback and sei whales [ratio of width of upper half of condyle
to lower half 185 to 110 in case of immature finback and 230 to 200
in case of adult sei whale; groove present on internal and external
faces of condyles of finback and sei whale mandibles ; external and in-

Fig. 10. Balaenoptera floridana, posterior view of condyle, no. 17882, M.C.Z.

ternal grooves on condyle of immature finback (no. 16039, U.S.NJM.)
connected by deep transverse furrow, dividing condyle into a large
elongated dorsal articular surface and a relatively small ventral sur-
face; transverse groove on condyle of adult finback (no. 237566,
U.S.N.M.) somewhat shallower than on immature mandibles].

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 465

Measurements (in millimeters) :

Greatest length of mandible in a straight line 2123

Greatest length of mandible along outside curvature 2256

Distance from anterior end to level of center of coronoid process

in a straight line 1 750

Greatest vertical diameter 100 mm. behind anterior end of ramus 127
Greatest transverse diameter 100 mm. behind anterior end of

ramus 47.8

Greatest vertical diameter 300 mm. behind anterior end of ramus 108.5
Greatest transverse diameter 300 mm. behind anterior end of

ramus 57.8

Greatest vertical diameter 1000 mm. behind anterior end of ramus 150 =*=
Greatest transverse diameter 1000 mm. behind anterior end of

ramus 74 =*=

Greatest vertical diameter 1600 mm. behind anterior end of ramus 117.5
Greatest transverse diameter 1600 mm. behind anterior end of

ramus 84

Greatest vertical diameter of ramus through coronoid process 163

Greatest transverse diameter of ramus at level of coronoid process 77
Least vertical diameter of ramus between coronoid process and

condyle 117

Horizontal distance between center of coronoid process and hinder

face of condyle 392

Distance from hinder face of condyle to anteriormost free rim of

entrance to dental canal 227

Greatest dorso-ventral diameter of condyle 1 79

Greatest transverse diameter of condyle 87

Remarks. The right mandible (pi. 6) of this fossil balaenopterine
whale is somewhat shorter and slenderer than the mandibles of either
of the two immature finbacks in the U. S. National Museum, and
differs also in other details of conformation. If this mandible belonged
to an adult, this extinct whale was somewhat smaller than the Recent
finback, Balaenoptera physalus, and considerably larger than the little
piked whale, Balaenoptera acutorostrata. No marked alterations in the
general conformation of the mandibles of immature and adult balae-
nopterine whales have been observed and it is therefore difficult to
estimate the age of extinct mysticetes represented solely by mandibles.

The length of this fossil mandible in a straight line is 2m 123 mm.
The right mandible of a Recent immature finback (no. 16039,
U.S.N.M.) the length of whose skeleton is unknown, measures 2m
610 mm. in a straight line. The length in a straight line of a right
mandible belonging to a 47 ft. 7 in. finback skeleton (no. 16045,

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U.S.N.M.) is approximately 3 m. It should be noted also that the
right mandible of an old male sei whale, Balaenoptera borealis (no.
239307, U.S.N.M.), about 45 feet in length, measures in a straight
line 3m 290 mm.

The right mandible of Balaenoptera floridana differs in essential
structural details from the mandibles of most of the fossil mysticetes
(Strobel, 1881) described from Pliocene horizons in Italy, although it
does resemble in certain features the left mandible described and
figured by Portis (1885, pp. 44-46, pi. 4, figs. 42-47) under the name
Balaenoptera cortesii. This last mentioned mandible, measuring 2m
38 mm. in length, was collected by Gastaldi in 1874 in a middle Pliocene
(Astian) sand bank at Montafia, Piemonte, Italy. The type skeleton of
Balaenoptera cortessii (Fischer), measuring 4m 50 mm. in length, was
found, however, by Cortesi in 1816 in the lower Pliocene (Plaisancian)
sandy blue clay of a stream which descends from Montezago and
empties into the Chiavenna River, a tributary of the Po River,
Piemonte, Italy .The mandible belonging to the type skeleton measures
lm 150 mm. in length and possesses characters that distinguish it frcm
the Montafia mandible.

The mandible from Montafia has a low triangular coronoid process,
the condyle compressed from side to side and obliquely truncated
when viewed from the side, the distal end of ramus dorso-ventrally
expanded, the ramus constricted dorso-ventrally behind distal expan-
sion and also in front of coronoid process, and the ledge above inferior
groove on internal surface of anterior end of ramus essentially similar
in position. Notwithstanding these points of resemblance, the Monta-
fia mandible seems to lack the subsidiary process behind and below
base of the coronoid process, the inner surface of anterior end of ramus
is roughened for insertion of connecting ligaments, the distal end is
noticeably widened from side to side, the posterior aspect of the con-
dyle is quite different in conformation, and the groove on the external
face of the condyle is connected by a transverse furrow with the corre-
sponding groove on the internal face of the condyle.

Among the skeletal remains of mysticetes found in the lower Pliocene
(Diestian) sands of the Antwerp Basin, Belgium, Beneden (1885, pt. 4)
has recognized several species, three of which, Plesiocetus brialmontii,
P. dubius, and P. hupschii, are represented by portions of mandibles.
Only one of these, P. dubius, exhibits a close resemblance to the Florida
mysticete. Notwithstanding the close similarity between the size and
conformation of the anterior end of the mandible of P. dubius and the
corresponding portion of the mandible of B. floridana, the greater

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 467

width of the ramus and the position of the external mental foramina
readily distinguish the former from the latter.

From the middle Pliocene (Sealdisian) sands of the Antwerp Basin,
Belgium, Beneden (1882, pt. 3) has figured the whole or portions of
mandibles identified as Balaenoptera borealina, B. musculoides, B. ros-
tratella, Burtinopsis minutus, and B. similis. It should be noted that
Beneden (1882, pt. 3, p. 651) has stated that the cetacean, which he
(1859, p. 141) had dedicated to Van Gorp under the name of Plcsioeetus
garopii, was the same as Balaenoptera musculoides (Beneden, 1880,
p. 15). The mandibles of the fossil mysticetes allocated to the genus
Balaenoptera by Beneden are distinguishable from Balaenoptera flori-
dana by the conformation of the condyle, the dental foramen, and the
ramus, especially the anterior end. The mandible of B. floridana is
likewise readily separable from those of Burtinopsis minutus and B.
similis by the shape of the condyle.

Four names have been bestowed by Owen (1846, pp. 531-534) on
incomplete mysticete bullae found in the upper Pliocene Red Crag
nodule bed at Felixstow, Suffolk, England. Until comparable material
representing the Florida mysticete is collected, the relationships of
these British species to B. floridana can not be ascertained.

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LITERATURE CITED

Abel, O.

1899. Untersuchungen liber die fossilen Platanistiden des Wiener

Beckens. Denkschr. math.-naturwiss. Kl. Kais. Akad. Wissensch.

Wien, 68, pp. 839-874, pis. 4.
1905. Les odontocetes du Bolderien (Miocene superieure) d'Anvers.

Mem. Mus Roy. d'Hist. Nat. Belgique, Bruxelles, 3, pp. 1-155,

figs. 27.
1909. Der Schadel von Saurodelphis argentinus aus dem Plioziin Argen-

tiniens. Sitz.-ber. kais. Akad. Wissensch. Wien, math.-naturw. Kl.
118, Abt. 1, pp. 255-272, pi. 1, fig. 1. March, 1909.

Allen, Glover M.

1921. Fossil cetaceans from the Florida phosphate beds. Jour. Mam-
malogy, 2, no. 3, pp. 144-159, pis. 9-12. August, 1921.
1941. A fossil river dolphin from Florida. Bull. Mus. Comp. Zool.
Harvard College, 89, no. 1, pp. 1-8, pis. 3. October, 1941.

Ameghino, Florentino.

1891a. Caracteres diagn6sticos de cincuenta especies nuevas de mamiferos
f6siles argentinos. Rev. Argentina Hist. Nat., Buenos Aires, 1,
entr. 3a, pp. 129-167, figs. 26-75. June 1, 1891.

1891b. Mamiferos y aves f6siles argentinos. — Especies nuevas, adiciones
y correcciones. Rev. Argentina Hist. Nat., Buenos Aires, 1, entr.
4a, pp. 240-259, fig. 1. August 1, 1891.

1898. Sinopsis Geol6gico-Paleontol6gica (Formaciones cenoz6icas y
creticeas). Segundo Censo de la Republica Argentina Mayo 10 de
1895, Buenos Aires, 1 (Territorio), pp. 113-255+ [l], figs. 104.

BENEDEN, P. J. VAN.

1859. Ossements fossiles decouverts a Saint-Nicholas, en 1859. Bull.

Acad. Roy Sci. Belgique, Bruxelles, ser. 2, 8, no. 11, pp. 123-148.
1880. Les Mysticetes a courts fanons des sables des environs d'Anvers.

Bull. Acad. Roy. Sci. Belgique, Bruxelles, ser. 2, 50, pp. 11-25.
1882. Description des ossements fossiles des environs d'Anvers. Pt. 3,

Cetaces. Genres: Megaptera, Balaenoptera, Burtinopsis et Er-

petocetus. Ann. Mus. Roy. Hist. Nat. Belgique, Bruxelles, ser.

Paleont., 7, pp. 1-90, pis. 40-109.

1885. Description des ossements fossiles des environs d'Anvers. Pt. 4,
Cetaces. Genre: PI esiocetus. Ann. Mus. Roy. Hist. Nat. Belgique,
Bruxelles, ser. Paleont., 9, pp. 1-40, pis. 1-30.

1886. Description des ossements fossiles des environs d'Anvers. Pt. 5,
Cetaces. Genres: Amphicetus, Heterocetus, Mesocetus, Idiocetus
et Isocetus. Ann. Mus. Roy. Hist. Nat. Belgique, Bruxelles, ser.
Paleont., 13, pp. 1-139, pis. 75.

KELLOGG: FOSSIL CETACEANS FROM THE FLORIDA TERTIARY 469
BENEDEN, P. J. VAN, AND P. GERVAIS.

1880, Osteographie des Cetaccs vivants et fossiles. Paris, Text, pp.
1-605; Atlas, folio pis. 1-64.

Brandt, J. F.

1873. Untersuchungen uber die Fossilen und Subfossilen Cetaceen
Europa's. Mem. Acad. Imp. Sci. St.-Petersbourg, s6r. 7, 20, no. 1
pp. viii + 372, pis. 34.

BURMEISTER, HERMANN.

1871. On Saurocetes argentinus, a new type of Zeuglodontidae. Ann.

and Mag. Nat. Hist., ser. 4, 7, no. 37, pp. 51-55, pi. 1. January,

1871.

1891a. La Prensa, Buenos Aires, June 26, 1891.

1891b. Nuevos objetos en el museo nacional. Anal. Soc. Cient. Argentina,
Buenos Aires, 32, entr. 4, pp. 161-163. October, 1891.

1892. Continuacion a las adiciones al examen crltico de los mamiferos
f6siles terciarios. Anal. Mus. Nac. Buenos Aires, 3, entr. 18, pp.
viii + 401-488, pis. 7-10.

