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YALE PEABODY MUSEUM

or Natura History

Number 51 June 27, 1961 New Haven, Conn.

THE CRESTED LIZARD HAWK (AVICEDA JERDONI)
IN THE PHILIPPINES

KENNETH C. Parkes

CarNnecitr Museum, PirrspurGH, PENNSYLVANIA

The genus Aviceda contains about five species of small hawks
distributed through the Old World tropics. Two forms of this
genus have been described from the Philippines: Hyptiepus
magnirostris Kaup, 1847, and Baza leucopias Sharpe, 1888.
The latter was based on a single specimen from the island of
Palawan. The application of these two names to the Crested
Lizard Hawks of the Philippines has varied. Modern authors
agree that the Philippine birds belong to the Indo-Malaysian
species Aviceda jerdoni. Peters (1931: 196) used the name
A. j. magnirostris for the populations of Luzon and Mindanao,
and attributed those of the islands of Romblon, Samar, and
Palawan to a subspecies to which he attached Sharpe’s name
leucopias. He was followed in this treatment by Hachisuka
(1984: 34-35) and Swann and Wetmore (1986: 294-295) ; the
former author adopted the emendation to “lewcopais” pub-
lished by Whitehead (1890: 43) at Sharpe’s request.

The most recent manual on Philippine birds (Delacour and
Mayr, 1946: 40) abandons the concept of two subspecies,
using the name magnirostris for the birds of all five islands
(Palawan, Luzon, Romblon, Samar, and Mindanao) from
which Aviceda has supposedly been recorded. The description

2 Postilla Yale Peabody Museum No. 51
of the “immature” A. j. magnirostris given by Delacour and
Mayr, however, is based on the unique type of lewcopias, which
was described as a “young female” by Sharpe.

I have examined the type of lewcopias, now in the American
Museum of Natural History (Rothschild Collection), which
remains to this writing the only specimen of Aviceda known
from Palawan. It is my belief that this bird (figured in color
by Whitehead, 1890: pl. 2) is albinistic, and that there is no
definite evidence that the lewcopias coloration represents a
normal “immature” plumage of magnirostris as described by
Delacour and Mayr. Truly “immature” specimens of Aviceda
of any species are rare in collections, and examination of the
entire series of this genus in the American Museum suggests
that this is because such plumages are held very briefly, unlike
the situation in many other Accipitridae. Two juvenile speci-
mens of A. jerdoni ceylonensis in the American Museum are
similar to definitively plumaged adults, but have the feathers
of back, crown, wing coverts, etc., edged with white, and the
upper breast streaked longitudinally. It seems very unlikely
that A. j. magnirostris, of which the adults rather closely
resemble other subspecies, would have a juvenal plumage as
radically different as that represented by lewcopias. As men-
tioned above, other Aviceda hold the juvenal plumage only
briefly; the type of lewcopias is, in part, badly worn and
bleached to an extent that juvenile Aviceda would not normally
have time to attain before molting. It thus seems probable that
the type of leuwcopias represents a “dilute” or albinistic adult
female. The fate of the name depends on whether a case can be
made for two subspecies of Aviceda jerdoni in the Philippines.

The presence of the small island of Romblon in the list of
Philippine localities from which Aviceda has been taken rests
on the record of Bourns and Worcester (1894: 33), who re-
ported “Baza leucopais Sharpe” from Romblon and Samar.
These authors had not seen the unique type of leucopias, and
identified their specimens from descriptions only. Their single
Romblon specimen is now in the collection of Carnegie Museum
(no. 137904). It is not an Aviceda at all, but an immature
female Spilornis cheela, collected September 7, 1892, just seven
days before Bourns and Worcester collected an adult female

June 27, 1961 The Crested Lizard Hawk 3

(C.M. 137902) of the same species on Romblon. The very small
size of these two specimens (flattened wing 308, 305 mm) and
the pale color of the adult incidentally confirm Rand’s identi-
fication of his single Romblon specimen as Spilornis cheela
panayensis (Rand, 1951: 578). The island of Romblon may
thus be removed from the known range of Aviceda jerdoni in
the Philippines.

For many vears the status of Aviceda on the island of
Samar was precisely like the Romblon status described above,
resting on a single Bourns and Worcester specimen. This bird
was collected at Catbalogan, Samar, on August 12, 1892, and
is now in the collection of the Minnesota Museum of Natural
History, University of Minnesota (no. 11416). It, too, is an
immature Spilornis cheela (male, wing 312 mm). There is,
however, a bona fide record of Aviceda jerdont from Samar:
an adult male was collected by D. S. Rabor on Mt. Capoto-an,
May 9, 1957. The statement of Rand and Rabor (1960: 374)
that A. j. magnirostris had been “collected previously [on
Samar| by Bourns and Worcester” is thus erroneous, and
theirs is the first specimen from that island.

Leaving out of consideration the single problematical Pala-
wan specimen (the type of lewcopias), we are left with three
islands from which Aviceda jerdoni is alleged to have been
recorded: Luzon, Samar, and Mindanao. The name Hyptiopus
magnirostris Kaup was based on Baza magnirostris Gray, (List
Bds. Brit. Mus., 1844: 19), a nomen nudum, which in turn
was based on a single specimen received by the British Museum
in 1842 among some 175 Philippine specimens collected by
Hugh Cuming. Mr. R. W. Sims has kindly examined this type
specimen for me, and writes as follows (letter of July 16,
1960) : “Cuming never tied collectors’ labels on his specimens,
hence the only label is that secured in the time of G. R. Gray.
The information there reads: ‘Baza magnirostris, locality Phil-
ippines, ex. coll. Cuming.’”” At some later date (apparently),
the additional locality “Island of Manilla, South” was entered
against this specimen in the British Museum Register (Sharpe,
18938: 555). The source of this additional locality information
is problematical. It is highly unlikely that it was supplied by
Cuming, the collector; Mr. S. P. Dance of the Mollusca Sec-

4 Postilla Yale Peabody Museum No. 51

tion of the British Museum, who is working on the life and
collections of Hugh Cuming, informs me that Cuming never,
to his knowledge, employed the expression “Island of Manilla,”
whereas “Island of Luzon” was frequently used by Cuming.
Mr. Sims, of the Bird Room, writes, “I gather from Mr. Dance
that in his later years Cuming employed a secretary to do all
his writing for him so it is possible that somewhere either on
the part of the secretary or perhaps on the part of G. R. Gray
there was an error in transcription and that the type of magni-
rostris was collected at some other locality.”” Mr. Dance (letter
of August 29, 1960) says that he ‘tshould be inclined to treat
any label on a Cuming specimen with circumspection . . .”

Sharpe (1893: 556) pointed out that nobody since Cuming’s
time had recorded Aviceda from Luzon, and this statement
is equally true 68 years after Sharpe’s words were printed.
Although new species are still being discovered on Luzon, it
seems highly unlikely that a bird the size of an Aviceda would
escape detection on this, the most heavily populated Philippine
Island, particularly since most Luzon birds collected in Cum-
ing’s time, some 130 years ago, came from the vicinity of
Manila. In view of the great uncertainty attached to the va-
hidity of the locality “Island of Manilla, South,’ I would
suggest that the type locality of Hyptiopus magnirostris Kaup
be designated as Davao, Mindanao, a source of some early
Philippine collections and a locality at which the species is
definitely known to occur. A precedent for such an action with
respect to a Cuming specimen was set by Rand and Rabor
(1960: 429).

An additional published record of Aviceda jerdoni “‘leuco-
pais” must be mentioned here. Hachisuka (1941: 72) lists five
“heht creamy birds” from Mr. Hirazawa’s collection as “leu-
copais.” As these were collected in Mindanao, where magni-
rostris is also known to occur, Hachisuka stated that the two
forms must be separate species unless further investigation
should prove them to be age or individual variants within a
species. One would suspect that this might serve to confirm
Delacour and Mayr’s identification of the lewcopias type as
the “immature” plumage of magnirostris. This is not the case,
however. One of the Hirazawa specimens was obtained by

—_

June 27, 1961 The Crested Lizard Hawk 5

Hachisuka, and is now in the personal collection of S. Dillon
Ripley at the Peabody Museum, Yale University, where I have
examined it. This specimen (Hachisuka 3751, SDR 337) was
collected near Davao, Mindanao, sometime in 1927-1928. It is
neither Aviceda nor Spilornis, but a pale, very worn example
of Pernis celebensis steerei! The latter species of Honey Buz-
zard, incidentally, is highly variable in color, as is its congener
P. ptilorhynchus, as amply illustrated by the series in the
Peabody Museum collection; one would not suspect this vari-
ability from the description given by Delacour and Mayr
(1946: 42).

An additional specimen in the Hachisuka collection (Hachi-
suka 3752, SDR 338) was collected at Bitogan, Sigaboy,
Mindanao in 1930 (no date given). It was first identified as
Spizaétus limnacetus, then reidentified as Aviceda jerdoni mag-
nirostris. It is, in fact, another specimen of Pernis celebensis
steerei. I do not know that Hachisuka ever specifically referred
to this specimen in print. As for the rest of the Hirazawa
series of “lewcopais,’ I have been unable to learn of their
present whereabouts, if, indeed, the Hirazawa collection sur-
vived World War II. One can but assume that all five speci-
mens were of one species, in which case they were not Aviceda
but Pernis, as illustrated by the example seen.

Rand and Rabor (1960: 374) stated that their one Samar
specimen of Aviceda jerdoni magnirostris, a male, had a defi-
nitely shorter wing than two females and an unsexed specimen
from Mindanao. There probably is no geographic significance
in this difference. The Samar male has a flattened wing of
294 mm; measurements of a Mindanao series run as follows:
male 299, 311; female 298, 299, 308, 314, 321, 324. It
would appear that males average slightly smaller than females.
There is a spread of 26 mm between the smallest and largest
females measured, and only 12 mm between the two Mindanao
males. The Samar male would undoubtedly fall within the range
of variation in size of a larger series of Mindanao males.

The unsexed specimen from Mindanao mentioned by Rand
and Rabor is readily identified as a female on the basis of
plumage color. Although this is not suggested in the descrip-
tion by Delacour and Mayr (1946: 40), Aviceda jerdoni is

6 Postilla Yale Peabody Museum No. 51

markedly sexually dimorphic in color. The difference is well
described for A. 7. jerdoni by Stuart Baker (1928: 174),
but the dimorphism of magnirostris does not appear to have
been correctly described. The descriptions of the “nearly adult
female” and ‘tadult male” given by McGregor (1909: 236-237)
and Hachisuka (1934: 34) are copied directly from Sharpe
(1874: 356), with and without credit, respectively. But, as
Sharpe himself pointed out (1893: 555-556), the “nearly adult
female” proved to have been a victim of a labeling error, and
was not in fact from the Philippines at all. The figure in
Delacour and Mayr (1946: 41) may be a composite of the
two specimens in the American Museum of Natural History,
as it partakes of the characteristics of both sexes. In males
the upper breast is more or less clear gray; in females this
area is mixed with rufous. The broad brown bars of the pos-
terior underparts are a darker, less rufescent brown in males.
The cheeks of males are gray with black shaft-streaks: this
area in females is buffy or pale rufous with darker-brown shaft-
streaks. Females are much more rufescent dorsally; this is
most striking on the crown and nape, which have bright rufous
feathers with darker centers. In males the crown is virtually
black, with narrow, paler feather-edges, and the nape nearly
lacks rufescence. On the outer rectrix of males, the dark bars
of the outer web are approximately opposite those of the inner
web, whereas in females the dark bars are offset from one an-
other, sometimes to the degree that (as in C.N.H.M. 184011)
the dark bars of the outer web are opposite the light bars of
the inner web. Incidentally, the figure in Delacour and Mayr
(1946: 41) shows the outer rectrices with only a narrow pale
tipping, whereas there is actually a rather broad pale terminal
band (20-28 mm).

The four Mindanao females now before me are rather uni-
form in dorsal color, making allowances for relative wear and
museum age. Only two males are at hand, one of which is the
Samar bird reported by Rand and Rabor (1960: 374). This
specimen differs slightly in color from a single Mindanao male
in being of a darker, colder tone of brown dorsally, lacking
almost completely the rufescent tinge on the nape of the latter
bird. The Samar specimen also has dark-gray rather than

-_

June 27, 1961 The Crested Lizard Hawk

dark-brown edgings to the black feathers in the center of the
crown. Further material must be compared before any geo-
graphical significance can be attached to these differences.
The Crested Lizard Hawk is a rare bird in the Philippines.
Dr. Rabor writes me (letter of September 17, 1960) that the
Samar male is the only specimen he has taken in 26 years of
collecting. I have attempted a census of Philippine specimens
of Aviceda jerdoni, and have managed to locate the following:

British Museum (Natural History)
6 “Philippines,” Cuming coll. (type of magnirostris).

? Davao, Mindanao, February, 1905.

American Museum of Natural History
2 Kalusian, Palawan, October 5, 1887 (type of leuco-
pias).
Davao, Mindanao, May 17, 1889.
2 Davao, Mindanao, September 8, 1903.

Chicago Natural History Museum

[ 2] Culamen, Caburan, Davao, Mindanao, January 25,
1947.

2 Mt. Busaw, Caburan, Davao, Mindanao, January 28,
1947.

a

2 Mati, Digos, Davao, Mindanao, February 9, 194
Mt. Capoto-an, Samar, May 9, 1957.

Os

University Zoological Museum, Copenhagen, Denmark
2 Talacogan, Upper Agusan, Mindanao, March 8, 1952.
2 Pulangi River, Bukidnon, Mindanao, November 20,
1952.

National Museum of the Philippines, Manila
6 Limot, Mati, Davao, Mindanao, April 10, 1949.
[2 Bunauan, Agusan, Mindanao, December 17, 1909: Me-
Gregor, 1910. This specimen was lost with the destruction of
the Museum in 1945. ]

8 Postilla Yale Peabody Museum No. 51

These thirteen specimens (one lost) are the only examples
of Aviceda to have been collected in the Philippines to my
present knowledge. I have examined the seven now in museums
in the United States.

Lint and Stott (1948: 42) published sight records of this
species from three localities in Zamboanga, western Mindanao,
all in October, 1945. In view of the rarity of Aviceda jerdoni,
however, the possibility must be considered that some similar
species was involved (several Philippine raptors have similar
patterns), particularly as a “bare tree within half a mile of the
center of the city of Zamboanga” seems a most unlikely situa-
tion for a species usually described as a shy resident of original
forests. The species has not been collected in Zamboanga.

It is obvious that the principal Philippine range of this
rare bird is central Mindanao; only two specimens are known
to have been collected elsewhere. It is Just possible that a recog-
nizable subspecies may inhabit Samar, but the status of the
Palawan population cannot be determined from the single
specimen known, which appears to be a freak individual.

ACKNOWLEDGEMENTS

Specimens were borrowed through the kind cooperation of
Dr. A. L. Rand of the Chicago Natural History Museum,
Dr. Dean Amadon of the American Museum of Natural His-
tory, and Mr. Robert W. Dickerman of the University of
Minnesota. Mr. Dickerman and Dr. Dwain Warner were instru-
mental in arranging the exchange of a portion of the Bourns
and Worcester Philippine collection to the Carnegie Museum.
Specimens were examined at the Peabody Museum of Natural
History, Yale University, through the courtesy of Drs. S.
Dillon Ripley and Philip S. Humphrey. Data on specimens in
the British Museum (Natural History) were supplied by Mr.
R. W. Sims, and information about Cuming’s collecting local-
ities by Mr. 8S. P. Dance. Data on specimens in the National
Museum of the Philippines were sent by Mr. Telesforo Oane,
and on those in the University Zoological Museum, Copen-
hagen, by Mr. Arne N¢rrevang.

June 27, 1961 The Crested Lizard Hawk 9

SUMMARY

The Crested Lizard Hawk, Aviceda jerdoni, is known from
the Philippines by thirteen or more specimens of the endemic
subspecies 4. j. magnirostris. The type locality “Luzon” gen-
erally given for this subspecies is almost certainly erroneous,
and Davao, Mindanao is substituted. A second supposed race,
A. j. leucopias, is based on a single specimen from Palawan
which is probably an aberrant individual. Other records of
“leucopias”’ from Romblon, Samar, and Mindanao are based
on misidentified specimens. All but two specimens of known
origin of magnirostris are from Mindanao; these are the
type of “leucopias” from Palawan and one from Samar which
may represent a recognizable race. Sexual dimorphism in
magnirostris 1s described for the first time, and all known
specimens listed.

10 Postilla Yale Peabody Museum No. 51

LITERATURE CITED

Baker, E. C. Stuart. 1928. Fauna of British India. Birds 5. Taylor &
Francis, London. 469 p.

Bourns, F. S. and D. C. Worcester. 1894. Preliminary notes on the birds
and mammals collected by the Menage scientific expedition to the
Philippine Islands. Occ. Pap. Minn. Acad. Nat. Sci. 1: 1-64

Delacour, J. and E. Mayr. 1946. Birds of the Philippines. Macmillan Co.,
New York. 309 p.

Hachisuka, M. 1934. The birds of the Philippine Islands 2 (4). Witherby,
London. 256 p.

Hachisuka, M. 1941. Further contributions to the ornithology of the
Philippines. Tori 11: 61-89.

Lint, Kk. C. and Kk. Stott, Jr. 1948. Notes on birds of the Philippines. Auk
65: 41-46.

McGregor, R. C. 1909. A manual of Philippine birds. Bureau of Science,
Manila. 769 p.

McGregor, R. C. 1910. Additional notes on birds from northern Mind-
anao, Philippine Islands. Phil. J. Sci. 5, see. D: 197.

Peters, J. L. 1931. Check-list of birds of the world 1. Harvard Uniy. Press,
Cambridge. 345 p.

Rand, A. L. 1951. Birds of Negros Island. Fieldiana: Zool. 31: 571-596.

Rand, A. I... and D. S. Rabor. 1960. Birds of the Philippine Islands:
Siquijor, Mount Malindang, Bohol, and Samar. Fieldiana: Zool. 35:
221-441.

Sharpe, R. B. 1874. Catalogue of the Accipitres, or diurnal birds of prey,
in the collection of the British Museum. British Museum, London. 480 p.

Sharpe, R. B. 1893. Bornean notes. Ibis [85]: 546-563.

Swann, H. k. and A. Wetmore. 1936. A monograph of the birds of prey
2 (13): 257-352.

Whitehead, J. 1890. Notes on the birds of Palawan. Ibis [32]: 38-61.

5
Rig

aes

pila

YALE PEABODY MUSEUM

oF NaTuRAL History

Number 52 June 28, 1961 New Haven, Conn.

AEGEAN BIRD NOTES I

DESCRIPTIONS OF NEW SUBSPECIES
FROM TURKEY

GeEorRGE E. Watson

Preasopy Museum, Yate UNIversITy

INTRODUCTION

In the course of gathering material for a study of the Aegean
avifauna, I was able to collect during March and April, 1960,
in south and west Asia Minor. The areas visited extended from
Tarsus, Icel, and Pozanti, Seyhan, along the south coast to
Mugla and up the west coast to Bursa. Some of the regions
visited have been little collected during the early spring. This
fresh material revealed that the populations of some species
in this area were markedly different from other known popula-
tions nearby. Therefore, the following descriptions of new
subspecies are presented. The distinctness of some of these
populations suggests that biogeographic studies of the Asia
Minor avifauna may be rewarding in terms of elucidating east-
ern Mediterranean late Pleistocene refugia (cf Kosswig, 1955,
Syst. Zool. 4: 49-73, 96).

2 Postilla Yale Peabody Museum No. 52

Prinia gracilis

The population of the Streaked Prinia inhabiting south
coastal Asia Minor was found to be consistently different from
its nearest geographical relatives and warranted description as
a new subspecies. This difference was suggested by the infor-
mation available to Hartert (1910, Vég. der pal. Fauna: 609)
and Zedlitz (1911, Journ. f. Orn. 59: 610) but overlooked by
them. I therefore propose the name:

Prinia gracilis akyildzi subsp. nov.

Type: Adult 6 (Y.P.M. No. 59196) collected in Antalya,
Turkey, March 31, 1960, by George E. Watson.

Diacnosis: Nearest to P. g. deltae and P. g. paldstinae but
darker above, with even broader dark-brown shaft-streaks on
the back: underparts brighter and more suffused with buff.
This character is most noticeable in fresh unworn plumage,
but even in late March south Asia Minor birds are separable
from spring specimens from Palestine, Syria, and the Nile of
Egypt. Brown shaft-streaks are present on the feathers at the
sides of the upper breast and form an indistinct necklace.
On some specimens shaft-streaks are also present on the
flanks. This character clearly separates this race from all
others of the species except P. g. deltae in which side and
breast-streaking is present in some specimens. Tail with less
well-defined and narrower black subterminal bars than either
P. g. paldstinae or P. g. deltae; tip buffish not whitish as in
P. g. paldstinae. In the reduced width and lack of definition
of the subterminal bars this form resembles P. g. trakensis, but
the back-streaking is far more emphasized and the general
color tone is darker, more brown. The culmen averages mark-
edly shorter than in any of the other three races.

MerasurEeMENts: Type, wing, 44; tail, 66; culmen (from
skull), 10.6 mm; weight, 6.5 gr. Six other specimens from An-
talya, Tarsus, and Adana, south Turkey: 2 é é wing, 42,

42; tail, 65, 60; bill, 11.2, 11 mm; weight, 6.7 gr; 4 2 2 wing,

June 28, 1961 Aegean Bird Notes from Turkey 3

Mpls. 6251 tall, 99, 009,100, 05 > bill, 1025, 11, Fl, 12 mm;

weight, 6.7, 6.5, 6.5 gr.

fon)

CoLor or sorr parts: Iris light red-brown, light-brown, or
cream; bill flesh or upper mandible brown, lower mandible
cream or yellow-cream; feet buffish-flesh or flesh. Apparently
the bill color in this species changes to black during the
breeding season.

The type series consists of six specimens, two males and
four females collected between March 2 and March 381, 1960,
at Antalya, and Tarsus, Icel, in south Turkey. In addition,
a single male from the American Museum collected in nearby
Adana, January 1, 1879, was also examined, A male collected
on March 3 is renewing feathers on the center throat. None
of the specimens had gonads enlarged for breeding, but males
were calling loudly from exposed perches.

Rance: This race is confined to the coastal fringe of south-
ern Turkey. It extends on the west as far as Antalya and on
the east to Adana and probably as far as the Arsuz plain
where Kizil Dag (Mount Amanus) may form a barrier between
this race and P. g. paldstinae, The species has been seen a
little farther north at Osmaniye (Danford, 1880, Ibis 22:
eek no American Museum specimens from Kara Suleiman,

Syria” (=—Karasiileyman, Maras, Turkey?), approach this
fae in darker-brown upperparts, broad shaft-streaks, short
bill, and reduced black subterminal tail bar; the underparts,
however, are very pale, as in P. g. trakensis.

Remarks: This population, the darkest, brownest, and most
heavily streaked of the species, is also the northernmost ;
Antalya is the farthest west that the species has been found in
Asia Minor. The population, therefore, comprises the end
points of east-west and south-north clines of increasing color
saturation from India and Arabia. Another saturated brown
population, P. g. deltae, occurs along the lower Nile and toward
Suez. Southern Turkey is also the western end point of the pop-
ulations which have narrow or indistinct subterminal tail bars.
These range from the Brahmaputra westward across India,

4, Postilla Yale Peabody Museum No. 52

Pakistan, southern Afghanistan, southern Iran, southeast Ara-
bia, Iraq, northern Syria (?) to south coastal Asia Minor.

This new race is named for Zubeyir Akyildiz who shared in
my explorations of southern Turkey.

In studying the Asia Minor population of Prinia gracilis,
I had the opportunity to examine some specimens from Arabia,
including the type of P. g. anguste Ripley from Bahrein
Island and part of the type series of P. g. carpenteri de
Schauensee and Ripley from Oman. P. g. anguste, based on a
single worn July specimen, is said to differ from P. g. hufufae,
the population of the adjacent mainland, in being darker and
more brownish-gray with narrower shaft-streaks. The only
specimens of P. g. hufufae available to me are five freshly
molted November birds of the Cox-Cheeseman type series in
the American Museum of Natural History. I can find no char-
acter in the type of P. g. anguste compared with this series
which I would not attribute to the birds being collected at
different times of the year. Meinertzhagen (1954, Birds of
Arabia: 219) states that a specimen he collected on Bahrein
is identical with mainland birds.

P. g. carpenteri from Muscat, Oman, is a well-marked race.
It differs sharply from P. g. hufufae to the north in its nar-
rower and less distinct subterminal tail bars; from P. g. ye-
menensis to the west in its more distinctly cross-rayed tail
and more finely pencilled head and back-streaking; and from
P. g. trakensis from Iraq and southwestern Iran and P. g.
lepida from southern Iran, Afghanistan, and northwestern In-
dia in its darker back coloration and more prominent sub-
terminal tail bars. I cannot agree with Meinertzhagen (ibid:
220) and lump Oman specimens with P. g. lepida.

II

Erithacus rubecula

The Robin varies clinally and occasionally markedly over
its continental Palearctic range. As shown by Lack (1946,
1947, Bull. Brit. Orn. Cl. 66: 55-65; 67: 51-54: and 1951,

June 28,1961 Aegean Bird Notes from Turkey 5

Ibis 93: 629-30), there is a widespread north and central
European form (E. r. rubecula) with olive-brown upperparts,
brownish tail and upper tail coverts, and reddish-orange throat
and breast. To the east, populations occur in Caucasia (EF. r.
caucasicus) and Iran (EF. r. hyrcanus) with darker backs
and breasts and with a strong rufous tinge to the sides of
the tail and upper tail coverts. In the Urals there breeds
a somewhat reddish-tailed, but strikingly pale gray-backed
form (E. r. tataricus). In Spain and Italy Robins tend
to be darker, approaching the north African form (E. r.
witherbyi). An isolated slightly grayish-backed population
(E. r. atlas) is found in northwest Morocco. On the other
hand, a different clinal tendency starts in Yugoslavia. Ex-
tending through southern Greece and across Bulgaria to Asia
Minor, Robins are markedly gray above and paler below,
but even Turkish specimens have the brownish tail of the

European populations.

{ e Ulu Dag

>

BS Turkey

?

HHS

e@ Boz Das
de> e@ Kara Suleyman
SoA Si
J

@ Adana

Ak Dage Antalya Se

SF
ef Lo TN) y
F f f pe
= mae Sen \ = Kizil Dae
z )
/

So Rhodes ~~

f

6 Postilla Yale Peabody Museum No. 52

Lack refrained from giving this southeast population a
name but noted its existence, especially pronounced in Bul-
garia, although he later (1951, Ibis 93: 629-630) believed
that the grayishness was merely due to wear rather than valid
geographic variation. Stresemann (1920, Avifauna Macedo-
nica: 179-180) also noted gray spring specimens in Macedonia.
Examples of the gray form have been found near Bursa in
northwest Turkey by Mrs. Scott-Neuhauser (1948, Senchen-
bergiana 28: 177), but she collected what appears to be E. r.
caucasicus in Rize. Koller (1948, Senchkenbergiana 28: 177)
collected a dark specimen (“H. r. xanthothoraz” = E. r.
caucasicus on passage towards the Aegean?) in Bolu.

The lack of a nomenclaturally recognized population breed-
ing in Asia Minor has led to confusion about the identity and
origin of the wintering populations of the Aegean (e.g. what
to do with E. r. vanthothorax). The very marked gray char-
acter of Asia Minor birds and the reddish tail and upper tail
coverts of Caucasian and other eastern populations makes
them both equally separable from the brownish-backed and
tailed Kuropean populations. The nomenclatural recognition
of this Balkan gray cline will perhaps lead to more meaningful
discussion of geographic variation in the eastern part of the
species range and will certainly make the nomenclature better
balanced, since all other clinal trends are named or overnamed.
Therefore, the most extreme population of this gray cline is
given the name:

Erithacus rubecula baleanicus subsp. nov.

Type: Adult 6 (Y.P.M. No. 59198) collected on Boz Dag,
Odemis, Izmir in western Turkey at 4,200 feet on April 20,
1960, by George E. Watson.

Diacnosts: Differs sharply from all other subspecies in its
concolorous olive-gray back and rump, brownish flanks, pale-
orange underparts, and grayish brown-edged rectrices. The
grayish upper tail coverts are most diagnostic of this form.
The population tends toward nominate FE. r. rubecula in north-
ern Yugoslavia and is

grayest in north and west Asia Minor.

June 28,1961 Aegean Bird Notes from Turkey 4

No difference from the nominate form in length of wing, tail,
and culmen, or weight.

MEASUREMENTS: Type, wing, 70; tail, 56; culmen, 15 mm;
weight, 17.5 gr; 2 ¢ $ wing, 71, 71; tail, 59, 58; culmen, 14,
14.5 mm; weight, 18.3, 18.7 gr; 4 2 2 wing, 70, 68, 71.5,
leo tail. 50, 56, 56. 58> culmen, 15.2, 14.2, 13.8, 14 mm:
weight, 22 (laying), 21.5 (layimg), 17, 17.

The type series consists of seven specimens, one breeding
male and two laying females from Boz Dag (March 20, 21),
and four wintering specimens from Antalya, south Turkey,
and the Aegean islands of Icaria and Andros (February and
March). In addition, nine breeding and wintering specimens
were examined from Bosnia and Herzegovina.

Rance: Breeds from northern Yugoslavia south through the
Balkans to southernmost peninsular Greece (rare in the Pelo-
ponnesus) and across Bulgaria to northern and western but
apparently not southern Asia Minor. At the eastern limits of
its range in Asia Minor it may meet a reddish-tailed popula-
tion of FE. r. caucasicus probably near Samsun or 'Trabzond,
but breeding specimens are lacking from this area. In the
southern part of its range, breeding is confined to the moun-
tains, both in conifer (Abies) and deciduous forests and open
woods. In winter, individuals move to lower altitudes, and some
birds may spread into the Aegean islands (Andros, Tearia )
and to the south coast of Turkey (Antalya).

Remarks: It is possible that a discontinuity exists between
the birds of southern Bulgaria and those of northwest Asia
Minor. Nothing would be gained, however, in further sepa-
rating the cline on this slight difference. In the Aegean is-
lands and in the southern Balkan Peninsula EF. r. rubecula
is also found wintering. See below for a discussion of “FE. r.
vanthothorax.”

This Balkan cline probably results from the northwestward
post-glacial expansion of an isolated forest population from
the Mediterranean “pluvial’” refuge into which the birds re-

olacial advance.

treated during the Wirm g

8 Postilla Yale Peabody Museum No. 52

I propose this new subspecies fully aware of the “tyranny
of subspecific names” (Lack 1946, Bull. Brit. Orn. Cl. 66: 63)
in the Robin. Partially following Dr. Lack’s admonition, I
have given this population the most appropriate geographic

name available for the entire range.

Many authors have questioned the validity of E. r. vantho-
thorax Salvadori and Festa (1913, Bull. Mus. Zool. Anat.
Comp. Torino 28: 15). This name is based on a series of five
birds collected during February and March in Rhodes in the
south Aegean. The characters cited in the description clearly
mark the birds as belonging to a reddish-tailed southeastern
form. Robins are common on Rhodes during the winter. Most
had left but I located a few on March 30, 1959; following that
date no more were seen. A Rhodian forest guard, who was
well aware of the identity of the bird, told me that no Robins
would be seen again until October. The type series in Turin
is too old and dirty to distinguish much color definition, but
on the basis of the type description and the following measure-
ments (personally taken) including the type, the series belongs
with the shorter-billed Caucasian birds, rather than with the
Iranian form: 1 ¢ (type) wing, 72; tail, 51; culmen, 14.5
mm; 2 22 wing, 71, 70. *tail, 53.5,)oa00- culmen, los elas
mm; 1 o wing, 69; tail, 53; culmen, 14.5 mm; I therefore en-
dorse Vaurie’s (1958, Birds Pal. Fauna: 376) decision in syn-
onomizing the name FE. r. wanthothoraax with E. r. caucasicus.

JUL
Prunella modularis

A very dark gray population of Hedge Sparrow breeds in
the fir forest up to the tree limit on Ulu Dag above Bursa
in northwest Turkey. The two males collected necessitated
comparison with series from nearby breeding localities. This
comparison revealed that the Asia Minor birds differ sharply
from nearby Balkan and Caucasian and Iranian birds and

June 28, 1961 Aegean Bird Notes from Turkey 9

further suggested that the Balkan population P. m. meinertz-
hagent Harrison and Pateff is merely the extremely dark end
point on a continental cline of increasing grayness.

There are two well-marked groups of populations of Hedge
Sparrows in southeast Kurope and southwest Asia. The gray-
ish nominate group modularis, with black back-spotting, breeds
over most of Europe down to southern Bulgaria and northern
Greece (Peus, 1957, Mitt. Zool. Mus. Berlin 33: 275). The
browner obscura form, with brown back-spotting breeds in the
Caucasus and northern Iran.

The Balkan population metnertzhageni is an extreme form
of the modularis group. The gray of the underparts is darker
and more extensive than in the nominate race; whitish flecking
is reduced or absent. Above, the bird is a trifle grayer with
larger darker spots. Most wintering Hedge Sparrows from
the Aegean are separable into two categories. Most of those
collected in the north, in Epirus and Macedonia and in the
Asia Minor islands, belong to the metnertzhageni population,
and most of those from the south and in the Cyclades are
northern European P. m. modularis. Iranian and Caucasian
birds differ from each other mainly in degree. Caucasian birds
are darker and the gray of the underparts is more extensive.

On the other hand, the two breeding males from Ulu Dag
show some blending of characters but differ from both these
forms in several characters and I therefore propose the name:

Prunella modularis euxina subsp. nov.

Type: Adult ¢ (Y.P.M. No. 59297) collected on Ulu Dag
(= Asiatic Mount Olympus of some authors), Bursa, north-
west Turkey, April 29, 1960, by George E. Watson.

Dracnosts: Head light ashy-gray heavily streaked with
brown: superciliary buffy-gray; back light reddish-brown with
darker-brown spots; rump and upper tail coverts brownish-
gray; underparts dark ashy-gray with white on the center of
the abdomen, some of the lower breast feathers tipped with
white, sides of upper breast washed with brown; flanks mod-
erately streaked with the same brown as the back; under tail

10 Postilla Yale Peabody Museum No. 52

coverts gray-brown. In dorsal coloration, this form is closest
to P. m. obscura in having brown not black spotting, but it
differs from eastern birds mainly in its grayer and darker
underparts and the lesser extent of white on the abdomen. It is
even darker gray below than the darkest individuals of P. m.
modularis from the Balkans. From both subspecies it differs
in having gray, not brown rump and upper tail coverts and in
having the flanks much less strongly marked with brown.

P.m. modularis Pom, euxina P.m. obscura
Crown gray with brown light ashy-gray gray-brown with
streaks with light- brown streaks
brown streaks
Superciliary dark-gray buffy-gray gray-buff
Dorsal spotting black brown brown
Rump andupper — brown gray brown

tail coverts

Underparts gray dark-gray light brownish-
gray
Flanks heavy brown restricted heavy brown
streaks streaking streaks
Under tail dark brownish- light-gray brownish-gray
coverts gray

MerasurEMENTs: Type and one other é: wing, 69, 68.5;
tail, 57, 56; culmen (from skull), 13, 14 mm; weight, 22, 21 gr.

Rance: Mountainous fir forests of northern Asia Minor.
Eastern limits of range unknown but probably meets P. m.
obscura in Transcaucasia or Turkish Armenia. Probably some-
what migratory (wing length and formula same as other mi-
gratory populations) but winter range unknown.

Remarks: Although there is some introgression of charac-
ters of the eastern populations into this north Asia Minor
population, the much darker-gray underparts and gray rump
suggest that it is on a separate evolutionary line, one that has
perhaps developed in situ in the thick fir forests of the rainy
north coast. The fact that the Bursa population is also so
sharply different in several characters from the Balkan form
further suggests that this population is isolated from the

June 28,1961 Aegean Bird Notes from Turkey ig

European birds as well. And in fact there are no high conif-
erous forests in European Turkey. The Ulu Dag population
therefore constitutes the end point of an Asia Minor cline
running westward from the Caucasus.