Cabrera, Angel.

1926. Cetaceos f6siles del Museo de la Plata. Rev. Mus. La Plata, 29,
pp. 363-411, figs. 19.

Cooke, C. W., and S. Mossom.

1929. Geology of Florida. 20th Ann. Rept. Florida State Geol. Surv.,
pp. 29-227.

Cuvier, Georges.

1825. Recherches sur les ossements fossiles, Paris, 3rd ed., 5, pp. 1-405,
pis. 27.

Fischer, Joanne Baptista.

1829. Synopsis Mammalium, Stuttgart, pp. xlii + 752.

Gervais, Paul.

1859. Zoologie et Paleontologie francaises, Paris, ed. 2, pp. viii + 544;
atlas, pp. xii, pis. 84.

Hanke, Herbert.

1914. Ein Beitrag zur Kenntnis der Anatomie des ausseren und mittleren
Ohres der Bartenwale. Jenaischen Zeitschr. f. Naturwiss., 51, no.
3, pp. 487-524, pis. 3.

Holl, Friedrich.

1829. Handbuch der Petrefactenkunde, Dresden, Erstes Bandchen, pp.
viii + 115. (Allgemeine Taschenbibliothek der Naturwissen-
schaften, Theil 9.)

470 bulletin: museum of comparative zoology

Kadic, Ottokar.

1907. Mesocetus hungaricus Kadic, eine neue Balaenopteridenart aus
dem Miozan von Borbolya in Ungarn. Mitteil. Jahrb. Kgl. Un-
garischen Geol. Anstalt, Budapest, 16, no. 2, pp. 23-91, figs. 70,
pis. 3.

Kellogg, Remington.

1925. Additions to the Tertiary history of the pelagic mammals on the
Pacific Coast of North America. Carnegie Instn. Washington
Publ. 348, pp. iii + 120, pis. 13. April 22, 1925.

LlLLIE, D. G.

1910. Observations on the anatomy and general biology of some mem-
bers of the larger Cetacea. Proc. Zool. Soc. London for 1910, pt.
3, pp. 769-792, figs. 69-78, pi. 74. October 11, 1910.

Owen, Richard.

1846. A history of British fossil mammals and birds, London, pp. xlvi +
560, figs. 236.

PORTIS, ALESSANDRO.

1885. Catalogo descrittivo dei Talassoterii rinvenuti nei Terreni Ter-
ziarii del Piemonte e della Liguria. Mem. R. Accad. Sci. Torino,
ser. 2, 37, pp. 247-365, pis. 9.

ROVERETO, CAYETANO.

1915 Nuevas investigaciones sobre los delfines longirostros del Mioceno
del Parang (Republica Argentina). Ann. Mus. Nac. Buenos Aires,
27, pp. 139-151, pis. 2-4. September 9, 1915.

Simpson, G. G.

1930. Tertiary land mammals of Florida. Bull. Amer. Mus. Nat. Hist.,

69, pp. 149-211. June 5, 1930.
1932. Fossil Sirenia of Florida and the evolution of the Sirenia. Bull.

Amer. Mus. Nat. Hist., 59, pp. 419-503, figs. 23. September 6,

1932.

Strobel, P.

1881. Iconografia comparata delle ossa fossili del Gabinetto di Storia
Naturale dell' Universita di Parma. Fasc. 1, Balenotteride
(Cetoterio e Megattera), Parma, pp. 1-32, pis. 5.

True, Frederick W.

1909. A new genus of fossil cetaceans from Santa Cruz Territory, Pata-
gonia; and description of a mandible and vertebrae of Prosqualo-
don. Smithson. Misc. Coll., 62, pt. 4, pp. 441-456, pis. 43-45.
August 7, 1907.

KELLOGG: FOSSIL CETACEANS FROM Till-: FLORIDA TERTIARY \< !

1910. Description of a skull and some vertebrae of the fossil cetacean
Diochotichus vanbenedeni from Santa Cruz, Patagonia. Bull.
Amer. Mus. Nat. Hist., 28, art. 4, pp. 19-32, pis. 1-5. March 22,

1910.

1912. A fossi] toothed cetacean from California, representing a new genus
and species. Smithson. Misc. Coll., 60, no. 11, pp. 1-7, pis. 2.
November 1, 1912.

Valenciennes, A.

1862. Sur une machoire inferieure de Dauphin fossile envoyee par M.
Thore, de Dax (departement des Landes). Comptes Rendus Acad
Sci., Paris, 54, no. 14, pp. 788-790.

White, T. E.

1942. Additions to the fauna of the Florida phosphates. Proc. New Eng-
land Zool. Club., 21, pp. 87-91. November 14, 1942.

Wilson, Leslie E.

1935. Miocene marine mammals from the Bakersfield Region, Cali-
fornia. Peabod}' Mus. Nat. Hist., New Haven, Bull. 4, pp. 143,
figs. 23.

PLATES

PLATE 1

Kellogg — Fossil Cetaceans from Florida Tertiary

PLATE 1

Fig. 1. Goniodelphis hudsoni, dorsal view of symphyseal region of mandibles,
no. 17881, M.C.Z.

Fig. 2. Goniodelphis hudsoni, lateral view of symphyseal region of right
mandible, no. 17881, M.C.Z.

BULL. MUS COMP. ZOOL.

Kellogg. Fossil Cetaceans. Plate 1.

PLATE 2

Kellogg — Fossil Cetaceans from Florida Tertiary

PLATE 2

Fig. 1. Goniodelphis hudsoni, external view of right mandible, no. 17879,
M.C.Z.

Fig. 2. Megalodelphis magnidens, external view of left mandible, no. 17883,
M.C.Z.

BULL MUS. COMP ZOOL.

Kellogg. Fossil Cetaceans. Plate 2.

''' -

I

I i

■

-V

- •

PLATE 3

Kellogg — Fossil Cetaceans from Florida Tertiary

PLATE 3

Fig. 1. Megalodelphis magnidens, ventral view of rostral fragment, no.
17880, M.C.Z.

• Fig. 2. Megalodelphis magnidens, dorsal view of hinder end of symphysis
and of left mandible, no. 17883, M.C.Z.

BULL MUS COMP ZOOL.

Kelloss. Fossil Cetaceans. Plate 3.

PLATE 4

Kellogg — Fossil Cetaceans from Florida Tertiary

PLATE 4

Fig. 1. ? Mesocetus, species indet., external (lateral) view of right bulla,
no. 17885, M.C.Z.

Fig. 2. ? Mesocetus, species indet., ventral view of right bulla, no. 17885,
M.C.Z.

Fig. 3. ? Mesocetus, species indet., posterior view of right bulla, no. 17885,
M.C.Z.

Fig. 4. ? Mesocetus, species indet., dorsal view of left bulla, no. 17885.
M.C.Z.

Fig. 5. ? Mesocetus, species indet., ventral view of left bulla, no. 17885,
M.C.Z.

Fig. 6. ? Mesocetus, species indet., anterior view of left bulla, no. 17885,
M.C.Z.

BULL MUS. COMP ZOOL.

Kello:;g. Fossil Citaceans. Plate 4.

^*.-*-*-

fLATI 5

Kellogg — Fossil Cetaceans from Florida Tertiary

PLATE 5

Fig. 1. ? Isocetus, species indet., dorsal view of right bulla, no. 17884,
M.C.Z.

Fig. 2. ? Isocetus, species indet., ventro-external view of right bulla, no.
17884, M.C.Z.

Fig. 3. ? Isocetus, species indet., anterior view of right bulla, no. 17884,
M.C.Z.

Fig. 4. ? Isocetus, species indet., external (lateral) view of left bulla, no.
17884, M.C.Z.

Fig. 5. ? Isocetus, species indet., ventral view of left bulla, no. 17884, M.C.Z.

Fig. 6. ? Isocetus, species indet., posterior view of left bulla, no. 17884,
M.C.Z.

BULL MUS. COMP ZOOL.

Kellogg. Fossil Cetaceans. Plate 5.

?

PLATE 6

Kellogg — Fossil Cetaceans from Florida Tertiary

PLATE 6

Fig. 1. Balaenoptera floridana, internal view of right mandible, no. 17882,
M.C.Z.

Fig. 2. Balaenoptera floridana, dorsal view of right mandible, no. 17882,
M.C.Z.

BULL. MUS. COMP. ZOOL.

Kellosg. Fossil Cetaceans. Plate 6.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 10

LATE PALEOZOIC XIPHOSURANS

By Percy E. Raymond

With Two Pl\tes

_ 5he Library <

H&lseaB ©f Comparative Zc.
^"S Barnard UMversItv

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

November, 1944

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After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 10

LATE PALEOZOIC XIPHOSURANS

By Percy E. Raymond

With Two Plates

of Compa;
Harvard

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

November, 1944

No. 8— Late Paleozoic Xiphosuratis
By Percy E. Raymond

The Xiphosura from the Carbondale series (mid-Pennsylvanian) of
Mazon Creek have long been known, but never critically studied.
The collections recently made by Mr. Frederick Thompson contain
several unusually good specimens presenting new points of interest,
so the time seems opportune to restudy the material from that region,
and to make comparisons with other related forms recently described.

The specimens from Mazon Creek and vicinity are in a peculiar
state of preservation. Like the plants, they are found in clay-ironstone
concretions in which they are comparatively little crushed. Such
distortion as is present is probably due largely to the thinness of the
test, which appears to have been much less strong than that of modern
limulids.

The specimens seem at one time to have been hollow molds, with a
space between the tergal and sternal tests. This space in some was
later partially rilled with pyrite; in others it contains a soft white
platy substance which Professor Cornelius S. Hurlbut, Jr., identified
for me as kaolin. Some specimens retain a thin coating of pyrite,
others have crystals of the same mineral imbedded in the kaolin, and
still others have nothing but kaolin. This substance can be removed
mechanically from such specimens as have a hard matrix, and good
molds of the dorsal and ventral surfaces secured.

I am indebted to Dr. Carl O. Dunbar for the loan of many specimens
from the Peabody Museum at Yale University, to Dr. Ray S. Bassler
for casts of types in the United States National Museum, and to Dr.
Christina Lochman for a specimen. For the photographs, thanks are
due to Prof. Frank M. Carpenter, who collected the Permian speci-
mens. The drawings were made by Dr. Robert R. Wheeler. Mr.
Frederick Thompson has generously presented many specimens to
the Museum of Comparative Zoology.

TERMINOLOGY

Dorsal Surface

Authors have used various terms for the parts of the test of xipho-
surans. There has been little usage of the same term with different
meanings, so there is little real confusion, but it may be well to explain
the terms used in this paper. I shall follow the usual custom of naming
parts so far as possible, by analogy with the trilobite.

The anterior shield is a cephalothorax, since it has six pairs of

476 bulletin: museum of comparative zoology

appendages. The term prosoma, now in general use, applies well to it.
The terminal segment of the body is so generally known as the telson
that no better name need be sought. But the median portion presents
problems.

In the primitive xiphosurans of the Silurian and early Devonian,
this part of the body consists of free segments. One is tempted to
call it a thorax, but this can not be done, for the last segment bears
the anal opening, showing that the telson is not homologous with the
pygidium of a trilobite. In cases where all the segments are free it
seems best to follow the general usage and employ the non-commital
term trunk to this portion of the body.

Modern xiphosurans have a single median shield. The anterior
portion bears six pairs of appendages, the posterior part none. The
anterior section is called the mesosoma, the posterior the metasoma.
This would be an excellent usage, were it not for the fact that it is
impossible to correlate the parts with those of Carboniferous members
of the group. The consolidation of the trunk segments into a single
shield appears to have begun at the posterior end. Some species of
Belinurus seem to have five, others seven, or even so few as four free
segments in front of a fused portion next to the telson. Hence the
obvious subdivisions of the trunk do not correspond exactly to the
meso- and metasoma. The application of these terms to fossils is
involved with theories. Moreover, a single word is needed to designate
the single median shield of the later members of the group. Packard
called it the urosome, an unfortunate term, for it is not a tail. Neither
is it an abdomen in any sense of the word, although often so desig-
nated. Dix and Pringle have lately revived Sir Richard Owen's term
thoracetron, a usage which will be followed here. They applied the
same name to the trunk of Belinurus, but it seems better to restrict
the term to shields which are completely consolidated. This leaves us
with no satisfactory terms for the parts of the trunk of those forms in
which fusion was in progress. Perhaps promesosoma and prometasoma
will be satisfactory.

A distinctive feature of the dorsal surface is the presence of a pair
of longitudinal furrows, limiting a relatively narrow median lobe on
both shields. These suggest comparison with the dorsal furrows of
trilobites and the inference that the median lobes are homologous in
the two sorts of animals. This is almost certainly true of the anterior
part of the thoracetron, for entopophyses for the attachment of
muscles project downward beneath them, just as do the appendifers
of trilobites. Whether the median lobe of the prosoma is homologous

RAYMOND: LATE PALEOZOIC XIPHOSURANS 477

with the trilobitan glabella is still debatable. Packard called the
median lobe of the thoracetron the cardiac lobe; Dunbar has used the
same term for its prolongation on the prosoma, and it seems logical
to adopt the term for the entire median lobe.

Fully as conspicuous as the dorsal furrows on the prosoma are the
longitudinal ridges behind the eyes. In Limulus they extend only a
short distance in front of the eyes, but in many of the Paleozoic forms
they converge forward, meeting on the median line. These are the
opthalmic ridges; the area enclosed within them, including the cardiac
lobe, was called the cardio-opthalmic region by Packard, the glabella
by H. Woodward, and the cranidium by Willard and Jones. The first of
these is preferable, since it is non-commital as to homology. For con-
venience it may be shortened to cardiopthalmic. Dunbar subdivided
this region into the median cardiac lobe and the lateral "glabellar"
lobes, but it would seem better to avoid implications as to homology
by applying some such term as intra-opthalmic to the areas between
the dorsal furrows and the opthalmic ridges.

These intra-opthalmic areas, in Limulus, have obscure transverse
depressions which alternate with thickened areas in the test. The
transverse furrows on the surface find expression on the inside of the
shell in curved ridges which bound the margins of muscular areas.
The muscles attached underneath the convex areas between the
depressions are those which extend either vertically or diagonally
downward to the borders or outer ends of the coxal segments of the
second to sixth legs on the prosoma. Most of the scars are adjacent
to the outer side of the ridge beneath each of the longitudinal furrows
bounding the cardiac lobe. The transverse depressions are inter-
muscular, and probably correspond in origin to the glabellar furrows
of a trilobite. In the latter animal, however, the furrows extend
mesially from the dorsal furrows in all except specialized members of
the group (e.g., Acidaspidae, Lichadidae). Hence it seems doubtful if
the whole cardiopthalmic area is homologous with the true glabella of
the trilobite.

Ventral Surface

The ventral membrane of Limulus is sufficiently chitinized to retain
a definite shape. The marginal portion, surrounded by a thickened
edge, is approximately horizontal. This portion corresponds with the
doublure of a trilobite, and will herein be mentioned by that name. It
is bounded posteriorly by ridges which curve backward abruptly near
the front and meet in a mesial spine. The roughly triangular region

478 bulletin: museum of comparative zoology

(sub frontal area) thus set off is beneath the stomach. From the
thickened ridges the ventral membrane rises in broad vaults, approxi-
mately parallel in convexity to the dorsal shield. The membrane is
not fully chitinized toward the summit of the vault, hence has con-
siderable flexibility. The uppermost edges are escaloped, with thick-
ened buttresses which unite to form fulcra (coxal attachments) for
movable articulation of the five appendages behind the chelicerae.

The median portion of the ventral surface is occupied by the
appendages. As seen from the inside, these appear as five pairs of
transversely elongated openings, each bounded by a thickened frame-
work somewhat complicated at the outer (proximal) ends. These
openings allow muscles from above to enter the basal segments of the
legs throughout their entire extent. The transverse areas between the
frames of the appendages are covered by membrane which is thin but
somewhat chitinous. Apparently the basal segments of the legs have
but little motion in any direction.

Chelicerae are attached near the median line back of the proximal
ends of the second, and about on a line with those of the third append-
ages. Two long thickenings, incurved at the posterior end, support
them. These might leave impressions on casts of the interior in the
fossil state. Near their anterior ends, on the median line, is a small
subcircular thickening, the subfrontal scleritc. Between the mesial ends
of the coxae of second pair of appendages, and just in front of the
mouth is the camerosome, a narrow, highly convex, keeled plate. The
bases of the chelicerae embrace its anterior moiety. Behind the
mouth is another plate, narrow in front and wide behind, the promeso-
stemite. This and the camerosome may represent the metastoma and
hypostoma of the trilobite.

Class ARACHNIDA
Subclass MEROSTOMATA Woodward

Order XIPHOSURA Gronovius

Suborder SYNXIPHOSURA Packard

Xiphosura with all the segments of the trunk freely movable.

This suborder contains many genera about which little is known.
The only ones which seem to be close relatives of the later limuloids
are Neolimulus Woodward, from the Upper Silurian, and Weinbergina
R. and E. Richter, from the Lower Devonian.

RAYMOND: LATE PALEOZOIC XIPHOSURANS 479

Suborder LIMULADA R. and E. Richter

Xiphosura with some or all of the trunk segments anchylosed.
Prosoma with opthalmic ridges, at least at the posterior margin.

This is only a part of the original definition of the suborder1, but it
seems inadvisable, in our present state of ignorance, to include state-
ments about the appendages or other morphological features as yet
unknown. For example, the Richters included as the first character-
istic in their diagnosis "Prosoma mit verlangerten Hinterecken,"
which is not true of any member of a new family to be described in
this paper.

The second sentence of the present definition is introduced in view
of the fact that anchylosis of segments is a common feature of various
lines of arthropods, and it is not unlikely that it took place in various
groups of the Synxiphosura.

Superfamily BELINURACEA nov.

Limulada with some of the anterior segments of the trunk movable,
two or more at the posterior end anchylosed.

Eller has recently reviewed the members of the genus Belinurus
and assembled figures of most of the described species.2 Without
actual material it would be unsafe to make a revision of the group, but
it is obvious that more than one genus is involved. Judging from the
terminal portion of the axial lobe of the thoracetron, Belinurus
metschetnensis and B. iswarinensis Tchernechev are almost certainly
species of Euproops, and one suspects that B. stepanovi of the same
author is another. Tchernechev may have been using Packard's
totally erroneous definition of Belinurus. It would be difficult for
anyone to write a paper with more mistakes to the page and plate
than that of Packard,3 and unfortunately it is so beautifully illustrated
that it is bound to cause confusion for years to come.

Turning to the other species of Belinurus, the figures seem to indicate
that the genus contains prototypes of the two sorts of prosomae
present in the Euproopacea and the Limulacea. Most of the species
have the posterior branch of the opthalmic ridge practically parallel
to the axis of the prosoma, extending straight forward to the eye.

1 Senckenbergiana., 2, 1929, p. 206.

2 Ann. Carnegie Mus., 27, 1938, pp. 129-150, pis. 9-14.

3 Mem. Nat. Acad. Sci., 3, 1886.

480 bulletin: museum of comparative zoology

This group includes Belinurus reginae Baily, B. arcuatus Baily, B.
concinnus Dix and Pringle, B. grandaevus Jones and Woodward, B.
alleghanyensis Eller, and some specimens which have been referred
to the type of the genus, B. bellulus Koenig (Eller's pi. X, figs, 7, 8).
This is the type of opthalmic ridge characteristic of the Limulacea.

Other species, not so numerous, have the type of ridge characteris-
tic of the Euproopacea, that is, a curved opthalmic ridge bearing out-
ward to the eye. This is probably characteristic of the true Belinurus
for it is shown in Koenig's original figure of the genotype (Petrificata
Derbiensia, 1809) and in specimens figured later by other writers
(Eller's pi. X, figs, 3, 4, 6). It is present also in Belinurus truemani
Dix and Pringle, B. morgani Dix and Pringle, and B. -pustulosus Dix
and Pringle.

It is also important to note that the species with the parallel opthal-
mic ridges have, on the whole, triangular trunks, whereas those with
curved ones are of a much more rounded type. It seems probable that
the Devonian ancestor of the Limulacea was not unlike B. alleghani-
ensis and that the Devonian ancestor of the Euproopacea was more
like the Carboniferous B. truemani.

Since these two lines are so clearly marked, it may be helpful to
make a new genus for the forms with parallel opthalmic ridges.

Family BELINURIDAE PACKARD (restricted)
Diagnosis, for the present, the same as for the super family.

Genus Belinurus Koenig

Belinuridae with the posterior portions of the opthalmic ridges
curved and directed outward. Trunk rounded, ovoid to semicircular
in outline, with two or more anchylosed segments at the posterior end.
Genotype, Belinurus bellulus Koenig.

Genus koenigiella genus nov.

Belinuridae with the posterior portions of the opthalmic ridges
parallel. Trunk subtriangular in outline, with two or more of the
segments anchylosed at the posterior end. Genotype, Belinurus
reginae Baily.

Species definitely assigned to this genus are Koenigiella alleghani-
ensis (Eller), K. reginae (Baily), K. arcuata (Baily), and K. koenigi-
ana (Woodward). The others mentioned above as being of this type

RAYMOND: LATE PALEOZOIC XIPHOSURANS 481

probably belong here, but in the absence of actual material it would
be unsafe to make a definite decision. I would however, venture to
predict that Prestwichia randalli Beecher from the Upper Devonian
will prove to belong to this family and possibly to this genus when the
trunk is found.

Superfamily EUPROOPACEA nov.

Limulada with broad, rounded thoracetron, with or without lateral
spines; posterior portions of opthalmic ridges, if present, turn outward
to the eyes.

This superfamily is proposed to include a branch of the Limulada
which seems not to have survived the Paleozoic. The thoracetron
approximates the shape of a circle, truncated where connected to the
prosoma. In a specialized family described below, (Liomesaspidae)
the lateral spines are lost, along with more or less of the longitudinal
and transverse furrows, producing relatively smooth forms. This is
the same sort of "smoothing out" that is so characteristic a feature of
various phyletic lineages of trilobites.