In habitat the north Asia Minor population differs some-
what from northern and especially western European birds.
It is found confined to the fir forest and tree line scrub in
Turkey, but the same species is a bird of deciduous gardens
and hedgerows in western Europe, and also open conifer woods
in north central Europe. Northern birds wintering in the
Aegean are usually to be found in lowland macchia or open
woods undergrowth.

A single female collected March 7, 1960, above Tarsus in
the foothills of the Cilician Taurus Mountains is clearly a
wintering example of the nominate race from northern Europe.
It is impossible to separate from the paler winter specimens
from the Aegean. Danforth also collected winter P. m. modu-
laris m the Taurus in 1879 (Cat. Birds Brit. Mus. 7: 652).
Apparently, only light-colored migrants with white-flecked
breasts were collected in northern Asia Minor by Kummer-
lowe and Niethammer (1935, Journ. f. Orn. 83: 40), Rossner
(19384: Akad. Wiss. Wien math-natur. Klasse Sitz. 1944: 307),
and Mrs. Scott-Neuhauser (1948, Senchkenbergiana 28: 178),
although the first pair of coliectors undoubtedly earlier ob-
served breeding birds. The new subspecific name is derived
from the classical epithet for the present-day Black Sea which
marks the northern limit of this population.

IV
Montifringilla nivalis

Three well-marked groups of populations of Snow Finch
occur in the Palearctic. The nivalis group of southern Kurope
has brown upperparts and a gray head; the variable alpicola
group of central Asia has a brown head and back; and the
henrict group of Tibet is uniform brown above but washed
with gray-brown below. Nine specimens from south Asia Minor
belong to the alpicola group, in which there are four fairly

Le Postilla Yale Peabody Museum No. 52

well-marked races. M. n. alpicola has an extensive range from
the Caucasian Mountains south into Turkish Armenia, where
the exact western limits of the range are unknown, and east-
ward across northern Iran towards Afghanistan and into the
Tian Shan range. In southern Iran, in the Zagros, a markedly
paler race M. n. gaddi occurs; in Mongolia another pale race
M. n. groum-grzimaili is found. These three races are long-
billed. A very pale-sandy and short-billed race M. n. kwen-
lunensis occurs in the Kun Lun and Astin Tagh ranges. The
south Asia Minor population, which is isolated from these four
populations, differs in being lighter in color and showing char-
acters which approach the European nivalis group. For this
isolated southern population I propose the name:

Montifringilla nivalis fahrettini subsp. nov.

Tyre: Adult ¢ (Y.P.M. No. 59445) collected at 6,400 feet
on Ak Dag, Kas, Antalya, southern Turkey on March 25,
1960, by George E. Watson.

Diacnosis: Closest to M. n. gaddi but much paler and grayer
on the back, less reddish-tan and with a pronounced grayish
tinge on the crown, whereas Zagros birds have the head essen-
tially the same color as the back. Wing and bill in the spring
adult much shorter than in the Zagros population at the same
season. Differs from M. n. alpicola in its markedly lighter and
less brown upperparts and in having a shorter bill and wing.
Differs from M. n. nivalis in much lighter upperparts, less
distinctly gray head, shorter wing, and lighter weight.

MEAsurEMENTs: Type, wing, 112; tail, 68; bill from skull,
14.5 mm; weight, 33 gr, 4¢ 6 wing, 111, 117, 115, 115,
(mean of 5: 114); tail, 68, 71, 69, 70 (69); bill, 13.5, 14,
14.5, 14.1 (14.1); weight, 31, 33, 34, 35.5 (33.3) Alon 9
wing, LVO, 111, 116; 106; CLIT .3)) 5 tanl, G5, 62, 75, 640066):
bill, 14, 14.5, 14, 14 (14.1); weight, 32.5, 32.5, 33, 28.5
(31.6).

M.n. nivalis (Greece), 4 3 ¢ wing, 116-120 (119) ; tail, 67-72
(69.5) ; bill, 13.5-14 (13.6) ; weight, 37-40 (38.1); 1 2 wing,

116; tail, 64; bill, 13.5 mm; weight, 37 gr.

June 28,1961 Aegean Bird Notes from Turkey 13
M. n. alpicola, 10 3 ¢ wing, 117-1
66-71 (68.9); 10 3 ¢ bill, 16-17.5 (
len Cll?) : 32 Potail, 62; 66, 72 ¢
(15.88).

T2A (USS): 7% 1 vd thal,
]

M.n. gaddi, 19 3 ¢ wing, 116-126 (119.3); 18 ¢ ¢ tail, 68-
76 (71.5); 18 ¢ 3 bill, 15-17.5 (16.2); 4 2 2 wing, 112-116
mite.) = ibe? 2 tail, 65-71 (68.5)5,5 2 2 bill, 15al6 (15:6).
Measurements of M. n. alpicola and M. n. gaddi are taken
from Vaurie (1949, Amer. Mus. Novitates 1406: 29).

Summer collected specimens of this species tend to have
shorter and more worn bills than winter birds (Stegmann 1982,
Journ. f. Orn. SO: 99). This is perhaps related to a change in
diet from insects picked off snow to seeds picked and scratched
from bare rocky soil. It should be pointed out, however, that
late March M. n. fahrettint were compared with late March
M. n. gaddi. The bills were found to be markedly longer and
more attenuated in the Zagros birds. A young January speci-
men from the north Zagros does have a short bill, but not as
short as that of the longest-billed Asia Minor bird. Further-
more, Vaurie’s measurements were taken on birds collected
at all times of the year, including January and February as
well as spring and summer. His shortest measurements for
M.n. gaddi and M. n. alpicola do not overlap at all the longest
measurements of the Asia Minor population. Taking the bill
measurements of five examples of each of the four populations
(data from Vaurie tbid.: 28) and of the two populations
M. n. gaddi and M, n. fahrettini and comparing them gives
the following variance ratio table:

df Ss ms VR

Between

AE ROMS) wean coco bomb obudT 3 25.8 8.6 21.5

7 ARON Slo aca dono oaumoadac 1 ied) fo) 46.6
Within

Al: (HOIS Gh Sbaccvoogeeeds c 16 6.5

ZOE OUP Simmeters sepals oicierarcl ic 8 14 al its
Total

AST OUP S) ton 2 5) eis eiete sissies ene 19 2.27

POU Stecrta eeycckake serdocis. tote 9 9.5

14 Postilla Yale Peabody Museum No. 52

Had all the measurements available for M. n. alpicola and
M. n. gaddi populations been taken during the same season,
the statistical results would have been even more striking.
The type series consists of nine birds collected on March 25
and 29 at from 6,000 to 7,300 feet on Ak, Kohu, and Manearli
Daglari near Elmali, Antalya, in south Turkey. The gonads
were little enlarged and the birds were still feeding in flocks.

Rance: Occurs on the highest mountain tops in the Bey
and ‘Taurus mountain ranges of south Turkey and probably
on Mounts Lebanon and Hermon; resident but descending to
about 6,000 feet in winter.

Remarks: The only other record of this species from south
Turkey is that of Danford (1878, Ibis 20: 23) who included
the species in his list ‘twith some hesitation”? on the basis of a
sight record on Anas Dag in the Taurus north of Adana. Snow
Finches were observed by Tristam (1868, [bis 10: 208) on the
peaks of Mounts Lebanon and Hermon. I failed to find the
species on Boz Dag, Izmir (ca. 7,100 feet), or Ulu Dag (Burs:
8,343 feet) in mid- and late April. It is doubtful whether there
are any other mountains in western and northwestern Asia
Minor high enough to support breeding populations of Snow
Finches, so that M. n. fahrettini is separated from the Euro-
pean populations of the species. On the other hand, it may
well occur through the anti-Taurus and in Kurdestan and
therefore be continuous with the Iranian populations, but it
has not yet been found in southeastern Turkey or northern
Iraq.

Five adult and two juvenal specimens of Snow Finch, M. n.
nivalis, from Greece constitute only the second record of this
species from that country verified by specimens. Reisor (1895,
Ornis Bal. 3: 23) found the bird on Vardusia, Makatsch (1950,
Die Vogelvelt Macedoniens: 117) probably saw the bird on
Olympus, and Flach (in litt.) recorded Snow Finches on Par-
nassos. The specimens are from Vardusia and Parnassos. The
species was not found on any of the Peloponnesian mountain
tops in spite of repeated searches.

In their distribution the populations of Snow Finches show
the type of disjunction common to glacial relicts. It is possible

June 28,1961 Aegean Bird Notes from Turkey iS

that when the climate of the Eastern Mediterranean was more
rigorous during the retreat of the Wiirm glaciation, many of
the mountains throughout the Aegean basin may have har-
bored populations of these Snow Finches. As the climate grew
warmer and the snow disappeared from all but the highest
peaks, extensive suitable habitats disappeared and the disjunct
distribution of today came about. The south Turkey popula-
tion, which shares the tendency toward a grayish head and a
short bill with the European population, is evidence that such
must have been the past history of the species in the eastern
Mediterranean.

The new subspecies is named for Fahrettin Ozgecil of the
Turkish Forest Department, who accompanied me on my trav-
els in the western sector of his country.

ACKNOWLEDGEMENTS

I am grateful to the following persons for aid in arranging
this trip and for hospitality received while making the collec-
tion in Turkey: Ambassador Ellis O. Briggs, Athens; William
A. Helseth, Ankara; Orhan Sagnak, forest director of Icel;
Zubeyir Akyildiz, forest engineer in Ankara; and Fahrettin
Ozgecil, chief forest engineer in Antalya. Without their gen-
erous aid the trip would not have been so fruitful. I have
borrowed specimens from the American Museum of Natural
History and have discussed taxonomic points with Dr. Charles
Vaurie. Dr. S. Dillon Ripley has examined some of the material
with me. This research was conducted while I was a National
Science Foundation predoctoral fellow.

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YALE PEABODY MUSEUM

or NATURAL History

Number 53 June 380, 1961 New Haven, Conn.

THE SKULL OF SCIURAVUS NITIDUS, A MIDDLE
EOCENE RODENT

Mary R. Dawson

SmirH CoLiece, NorTHAMPTON, Mass.

Primitive rodents of the family Sciuravidae are known pri-
marily from the Eocene of western North America, with the
genus Scituravus itself ranging from late early Eocene (Gazin,
in press) through the late Eocene (Wilson, 1949b, pp. 81-82).
The type specimen of Sciuravus nitidus Marsh, 1871, the type
species, is an incomplete maxilla with M'* from the middle
Eocene Bridger formation of Wyoming. The type specimen is
from the lower Bridger, but specimens referred to S. nitidus
occur throughout the Bridger formation (Wilson, 1938, pp.
130-1382). The skull of §. nitidus was described briefly by
Matthew (1910, pp. 59-60). More recent collections from the
Bridger have yielded more complete specimens of this species
than were available to Matthew. These specimens allow an
amplified description to be made of the skull of S. nitidus,
the only Bridger sciuravid whose essentially complete skull
is known.

ACKNOWLEDGEMENTS

Grateful thanks are given to Dr. C. L. Gazin of the United
States National Museum (U.S.N.M.) for valuable stratigra-
phic information and for permission to study specimens of

bo

Postilla Yale Peabody Museum No. 53

Sciuracus nitidus; to Drs. J. T. Gregory and E. L. Simons
for access to the collections and for facilities for study in the
Yale Peabody Museum (Y.P.M.); to Drs. C. C. Black of the
Carnegie Museum (C.M.) and M. C. McKenna of the American
Museum of Natural History (A.M.N.H.) for the loan of
specimens. Plates I, III, IV, and V, figure 1 are prepared
from photographs furnished by courtesy of the Smithsonian
Institution, and plate V, figure 2 is from a photograph by
J. Howard, Yale Peabody Museum. This study, supported by
National Science Foundation Grant G14254, was completed
while the author was a Research Associate of the Division of
Vertebrate Paleontology, Yale Peabody Museum.

DESCRIPTION

The following description of the skull and lower jaw of
Sciuravus nitidus is based in large part on U.S.N.M. No.
22477, a nearly complete skull with associated lower jaw and
postcranial fragments from near Millersville, Wyoming. The
specimen was collected by C. L. Gazin and F. L. Pearce in
1946. The skull is distorted by some anteroposterior compres-
sion and the left side is pushed forward relative to the right.
Tooth wear indicates that the individual was adult. Other
specimens used are: U.S.N.M. No. 18100, partial skull and
jaws of a young individual having dP*; A.M.N.H. No. 12551,
skull flattened dorsoventrally and jaws figured by Matthew
(1910, figs. 18-15) ; A.M.N.H. Nos. 12531 and 18101, partial
skulls. Less complete specimens include Y.P.M. No. 138458,
C.M. No. 9683, U.S.N.M. Nos. 17697 and 17700. All specimens
are from the lower Bridger.

Skull. The overall appearance of the skull is that of a rather
generalized, primitive rodent. Postmortem distortion of the
specimens prevents an entirely clear picture of the skull shape.
Thus, the doming of the parietals in U.S.N.M. No. 22477 is
due at least in part to anteroposterior compression, A.M.N.H.
No. 12551 is flattened dorsoventrally, U.S.N.M. No. 18100
and A.M.N.H. No. 13101 are compressed transversely.

Viewed dorsally (pl. I) the skull widens from the anterior
tips of the nasals to the anterior zygomatic roots, has its

June 30,1961 The Skull of Sctwravus Nitidus 3

greatest width across the zygomatic arches, which essentially
parallel one another, and is narrower posterior to the arches.
The nasals are slightly convex dorsally, especially anterodors-
ally, and extend farther forward than the anterior surface of
the incisors. The nasals terminate posteriorly about on a line
with the anterior orbital border. The frontals exhibit a dis-
tinct interorbital constriction, anterior to which there is a
protrusion on each side possibly similar to the “rudiments of
processes” in Pseudocylindrodon (Burke, 1988, p. 260). The
protrusion is less in evidence in A.M.N.H. No. 12551 and
U.S.N.M. No. 18100 than in U.S.N.M. No. 22477. The large
interparietal, shown in Nos. 12551 and 18100, has a convex
anterior margin and widens posteriorly. Lateral to it the
parietal extends to the occipital bone. There is no sagittal
crest. The squamosal is incomplete in all specimens but seems
to have had a narrow dorsal exposure. The occipital bone
extends onto the dorsal skull surface for a very short distance,
and the nuchal crest on that bone forms the posterodorsal
margin of the skull.

Maxillary, jugal, and squamosal components form the
zygoma. The jugal contacts the maxilla and lacrimal anteri-
orly, tapers posteriorly, and terminates bluntly below the
posterior edge of the zygomatic process of the squamosal.

The anterior surface of the anterior zygomatic root 1s
essentially vertical and is in a line anterior to P® (pls. III, IV).
The root is farther forward relative to the cheek teeth than in
Paramys delicatus (Matthew, 1910, fig. 2). The rounded
infraorbital foramen, which is situated in the maxilla medial
to the anterior zygomatic root, is relatively a little larger
than in Aplodontia. From a ventromedial knob of bone on
the anterior root a ridge passes laterad, curves backward and
upward onto the lateral surface of the jugal, and continues
toward the posterior end of that bone, diminishing posteriorly.
As preserved in U.S.N.M. No. 22477, the outer surface of the
zygoma faces laterally above and ventrolaterally below the
ridge. The origin of the masseter muscle seems to be limited
to the zygoma, the “primitive sciuromorph” or “protrogo-
morph” condition. There is no indication that any of the

4 Postilla Yale Peabody Museum No. 58

masseter muscle originated on the rostrum anterior to the
anterior zygomatic root. The ridge on the zygoma probably
marks the dorsal edge of the origin of the masseter lateralis
muscle. That the anterior part of the masseter may have been
strengthened is suggested by the ventromedial knob and the
ridge, which is more distinct anteriorly.

The orbit is moderately good-sized, about comparable in
proportions to that in Cavia but relatively smaller than in
Cynomys. The lacrimal bone forms part of the anteromedial
orbital border. The surface of this bone is lightly pitted where
it extends onto the face anterior to the orbit. The most com-
plete orbital wall occurs in U.S.N.M. No. 18100, although
cracking obscures details. The frontal forms most of the
anteromedial orbital wall. Near the dorsal rim of the orbit
and posterior to the supraorbital protrusion is a small fora-
men. The sphenopalatine foramen (terminology of foramina
follows Hill, 1935) is ventromedian in the orbit, approxi-
mately in line with M*, and is bounded anteriorly, dorsally,
and ventrally by the maxilla. Boundaries of the palatine bone
are not clear, and either this bone or the maxilla borders the
sphenopalatine foramen posteriorly. Several foramina are
present in the maxilla posterolateral to the sphenopalatine
foramen. A plate of bone posterior to the maxilla and frontal
in the orbit seems to be the orbitosphenoid. More posteriorly,
in a line behind M®, the posteromedial orbital wall is depressed
and overhung by more dorsal parts. A dorsoventrally elongated
slit in the depression may be the sphenoidal fissure. The
alisphenoid borders the slit posterolaterally. Anterodorsal to
the slit and partly separated from it by a bony bar is a
rounded gap. It is suggested tentatively that the optic foramen
may be in the gap. Parietal and squamosal form part of the
posterior wall of the orbit.

On the palate (pls. II, III) the elongate incisive foramina
are separated from one another by a median septum and are
within the premaxilla except posteriorly, where they are
bounded by the maxilla. A slight concavity of the maxilla
P#,

occurs anterior to The two rows of teeth essentially

parallel one another. Between them the palate is nearly flat.

June 30,1961 The Skull of Sctwravus Nitidus 5

The palatine extends forward about to a line with the anterior
or middle of M', where the jagged maxilla-palatine suture
crosses the palate transversely. The foramen for the palatine
artery is present approximately in line with the anterior half
of M>. The posterior margin of the palate, about opposite
the middle of M*, has a posteriorly directed median projection,
on each side of which a small flange of bone projects antero-
ventrally below the palatal surface.

The region of the pterygoid plates is incompletely preserved
in all specimens but can be reconstructed in part. A ridge
extends posterad from behind M*, becomes thicker, and leads
into entopterygoid and ectopterygoid plates. A shallow ptery-
goid fossa is enclosed by the plates. The specimens suggest
that the entopterygoid plate lacked a contact with the bulla,
but the ectopterygoid plate continues posteriorly and contacts
the lateral side of the bulla. In Paramys delicatus (Matthew,
1910, fig. 2) the ectopterygoid plate is absent, and the ptery-
goid fossa is less distinct than in S, nitidus. A small foramen
between the plates in A.M.N.H. No. 12551 may represent
the posterior opening of the sphenopterygoid canal. Postero-
lateral to this foramen is a second foramen, probably the
foramen ovale. The ectopterygoid plate forms a bridge across
this foramen, which resembles the foramen ovale in Paradji-
daumo (Wilson, 1949a, pp. 38-39 and fig. 1C) and in Cynomys.
The position of another foramen is indicated by a notch in
the basisphenoid at the anteromedial edge of the tympanic
bulla.

The tympanic bulla in Sciwravus nitidus is ossified but
seems to have been loosely attached. Out of seven otic regions,
in U.S.N.M. Nos. 22477 and 18100 and A.M.N.H. Nos. 12531
and 12551, the bulla is in place in two cases, on one side in
Nos. 22477 and 12581. A suture seems to be traceable between
bulla and periotic, and where the bulla is missing there is no
clear broken area on the periotic. The rounded bulla is not
dissimilar in general appearance to that in Neotoma. The bulla
has a large opening but lacks an elongated bony meatus. The
anterodorsal lip of the opening is thickened. The bulla in
A.M.N.H. No. 12531 may have been incomplete dorsally, and

6 Postilla Yale Peabody Museum No. 53

a process of the squamosal curves down over this possibly
incomplete closure. Viewed ventrally, the bulla has a short
anteromedial protrusion, The bulla does not completely cover
the petrosal, which is exposed medial to the bulla. A somewhat
similar exposure of the petrosal has been reported in Paradji-
daumo (Wilson, 1949a, p. 39). Where absence of the bulla
exposes the periotic, the auditory prominence appears as a
distinct, raised structure. The fenestra vestibuli faces dorso-
laterally and the fenestra cochleae posterolaterally. The mas-
toid process is short and blunt, and the stylomastoid foramen
seems to be represented by a notch medioventral to the process.

Between the bullae the basioccipital exhibits a_ distinct
longitudinal ridge. The elongate jugular foramen is between
basioccipital and periotic, posteromedial to the bulla. The
hypoglossal foramen pierces the condyle posterior and medial
to the jugular foramen.

The occipital surface (pl. V, fig. 1) is convex dorsally. The
nuchal crest leads into two ridges, one on the lateral surface
of the mastoid and the other along the mastoid-exoccipital
contact. The mastoid is exposed on the occipital surface near
the dorsal part of the latter ridge. The short paroccipital
process does not contact the bulla.

Lower jaw. On the lateral surface of the horizontal ramus
(pl. IV) a large mental foramen is present about in line with
the anterior wall of Py. A smaller foramen or foramina occur
shghtly farther posteriorly. The masseteric fossa is bounded
by dorsal and ventral ridges that meet in a line approximately
between M, and M,. The masseteric fossa in Paramys delica-
tus does not extend so far anteriorly (Matthew, 1910, fig. 1).
The nearly complete lower jaw of C.M. No. 9683 shows that
the coronoid process rises steeply and hooks backward toward
the lower condyloid process. The lateral surface of the coro-
noid process is slightly concave for the insertion of the tempor-
alis muscle. On the medial surface of the jaw (pl. V, fig. 2)
a ridge extends posterodorsad from the posterior end of the
row of cheek teeth toward the condyle. The dental foramen
is in the dorsal side of the ridge, approximately in a line below

the posterior edge of the coronoid process. Ventral to the

pee

June 30,1961 The Skull of Sciwravus Nitidus Tf

ridge is the concavity for insertion of the internal pterygoid
muscle; unfortunately, the area for pterygoid insertion is not
completely preserved in any of the specimens. A slight con-
cavity dorsal to the ridge on the condyloid process probably
marks the insertion of the external pterygoid muscle.

Measurements (in millimeters) of U.S.N.M. No. 22477:

Gueatest: leneth ‘ofmskulll ~,.!2..039..1..: 39.4
Anterior width across zygomatic arches .... 21.5
Posterior width across zygomatic arches .... 22.2
ELeIo iL sOCCID UL: Aaee ety Ae Bebe en 12.8
Length diastema, posterior of Ito P® ..... 8.1
Meno iaaiotts ee Ab8 2 ey Capt Gates See 9.6
enetho lett Pz NE.) eras seh oh ee 9.5
Outside depth of lower jaw at M, Reni ech 6.2
DISCUSSION

Sciuravids have been suggested, mostly on the basis of dental
characters, as possible ancestors for various later rodent
groups including muroids and geomyoids (Wilson, 1949a,
p. 47, 1949b, pp. 97-98; Wood, 1959, p. 358). As described
here the skull of Sciwravus nitidus does not seem to add much
in the way of positive evidence on relationships. Primitive
features include the large jugal, zygomasseteric structure,
seemingly shallow pterygoid fossa, ossified but loosely attached
bulla, short paroccipital processes. The incomplete medial
covering of the petrosal by the bulla may also be primitive,
a feature that §. nitidus shares with Paradjidaumo, an eomyid.
The masseter muscles may have been somewhat better devel-
oped for gnawing than in some contemporary paramyids, such
as Paramys delicatus. This is suggested by the more anterior
position of the anterior zygomatic root and of the mandibular
masseteric fossa, and by the distinct anterior knob and ridge
on the zygoma. A more distinct pterygoid fossa than in
P. delicatus suggests some strengthening of the pterygoid
muscles as well. The zygomasseteric structure is still, however,

8 Postilla Yale Peabody Museum No. 53

that of a primitive sciuromorph. Whether the indications of
muscle strengthening show that the sciuravids were experi-
menting along lines leading toward more advanced zygomass-
eteric conditions remains a matter of speculation. At any
rate, the skull of S. nitidus seems to lack any specializations
countering the suggestions that sciuravids gave rise to some
of the advanced rodent groups.

LiIreERATURE CITED

Burke, J. J., 1938. A new cylindrodont rodent from the Oligocene of
Montana. Ann. Carnegie Mus., vol. 27, pp. 255-274.

Gazin, C. L., in press. New sciuravid rodents from the lower Eocene
Knight formation of western Wyoming.

Hill, J. E., 1935. The cranial foramina in rodents. Jour. Mammal., vol. 16,
pp. 121-129.

Matthew, W. D., 1910. On the osteology and relationships of Paramys, and
the affinities of the Ischyromyidae. Bull. Amer. Mus. Nat. Hist., vol. 28,
pp. 43-72.

Wilson, R. W., 1938. Review of some rodent genera from the Bridger
Eocene. Amer. Jour. Sci., 5th ser., vol. 35, pp. 123-137, 207-222, 297-304.

, 1949a. On some White River fossil rodents. Carnegie Inst. Wash.

Publ. 584, pp. 27-50.

, 1949b. Early Tertiary rodents of North America. Carnegie Inst.
Wash. Publ. 584, pp. 67-164.

Wood, A. E., 1959. Eocene radiation and phylogeny of the rodents. Evolu-
tion, vol. 8, pp. 354-361.

June 30,1961 The Skull of Sciuwravus Nitidus 9

Plate I
Dorsal view of skull of Sciuravus nitidus, U.S.N.M. No. 22477, approx. x 3.

10 Postilla Yale Peabody Museum No. 538

Plate II

Ventral view of skull of Sciuravus nitidus, approx. x 3. Based on U.S.N.M.
No. 22477; restored in part from A.M.N.H. Nos. 12551 and 12531. 1, inci-
sive foramen; 2, sphenopterygoid foramen; 3, entopterygoid plate; 4, fora-
men in basisphenoid; 5, auditory prominence; 6, mastoid process; 7,
paroccipital process; 8, fenestra cochleae; 9, hypoglossal foramen; 10,
jugular foramen; 11, stylomastoid foramen; 12, tympanic bulla; 13, foramen
ovale; 14, ectopterygoid plate; 15, foramen for palatine artery; 16, infra-
orbital foramen.

June 30,1961 The Skull of Sctwravus Nitidus 11

Plate III
Ventral view of skull of Sciwravus nitidus, U.S.N.M. No. 22477, approx. x 3.

Postilla Yale Peabody Museum No. 53

Plate IV

Lateral view of skull and lower jaw of Sciwravus nitidus, U.S.N.M.
No. 22477, approx. x 3.

June 30,1961 The Skull of Sctwravus Nitidus 13

Plate V

Sciuravus nitidus, approx. x 3. Figure 1. Occipital view of skull, U.S.N.M.
No. 22477. Figure 2. Medial view of right lower jaw, C.M. No. 9683.

Priile

YALE PEABODY MUSEUM

oF NaTurRAL History

Number 54 October 30, 1961 New Haven, Conn.

THE DENTITION OF OURAYIA: —ITS BEARING
ON RELATIONSHIPS OF OMOMYID PROSIMIANS

E.wyn L. Smvons

Gazin (1958) has established for the North American
anaptomorph, or so-called *ttarsioid” prosimian Primates two
families: the Anaptomorphidae, containing seven early and
middle Kocene genera; and the Omomyidae, to which eleven
Eocene and one early Oligocene genera are assigned by him.
It has long been recognized that members of these two families
differ distinctly from the more clearly lemur-like North
American prosimians of the subfamily Notharctidae, which
has been suggested, originally by Leidy (1873:90) and more
recently by W. K. Gregory (1921:220), as being plausibly the
group from which the South American platyrrhine Primates
arose. Recently, however, the candidacy of Notharctus and its
allies for such an ancestral position has been increasingly
questioned. This is because, although generalized in many
ways, notharctids already show a number of features that are
unlike Platyrrhini. The greater expression of the hypocone
and mesostyle, together with a tendency toward doubling of
the outer cusp of the fourth upper premolar — all non-platyr-
thine features, but seen in the successively later species of
Notharctus—indicatea dental pattern that was diverging
from, and not approximating that which typifies the Platyr-

2 Postilla Yale Peabody Museum No. 54

rhini. Moreover, both Notharctus and Smilodectes exhibit up-
per third molars that, relative to the other cheek teeth, are
much larger and more complex than those of the earliest known
notharctine, Pelycodus, as well as of those of Oligocene-Recent
South American Monkeys.

Some current students regard omomyid prosimians as al-
ternative, or better, candidates for the ancestry of Ceboidea;
for instance, see Gazin (1958:100). This idea appears to have
had its origin in comments by J. L. Wortman (1904:242) in
his imaginative but often overlooked studies on Eocene Pri-
mates at the Yale Peabody Museum. Probably the main reason
the case for a possible ancestor-descendant relationship be-
tween omomyids and ceboids has not previously been considered
in detail is that no really complete dentitions of members of
this prosimian family have been described. Without better
knowledge of the anterior dentition most of those acquainted
with the problem appear to have hesitated in expressing opin-
ions as to the phyletic relationships of this group. The com-
pleteness of the dentition in one omomyid, a specimen of
Ourayia uintensis, described below, largely obviates this diffi-

culty.

ACKNOWLEDGMENTS

The writer is greatly indebted to Dr. Glenn L. Jepsen of
the Department of Geology, Princeton University, who gener-
ously made available for study and description here the
unequalled specimens of Ourayia in his charge collected for
Princeton by J. B. Hatcher, O. A. Peterson [both formerly
associated with the Yale Peabody Museum], and by later
field expeditions of Princeton University. Thanks are also due
Dr. M. C. McKenna of the American Museum of Natural
History for making available for study the type specimen of
Ourayia wintensis (Osborn) and to Dr. C. L. Gazin of the
U.S. National Museum for kindly discussing with me some
problems relating to the Omomyidae. Figures were prepared
by Margaret EK. Freeman of New Haven and the early com-
pletion of this paper was facilitated by a grant from the
Wenner-Gren Foundation for Anthropological Research of

New York.

Oct. 30, 1961 The Dentition of Ourayia 3

ABBREVIATIONS

A.M.N.H. . American Museum of Natural History, New York.

Pim Oem yet eters is. ce ... Carnegie Museum, Pittsburgh.
Paw Princeton Universi'y, Princeton
Wor MM. . Yale Peabody Museum, New Haven

DESCRIPTION AND RELATIONSHIPS OF OURAYIA

The single species of this genus, Ourayia uintensis, is ap-
parently retricted to Uinta (B) horizon of the late Eocene.
It was originally described by Osborn (1895:77) as a species
of Microsyops. Later, Osborn recognized that the assignment
of this species to Microsyops was in error, remarking
(1902 :202): “Its nearer reference is either to the Anapto-
morphidae or to some member of the Notharctidae.”’? A further,
but incomplete, step in the direction of a more correct taxo-
nomic assignment for this prosimian species was made by
Wortman (1904:232) who referred it to the genus Omomys.
However, it remained for Gazin (1958:70) to recognize that
this species belongs to a distinct omomyid genus for which he
coined the name Ourayia, after the village of Ouray to the
north of the “White River pocket,” Utah, from which it seems
most, if not all, known specimens of this species have been
vecovered. Gazin (1958) noted that this genus is close to later
Eocene omomyids such as Washakius, Hemiacodon, and
Stockia and pointed out that the species differs greatly from
any assigned to Notharctus which it resembles only in its
comparatively large size. The specimens from the Princeton
collections, described here, are much more complete than the
American Museum materials available to Gazin and further
serve to emphasize the distinctness of the dentition of Ourayia
from that of any notharctid. In view of the completeness of
the specimen P.U. 16431 it is now possible to define much
more adequately the structure and relationships of this primi-
tive prosimian.

4 Postilla Yale Peabody Museum No. 54

Order PRIMATES Linnaeus 1758
Suborder PROSIMIT Illiger 1811
? Infraorder LEMuRIFoRMEs Gregory 1915
Family Omomyidae Gazin 1958

Subfamily Omomyinae Wortman 1904
OURAYIA Gazin, 1958

Type species: Ourayia uintensis (Osborn), 1895.

Included species: Owrayia wintensis.
Distribution: Late Eocene, Uintan stage; White River pocket, Uinta
Basin, Uintah County, Utah.

Generic distinctions: In addition too having lower and more anteropos-
teriorly elongated trigonids and comparatively larger talonid basins than
in Hemiacodon, as mentioned by Gazin (1958-71) for Ourayia, the hy-
poconid is less anteroposteriorly conpressed than in Hemiacodon and does
not project as far laterally from the main body of M, as it does in Stockia
and Hemiacodon. M,-.. hypoconulids are not distinctly set off as in Hemi-
acodon and Washakius. Resembles [Temiacodon and differs from Washakius
in lacking molar metastylids. In Ourayia the apices of molar paraconids are
situated somewhat more internally than in Stockia and M, hypoconulids are
comparatively smaller than in Hemiacodon and Washakius. Second and
third molars above and below are larger compared to first molars than in
Washakius and Hemiacodon. Ourayia agrees with species of these two gen-
era in haying crenulate tooth surfaces, but unlike Washakius has only a
single metaconule. Differs from //emiacodon in having a much less sharply
broken crest between para- and metacones and no appreciable development
of P* parastylar cuspule. Anterolingual cingular region of M'? shows a
pericone variably present as is the case also in Hemiacodon and Omomys
among omomyids and in the living platyrrhine Samiri sciurea.

Discussion: One difference between Ourayia and Hemiaco-
don, noted by Gazin (1958:71), that the Ms paraconid of the
former is nearly obscured in the anterior trigonid crest, is now
seen to be a feature of the type specimen alone. The M, para-
conid on both sides in P.U. 11236 and P.U. 16431 shows dis-
tinetly. This difference, together with slightly smaller size and
less oval My outline in the latter two specimens might suggest a
species distinction for the Princeton materials were they not
from the same horizon and area as the type. Gazin pointed
out that the lower dentition of this primate does not resemble
Notharctus. This conclusion is amply confirmed in the upper
dentition of Ourayia where, apart from basic differences in

cheek tooth crown patterns, the upper incisors are seen to be

Oct. 30, 1961 The Dentition of Ourayia 5

comparatively huge and the canine much reduced. These fea-
tures, together with the loss of P=, are in marked contrast to
the situation in notharctines which have small upper incisors
and long canines. The difference here is of the same order of
magnitude as that separating Malagasy lemur from loris
dental patterns and amply justifies reference of Ourayia and
Notharctus to different families.

As regards the phyletic position of Ourayia among omo-
myids I suspect that the genus may have been derived directly
from Hemiacodon, particularly since the Ms paraconid is not
as unlike that of Hemiacodon as was supposed from the type
specimen alone.

Possible relationships with the early Oligocene Macrotarsius
montanus are less certain, but direct derivation of this species
from Ourayia is not out of the question. Crenulations of the
enamel resembling those of both Hemiacodon and Ourayia,
although less pronounced, are evident in the talonid basins
of the Oligocene form. Paraconids are situated slightly more
laterally than in species of the latter two genera, but this
difference need not rule out a close relationship between them
and Macrotarsius. Present knowledge of the relative size, num-
ber and positioning of the anterior lower teeth in several omo-
myid genera, Omomys, Washakius, Chlororhysis, Hemiacodon,
and Ourayia strongly suggests that Clark (1941 :562) was
correct in interpreting the lower dental formula in this Oli-
gocene omomyid as 2.1.3.3, the typical formula for the group.
If P, is lost and Ps single-rooted as in other Omomyidae, then
the small anteriormost remaining tooth in Macrotarsius must
be the base of a reduced canine (see fig. 1). This canine, how-
ever, is not less prominent than in any other omomyid as Clark
suggested, although the lateral incisors do appear to reach
an extreme of reduction, judging from the alveolus. In Oura-
yia, moreover, the lower incisors are more procumbent. ‘To
the extent that the species of Hemiacodon, Ourayia, and
Macrotarsius are in, or close to, a single progressing phylum,
the suggested trends, apart from size increase, were toward
deepening and shortening of the ramus mandibuli, together
with a size reduction and more vertical implacement of the
lower incisors. Perhaps the latter changes are to be correlated

6 Postilla Yale Peabody Museum No. 54

with increasing use of the hands in feeding and with facial

foreshortening.

OURAYIA UINTENSIS (Osborn), 1895
Figures 1, 2, 3.