In the main family, Euproopidae, the opthalmic ridges have a
typical course, a curved posterior portion turning outward from the
posterior margin forward to the eye, then forward and inward, to join
the anterior end of the cardiac lobe. This course is that seen in many
species of belinurids, and differs from that in the Limulacea, in which
the posterior portion of the ridge extends straight forward or slightly
inward to reach the eye. This same course is characteristic of such of
the Liomesaspidae as retain the ridges, but the dorsal surface of the
prosoma in the Elleriidae and Prolimulidae is unknown.

A characteristic feature of the Euproopidae and Liomesaspidae,
but not the Elleriidae, is the nature of the posterior end of the axial
lobe of the thoracetron. In this region there are no transverse furrows,
but the surface rises into a high boss with a low conical spine at the
top. Behind the spine the surface drops abruptly in a concave slope
to a low, smooth posterior area. Some specimens show an impressed
line on the back slope of the boss, rising to an inverted A just behind
the spine.

There is no indication of articulated spines on the thoracetron of
any member of this superfamily, or any suggestion of the trapezoidal
shape of the thoracetron of the Limulacea. It seems to have been an
evolutionary line entirely different from that which led to modern
Limulus.

482 bulletin: museum of comparative zoology

Family EUPROOPIDAE Eller

Euproopacea with dorsal furrows on the prosoma and marginal
spines on the thoracetron.

Genus Prestwichianella H. Woodward

Limulus Prestwich, Trans. Geol. Soc, 2d ser., 1840, vol. 5, pi. 41, figs. 5, 6. —
Belinurus Baily, Ann. and Mag. Nat. Hist., 1863, ser. 3, vol. 11, p. 113. —
Prestwichia H. Woodward, Quart. Jour. Geol. Soc. London, 1867, vol. 23,
p. 32, pi. 1, fig. 2; Paleontological Soc, London, 1878, p. 244, pi. 31, fig. 5;
Geol. Magazine, 1868, vol. 5, p. 2. Meek and Worthen, Pal. Illinois, 1868,
vol. 3, p. 547, fig. B, p. 548. Packard, Nat. Acad, of Sci., Mem. 16, 1886,
p. 148, fig. 10— Dunbar, Am. Jour. Sci., 1923, ser. 3, vol. 5, p. 451 —
Prestwichinella, H. Woodward, Geol. Mag., 1918, ser. 6, vol. 5, p. 469.
Dix and Pringle, Summary of Progress Geol. Sur. Great Britain for 1928,
1929, p. 92, 101. Pruvost, Mem. du Mus6e Roy. d'Hist. Nat. Belgique,
No. 44, 1930, p. 200.

Euproopidae with a short, wide cardiopthalmic area, subdivided
into four parts. Genotype, Limulus rotwidatus Prestwich.

Euproops and Prestwichia were described in the same year, 1867,
and no satisfactory distinction has ever been drawn between them.
Meek, in his original description of Euproops said: "From Prestwichia
with which it more nearly agrees in general form as well as in its
anchylosed segments, it differs in having the area enclosed by the eye-
ridge (glabella) comparatively small, and of a quadrangular form,
with the eyes situated far forward on its anterior lateral angles." He
also called attention to the fact that the cardiopthalmic area of
Prestwichia was proportionately larger than in Euproops, and was
transversely elliptical rather than quadrangular in outline.

Henry Woodward long refused to accept Euproops as a distinct
genus, which is not surprising, for after his original definition of
Prestwichia he cited as typical species, first, Limulus anthrax Prest-
wich, and second, L. rotundatus Prestwich. Fifty-one years later, in
1918, he admitted that L. anthrax was a Euproops, hence it is evident
that the original definition included both genera. As the first species
mentioned under the generic name Prestwichia, P. anthrax might have
been selected as the genotype, in which case Euproops would have
become a synonym. Fortunately it was not, and when in 1918, Wood-
ward learned that the term Prestiwichia had been used before 1867,
he designated Prestwichia rotundata as the type of a genus under the
new designation Prestwichianella. On this occasion he stated of P.

RAYMOND: LATE PALEOZOIC XIPHOSURANS 483

rotundata: "the glabella [cardiopthalmic area] is divided along the
center by the axial furrow, and by two other slightly diverging parallel
lines on either of the axis, reaching nearly half-way to the frontal
border, where they are arcaded, forming a raised confluent line in
front of the glabella. The circular line seen outside the border of the
glabella may indicate the impression of the line of the broad incurved
undermargin of the head-shield." The first part of this statement is
incomprehensible to the writer, for he has seen no other xiphosuran
with a median furrow on the cardiac lobe. It is, however, quoted by
Dix and Pringle in 1929, apparently with approval, although the
new species which they describe is said to have a narrow raised median
ridge. According to Dix and Pringle, the eyes of Prestwichianella are
situated at the anterior lateral angles of the cardiopthalmic area, as
in Euproops. Woodward made no definite statement as to the position
of the eyes in P. rotundata except that they are on the raised lateral
border, nor are they shown in any figure.

Dix and Pringle were not able to find any further specimens which
they could positively identify as P. rotundata, hence it is necessary to
draw our conclusions as to the generic characteristics from the type.
The original specimen in the Prestwich collection has been properly
figured twice, once by that author, and again by Woodward (1878).
A diagramatic figure by Woodward (1867) has been widely copied, but
is incorrect in many particulars. Inaccurate as this figure is, it does
bring out what seems to be a unique characteristic of P. rotundata,
that is, that the cardiopthalmic area is quadra-, not tripartite. This
is in itself a sufficient generic characteristic, and may for the present
stand as the most important feature.

Pruvost identified P. rotundata in the Westphalian of northern
France and adjacent portions of Belgium, but his figure does not show
the critical area of the prosoma. He reported the presence of the
species not only in continental deposits, but also in marine beds
associated with Productus carbonarius.

484 bulletin: museum of comparative zoology

Genus EUPROOPS Meek

Bellinurus Meek and Worthen, Proc. Acad. Nat. Sci. Philadelphia, 1865, p. 44.
Geol. Survey Illinois, vol. 2, 1866, p. 395.

Prestwichia Meek, Am. Jour. Sci., ser. 2, vol. 43, 1867, p. 257, Packard, Nat.
Acad. Sci., vol. 3, mem. 16, 1886, p. 146, 148, 150. Bergeron, Bull. Soc.
Geol. France, ser. 3, vol. 21, 1893, p. 342; vol. 23, 1895, p. 480. Zalessky,
Bull. Comite Geol. de St. Petersbourg, vol. 26, 1907, p. 423. Bolton,
Trans. Manchester Geol. Soc, vol. 34, 1915. Pruvost, Mem. Carte Geol.
de France, 1919, p. 333. Tchernechev, Bull. Comite Geol. de Leningrad,
vol. 47, 1928, p. 526. Euproops Meek, Am. Jour. Sci., ser. 2, vol. 43, 1867,
p. 394; Geol. Mag. vol. 4, 1867, p. 320. H. Woodward, Geol. Mag. 1868,
vol. 5, p. 2. Meek and Worthen, Geol. Survey Illinois, vol. 3, 1868, p. 547,
White, Geol. and Nat. Hist. Indiana, 13th Ann. Rept. for 1883, (1884),
p. 170. Packard, Am. Naturalist, vol. 19, 1885, p. 292. Ebert, Jahrb. d,
geolog. Landesanst, 1889, p. 218. Baldwin, Geol. Mag., Dec. 5, vol. 8,
1911, p. 75. H. Woodward, Geol. Mag. n.s., Dec. 6, vol. 5, 1918, p. 465.
Dix and Pringle, Summary of Progress, Geological Survey, Great Britain,
for 1928 (1929), pt. 2, p. 103. Pruvost, Mem. du Musee Roy. d'Hist. Nat.
Belgique, no. 44, 1940, p. 201. Kobayashi, Jap. Jour. Geol. and Geog.
Tokyo, vol. 10, 1933, p. 178. Willard and Jones, Proc. Penna. Acad. Sci.,
vol. 35, 1935, p. 127. Eller, Ann. Carnegie Mus., vol. 27, 1938, p. 152.
Prestwichianella, Tchernechev, Comite Geol. du Leningrad, vol. 46, 1927,
no. 5, p. 648, 653. — Anthracopeltis, Boulay, Ann. Soc. Scientifique,
Bruxelles, 4 annee, 1880, p. 277.

Euproopidae with a tripartite cardiopthalmic area, the cardiac lobe
bordered by dorsal furrows. Intergenal spines present. Genotype,
Bellinurus danae Meek and Worthen.

Euproops danae (Meek and Worthen)

Bellinurus danae Meek and Worthen, Proc. Acad. Nat. Sci., Philadelphia,
1865, p. 44; Illinois Geol. Survey, vol. 2, 1866, p. 395, pi. 32, fig. 2, 2a.

Prestwichia danae Meek, Am. Jour. Sci., ser. 2, vol. 43, 1867, p. 257. Packard,
Nat. Acad. Sci., vol. 3, 1886, Mem. 16, p. 146, pi. 5, figs. 3, 3a; pi. 6, figs.
1, la, 2, 2a.

Euproops danae Meek. Am. Jour. Sci. Ser. 2, vol. 43, 1867, p. 395; Geol. Mag.,
vol. 4, 1867, p. 320. H. Woodward, Geol. Mag. vol. 5, 1868, p. 2. Meek
and Worthen, Geol. Sur. Illinois, vol. 3, 1868, p. 547, text fig. A. White,
Geol. & Nat. Hist. Sur. Indiana, 13th Ann. Rept. for 1883 (1885), p. 170,
pi. 39, fig. 1.

Euproops colletti White, Geol. Nat. Hist. Sur. Indiana, 13th Ann. Rept. for
1883 (1885), p. 172, pi. 39, fig. 2.

RAYMOND: LATE PALEOZOIC XIPHOSURANS 485

For bibliography of numerous European specimens which have been
referred to this species, see Pruvost, Mem. du Musee Roy. d'Hist.
Nat. Belgique, no. 44, 1930, p. 203. Without specimens, it is impossible
to say whether or not any of them are conspecific with the American
forms.

This species has been so well described that no formal diagnosis is
necessary. Although specimens are common, few are really well pre-
served. The test appears to have been but slightly if at all impreg-
nated with calcium carbonate, perhaps because of the freshwater
environment. Because of its tenuous nature the shell wrinkled easily,
and I have seen no individual with the prosoma undistorted. The
thoracetron, however, is in many cases, rather well preserved. The best
figures are the photographs in Packard's article (specimen in U. S.
Nat. Mus. no. 38, 954).

The prosoma is from 2.5 to 3 times as wide, at the bases of the genal
spines, as it is long. The cardiopthalmic area is outlined by narrow
ridges which are concave outward back of the eyes, convex forward in
front of them, forming a double arch supported by the narrow anterior
prolongation of the cardiac lobe. The eyes are a little further apart
than are the posterior ends of the opthalmic ridges, and the cardiac
lobe is about one-fourth shorter than the distance between the eyes.
As pointed out by Meek and Worthen in their original description, the
cardiac lobe tapers rapidly forward, the anterior third being narrow,
almost linear. The better preserved specimens show a transverse bar
at the point where the taper ends.

An exceptional specimen (Yale Univ. Mus. No. 16,909) shows the
paired ocelli at the anterior end of the cardiac lobe, and three pairs of
faintly impressed crescentic pits in the dorsal furrows. The first of
these is immediately behind the transverse bar just mentioned, the
last at the base of the cardiac lobe. The intermediate ones are exceed-
ingly faint. These are doubtless scars of attachment of coxal muscles.

The thoracetron is divided by dorsal furrows into a median and
lateral lobes. The furrows are almost parallel but diverge near the
posterior end to embrace a blunt boss which is excavated behind.
Including this boss, there are six rings on the axial lobe; the first and
third bear rounded tubercles; the boss has a short thornlike spine.
In the dorsal furrows there are six pairs of depressions, indicating six
pairs of entopopheses like those of modern Limulus. The first pair is
beside the first ring, the last ones just in front of the boss. On the front
of the boss, inside the furrows, is a pair of shallow conical pits.

On the lateral lobes, each segment is set off by a narrow ridge at its

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posterior margin. Six of these are extended across the flattened border
into long spines directed radially backward. A seventh segment is
indicated by short spines which extend backward close to the telson.
Compared with Hamulus, there is little space between the posterior
appendages and the telson. The margins show no indications of
articulated spines.

Few specimens retain the entire telson, which is much longer than
is generally supposed. On the only individual on which it appears to
be complete (M.C.Z. 4686, obverse), it is 26 mm. long. The thoracetron
to which it is attached is 17 mm. long, and the prosoma 19 mm.

Young specimens differ in some respects from the adult. The
cardiac lobe tapers regularly forward instead of contracting to a
narrow ridge at the half-length, and the thoracetron is proportion-
ately wider, with shorter, much less strongly developed spines. That
the young, at least, had the power of enrollment, is shown by one
specimen (M.C.Z. 4673).

Measurements. The following are the measurements of the type
given by Meek and Worthen in their original article: "Entire length
from the extremity of the caudal segment to the anterior margin of
the cephalo-thorax, about 1.90 inches. Length of cephalo-thorax, 0.57
inch, breadth of do. to the extremities of the postero-lateral spines,
1.70 inches; length of area included within the ocular ridge, 0.50 inch;
greatest breadth of do. (which is the distance between the eyes,) 0.60
inch. Length of abdomen, 0.65 inch; breadth of do., exclusive of the
flattened margin, 0.94 inch, including it, 1.06 inch; breadth of mesial
lobe, 0.23 inch; length of caudal segment, about 0.60 inch."

Formation and locality. This is the most common xiphosuran in the
nodules in the Francis Creek shale in the Mazon Creek region in
Grundy and Wills counties, Illinois. Evproops colletti, described by
White from a poorly preserved specimen from Durkee's Ferry, Vigo
County, Indiana, probably belongs to this species.

Euproops thompsoni spec. nov.

Figs. 1, 2

Study of collections of Euproops from the vicinity of Mazon Creek
shows that the common forms represent two species. The chief differ-
ence is in the proportions of the prosoma. E. danae is a wide headed
form, the width at the bases of the genal spines being from 2.5 to 3
times the length. The other species is relatively more narrow headed,

RAYMOND: LATE PALEOZOIC XIPHOSURANS

487

with a definite ratio of width to length of two to one. Specimens of
this kind are rather common.

It might seem that the difference in proportion could be due to the
state of preservation, that is, that the flattened specimens would show
a proportionately wider prosoma than an uncrushed one. One does
find some variations in measurements due to this cause, but I have
found in the Harvard and Yale collections both young and full grown
flattened and fully convex specimens of both the wide and the narrow
headed forms, and am convinced that the difference is not one of con-
dition of preservation. The prosomas, if well preserved, can easily be
distinguished by the difference in the form of the cardiac lobe.

Fig. 1. Euproops thompsoni Raymond — a composite figure to show the
writer's interpretation of the structure. The prosoma is based on a not-quite-
fullgrown individual (M.C.Z. no. 4683) and the remainder on the holotype
(M.C.Z. no. 4669). x 2.

The prosoma, neglecting the genal spines, is semi-circular in outline,
moderately convex in the adult, highly vaulted in the halfgrown

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individuals. The cardiac lobe differs from that of E. danae in its
uniform taper forward: in some specimens there is expansion in front
of the mid-length. Since the prosoma is proportionately longer, the
cardiopthalmic area appears to be somewhat smaller than in E. danae.

Fig. 2. Euproops thompsorii Raymond. The prosoma of a young individual
with especially well preserved genal and opthalmic spines, x 3. M.C.Z.
no. 4684.

Young specimens

Several young individuals in the collection are unusually well pre-
served. The prosoma is more highly convex than in the adult, and the
cardiac lobe has less anterior expansion. There is a short spine at the
posterior end of the cardiac lobe, and a long one at the end of each
opthalmic ridge; each spine has a narrow dorsal carina. The intra-
opthalmic areas, between the cardiac lobe and the opthalmic ridges
have shallow transverse furrows, limiting six pairs of ill-defined lobes.
The genal spines of these uncrushed specimens are directed almost
straight backward suggesting that the flare so common in the adults
is due to flattening. Two specimens are casts of the under side, and
show in front of the cardiac lobe a longitudinal depression (cardiac
furrow) occupied by the anterior ventral portion of the stomach.
The doublure is narrow, with what appears to be a median plate.
The specimen also shows the proximal segments of three of the
appendages. They indicate that the coxae were short and attached
just outside the dorsal furrows.

RAYMOND: LATE PALEOZOIC XIPHOSURANS 489

The thoracetron is proportionately shorter than in the adult, but
has 6 pairs of ribs, 5 rings on the cardiac lobe, and 6 pairs of entopo-
physes.

Measurements. The holotype (M. C. Z. 4669), is 57 mm. long,
including the telson. The prosoma is 20 mm. long, 38 mms. wide
at bases of genal spines: the cardiac lobe is 11 mm. long, the width
between the eyes is 14 mm., and the posterior ends of the opthalmic
ridges are 14 mm. apart. The thoracetron is about 20 mm. long, and
about 24 mm. in greatest width. The telson is 23 mm. long, but is
incomplete at the posterior end. The prosoma of a young specimen
(M.C.Z. 46S2), is 10 mm. long, 21 mm. wide; the cardiac lobe is
6.5 mm. long, the width between the eyes 9 mm., and the poste-
rior ends of the opthalmic ridges are 7 mm. apart. The thoracetron of
another young specimen (M.C.Z. 4670), is 8 mm. long and 12 mm.
wide. Apparently the thoracetron increases more rapidly in length
than in width during growth.

Formation and locality. The specimens here figured are from the
Francis Creek shale of the Carbondale series at the Wilmington Strip
Mine, Wills Co., Illinois. They were donated by Mr. Frederick
Thompson, for whom the species is named. The holotype is M.C.Z.
4669 and the paratypes M.C.Z. 4670, 4676, 4682, 4683, and 4684.

Euproops darrahi spec. nov.
PI. 2, fig. 4

The specimen on which this species is founded is unquestionably a
young Euproops of the E. danae type, the prosoma being about 2.5
times as wide as long. It differs from E. danae, however, in that the
cardiopthalmic area and the cardiac lobe are proportionally longer,
and in that the thoracetron is proportionately considerably larger.

The specimen appears to show the interior of the test, and 6 pairs
of entopophyses are well shown on the thoracetron. Because it is a
young individual, the marginal spines are short. The cardiac lobe
tapers gradually forward from the posterior to the anterior end, a
characteristic of the young of E. danae and the adult of E. thompsoni.

Dimensions. Length of holotype without telson, 10 mm.; length of
prosoma, 5 mm., width at bases of genal spines, 13 mm.; length of
cardiac lobe, 4 mm.; length of thoracetron, 5 mm., width, about 9.5
mm.

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Formation and locality. The holotype (M.C.Z. 4691) was found in
the Mason shale beneath the Brush Creek limestone in the lower part
of the Conemaugh, at Fair Oaks, Ambridge, Pennsylvania. It was col-
lected and donated by Mr. W. C. Darrah, for whom it is named.

0

Euproops laevicula spec. nov.

Fig. 3

The prosoma of this species is like that of E. thompsoni; but the
thoracetron differs in having the lateral lobes almost smooth, with
only the faintest traces of transverse ridges. The axial lobe has only

Fig. 3. Euproops laevicula Raymond.
Mus., no. 16912).

The holotype. x 3. (Yale Univ.

faint transverse furrows, and although there are nodes on the third and
last rings they are less well developed than in other species. There
appears to be an extra pair of lateral spines at the anterior end of the
thoracetron, a condition which may have some significance in connec-
tion with the position of the articulation between the prosoma and
thoracetron.

Dimensions. The holotype, a young specimen, has a prosoma 8.5
mm. long and about 18 mm. wide. The thoracetron is 7 mm. long and
about 14 mm. wide.

Formation and locality. From the Francis Creek shale at Mazon
Creek, Grundy Co., Illlinois. The holotype is 16912 and the paratype
16916 in the Yale University Museum.

RAYMOND: LATE PALEOZOIC XIPHOSURANS 491

Euproops laticephalus spec. nov.

Fig. 4

This species is characterized by the great width of the prosoma
and the relatively short wide cardiopthalmic area. It differs in the
latter respect from all the other American species. The cardiac lobe is
like that of E. danae, that is, it tapers abruptly forward and is then
continued as a narrow median ridge. There is a small median pustule

Fig. 4. Euproops laticephalus Raymond. The holotype. Natural size.

at the posterior end. Only the proximal portion of one genal spine is
preserved, but it indicates that the full length was considerable.

Euproops islwyni Dix and Pringle4 has the same broad form and
short wide cardiopthalmic area. It may be related.

Dimensions. The prosoma is 17 mm. long and about 40 mm. wide.
The cardiopthalmic area is 9 mm. long on the median line, 11 mm.
wide at the posterior margin, and the width between the eyes is 13 mm.

The species is probably most closely allied to E. danae, differing in
having a shorter cardiopthalmic area.

Formation and locality. The holotype was collected many years ago
by Leo Lesquereux from roof shale of the Salem coal, high in the
Pennsylvanian, near Potts ville, Penna. W. C. Darrah has identified
the following plants on the same slab: Asterophyllites sp., Spheno-
phyllum filiculme Fontaine and White, Pecopteris arborescens, Schlot-
heim, P. feminaeformis Schlotheim, Mariopteris ribeyroni Zeiller, and
Neuropteris plicata Brongniart. This flora is, Mr. Darrah informs me,
characteristic of the lower beds of the Monongahela series of the
Pennsylvanian. The holotype is M.C.Z. 4692.

Euproops longispina Packard

Euproops longispina Packard, Am. Naturalist, vol. 19, 1885, p. 292.
Prestivichia longispina Packard, Nat. Acad. Science, Mem. vol. 3, 1886, p. 147,
pi. 5, fig. 4, (not pi. 6, fig. 3).