Microsyops wintensis Osborn 1895, Bull. Amer. Mus. Nat. Hist. No. 7, pp.

Oy 105 Ue
Omomys uintensis Wortman 1904, Amer. Jour. Sci. 4th Ser., Vol. 17, pp.

134, 135.
Ourayia uintensis Gazin, 1958, Smithsonian Misc. Coll. Vol. 136, No. 1, pp.

70-72, pl. 13, fig. 8.

Type of genotypic species: A.M.N.H. No. 1899; left mandibular ramus
with P,-M..

Type locality: Late Eocene, Uinta (B), White River, Utah.

Hyopdigm: Type and A.M.N.H. 1900, mandibular fragment with right
M,; P.U. 11236, left and right mandibular rami with left P,-M, (lacking
trigonid of M., alveoli of P.; right P,-M, (trigon of M, missing), alveoli
of I,-P,; P.U. 11288, edentulous right mandibular ramus; P.U. 16431, max-
illae with upper dentition excepting left P°, mandibular fragments with
left I,, P;-,, damaged M,, M.-M,; right I,, damaged P,, M,-M., trigonid
of Mj.

Horizon and locality: Lower Uinta [B], upper Eocene, White River
Pocket, Utah. P.U. 16418, Section 2, T. 9 S., R. 20 E. Uintah County, Utah;
P.U. 11236, 11288 Uinta |B], Kennedy’s Hole, White River, Utah.

Specific diagnosis and description:! A moderate sized prosimian; com-
parable parts approximately with'n the size range of the living Perodicticus
potto. Dental formula 2.1.3.5; I, spatulate, and somewhat procumbent,

2.1.3.3
posteriorly with median vertical ridge and basal cingulum rising internally
halfway toward crown; I, smaller than I, (alveolus); © probably smaller
than I, (root), no diastema; P. single-rooted; P, with internal cingulum
lacking on P,; P;-, of equal height, paraconid, metaconid, and external cin-
gulum present in P,, -lacking in P;, heel of P;-, with single cusp situated
laterally. Surfaces of cheek teeth, particularly, bearing wrinkled or crenu-
late patterns. M, slightly larger and more oval in outline than M,. Molar
paraconids distinct (except in M,. of type), situated only slightly less
laterally than metaconids and connected with protoconids by an arcuate
crest. Hypoconulids not sharply set off on M,... M, hypoconulid not as

‘I fail to grasp fully the logic behind the increasingly popular practice
of neglecting to distinguish between generic and specific diagnoses in
treatments of monospecific genera. Generic characters, in this case, are
those features which prevent reference of such species to other genera;
the specific, those attributes which, combined, characterize a given, and
no other, species. The two suites of features are not indistinguishable,
although I suspect that the failure to see that they are not may be one
prime contributor to the production of unnecessary or invalid genera.

Oct. 30, 1961 The Dentition of Ourayia 7

large as in most earlier omomyids. Upper incisors large, I°* sub-equal in
size, spatulate; upper canine reduced, premolariform, smaller than P?; P?
small, lacking lingual cusp; P** with s‘ngle inner and outer cusps. M'?
with para- and meticonules, varying expression of cuspules on lingual pro-
tocone cingulum in positions of hypocone and pericone, pronounced labial
cingula with cuspule not seen in /Temiacodon in position of mesostyle.

Discussion: OQurayia uintensis is of greater size than are
other known omomyids except for the considerably younger
species Macrotarsius montanus of Chadronian age, which is
about ten percent larger in comparable parts. Among omo-
myids earlier or contemporary with Ourayia, Hemiacodon
gracilis most nearly rivals it in size, being about eighty-five
percent as large in most measurements. The remote possibility
that canines were lost in O. uintensis and that there were
actually four premolars above and below in this species has
been considered but rejected. In spite of its premolariform
appearance, the upper canine apex is directed forward while
that of P* has a distinct backward tilt which can hardly be
due to crushing. Moreover, occlusion of the teeth in P.U.
16431 shows that the lower canine (which has a larger root
than the teeth adjacent to it) les in front of the upper canine,
as would be expected.

The maxillae of P.U. 16431 are crushed, but some features
of interest are still to be observed (see fig. 2). Much of the
premaxilla of the right side is preserved, and a wavy suture
just in front of the canine and arching backward is indicated
on both sides. Both narial margins of the premaxillae are
evident anteriorly. The thin, fairly long right nasal has drop-
ped down between the premaxillae. In the orbital region the
jugal is missing and the orbital border of the right maxilla
much eroded, so that it is not possible to determine whether
the malar contacted the lachrymal. However, it is difficult to
avoid the conclusion that the orbit was quite large relative to
the size of the rostrum. This possibility is also indicated by
a specimen of Hemiacodon figured by Gazin (1958: ple 4, fig:
4) in which the supraorbital border of both orbits is preserved.
Since this specimen and P.U. 16431 are the only individuals of
any species of omomyid primate which preserve part of the
skull other than maxillae or dentaries, they deserve special
comment. It appears that in Ourayia the depth of the rostrum

8 Postilla Yale Peabody Museum No. 54

above the canine was greater than the horizontal distance from
the narial to orbital borders, so that this primate shows the
progressive feature of being comparatively short snouted,
evidently much more so than in Smilodectes or Notharctus.
On the other hand, this ‘tadvanced’” omomyid feature is
balanced by the presence of a metopic suture between the
frontals in Hemiacodon figured by Gazin as is generally the
case in “primitive” or non-tarsioid prosimians.

The reduced upper canine of O. wintensis, taken together
with possession of a lower canine root that is hardly larger
than Py in cross-sectional area, indicates a small, premolari-
form lower canine. Both Matthew and Granger (1915: fig. 24)
and Gazin (1958: pl. 8) illustrated specimens of the closely
related Hemiacodon gracilis which preserve broken lower
canines that are indeterminate as to the height of this tooth.
A specimen of H. gracilis recently located in the Yale collec-
tions, Y.P.M. 16253 from Henry’s Fork, Wyoming, shows
that the entire tooth was extremely premolariform and only
barely higher than P., (see fig. 1). In view of an overall simi-
larity in general size and proportions of the other teeth
between O. wintensis and H. gracilis, the reduced upper canine
of the former almost certainly opposed a premolariform tooth
below. In earlier omomyids, Omomys and Chlororhysis, the
lower canine is relatively larger (fig 1). Such canine reduction
in the later omomyids practically eliminates them from con-
sideration as being ancestors of any Ceboidea.

Although O. wintensis may be derived from H. gracilis, ref-
erence of the former species to the genus Hemiacodon seems
out of the question. The primary distinctions between these
two genera have been cited in the diagnosis given above (page
4), and they are considerably greater than those which have
been proposed as separating the genera Omomys, Loveina, and
Chlororhysis. No doubt, known omomyid species could be
lumped under fewer genera, but in the fragmentary state of
present knowledge concerning them, limited almost entirely
to dentitions, this would serve no useful purpose and in any
event cannot be attempted here.

Curiously, wear on M of O. uintensis, P.U. 16431, is dis-

: 2 2 : ae
tinctly less than on M-,. That this could be due to misinter-

Oct. 30, 1961 The Dentition of Ourayia 4)

pretation of the dental formula, with molariform P——and loss
of one of the molars seems impossible, particularly because in
the closely related H. gracilis the normal sequence of molar
wear can be established for numerous specimens.

MEASUREMENTS IN MILLIMETERS OF OURAYIA UINT'ENSIS
LOWER DENTITION

PAU: Telefe
No. No. A.M.N.H.
16431 11236 1899
Hength Vy3— Ms, 2. 2. es ce wes 27.6
Length P, —M, Se ee 18.0 18.8
Length P. — Ms . len”) 14.6 15.0
Depth of jaw below M, . 6.2 6.8 deO
Transverse diameters: I, ......... 2.0 1.9
| ee eae
Cyne eee Tae :
ots. ats Thigh (Ce) See ae
| eae ee ee ee 2.0 2.3 2.3
li ee 2a 2.5 2.6
M, (trigonid) 3.0 2.8 2.9
M. 3.1 Ball 3.4
Mine ee eer. 2.9 3.0
M, (talonid) 3.5 3.3 3.6
i Ce eee 3.6 3.3 3.8
MG ae base ee 3.2 3.0
Anteroposterior diameters
| [Ean he Se ee 2.3 2.1
JI es et
lek: ee neers 1L.8(?) :
Re 7538) 3.1 2
P, Meee tse ar
M, _ 4.0 4.2 4.6
M. 4.2 4.2 45
M, 5.2 4.6

10 Postilla Yale Peabody Museum No. 54

UPPER DENTITION

1e51 Ole
No.
16431
Lenght Ne —— Ma cnn en ee 28.9
Dengthie == MO ok) Re oe 175
Transverse diameters: I’ ......... re
| Cae ons. esr
Cone tee 1.4
Pes ie. ce cele 1.4
| ee ee 3.1
Ph 2 ere eae 3.8
Vie oer et ie:
MiG ieeh ca cae 4
NIG ae a eee AES
Anteroposterior diameters :
Nas Je. pence 3.4
| (Ber Re ol ee 3.1
Ge 2 60g eee
| eat Rr F283
Pe ne wenn ool
Dire 22. te ee ak 3.0
1. Eaaereeeyh Bras a 3.8
Ny (ee coe a Mees 4.1
Mee ten ee 3.8

Measurements for a possibly associated right P. of P.U.
16431 have been given above. Although there is no definite
contact between this tooth and the right mandibular fragment,
it is the proper size and shape for an omomyid P., judging

from morphology of Ps in Omomys and Hemiacodon.

RELATIONSHIPS OF THE OMOMYIDAE
The hypothesis that omomyid prosimians may be near the
ancestry of the platyrrhine monkeys is based on several points

of direct and indirect evidence, most of which are rather

Oct. 30, 1961 The Dentition of Ourayia il

equivocal, as is often the case with such phyletic conjec-
tures. For those who do not favor a notharctid derivation for
platyrrhines these small Primates remain as the only other
group now known in North America which contains mem-
bers that are early and generalized enough to admit of
such a relationship. Other early Cenozoic families of North
American Primates including Plesiadapidae, Carpolestidae,
Anaptomorphidae, Phenacolemuridae, and Paromomyidae (if
the latter two are regarded as distinct) exhibit extreme tooth
specialization or reduction which entirely eliminates their
known members from any ancestral relation to the South
American Monkeys. It is clear, however, that if the dental
form and arrangement of Ourayia uintensis was at all typical
of the later Omomyidae as a whole, it would require a reversal
of the trend toward reduction of the canines, seen in this
species in order to reach the condition typical of the ceboid
monkeys. Such a possibility seems at best rather unlikely.

The question of the ancestry of the platyrrhine monkeys
bears rather crucially on the interpretation of a number of
more general assumptions inherent to the currently accepted
higher categories of Primates, particularly the concept of the
suborder Anthropoidea, as well as to the widely accepted
succession of grades, from lemuroid to tarsioid, to monkey
and etc., through which the ancestors of man and the other
Higher Primates are commonly supposed to have passed.
Perhaps the problem is largely semantic, resulting from the
all too human tendency to superimpose an idealistically sub-
divided terminology on what are actually continua of evolving
lineages. Nevertheless, most of the various named higher
‘ategories of Primates have been, and presumably will continue
to be, useful in talking about evolutionary relationhips within
the order. One possible reaction might be to suspend judge-
ment or discussion of relationships between early and late
Cenozoic Primates in view of the partial and fragmentary
evidence now available, but within the framework that has
been set up by previous research it seems advisable to follow
out some earlier suggestions to their logical conclusion. For
instance, if platyrrhine monkeys were actually derived from
anything like the Notharctus group or even from the omomyid

12 Postilla Yale Peabody Museum No. 54

prosimians, it is difficult to see how they could have passed
through a grade of organization that need be qualified as
tarsioid. If either of these groups are actually ancestral to the
South American Monkeys, but not to the Old World Higher
Primates, then it is also necessary to conclude that those
features which are shared by Old and New World Higher
Primates are the result of parallel evolution and that these
two groups were independently derived from the Prosimii.

In the latter case the taxon Anthropoidea consists of a
grade in the sense of Huxley (1958) rather than a clade, in
which the common ancestor of subsequent derivative stocks
shares something of the definition which justifies the associa-
tion of such subsequent groups within a single taxon. If Catar-
rhini and Platyrrhini were derived from independent stocks of
Prosimu, then Anthropoidea have a polyphyletic origin, even if
such stocks belonged to the same major division of prosimians.

When considering the various alternatives for the derivation
of the Platyrrhini it may be noted that latest species of the
genus Notharctus, and of Smilodectes as well, are rather large
prosimians, having approximately the body size range seen in
species of the living Malagasy genus Lemur or in the domestic
cat. It seems implausible, although not impossible, that forms
such as the pigmy marmoset could have descended from ances-
tors of the size range of known notharctids. The smallest
notharctids, species of the early Eocene genus Pelycodus, in
comparable parts, have about twice the linear dimensions of
the smaller species of Callithrix and are even larger compared
to Cebuella, should the latter genus be sustained as distinct
from Callithrivx | Hapale|. Inasmuch as known species of Pely-
codus give every evidence of being close to the origin of the
taxon Notharctidae, there is little reason to posit that there
ever were unknown smaller members of this subfamily from
which marmosets such as Cebuella might more plausibly have
been derived without marked size decrease at some intermediate
period. One is therefore faced with the supposition that, if
Notharctidae are in or near the ancestry of platyrrhines, mar-
mosets have undergone a size reduction since their initial dif-
ferentiation. This view has sometimes been put forward, but

to date there is no paleontological evidence for it.

Oct. 30, 1961 The Dentition of Ourayia 13

In conclusion, it is possible to say that in spite of the fact
that late Eocene and Oligocene omomyids were specializing
along distinct lines of their own, not foreshadowing Platyrrhini,
it seems probable that Omomys and its immediate forebears are
the most likely early Cenozoic prosimians to have a direct
relationship to the rise of the South American Monkeys.
Among principal evidences supporting this view are the obser-
vations that Omomys, or one or more forms allied to it, was
smaller than any known ceboids, had suitably unspecialized
molar crown patterns together with small third molars, shared
with some ceboids the otherwise nearly unique possession of a
pericone cusp, and belongs to a group showing trends away
from the primitive prosimian condition toward foreshortening
of the rostrum, orbital enlargement, and vertical incisor em-
placement. Moreover, Omomyidae are the only known family
of ancient and undoubted Primates now known which possessed
exactly the same dental formula as do the living Cebidae.
Nevertheless, only in earliest omomyids are relative sizes of
respective tooth types reasonably satisfactory for derivation
of the tooth morphology characteristic of Oligocene-Recent
South American Monkeys.

14 Postilla Yale Peabody Museum No. 54

Figure 1.

Diagrammatic reconstructions of a sequence of representative omomyid
species, approx. x 3. Dotted outlines hypothetical.

These species may not represent a single phylum, but each is typical of
the successive Epoch substage to which it belongs. Specimens on which
this chart is based are as follows: Macrotarsius montanus (type) C. M.
9592 (reversed) ; Ourayia uintensis, P.U. 16431 (P, and M, reversed from
right ramus): Hemiacodon gracil’s (composite), part A—A.M.N.H.
12037, part B—Y.P.M. 16253, part C—Y.P.M. 12987-1; Omomys carteri
(composite), part A—A.M.N.H. 12600, part B—Y.P.M. 16287 (reversed),
part C—Y.P.M. 13219-2 (reversed).

Oct. 30, 1961

---~\

EARLY OLIGOCENE

EXQICIE NIE

CA TE

WwW
Zz
WW
Oo
(e)
W

MIDOLE

The Dentition of Ourayia

MACROTARSIUS

OMOMYS

16 Postilla Yale Peabody Museum No. 5k

Figure 2.

Occlusal views of right upper, and left lower dentitions of Ourayia win-
tensis, P.U. 16431, (M, reversed from right side), approx. x 5.5.

Oct. 80, 1961 The Dentition of Ourayia ily

18 ; Postilla Yale Peabody Museum No. 54

Figure 3.

Lateral view of right maxilla of Ourayia uintensis, P.U. 16431, approx.
x 5.5.

Oct. 30, 1961

The Dentition of Ourayia

ig

20 Postilla Yale Peabody Museum No. 54

REFERENCES

Clark, J., 1941. An anaptomorphid primate from the Oligocene of Montana.
Journ. Paleo., v. 14, no. 5, pp. 562-563, 1 fig.

Gazin, C. L., 1958. A review of the Middle and Upper Eocene Primates of
North America. Smiths. Mise. Coll. v. 136, no. 1, pp. 1-112, 14 pl.

Gregory, W. K., 1921. On the structure and relations of Notharetus, an
American Kocene primate. Mem. Amer. Mus. Nat. Hist., v. 3, pt. 2,
pp. 49-243, 84 fig., 36 pl.

reference to grades. Uppsala Univ. Arsskrift, No. 6, pp. 21-39.

Leidy, J., 18783. Contributions to the extinct vertebrate fauna of the
western territories. Rep. U.S. Geol. Surv. Terr. (Hayden), v. 1, pp.
1-358, 33 fig., 37 pl.

Matthew, W. D. and W. Granger, 1915. A revision of lower Eocene Wasatch
and Wind River Faunas. Part IV., Bull. Amer. Mus. Nat. Hist., v.
34, pp. 429-483, 52 fig., 1 pl.

Osborn, H. F., 1895. Fossil mammals of the Uinta Basin. Expedition of
1€94. Bull. Amer. Mus. Nat. Hist., v. 7, pp. 71-105, 17 fig.

—, 1902. American Eocene Primates, and the supposed rodent
family Mixodectidae. Bull. Amer. Mus. Nat. Hist., v. 16, pp. 169-214,
40 fig.

Wortman, J L., 1903-1904. Studies of Eocene Mammalia in the Marsh Col-
lection, Peabody Museum. Part 2. Primates. Amer. Journ. Sci., 4th Ser.,
v. 15, pp. 163-176, 399-414, 419-436, v. 16, pp. 345-368, v. 17, pp. 23-33,
133-140, 203-214, 48 fig., 2 pl.

LE

YALE PEABODY MUSEUM

oF NATURAL History

Number 55 November 6, 1961 New Haven, Conn.

NOTES ON SAGITT'A FRIDERICI RYVTER-ZAHONY
COLLECTED OFF PERU*

Taxast Toxioka
Sero Marine Brotocicat LABORATORY

SIRAHAMA, JAPAN

In a paper on the chaetognath fauna off Peru, Bieri (1957)
remarks, “In the 1941 Peru material a complete set of inter-
grades exists between 8. friderici to the south and S. tenuis to
the north. The same situation has been observed by the author
in samples taken off Lower California except that there the
tenuis-like form is to the south and the friderici-like form to
the north” (pp. 261-262, fig. 13). He showed that Sagitta
bipunctata described by Michael (1911) was identical with the
friderici-like form and that S. tenuis and S. friderict were eco-
typic variants of a single interbreeding population or species
(p. 261). The smaller tenwis-form was considered a warm water
form at that time.

In Bieri’s 1952 material, however, only the larger friderici-
form was found at stations as far north as the Gulf of Guaya-
qui. He then changed his opinion, admitting the validity of
S. friderict and characterizing it as neritic or nearshore. He
noted that the “distribution of Sagitta friderici off California,

* Contributions from the Seto Marine Biological Laboratory, No. 359.

2 Postilla Yale Peabody Museum No. 55

Peru and Chile, and North and South Africa, is correlated with
upwelling and suggested that temperature and salinity are re-
lated to the distribution pattern of this species” (Bieri, 1959,
pp. 14-18, fig. 17). He re-examined the specimens of Michael’s
S. bipunctata collected off California and found that they were
in fact S. friderict (p. 14, footnote). The northernmost record
of §. friderici in the eastern Pacific is evidently Monterey Bay
where Bigelow and Leslie (1930) found Michael’s S. bipunctata
to be common (pp. 552-553). While there is no published
record of §. friderict in Chilean waters, Bieri believes that the
distribution of this species extends to the waters off Chile.

Sund (1959a) considers S. friderict as a synonym of S.
tenuis and showed during the Eastropic Expedition (1959b)
that 8. tenuis occurred in only a limited area of the Gulf of
Panama.

I have examined S. tenuis from Scammon’s Lagoon and
Manuela Lagoon, Baja California, and S. friderici collected in
the blue-green water along the southern California coast near
San Diego (1959). I have also found many specimens refer-
able to 8. friderict in collections of the Transpac and Shellback
expeditions. Most of them were identified without any hesitation
as §. friderici, but some doubtful specimens from the offshore
waters in the Shellback Area were placed in Groups A and B
(pp. 360-364, table 7, fig. 5). As these specimens were mostly
found in more or less imperfect states of preservation, I could
not examine the exact structure of the seminal vesicle or the
corona ciliata.

Fortunately, however, I have had a chance to examine some
excellently preserved specimens referable to §. friderict from
the collection of the 1958 Yale Peruvian Expedition which were
submitted to me for examination by Dr. G. B. Deevey of the
Bingham Oceanographic Laboratory, Yale University, to whom
I want to express my hearty thanks for her kindness.

These specimens include ten individuals, 8.9 mm - 13.3 mm in
length, collected on April 2 at Station 34, 4° 3’ S. Lat., 81°
10’ W. Long. ; and 12 individuals, 10.0 mm - 13.5 mm in length,
collected on April 14 at Station 81, 3° 36’ S. Lat., 80° 47’ W.
Long. Measurements of the specimens from the two stations are

given in tables 1 and 2 and resemble each other closely except

Nov. 6,1961 Sagitta friderici Ritter-Zahony 3

for the TC value (ratio of anterior part of the posterior fin
along the trunk to posterior part of the posterior fin along
the caudal segment times 100). All the morphological charac-
teristics mentioned below are common to the specimens from
both stations.

SbvGsp ona k

Sagitta friderici, armature formulae of individuals from Sta. 34.

Body Caudal segment Anterior Posterior

length in per cent* Hooks teeth teeth TC-value

8.9 mm 27.0 8-9 6-6 15-16 81.6-84.7
STS 26.4 7-7 9-9 20 - 21 80.4-88.9
HES 25.6 7-7 39 22 - 22 73.3-80.0
11.9 26.4 7-7 8-9 20 - 20 87.4-87.5
12.0 26.4 7-7 9-9 18 - 21 86.9-88.7
12.3 25.2 7-8 8-9 20 - 20 76.9-82.7
12.5 25.8 7-7 8-8 21 - 22 78.6-83.2
12.7 25.3 7-7 9-10 20 - 20 84.7-85.9
13.0 27.4 7-7 9-10 21 - 23 69.5-75.8
13.3 25.7 7-7 8-8 20-21 80.5-84.8

* Caudal fin included.

TABLE 2
Sagitta friderici, armature formulae of individuals from Sta. 81.

Body Caudal segment Anterior Posterior

length in per cent* Hooks teeth teeth TC-value
10.0 mm 25.5 9-? 8-8 18-19 67.3-76.8
10.6 26.6 8-8 8-8 18-19 67.5-71.4
te 26.6 7-8 8-8 19-19 77.8-83.3
11.5 27.4 8 8 18 78.6-80.7
11.6 26.2 7-7 9-10 20 - 22 83.8-87.0
aE 27.4 7-8 10-10 23 - 23 87.3-91.4
11.9 26.7 7-7 10-10 22 - 23 83.1-85.2
12.0 26.5 8-8 8-9 21-21 64.2-76.5
12.3 26.9 7 11 23 85.7-91.1
12.4 25.4: 8-8 8-8 21 - 22 70.9-79.5
12.6 27.1 7-8 9-9 21 - 22 79.5-80.0
13.5 26.7 7-8 9 22 66.4-71.0

* Caudal fin included.

4 Postilla Yale Peabody Museum No. 5

Or

DESCRIPTION

Length up to 13.5 mm; the tail segment occupies 25.2% to
27.4% of the whole body length including the caudal fin, with
an average of 26.4% for 22 measurements. The body is
moderately rigid or rather soft and so translucent that the
whitish and opaque intestine can be seen. It is usually widest in
the posterior part of the trunk in the region of the posterior
fins; there is no constriction at the trunk-tail septum. The an-
terior fin usually begins at the level of the posterior end of the
ventral ganglion, although in a few specimens there may be a
short distance, less than half of the ganglion’s length, between
the posterior end of the ventral ganglion and the anterior end
of the anterior fin. The posterior fin is very slightly shorter
than the anterior one; the ratio for anterior fin to posterior fin
times 100 is 86-113, with an average of 105 for 22 individuals.
The value is slightly higher in individuals from Sta. 34 than in
those from Sta. 81, namely 100-113 (with an average of 108 for
ten individuals) compared with 86-113 (with an average of
102 for 12 individuals). The fin is widest behind the trunk-tail
septum. Both anterior and posterior fins are fully rayed; the
rays are set vertically to the base in a small anterior part of
each fin. The TC value is 69.5-88.9, with an average of 82.1
for ten individuals from Sta. 34: while it is 66.4-91.4, with an
average of 78.6 for 12 individuals from Sta. 81. The distance
between the anterior and posterior fins is highly variable, with
an average value of 1/3.75 of the length of the anterior fin for
22 individuals; the distance is slightly shorter in individuals
from Sta. 81 than in those from Sta. 34, the denominator be-
ing 3.1-6.0 (an average of 4.18 for 12 individuals) as against
2.3-4.1 (3.23, average for ten individuals). The collarette is
distinct around the neck and diminishes in thickness posteriorly,
reaching one half to two thirds of the distance from the neck
to the ventral ganglion. The eye pigment (figs. 5-6) is small to
medium, roundish or oval in shape. and the eyes are situated
rather widely apart. The distance between the eyes is 24.7 %-
36.8% (average, 32.7% for five specimens) of the width of the
head at the level of the eyes. The corona ciliata (figs. 2-4)* is
* Heydorn (1959) described the corona ciliata as starting just behind or

in front of the eyes in South African specimens, but I have never seen

any specimen with the corona beginning just behind the eyes.

Noy. 6,1961 Sagitta friderici Ritter-Zahony 5

elongate. It begins just behind the brain and extends poster-
iorly for two-thirds of the distance from the neck to the ventral
ganglion, or for cne and one-half to three times and most fre-
quently two and a half times the head length (see fig. 1). The
corona has a pair of very prominent sinuses behind the eyes,
but posteriorly it is only slightly sinuous. Usually six pairs of
tufts of large tactile setae are found along the corona, although
they fluctuate in number from five to seven. Intestinal diverti-
cula are absent. Hooks number seven or eight, rarely up to
nine; anterior teeth number six to ten, rarely up to 11; and
both rows meet each other at an acute angle. There are 15 to 28
posterior teeth.

The 8.9 mm specimen from Station 34 is devoid of ovaries
and seminal vesicles, but all others are provided with these
structures in various stages of development. The anterior end
of the ovary is situated most frequently near the anterior end
of the posterior fin, but it may occur a considerable distance
from this level; in five individuals the ovaries extend far beyond
the anterior end of the posterior fin and attain the level of the
middle of the anterior fin. Immature ova are only 83 / in long
diameter for an average of 11 measurements, whereas mature
ones are 210-240 4 in long diameter. The seminal vesicle (figs.
7-14) is situated just at the base of the caudal fin; the poster-
ior fin also ends very close to the vesicle. In earlier stages of
development, the anterior glandular portion, which is some-
what rounded and walled with a tall epithelium that secretes
the mucus for agglomerating sperm, is very prominent as com-
pared with the low and inconspicuous saccular portion. How-
ever, the saccular portion becomes very prominent in advanced
stages when it swells outwards more than the anterior glandular
portion. The rupture seems to occur along the lateral side of
the glandular portion at maturity.

REMARKS

The difference in TC value between the specimens from
Station 34 and Station 81 is 3.5 when average values are com-
pared. However, the value varies considerably since it is much

6 Postilla Yale Peabody Museum No. 55

EXPLANATION OF PLATE

Sagitta friderici Ritter-Zahony from Peruvian waters.

1. Dorsal side of specimen 12.3 mm in length from Sta. 81.

2. Corona ciliata of the same individual.

3. Aberrant corona ciliata of specimen 10.6 mm in length from Sta. 81.
4. Aberrant corona ciliata of specimen 11.3 mm in length from Sta. 81.
5. Eyes of specimen 11.9 mm in length from Sta. 81. 300x.

6. Eyes of specimen 12.0 mm in length from Sta. 81. 300x.

7-14. Seminal vesicles arranged in order of developmental stages. 100x.
7. Specimen 12.0 mm in length from Sta. 34.
8. Specimen 11.7 mm in length from Sta. 34.
9. Specimen 12.3 mm in length from Sta. 34.
10. Specimen 13.0 mm in length from Sta. 34.
11. Specimen 12.5 mm in length from Sta, 34.
12. Specimen 10.0 mm in length from Sta. 81.

13. Right vesicle of specimen 11.9 mm in length from Sta. 34;
dorsal view.

14. Left vesicle of same individual; ventral view.

-Zahony

Sagitta friderict Ritter

7» L96L

6

Nov.

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an
ae

& sh
he

sieiieagll

as

Pee ROR S SDT ITED TYTN TOOT OE FATE ETT

SXQKKKCCGSS”

8 Postilla Yale Peabody Museum No.

wr

5

affected by contraction or bending of the body. Even for the
same individual, the value can be different between the right and
left sides. For the 22 specimens examined here, the difference
in value found between the two sides of the same individual
Huctuates from 0.1 to 12.3, most frequently in the range from
1 to 7; and the average difference is 4.7, This is clearly greater
than the difference noted between the samples from the two
stations and suggests that the difference in TC value found for
these specimens is insignificant. Thus, individuals from Station
34 may be considered to be identical with those from Station 81,
and all these are identical with those of my Group A (Tokioka,
1959, p. 361). The latter vary up to 13.8 mm in body length,
are armed with seven to nine hooks, have up to 11 anterior
teeth and up to 25 posterior teeth, and show a TC value fluc-
tuating from 80 to 91.1 (average, 86.3). The differences be-
tween Group A and Group B mentioned in the same paper are
too great for these groups to be merged.

The appearance and structure of the specimens from the
Bingham Oceanographic Collection resemble closely those of
Sagitta friderici as described repeatedly in many previous
papers, except for the armature formulae. The maximal num-
bers of anterior teeth (11) and posterior teeth (28) exceed
those known previously for Sagitta friderict. Thus it is neces-
sary to review the previous descripticns of 8. friderict in order
to establish the range of variation. There follows a list of di-
mensions for §, friderici published by previous authors. As the
species was established by Ritter-Zahony (1911) and examined
and described most exactly by Faure (1952) and Furnestin
(1957), I shall begin with the data of these authors.

Ritter-Zahony (1911)
Loc.: The surface layer off South-West Africa and Cape
Verde.
Body length: up to 13 mm.
Hooks: 8 - 9.
Anterior teeth: up to 9.

Posterior teeth: up to 22.

Nov. 6,1961 Sagitta friderict Ritter-Zahony 9

Faure (1952)
Loc.: The neritic waters off Morocco.
Body length: up to 15 mm.
Hooks: 5-9, most frequently 7 - 8.
Anterior teeth: up to 8, most frequently 6.
Posterior teeth: up to 21, most frequently 12 - 13.

Furnestin (1953)
Loc.: Israel.
Body length: up to 10.2 mm.
Hooks: 6-8.
Anterior teeth: 4-8.
Posterior teeth: 8 - 14.

Furnestin (1956)
Loc.: Tangier Bay and the west entrance to the Gibraltar
Straits.
Body length: up to 12.5 mm.

Furnestin (1957)
Loc.: The neritic waters off Morocco.
Body length: up to 15.1 mm.
Hooks: 5-9, most frequently 7 - 8.
Anterior teeth: up to 8.
Posterior teeth: up to 17.

Michael (1911), as Sagitta bipunctata
Loc.: San Diego region.
Body length: up to 17 mm.
Hooks: 7 - 8.
Anterior teeth: 5-7.
Posterior teeth: 12-14.

Seaccini and Ghirardelli (1941)
Koc Kiowe-Oro:
Body length: 7 mm-10 mm.
Hooks: 7 -8.
Anterior teeth: 6-8.
Posterior teeth: 14 - 20.

10 Postilla Yale Peabody Museum

Vannucci and Hosoé (1952)

Loc.: Near Trinidad in the South Atlantic.
Body length: 8.2 mm - 8.5 mm.

Hooks: 8.

Anterior teeth: 7.
Posterior teeth: 12.

Colman (1959)

Loc.: Eastern Central Atlantic.
Body length: 6.8 mm and 12.7 mm.
Hooks: 7-8.

Anterior teeth: 6-8.

Posterior teeth: 12-18.

Heydorn (1959)

Loc.: The neritic waters off South West Africa.

Body length: up to 18 mm.
Hooks: up to 9.

Anterior teeth: up to 8.
Posterior teeth: up to 20.

Tokioka (1955)
Loc.: The neritic waters off Morocco.
Body length: up to 11.6 mm.
Hooks: 7 -9.
Anterior teeth: 5-7.
Posterior teeth: 11-17.
TC value: ‘71.1 - 91.0; av. 82.0.

Tokioka (1959)

Loc.: Blue-green water off lower California.
Body length: up to 19 mm.

Hooks: 7-9.

Anterior teeth: up to 6.

Posterior teeth: up to 12.

TC value: 83.3 - 134.0; av. 107.1.

No. 55

Noy. 6,1961 Sagitta friderict Ritter-Zahony BI

Tokioka (1959), Group B
Loc.: The waters off central and northwestern South
America.
Body length: up to 9.4 mm.
Hooks: 5-6.
Anterior teeth: 6-9.
Posterior teeth: 13-19. <
TC value: 88.4- 114.3; av. 96.6.

The present specimens and Tokioka’s Group A are charac-
terized by the following data:

Loc.: The waters off central and northwestern South
America.

Body length: up to 13.8 mm.
Hooks: 7 to 9.

Anterior teeth: up to 11.
Posterior teeth: up to 25.
TC value: 78.6 to 86.8.

As nine anterior and 22 posterior teeth are already recorded
for Sagitta friderici, the existence of 11 anterior and 25
posterior teeth is not unreasonable; these may be accepted as
the upper limits in the species. Group B is characterized by
fewer hooks than usual, but this low number of hooks is shared
by some specimens from neritic waters off Morocco and Israel.
Thus Group B may most probably be included in S. friderict
as an unusual group. The maximum body length is 19 mm,
found in the collection from the blue-green water along South-
ern Califorma (Tokioka, 1959, table 27 on p. 390).

Detailed comparisons have been made between Sagitta fri-
derict and S. bipunctata by Faure (1952) and Furnestin
(1957), between S. friderici and S. setosa by Furnestin (1957,
1958) and between S. friderict and S§. hispida by Furnestin
(1957). However, the most serious problem concerns the sep-
aration of S. friderict from S. tenuis. Pierce (1951) considers
these two species as ecological forms of S. tenuis, and Sund
(1959a) seems to agree without giving any clear reasons. On
the other hand, Fraser (1952), Furnestin (1957), Bieri

12 Postilla Yale Peabody Museum No. 55

(1959), Colman (1959), and the present writer (Tokicka,
1955, 1959) ae the validity of S§. friderici. S. tenuis can be
separated from §. friderici by the smaller size of mature in-
dividuals (less than 10.7 mm), the slightly larger number of
anterior and posterior teeth (up to eight anterior and up to
19 posterior teeth for individuals less than 8 mm), and the
comparatively smaller TC value (55.3 - 84.5, with an average
of 64.7, given by Tokioka, 1955; 29.9 - 72.4, usually 40.8 - 61.3
as reported by Colman, 1959). Furthermore, Fraser (1952)
mentions the difference in general appearance between S. tenuis
and §. friderici.