* Geol. Survey of Great Britain, Summary of Progress for 1928, 1929, p. 107, fig. 12.

492 bulletin: museum of comparative zoology

This species was badly described and figured by Packard. Dr.
Bassler has been kind enough to send me a cast of the specimen on
which it was founded. In this case there is a real holotype, (U. S. Nat.
Mus., 38, 857), for the author definitely stated that the species was
based on this individual.

The general characteristics of the species are those of E. danae, the
prosoma being short and wide. The direction of the opthalmic ridges
is the same as in that species, but the cardiac lobe tapers gradually
forward as in E. thompsoni. Packard's figure of the prosoma may be
entirely ignored, for the cardiopthalmic area does not taper forward,
and the genal spines are not particularly long.

The thoracetron is like that shown by Packard only in the length of
the marginal spines. The cardiac lobe is definitely outlined, has five
rings in front of the large posterior one, and there are the usual ele-
vated ribs on the plural lobes.

This species differs from Euproops danae in having much longer
spines on the thoracetron; from E. thompsoni in this respect and in the
wider prosoma ; from E. laticephalus in having a longer cardiopthalmic
area, and from E. packardi and E. laevicula in having more distinct
ribs in the pleural lobes, as well as in the long spines, which are the
most distinctive feature.

Formation and locality. This specimen is from the shale over Coal
E, (Mammoth vein), at the Butler mine, Pittston, Pennsylvania, and
hence is presumably of Upper Allegheny age. Holotype in the U. S.
National Museum, no. 38,857.

Euproops spec. ind.

Prestwichia longispina Packard (partim), Nat. Acad. Sciences, Mem. vol. 3,
1886, p. 147, pi. 6, fig. 3.

So far as one can judge, this rather poorly preserved specimen
belongs to a species closely allied to E. thompsoni. As mentioned in
the discussion of E. longispina it was mentioned by Packard as an
"additional specimen," and so should not be listed as a cotype as it
has been in the U. S. National Museum.

Formation and locality. The exact zone from which this specimen
was obtained is not known, as it was found on the dump at the Oak-
wood colliery, Wilkes-Barre, Penna. It is supposed to have come from
Upper Allegheny or Lower Conemaugh strata.

RAYMOND: LATE PALEOZOIC XIPHOSURANS 493

Euproops amiae H. Woodward

Euproops Amiae H. Woodward, Geol. Magazine, ser. 6, vol. 5, 1918, p. 465,
figs. 2, 3, 4.

Euproops amiae is closely allied to both E. danae and E. thompsoni.
The specimen is small, the prosoma only 12 mm. long, and hence
probably not adult. The width at the bases of the genal spines is 29
mm., nearly 2.5 times the length, hence the proportions suggest those
of E. danae. The species differs from both E. danae and E. thompsoni,
however, in having a wider cardiac lobe.

The original specimens were from the Glace Bay mines, Cape
Breton, Nova Scotia.

Euproops packardi Willard and Jones

Euproops packardi Willard and Jones, Penn. Acad. Sci., Proc. vol. 9, 1935, p.
127, figs. 1, 2.

This species was founded on what appears to be a young individual
of the E. danae type. It has a wide prosoma, showing relationship to
E. danae, and a short wide thoracetron indicating that it is young (the
length of the specimen without telson is 17.5 mm.). The cardiac lobe
is particularly like that of E. danae in coming to a point at about the
half length, and being continued as a narrow7 ridge. For details of the
prosoma the photograph rather than the drawing should be consulted.

The real difference from E. danae is in the thoracetron. The ridges
which indicate the segmentation do not cross the margins onto the
spines, nor do they reach the axial lobe. In this respect the species is
more closely allied to E. laevicula, described above.

The specimen wras collected from a culm pile at the Baltimore mines
near Parsons, Penna., and is probably of Allegheny age.

A specimen in the Lesquereux collection (M.C.Z. 4693) from an
undetermined zone "Low in the Coal measures" at or near Wilkes-
Barre, Penna., may be an adult of E. packardi. It is preserved in
pyrite coated with carbonaceous mud, and is exposed from the inside.
It agrees with E. packardi in having incomplete ribs on the pleural
lobes of the thoracetron, and in proportions. The prosoma is 12 mm.
long and about 30 mm., wide, and the cardiopthalmic area has a mid-
length of 8.5 mm. The thoracetron is 9.5 mm. long and about 20 mm.
wide, hence proportionally a trifle narrower than that of the younger
individual which is the holotype.

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Euproops lacoei (Packard)

Belinurus lacoei Packard. Am. Naturalist, vol. 19, 1885, p. 292: Nat. Acad.
Sciences, Mem. vol. 3, 1886, p. 149, pi. 5, fig. 5.

As Dunbar has already pointed out, this species cannot possibly be
included in Belinurus. It is a Euproops, closely related to E. danae.

Schuchert, in cataloging the types in the United States National
Museum1 remarked of this species : "Probably the same as Prestwichia
danae. Probably a sexual difference." Even before I saw this state-
ment I had inferred from the figure that Belinurus lacoei was a young
Euproops danae, but curiosity about the number of segments in the
thoracetron led me to seek information about the types. Fortunately
they are in the National Museum, and Dr. Ray S. Bassler has supplied
me with casts of all of them.

Packard listed as "types," specimens in Mr. Lacoe's collection
numbered 210m, 210y, 210wx, 212ab, and 213a. All were, therefore,
cotypes. In the Catalogue already referred to, however, Schuchert
listed only one of them, no. 210y, and designated it as the holotype.
His reason for doing this is probably explained by the label now with
the specimen, and supposed to be in Mr. Lacoe's handwriting, stating
that it is the original of Packard's plate 5, figure 5.

Without the label, no one would have suspected that this was the
type, for the middle portion of the prosoma is entirely broken away,
no traces of opthalmic ridges, eyes, or cardiac lobe remaining. The
damage is so great that it is difficult to measure the length, but it is
about 10 mm. Since the figure is labeled as twice natural size, this
measurement checks. The width at the bases of the genal spines is
about 24 mm. which is only a little less than that shown by the figure.
The thoracetron is poorly preserved, is shorter and wider than in the
figure, and the cardiac lobe so damaged that the number of rings can
not be surely counted. There is no reason to think that there are more
than in E. danae. The telson is not completely exposed, the distal
portion being under the matrix. The part shown is 12 mm. long,
whereas, according to the figure, it should be 30 mm. long. From the
part visible and the rate of taper, a total of 20 mm. would be the
maximum length to be expected. In other words, the "holotype"
shows none of the characteristics of the figure except correspondence
in length of the prosoma.

'U. S. Nat. Mus. Bull. 53, 1905, p. 96.

RAYMOND: LATE PALEOZOIC XIPHOSURANS •!!).")

There is, however, a question whether this specimen, although it was
the basis of the figure, was really the most important of the cotypes,
for Packard's measurements of what he calls the "best preserved speci-
men" are those of a much smaller individual, with the prosoma and
thoracetron together 15 mm. long. There is no individual among the
cotypes of exactly this size, and but one which closely approximates it.
This is the one numbered 210hl, which occupies first place in Packard's
list. The prosoma and thoracetron are 14 mm. long in this specimen.
The prosoma is 7.5 mm. long and 17.5 mm. wide, and the telson,
which is incomplete, is 11 mm. long. The entire length might have
been as much as 14 or 15 mm., or about the same length as the body.
In all probability this is the measured individual, but it is so poorly
preserved that it could not have furnished the information for the
figure. The cardiac lobe of the thoracetron is especially poor, and the
number of segments cannot be counted.

Packard's figure is admittedly composite, but the types justify none
of its important characteristics. Two of them have already been dis-
cussed. Lacoe's no. 210^ is, perhaps, the best preserved of the lot.
It has the same sort of cardiac lobe on the prosoma as E. danae, and
the same number of rings on the thoracetron, but the lateral lobes of
that shield are about as smooth as in E. laevicula. No. 212ab has the
same sort of thoracetron, and a badly damaged prosoma. Neither
shows a great deal of the telson. No. 213a is a small, badly pre-
served individual which may be a specimen of Liomesaspis laevis. In
addition to these, the National Museum has another specimen not
mentioned by Packard, Lacoe's No. 212° (U. S. Nat. Mus. no. 38,861).
This is much better preserved than any of the cotypes, but is a typical
young E. danae, with the cardiac lobe continued forward from the mid-
length as a narrow ridge, and typical thoracetron. The prosoma is
10 mm. long and 25 mm. wide.

It seems then, that Euprobps lacoei was founded upon a misinterpre-
tation of five poorly preserved specimens, the least obscure of which is
the one Schuchert designated as the holotype. Such evidence as exists
indicates that it is a young individual of E. danae, and the name may
as well be dropped.

Formation and locality. All the specimens are said to have been
found at Mazon Creek, Grundy County, Illinois.

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Genus Anacontium genus nov.

Euproopidae with tripartite cardiopthalmic area, but without inter-
genal spines. Type, Anacontium carpenteri, spec. nov.

Anacontium carpenteri spec. nov.
Fig. 5
Prosoma rounded in outline, less than twice as wide as long, evenly
convex. The genal spines are small, vestigal. The cardiac lobe is rela-
tively short, wide at the posterior end, tapering forward, ending in a
prominence behind the point at which the preocular ridges meet. The
dorsal furrows are shallow but distinct. The intra-opthalmic areas

Fig. 5. Anacontium carpenteri Raymond, x 4.
Fig. 6. Anacontium brevis Raymond, x 4.

show no furrows. The eyes are small, well forward and far apart, so
that the cardiopthalmic area occupies a large portion of the prosoma.
The genal spines are minute, and probably would not be seen on poorly
preserved specimens. The posterior margin of the prosoma is turned
down vertically, as in modern Limulus.

Measurements. The holotype (prosoma) is 7 mm. long, 12 mm.
wide; the width between the eyes is 7.50 mm. and the posterior ends
of the opthalmic ridges are 5.75 mm. apart. The cardiac lobe is 4.5
mm. long, and 3 mm. wide at the posterior end. The cardiopthalmic
area is 5 mm. long, on the median line. The paratype (prosoma) is 5
mm. long, about 9 mm. wide and the width between the eyes is 5.50
mm.

Formation and locality. Two specimens of the prosoma were collected
from the Wellington formation on the southwest quarter of the north-
west quarter of section 2 of township 21 north, range 1 west, in Noble
County, Oklahoma, where they were associated with numerous speci-
mens of Conchostraca and insects. The holotype is M.C.Z. 4724, and
the paratype M.C.Z. 4725. It is named for Professor F. M. Carpenter
who found the specimens, and whose studies and collections have
greatly enlarged our knowledge of Permian arthropodan faunas.

RAYMOND: LATE PALEOZOIC XIPH0SURA\> 497

Anacoxtium brevis spec, no v.
Fig. 6

This species differs from the preceding in having well developed
genal spines, the eyes further forward, and particularly in the unusually
short cardiac lobe. It might at first sight be confused with Paleolim id us
avitus, because of the lack of intergenal spines and the general configur-
ation of the prosoma. The postocular ridges curve inward, however,
and the cardiac lobe is surprisingly short.