However, according to Furnestin’s (1959) descriptions of
the variability of S, friderici, the specimens from the waters
along the Senegal coast sometimes look more massive and have
Icnger ovaries than those from the Gulf of Guinea which seem
to be weaker in body appearance and attain maturity more
rapidly than those of Morocco waters. Colman (1959) records
that individuals of S. tenzis from Cedar Keys, Florida, differ
from those of British Guiana in having a somewhat shorter
caudal segment (26%-29% v. 27%-34%), sueuely fewer hooks
(7-8 v. 7-9), and a higher TC value (53.8%-92.3°% v. 29.9%
72.4%). Also, the anterior fin begins more posteriorly and
consequently slightly shorter. The massive body appearance
of S. friderict from the waters along the Senegal coast some-
what resembles that of S. tenuis, and the contrarily higher TC
value (53.8%-92.3%) of S. tenuis from Cedar Keys, Florida,
lies within the range of variation of TC values for §. friderici.
These two points are noteworthy, although they do not pro-
vide sufficient evidence to combine S. friderict with S. tenuis
completely. Usually 8. tenuis is found in embayments or in
areas more or less protected from the open sea (Suarez, 1955;
Pierce, 1958; Colman, 1959: and Tokioka, 1959), although its
distribution extends more than five miles offshore in the coastal
waters of western Florida. On the other hand, the distribution
of S. friderict is confined to the neritic water mass according
to Scaccini and Ghirardelli (1941), Faure (1952), Furnestin
(1956, 1957, 1959, 1960), Bieri (1957, 1959), Sund (1959b),
Heydorn (1959), and Tokioka (1959). The reported occur-
rences of §. friderici near Trinidad Island in the South Atlantic

Nov. 6,1961 Sagitta friderict Ritter-Zahony 13

(Vannucci and Hosoé, 1952) and in the eastern Central Atlan-
tic (Colman, 1959) indicate that this species is not confined
to neritic waters as suggested by Faure. It is probable, how-
ever, that the specimens found by these authors are drift forms
carried far offshore from neritic waters.

Finally, it may be noted that the individuals of S. friderict
occurring in the northern part of the range of the species in the
Eastern Pacific and those occurring in the southern part differ
considerably from each other in tooth number.

SUMMARY

A collection of chaetognaths from the waters off Peru is
described; all are referred to Sagitta friderict Ritter-Zahony.
The range of variation within this species and the distinctness
of 8. tenuis Conant from it are discussed. §. fridericit is found
in neritic water masses, while §. fenwis occurs in embayments
and areas more or less protected from the open sea.

POSTSCRIPT

After I had sent the manuscript of this paper to the editor,
five more papers including descriptions or notes on Sagitta
friderici, S. tenuis or on some forms allied to them were
published.

Bainbridge, V. (1960: The plankton of inshore waters off
Freetown, Sierra Leone. Colonial Off. Fish. Publ. No. 13,)
mentions that 8. friderict and S. hispida were the most import-
ant chaetognaths in that area, while the occurrence of S. tenuis
was sporadic,

Sudrez-Caabro, J. A. and Madruga, J. E. (1960: The Chae-
tognatha of the northeastern coast of Honduras, Central
America. Bull. mar. Sci. Gulf Carib., 10: 421-429.) found a
small number of §. tenuis near the entrance to Caratasca La-
goon on the northeastern coast of Honduras. These specimens
were 4 mm-8 mm in length with the tail segment 26.7 %-28.5%
as long as the body length, were armed with 6-8 hooks, and had
4-7 anterior and 6-8 (4 mm-5 mm long individuals) to 10-18
(7 mm-8 mm long individuals) posterior teeth. The TC-value

14 Postilla Yale Peabody Museum No. 55

measured on Fig. 3A is 86.4. This is unusually large for §.
tenuis if the figure is made quite accurately.

Fraser, J. H. (1961: Nigerian Chaetognatha-Sagitta fri-
derict R. Z. Ann. Mag. nat. Hist. (13) 3: 289-290.) examined
S. friderict found in four tubes of plankton taken during half-
hour hauls made in August - October 1957 in the estuary of
the Bonny River, off Port Harcourt, Nigeria. Only the present
species of chaetognath was found in the collections. The maxi-
mum length was 9 mm in the August sample, 13 mm in Septem-
ber, and 11.5 mm in October. The annual temperature and
salinity ranges are 26 - 30°C and 12 - 23 0/00, but the salinity
dropped to 11.5 0/00 in September 1957.

Alvarino A. (1961: Two new chaetognaths from the Pacific.
Pacif. Sci. 15: 67-77.) established a new species, Sagitta
euneritica, which occurs close to shore from Cape Mendocino
to Punta Eugenia in Baja California. She considers this new
species as identical with the form which Bieri (1957 and 1959)
recorded as S. friderici (?) in the same area along the coast
of North America and extending south to the waters of Peru
and Chile. And very probably many of the S. friderici collected
and examined by me in the blue-green water along southern
California (1959) are to be included in the new species accord-
ing to her opinion. She compared S§. ewneritica with S. friderici
and §. setosa, but the last species differs distinctly from S.
friderict and S. euneritica in the fully matured state as the
seminal vesicle is far from the tail fin. Thus, the most important
point is the distinction between §. ewneritica and S. friderici.
As far as I am aware, two of the characteristics given by
Alvarino for S. euneritica seem to be significant. One is the
position of the posterior fin which les more on the trunk than
on the tail; the other is the structure of the seminal vesicle.
However, the TC-value varied from 76 to 141.7 in 33 specimens
of §. friderict collected in the blue-green water and vicinity and
therefore the former character cannot be a definite one. The
latter character may be the only one differentiating §. ewneri-
tica from S. friderici. The author of 8. ewneritica mentions that
the seminal vesicle of her species is like that of §. neglecta and
her Fig. 9 shows that it is devoid of any glandular portion at
the anterior end of the vesicle. But I observed seminal vesicles

Noy. 6,1961 Sagitta friderict Ritter-Zahony 15

just like those of typical S. friderict in some individuals col-
lected in the blue-green water. Moreover, it is noteworthy that
the outline of the vesicle shown by Alvarifo in Fig. 9 resembles
closely some of those shown in Figs. 7-14 of this paper. It is
not impossible that S. ewneritica is nothing but an intraspeci-
fic variant of S. friderici.

Sund, P. N. (1961: Two new species of Chaetognatha from
the waters off Peru. Pacif. Sci. 15: 105-111.) established two
new species, Sagitta peruviana, distributed rather widely in the
coastal waters of Peru, and Sagitta popovicti, found only near
the entrance to the Port of Talara, Peru. Evidently S. peruwvi-
ana is identical with the individuals of SS. friderict treated in
this paper, and S. popovicii seems to be identical with S. tenuis.

Larerarure Crrep

Bieri, R., 1957. The chaetognath fauna off Peru in 1941. Pacif. Sci. 77:
255-264.

, 1959. The distribution of the planktonic Chaetognatha in the
Pacific and their relationship to the water masses. Limnol. Oceanogr.
4: 1-28.

Bigelow, H. B. and M. Leslie., 1930. Reconnaissance of the waters and
plankton of Monterey Bay, July, 1928. Bull. Mus. comp. Zool. Harv.
70; 427-581.

Colman, J. S., 1959. The “Rosaura” Expedition, 1937-1938. Chaetognatha.
Bull. Brit. Mus. (nat. Hist.) Zool. 5: 219-253.

Faure, M.-L., 1952. Contribution 4 étude morphologique et biologique de
deux chaetognathes des eaux atlantiques du Maroc: Sagitta friderici
Ritter-Zahony et Sagitta bipunctata Quoy et Gaimard. Vie et Milieu 3:
25-43.

Fraser, J. H., 1952. The Chaetognatha and other zooplankton of the Scot-
tish area and their value as biological indicators of hydrographical
conditions. Mar. Res. Dept. Agric. Fish. Scot. 1952 (2): 52 pp.

Furnestin, M.-L., 1953. Sur quelques chaetognathes d’Israél. Bull. Sea Fish.
Res. Sta. (Minist. Agric. Div. Fish. Israel) 6: 411-414.

- , 1956. Chaetognathes de la Baie de Tanger et de lentrée
occidentale du Détroit de Gibraltar. Rapp. Comm. int. Mer. Médit. 73:
213-217.

, 1957. Chaetognathes et zooplancton du secteur atlantique
marocain. Rev. Trav. Inst. Péches marit. 22; 1-356.

= , 1958. Les variations morphologiques de Sagitta setosa
Miiller et ses rapports avec deux espéces voisines. Rev. Tray. Inst.
Péches marit. 22: 211-223.

, 1959. Campagne de la Calypso dans le golfe de Guinée
et aux iles Principe, Sao Tomé, Annobon (1956). 8. Chaetognathes.
Ann. Inst. océanogr. Monaco 37; 219-233.

16 Postilla Yale Peabody Museum No. 55

» 1960. Observations sur quelques échantillons de zooplanc-
ton d’ Afrique occidentale. Bull. Inst. france. Afr. noire. 224: 142-151.

Heydorn, A. E. F., 1959. The Chaetognatha off the west coast of the Union
of South Africa, July, 1954- June, 1955. Invest. Rep. Div. Fish. S.
Afr. 36: 56 pp.
Michael, EK. L., 1911. Classification and vertical distribution of Chaetog-
natha of the San Diego region. Univ. Calif. Publ. Zool. 8: 21-186.
Pierce, E. L., 1951. The Chaetognatha of the west coast of Florida. Biol.
Bull. Woods Hole /00: 206-228.

, 1958. The Chaetognatha of the inshore waters of North
Carolina. Limnol. Oceanogr. 3: 166-170.

Ritter-Zahony, R., 1911. Revision der Chaetognathen. Dtsch. SiidpolExped.
Tis tal Ge lett

Seaccini, A. and E. Ghirardelli, 1941. Chetognati raccolti lungo le coste del
Rio de Oro. Note Ist. Biol. mar. Rovigno 2 (21): 3-16.

o

Suarez Caabro, J. A., 1955. Quetognatos de los mares cubanos. Mem. Soc.
cubana Hist. nat. 22: 125-180.

Sund, P. N., 1959. A key to the Chaetognatha of the tropical eastern
Pacific Ocean. Pacif. Sci. 73: 269-285.

Sund, P. N. and J. A. Renner, 1959. The Chaetognatha of the Eastropic
Expedition, with notes as to their possible value as indicators of hydro-
graphic conditions. Bull. inter-Amer. tropical Tuna Comm. 3: 393-436.

Tokioka, Takasi, 1955. Notes on some chaetognaths from the Gulf of Mex-
ico. Bull. mar. Sci. Gulf Carib. 5: 52-65.

, 1959. Observations on the taxonomy and distribution of
chaetognaths of the North Pacific. Publ. Seto mar. biol. Lab. 7:
349-456.

Vannucci, M. and K. Hosoé, 1952. Resultados cientificos do cruzeiro de

“Baepandi” e do “Vega” a I. Trinidade. Chaetognatha. Bol. Inst.
oceanogr. S. Paulo 3; 5-30.

patella

YALE PEABODY MUSEUM

or NaTuRAL History

Number 56 November 20, 1961 New Haven, Conn.

NOTES ON AMPHISBAENIDS
(AMPHISBAENIA; REPTILIA)

2. Amphisbaena occidentalis Cope from the Coastal Plain of
Northern Peru.

Cart GANS

DEPARTMENT OF BioLtoGy, UNiversIty OF BUFFALO
Burrato, New York

Amphisbaena occidentalis was described by Cope (1876, p.
176; 1885, pl.) from four specimens collected in the “Valley
of Jequetepeque.” Elsewhere (p. 159) in the first paper he
described the locality as extending “from the Cordillera of
Caxamarca to near the coast of Pacasmayo” in western Peru.
The description did not include counts of body annuli nor the
number of segments at midbody, so that Boulenger (1885,
p- 4483; also Strauch, 1881, col. 54) omitted the species from
his catalog as it was “not sufficiently characterized.” Boettger
later (1889, p. 311) provided counts for five specimens col-
lected in the immediate vicinity of Pacasmayo.

Boettger’s paper appears to have been overlooked by Ste)-
neger (1911, p. 203), who relied on a miscount (179 instead
of 279 body annuli) of one of the types and described a single
specimen from Piura as the new species Amphisbaena town-

sendi. Parker (1932, p. 178) obtained the body and tail counts

2 Postilla Yale Peabody Museum No. 56

Figure 1. Map of northwestern Peru to show localities
mentioned in the text.

Noy. 20, 1961 Notes on Amphisbaenids 3

of Cope’s types and placed townsendi into the synonymy of
occidentalis in a note that also listed data for four specimens
from Lobitos, Talara. He remarked that there seemed to be
a north-south gradient in the number of caudal annuli within
the composite sample.

The present redescription was prompted by the discovery
of 14 specimens collected in March 1864 by C. F. Winslow at

La Huaca, River Chira, Peru and now in the collection of the

Chimbote

Chiclin [ae]

-

Pacasmayo a

Jequetepeque

Piura Jig

La Huaca | tl

Negritos L |
AUTOTOMY

Talara nee | |

Lobitos [ |

CAUDAL ANNULI

Figure 2. Amphisbaena occidentalis. Diagrammatic comparison of the
number of caudal annuli for specimens from the several localities. The
annulus at which autotomy occurred is indicated. One of the Pacasmayo
specimens may have a damaged, but not autotomized, tail.

+ Postilla Yale Peabody Museum No. 56

Peabody Museum at Yale University. These and some others
tripled the available number of specimens (all of which have
been reexamined) and doubled the known range of the species.
The data for all specimens are given in the table.

Analysis of these specimens indicated that Parker was cor-
rect, in suggesting that Amphisbaena occidentalis was poly-
morphic, and disclosed a number of other characters in which
the populations differed. The species consists of two clearly
defined forms, ranging from Lobitos to Piura, and from Jeque-
tepeque to Chimbote respectively (Fig. 1). These populations
are differentiated by caudal counts (Fig. 2), body proportions
(Fig. 3), chin shield arrangements (Figs. 4 and 5), and nature
of caudal autotomy (Figs. 6 and 7).

a A.o.occidentalis

Teac

: ar " SYNTYPES °
' o A.o.townsendi
< ; - HOLOTYPE
D> 24 °
S °
©
a (ee) ® A A
0 ® @
ro)
20 © oO
Oo ha A
(2) A
A
@) (e) A A
16 fe)
° A
°
12 fe}
°
°

10 14 18 22 26

Snout - Vent Length - cm

Figure 3. Amphisbaena occidentalis. Scatter diagram of tail versus
snout-vent length for samples of the two races.

Nov. 20, 1961 Notes on Amphisbaenids 5

The lack of material from the region between Piura and
Jequetepeque does not permit a closer definition of the “break”
nor are there intergrade specimens. The decision to call these
two forms races rather than full, geographically replacing,
species has been taken with considerable reservations. It was
based on their general similarity in head shape, head segment
arrangement, counts of body annuli, and the remarkable and
unique color pattern (Fig. 8). Yet the difference between them
is considerably more than that observed in other subspecies
situations in amphisbaenids (Vanzolini, 1951; Gans and Alex-
ander, ms.). There is also some parallel to the situation of
A. silvestrii and A. neglecta in Mato Grosso (Gans, ms.),
involving two possibly sympatric “sibling” species again dis-
tmnct from other forms. These two situations seem made to
order for the attention of local cytotaxonomists.

It may well be true that the hiatus between the ranges of
the two forms of A. occidentalis is not presently inhabited.
Amphisbaenids seem to be highly dependent upon substrate
humidity and all localities from which specimens of A. occi-
dentalis have been examined lie along coastal river courses.
The Desierto de Sechura between Jequetepeque and Piura is
a region without permanent water, in which the water table
may be low enough to prevent the survival of amphisbaenids.

It is a pleasure to acknowledge permission of the following
curators to examine material stored in their institutions (re-
ferred to by the abbreviations in parentheses throughout the
text): Mr. Charles M. Bogert, The American Museum of Nat-
ural History (AMNH); Dr. James E. Bohlke, The Academy
of Natural Sciences of Philadelphia (ANSP); Miss A. G. C.
Grandison, British Museum (Natural History) (BM); Dr.
Robert F. Inger, Chicago Natural History Museum (CNHM) ;
Dr. Philip S. Humphrey, Peabody Museum of Natural History
at Yale University (PMY); Dr. Konrad Klemmer, Sencken-
bergische Naturforschende Gesellschaft (SMF); Dr. Doris
M. Cochran, United States National Museum (USNM); Dr.
Heinz Wermuth, Zoologisches Museum der Universitat, Berlin
(ZMU). I am indebted to Dr. Virginia Cummings for her care
in preparing the drawings. Opportunity to visit several Euro-
pean Institutions was afforded by a grant from the estate of

6 Postilla Yale Peabody Museum No. 56

Leo Leeser and the overall project owes its support to grant
G-9054 from the National Science Foundation.

Amphisbaena occidentalis Cope, 1876.

A small species of Amphisbaena with a dark purplish-brown
dorsal, and light ventral coloration. The dorsal color is pro-
duced by pigmentation of the segments and drops out by
segments along the sides (occasionally on the back) producing
a characteristic mottled effect. The dorsal surfaces of head
and tail are more densely and solidly pigmented than those
of the body. The head is flattened and there is considerable
dorsoventral compression of the pectoral region. The muscle
masses lying over the parietal portion of the skull do not
change the outline of the head in large specimens. The rostral
is small, scarcely visible from above. Pairs of nasals, pre-
frontals and frontals form a suture along the head, with the
posterior edge of the frontals lying somewhat anterior to the
angle of the mouth (for nomenclature see Gans and Alexander,
ms.). The occipitals are no larger than the dorsal segments.
Four supralabials, the second largest. Three infralabials plus a
small fourth scale that also forms the angle of the gape and pro-
jects dorsad around the posterior edge of the last supralabial.

261-79 body annuli from the back of last infralabial to and
including the pore-bearing precloacals. The first four to six
annuli of the neck region shorter than body annuli. Since the
posterior edge of the frontals lies anterior to the fourth infra-
labial the annuli curve forward dorsally. This may be com-
pensated for by an elongation of the dorsal segments of the
7th to 12th annuli (approximately). These annuli thus appear
more or less V-shaped in dorsal view. Three to four dorsal half
annuli (not included in the counts) are generally present in
this region and there is a tendency toward considerable irregu-
lar interdigitation and complexity in the ventral portion of this
(the pectoral) region. The six precloacal annuli also tend
toward irregularities and asymmetries. Segments per midbody
annulus generally 16 or 18 dorsals, 24 or 26 ventrals, 42 or
44 total. Ranges are 16-19 dorsals, 22-27 ventrals, 38 -
47 totals.

Nov. 20, 1961 Notes on Amphisbaenids 7

Dorsal and ventral folding lines (grooves) are present but
not differentiated. The lateral lines may be faintly indicated
in approximately 50% of the specimens, on others they are
only noticeable as aligned intersegmental sutures.

There are four precloacal pores in all specimens, followed
by six to nine precloacal segments, of which the central six
are often large and of equal size. The post-cloacal segments
are generally subject to splitting, yielding 11 to 16 very ir-
regular radial segments. The pores of males are large and
generally pigmented, those of females smaller, faintly indicated
and nonpigmented. No other characters show significant sexual

dimorphism.

KEY TO SUBSPECIES OF A. occidentalis

1. 18 to 21 caudal annuli; no autotomy constriction; no
specimens with autotomized tail; tail shorter (fig. 2); 3
post-genials in first row; little if any elongation of dorsal
segments of trunk annuli7 to1l2....... A. o. occidentalis

2. 22 to 26 caudal annuli; the seventh and/or eighth caudal
annulus narrower, often with pigmented ventral segments,
and with tail constricted at this autotomy level; some
specimens with autotomized tail; tail longer; 4 to 5 post-
genials in first row; marked elongation of dorsal segments
oitrunk-annuly 7 tovlZ ) 2.6 fy.28 ce nae: A, o. townsendi

Amphisbaena occidentalis occidentalis Cope, 1876

Amphisbaena occidentalis Cope, 1876, p. 176. Terra typica:
“Valley of Jequetepeque,” James Orton, col. Syntypes:
ANSP 11355-8.

Locality records:—Peru:—ZMU 9631. Departamento An-
cash: Chimbote CNHM 5661. Departamento La Libertad:
Chiclin CNHM 34290-1. Pacasmayo (Boettger, 1889, 1893;
Boulenger, 1890; Parker, 1932); BM 1889.7.19.1; SMF
11815-18. “Valley of Jequetepeque” (Cope, 1876, 1885, 1892;
Boulenger, 1885; Stejneger, 1911; Camp, 1923; Parker, 1932;
Burt and Burt, 1933) ; ANSP 11355-58.

8 Postilla Yale Peabody Museum No. 56

Amphisbaena occidentalis townsendi Stejneger, 1911,
new combination.

Amphisbaena townsendi Stejneger, 1911, p. 283. Terra typica:
“Piura, Peru,” C. H. T. Townsend, col. Holotype: USNM
47987.

Locality records: — Peru:— Departamento Piura: Piura
(Stejneger, 1911; Burt and Burt, 1930, 1933; Parker, 1932) ;
USNM 47087. La Huaca, Rio Chira PMY 508-1 - 14. Negri-
tos CNHM 5725, 41554. Near Negritos CNHM 38681. Que-
brada Parifas, near Negritos CNHM 8361, 8385. Talara
AMNH 66642. Parifias Valley, northeast of Talara CNHM
8450-51. Lobitos, Talara (Parker, 1932) ; BM 1929.12.12.2-5.
30 miles from Lobitos BM 19382.9.5.9-10.

REFERENCES

Boettger, O., 1889. Herpetologische Miscellen. xi. Nordwest-Peru. Ber.
Senckenberg. naturf. Ges., p. 267-316.

—, 1893. Katalog der Reptilien-Sammlung im Museum der Senck-
enbergischen naturforschenden Gesellschaft in Frankfurt-am- Main.
Frankfurt, ix + 140 p.

Boulenger, G. A., 1885. Catalogue of the lizards in the British Museum
(Natural History). 2nd ed. London, vy. 2, xiii + 497 p.

, 1890. First report on additions to the lizard collection in the
British Museum (Natural History). Proc. Zool. Soc. London, p. 77-87,
(p. 79).

Burt, C. E., and M. D. Burt, 1930. The South American lizards in the
collection of the United States National Museum. Proc. U. S. Nat.
Mus., v. 78, no. 6, p. 1-52.

, 1933. A preliminary check list of the lizards of South Amer-
ica. Transact. Acad. Sci. St. Louis, v. 28, nos. 1-2, p. 1-104.

Camp, C. L., 1923. Classification of the lizards. Bull. Amer. Mus. Nat.
Hist., v. 48, art. 11, p. 289-481.

Cope, E. D., 1876. Report on the reptiles brought by Professor James
Orton from the middle and upper Amazon, and western Peru. Jour.
Acad. Nat. Sci. Philadelphia, ser. 2, v. 8, no. 6, p. 159-83.

, 1885. Twelfth contribution to the herpetology of Tropical
America. Proc. Amer. Philos. Soe., v. 22, p. 167-94.

, 1892. On degenerate types of scapular and pelvic arches in
the Lacertilia Jour. Morphol., v. 7, p. 223-44.

Gans, C., 1962. Notes on amphisbaenids. 3. Redefinition and description of
the Brasilian reptiles dA mphisbaena silvestrii Boulenger and A. neglecta
Dunn and Piatt. Copeia, (in press).

Nov. 20, 1961 Notes on Amphisbaenids 9

Gans, C. and A. A. Alexander, 1962. Studies on amphisbaenids (Amphis-
baenia, Reptilia). 2. On the amphisbaenids of the Antilles. Bull. Mus.
Comp. Zool., (in press).

Parker, H. W., 1932. The status of two Peruvian lizards. Copeia, no. 4,
p- 178.

Stejneger, L., 1911. Description of a new amphisbaenoid lizard from Peru.
Proc. U. S. Nat. Mus., v. 41, no. 1856, p. 283-4.

Strauch, A., 1881. Bemerkungen iiber die Eidechsenfamilie der Amphis-
baeniden. Mel. Biol. Acad. Imp. Sci. St. Pétersbourg, v. 11, p. 355-
479; also in: Bull. Acad. Imp. Sci. St. Pétersbourg, v. 28, no. 8, cols.
45-131.

Vanzolini, P. E., 1950. Contribuicdes ao conhecimento dos lagartos brasi-
leiros da familia Amphisbaenidae Gray, 1825. 1. SObre uma nova sube-
spécie insular da Amphisbaena darwinitti D. & B., 1839. Pap. Avul.
Dept. Zool. (Sao Paulo), v. 9, p. 69-78.

, 1951. Contributions to the knowledge of the Brasilian lizards
of the family Amphisbaenidae Gray, 1825. 6. On the geographical dis-
tribution and differentiation of Amphisbaena fuliginosa Linné. Bull.
Mus. Comp. Zool., v. 106, p. 1-67.

10 Postilla Yale Peabody Museum 7 No. 56

TABLE OF DATA FOR ALL SPECIMENS

Museum Seg- Chin

Number Sex Annuli ments Shield Length
ZMU 9631 a 267+ 3 +17S 46 3 213+ 18
CNHM _ 5661 Q 262+ 3 +20S 18/24 3 160+ 15
CNHM = 34290 Q 266+ 3/4+17S 41 3 192+17
CNHM = 34291 Q 2638+ 3 +20S 16/26 3 243 + 22
BM 1889.7.19.1 2 272+4/5+18S 18/26 3 220-+- 18.5
SMF 11815 a 265+ 4 +198 43 3 223-+- 19
SMF 11816 juv 268+ 3 +195 47? 3 83+ 8.5
SMF 11817 2 272+ 5 +144 18/26 3 243 + 16?
SMF 11818 2 272+ 4 +208 20/28 3 226+ 19
ANSP 11355 3 275+ 3 +18S 18/26 3 ?260- 22
ANSP 11356 Q 270+ 4 +198 44 3 237+ 19
ANSP 11357 é 266+ 4/3+19S 16/24 3 215+17
ANSP 11358 é 269+ 4 +185 44, 3 217+ 18
USNM —§ 47087 a 275+ 2 +(8)24S 18/26 4 208+ 21
PMY 508-1 Q 267+3/4+(7)23S 16/26 4 187+ 20
PMY 508-2 Q 271+ 4 + (8)23S 16/26 4 139+ 14
PMY 508-3 Q 2644+4+ 7A 19/26 5 211+x
PMY 508-4 A 263+ 3 + (8)25S 44, 4 183+ 21
PMY 508-5 Q 272+ 34+ 7A 42 4 191+x
PMY 508-6 3 261+3/2+(8)24S 42 5 145417
PMY 508-7 Q 267+ 3 +(7)23S 44 4 182+ 19
PMY 508-8 A 265+ 3 +(8)22S 16/24 5 165+17
PMY 508-9 Q 266+ 3 + (8)23S 42 4 182+ 19
PMY 508-10 S$ 264+ 34(7)2385 16/26 4 197 +21
PMY 508-11 Q 263+ 3 +(7)24S 18/26 4 195+21
PMY 508-12 é 2714+ 3 +(8)24S 18/26 4 183 + 20
PMY 508-13 Q 266+ 3/4+ ( )24S 44 4 171+ 20
PMY 508-14 juv 272+ 3 4+(8)25S 42 5 89+ 10
CNHM 5725 g 2794+ 8 + (8)28S 16/22 5 219 + 22
CNHM 41554 3} 273+ 2+8A 16/26 5 230+ x
CNHM _ 38681 Q 269+3/4+(7)25S 18/26 5 197 + 23
CNHM _ 8361 ? 271+ 3 +(8)25S 18/26 4 132+ 15
CNHM — 8385 Q 271+ 4 +(7)24S 18/26 4 180 + 18.5
AMNH __ 66642 2 277+ 3 +( )245 18/25 4 153+ 16
CNHM 8450 $ 275+ 34(8)24S 17/24 4 198 + 22
CNHM = 8451 as 277+ 3 + (9)26S 47 5 202 + 22?
BM 1929.12.12.2 Q 271+ 3 +(8)25S 18/26 5 252+ 28
BM 1929.12.12.3 Q 273+ 4 + (8)24S 16/25 5 227 + 24
BM 1929.12.12.4 juv 272+ 5 +(8)23S 18/28 5 111+ 11
BM 1929.12.12.5 juy 278+ 3 +(8)25S 18/26 5 116+ 12
BM 19382.9.5.9 3 269+ 4 +(8)238S 16/24 5 218 + 22
BM 1932.9.5.10 9 279+4/5+ (9)238S 17/26 5 204 + 20.5

Nov. 20, 1961 Notes on Amphisbaenids 1

a
Ss |
Boral
ae
a
rL
LT
1
ei
i

oCH “ee
QUEge oe Se
SCE EE

Figure 4. Amphisbaena o. occidentalis. Dorsal, lateral and ventral views
of the head and neck of CNHM 34290 from Chiclin, Dept. La Libertad.
The line equals 1 mm to scale. (V. Cummings, del.)

12 Postilla Yale Peabody Museum No. 56

Figure 5. Amphisbaena o. townsendi. Dorsal, lateral and ventral views of
the head and neck of PMY 508-9 from La Huaca, Dept. Piura. The line
equals 1 mm to scale. (V. Cummings, del.)

Nov. 20, 1961 Notes on Amphisbaenids 13

i eed ee

ASR

1 See

[Sad

=
L—I =

14 Postilla Yale Peabody Museum No. 56

Figure 6. Amphisbaena o. occidentalis. Ventral view of cloaca and tail
of same specimen as figure 4. Note lack of autotomy level. The line equals
1 mm to scale. (V. Cummings, del.)

Figure 7. Amphisbacna o. townsendi. Ventral view of cloaca and tail of
same specimen as figure 5. Note the narrowing of the eighth postcloacal
annulus indicating the preferred autotomy level. The line equals 1 mm to
scale. (V. Cummings, del.)

Nov. 20, 1961 Notes on Amphisbaenids

ICT
(TTS
WT
(TTT
WT
LTT
WU
et
nrgent

16 Postilla Yale Peabody Museum No. 56

Figures 8-13. Amphisbaena occidentalis. Photographs demonstrating the
color patterns. Figs. 8-9 show 4. 0. occidentalis, and figs. 10-13 show
A. 0. townsendi.

Figure 8. Lateral view of the head of CNHM 34290 from Chiclin,
Dept. La Libertad. Figure 9. Dense dorsal and sharply delimited lighter
ventral coloration shown in three-quarter view of the midbody of the
same specimen. Figures 10-12 show the various stages of lightening
on the dorsal color (by the lack of pigment on an increasing number of
dorsal segments) within a single series, on midbody dorsal views of
PMY 508-4 (Fig. 10), PMY 508-3 (Fig. 11) and PMY 508-9 (Fig. 12)
all from La Huaca, Dept. Piura. Figure 13. Ventral view of cloaca and
tail of CNHM 8385 from Quebrada Parifas, Depto. Piura. Note the
sharp pigmentation of the reduced postcloacal annulus, which marks
the preferred level of autotomy. Also the extremely faint, non-pigmented
precloacal pores in this female specimen.

ile)

Notes on Amphisbaenids

1961

Nov. 20,

ae

SAEs ST spate

DADE,

YALE PEABODY MUSEUM

oF NaTuRAL Hisrory

Number 57 November 30, 1961 New Haven, Conn.

THE PHYLETIC POSITION OF RAMAPITHECUS

.

Etwyn L. Srmowns

Recent discoveries of early Pleistocene hominids at Olduvai
gorge, Tanganyika, by expeditions under the direction of Dr.
L. S. B. Leakey have pushed back certain knowledge of fossil
man almost to the beginning of this epoch. To the extent that
the K-A date suggested for these early men, 1.75 million years,
(Leakey et al. 1961) is accurate, the beginning of the ‘Villa-
franchian” provincial age, and thus of the Pleistocene itself,
is shown to be considerably earlicr than most previous esti-
mates. It therefore scems appropriate that renewed attention
be drawn to the only Plocene fossil primate specimen known
to this writer, which can be defended as being within, or near,
the population ancestral to Pleistocene and subsequent hom-
inids, the type maxilla of Ramapithecus brevirostris at Yale
Peabody Museum.

This maxilla, Peabody Museum No. 13799, was collected
August 9, 1932 by the Yale North India Paleontological Ex-
pedition under Dr. G. E. Lewis (Fig. 1). The geologic occur-
rence of R. brevirostris was first given by Lewis (1934) as
“Hither latest Middle Siwalik [Dhok Pathan Zone] or basal
upper Siwalik [Tatrot Zone ].” However, Lewis (1937) later
determined the horizon of Y.P.M. 18799 as being within the

Nagri zone, which is of Pliocene early Middle Siwalik age.

2 Postilla Yale Peabody Museum No. 57

Gregory et al. (1937) also indicate the level of this specimen
as Nagri.

Consequently, Hooijer and Colbert (1951) seem to have
erred in listing Ramapithecus as occurring only in the Tatrot
zone fauna which they suggest as being very close to the Plio-
Pleistocene boundary. Regardless of these published differences
in age determination the provenance of the specimen is known,
so that, at least potentially, its temporal position can be veri-
fied. Faunal correlations indicate that, even in the unlikely
event that Ramapithecus occurs as late as the Tatrot horizon,
this primate is distinctly older than the “Villafranchian” hom-
inids of Olduvai gorge.

In spite of the significance of Y.P.M. 13799, as being pos-
sibly the earliest known hominid, it has been largely overlooked,
or briefly dealt with in the more recent summaries of hominid
evolution, a common conclusion being that the type is too
fragmentary to permit taxonomic assignment. Actually, such
a conclusion is incorrect and misleading. This right maxilla
provides at least some information as to shape, size or posi-
tioning of the entire upper dentition except for M”’, in that
alveolae of I'*, C are preserved as well as the series P* through
M’. Moreover the base of the nasal aperture can be seen above
the incisors, and, contra Hrdli¢ka (1935), the dental arcade
can be determined as parabolic and not U-shaped, as was
correctly stated by Lewis (1934) in the original description
of this form (see Fig. 2). Some may think (as Hrdliéka did)
that extrapolating from the right maxilla alone, in order to
determine that the disposition of the upper cheek teeth is in
an arcuate line, instead of being arranged in the parallel series
seen in all pongids, is a rather uncertain procedure. However,
at one point (see arrow 1, figure 2) the maxilla reaches nearly,
if not entirely to the point of the palatal intermaxillary suture.
Since we may safely assume that Ramapithecus, like other
vertebrates, was bilaterally symmetrical, if the right maxilla
and its mirror-image are pivoted around this point the amount
of posterior divergence of the cheek tooth rows cannot be
further decreased beyond the arrangement shown in figure 2
without assuming an impossibly long basal diameter for the

Nov. 30, 1961 Phyletic Position of Ramapithecus 3

central incisor pair (figure 2, arrow 2). In fact, the space
allowed for these teeth in figure 2 (in order to be on the safe
side) is intentionally made greater than it is likely to have
been. Preservation of the entire length of the alveolar cavity
of the right central incisor allows for comparative measure-
ments as to its size. The central incisor root of Ramapithecus
is only about half as long as it is in a series of chimpanzees
examined in this connection and which had cheek teeth of the
same absolute size as Y.P.M. 13799. In orangutans the central
incisors have, comparatively, still longer roots than does Pan.
As is well known, possession of large incisors relative to cheek
teeth is a general feature distinguishing both fossil and_liv-
ing pongids from known hominids. In this feature of central
incisor size, as in others, such as the highly arched palate,
Ramapithecus agrees more closely with Hominidae than with
Pongidae.

It is evident that most of the misapprehensions regarding
Ramapithecus now current trace back to Hrdli¢ka’s discussion
of the specimen (1935) in which he insisted that the form
could not be a hominid. Even a casual examination of this
paper is sufficient to show that it bears every evidence of being
a controversial and non-objective contribution. In contrast to
this, all of the hominid resemblances cited for Y.P.M. 13799
by Lewis (1934) appear to this writer to have been correctly
drawn, and these are reinforced by the additional hominid
features called to attention here.

However, another possible source of uncertainty regarding
the genus may derive from a mandible, Peabody Museum No.
13807, assigned by Lewis (1934) to Ramapithecus, but to a
different species R. hariensis. This mandible shows hetero-
morphy in the lower premolars of the sort characteristic of
pongids but which is not known in undoubted Hominidae. In
view of this heteromorphy, not indicated in P** of R. brevi-
rostris and inasmuch as the mandible of R. hariensis comes
from a different locality, and from a horizon that may be
considerably lower in the section, I see no convincing reason
for associating generically the form it represents with that of
the maxilla of R. brevirostris.