Measurements. The only known specimen is a prosoma which is not
well preserved on the anterior margin, and in which the cardiac lobe
has been crushed so that it is concave, instead of being convex. The
measurements are therefore approximate.

Length, about 6.00 mm., width about 10.00 mm. The width between
the eyes is 3.00 mm., the cardiac lobe is about 2.00 mm. long, and the
point where the preocular ridges meet on the median line is 4.25 mm.
from the posterior margin.

Formation and locality. The holotype, M.C.Z. 4726 was collected by
Dr. F. M. Carpenter from the Wellington formation on the southwest
quarter of the northwest quarter of section 2, township 21 north,
range 1 west, in Noble County, Oklahoma.

Family Elleriidae fam. now

Euproopacea in which the primitive segmentation of the posterior
portion of the axial lobe of the thoracetron is not obscured.

Genus Elleria genus now
Elleria morani (Eller)
Euproops morani Eller. Ann. Carnegie Mus., vol. 27, 1938, p. 151, fig. 1.

Eller himself hesitated to make a new genus for this species, because
onlv the thoracetron was known. However, it is not a Euproops, and
to assign it to that genus gives the impression that Euproops occurs
in the Devonian, which is misleading. I am therefore, naming it for
Dr. Eller, and expressing the hope that he will find the prosoma which
belongs with the species.

Elleria morani, as preserved, lacks the anterior part of the thorace-
tron but even though it had no more lateral area than is shown on the
right-hand side of the original figure, it must have had two more rings
on the axial lobe, making eight in all. Euproops has five distinct rings

498 bulletin: museum of comparative zoology

and six lateral ridges, one of them springing from the sides of the
composite sixth ring. To make a homology, it would be necessary to
postulate that the last three rings on E. morani corresponded with the
anchylosed area at the posterior end of the axial lobe of Euproops.
Whatever the actual structure, it lacks the characteristic expression
of the posterior end of the thoracetron of the Euproopidae.

It is, however, an interesting and important specimen, being the
oldest (Upper Devonian) xiphosuran with a fully anchylosed thorace-
tron. It is primitive, in that the median elements of all three of the
prometasomatic segments remain distinct. In all probability there
would be, in a complete specimen, eight rings on the axial lobe. The
deep emargination at the posterior end may also be primitive, for
Euproops shows less of this feature and Liomesaspis none of it, and
the latter is certainly a specialized genus.

Formation and locality. The holotype is from the Salamanca sand-
stone of the marine Upper Devonian at North Warren, Penna.

Family Liomesaspidae fam. nov.

Euproopacea without true dorsal furrows on the prosoma or lateral
spines on the thoracetron.

Genus Liomesaspis genus nov.

Liomesaspidae with rounded prosoma and without genal spines or
defined cardiac lobe in the adult. Axial lobe of thoracetron clearly
defined, but obscurely segmented. Genotype, Liomesaspis laevis spec,
nov.

RAYMOND: LATE PALEOZOIC XIPHOSURANS

499

Liomesaspis laevis spec. nov.

Figs. 7, 8, 9, 10

Prosoma evenly convex, from one-third to one-half wider than
long, with a narrow flattened brim, only traces of which have been
seen. The cardiac lobe is not outlined and the intraopthalmic area is

Fig. 7. Liomesaspis laevis Raymond. A much wrinkled specimen, with little
trace of segmentation, x 3.

Fig. 8. The same species. A paratype showing more divisions of the cardiac
lobe of the thoracetron and retaining the telson. Yale Univ. Mus., no. 16913.
x3.

nearly smooth, except for two divergent furrows which extend forward
and outward from the posterior margin. On specimen M.C.Z. 4696 (fig.
7) the opthalmic ridges project as short spines at the posterior margin,
but their full course can not be traced. Eyes are probably present,
but no specimen is well enough preserved to give absolute proof of
their position. They are probably a little in front of the middle, and
widely separated. In front of the putative positions of the eyes, each
opthalmic ridge arches around to a reentrant on the median line, as in
Euproops. An immature specimen in the Yale University Museum
(no. 16,914) shows small genal spines, placed well forward at the sides.

500

bulletin: museum of comparative zoology

This is of interest since it indicates in these animals the same tendency
as in the trilobites for genal spines to move forward and disappear.

The test of all specimens of the prosoma was evidently weak and
thin, for all are considerably distorted and it is impossible to be sure
of the original configuration. Since the specimens are small, this is in
curious contrast to the condition in Euproops, in which many of the
young are well preserved, whereas the adults are distorted.

Fig. 9. Liomesaspis laevis Raymond. An unusually well preserved indi-
vidual, except for the telson. The holotype. x 2.5.

Fig. 10. The same species. A thoracetron, exposing the lower surface. At
the posterior end are processes for articulation with the telson. Yale Univ.
Mus., no. 16915. x 2.

The thoracetron, on the other hand, is commonly well preserved.
The axial lobe is narrow, but expands at the posterior end where it
rises into a high blunt spine. On the best preserved specimens this
spine is excavated behind, having exactly the same shape as in
Euproops danae. The best preserved axial lobes show three complete
rings and a half -ring in front of the terminal enlarged portion, but the
transverse furrows are so shallow that the lobation is not conspicuous.
As a rule it shows better on the cast of the interior of the shell than on
the exterior. Three pairs of short linear grooves indicate the position
of the entopophyses.

The lateral lobes are smooth, flat on the upper anterior surfaces,
abruptly turned downward at the sides and back.

The telson is long and slender, slightly over two-fifths of the total
length in the two specimens in which it has been possible to excavate
the whole of it. Two processes extending backward from the underside
of the thoracetron prevented its being turned downward, hence it was

RAYMOND: LATE PALEOZOIC XIPHOSURANS 501

of no value as a pushing organ, and it must have been used principally
in righting the animal when accidentally overturned. Most of the
specimens show no trace of it, but in the three in which parts of it are
preserved, it is turned upward.

These animals seem to be particularly well adapted for enrolment,
since the posterior part of the prosoma fits the anterior margin of the
thoracetron, and the sides of the adjacent portions of the two shields
are so moulded as to fit against one another. Specimen M.C.Z. 4697
may be such an enrolled individual.

Measurements. The holotype (fig. 9) is 10.50 mm. long without the
telson. The prosoma is 9.25 mm. long and 15 mm. in greatest width.
The thoracetron about 7 mm. long and 12 mm. in greatest width. The
thoracetron is partially overlapped by the prosoma. A complete
specimen (Yale Univ. Mus. no. 16,913), is 24 mm. long; the prosoma
8 mm., the thoracetron about 6 mm., and the telson 10 mm. long. A
paratype (M.C.Z. 4696) is 17 mm. long without the telson, the prosoma
10 mm. long and 13.5 mm. wide, the thoracetron 7 mm. long and 12
mm. wide. An isolated thoracetron (M.C.Z. 4697) is 11 mm. long
and 15 mm. wide: an immature specimen (Yale Univ. Mus.) is 13 mm.
long, the prosoma 4 mm., the thoracetron 3 mm. and the telson 6 mm.
long.

Formation and locality. It is curious that this little form has not
been described previously, for it seems to be fairly common in the
Francis Creek shale at Mazon Creek, Illinois. The holotype is M.C.Z.
4698, the paratype shown in fig. 7 is M.C.Z. 4696; those shown in figs.
8 and 10 are in the Yale University Museum, where there are several
excellent specimens.

It is probable that these specimens have been mistaken for young
or incomplete individuals of Euproops danae, but the collection studied
contains many young of that species which are so like the adult that
there is no justification for such identification.

Genus Pringlia genus nov.

Liomesaspidae with the cardiac lobe well developed. Genotype,
Prestwichia birtwelli H. Woodward.

Pringlia birtwelli (H. Woodward)

Prestwichia Birtwelli H. Woodward, Geol. Magazine, vol. 9, 1872, p. 440, pi. 10,

figs. 9,10; Paleontographical Soc. London, 1878, p. 247, pi. 31, figs. 7a, b, c.

Euproops Birtwelli H. Woodward, Geol. Magazine, Ser. 6, vol. 5, 1918, p. 468.

502 bulletin: museum of comparative zoology

Woodward described this species originally as without spines on
the border of the thoracetron, but in his last paper listed above he
stated that in all probability spines were present, but hidden in the
matrix. In view of what is known of the numerous specimens of
Liomesaspis that seems highly unlikely.

The general configuration of Pringlia birtwelli is almost identical
with that of Liomesaspis, but although there are no true dorsal fur-
rows on the prosoma, the cardiac lobe is outlined for its entire length.
Moreover, there are distinct genal angles, with a trace of a minute
spine. It is not at all likely that the two small spots midway in the
head, which Woodward identified as eyes are really such, for they are
outside the opthalmic ridges. It is more probable that the eyes are
far forward, where the opthalmic ridges turn inward.

The thoracetron shows five rings on the cardiac lobe, each with a
small median pustule, and behind them is a longer spine-bearing
terminal portion, as in Liomesaspis. The lateral lobes show traces of
segmentation.

Measurements. Woodward gives the following measurements:
(one line equals about 2 mm.). Entire body; length, 8 lines, greatest
breadth, 8 lines. The prosoma is 4 lines long, the thoracetron 4 lines,
the telson 4 lines. One would judge from the figure that the telson
is incomplete. The proportions are, therefore, about the same as in
Liomesaspis laevis.

Formation and locality. Only two specimens have ever been found,
so far as I can learn. They came from the Coal Measures at the
Cornfield Pit, on the south bank of the River Calder, Padiham,
Lancashire, England. The generic name is for Dr. John Pringle, in
recognition of his years of study of the Coal Measures of Great
Britain.

RAYMOND: LATE PALEOZOIC XIPHOSURANS

503

Pringlia bispinosa spec. nov.
Fig. 11

Only a single prosoma is known. It is roughly subcircular, depressed,
without genal spines, but with strong spines at the posterior ends of
the opthalmic ridges. The course of each of these ridges is forward

Fig. 11. Pringlea bispinosa Raymond. The holotype.
no. 16911. x3.

Yale Univ. Mus.,

and outward to the eye, which is at about midlength, then forward
and inward to the median line, where the two ridges unite at the
anterior end of the cardiac lobe. The latter is raised but slightly
above the general surface, and tapers uniformly forward.

The greatest width of the prosoma is at the genal angles which are
pointed, but without spines.

This species differs from P. birtivelli chiefly in that the eyes are
further back on the shield.

Measurements. Length of prosoma, 10.5 mm., width at genal
angles, 15.5 mm. Length of cardiac lobe, 7 mm., width between
eyes, 10 mm.

Formation and locality. The holotype, from the Francis Creek shale
at Mazon Creek, Illinois, is no. 16911 in the Yale University Museum.

Genus Prolimulus Fritsch

Prolimulus Fritsch, Geol. Magazine, dec. 4, vol. 6, 1899, p. 58, fig., Fauna der
Gaskohle und der Kalksteine der Permformation Bohems., vol. 4, 1899-
1901, p. 64, figs. 369, 370, pi. 155.

All specimens of the genotype, Prolimulus woodardi Fritsch are
so badly preserved that this genus can hardly be said to have any

504 bulletin: museum of comparative zoology

generic characteristics. The one outstanding feature that is significant
is that the prosoma has no genal spines, which may indicate relationshp
to Liomesaspis, with which genus it agrees further in lacking spines
on the thoracetron. It may therefore be placed provisionally in the
Liomesaspidae.

According to Fritsch, the prosoma is about 1.5 times as wide as
long, the thoracetron somewhat wider in proportion. A specimen in
the Museum of Comparative Zoology (M.C.Z. 4694) has the following
dimensions: length of prosoma, 9.5 mm., width, about 14 mm.; length
thoracetron, 7 mm., width 11 mm.

Formation and locality. This species is common in the Permian
Gaskohle at Nyran in Bohemia.

Superfamily LIMULACEA nov.

Limulada with the posterior portions of the opthalmic ridges parallel;
thoracetron trapezoidal, with movable lateral spines.

Family PALEOLIMULIDAE fam. nov.

Limulacea with opthalmic ridges meeting in front of the eyes,
with a narrow doublure on the prosoma, and axial rings on the thor-
acetron.

Genus Paleolimulus Dunbar

Paleolimulidae with conspicuously lobed intra-opthalmic areas
and with lateral lobes of the thoracetron smooth except for rows of
nodes near the margin. Genotype, Paleolimulus aiitus Dunbar.

Dunbar considered the lobation of the "glabella" (intra-opthalmic
areas) as the most important characteristic of this genus. This is
only partially true, for some almost fully grown specimens of Euprobps
thompsoni show it, as do many individuals of the Upper Jurassic
Limulus walchi. In fact, it is not difficult to find specimens of the
modern Limulus polyphemus which show lobation.

It may be, since only one genus is known, that I have included in
the family characteristics some features which are confined to the
genus. As mentioned above, in connection with the Belinuracea, it is
probable that Prestwichia randalli is a member of a more primitive
genus than Paleolimulus, to which it has been tentatively referred
by Dunbar.

RAYMOND: LATE PALEOZOIC XIPHOSURANS 50&.

Paleolimilus avitits Dunbar
PI. 1, pi. 2, figs. 1,2,3.

Paleolimulus- avitus Dunbar. Am. Jour. Sci., vol. 5, 1923, p. 444, pi. 2, fig. 1,
text figs. 2-6.

Dr. Frank M. Carpenter has collected several specimens of this
species from the typical locality, Elmo, Kansas, where it is a relatively
rare fossil associated with insects and plants in the Lower Permian
Wellington shales. They permit me to add a few details to Professor
Dunbar's excellent description.

The dorsal surface

To one who has been studying the Euproopacea, the most striking
features of the prosoma are the Limulus-like characteristics of parallel
post-ocular opthalmic ridges without spines at the posterior end, the
downward flexure of the test at the posterior margin, and the concave
areas between the posterior ends of the opthalmic ridges and the
tips of the genal spines.

The lateral extension of the first half-segment of the thoracetron
is also prognostic of Limulus, and entirely unlike anything seen in
the Euproopidae. It shows, however, a much more primitive condition
than in modern Limulus in that the distal spines are turned downward
instead of upward, and extend out beyond the greatest width of the
remainder of the shield. They are, in fact, somewhat longer than
shown in Dunbar's restoration, whereas in Limulus polyphemus they
are short. In the Jurassic L. walchi they appear to be in a somewhat
intermediate condition, less strongly developed.

No specimen in our collection shows the movable spines, or stylets,
but that they were present is shown by mounds for their attachment.
Such can be seen along the margin in figure 2, pi. 2. It is probable
that the stylets were in general larger than the two figured by Dunbar.

A peculiar feature, and one which can not be satisfactorily inter-
preted from the material at hand, is the apparent presence of a free
segment behind the thoracetron and above the anterior end of the
telson. This is shown on four specimens (M.C.Z. 4659, 4660, 4664,
4668), but all of them leave something to be desired as to detail.
The natural interpretation of this segment would be that it corre-
sponds with the transverse process on the anterior end of the telson
of modern Limulus. To this process are attached the dorsal muscles
which lift the telson. In modern Limulus this process is partly or

506 bulletin: museum of comparative zoology

entirely concealed when the telson is horizontal in position, and is
pulled forward entirely beneath the thoracetron when the telson is
lifted. In Paleolimulus, however, the transverse bar does not move
under the thoracetron, but is entirely behind it, and, moreover, it
has a lateral lappet on each side, so that it has the appearance of a
full segment. It may be that it is connected with the telson as in
Limulus, in which case it merely represents a primitive condition of
the transverse process. Even so, it does suggest that that process
was originally a segment of the trunk that has become attached to
the telson. The subject is of considerable interest as bearing upon
what has become of the posterior (six?) segments of the trunk in the
Xiphosura. For illustrations, see pi. 2, figs. 1, 2. It is perhaps best
shown by an unfigured specimen (M.C.Z. 4668), where it is definitely
free from the thoracetron, and probably free from the telson.

The doublure of the prosoma is narrow. It widens somewhat in
the middle of the front, but the posterior edge makes a smooth curve,
there being no such angulation as in modern Limulus.

Appendages

All ten of the entire specimens in the Museum of Comparative
Zoology retain more or less well preserved appendages, in most cases
pressed flat against the inner surface of the test. Yet so well did
Professor Dunbar describe them from his one specimen that com-
paratively little can be added.

As in modern Limulus, the coxae were elongate, fixed to the ventral
membrane, with the dorsal side open for the intrusion of muscles,
the aperture being outlined by a thickened frame. As in Limulus,
the five pairs spread outward and forward from the position of the
mouth, which presumably was at about the mid-length of the cardiac
lobe. The coxae seem proportionally as long as in the modern forms,
and they occupy as much space under the cardiopthalmic areas.
Their inner ends are but poorly preserved and show no traces of
gnathal spines.

Only four specimens (M.C.Z. 4658, 4664, 4665, and 4667) show
traces of the chelicerae. They are best preserved in specimen no. 4664,
wher they can be seen to be attached beside the posterior end of the
camerosome. Two segments project forward and outward, but the
pincers are not shown. Specimen no. 4665 shows a pair of pincers
ahead of the other four pincer-bearing appendages. They are turned
outward, but much nearer the median line than are those behind.

RAYMOND: LATE PALEOZOIC XIPHOSURAXS 507

It appears then, that the chelicerae were short, not recurved, and
had two segments in addition to a small pair of pincers.

The walking legs are more or less well preserved on all the specimens,
but it is impossible to make out the details of the segments. There
appear to be three segments in addition to the pincers, as in modern
Limulus. They are, perhaps, best shown on M.C.Z. 4662 (PI. 2, fig. 2).
The pincer segment is long, the one proximal to it shorter, apparently
almost square when crushed. The details of the one which articulates
with the coxa are vague in outline in all specimens. The pincers them-
selves are best shown on M.C.Z. 4665, which retains all four on the
left side and one on the right. The actual pincer part of the outer
segments is progressively larger from the second to the fifth appendage,
the increase being from a length of 1 mm. to that of 2 mm. The
prosoma of this specimen is 8.5 mm. long. The pincers themselves
are slender and gently curved, like those of modern Limulus.

The walking legs, although built on the same plan, seem to be more
slender, and much less specialized for digging than those of the modern
species. They show much less modifications for a downward-turned
position.

The sixth pair of legs, the "pushers" are more complete than those
on Dunbar's specimen since they show a long slender segment beyond
the one with the whorl of flattened setae. Whether or not this segment
bears pincers I am not sure, for they are not present on the most
complete leg (M.C.Z. &665). The whorl of blades on the pusher is
best shown on M.C.Z. 4662 (PI. 2, fig. 2).

Several specimens show traces of gills in the thoracetron, the best
of them being M.C.Z. 4660 (PI. 2, fig. 1). They express themselves
as concentric curved lines beneath the test on the lateral lobes.
Apparently the gross structure is the same as in the modern relatives.
The area occupied by the gills extends much farther back, however,
almost to the posterior end.

The camerosome is shown by only one specimen, M.C.Z. 4664,
and by that only in dorsal outline. It is elongate, narrow, somewhat
constricted at the posterior end. Although not fully cleaned out, it
appears to be canoe shaped, but not so deeply keeled as in Limulus.

So far as the specimens can be interpreted, the appendages of
Paleolimulus differ from those of modern forms only in being some-
what less specialized for digging and in lacking spinose outgrowths.
They are by no means suggestive of any primitive condition.

The appendages of Euproops are less well known, the only really
good specimen retaining them being the one described by Packard.1

> Nat. Acad. Sci.. Mem. vol. 3, 1886, p. 146, pi. 5, fig. 3a; pi. 6. figs. 1, la.

508 bulletin: museum of comparative zoology

This specimen lacks the coxae, but they are preserved, in part at least
upon a prosoma of Euproops thompsoni (M. C. Z. 4682), a young
specimen of E. laevicula in the Yale University Museum (no. 16916),
and a young individual of E. danae (Yale, no. 16910).

The young specimen of E. thompsoni has the appendages so im-
perfectly preserved that no definite conclusions can be drawn. It
appears, however, that the coxae were as elongate as in Paleolimulus.
The young E. laevicula is more important. The coxae have the same
direction as in Paleolimulus, but each has a long slender process
extending inward under the cardiac lobe. The young E. danae have
the same radial arrangement of slender coxae under the intra -opthalmic
areas as is present in Paleolimulus and Limulus.

So far as one can judge from the photograph, the outer appendages
of Euproops are about as restored by Packard. All are exceedingly
slender, even more so than in Paleolimulus, and hence even less
adapted for digging. The last pair are probably incomplete, as they
show no segment beyond the whorl of three short blades. Such a
segment is, however, mentioned in the text. The pincers of the
walking legs appear to be slender.

PLATES

PLATE 1

Raymond — Late Paleozoic Xiphosurans

PLATE 1

Palaeolimulus avitus Dunbar. A complete specimen, showing many of the
appendages of the prosoma. M.C.Z., no. 4658. x 6.

BULL. MUS. COMP. ZOOL.

Raymond. Late Paleozoic Xiphosurans. Plate 1.

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PLATE 2

Raymond — Late Paleozoic Xiphosurans

PLATE 2

Fig. 1. Palaeolimulus avihis Dunbar. A nearly complete specimen, showing
pincers on the prosoma, remains of gills on the thoracetron, and a segment
between the mid-shield and the telson. M.C.Z. no. 4660. x 12.

Fig. 2. The same species. A specimen with unusually well preserved ap-
pendages on the prosoma. It also shows the points of insertion of the stylets on
the thoracetron, and the segment in front of the telson. M.C.Z. no. 4662. x 4.

Fig. 3. The same species. An unusually well preserved prosoma, showing
the eyes and, at the front, the doublure. M.C.Z. no. 4161. x 3.

Fig. 4. Euproops darrahi Raymond. The holotype. Note the long narrow
anterior portion of the cardiac lobe. M.C.Z. no. 4691. x 3.6.

BULL. MUS. COMP. ZOOL.

Raymond. Late Paleozoic Xiphosurans. Plate 2.

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