4 Postilla Yale Peabody Museum No. 57

What then can be stated as fact regarding the type maxilla
of Ramapithecus? As the species name implies, and as Lewis
originally stated, this primate exhibits a reduction in prog-
nathism, upper incisor size, and in length from the alveolar
border above the incisors to the base of the nasal opening,
when compared to pongids of its general size, whether living
or fossil. This length from nasal aperture to T° in Ramapithe-
cus is approximately 44 per cent of the length of P* - M* (see
arrows, figure 1) while corresponding percentages in a series of
specimens of Pan range from 70 to 98. Specimens of Pongo
and Gorilla examined fall within the range of Pan, in this

proportion.

In addition to the foregoing differences, the upper incisors
and canine, judging from their alveolae, cannot have been as
large as they typically are in even the smallest Great Apes,
a fact also pointed out by Lewis (1934), who remarked: “The
face is very slightly prognathous, as contrasted with recent
Simiidae. There are no diastemata in the dental series. The
canine is small, not an antero-posteriorly elongated trenchant
tusk but a hominid type with a transverse dimension exceeding
the antero-posterior dimension.” Lewis (1934: 163-166) fully
discussed the dental characters of Y.P.M. 13799, consequently
it is unnecessary to repeat this description here. In general,
crown patterns resemble both Dryopithecus and Australopi-
thecus about equally.

Without further extending the polemical atmosphere sur-
rounding this specimen, so unfortunately initiated by Hrdli¢ka,
this writer will simply call attention to his final statement
regarding Ramapithecus, since he appears to be the only per-
son to have studied the actual specimen who has published
doubts as to its hominid status. The significance of this remark,
in the light of modern understanding of the australopithicines as
hominids, seems to have been overlooked. Hrdli¢ka (1935 :36)
observed: “The genus [Ramapithecus |, although in the upper
denture, in general, nearer to man than are any of the Dryopi-
theci or the Australopithecus cannot ... be legitimately estab-
lished as a hominid, that is a form within the direct human
ancestry.” This curious statement, indicates that Hrdlicka

Nov. 30, 1961 Phyletic Position of Ramapithecus 5

would now have to place the genus in the Hominidae since he
regarded it as more man-like than Australopithecus, a genus
universally accepted today by competent students as belong-
ing to this family. Evidently if there are convincing reasons
why Ramapithecus brevirostris should not be regarded as
representing the earliest known hominid they have not been
demonstrated to date.

To contend that the specimen is too inadequate for definite
taxonomic assignment implies that pongids and hominids can-
not be distinguished, even when reasonable information is
available regarding the size, emplacement, structure and ar-
rangement (whether arcuate or parabolic) of nearly all of
the upper dentition, together with several characters of palate
and face as well. Postcranial remains, if found, might make
it easier to assign this primate taxonomically, but the six
or seven distinct approximations to hominid morphology dis-
cussed here for Y.P.M. 13799 provide an adequate basis for
associating it with the latter family. It seems illogical to
choose the alternative of regarding this form as belonging to
an otherwise unknown group of apes, parallelistic toward
hominids but not closely related to them, when it occurs in the
proper time and place to represent a forerunner of Pleistocene
Hominidae.

REFERENCES

Gregory, W. K., M. Hellman and G. E. Lewis, 1937. Fossil anthropoids of
the Yale-Cambridge India Expedition of 1935. Carnegie Inst. Wash.
Publ. No. 495, pp. 1-27, 8 pl.

Hooijer, D. A. and E. H. Colbert, 1951. A note on the Plio-Pleistocene
boundary in the Siwalik Series of India and in Java. Amer. Journ.
Sci., v. 249, pp. 533-538.

Hrdli¢ka, A., 1935. The Yale fossils of anthropoid apes. Amer. Journ. Sci.,
v. 229, pp. 34-40.

Leakey, L. S. B., J. F. Evenden and G. H. Curtis, 1961. Age of Bed I,
Olduvai gorge, Tanganyika. Nature, v. 191, pp. 478-479.

Lewis, G. E., 1934. Preliminary notice of new man-like apes from India.
Amer. Journ. Sci., v. 227, pp. 161-179, 2 pls.

Lewis, G. E., 1937. Taxonomic syllabus of Siwalik fossil anthropoids. Amer.
Journ. Sci., v. 234, pp. 139-147.

Postilla Yale Peabody Museum No. 57

Figure 1

Occlusal view (A) and lateral view (B) of right maxilla of type of
Ramapithecus brevirostris, Y.P.M. 13799.

Nov. 30, 1961 Phyletic Position of Ramapithecus

Postilla Yale Peabody Museum No. OG

Figure 2

Ramapithecus brevirostris, right maxilla, Y.P.M. 13799, and reverse
of same, showing arcuate arrangement of teeth.

Nov. 30, 1961 Phyletic Position of Ramapithecus

Pa F

Bale

YALE PEABODY MUSEUM

oF NaTruraAu Hisrory

Number 58 April 16, 1962 New Haven, Conn.

A NEW DEVONIAN PELECYPOD FROM ALASKA
AND ITS BEARING ON PTERIOID PHYLOGENY

A. Ler McALestTer

One of the most problematic aspects of a recent revision of
some New York Upper Devonian pelecypods (McAlester,
1962a) was the generic status and phylogenetic position of
the well-known Chemung stage pterioid species Cornellites
(formerly Pterinea) chemungensis (Conrad). This species is
known only from the northern Appalachian region where it
has long been considered to be an index fossil to the “type”
Chemung (Chemung stage of Cooper, 1942; see also Williams,
1907, and Chadwick, 1935). Like most “index fossils,” C.
chemungensis first appears fully-developed in the stratigraphic
record and then shows no evolutionary change before it disap-
pears in late Chemung time. It is a common fossil at several
localities which expose the middle Chemung stage horizons, but
it is rare in the lowest and highest horizons of the stage. Most
New York Upper Devonian clams have closely related species
in the prolific earlier faunas of the New York Middle Devonian,
but no probable ancestral or closely related species have been
discovered for C. chemungensis. Furthermore, it is a morpho-
logically distinctive species that can only with difficulty be
forced into any of the standard genera of Devonian pterioids,

c MITH

of

2 Postilla Yale Peabody Museum No. 58

and for this reason it was very tentatively assigned to the
Lower and Middle Devonian genus Cornellites in the revision of
the Chemung faunas. This morphologic distinctiveness and ap-
parent temporal and spatial isolation have combined to make
the origins and relations of C. chemungensis most puzzling.

“Cornellites’ chemungensis

UPPER DEVONIAN
Finger Lakes

|

2
c ov
oom
fe ie
=
oo
Ee

Actinopteria taberi

MIDDLE DEVONIAN
Tioughnioga
stage

Cazenovia
stage

Figure 1. Suggested phylogenetic position of Actinopteria taberi. The
shaded areas show known ranges and relative abundances of Actinopteria
boydi and “Cornellites” chemungensis. The drawings of the species are
schematic and are not intended to represent particular specimens.

Apr. 16, 1962 Devonian Pelecypod From Alaska 3

Recently some unexpected new light has been thrown on this
problem by several collections of poorly-preserved fossils found
at Paleozoic outcrops which were exposed during highway
construction operations near Livengood, Alaska. These speci-
mens were collected by Bond Taber of the U. S. Geological
Survey in the course of stratigraphic investigations in the
Livengood region and were kindly sent to me for identification
by R. B. Neuman and J. T. Dutro of the Geological Survey.

Pelecypods are the dominant element in the faunas from
these outcrops, although some fragmentary gastropods,
brachiopods, and crinoid columnals are also found. The pele-
cypods are of several types including schizodont, grammysioid,
mytiloid, and other forms, but by far the most abundant and
well-preserved element in these collections is a_ distinctive
pterioid pelecypod which is described below as the new species
Actinopteria taberi. This form is morphologically intermediate
between the enigmatic C. chemungensis and the common Ap-
palachian Middle Devonian species Actinopteria boydi. It
therefore suggests that the Upper Devonian species ‘“Cornel-
lites” chemungensis may not be related to the Lower and
Middle Devonian Pterinea-Cornellites stock as has long been
assumed, but may instead have had an independent origin from
an Actinopteria stock. The probable morphologic changes and
time relations in this suggested phylogeny are shown schemati-
cally in fig. 1.

Mr. Taber anticipates further stratigraphic studies and fos-
sil collecting in these rocks, and we hope that this work will pro-
vide additional specimens and data for a full description of the
less common pelecypods of this significant fauna.

Actinopteria taberi, n. sp.

Figures 3-18

Description. Shell of medium size (median length of 14 measurable
specimens 20 mm), inequivalve, left valve moderately convex, right valve
slightly convex in umbonal region, becoming flattened towards margin.
Shape variable, height ranging from 75 to 100 per cent of length (mean
of 12 measurable specimens 80 per cent). Prominent anterior auricle and
posterior wing on both right and left valves, relative length of both auricle
and wing variable. Prominent radial surface sculpture on body of left

4 Postilla Yale Peabody Museum No. 58

valve, becoming somewhat finer on posterior wing (figs. 3, 4). Well-preserved
fragments of left valves show traces of very fine concentric sculpture
(fig. 5). Surface sculpture of right valves consisting of strong radial
ridges on posterior wing only, rest of valve showing faint radial and con-
centric sculpture making up fine reticulate network which is most prom-
inent in umbonal region. Dentition and hinge features unknown. Ligament
unknown. Musculature and interior features unknown. Shell material
unknown.

Types. Holotype: U. S. National Museum No. 140873. Type locality:
locality A (see below), near Livengood, Alaska. Stratigraphic position:
probably near Middle-Upper Devonian boundary as defined in New York
(see below). Additional specimens: U. S. National Museum Nos. 140874-
140896 (No. 140897 questionably this species).

Material. The species is based on 18 relatively complete specimens (11
left and 7 right valves) and about 30 fragments. All of the specimens are
preserved as predominantly external “composite molds” (see McAlester,
1962b). During diagenetic alteration the composite molds have become
colored by a bright orange limonitic stain which clearly distinguishes the
outlines of the former shell material against the much darker matrix. All
of the specimens are permanently deposited in the U. S. National Museum,
Washington, D. C.

Geographic occurrence. The species is known with certainty only from
specimens found at the following two localities in the vicinity of Livengood,
Alaska. Locality A (U. S. Geological Survey field locality number

60ATb85): medium bedded shale and siltstone from borrow pit on north
side of Livengood-Eureka road approximately 7 miles southwest of Liven-
good, Alaska, Lat. 65° 27’N, Long. 148° 43’W, (N13.9 inches, E8.5 inches
from SW corner measured parallel and perpendicular to west margin of
Livengood B-4 quadrangle, Alaska, 1953 edition). Twelve relatively com-
plete specimens and about 20 fragments were found at this locality.
Locality B (U.S. Geological Survey field locality number 60ATD500) : thin-
bedded shales and siltstones with minor medium-bedded, medium to coarse-
grained sandstone from borrow pit approximately 7.7 miles east of Liven-
good, Alaska, at approximately mile 61.4 on Elliott Highway (U.S. 97),
Lat. 65° 29.1’N, Long. 148° 21.7’W, (N16.2 inches, E3.95 inches from SW
corner measured parallel and perpendicular to west margin of Livengood
B-3 quadrangle, Alaska, 1954 edition). Six relatively complete specimens
and about 10 fragments were found at this locality.

Pterioid fragments also occur less commonly at several other borrow
pits along the Livengood-Eureka road west of locality A. Unlike A. taberi,
some of these specimens show right valves with strong, imbricate, concentric
sculpture. Pending discovery of more material which would permit an
analysis of variation in pterioids from these localities, only the specimens
from localities A and B are included in 4. taberi.

Stratigraphic occurrence. The following discussion of possible strati-
graphic relations is based upon comments generously supplied by Mr. Taber
who notes that there is considerable doubt about the age of the exposures
from which the pelecypod faunas were collected. Mertie (1937) mapped the

Apr. 16, 1962 Devonian Pelecypod From Alaska 5

outcrop areas of localities A and B as Devonian non-carbonate or in-
trusive rocks (Dne, Dbi). The two nearest fossil localities mentioned by
Mertie (p. 102 and 120) lie between localities A and B; Mertie’s locality
16AMt64a was about 11% miles southeast of Livengood, and his locality
21AMt128 was about 11% miles south of Livengood. Both collections were
made from limestone beds within essentially clastic sections that Mr. Taber
feels are equivalent to each other and also approximately equivalent to the
clastic sections in which the pelecypods were found. The fossils from
Mertie’s two localities (apparently mostly corals and brachiopods) were
identified by Edwin Kirk, who regarded collection 16AMt64a to be Middle
Devonian (Mertie, p. 103), while 21AMt128 was identified less definitely as
“Devonian or Mississippian, more likely the former” (unpublished U. S.
Geological Survey memorandum). Since Mertie’s localities and the pelecy-
pod bearing clastics may be equivalent, these identifications suggest a
Middle Devonian or, perhaps, younger age for 4. taberi.

The proposed phylogenetic position of A. taberi further strengthens
this suggested age. The exact first appearance in the Appalachian Middle
Devonian of Actinopteria boydi, the proposed ancestral form, has not yet
been established, but it is extremely abundant in some lower Middle
Devonian horizons, particularly the Delphi Station member of the Skaneate-
les formation (see Cooper, 1930, p. 219; also Cooper, 1942). It remains a
locally common form throughout the New York Middle Devonian, but it
becomes rare in the Upper Devonian where it is last known from only a
few Chemung stage specimens. As mentioned earlier, the probable descend-
ent species, “Cornellites” chemungensis, is known only from the Upper
Devonian Chemung stage of the New York region. These relations, which
are summarized in fig. 1, are fully consistent with a late Middle or early
Upper Devonian age for the Alaskan localities which contain the transitional
A. taberi. It should be noted, however, that the Alaskan specimens could
represent a later local survival of the transitional evolutionary stage and
hence be contemporaneous with, or even younger than, the C. chemungensis
bearing rocks of New York. On the other hand, a pre-Middle Devonian age
would be most improbable for these localities if the proposed phylogeny
is correct.

As noted below, slight morphologic differences in the specimens also
vaguely suggest that locality A is the older of the two 4. taberi occurrences.

Comparisons. 4. faberi differs in the following ways from its nearest
probable relatives:

Actinopteria boydi (Conrad): Left valves of A. taberi differ in
showing coarser and more prominent radial sculpture and much finer
concentric sculpture. Right valves of A. taberi show finer and more
regular concentric sculpture, somewhat stronger radial sculpture on
the posterior wing, and obscure traces of radial sculpture on the
body. Both valves differ from A. boydi in having smaller but more
sharply defined anterior auricles and a generally more upright shape.

“Cornellites” chemungensis (Conrad): Left valves of A. taberi
differ in having faint concentric sculpture and less widely spaced
radial sculpture. Right valves of A. taberi have more prominent con-
centric sculpture and finer, less widely spaced radial sculpture on
the posterior wing. Both valves have smaller anterior auricles and a
less upright shape than C. chemungensis.

6 Postilla Yale Peabody Museum No. 58

Discussion. “Cornellites’” chemungensis and Actinopteria
boydi are dissimilar enough so that the evolution of one from
the other would not seem very likely were it not for the transi-
tional Alaskan specimens. The evolution of A. boydi into C.
chemungensis would primarily require an increase in size and
erectness as well as a loss of concentric sculpture, a
strengthening of radial sculpture, and a slight deepening of
the “byssal notch” resulting in a more clearly-defined anterior
auricle. An enlargement of the external ligament area and a
strengthening of the dentition may have also been necessary,
although too little is known of the hinge area in A. boydi to
evaluate this possibility. As illustrated in fig. 2, the Alaskan

“ 4 " RSS
Cornellites chemungensis ~~~
SS
KS actinopteria taberi —~

Sactinopteria boydi SSS
E = A = median value

60 70 80 90 100 110 120 130 140 150
seas Height x 100
Length

Figure 2. Range of shape variation in Actinopteria taberi, Actinopteria
boydi, and “Cornellites” chemungensis. The data for A. boydi were based on
27 Yale Peabody Museum specimens from Pratts Falls, New York. The C.
chemungensis measurements were taken from the 41 complete specimens
described in McAlester, 1962a.

specimens are intermediate in erectness. They also show the
expected strengthening of radial sculpture and weakening
of concentric sculpture, as well as a more clearly-defined
anterior auricle. 4. taberi is not, however, completely ideal as
a morphologic intermediate because several of the specimens
show anterior auricles that are somewhat smaller than those
of either A. boydi or C. chemungensis. This feature and other
more minor differences caution that A. taberi may be some-
what removed from the direct evolutionary line between A.
boydi and C, chemungensis.

Apr. 16, 1962 Devonian Pelecypod From Alaska |

There also appear to be morphologic differences in the
material from each of the Alaskan localities, although too few
good specimens are available to fully evaluate the consistency
of this variation. Specimens from locality A (figs. 3-6, 8, 11-14,
16-18) are generally smaller, less erect, and show slightly finer
sculpture than those from locality B (figs. 7,9, 10,15). The
specimens from locality A are therefore most like the suggested
stem form, 4. boydi, whereas the locality B specimens are
closer to C. chemungensis, the proposed descendent form. This
tentatively suggests that locality A is the older of the two
localities. More material might show these differences, if con-
sistent, to be great enough for two specific names, but it now
seems preferable to consider the specimens as one variable
species. A single right valve found at locality B (fig. 19) is
very erect and has a much larger anterior auricle than any of
the other specimens from either locality. This specimen differs
from the few known right valves of C. chemungensis only in
having finer radial sculpture on the posterior wing with traces
on the body of the valve. For this reason the specimen is doubt-
fully identified as A. taberi and was not included in the above
species description. It may have come from a younger horizon
than the other specimens found at locality B.

The dominance of A. taberi at these localities suggests simi-
lar associations dominated by “Cornellites” chemungensis in the
New York Devonian which have been interpreted as “‘patch-
reef” epifaunal assemblages adapted for life on hard shelly
bottoms (McAlester, 1960). On the other hand, the frag-
mentary preservation of most of the Alaskan specimens also
indicates probable strong current or wave action with
transportation and perhaps mixing of the pelecypod faunas.
Further ecologic interpretation may become possible with the
discovery of new material and the description of the several!
less common pelecypods found at these localities.

No likely descendants of C. chemungensis are now known
and therefore A. taberi and C. chemungensis may well form a
compact and isolated evolutionary group. Normally it would
be appropriate to propose a new generic name for these forms
in view of their morphologic distinctiveness and probable evolu-
tionary isolation. Because, however, of the current generic

8 Postilla Yale Peabody Museum No. 58

chaos in the Paleozoic pterioid pelecypods, I prefer to avoid
adding still another generic name without undertaking a more
complete study of at least the Devonian pterioids. In any event,
such nomenclatural matters are of minor importance in rela-
tion to the more significant evolutionary facts that they are
intended to express. The transitional Alaskan specimens, which
are clearly specifically distinctive, may of course with equal
justification be assigned to the stem genus Actinopteria since
they are morphological intermediates. This course is followed
here pending more complete generic revision. It should be noted,
however, that such revision will almost certainty result in a
new generic name for “Cornellites” chemungensis.

Only one other New York Middle Devonian species is at all
likely to have been ancestral to C. chemungensis. This is the
common species Cornellites (formerly Pterinea) flabella
(Conrad), which is undoubtedly closely related to the Rhenish
Lower Devonian type species of the genus Cornellites. C.
flabella shows some morphologic similarities with C. chemung-
ensis (particularly the upright shape and the distribution of
the sculpture on the right valve) but, in general, I regard the
differences between these two forms to be even greater than
those which separate C. chemungensis from A. boydi. Among
the significant dissimilarities are the very strong reticulate
sculpture, the proportionately larger and differently shaped
auricles, and the strongly inflated left valve and strongly con-
‘ave right valve. In particular the strong convex-concave shape
appears to be an extreme specialization away from the more
typical pterioid pattern of a slightly flattened right valve and
a more gently convex left valve which is the pattern of both
A. boydi and C. chemungensis. The evolution of C. chemung-
ensis from C. flabella therefore now seems to me to be most
improbable, based on our admittedly imperfect knowledge of
New York Middle Devonian pelecypods.

The species is gratefully dedicated to Mr. Bond Taber of
the U.S. Geological Survey to whom I am indebted not only
for collecting the specimens, but also for valuable suggestions
and enthusiastic cooperation. I also wish to express my appre-
ciation to: R. B. Neuman and J. T. Dutro of the Geological
Survey for first sending me the material; to my colleague K.

Apr. 16, 1962 Devonian Pelecypod From Alaska 9

M. Waage for discussions and advice; and to John Howard and

Bruce Umminger who carefully prepared the photographs and
line drawings.

Lirerature Crrep

Chadwick, G. H., 1935. Faunal differentiation in the Upper Devonian:
Geol. Soc. America Bull., vy. 46, p. 305-341.

Cooper, G. A., 1930. Stratigraphy of the Hamilton group of New York,
Pt. 2: Am. Jour. Sci., 5th Ser., v. 19, p. 214-236.

—, chairman, 1942. Correlation of the Devonian sedimentary
formations of North America: Geol. Soc. America Bull. v. 53, p.
1729-1793.

McAlester, A. L., 1960. Pelecypod associations and ecology in the New York
Upper Devonian (abs.): Geol. Soc. America Bull. v. 71, p. 1924.
————., 1962a. Upper Devonian pelecypods of the New York Chemung

stage: Yale Univ. Peabody Mus. Nat. Hist. Bull. 16, 88 p., 32 pl.
, 1962b. Mode of preservation in early Paleozoic pelecypods
and its morphologic and ecologic significance: Jour. Paleontology, v.
36, p. 69-73.
Mertie, J. B., 1937. The Yukon-Tanana region, Alaska: U.S. Geol. Survey
Bull. 872, 276 p., 15 pl.
Williams, H. S., 1907. The Devonian section of Ithaca, New York: Jour.
Geology, v. 15, p. 93-112.

10 Postilla Yale Peabody Museum No. 58

Figures 3-11. Actinopteria taberi, n. sp. All figures are left valves magni-
fied x 1.5.

Figure 3. Holotype, USNM 140873, loc. A. Figure 4. USNM 140874,
loc. A. Figure 5. Latex cast of USNM 140875, loc. A, a fragment of
a left valve showing the concentric sculpture. Figure 6. USNM
140876, loc. A. Figure 7. USNM 140877, loc. B. Figure 8. USNM
140878, loc. A. Figure 9. USNM 140879, loc. B. Figure 10. USNM
140880, loc. B; the posterior and ventral regions are preserved only
as a fragmentary negative composite mold and are therefore illus-
trated from a latex cast; the umbonal and anterior regions preserve

the original positive composite mold. Figure 11. USNM 140881, loc. A.

BI

L

‘
c

Alask

ypod From

in Pelecy

‘
c

evoni

D

Apr. 16, 1962

4

12 Postilla Yale Peabody Museum No. 58

Figures 12-18. Actinopteria tuberi, n. sp. All figures are right valves
magnified x 1.5.
Figure 12. USNM 140882, loc. A. Figure 13. USNM 140883, loc. A.
Figure 14. USNM 140884, loc. A. Figure 15. Latex cast of USNM
140885, loc. B. Figure 16. Latex cast of USNM 140886, loc. A.
Figure 17. USNM 140887, loc. A. Figure 18. USNM 140888, loc. A.

Figure 19. ?Actinopteria taberi, n. sp. Right valve magnified x 1.5.

Figure 19. USNM 140897, loc. B, a doubtfully identified specimen
(see text).

Apr. 16, 1962 Devonian Pelecypod From Alaska 13

_ a Bn ee a

Arnie

YALE PEABODY MUSEUM

oF Natura History

Number 59 April 18, 1962 New Haven, Conn.

BIRDS OBSERVED IN THE IMHOFF GARDENS OF
DHAHRAN, AND AT RAS TANURA, SAUDI ARABIA

1959—1962

Mary G. Eppy*

Dhahran is an oil town with a population of about 3,000
United States citizens, established in 1937 by the Arabian-
American Oil Company as headquarters for its overseas staff.
About ten miles inland from the Persian Gulf coast and sur-
rounded by desert, its gardens have flourished by reason of the
large quantity of natural, slightly brackish water obtained
from wells. Gardens, lawns, hedges of mock honeysuckle, and
trees, date palms, acacia, tamarisk all now grow freely as well
as many ornamental plants such as oleanders and bougainvillea.

Water is used in vast quantities, especially for nearly six
months of air conditioning in the settlement. All of the sewage
water flows into the desert several miles to the east where it 1s
treated and purified. Here at the Imhoff gardens several smal!
ponds have been created, surrounded partly by cultivation,
partly by rank herbage and rushes. This new environment,
where only desert existed previously, has naturally altered the

*Mrs. William Eddy, Beirut, Lebanon.

2 Postilla Yale Peabody Museum No. 59

bird population radically. It was first mentioned by Ripley
(1951) in his paper on a visit to Dhahran and Bahrein in the
summer of 1950. It was of course totally unknown to Ticehurst
and Cheesman (1925) in the only other published short paper
on the area.

In winter the Imhoff gardens with their lakes become a
stopping place for many migrants. In summer, the tamarisk
trees and garden atmosphere provide a welcome shelter from
the burning heat of the open desert, though temperatures may
reach 120 degrees Fahrenheit in the shade. The gardens are
now surrounded by a fence which gives added protection to the
bird fauna, although poachers attempt to shoot at the duck
during the winter migration.

My first visit to these gardens occurred in July, 1950
accompanied by Mr. and Mrs. 8. Dillon Ripley. Since then I
have always visited the Imhoff gardens as well as the sand
spits along the coast near Ras Tanura, the oil refinery forty
miles north along the coast, whenever my husband and I visited
eastern Saudi Arabia during the ensuing years. For texts I
have used Peterson, Mountfort and Hollom’s Field Guide to
the Birds of Europe, and Meinertzhagen’s Birds of Arabia,
both invaluable aids. The following notes should be of mterest
when compared with the earlier records of Ripley, as well as
Meinertzbagen’s book. It is obvious that migrants as well as
local resident species are able to exploit rapidly such a newly
emerging favorable environment such as that at Imhoff. It is
my sincere hope that the following list will encourage further
observations by others in this fascinating spot.

Annotated List from the Imhoft Gardens*

Podiceps nigricollis, Black-necked Grebe. January, 1960 and
1961, and April, 1960. Meinertzhagen (op. cit. p. 438) re-

93

cords this species as “‘scarce

Podiceps ruficollis, Little Grebe. January, 1960 and 1961;
November, 1960. Meinertzhagen (op. cit., p. 439) lists two
sight records for the species, from Aden and Bahrein.

*Editorial comments are by S. Dillon Ripley.

Apr. 18, 1962 Birds in the Imhoff Gardens 3
Phalacrocorax carbo, Cormorant. March, 1959; January,
April, June and November, 1960.

Ardea cinerea, Grey Heron. Seen commonly in January, April,
May, June and November, 1960 and 1961. I thought that
there were immatures among the four or more birds seen on
each visit.

Ardea purpurea, Purple Heron. Seen once in late May-early
June, 1961. The neck feathers appeared buffy; perhaps a
sub-adult.

Nycticorax nycticorax, Night Heron. Seen only once in Jan-
uary, 1960.

Txobrychus minutus, Little Bittern. More than two birds were
seen from May 27 to June 3, 1961 in heavy reeds. Breeding?

Milvus migrans, Black Kite. January, 1961.
Falco tinnunculus, Kestrel. Seen at all seasons.
Falco naumanni, Lesser Kestrel. Seen migrating in March.

Falco peregrinus, Peregrine. April, 1960, and in late May,
1951, a very late date unless the bird was an escape.

Phoenicopterus ruber, Flamingo. Seen once, a single bird in
fon) f=)

late May, 1961.
Anas platyrhynchos, Mallard. January and November, 1960.
Anas crecca, Teal. January, 1960 and 1961.
Anas clypeata, Shoveler. January, April and November.
Anas acuta, Pintail. January and November.

Aythya nyroca, White-eye or Ferruginous Pochard. January,
November and June. The June record was repeated in 1960
and again in 1961 when a pair was seen every day for a week.
This is not only the southernmost record for the species in
Arabia, but also seems exceptionally late in the year.

Aythya ferina, Pochard. Numerous in January and November.

Fulica atra, Coot. Common in January, April and November.

4 Postilla Yale Peabody Museum No. 59

Charadrius squatarola, Grey Plover. January and November.

Charadrius apricarius, Golden Plover. April, 1960. (A valu-
able record, as sight records or specimens from Arabia are
almost nonexistent, Ed.)

Charadrius alewandrinus, Kentish Plover. Resident, seen dis-
playing in late May-early June in the Gardens.

Charadrius dubius, Little Ringed Plover. Seen in January,
April, May-June and November.

Charadrius hiaticula, Ringed Plover. January, April, June
and November.

Calidris alpina, Dunlin. January, April and November.
, i

Calidris testacea, Curlew Sandpiper. Six seen in the Gardens
in late May-June, 1961.

Calidris minuta, Little Stint. Very common in January and
November. A few seen in April, and one or two still lingering
as late as late May-June, 1961.

Tringa totanus, Redshank. January, April and May, (May 3,
vee

Tringa nebularia, Greenshank. January, April and November.
T'ringa stagnatilis, Marsh Sandpiper. January, November.

Tringa ochropus, Green Sandpiper. January. March, April,
and late May-June, 1961. Not recorded so late in Arabia
before.

Tringa glareola, \WWood Sandpiper. January, late May-June,
1961 (a new late record for Arabia, Ed.) and November.

Tringa hypoleucos, Common Sandpiper. January, April, late
May-June, 1961 (a new late record for Arabia), and
November.

Tringa erythropus, Spotted Redshank. January, 1960, and
May 8, 1957. (A first winter record for Arabia, Ed.)

Apr. 18, 1962. Birds in the Imhoff Gardens 5

Calidris canutus, Knot. April, 1960. A first record for Saudi
Arabia, although the species has been recorded from Muscat.

Limosa limosa, Black-tailed Godwit. January, April and
September in various years from 1956 to 1960. Records are
scarce for Arabia.

Himantopus himantopus, Stilt. April, 1960. Called a “rare
straggler” by Meinertzhagen (op. cit., p. 508).

Capella gallinago, Snipe. January, April and November.
Cursorius cursor, Cream-colored Courser. January.
Glareola pratincola, Pratincole. A single bird, late May, 1961.

Larus argentatus (expanded L. fuscus of Meinertzhagen, Ed.)
Herring Gull. January, March, April, and June.

Larus fuscus, Lesser Black-backed Gull. March, April and
November, 1959-60.

Larus ridibundus, Black-headed Gull. January and April.

]
Py,

Chlidonias leucoptera, White-winged Black Tern. Up to eight
individuals in late May-June, 1961; also a single bird in
June, 1960. Third record for Arabia vide Meinertzhagen
(op: cit, p..o17).

Pterocles alchata, Pin-tailed Sandgrouse. Several seen in
November, 1960. Rarely recorded in Arabia, but probably
straggles erratically in the area.

Streptopelia turtur, Turtle Dove. A single dove seen in the
Gardens in late May-June, 1961, Dhahran, April, 1957.

Psittacula krameri, Rose-ringed Parakeet. Now established at
Dhahran and a new record for Saudi Arabia. (Known pre-
viously only from Oman and Muscat, see de Schauensee and

Ripley, 1953, Ed.)

Apus apus, Swift. Several seen in late May-June. (An attempt
should be made to find the Swift as a breeding species in

Arabia, Ed.)

Merops apiaster, Bee-eater. Seen in January and May.

6 Postilla Yale Peabody Museum No. 59

Merops superciliosus, Blue-cheeked Bee-eater, March and April.
Hirundo rustica, Swallow. March, April, June.

Hirundo d. daurica, Red-rumped Swallow. A migrating flock of
twenty birds seen in February, 1962.

Riparia riparia, Sand Martin. Commonly seen in May-June,
far later than recorded by Meinertzhagen, (op. cit., p. 279-

80).
Delichon urbica, House Martin. January, April.

Upupa epops, Hoopoe. Seen in March and April, as well as

July (Ripley, 1951, p. 5).

Lanius excubitor, Great Grey Shrike. Seen throughout the
year. (Subspecies? Ed.)

Lanius minor, Lesser Grey Shrike. May, 1957, a new spring
record for Arabia.

Lanius senator, Woodchat Shrike. January, March, May.

Lanius collurio, Red-backed Shrike. Dates in late May-June are
later than recorded by Meinertzhagen (op. cit., p. 171-2).

Lanius isabellinus, Isabelline or Pale Brown Shrike. Seen in
January, 1960 and 1961.

Galerida cristata, Crested Lark. Seen at every month of the
year; (probably magna, Ed.)

Certhilauda alaudipes, Bifasciated Lark. Seen at various
seasons.

Alauda arvensis, Skylark. (Not unexpected in January, Ed.)

Eremopteryx nigriceps, Black-crowned Finch-lark. Lives on
the desert fringes of the Gardens and Dhahran town.

Sturnus vulgaris, Starling. Seen in January and March at
Dhahran.

Oriolus ortiolus, Oriole. Three to four birds seen singing in late
May-June form a most unusual record for this species which
is not known to breed south of southern Iran.

Apr. 18, 1962 — Birds in the Imhoff Gardens

~t

Corvus corax, Brown-necked Raven. March, April, July.

Pycnonotus leucotis, Bulbul. Common in the Gardens, but now
far less common in Dhahran town since heavy doses of DDT
were given to the lawns to control worms. Thus even in
Arabia pesticides have had an effect on the birds which are
attracted by human habitation. (In spite of Meinertzhagen’s
doubts about this population, op. cit., p. 180, I find Dhahran
birds still paler and larger than mesopotamiae from Iraq,

Ed.)

Muscicapa striata, Spotted Flycatcher, Seen in late May-June
and at the normal spring and fall migration dates. Are some
individuals resident?

Musicapa albicollis, Collared Flycatcher. Seen in March, 1957.

Sylvia curruca, Lesser Whitethroat. April, 1960 (probably
minula which has been taken in Oman and the Hadramaut

in March, Ed.)

Sylvia atricapilla, Black Cap Warbler. April and May, 1957,
seen both at Imhoff and in Dhahran.

Sylvia borin, Garden Warbler. January, April.

Sylvia nana, Desert Warbler. Seen in the desert in late May-
June.

Hippolais languida, Upcher’s Warbler. Seen in late May-June.
Breeding?

Hippolais pallida, Olivaceous Warbler. Seen in late May-June.
This warbler may breed in Arabia, but it is not at present
known to do so.

Hippolais icterina, Icterine Warbler. Seen once in late May-
June. A second record for Arabia.

Phylloscopus trochilus, Willow Warbler. January, March,
April, June and November.

Phylloscopus collybita, Chiffchaff. January, March, April,

June and November.

8 Postilla Yale Peabody Museum No. 59

Phylloscopus sibilatrix, Wood Warbler. Seen once in April,
1960.

Acrocephalus schoenobaenus, Sedge Warbler. Very secretive,
but the churring insect-like call reveals its whereabouts in
the cut-over tamarisk trees near the lake. Seen in late May-

June, a very late record for the species.

Lusciniola melanopogon, Moustached Warbler. More than one
specimen seen on several occasions walking over the rocks
near the lake edge close to bushes and trees. The eye-stripe
was very clear indicating that this was not the Sedge
Warbler. Seen in late May-June.

Prinia gracilis, Graceful Warbler. Resident and seen at all

seasons.

Erithacus megarhynchos, Nightingale. Observed once in late
May-June.

Erithacus svecicus, Bluethroat. Several seen in January and

April.

Phoenicurus phoenicurus, Redstart. January, April, late May-
June, and November, (Not previously recorded from eastern

Arabia, Ed.)
Phoenicurus ochruros, Black Redstart. January and April.
Saxicola rubetra, Whinchat. January, March.
Saxicola torquata, Stonechat. January, March and November.

Oenanthe oenanthe, Wheatear. Seen in January and April.
Oenanthe hispanica, Black-eared Wheatear. April, 1960.
Oenanthe leucomela, Pleschanka’s or Pied Wheatear. March

April records are useful as they are the first from eastern
Arabia,

Oenanthe isabellina, Isabelline Wheatear. January, March,
April, May and November.

Oenanthe finschii, Finsch’s Chat. March, April.

Apr. 18, 1962 — Birds in the Imhoff Gardens 9

Oenanthe vanthoprymna, Red-tailed Chat or Wheatear. Seen
in November, 1960.

Monticola sawatilis, Rock Thrush. January, March and May.

Turdus ericetorum, Song Thrush. Seen in January, 1960 and

1961.

Turdus ruficollis atrogularis, Black-throated thrush. Two
birds seen in gardens of Dhahran, February, 1962.

Anthus pratensis, Meadow Pipit. Seen in January, 1960,
1961, March, 1959 and April, 1960. A first sight record for
Arabia.

Anthus spinoletta, Water Pipit. January, 1960.
Anthus campestris, Tawny Pipit. January and April.

Anthus cervinus, Red-throated Pipit. January, March, April
and May.

Anthus trivialis, Tree Pipit. January, April and November.
Motacilla alba, White Wagtail. January, March, April and
November.

Motacilla flava, Blue-headed Wagtail. March and April.

Motacilla flava, Yellow form or Yellow Wagtail (lutea? Ed.)
March and April.

Motacilla flava feldegg, Black-headed Wagtail. Seen in April,
1960 and again a pair in late May-June. One was panting in
the heat. (A late date for these migrants, Ed.)

Motacilla cinerea, Grey Wagtail. A single record in March,
1959.

Passer domesticus, House Sparrow. Extremely common espe-
cially near the horse corrals.

Carduelis spinus, Siskin. November 13, 1959, a flock of Siskin
in the thickets of Imhoff. I am very familiar with the Siskin
cn migration in Lebanon and I remember that I looked up

10 Postilla Yale Peabody Museum No. 59

Meinertzhagen to see what he had to say and found the Siskin
not mentioned in his Birds of Arabia. The Bannermans in
Birds of Cyprus (1958) call the Siskin a winter visitor and
passage migrant. (New record for Arabia, Ed.)

Emberiza calandra, Corn Bunting. Seen in January, 1960 and
1961. (A new winter record for Arabia, Ed.)

Emberiza hortulana, Ortolan Bunting. April, 1960.

Lirerature Crrep

Bannerman, D. A. and W. M. Bannerman, 1958. Birds of Cyprus. Oliver
and Boyd, Edinburgh and London. 384 pp.

de Schauensee, R.M. and S. Dillon Ripley, 1953, Birds of Oman and Mus-
cat, Proc. Acad. Nat. Sci. Philadelphia, vol. 105, p. 71-90.

Meinertzhagen, R., 1954. Birds of Arabia. Oliver and Boyd, Edinburgh and
London, 624 pp.

Peterson, R., G. Mountfort and P.A.D. Hollom, London, 1954, r.d.
London. A Field Guide to the Birds of Britain and Europe.

Ripley, S.D., 1951. Birds collected and noted round Dhahran, Saudi
Arabia, and Bahrein Island. Yale Peabody Museum of Natural History
Postilla, no. 9, pp. 1-11.

Ticehurst, C. B. and R. E. Cheesman, 1925. The Birds of Jabrin, Jafura,
and Hasa in Central and Eastern Arabia and of Bahrain Island,
Persian Gulf, pp. 1-31.

- - 7

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7 '
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Die

YALE PEABODY MUSEUM

or NaTurAL History

Number 60 May 7, 1962 New Haven, Conn.

NOTES ON A COLLECTION OF BIRDS
FROM SURINAM

Puiuie S. HumpuHrey Anpd Rupour Freunp

During the period November 18 to December 17, 1961, Mr.
and Mrs. Rudolf Freund traveled in Surinam and made small
collections of vertebrates and insects for the Yale Peabody Mu-
seum of Natural History. They collected birds at three locali-
ties in southern Surinam. One of these localities (Kayserge-
bergte Airstrip) had been visited in October of the same year by
Mr. Harry A. Beatty who collected birds there for the Chicago
Museum of Natural History (Blake, 1961). To our knowledge
no previous collections of birds have been made at the two
other localities visited by the Freunds (Sipaliwini Airstrip and
Paru Savannah).

The Freunds collected a total of 117 birds which they pre-
served in 10 percent buffered Formalin. When the specimens
arrived at the Peabody Museum they were washed in water and
then stored in 70 per cent alcohol. All specimens were dried
with a jet of compressed air and then identified ; 32 specimens
were prepared as study skins by Mr. David H. Parsons.

ACKNOWLEDGMENTS

The Freunds’ trip to the interior of Surimam would have
been impossible without the generous assistance of the govern-

2 Postilla Yale Peabody Museum No. 60

ment of Surinam. Mr. and Mrs. Freund and the Peabody
Museum of Natural History are especially grateful to the
following people and institutions: Mr. Harold Nassy, Director
of the Government Information Service; LARECO, the Land
Reclamation Corporation; Mr. Fritz Barends, Mayor of Para-
maribo; Dr. D. C. Geijskes, Director of the Surinam Museum;
Corporal H. P. Pijpers of the Surinam Army; Mrs. Dorothy
Badger, Surinam Tourist Bureau, New York; Surinamse Luft-
vaartmaatschappij (Surinam Airways); and Pan American
Airways.

We are also grateful to Mr. Emmet R. Blake, Chicago
Museum of Natural History; Mr. James Bond, Philadelphia
Academy of Natural Sciences; Dr. Kenneth C. Parkes, Car-
negie Museum; and Dr. Dean Amadon, American Museum of
Natural History, for use of specimens in their care. We are
especially indebted to Mr. F. Haverschmidt for calling to our
attention a species misidentified in the manuscript of this note
and for bringing up to date our comments on three species.
Yr. Wesley E. Lanyon end Mr. George E. Watson kindly
examined specimens for us at the American Museum of Nat-
ural History. Dr. S. Dillon Ripley, Mr. Peter Ames, and

others have given us advice or assistance.

COLLECTING LOCALITIES

Kaysergebergte Airstrip. The Freunds collected 58 birds at
Kaysergebergte Airstrip (see map) from November 23 through
December 6, 1961. This airstrip is one of seven recently built
during “Operation Grasshopper,” a program which is opening
the previously almost inaccessible forested interior of Surinam
to travel. Kaysergebergte Airstrip is a clearing of approxi-
mately 200 by 2000 yards in the tropical forest. Most of the
specimens were collected near or at the forest edge bordering
the airstrip; some were collected along trails from the airstrip

to nearby Oranje Creek.

Sipaliwini Airstrip and Paru Savannah. he Freunds col-
lected 58 birds at Sipaliwini Airstrip and Paru Savannah
December 7 through 11, 1961. Sipaliwini is the southernmost
of seven airstrips constructed during ‘Operation Grasshopper”

May 7, 1962 Birds From Surinam 3

(see map). It is close to the vast Paru Savannah which ex-
tends from British Guiana through southernmost Surinam and
French Guiana down into northern Brazil where it forms the
drainage basin of the northern tributaries of the Amazon.

Suriname

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Map of Surinam showing locations of airstrips and airfields. Black dots
mark localities visited by Mr. and Mrs. Freund.

From Sipaliwini the Freunds traveled by dugout southeast up
the Sipaliwini River to its junction with Vier Gebroeders Creek :
from there they went on foot to a camp established in March,
1961, by Dr. Geijskes and Corporal Pijpers. The camp is

4. Postilla Yale Peabody Museum No. 60

located on Vier Gebroeders Creek which at this point forms
the meeting place of the tropical forest and the Paru Sa-
vannah. In December the Paru Savannah is criss-crossed with
dry creek beds lined with Mauritius Palms, shrubs and lux-
uriant growth of grasses and sedges three feet in height. All
birds were collected in vegetation bordering the dry creek beds.

LIST OF SPECIES

Harpagus bidentatus bidentatus (Latham). 2 skin (J 927),
Sipaliwini River, December 11, 1961. Skin around eye light
gray; cere green; beak black; feet yellow. Bird seen when
it flew across trail in forest and landed forty feet up in
tree at edge of trail; this individual was quite tame.

Buteo magnirostris magnirostris (Gmelin). Specimen in For-
malin (J 776), collected along the roadside of “Garnizoen
Pad,” twenty-five kilometers from Paramaribo, going west
to the Saramacca River, November 19, 1961.

Colinus cristatus sonnint (Temminck). ¢ 2 skins (J 8835,
J 884), Sipaliwini Airstrip, December 8, 1961. These speci-
mens were collected at edge of airstrip in an area of dry,
sandy clay soil with patches of grass and sedges varying
in height from six to eighteen inches. Freund first saw these
birds in a flock of six to eight which did not flush until

after his second shot.

Actitis macularia (Linnaeus). Specimen in Formalin (J 837),
Kaysergebergte Airstrip, December 1, 1961, collected
10:30 p.m. in a mist net set over a drainage ditch at the

edge of the airstrip.

Columbigallina passerina griseola (Spix). Four specimens in
Formalin (J 804, J 809, J 810, J 813), Kaysergebergte
Airstrip, November 23 and 24, 1961. Freund found a nest
on November 23 containing two eggs which hatched Decem-
ber 5. The nest was on the ground in a patch of grass in

an area of dry, sandy soil at edge of airstrip.

May 7, 1962 Birds From Surinam 5

Columbigallina minuta minuta (Linnaeus). Two specimens in
Formalin (J 900, J 901), Paru Savannah, December 10,
1961.

Leptotila rufavilla rufaxilla (Richard and Bernard). Three
specimens in Formalin (J 917, J 925, J 926), Sipaliwini
Airstrip, December 10 and 11, 1961.

Ara manilata (Boddaert). é skin (J 835), palm tree at edge
of Kaysergebergte Airstrip, December 1, 1961; specimen in
Formalin (J 899), Paru Savannah, December 10, 1961.

Pyrrhura picta picta (P.L.S. Miller). Specimen in Formalin
(J 866), trail to Oranje Creek, Kaysergebergte Airstrip,
December 5, 1961.

Piaya melanogaster melanogaster (Vieillot). Specimen in For-
malin (J 898), junction of Sipaliwint River and Vier
Gebroeders Creek, December 9, 1961. Bill brilliant red;
patch of skin around eye yellow and blue. Haverschmidt
(in litt.) notes that he has collected several of these cuckoos
since 1955 and that the species is ‘tnot common.”

Caprimulgus nigrescens nigrescens Cabanis. 6 skin (J 824),
Kaysergebergte Airstrip, November 25, 1961. This night-
jar was first seen in the beam of Freund’s headlamp at
9:00 p.m.; its eyes glowed red. The bird was perched on
a low branch of shrub at the beginning of a forest trail.
? skin (J 941), Sipaliwini, December 11, 1961. This bird
was collected at 4:00 p.m. at the forest edge. Haverschmidt
(1955:70)lists this species as rather rare on savannahs.

Polytmus theresiae theresiae (Da Silva Maia). Sex? skin
(J 868), Kaysergebergte Airstrip, December 6, 1961.

Heliothryx aurita aurita (Gmelin). Specimen in Formalin
(J 943), Sipaliwini, December 11, 1961. Considered rare
by Haverschmidt (1955:76) who lists Neger Kreek as

the only recent locality.

6 Postilla Yale Peabody Museum No. 60

Trogon viridis viridis Linnaeus. Specimen in Formalin (J 935),
Sipaliwini, December 11, 1961. Flesh around eye pale blue:
| ’ ’ ye |
bill gray-black.

Brachygalba lugubris lugubris (Swainson). Specimen in For-
malin (J 920), Sipaliwimi Airstrip, December 10, 1961.

Galbula leucogastra leucogastra Vieillot. Specimen in Formalin
(J 888), Sipaliwini, December 8, 1961; 2 skin (J 933),
Sipaliwini, December 11, 1961. J 888 was collected in a
bamboo thicket.

Bucco tamiata tamiata Gmelin. Sex? skin (J 942), Sipaliwini,
December 11, 1961. Considered by Haverschmidt (1955:
80) to be “rather rare in the savanna belt.”

Monasa atra (Boddaert). Specimen in Formalin (J 873),

Oranje Creek, Kaysergebergte Airstrip, December 6, 1961.

Chelidoptera tenebrosa tenebrosa (Pallas). Four specimens in
Formalin (J 806, J 828, J 829, J 830), Kaysergebergte
Airstrip, November 23 and 28, 1961. Specimen in Formalin
(J 860), trail to Oranje Creek, Kaysergebergte, December
5, 1261. Two specimens in Formalin (J 893, J 939), Sipali-
wini, December 8 and 11, 1961, (J 898, Surinam Museum).

Picumnus exilis buffoni Lafresnaye. Specimen in’ Formalin
(J 885), Sipaliwini, December 8, 1961.

Celeus elegans subsp.? 2 skin (J 819), Kaysergebergte <Air-
strip, November 25, 1961; eve reddish brown; bill bluish

ivory.

Dryocopus lineatus lineatus (Linnaeus). Specimen in Formalin
(J 807), Kaysergebergte Airstrip, November 24, 1961.
Specimen in Formalin (J 906), Sipaliwint Airstrip, De-
cember 10, 1961.

Campephilus rubricollis rubricollis (Boddaert). Three speci-
mens in Formalin (J 826, J 827, J 867), Kaysergebergte
Airstrip, November 26 and December 5, 1961.

May 7, 1962 Birds From Surinam 7

Glyphorynchus spirurus spirurus (Vieillot). ¢ skin (J 825),
Kaysergebergte Airstrip, November 26, 1961.

Niphorhynchus guttatus polystictus (Salvin and Godman).
Specimen in Formalin (J 872), trail to Oranje Creek,
Kaysergebergte Airstrip, December 6, 1961.

Lepidocolaptes albo-lineatus albo-lineatus (uatresnaye). &
skin (J 842), specimen in Formalin (J 871), Kayserge-
bergte Airstrip, December 3 and 6, 1961. Specimen in
Formalin (J 907), Sipaliwini Airstrip, December 10, 1961.

Formicivora rufa subsp. Sex? skin (J 904), Paru Savannah,
December 10, 1961. To our knowledge this species has
not been reported previously for Surinam. Collected along
a dry creek bed lined with Mauritius Palms and heavy
shrubs. This specimen is in streaked female plumage; it is
darker ventrally than most specimens of F. r. chapmani
but is matched by extreme specimens in series at the Amer-
ican Museum of Natural History, and a series before us
from Carnegie Museum, Chicago Museum of Natural His-
tory, Philadelphia Academy, and Yale Peabody Museum
of Natural History.

Thamnophilus punctatus punctatus (Shaw). Specimen in For-
malin (J 913), Sipaliwini Airstrip, December 10, 1961.

Tityra cayana cayana (Linnaeus). Two specimens in Formalin
(J 846, J 847), Kaysergebergte Airstrip, December 3,
1961.

Tyrannus melancholicus despotes (Lichtenstein). Five speci-
mens in Formalin (J 817, J 823, J 831, J 832, J 858),
Kaysergebergte Airstrip, November 25, 28, 29, and De-
cember 5, 1961. Specimen in Formalin (J 886), Sipaliwini
Airstrip, December 8, 1961.

Tyrannus albogularis Burmeister. 2 2 skins (J 805, J 833).
Kaysergebergte Airstrip, November 23 and 29, 1961. Not
previously reported for Surinam. Both specimens were col-

lected at forest edge bordering the airstrip.

8 Postilla Yale Peabody Museum No. 60

Megarhynchus pitangua pitangua (Linnaeus ).wo specimens
in Formalin (J 928, J 929), Sipaliwini, December 11, 1961.

Myiozetetes cayanensis cayanensis (Linnaeus). Specimen in
Formalin (J 874), Kaysergebergte Airstrip, December 6,
1961. Specimen in Formalin (J 896), Paru Savannah,
December 9, 1961.

Myiarchus ferox ferox (Gmelin). Specimen in Formalin (J 910),
Sipaliwini Airstrip, December 10, 1961.

Myiarchus sp? Sex? skin (J 934), Sipaliwini, December 11,
1961. Dr. Wesley E. Lanyon very kindly examined this
specimen; he says (in litt.) that it is an immature bird

and belongs to the feroa-swainsoni complex.

Todirostrum cinereum cinereum (Linnaeus). Specimen in For-
malin (J 889), Sipaliwim, December 8, 1961.

Colopteryx galeatus (Boddaert). 2 skin (J 921), Sipaliwini
Airstrip, December 10, 1961.

Myiopagis gaimardii guianensis (Berlepsch). Specimen in For-
malin (J 940), Sipaliwim, December 11, 1961.

Hirundo rustica erythrogaster Boddaert. Two specimens in
Formalin (J 811, J 816), Kaysergebergte Airstrip, No-
vember 24 and 25, 1961. Listed by Haverschmidt (1955:
118) as a “numerous migrant ... 1m open country in the

coastal area.”

Turdus leucomelas albiventer Spix. 2 skin (J 897), Paru Sa-
vannah, December 9, 1961.

Polioptila plumbea plumbea (Gmelin). Four specimens in For-
malin (J 864, J 865, J 869, J 870), trail to Oranje Creek,
Kaysergebergte Airstrip, December 5 and 6, 1961. Three
specimens in Formalin (J 908, J 909, J 937), Sipaliwini
Airstrip, December 10 and 11, 1961.

May 7, 1962 Birds From Surinam 9

Cyanerpes cyaneus cyaneus (Linnaeus). Three ¢ and one 2
skins (J 845, J 848, J 849, J 850) and one specimen in
Formalin (J 851), Kaysergebergte Airstrip, December 3
(J 845) and 4, 1961. Specimen in Formalin (J 892),
Sipaliwini, December 8, 1961 (Surinam Museum).

Dacnis cayana cayana (Linnaeus). Specimen in’ Formalin
(J 863), Kaysergebergte Airstrip, December 5, 1961.
Specimen in Formalin (J 931), Sipaliwini, December 11,
1961.

Cacicus cela cela (Linnaeus). Specimen in Formalin (J 818),
Kaysergebergte Airstrip, November 25, 1961; bill greenish

ivory, eve light turquoise blue.

Scaphidura oryzivora oryzivora (Gmelin). Specimen in For-
malin (J 859), Kaysergebergte Airstrip, December 5, 1961.
For use of Scaphidura see Parkes (1954: 229).

Icterus cayanensis cayanensis (Linnaeus). Specimen in For-
malin (J 932), Sipaliwini, December 11, 1961. Haver-
schmidt (in litt.) notes that he has collected several since
1955 and that the species is ‘not common” in Surinam.

Tersina viridis occidentalis (Sclater). 6 skin (J 890), sa-
vannah at Sipaliwini, December 8, 1961. Third specimen
reported for Surinam (cf. Blake, 1961:182).

Tanagra violacea violacea (Linnaeus). Four specimens in For-
malin (J 852, J 853, J 861, J 862), Kaysergebergte Air-
strip, December 4+ and 5, 1961.

ba palmarum melanoptera (Sclater). Three specimens

1 Formalin (J 854, J 855, J 856), Kaysergebergte Air-

nee December 4, 1961. Two specimens in’ Formalin
(J 930, J 938), Sipaliwini Airstrip, December 11, 1961.

Tachyphonus surinamus surinamus (Linnaeus). Two specimens
in Formalin (J 911, J 918), Sipaliwini Airstrip, December
10, 1961.

10 Postilla Yale Peabody Museum No. 60

Tachyphonus phoenicius Swainscn. 6 & skins (J 919, J 936),
Sipaliwini Airstrip, December 10 and 11, 1961; specimen
in Formalin (J 944), Sipaliwint Airstrip, December 11,
1961. Upper bill black, lower bill pale gray, feet black in

male and female.

Hemithraupis guira nigrigula (Boddaert). Specimen in For-
malin (J 844), path to Oranje Creek, Kaysergebergte Air-
strip, December 3, 1961.

Hemithraupis flavicollis flavicollis (Vieillot). ¢ skin (J 843),
path to Oranje Creek, Kaysergebergte Airstrip, December
3, 1961. Third record reported for Surinam (cf. Blake,
1961: 182).

Schistochlamys melanopis melanopis (Latham). ¢ skin (J 914),
Sipaliwini Airstrip, December 10, 1961.

Zonotrichia capensis tocantinsi Chapman. 4 skin (J 903),
Paru Savannah, December 10, 1961. The occurrence of
this form in southernmost Surinam confirms Chapman’s
(1940:400) suspicion that the species occurs from lower

Amazonia to the range of nominate capensis.

Sporophila lineola (Linnaeus). 46 skin (J 808), Kayserge-
bergte Airstrip, November 24, 1961. Listed by Haver-
schmidt (1955:186) as “not common in open places in

the coastal area.”

Sporophila minuta minuta (Linnaeus). 2 skin (J 902), Paru
Savannah, December 10, 1961.

Myiospiza humeralis humeralis (Bose). Specimen in Formalin
(J 905), Paru Savannah, December 10, 1961; specimen
in Formalin (J 912), Sipaliwini Airstrip, December 10,
1961.

Emberizoides herbicola sphenurus (Vieillot). Two @ skins
(J 887, J 916), Sipaliwini Airstrip, December 8 and 10,
1961; two specimens in Formalin (J 891, J 915), Sipah-
wini Airstrip, December 8 and 10, 1961.

May 7, 1962 Birds From Surinam 1

Lirerature CIrep

Blake, Emmet R. 1961. New Bird Records from Surinam. Ardea, 49:
178-183.

Chapman, Frank M. 1940. The Post-glacial History of Zonotrichia capensis.
Bull. Amer. Mus. Nat. Hist., 77:381-438.

Haverschmidt, F. 1955. List of the Birds of Surinam. Publications of the
Foundation for Scientific Research in Surinam and the Netherlands An-
tilles, Utrecht; no. 13: 1-153.

Parkes, K. C. 1954. The Generic Name of the Rice Grackle. Condor,

56 :229.

ee

YALE PEABODY MUSEUM

or NaTuRAL History

Number 61 May 10, 1962 New Haven, Conn.

THE AFFINITIES OF THE PINK-HEADED DUCK
(RHODONESSA CARYOPHYLLACEA )

Puitie S. HuMPHREY AND S. Ditton RIPLEY

The rare or extinct Pink-headed Duck of India has had a
checkered taxonomic history, having been placed at one time
or another with the perching ducks (Cairinini), the dabbling
ducks (Anatini), and the pochards (Aythyin1). Delacour and
Mayr (1945 :23-24) in their brilliant revision of the family
Anatidae considered Rhodonessa as belonging to the tribe
Anatini because of similarities in display and posture. Later,
Delacour (1956:197) stated that it “is probably related to
Anas more nearly than to any others, but it may also have
some connection with the pochards, as it somewhat approxi-
mates in proportions the species of Netta, and it has a similar
trachea. It certainly shows no close relationship to the Wocd
Ducks (Cairinini).” Verheyen (1955:22) places Rhodonessa
with the pochards, an alliance which had been suggested earlier
by Garrod (1875 :153-154) on the basis of the trachea. Peters
(1931:170) put Rhodonessa between Malacorhynchus (Pink-
eared Duck) and Aix (Wood Duck) in his subfamily Anatinae,
which also included the dabbling ducks and a variety of other
forms. Phillips (1922:90-938) and Salvadori (1895 :61-63)
placed the genus in a subfamily Plectropterinae among genera
which are now considered to be perching ducks (Cairinini).

SMITHSONIAN aq yp

INSTITUTION

2 Postilla Yale Peabody Museum No. 61

Most recently Johnsgard (1961:78,80) has recognized the
aythyine affinities of Rhodonessa; he considers Rhodonessa a
connecting link (along with Marmaronetta) between the Ay-
thyini and the Anatini. Woolfenden (1961:114), using several
osteological features as evidence, has placed Rhodonessa in the
tribe Aythyini.

Rhodonessa has had an uncertain status because it combines
some of the characters of two very different groups of water-
fowl: the pochards on the one hand, and on the other, ducks
which are better adapted for a more terrestrial existence,
namely the dabbling ducks and perching ducks. This combina-
tion of characters has led some workers to suggest that the
Pink-headed Duck might be a “link” relating in a phylogenetic
sense the pochards and the dabbling ducks.

Humphrey’s interest in this problem was aroused when it
was noted that the trachea of the male Pink-headed Duck is
very similar to tracheae of males of species in the tribe Ay-
thyini, differing from these only in small details. This striking
morphological similarity and Ripley’s interest in the curious
distribution of Rhodonessa and other Indian birds have
prompted us to investigate further the affinities of this puzzling
genus.

ACKNOWLEDGMENTS

Humphrey’s part of this study was supported by the Na-
tional Science Foundation as part of a project on the anatomy
of the trachea of ducks and the classification of that group.

Anatomical specimens examined during the course of this
study were obtained through the generosity of M. C. Downes
(Department of Fisheries and Game, Australia), H. J. Frith
(Wildlife Survey Section, C.S.I.R.O, Australia), R. P. Gros-
senheider, G. S. Hunt, G. V. T. Matthews (The Severn Wild-
fowl Trust), K. C. Parkes, W. H. Partridge, Peter Stetten-
heim, the Departments of Conservation of the states of Michi-
gan and Washington, the Department of Animal Pathology
(Cambridge, England), and the University of Washington
Marine Biological Station at Friday Harbor; this study would
not have been possible without the invaluable help of these
people and institutions. We are indebted to the British Museum
of Natural History, the American Museum of Natural History,

May 10, 1962 The Pink-Headed Duck 3

the United States National Museum and the University of
Michigan Museum of Zoology for the use of specimens in their
care. Many of the specimens examined during the course of
this study were first obtained while Humphrey was at the
University of Michigan Museum of Zoology: we are especially
grateful to R. W. Storer and the late Josselyn Van Tyne for
allowing this material to be incorporated in the collections of
the Yale Peabody Museum of Natural History.

Our thanks must also go to Mrs. Shirley Hartman for pre-
paring the illustrations and to Mr. George E. Watson for

many helpful suggestions.

STRUCTURAL FEATURES

Little is known of the anatomy of the Pink-headed Duck.

Garrod (1875 :153-154) described and figured the trachea and

syrinx of both sexes; Verheyen (1955:22) and more recently

Woolfenden (1961 :14, 41, 52, 54, 114) have commented on
the osteology of the species.

Plumage pattern and general appearance. The Pink-headed
Duck was a long-necked, rather awkward looking bird and at
first sight, little like a pochard in bodily proportions. The pos-
ture of the species and its display habits (discussed below) have
led some authors, notably Delacour and Mayr (1945), to ally
it with the dabbling ducks.

A general comparison of the plumage patterns of the Pink-
headed Duck and all other waterfowl leaves us with the impres-
sion that the species has in this character more in common
with the pochards than with any other group. The coloration of
the Pink-headed Duck (apart from the pink head and neck,
which are in color unique among waterfowl) is very much like
that of the pochards. The similarity is especially noteworthy in
the pattern of coloration of the wing (Ripley, 1950 :903-904).
Rhodonessa lacks an iridescent speculum, and in fact has sec-
ondaries which are practically identical in markings with those

of many pochards.

Trachea. The trachea of the male Pink-headed Duck is in
its general features indistinguishable from the tracheae of
males of the tribe Aythyini. However, it differs in the ag-

4 Postilla Yale Peabody Museum No. 61

gregate of several details of structure from the tracheae of
males of any of the pochards.

We have examined specimens or figures of tracheae of males
of all species in the tribe Aythyini except Netta erythroph-
thalma, Aythya nyroca, and Aythya novae-seelandiae.

Through the courtesy of Dr. James D. McDonald of the
British Museum of Natural History we have been able to study
a specimen of the caudal part of the trachea of a male Pink-
headed Duck. Males of Rhodonessa and the various species of
Aythyini are alike in general conformation of the syringeal
region. The similarity is most striking in’ 1) the form of the
partly bony, partly membranous swelling or dilatation to the
left, and 2) the conformation of the laterally expanded,
partly fused rings antericr to the tracheo-bronchial junction
(Figure 1).

In form of the dilatation to the left, male Rhodonessa differs
from males of the tribe Aythyini as follows:

1) the membrane-covered fenestrae are poorly developed.

2) the dilatation is not as strongly laterally compressed as
in the Aythyini.

3) the dilatation does not extend as far anteriorly (cepha-
lad) as in the Aythyini.

4) the lateral plane of orientation of the dilatation is dorsal
to ventral not dorso-medial to ventro-lateral as in the Aythyini.

5) the dilatation is more expanded or swollen caudally than
in the Aythyini.

In every respect save the last, the dilatation of the caudal
end of the trachea of Rhodonessa is less well developed than in
the Aythyini. In Rhodonessa this dilatation is clearly a some-
what less elaborate version of the same structure in the Ay-
thyini. Rhodonessa resembles no other group of ducks in this
respect.

According to Garrod (1875:154), the trachea of the male
Pink-headed Duck has ‘ta shght fusiform dilatation” anterior
to the syringeal region. Mid-tracheal swellings of one kind or
another occur commonly in only two of the major groups of
waterfowl, the Mergini (Bucephala, Mergus, Histrionicus,
Melanitta) and the Aythyini (several species of Aythya,

May 10, 1962 The Pink-Headed Duck 5

Figure 1. Caudal ends of tracheae of males of A) Rhodonessa caryophyl-
lacea, B) Metopiana peposaca, and C) Aythya affinis; each specimen
drawn in the following views: 1) dorsal, 2) left lateral, 3) caudal,
and 4) ventral. Magnification x 1.18.

6 Postilla Yale Peabody Museum No. 61

Netta). One, and possibly more, species of Anas (Anas versi-
color, but not all species are known anatomically) have a mid-
tracheal swelling.

Garrod (1875:154) describes the caudal end of the trachea
just anterior to the syringeal region of the male Pink-headed
Duck as follows: ‘tthe lower end of the trachea is hardly con-
tracted at all. There is, however, a slight thinning of the
anterior portions of some of the inferior tracheal rings... . a
small, transverse, anterior fenestra being the result.” Garrod’s
figure of the trachea of the male of this species illustrates 13
such fenestrae. A similar modification of the ventral parts of
some of the more caudal tracheal rings occurs in male Clangula
(tribe Mergini), but there are in that species only seven fen-
estrae. Males and females of Sarkidiornis (Cairinini) have fen-
estrae of this kind in the caudal part of the trachea; these
fenestrae are fewer and less well developed in the females.

The structure of the syringeal region of the tracheae of
males of the genera Aythya, Netta, and Metopiana is peculiar
to the group. From the standpoint of the structure of the male
syrinx, Rhodonessa clearly belongs in the tribe Aythyini. The
syringes of males of the tribes Anatini and Cairinini have much
in common structurally and differ significantly from those of
Rhodonessa, Aythya, Netta, and Metopiana.

Humerus. Woolfenden (1959:184) has described a method
of distinguishing “tthe humeri of the Anatinae from those of
the Aythyinae [ classification of Peters, 1931], based on certain
characters of the pneumatic fossa .... In the Anatinae the
fossa is deeper and partially excavates the medial bar. The
construction is such that the palmar surface of the bar is not
completely visible. Furthermore, the fossa usually possesses
many bony struts. In the Aythyinae the pneumatic fossa is
shallower, and the medial bar is essentially continuous with
the shaft, exposing its palmar surface. Struts within the fossa
are rare; in most cases the wall is solid.”

Woolfenden found that the pneumatic fossa of Metopiana
peposaca agrees “in all respects with those of the Anatinae.”
He says further that “this deviation from what seems a reliable
method of distinguishing the two subfamilies may be of phylo-

May 10, 1962 The Pink-Headed Duck i

gentic significance, for Delacour and Mayr (1945:25-26)
consider Metopiana, along with Netta rufina and Aythya
erythrophthalma, to ‘constitute a bridge between the river ducks

S95,

and the more specialized pochards of the genus Aythya....

We have examined the pneumatic fossa of the humerus of
Anas platyrhynchos, A. fulvigula, A. falcata, A. poecilorhyncha,
Aythya marila, A. fuligula, A. ferina, Metopiana peposaca,

Figure 2. Head of left humerus, anconal view; a) Rhodonessa caryophyl-
lacea, b) Aythya marila, c) Metopiana peposaca, a) Mergus serrator,
and e) dnas platyrhynchos. Magnification x 1.75.

Rhodonessa caryophyllacea, Histrionicus histrionicus, Mela-
nitta fusca, M. nigra, Somateria mollissima, Bucephala albeola,
Mergus serrator, M. merganser, and Aix sponsa. Using Wool-
fenden’s criterion for classifying pneumatic fossae into “aythy-
ine” or “tanatine” types, we find the following:
“Anatine” pneumatic fossa: Anas, Metopiana, Rhodonessa,
> > an MES ~~)
Mergus, Aix. (Figure 2.)
“Aythyine” pneumatic fossa: Aythya, Bucephala, Histrion-
icus, Melanitta, Somateria.

8 Postilla Yale Peabody Museum No. 61

The fact that Mergus has an “anatine” pneumatic fossa
suggests to us that this character has undoubtedly arisen in-
dependently in three or more different groups (Mergini, Aythy-
im, Anatini). Delacour and Mayr (1945) have pointed out the
close relationship between Bucephala and Mergus; Humphrey
(1955) has shown that Mergus probably evolved from a
Bucephala-like ancestor. We feel that the “taythyine” pneuma-

tic fossa of Bucephala is evidence that the ‘

‘anatine”’ pneumatic
fossa of Mergus is a derived condition which in no way indicates
relationships with the tribe Anatini. Therefore, the Anas-like
condition of the pneumatic fossa of Metopiana does not neces-
sarily indicate that this genus has any close affinity to the
Anatini. In view of the foregoing, the “anatine” condition of the
pneumatic fossa of Rhodonessa cannot be used as evidence to

clarify the relationships of this genus.

Feet. The feet of the Pink-headed Duck have a number of
characters in common with those of Anas, e.g., lack of a lobe
on the hallux, digits III and IV approximately equal in length
or digit III slightly longer, digits relatively short compared
to length of humerus. However, there are some features of the
foot of Rhodonessa which suggest that its resemblance to the
feet of dabbling ducks is secondarily derived.

The fact that the Pink-headed Duck, by its tracheal anatomy
cbviously a pochard, has feet like a dabbling duck prompted us
to compare the feet of dabbling ducks and pochards. To this
end we measured skeletal elements of the feet of the following

species:
Anas discors 4 Aythya collaris 4
” acuta 3 cc americana ¢
” querquedula ° ee ferina
” gibberifrons 4 ‘i valisineria 6
” cyanoptera 4 ‘ affinis 8
” rubripes 19 ¢ Metopiana peposaca 4
Aythya marila é Netta rufina
7 fuligula ¢ Rhodonessa caryophyllacea &

e nyroca @

May 10, 1962 The Pink-Headed Duck 9

Measurements of these specimens are presented in Table 1
which also includes means, standard deviations, minima and
maxima of the measurements of samples of Anas rubripes and
Aythya affinis.

As is apparent from examination of specimens of dabbling
ducks (Anas) and pochards (Aythya), these two groups of
waterfowl differ in size of foot, the pochards having relatively
much larger feet than the dabbling ducks. Using greatest length
of humerus as an index of general body size, the greatest length
of the tarsometatarsus and of each digit (minus ungual pha-
lanx) was expressed as a per cent of humerus length. These
ratios are presented in Table 2 where it can be seen that in
every case except length of tarsometatarsus the ratios of the
elements of the foot of Rhodonessa are much smaller than
those of Aythya and fall among those of the Anas group. The
tarsometatarsus of Rhodonessa is relatively somewhat shorter
than that of any dabbling duck but rather long for a pochard,
although those of some pochards (Aythya nyroca, A. valisi-
neria, Metopiana) are about the same relative length or slightly
longer. In common with the dabbling ducks and Rhodonessa,
Metopiana has relatively short digits; except for the hallux,
the digits of Netta rufina are also very short. The relative
length of the tarsometatarsus of Netta is short like that of the
more typical pochards.

Possibly there is a difference in the relative lengths of the
humeri of dabbling ducks and pochards correlated with dif-
ferences in the flying abilities of the two groups. Although we
know of no way of testing for this possibility, we doubt that
there is enough of an adaptive difference in relative length of
the humerus in the two groups to invalidate using length of
this element as a measure of body size and as a means of com-
paring the relative lengths of the elements of the foot.

There is a clear cut difference between the species of Anas
and those of Aythya in the relative lengths of digits III and
IV. In Aythya (and Netta rufina) digit IV is longer than digit
III (see Table 3). In Anas digit III is usually longer but may
be equal to or slightly shorter than digit IV. In Anas rubripes
(sample of ten males) digit IV is usually slightly shorter (up
to 3.6 per cent shorter) than digit IIT; in five out of ten speci-

No. 61

Postilla Yale Peabody Museum

10

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152 Postilla Yale Peabody Museum No. 61

mens, digit IV was about the same length as or slightly longer
(up to 1.4 per cent longer) than digit IIL.

Rhodonessa is Anas-like in this character; digit IV is 0.9 to
2.1 per cent (three specimens) shorter than digit III.

We have compared the relative lengths of the phalanges of
the feet of Rhodonessa, Netta, Metopiana, and the various
species of Anas and Aythya using the proximal phalanx of
digit III as the basis for the intramembral proportions. The
relative lengths of the phalanges of Rhodonessa as compared
with those of the pochards and the dabbling ducks lead us to
believe that the dabbling-duck-like foot of Rhodonessa has
evolved from a typical pochard foot. Unfortunately, lack of
material makes it impossible to analyze the variability of the
relative lengths of the phalanges of Rhodonessa. We suspect,
however, that the variability is of the same order of magnitude
as found in a sample of eight Aythya affinis. If this is true, we
see no other explanation for the peculiar phalangeal propor-
tions of the foot of Rhodonessa than that they are the result
of modification of an ancestral pochard-like foot.

The relative length of the proximal phalanx of digit IT of
Rhodonessa is greater than the maximum found for Anas and
well within the minimum range for Aythya. Allowing for
variability, one could safely say that this element is on the
large side for Anas or on the small side for Aythya. The distal
phalanx of digit Il of Rhodonessa is relatively rather short
for either Anas or Aythya. In digit III of Rhodonessa the
second phalanx is relatively shorter than in any of the eight
pochards studied and is slightly below the average of the six
dabbling ducks. The distal phalanx is relatively smaller than
the smallest of Anas and much smaller than in Aythya. The
hallux (digit I) of Rhodonessa is relatively longer than in Anas
and among the shorter of Aythya. (See Table 4.)

In summary, the foot of Rhodonessa is Anas-like in’ 1) the
relative shortness of digits II, WI and IV, 2) the absence of a
lobe on the hallux and 3) the relative lengths of digits III and
IV; it is more pochard-like (but perhaps intermediate) in 1)
length of tarsometatarsus and 2) length of hallux. The
intramembral proportions of the foot of Rhodonessa suggest
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16 Postilla Yale Peabody Museum No. 61

reduction, the more proximal phalanges being least affected
and the distal phalanges most affected. This is most apparent
in digit III but is suggested in digit IT. Digit IV must have
undergone the greatest reduction of all from the presumed
ancestral pochard condition of having been longer than digit
III. Miller (1937 :45) found in his studies of the feet of geese
that “increase or decrease in toe length takes place to a
greater degree in digits two and four than in three, and in-
crease or decrease in phalangeal length takes place to a greater
degree in distal (exclusive of ungual) than in proximal pha-

langes.”’
ges.

BEHAVIOR AND ECOLOGY

There has been little recorded about Rhodonessa beyond the
fact that it has always been observed uncommonly. The Pink-
headed Duck appears to have been a solitary species, occurring
as pairs in the breeding season or small groups in winter (Hume
and Marshall, 1881, 3:176) on isolated marshy ponds or
swampy lakes and not joining the large wintering concentra-
tions of migrant waterfowl. Delacour (1956:198) notes that
its behavior was much like that of dabbling ducks. The species
nested in April in long grass (Andropogon) or grass tufts
sometimes away from the water. The eggs were white and
uniquely spherical (Finn, 1909:25) and the nests well-formed
with no special lining.

The male had a whizzy whistle like a Mallard but lower and
weaker. The female had a low quack. Finn (1909 :86), however,
speaks of the male’s call as quite unlike that of any other duck,
resembling the syllables ‘twugh-ah.”

Males when together (in captivity) were noted to display like
Mallards but more simply. The head was drawn in between the
shoulders while the head feathers were puffed out. Following
this the neck was stretched upwards and the call uttered. How-
ever, as Delacour (1956:198) has observed and described the
display, it was so simple that it lacks any real relationship
to the display of a dabbling duck. It resembled equally well
the simplest forms of aythyine display such as that of
Metopiana for example, lacking only the angular position of the
bill pointed upwards at the climax of the neck-stretch. These

May 10, 1962 The Pink-Headed Duck 1%

observations, in captivity, may have been of birds insufficiently
stimulated to have produced a diagnostic display.

Flight of Rhodonessa is said to have been light and easy
and the habits like those of a true surface-feeder (Finn, 1909:
86). Finn (1915:25) observed an individual dive “as neatly
and long as a pochard.” Perching was never observed.

The Pink-headed Duck was a solitary, non-migratory species
which was local in distribution: the center of its restricted
distribution was the Gangetic area of the “terai” of northern
Bihar. The last record of its occurrence in nature was in 19385
(C. M. Inglis). Jerdon (in Hume and Marshall, 1881, 3:176)
reports that the birds remained out in the center of the pond
during the day and so presumably fed at night. Shillingford
(an Hume and Marshall) reports that a gizzard contained
water weeds and small shells. The duck was said not to have
been a palatable species, indicating an animal diet.

It is unfortunate that so little is known of the habits and
behavior of the Pink-headed Duck. The lack of information is
particularly aggravating considering that this now extinct or
nearly extinct species was at one time not uncommon in col-
lections of European aviculturists where it displayed but never
nested. All that can now be said is that the anatomy and pro-
portions of the foot of Rhodonessa indicate feeding and loco-
motor adaptations very much like those of dabbling ducks. The
sparse behavioral information available provides some if not
overwhelming support for this notion.

DISCUSSION

What sort of history of ecological changes can have pro-
duced the Pink-headed Duck, a pochard with strikingly ana-
tine modifications of the foot? The most likely sorts of environ-
mental changes would have involved a shift from the more
typical pochard habitat, that is, a marine or inland sea littoral
environment, to the present one of marshy, fresh water ponds.
It appears that just such a change took place in India during
Tertiary times.

The Neogene period, characterized by a general regression
in the middle Miocene, brought an isolation of the eastern

18 Postilla Yale Peabody Museum No. 61

Mediterranean into an inland sea. This eastern Mediterranean
province is described by Gignoux (1955 :554-5, 587-8, fig. 144).
Characteristically beginning with the upper Vindobonian or
upper Sarmatian, the inland sea thus formed began to show
isolated genera of invertebrates responsive to decreased salinity
with the outpourings of large fresh water rivers. The eastern-
most of these lacustrine basins is the Aral-Caspian which
stretched north to Samara and east far beyond the Aral Sea.
This eastern basin escaped the reinvasion of the Mediterranean
which occurred in the Quaternary with the opening of the
Dardanelles and the extinguishing of the Levantine and western
Caspian faunas.

The southern parts of this area of trapped inland sea in the
southern Caspian and south of the newly risen Caucasus re-
present part of the old Mesogean or Tethys Sea basin, con-
tinuous through to the Indian Ocean and cut at the beginning
of the Miocene.

Similarly in India, the Miocene system (Wadia, 1944 :256-
275) shows a series of clays and sandstones whose characters
suggest deposition in an estuary or the broad mouth of a river
(Gaj Series in Sind). This shows a regression of the sea border
and its replacement by the wide basin of an estuary. A good
example of this sequence is the Potwar trough which shows de-
posits of nummulitic form over Mesozoic rocks, overlain by
brackish-water sediments of Aquitanian and Burdigalian age
(lower Oligocene and upper Miocene) followed by the fluvia-
tile and sub-aerial Siwalik strata.

How far southward the Siwalik system lies is unclear; much
of it is probably buried under the recent alluvium of the Ganges.
Siwalik birds are interesting in that Phalacrocoraa, Pelecanus,
and Mergus are represented, genera found in India today, and
characteristic of very large lakes or large river systems.

The Pleistocene system in India reveals evidences of glacia-
tion in the form of moraines and polished and grooved rock
at low altitudes in the Himalayas, and also the desiccation
of the Tibetan lakes consequent on the disappearance com-
paratively recently of the glaciers of the ice age.

The plains of India north of the peninsula reveal another
aspect of the Pleistocene in the Indo-Gangetic depression filled

May 10, 1962 The Pink-Headed Duck 19

with thousands of feet of alluvium (Wadia, 1944, fig. 33). The
present river systems bear little relation to their past history
even during historic times. This is a vast area of wandering
rivers and repeated alterations of clays, sand and marls with
recurring layers of peat, lignite and some forest beds. Huge
deltaic areas as in Bengal cover parts cf the former Bay of
Bengal, or in the west, as in the Rann of Kutch, remains of
bays of the Arabian Sea.

All of this intervening plain except for occasional projections
of older rocks must represent the remains of an inland sea,
perhaps formerly continuous with the Aral-Caspian basin of
Miocene times. The whole recent history of this area would
appear to be one of shift from marine to brackish to lacustrine
to riverine conditions. Following this has been the sustained
recent desiccation of central India in historic times (Salm Ah,
1927 :833-861). Seventeenth century records show the range
of the great Indian rhinoceros for example as occurring up te
Peshawar and the foothills of the northwest in what is now a
semi-desert area.

In view of the above it seems very likely that certain animals
adapted for life in or about large bodies of water if capable
of the necessary evolutionary modifications, would have been
able to adjust to this radical change in environment and continue
to persist. Of the four endemic species of Indian birds discussed
by Ripley (1961: xxiii) two are endemic Indian genera whose
normal habitat is aquatic or semi-aquatic. One is the bristled
grass warbler, Chaetornis, a bird of the Gangetic drainage
system, local in habitat, found in moist grassy places. The
other is the Pink-headed Duck, which has been found only in
lakes or ponds in the are from Bihar to eastern Assam, al-
though there are scattered records of nearly a hundred years
ago indicating a wider distribution throughout the Indian
plains and spottily into the Peninsula. If this species indeed
shows affinities with the estuarine diving ducks as we believe
the anatomical evidence suggests, then it would appear to be
descended from a form which was plastic enough to respond
adaptively to long-term changes in the ecology of southern
Asia. It evolved gradually into a fresh water surface feeding
type of duck as the total water area diminished. Again, with the

20 Postilla Yale Peabody Museum No. 61

disintegration of the Aral-Caspian, it became isolated in the
furthest reaches of the increasingly dessicated Gangetic system.
The Pink-headed Duck ultimately became trapped, so to speak,
in the evironment to which it had become adapted and as a
sedentary relict species appears at present to have vanished
almost, if not completely, from the scene.

The evolution of the Pink-headed Duck appears to have been
related to the major environmental changes which took place
in India during Tertiary times. The principal occurrence in the
history of this peculiar genus was the development of dabbling-
duck-like locomotor adaptations, at least so we infer from the
curious modifications of the foot. The trachea, plumage pat-
tern and certain characteristics of the foot leave no doubt
about the aythyine relationships of Rhodonessa.

LIrerRATURE Crrep

Ali, Salim, 1927. The Moghul Emperors of India as Naturalists and Sports-
men. Jour. Bombay Nat. Hist. Soc., 31:833-861.

Delacour, Jean, 1956. The Waterfowl of the World, Vol. 2. Country Life
Limited. London.

Delacour, Jean, and Ernst Mayr, 1945. The Family Anatidae. Wilson Bull.,
57:3-55.

Finn, Frank, 1909. The Water Fowl! of India and Asia. Thacker, Spink and
Co., Caleutta. ix + 121 p.

Finn, Frank, 1915. Indian Sporting Birds. F. Edwards, London. xi + 280 p.

Garrod, Alfred Henry 1875. On the Form of the Lower Larynx in Certain
Species of Ducks. Proc. Zool. Soc. London, 1875:151-156.

Gignoux, Maurice, 1955. Stratigraphic Geology. W. H. Freeman, San
Francisco. xvi + 682 p.

Hume, A. O., and C. H. T. Marshall, 1881. The Game Birds of India,
Burmah, and Ceylon, Vol. 3. Calcutta.

Humphrey, Philip S., 1955. The Relationships of the Sea Ducks (Tribe
Mergini). Ph.D. Dissertation, Univ. Mich.

Johnsgard, Paul A., 1961. The Taxonomy of the Anatidae—A Behavioural
Analysis. Ibis, 103a (1961) :71-85.

Miller, Alden H., 1937. Structural Modifications in the Hawaiian Goose
(Vesochen sandvicensis), a Study in Adaptive Evolution. Univ. Calif.
Publ. Zool., 42:1-80.

Peters, James Lee, 1931. Check-List of Birds of the World, Vol. 1. Harvard
Univ. Press, Cambridge.

Phillips, John C., 1922. A Natural History of the Ducks. Vol. 1. Houghton
Mifflin. Boston.

May 10, 1962 The Pink-Headed Duck 21

Ripley, S. Dillon, 1950. Vanishing and Extinct Bird Species of India. Jour.
Bombay Nat. Hist. Soc., 50:902-906.

Ripley, S. Dillon, 1961. A Synopsis of the Birds of India and Pakistan.
Bombay Natural History Society.

Salvadori, ‘Tommaso, 1895. Catalogue of the Chenomorphae (Palamedeae,
Phoenicopteri, Anseres), Crypturi, and Ratitae in the collection of
the British Museum. Catalogue of the birds in the British Museum,
Vol. 27. London.

Verheyen, René, 1955. La Systématique des Ansériformes Basée sur
VPOstéologie Comparée (suite). Bull. Inst. Roy. Sci. Nat. Belgique, Vol.
31, no. 37:1-22.

Wadia, D. N., 1944. Geology of India. London.

Woolfenden, Glen E., 1959. A Pleistocene Avifauna from Rock Spring,
Florida. Wilson Bull, 71:183-187.

Woolfenden, Glen E., 1961. Postcranial Osteology of the Waterfowl. Bull.
Fla. State Mus., Vol. 6, no. 1:1-129.

56S) 13.

87
balla

YALE PEABODY MUSEUM

oF NaTuraL History

Number 62 June 29, 1962 New Haven, Conn.

THE CRANIAL CRESTS OF
HADROSAURIAN DINOSAURS

Joun H. Ostrom

INTRODUCTION

Ever since 1914 when Lawrence Lambe described and figured
the first of the hooded dinosaurs, there has been considerable
speculation and debate about the function of the hadrosaurian
cranial crest. Although the debate has subsided somewhat in
recent years, it has not been because general agreement has
been reached. Dozens of crested hadrosaurs have been collected
and described since that first discovery and one cannot help
but be impressed by the variety of crestal shapes and_ sizes
represented. There are the relatively small, almost incipient
crests of Procheneosaurus and Cheneosaurus, the large “Corin-
thian helmets” of Corythosaurus and Hypacrosaurus, the “top
hats” of Lambeosaurus, and the curved, tubular crests of
Parasaurolophus. But in spite of the number and diversity of
available specimens, the crestal function has remained quite
elusive.

In 1920, Lambe discovered that the hadrosaurian crest was
not a solid bony structure, but was instead constructed as a
thin and delicate bony sheath which enclosed folded and some-
times complicated passages and chambers. Prior to this dis-

2 Postilla Yale Peabody Museum No. 62

covery, most paleontologists had viewed these cranial prom-
inences simply as interesting decorative features. Lambe’s reve-
lation, however, made such an interpretation quite untenable
and a rash of hypotheses attempting to explain these structures
ensued. The aquatic adaptations of the hadrosaurs had already
been noted by this time and consequently many of these theories
related the complex crestal cavities to the probable semi-aquatic
habits of these ornithischian dinosaurs.

In spite of some popular appeal, and what may appear to
be a certain degree of adaptive significance, most of the crestal
hypotheses that are correlated with an aquatic mode of life
involve serious weaknesses that have generally been overlooked.
This situation, together with certain recently acquired infor-
mation, warrants a careful reconsideration of the hadrosaurian
crest problem.

ACKNOWLEDGMENTS

Several individuals have facilitated this study and I hereby
gratefully acknowledge their aid and advice. My thanks go to
Dr. Edwin H. Colbert of the American Museum of Natural
History, Dr. Rainer Zangerl of the Chicago Natural History
Museum, Dr. L. S. Russell and Mr. Charles M. Sternberg of
the National Museum of Canada, Dr Gordon Edmund of the
Royal Ontario Museum, and Dr. Joseph T. Gregory, formerly
of the Yale Peabody Museum and now of the University of
California. I am also indebted to Dr. Tilly Edinger of Harvard
University and Dr. Nicholas Hotton III of the United States
National Museum, who graciously reviewed the manuscript.

ABBREVIATIONS

Reference is made to specimens in the collections of several
institutions. and in such references the institutional names are
abbreviated as follows:

A.M.N.H. — American Museum of Natural History

C.N.H.M. — Chicago Natural History Museum
N.M.C. — National Museum of Canada, Ottawa
R.O.M. — Royal Ontario Museum, Toronto

Y.P.M. — Peabody Museum of Natural History, Yale

University.

June 29, 1962 Hadrosaurian Dinosaurs 3

Abbreviations used in the text figures are as follows:

AC. — Anterior chamber of nasal capsule
AS. — Anterior space of olfactory chamber
AT. — Anterior nasal tube of nasal capsule

CO. — Concha
CT. — Choanal tube

EN.— External naris
IN. — Internal naris
LC. — Lateral crest cavity

MC.— Medial crest cavity

OB: — Olfactory bulbs

OC. -— Olfactory chamber of nasal capsule
OS. — Antorbital space of olfactory chamber

PREVIOUS THEORIES

Wilham Parks was the first to succumb to the temptation
of theorizing about the functional significance of hadrosaurian
crests. In his description of Parasaurolophus walheri (1922
he argued rather persuasively that the crest of this species
had been joined to the neural spines of the anterior dorsal
vertebrae by means of a strong muscular or ligamentous con-
nection. By inference, at least, this would have facilitated move-
ment of the rather large and unwieldy head. Parks’ hypothesis,
however, was based almost entirely upon what appears to the
present writer to be a pathologic aberration of the sixth and
seventh dorsal spines.

In opposition to Parks’ interpretation, it must be noted that
no identifiable muscle or ligament scars can be seen on the crest
of P. walkeri, or on that of any other crested hadrosaur. It
might also be pointed out that other hadrosaurs, such as
Edmontosaurus and Kritosaurus, possessed far larger heads,
but failed to develop such supporting or leverage structures.
Furthermore, the crest of P. walkeri itself is largely responsible
for the unwieldy nature of the head and therefore it cannot
be seriously considered as an adaptation to counteract its own
disadvantageous effects. And finally, Parks’ hypothesis, pub-
lished two years after Lambe’s discovery, in no way accounts

for the passages within the crest.

4 Postilla Yale Peabody Museum No. 62

In 1924 Othenio Abel published the more fanciful suggestion
that hadrosaurian crests were defensive structures, perhaps
used in mating combat. In the same paper Abel very briefly
considered the possibility that the crest cavities were also
related to olfaction and he compared the crested hadrosaurs
with crocodiles and their somewhat intensified olfactory powers.
This paper was followed in 1929 by Nopesa’s sex character
hypothesis wherein he presumed the crests to be secondary sex
characters, the crested forms being the *tmales” and the non-
crested varieties the “females.” Again, with the exception of
olfaction, neither Abel’s nor Nopesa’s suggestion accounts for
the presence of crestal passages, and Nopesa’s hypothesis
proves somewhat inadequate on other grounds as well. (With
the exception of Parasaurolophus tubicen, only non-crested
“females” have been recovered from the Lance and equivalent
stratigraphic units. )

Carl Wiman’s resonating chamber theory (1931) was the
first to give serious consideration to the most striking feature
of hadrosaurian crests—the crestal cavities. That these cavities
may have served as resonating chambers similar to those found
in certain modern birds, appears entirely plausible, especially
in view of the notoriously noisy habits of modern crocodiles. In
fact, this suggestion is perhaps the best of any theory proposed
so far, but it does seem somewhat improbable that this activity
alone could have been responsible for such extensive cranial
modifications.

A. S. Romer (1933) initiated the aquatic adaptation school
with his comment that there may have been external narial
openings near the top of the crest through which the animal
could have breathed while almost completely submerged. An
elevated or dorsal position for the external nares is a well
known adaptation in certain living and extinct, air breathing,
aquatic animals, but to date no such condition has been verified
in the hadrosaurian crests, in spite of some comments to the
contrary.

A series of papers by Martin Wilfarth (1988, 1939, 1940,
and 1947) expanded the suggestion of Romer’s relating the
crests to an aquatic mode of life. In the first of these papers,
Wilfarth suggested that the flat-headed hadrosaurs were

June 29, 1962 Hadrosaurian Dinosaurs 5

proboscis-bearing reptiles in which there existed a large mus-
cular, “telescoping air tube” that rose from the preorbital fossae
of the snout. This device enabled the animal to “reach” up to
the surface for air without interrupting its underwater feeding.
The deep excavation of the snout is interpreted by Wilfarth
as the origin scar of the large proboscis muscles. In the crested
hadrosaurs, a similar muscular “snorkel” was pictured as at-
tached to the crest, but strangely enough, in the same paper,
Wilfarth accepts Parks’ ligamentous connection between the
dorsal vertebrae and the crest in Parasaurolophus. In later
papers, the “snorkel” interpretation is greatly elaborated by
Wilfarth. The crested hadrosaurs are represented with unique
“upper” external nares located high on the crests. The usual
narial openings situated rostrally on the snout are considered
as nonfunctional relicts closed in life by skin and other tissues.
The large surface area of the crest presumably provided large
areas for the origins of proboscis muscles and inspired air was
conducted down the flexible “breathing tube,” through dorsal
“nares” into the crest cavities and then to the mouth cavity
and trachea. It is further suggested that this remarkable
breathing tube may also have been prehensile and therefore
useful in feeding.

Wilfarth’s imaginative solution to the hadrosaurian crest
problem, appealing and exotic as it may be, is not supported
by any evidence. In the first place, not all of the hollow hadro-
saurian crests are characterized by the required external open-
ings which Wilfarth has interpreted as external nares. The
crest of Cheneosaurus tolmanensis, for example, is not pene-
trated by any lateral or dorsal openings. The same is also
true of Corythosaurus frontalis, Corythosaurus brevicristatus,
and Hypacrosaurus altispinus. And in spite of Wilfarth’s
implications to the contrary (1947), there are no terminal
openings in the crest of either Parasaurolophus walkeri or P.
tubicen and they appear to have been absent in P. cyrtocristatus
as well. Furthermore, the highly varied and irregularly shaped
crestal openings that do occur in some specimens, (Corytho-
saurus casuarius, Lambeosaurus lambei, and Procheneosaurus
praeceps) appear to be due to incomplete ossification at bone
margins, and the areas involved were probably closed by

6 Postilla Yale Peabody Museum No. 62

cartilage or membrane. Still more doubt is cast on Wilfarth’s
hypothesis by the lack of any discernible muscle scars on any
of the hadrosaurian crests. Although the preorbital fossae of
the non-crested hadrosaurs might be considered as excellent
scars left by proboscis muscles, it seems much more likely that
these excavations contained narial sphincter muscles together
with relatively large nasal organs.

Although perhaps not comparable, proboscis-bearing mam-
mals are generally characterized by a marked reduction in the
size of the nasal bones rather than an enlargement as is the
case in many of the crested hadrosaurs. Furthermore, the
nearest thing to a homologous structure, the elephant trunk,
is not used as a “snorkel,” but is a prehensile feeding mecha-
nism and it apparently has always been such a device. If the
hadrosaurian snorkel-like proboscis were also prehensile, as
suggested, it would hardly seem to be functionally advan-
tageous together with an expanded beak, as C. M. Sternberg
(1939) has already pointed out.

Two additional explanations that have received a much
greater deeree of acceptance are similarly correlated with the
semi-aquatic habits of the hadrosaurs. C. M. Sternberg (1935,
1939, 1942, and 1953) and L. S. Russell (1946) consid-
ered the crest as a trapping device which prevented the
entrance cf water into the narial passages and lungs while the
animal was submerged. According to this thesis, the S-shaped
narial loop is considered analogous to an inverted U-tube,
and the greatly elongated crest of Parasaurolophus is rep-
resented as the functional peak of this adaptation. In contrast
to the trap theory is the air storage hyothesis put forward by
E. H. Colbert (1945 and 1955) and Romer (1933 and 1945).
Here, the crestal cavities are interpreted as chambers for
retaining a reserve supply of air which would have enabled the
animal to remain submerged for longer periods. Both of these
capabilities would seem to be highly advantageous adaptations
for air breathing, aquatic animals. However, neither of these
functions seems possible for the structure involved.

The U-tube trap explanation is unsatisfactory for several
reasons. First, the mere presence of an inverted loop in the
narial passage could not in itself have prevented the entrance
of water into the narial passages, or even into the lungs. Water

June 29, 1962 Hadrosaurian Dinosaurs 7

does not rise in the inverted U-tube for the simple reason that
pressures at both ends of the tube are equal. In the case of
the hadrosaurian loop, equal pressures on opposite sides of the
loop could only occur in the emerged state. Even at shallow
depths, hydrostatic pressure would have exceeded the air pres-
sure within the crestal cavities and lungs. Such excessive
hydrostatic pressure would have resulted in compression of
crestal air and the entrance of water into the narial passages
—even in an upward path against the force of gravity. Only
excessive lung pressure (over hydrostatic pressure) could have
prevented water from rising within the narial loop with the
consequent admission of water into the nasal chambers and
possibly into the lungs as well. No modern air breathing verte-
brates, aquatic or semi-aquatic, rely solely on lung pressure
to prevent drowning. Moreover, when one considers that the
far simpler and more effective sphincter valves or vascularized
narial tissues have been developed repeatedly for this very
same function in such diverse aquatic animals as cetaceans,
sirenians, pinnepeds, rodents, crocodiles, lizards, snakes and
amphibians, the proposed water trapping function of the
hadrosaurian crests loses much of its appeal.

Storage of a reserve air supply, for the purpose of prolong-
ing the period of submergence, at first glance seems to be a very
plausible and practical adaptation for semi-aquatic, air breath-
ing animals. However, two quite unrelated factors make such a
thesis highly improbable—if not impossible. First, the available
volume of the crestal chambers appears totally inadequate in
comparison with the probable lung capacity. A very conserva-
tive estimate of the lung capacity of Corythosaurus casuarius,
(A.M.N.H. No. 5338) for example, is approximately 65,000
ec., or less than one fifth of the total volume of the rib cage.
If it be assumed that the entire crest volume of this species
could have been utilized as storage space, the total available
volume of the uncrushed crest probably did not exceed 2500 cc.
Thus in this particular species, the crest volume at best rep-
resented only about four per cent of the total lung capacity,
and it is more than likely that it actually represented a much
smaller fraction. Species with much smaller crests, such as
Procheneosaurus and Cheneosaurus, would have had an even

8 Postilla Yale Peabody Museum No. 62

less significant volume of reserve air. It therefore seems unlikely
that such extreme structural modifications were related to air
storage. Furthermore, it seems somewhat anomalous that a
seml-aquatic vertebrate should have undergone such extreme
modifications for this purpose when purely aquatic air breathers
have been so conservative with regard to such adaptations.

Finally, if we consider the manner in which the proposed
storage cavities would have been utilized, it is immediately ap-
parent that highly undesirable or totally impossible conditions
would have resulted. For example, in order to withdraw this
“reserve air supply” from the crest cavities and into the lungs,
something (either air or water) must have displaced it. Water
seems a most unlikely agent in view of its adverse effects and
the numerous effective precautions against just such circum-
stances that have been repeatedly developed in other aquatic
tetrapods. Air as a displacing substance, obviously was avail-
able only in an emerged state—in which case the “reserve air
supply” was unnecessary. If narial sphincter muscles or vas-
cularized narial valves prevented the entrance of water into
these narial passages, as seems probable, these very same
valves must have prevented air withdrawal from the crest dur-
ing submergence. Thus, in spite of the attractiveness of the air
storage theory, it seems quite improbable on the basis of
current evidence.

NASAL APPARATUS IN MODERN REPTILES

Partial or complete dissection of several hadrosaurian crests
has shown that the crestal cavities are continuous with the
rostrally situated external nares and demonstrates most clearly
that this structure was largely, if not entirely, related to the
nasal apparatus and to some phase of respiratory activity. It
is necessary therefore to examine the nasal anatomy of modern
reptiles and to consider the various functions of the tetrapod
nasal system.

The primary function of the nasal apparatus quite obviously
is the conduction of air from the exterior to the lungs. In addi-
tion, this principal activity requires other preparatory fune-
tions such as cleansing, warming (or cooling), and humidif ying
of the inspired air. (The trapping hypothesis relates to the first

June 29, 1962 Hadrosaurian Dinosaurs 9

of these.) It should not be necessory to consider each of these
functions in detail in order to arrive at the respective merits
of each as an explanation of the structures under consideration.
It is true that temperature regulation or humidification of in-
spired air may be the proper explanation, but all of these seem
improbable in view of the humid, temperate to subtropical con-
ditions indicated for the hadrosaurian environment.

A second major function involving the nasal apparatus is
that of olfaction. It is surprising that this activity, which is
associated with respiratory activities in all tetrapods, has not
been considered more seriously before this. Perhaps the reason
for this may he in the general impression that modern reptiles
do not possess a highly developed sense of smell. This is cer-
tainly true, if one compares them with mammals—particularly
with macrosmatic mammals, but the fact that reptiles do have
olfactory powers cannot be disputed.

Mammals which are characterized by a highly developed
sense of smell show a corresponding expansion of the olfactory
sensory epithelium. This is reflected in the expanded and com-
plex turbinal and ethmoid systems. There is of course no com-
parable ‘tethmoid-turbinal complex” in the hadrosaurs, but the
crestal cavities could well have contained analogous structures.
The question to be answered here is—what was the purpose of
the greatly elongated and complex path of the narial passages ?
Might not this expanded passage have been a means of increas-
ing the surface area of sensory epithelium? The form, pattern
and size of the cavities are quite variable, but in every crested
hadrosaur there has been some increase in the length of the
narial passages through which the inspired air traveled. This
is the only common denominator for all lambeosaurine crests.

Examination of the nasal capsule in modern reptiles reveals
some interesting morphologic evidence pertinent to this subject.
The reptilian nasal capsule (see fig. 1), which is largely car-
tilaginous, encloses the membranous nasal sac. The sac itself
consists of four major parts: the anterior nasal tube, the
anterior chamber, the olfactory chamber, and the choanal tube
(Beecker, 1903; Pratt, 1948; Bellairs and Boyd, 1950; and
Oelrich, 1956). Each of these regions may vary from one form
to another, but they are at least partly distinct from each

10 Postilla Yale Peabody Museum No. 62

other functionally as well as morphologically. The anterior
chamber is principally a preparatory mechanism (filtering,
humidifying, and temperature regulating), while the olfactory

Cr

Fig. 1. Cartilaginous nasal capsule of Clenosaura pectinata, A. Dorsal

view of capsule as seen in rostrally inclined section (see line a-a in dia-
gram C.) showing the anterior part of the nasal canal. B. Dorsal view as
seen in horizontal section through the center of the capsule (see line b-b
in diagram C.) showing the posterior and ventral parts of the nasal canal.
C. Lateral view of the capsule as seen in parasagittal section (see line ¢-¢
in diagram A.) For abbreviations see text.

June 29, 1962 Hadrosaurian Dinosaurs TL

chamber is in large part sensory in function, The olfactory
chamber is further divisible into three regions relating to an
epithelial swelling or protuberance of its lateral wall—the
concha. These are: the anterior space (anterior to the conchal
swelling), the conchal zone, and the antorbital space (posterior
to the concha) (Oelrich, 1956). Ventrally the olfactory cham-
ber (usually in the conchal zone) opens into the choanal tube
and thereby communicates with the oral cavity.

In Sphenodon (see fig. 2) a short anterior nasal tube passes
posteromedially from the laterally situated external naris to a

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Pig. 2. Lateral view of the skull of Sphenodon punctatum illustrating
the locations and relative sizes of the cartilaginous nasal capsule (stippled
pattern) and the olfactory structures of the brain. Notice the long ol-
factory stalks and the olfactory bulbs situated just behind the relatively
small nasal capsule.

short but slightly expanded anterior chamber adjacent to the
nasal septum. The anterior chamber is continuous with a larger
olfactory chamber behind. (Although the olfactory chamber
of Sphenodon is larger than the anterior chamber, it is rela-
tively smaller than the olfactory chamber in most lizards
[ Pratt, 1948].) Within the olfactory chamber, the anterior
space is restricted and the concha occurs as a very weakly
developed swelling of the lateral wall. The antorbital space is

also restricted. Sensory epithelium is limited to relatively small

12 Postilla Yale Peabody Museum No. 62

areas on the medial and dorsal surfaces of the restricted concha,
a relatively much smaller proportion of the available area as
contrasted with certain lizards (Pratt, 1948). Olfactory nerve
fibers arise from these small areas of olfactory epithelium and
pass posteriorly along the lateral wall of the antorbital space
and through the posterior wall of the nasal capsule (planum
antorbitale) where they join the olfactory bulb just behind.
It is important to note here that in spite of the relatively weak
development of the olfactory chamber, the small olfactory
bulbs are situated immediately behind the short nasal capsule
anterior to the orbits.

The relatively short preorbital length, together with the
large eyes of Sphenodon, limits the available space for the nasal
capsule and undoubtedly is related to the restricted develop-
ment of the olfactory chamber and concha. The latter ap-
parently is not related, however, to the limited olfactory sen-
sitivity of Sphenodon, because even the available surface area
of the olfactory chamber is not fully utilized for sensory func-
tions. The limited olfactory powers (Pratt, 1948) would ap-
pear to be in contradiction to the nocturnal, predatory nature
of the animal where an acute sense of smell would seem to be
advantageous, but on the other hand it is consistent with the
burrow-living (but possibly not fossorial) habits of Sphenodon.

Stebbins (1943 and 1948) and Pratt (1948) have demon-
strated that there is considerable variation in the respective
parts of the lizard nasal sac depending upon the mode of life.
For example, in deserticolous lizards the trapping and humidi-
fying functions are more critical and there is a corresponding
enlargement of the anterior chamber. In arboreal lizards, vision
is of greater importance than olfaction and the visual appara-
tus seemingly has expanded and limited the development of the
olfactory chamber. In cursorial forms, particularly in the non-
burrowing varieties, vision and olfaction are both important.
Thus there commonly exists a relatively large eye which limits
the expansion of the olfactory chamber in front. However, in
such cases, the concha is usually greatly expanded and it ap-
pears that conchal expansion is an alternative method of
maintaining or increasing the available area for olfactory
epithelium when other factors (orbital expansion or short snout

June 29, 1962 Hadrosaurian Dinosaurs 13

length) restrict expansion of the olfactory chamber itself
(Pratt, 1948).

In Ctenosaura (see figs. 1 and 3), the anterior nasal tube
extends for a short distance anteromedially from the antero-
laterally placed external naris. Adjacent to the nasal septum it
turns abruptly backward and upward, extending to a long and
expanded anterior chamber. This is continuous dorsomedially
with a narrow anterior space and inflated conchal zone. Pos-
teriorly, a large antorbital space extends ventrally and then
forward to the choanal tube. Within the olfactory chamber,
olfactory epithelium lines the medial and dorsal walls, the
medial and posterior surfaces of the concha, and the posterior
wall of the antorbital space (Oelrich, 1956). Olfactory nerve

Fig. 3. Lateral view of the skull of Clenosaura pectinata showing the
locations and relative sizes of the nasal capsule (stippled pattern) and the
olfactory centers of the brain.

fibers pass posteriorly from these areas, penetrate the planum
antorbitale and unite with the olfactory bulb situated anterior
and dorsomedial to the orbits.

Relative to Sphenodon, Ctenosaura possesses enlarged ol-
factory chambers and considerably more extensive olfactory
epithelium. As in Sphenodon, the olfactory bulbs are situated
far forward, immediately behind the planum antorbitale of the
nasal capsule and in front of the orbits.

In Alligator (see fig. 4), an extremely short anterior nasal
tube descends from the dorsally situated external naris. At its

14 Postilla Yale Peabody Museum No. 62

ventral limit it turns abruptly backward and opens into a very
long and slightly inflated anterior chamber that reaches cau-
dally to the anterior extremity of the palatine. Although
compressed dorsoventrally, the anterior chamber expands lat-
erally reaching its maximum width at about mid-length Pos-
teriorly, the anterior chamber is continuous with a rather large
olfactory chamber. A very prominent concha occupies most of
the olfactory chamber with the result that both the anterior
space and the antorbital space are restricted in volume, al-
though both are of moderate size. The total length of the nasal

Fig. 4. Skull of Alligator mississipiensis as seen in near-sigittal section,
showing the relationships between the olfactory centers of the brain and
the enlarged nasal capsule (stippled pattern). Notice the enlarged anterior
chamber and olfactory chamber as well as the elongated choanal tube.

capsule is approximately equal to the total preorbital skull
length, the planum antorbitale being situated just beneath the
anterior rim of the orbit.

Olfactory epithelium lines portions of the medial, dorsal, and
ventral walls of the olfactory chamber, the dorsal, medial, and
ventral surfaces of the concha, and the posterior wall of the
antorbital space. From these rather extensive sensory areas,
olfactory nerve fibers extend dorsoposteriorly to the moder-
ately sized olfactory bulbs that are placed dorsomedially at
the level of the anterior margins of the orbits. As in Sphenodon
and in lizards, the olfactory bulbs lhe immediately behind the
rear wall of the nasal capsule.

Generally speaking, there is a marked reduction in the ol-

factory sensitivities of aquatic animals, but this does not seem

Or

June 29, 1962 Hadrosaurian Dinosaurs ih

to be true of the Crocodilia. The long snout has provided space
for a relatively long nasal capsule that extends over the entire
preorbital length. Extensive development of sensory epithelium,
an expanded concha, and moderately sized olfactory bulbs all
indicate a significant (if not acute) level of olfactory sensitivity.

NASAL APPARATUS OF THE HADROSAURIDAE

Turning our attention back to the group in question, the
Hadrosauridae are subdivided into three subfamilies (four by
Lull and Wright [1942] and certain other workers) chiefly on
the basis of cranial anatomy. The three subfamilies recognized
here are the Hadrosaurinae, characterized by flat, non-crested
skulls; the Lambeosaurinae, possessing hollow bony crests com-
posed of the premaxillaries and nasals extending over the top
of the cranium; and the Saurolophinae, which bear less promi-
nent, solid, bony crests composed exclusively of the nasals. The
lambeosaurine crests have been referred to as “Strue narial”
crests because they are all hollow and enclose parts of the
elongated and sometimes complicated narial passages. The
various species of Procheneosaurus, Cheneosaurus, Corythosau-
rus, Lambeosaurus, Hypacrosaurus, and Parasaurolophus all
display variations of the true narial crest. Crests of the
Saurolophinae (Saurolophus, Prosaurolophus, and Brachylo-
phosaurus) have been labeled ‘“*pseudo-narial’’ crests, because
of their solid construction and the fact that they do not enclose
any part of the nasal apparatus (Ostrom, 1961b).

Correlated with these cranial conditions is the construction
of the hadrosaurian snout. Those hadrosaurs which bear true
narial crests have greatly reduced snouts, while all other hadro-
saurs are characterized by a long and inflated snout. As a re-
sult, all members of the Hadrosaurinae and the Saurolophinae
have conspicuous preorbital fossae, while the Lambeosaurinae
have virtually no preorbital space available in the snout. It is
quite reasonable to assume that the prominent preorbital ex-
cavation of the hadrosaurine and saurolophine premaxillaries
and nasals housed (at least in part) rather large nasal organs
as suggested by Versluys in 1936. In the Lambeosaurinae, on
the other hand, the snout is greatly compressed dorsoventrally

16 Postilla Yale Peabody Museum No. 62

and there is no large excavation of these bones. Instead, the
premaxillaries and nasals have been prolonged dorsocaudally
and inflated at a more dorsal and posterior level forming the
true narial crest.

In Procheneosaurus the inflated region occurs dorsal to the
maxillaries immediately in front of the orbits. In Cheneosaurus
it occupies a similar position, but has expanded dorsocaudally
and lies over the orbits as well. The inflated zone of Corytho-
saurus, Hypacrosaurus, and Lambeosaurus has expanded still

Cle

Fig. 5. Outline sketch of the skull of Edmontosaurus regalis (N.M.C.
No. 2288) illustrating the probable position of the cartilaginous nasal
capsule (stippled pattern) in the preorbital fossa. The preorbital position
of the olfactory bulbs suggested here is based upon the close proximity of
nasal capsule and olfactory bulbs found in all modern reptiles. Notice that
the expanded snout of the Hadrosaurinae provides ample space for a nasal
capsule and narial sphincter muscles.

further dorsocaudally so that in nearly every instance it hes
well above and behind the orbital level in a supracranial posi-
tion. And in Parasaurolophus, an extreme expansion has car-
ried the inflated region well behind the cranium, as well as
above it, to a postcranial position. These hadrosaurs would

seem to form a sequence of structural stages through which the

-~

June 29, 1962 Hadrosaurian Dinosaurs 17

nasal apparatus has expanded and shifted from a rostral posi-
tion on the snout to a preorbital, to a supraorbital, to a
supracranial, and finally to a partially posteramial position.
(See figs. 6-9.)

Considering the true narial crests in more detail, they all
involve a dorsal and caudal expansion of both the premaxil-

Fig. 6. Outline sketch of the skull of Procheneosaurus cranibrevis
(N.M.C. No. 8633) showing a reconstruction of the narial passages of the
crest. The left premaxillary and nasal have been partially removed to show
the paired lower canals and the “undivided” medial cavity and “choanal”
tube. The arrow indicates the passage of the “choanal” canal from the
crest. (From Ostrom, 1961b.)

laries and the nasals, the degree of expansion and the relative
participation of each varying considerably between different
species. Most conservative are those of the three species of
Procheneosaurus (P. praeceps, P. erectofrons, and P. crani-
brevis) and the only known species of Cheneosaurus (C. tol-
manensis). In these species the premaxillaries are split into

18 Postilla Yale Peabody Museum No. 62

superior and inferior rami or lobes lying above and below the
nares. Both lobes ascend caudally, almost completely enclosing
the anterior portion of the narial passage. The nasal bone
occupies a dorsal position between the upper extremities of the
premaxillary lobes and encloses the upper part of the narial
canal.

Fig. 7. Outline sketch of the skull of Corythosaurus excavatus (N.M.C.
No. 8676) showing a reconstruction of the narial passages of the crest.
The left premaxillary and nasal have been partially removed to show the
paired lower canals and lateral cavities and the “undivided” medial cavity.
The arrow indicates the approximate passage of the “choanal” canal from
the crest. (From Ostrom, 1961b.)

Within the crest of Procheneosaurus (as may be seen in P.
cranibrevis, N.M.C. No. 8633) the narial passages ascend as
separate canals from the rostrally situated external nares. (See
fig. 6.) This portion of the canal, although quite long, may be

analogous to the anterior nasal tube of modern reptiles. Dor-

June 29, 1962 Hadrosaurian Dinosaurs 19

sally, at the anterior limit of the nasal bone, the narial canals
loop forward and then upward in an S-shaped curve and finally
enter a common medial crest cavity anterior to and at the level
of the orbits. (This medial cavity quite probably was divided
by cartilage or membranous tissues in life, consistent with the
paired conditions of modern reptilian nasal structures.) Ventro-
posteriorly, a single (also probably paired in life) ‘‘choanal”
canal descends frem this medial crest cavity into the inter-
orbital region. The latter feature is certainly comparable to
the choanal tube of the modern reptilian nasal capsule and the
undivided cavity above it would appear to represent the loca-
tion of paired olfactory chambers. The S-shaped curve of the
nasal passage then seems to represent the anterior chamber.

The crest of Corythosaurus excavatus (N.M.C. No. 8676),
although differing in details, displays a very similar pattern
(see fig. 7). In this and most other species of Corythosaurus,
the nasal is a very prominent bone forming a major part of the
crest. Again, separate narial passages (anterior nasal tubes?)
ascend posteriorly from the external nares, enclosed for most
of their length by the two lobes of the premaxillaries. In front
of and shghtly above the level of the orbits, the two passages
turn forward and then upward and back again, forming sepa-
rate S-shaped curves (anterior chambers?) similar to those of
Procheneosaurus cranibrevis. Unlike the condition in that
species, however, the paired narial canals lead into paired
lateral cavities above and posterior to the S-shaped loops.
These lateral cavities (olfactory chambers?) occupy the lower
half of the crest, but dorsally they open into a common medial
cavity that extends well up into the crest. Although this latter
cavity could not be fully explored, it apparently extends ven-
trally as a single ‘‘choanal”’ tube between the lateral cavities
to the interorbital region just anterior to the olfactory canal
of the brain case. These relationships indicate a great expan-
sion of the lateral (olfactory) chambers back around the more
medially placed “choanal” tube.

Two specimens of Lambeosaurus (L. lambei, N.M.C. No.
2869, and L. clavinitialis, Y.P.M. No. 3222) display similar
narial patterns characterized by the independently ascending
passages (anterior nasal tubes?) and the S-shaped loops

20 Postilla Yale Peabody Museum No. 62

(anterior chambers?) (see fig. 8). In both of these specimens,
however, the canals remain separated for some distance behind
these loops before entering the large undivided medial cavity

in the posterior half of the crest. As in other forms, this medial

Fig. 8. Outline sketch of the skull of Lambeosaurus clavinitialis (Y.P.M.
No. 3222) showing a reconstruction of the narial passages of the crest.
The lateral elements of the crest have been partially removed to show the
paired condition of the lower canals and the “undivided” dorsal (posterior)
cavity and “choanal” tube. Organic material, perhaps representing cartila-
ginous tissues, found in the posterior cavity indicates this cavity may have
been divided by a median septum in life. The arrow indicates the passage
of the “choanal” canal from the crest. (From Ostrom, 1961b.)

crest cavity (site of the olfactory chambers?) is continuous

.

ventrally with an unpaired ‘tchoanal” tube leading to the
interorbital region just anterior to the brain case.
The crest of Parasaurolophus cyrtocristatus (C.N.H.M. No.

P27393) provides the most detailed information yet available

June 29, 1962 Hadrosaurian Dinosaurs 21

about hadrosaurian crestal cavities (Ostrom, Ms.) (see fig. 9).
Dissection of one side of the crest illustrates that the narial
canals extend as separate passages along the full length of the
crest. From the rostrally situated external nares, the paired
passages (anterior nasal tubes?) ascend in the upper half of the
tubular crest. At the caudal extremity, they loop down to the
lower half of the crest and pass forward to the roof of the skull.
Only at the base of the crest, Just above and behind the orbits
do these paired passages join in a common cavity. (‘These in-
ferior passages and the undivided basal cavity may represent
the location of the olfactory chambers.) Thin medial lamellae

y SoS qe oo
Tri ET

-_eoecrre

CS

Fig. 9. Outline sketch of the crest of Parasaurolophus cyrtocristatus
(C.N.H.M. No. P27393) with the lateral elements removed to show the
pattern of the crestal cavities. ‘The only “unpaired” portion of the passages
occurs at the base of the crest just above the orbits. The ‘“choanal” tube
extends ventrally (arrow) from the medial cavity into the interorbital
region. (From Ostrom, 1961b.)

of the premaxillaries form a continuous wall between the canals
over their entire length, and similar transverse bony walls
separate the upper and lower canals as well. The undivided
medial cavity at the base of the crest communicates with the
interorbital region by way of a ‘“tchoanal’’ tube as in other
crested hadrosaurs. The only subsidiary chambers discovered

22 Postilla Yale Peabody Museum No. 62

in this relatively simple crest plan were a pair of elongated
cavities situated between the upper and lower passages. These
extend caudally from the common cavity for about half of the
total crest length.

In each of these specimens, at least a part of the crest is
that part

of the narial cavities adjacent to the “‘choanal” canal. It is

occupied by a large undivided or unpaired chamber

quite probable that this common cavity, as well as the
“choanal” canal, was actually divided by cartilage or mem-
branous tissue. In support of this are numerous patches of
organic material dispersed throughout the matrix of these
crestal cavities of both Parasaurolophus cyrtocristatus and
Lambeosaurus clavinitialis. These appear to represent remnants
which

of thin sheets of organic material—probably cartilage
were originally situated within the crestal cavities. In view of
the cartilaginous nature of the modern reptilian nasal capsule,
it is quite possible that the hadrosaurian capsule was of similar
construction and that these bits if organic material are rem-
nants of this structure. No similar material could be discovered
in the matrix outside of the crestal cavities, but most of the
matrix had already been removed prior to the present study.
None of the other crests which were examined showed any
comparable material within the accessible parts of the crest.

OLFACTION IN THE HADROSAURS

That the hadrosaurian dinosaurs were like all other tetra-
pods in the possession of a nasal capsule, is not likely to be
disputed. And that this capsule was concerned with the same
fundamental activities as those of modern tetrapods is likewise
beyond doubt. It even seems safe to assume that the hadro-
saurian nasal capsule was probably of the same basic design
as that of modern reptiles. But as to the precise capsule form
and the relative importance of the several nasal functions there
is considerable uncertainty.

The restricted construction of the lambeosaurine snout in-
dicates that the nasal apparatus could not have been located
rostrally. The crest, on the other hand, not only provides a
likely site—it constitutes the only possible site for the lambeo-
saurine nasal capsule. Moreover, the construction of the crest

June 29, 1962 Hadrosaurian Dinosaurs 23

cavities seems to parallel in a very general way the basic pat-
tern of the capsule in modern reptiles, consisting of a long
narrow “anterior nasal tube,” a more inflated and sinuous
“anterior chamber,” a large, inflated “olfactory chamber,” and
a “choanal tube.”

The origin of this expanded nasal “capsule” cannot be ex-
plained by the primary nasal function—air conduction. Nor

Fig. 10. Outline sketch of the skull of Prochencosaurus cranibrevis
(N.M.C. No. 8633) illustrating the dorso-ventrally compressed snout of
the Lambeosaurinae and the inflated narial crest. The stippled pattern in-
dicates the proposed disposition of the nasal capsule within the crestal
cavities and its probable relationship to the olfactory bulbs.

does it seem probable that air preparation was a significant
causal factor. Olfaction, on the other hand, appears very
likely as a principal factor in the development of the lambeo-
saurine crest. As we have seen in the several modern reptiles

24 Postilla Yale Peabody Museum No. 62

considered here, appreciable variation occurs in the total
amount of olfactory epithelium, and the available surface area
may be expanded in certain forms by an invagination (concha)
of the olfactory chamber wall. In mammals, still more complex
folding of the epithelium has resulted in an increase in sensory
epithelial surface area and more acute olfactory powers. With-
in the lambeosaurine crest, the nasal passage has been elon-
gated and in some sections greatly inflated. But not only has
it been lengthened and inflated, the total surface area has also
been increased, perhaps as a parallel means of increasing the
total area available for olfactory epithelium. In support of
this interpretation, it should be noted that the largest of the
crestal cavities is always situated immediately adjacent to the
“choanal” canal and it is this segment of the mcdern reptilian
capsule that is usually involved with sensory activities.

If the large chamber of the lambeosaurine crest did in fact
house the olfactory chambers of the nasal capsule, how were
the contained areas of olfactory epithelium innervated? Several
specimens suggest an answer. Corythosaurus casuarius (R.O.M.
No. 1933), Corythosaurus excavatus (N.M.C. No. 8676),
Lambeosaurus lambei (N.M.C. No. 2869), and Lambeosaurus
clavinitialis (Y.P.M. No. 3222) display a uniform relationship
between the “choanal” passage from the crest and the olfactory
foramen at the anterior limit of the brain case. In each of

.

these specimens, the “choanal” passage opens into the inter-
orbital region immediately in front of the olfactory nerve
canal. The distance between the olfactory foramen of the brain
case and the “choanal” opening of the crest never exceeds 40 mm.
A fifth specimen, Parasaurolophus cyrtocristatus (C.N.H.M.
No. P27398) displays the ‘tchoanal’” opening located in a
similar position that must have been immediately anterior to
the brain case. Unfortunately, however, the brain case is not
preserved so the relationship cannot be verified in this par-
ticular case.

The proximity of these two openings, together with their
respective dimensions, indicate that the olfactory bulbs were
probably located up in the crest cavities and the olfactory
tracts passed downward out of the crestal cavities through

the “choanal”’ canal and then turned backward into the endo-

wor |

June 29, 1962 Hadrosaurian Dinosaurs 2!
5

cranial cavity (see figs. 10 and 11). In fact, no other suitable
site for the olfactory bulbs seems to exist. Additional support
for this interpretation was recently found in Lambeosaurus
clavinitialis (Y.P.M. No. 3222) where remains of what appears
to have been a cartilaginous extension of the walls of the olfac-

ml
WGTHCCAINUOTRILERTALELEA

So Sos

Fig. 11. Outline sketch of the skull of Lambeosaurus clavinitialis
(Y.P.M. No. 3222) showing the dorso-ventrally compressed snout and the
greatly inflated narial crest. The stippled pattern indicates the generalized
arrangement of the nasal capsule proposed here together with the probable
location of the olfactory bulbs.

tory nerve canal passes from the brain case up into the crestal

cavities through the ‘

‘choanal” canal. Strange as such a posi-
tion may appear, it conforms only with a crestal position of
the olfactory chambers and is entirely consistent with the mod-
ern reptilian condition where the olfactory bulbs are located

immediately adjacent to the nasal or olfactory sac.

26 Postilla Yale Peabody Museum No. 62

CONCLUSION

From all available evidence, it appears quite likely that the
lambeosaurine crest was an adaptation for increasing olfactory
sensitivity by providing an increase in the total surface area
available for olfactory epithelium. (It also seems probable,
in view of the very large and unique preorbital fossae, that the
non-crested hadrosaurs had also developed enlarged or modified
nasal apparatus situated in a more normal position in the in-
flated snout. See fig. 5.) But the inevitable question arises, why
should the hadrosaurs, rather than any other dinosaur, have
required superior olfactory powers? Of course this question
cannot be answered to the complete satisfaction of everyone
concerned, but some interesting possibilities are suggested by
reconsidering hadrosaurian ecology.

It is quite probable that hadrosaurs lived a rather passive,
perhaps even retiring existence as relatively slow moving, ter-
restrial or amphibious herbivores, That they were able to move
about over the land is verified by their skeletal construction
and that they may have been terrestrial browzers, at least in
part, is suggested by Krausel’s(1922) analysis of the “stomach
contents” of Anatosaurus (T'rachodon) annectens. Other evi-
dence (the large laterally compressed tail and the webbed
manus) points to a certain dependency upon an aquatic en-
vironment—perhaps for food in the form of soft aquatic plants
or perhaps as a place of refuge from terrestrial predators.

In comparing the hadrosaurs with other dinosaurian herbi-
vores, it is striking that they alone lack any obvious defensive
or protective adaptations. They possessed no horns, no claws,
no sharp teeth, they carried no clubbed or spiked tail, and they
had no bony armor. They certainly were not constructed for
rapid flight and they cannot be considered giants for their
time. In short, the hadrosaurs appear to have been quite
defenseless—a most improbable plight. As an alternative it
seems increasingly probable that they depended upon the rela-
tive security of lakes, swamps, or rivers and thereby escaped
from their enemies.

However, such inland waters represented only potential
safety as long as the individual was out on the land—potential
safety contingent upon adequate advance detection of the im-

bo

~
(

June 29, 1962 Hadrosaurian Dinosaurs

pending danger by one or more of the sensory systems. Here
is where acute olfactory sensitivity may have had significant
survival value for the hadrosaurs, just as it does for many
modern mammalian herbivores.

BrIBLioGRAPHY

Abel, Othenio, 1924. Die neuen Dinosauierfunde in der Oberkreide Canadas.
Jahrg. Naturwiss. Berlin, vy. 12, no. 36, p. 709-716, 12 fig.

Beecker, A., 1903. Vergleichende stilstik der Nasenregion bei den Saurien,
Vogeln und Siugethieren. Morphol. Jahrb. vy. 31, p. 565-619.

Bellairs, A., and J. D. Boyd, 1950. The lachrymal apparatus in lizards
and snakes. II. The anterior part of the lachrymal duct and its rela-
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Proc. Zool. Soc. London, y. 117, p. 81-108.

Christensen, Kermit, 1927. The morphology of the brain of Sphenodon.
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Colbert, Edwin H., 1945. The dinosaur book Amer. Mus. Nat. Hist., Man
and Nature Publ. no. 14, 156 p., illus.

, 1955 Evolution of the Vertebrates. New York, John Wiley and
Sons, 479 p., 122 fig.

Gilmore, Charles W., 1924. On the genus Stephanosaurus, with a descrip-
tion of Lambeosaurus lambei, Parks. Geol. Sury. Bull., Canada Dept.
Mines, no. 38, geol. ser. 43, p. 29-48, fig. 8-12, pl. 6-10.

Haas, Georg, 1937. The structure of the nasal cavity of the Chameleon.
Jour. Morph., vy. 61, p. 433.

Kappers, C. U. Ariens, G. Carl Huber, and Elizabeth Crosby, 1936. The
comparative anatomy of the nervous system of vertebrates, including
man. London, Macmillan and Co., 2 vols., 1845 p., illus.

Kriusel, Richard, 1922. Die Nahrung von Trachodon. Paleont. Zeitschr. v.
4, p. 80.

Lambe, Lawrence, 1914. On a new genus and species of carnivorous dino-
saur from the Belly River formation of Alberta, with a description
of Stephanosaurus marginatus trom the same horizon. Ottawa Nat.,
v. 28, p. 17-20, 1 pl.

, 1920. The hadrosaur Edmontosaurus from the upper Cretaceous
of Alberta. Geol. Sury. Mem., Canada Dept. Mines, no. 120, geol. ser.
102, p. 1-79, 39 fig.

Lull, Richard S., and Nelda E. Wright, 1942. Hadrosaurian dinosaurs of
North America. Spec. Paper, Geol. Soc. Amer., no. 40, 242 p-» 90 fig.
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Nopesa, Franz, 1929. Sexual differences in ornithopodous dinosaurs. Paleo-
biologica, v. 2, p. 187-201, 3 fig.

Oelrich, Thomas M., 1956. The anatomy of the head of Ctenosaura pecti-
nata (Iguanidae). Mise. Publ. Univ. Michigan Mus. Zool., no. 94, 122
[Os fae) ake

Osawa, G., 1898. Beitrige zur Lehre der Sinnesorgane der Hatteria punc-
tata. Arch. Mikros. Anat., v. 52, p. 268.

Ostrom, John H., 196la. A new species of hadrosaurian dinosaur from the
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28 Postilla Yale Peabody Museum No. 62

———, 1961b. Cranial morphology of the hadrosaurian dinosaurs of
North America. Amer. Mus. Nat. Hist. Bull. v. 122, art. 2, p. 37-186,
78 fig., 6 pl.

———- , (In press.) Parasaurolophus cyrtocristatus, a crested hadro-
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Papez, James W., 1929. Comparative neurology. New York, Thomas Y.
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Parker, G. H., 1922. Smell, taste, and allied senses in the vertebrates.
Philadelphia, J. B. Lippincott Co., 192 p., 37 fig.

Parks, William, 1922. Parasaurolophus walkeri, a new genus and species
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———,, 1923. Corythosaurus intermedius, a new species of trachodont dino-
saur. Univ. Toronto Studies, geol. ser. 15, p. 1-57, 13 fig., 6 pl.

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58 fig.

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Chicago Press, 687 p. 337 fig.

, 1956. Osteology of the reptiles. Chicago, Univ. Chicago Press,
772 p., 248 fig.

Russell, Loris S., 1946. The crest of the dinosaur Parasaurolophus. Roy.
Ontario Mus., Paleont., Contrib., no. 11, p. 1-5, 2 fig.

Stebbins, Robert C., 1948. Adaptations in the nasal passages for sand bur-
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2 pl.

————, 1948. Nasal structure in lizards with reference to olfaction and
conditioning of the inspired air. Amer. Jour. Anat., vy. 83 p. 183-221,
9 fig.

Sternberg, Charles M., 1935. Hooded hadrosaurs of the Belly River series
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52, p. 1-37, 2 fig., 7 pl.

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Paleont., v. 16, p. 133-134, 1 fig.

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Trost, Edeltraud, 1956. Uber die Lage des Foramen Parietale bei rezenten
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Versluys, Jan, 1923. Der Schiidel des Skelettes von Trachodon annectens
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p. 1-19.

——, 1936. Kranium und Visceralskelett der Sauropsiden. Handb.
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Watkinson, Grace B., 1906. The cranial nerves of Varanus bivittatus.
Morph. Jahrb., v. 35; p. 450-572, pl. 11-13.

June 29, 1962 Hadrosaurian Dinosaurs 29
Wilfarth, Martin, 1938. Gab
Deutsche Geol. Gesell., v. 90, p. 87-100, 4 fig., 1 pl.
, 1939. Die Nasenbasis der Lambeosaurinae, Zentralbl. Min. Geol.
Palaeont., Abt. B, p. 24-39, 22 fig.
, 1940. Der

es risseltragende Dinosaurier? Zeitschr.

Atemriissel der Hadrosauriden. Halle, p. 1-24, 9 fig.

, 1947. Riisseltragende Dinosaurier. Orion (Munich) y. 2, p. 525-
532, 8 fig.

Wiman, Carl, 1931. Parasaurolophus tubicen, n. sp. aus der Kreide in New
Mexico. Nova Acta R. Soc. Sci. Upsaliensis, ser. 4, v. 7, no. 5, p.
1-11, 3 pl.

SQ
me

ete

YALE PEABODY MUSEUM

oF NATURAL History

Number 63 July 9, 1962 New Haven, Conn.

BRIEF COMMENTS ON THE THRUSHES

S. Ditton RIPLEY

In Postilla no. 13, 1952, pp. 1-48 et seq., 1952, 1954, I
published a fairly extensive review of this large and sprawl-
ing subfamily of the Old World Muscicapidae. More recently I
have been gathering data on this assemblage for Peters’ Check-
list, and so it appears worthwhile to list here a few of the
changes and revisions which have been made subsequent to my
earlier publication.

Certain large general works have been of the greatest use
such as Chapin’s (1958) section on thrushes in his monograph
on birds of the Congo, Vaurie’s (1959) monograph on Palearc-
tic birds and Mackworth-Praed and Grant’s (1955) volumes
on birds of eastern Africa. In addition shorter papers have been
of value such as that of White (1961) on African thrush
genera. I have been very grateful for comments and help re-
ceived from Messrs. Clancey, Deignan, Dilger, Charles Hart-
shorne, Ivanov, Johansen, Mmes. B. P. Hall and E. Koslova,
Messrs. Lees-Smith, Meinertzhagen, Terence Oatley, George
Watson, White, and Winterbottom.

After correspondence and discussion with these various
authors, as well as with the editors of the Chech-list, Messrs.
Greenway, Mayr, and Paynter, it appeared wise to retain the
Prunellidae as a monotypic family close to the thrushes. I

= Postilla Yale Peabody Museum No. 63

believe my preliminary revisional work of 1952 has served a
good purpose, however, in raising comments and criticism.
Without such preliminary work and the subsequent eddies and
fluctuations of opinion aroused, little long-range revisional
work would be possible.

Erythropygia, Cercotrichas, and Pinarornis.

Following the suggestion of White (1961), I have grouped
Cercotrichas and Pinarornis next to Erythropygia. I do not
agree to merging Cercotrichas with Erythropygia. This is a
rather distinct species and such field workers as Meinertzhagen
(1954) would keep them decidedly separate.

Pogonocichla and Erithacus.

Moreau’s interesting paper (1951) offers evidence that
Pogonocichla with two included species has characters which set
it sufficiently apart to be kept as a separate genus. Little
published work has convinced me that the genus Erithacus can-
not be kept as an expanded genus. Japanese workers, among
them Morioka (in litt.), point out the close resemblance of the
Japanese robin, akahige to the European rubecula, contra Lack
(1954). More recently Hall (1961) has suggested adding the
species gabela, described from Angola in 1957, to the genus
Erithacus.

Cossypha and Nenocopsychus.

The reasons advanced by White (1961) for transferring
Nenocopsychus to Cossypha seem provocative, and I ac-
cordingly have followed this suggestion, having already in-
cluded Dessonornis (Bessonornis auct.) earlier.

Pseudocossyphus.

I have removed the species sharpeit and imerina from
Cossypha and placed them in Pseudocossyphus following Good-
win (1956) whose suggestions and evidence I have found most

helpful.

July 9, 1962 Brief Comments cn the Thrushes 3

Rhyacornis and Chaimarrornis.

I also follow Goodwin (1957) in keeping Rhyacornis as a
separate genus for the small tropical redstarts with such dis-
tinctive alternate plumages, and placing Chaimarrornis close
to Monticola following Oenanthe.

Grandala and Sialia.

It is not obvious to anyone apparently where Grandala
should be listed. I leave it near Stalia for lack of a better linear
arrangement suggestion, not out of any conviction on the basis
of anatomy or phenotypic characters.

Phaeornis.

From personal observation in the field and listening to
songs I have decided that my earlier feeling that Phaeornis
was close to Myadestes following Stejneger (1887) was wrong.
Phaeornis has two species on Kauai Island which overlap ecolo-
gically. The larger, a subspecies of obscurus, has a broad bill
which resembles a solitaire. I believe it is a secondary adapta-
tion to coexistence in an overlapping niche, the separate first
invader being small and possessing a much smaller bill. In
powerful song and apparently in habits these birds seem far
closer to Catharus especially the central American nightingale-
thrush group.

Stizorhina.

Stizorhina may well prove to be a flycatcher. Unpublished
work in this laboratory (Ames ms.) shows un-turdine-like indi-
cations in the syrinx of 8. fraseri although the evidence is by ne
means compelling, as Myadestes also has a very similar syrinx.

Myrmecocichla, Pentholaea and T'hamnolaea.

I follow White (1961) in combining Pentholaea with
Myrmecocichla, although I keep Thamnolaea distinct, both on

4 Postilla Yale Peabody Museum No. 63

account of slight plumage differences as well as partially of
habits, as Cave and Macdonald (1955) emphasize for T'. coro-
nata. I have moved the Buff-streaked Chat, bifasciata, to
Oenanthe on further examination, convinced that in plumage
and behavior it fits better with the wheatears.

Catharus and Platycichla.

A few species changes have been made in the arrangement
of Catharus on the advice of various tropical American special-
ists such as Messrs. Blake, Paynter, Skutch, and Wetmore. I
have placed Platycichla as a separate genus on the advice of
several authors including Messrs. Bond and Phelps (personal
communication ).

Turdus.

For advice on the large genus T'urdus I have been much
indebted to Dr. Chapin and Mrs. Hall on African species and
specimens in the British Museum. This huge genus with more
than sixty-five species is a difficult one to arrange and I have
felt constrained to list the species regionally beginning with
Africa and ending with the New World. I have valued Dr.
Wetmore’s comments on ignobilis and plebejus which I have
separated as two species. I have also moved swalesi near to our
North American robin, feeling that this species is merely a
relict robin.

Several problematical genera have been removed from the
arrangements of thrushes of earlier authors just as others
have been added to it. Some of these former are T'urnagra, the
New Zealand “thrush,” obviously not a thrush at all, v¢de Oliver
(1955), who places the genus in a separate family based on the
presence of mouth bristles, union of maxillo-palatines, long
first primary and lack of spotted young. Namibornis (or
Bradornis) herero,the south West African flycatcher, Achaé-
tops, another South West African genus, inadvertently as-
signed to the thrushes by the South African Ornithological
Society List Committee (1958), and “Cercomela” buryt which

Or

July 9, 1962 Brief Comments on the Thrushes

may or may not be a Parisoma, but certainly with its wing
structure appears to bea warbler rather than a chat, are all
problematical species which I would include in the Musci-

capinae, Timaliinae and Sylvinae respectively.

LireraATuRE CITED

Cave, F. O. and J. D. Macdonald, 1955. Birds of the Sudan, Edinburgh,

ne

p. 275.

Chapin, J. P., 1953. Bull. Amer. Mus. Nat. Hist. 75A: 480-593.

Goodwin, D., 1956. Bull. Brit. Orn. Club, 76: 143-4.

—, 1957. Bull. Brit. Orn. Club, 77: 110-113.

Hall, B. P., 1961. Bull. Brit. Orn. Club, 81: 45-6.

Mackworth-Praed, C. W. and C. H. B. Grant, 1955. African Handbook
Birds, ser. 1, 2: 227-333.

Meinertzhagen, R., 1954. Birds of Arabia, pp. 220-2, 266-7.

Moreau, R. E., 1951. Ibis, 93: 383-401.

Oliver, W. R. B., 1955. New Zealand Birds (2 ed.) Wellington, p. 524.

Second Report of the S. A. O. S. List Committee, 1958. Ostrich, vol. 29, (1),
p. 37.

Stejneger, L., 1887. Proc. U. S. Nat. Mus. 10: 90.

Vaurie, C., 1959. Birds Palearctic Fauna. London, pp. 333-419.

White, C. M. N., 1961. Bull. Brit. Orn. Club, 81: 117-119, 150-152, 164-168.

50S) 74
P37

Luli

YALE PEABODY MUSEUM

oF Natura History

Number 64 August 7, 1962 New Haven, Conn.

TWO NEW PRIMATE SPECIES FROM
THE AFRICAN OLIGOCENE

Exiwyn L. Smvons

One of the major objectives of the Yale 1961-1962 Paleon-
tological Expedition to the Fayum region of Egypt was to re-
cover a larger sample than previously known of the earliest
mammalian microfauna from the continent of Africa, that of
the Fluviomarine formation of the Fayum early Oligocene.
During the course of our investigations the expedition staff
succeeded in locating specimens assignable to two new species
of Primates. In view of the considerable interest in, and rarity
of, Old World Primates dating from this epoch it seems advis-
able to publish a preliminary description of these two forms
without delay, so that they will be available for consideration
by other authors. It is intended that a fuller analysis of their
morphology and relationships will be included in a study of the
Fayum mammalian microfauna now being prepared by the
writer.

ACKNOWLEDGMENT

I wish to express my sincere thanks to Dr. Osman Moharam,
Director of the Department of Geological and Mineralogical!
Research, Ministry of Industry, United Arab Republic; Dr.

2 Postilla Yale Peabody Museum No. 64

Riad Higasy, former Minister of Industry, and Dr. Y. Shawki
Moustafa, Curator of Fossil Vertebrates, Cairo Museum of
Geology, for generously supplying facilities and assistance
which contributed greatly to the success of our expedition to
the Fayum badlands. The field research of which this report is
an outgrowth was made possible by a grant in Geology of the
National Science Foundation. Illustrations were prepared by
Margaret E. Freeman of New Haven and their execution was
partly financed by a grant from the Wenner-Gren Foundation
of New York. The photograph (figure 3) was prepared by
John Howard of the museum staff.

ABBREVIATIONS

ALM-UNGET. _..American Museum of Natural History,
New York.

NEC-PE- -Yale-Cairo, Paleontological Expedition
(field numbers).

VOR AVer Yale Peabody Museum, New Haven.

SYSTEMATICS
Class MamMatLia

Order PRIMATES

Suborder ANTHROPOIDEA

Infraorder CATARRHINI
OLIGOPITHECUS,' new genus

Type: Oligopithecus savagei, new species

Generic characters; Lower dental formula 2?. 1. 2. 3., size of mandi-
ble approximately that of the living ceboid primate Leontocebus rosalius,
slightly smaller than its contemporary Propliopithecus haeckeli. Differs
from the latter in having a slightly shallower mandibular ramus, more

‘Named with reference to the Oligocene occurrence of this catarrhine.

August 7, 1962 Two New Primate Species 3

anteroposteriorly elongated P,, distinct paraconid on P,-M,, lower molar
external cingula less distinct and hypoconulid shifted much more lingually
than in Propliopithecus and not distinctly separated from entoconid.
Differs from Parapithecus fraasi in its larger size and in possession of
undoubted, large canine anterior to P,, in absence of metaconid cusp on
P,,—present in Parapithecus, in possession of anteroposteriorly elongated
P., not seen in the latter genus, and in having a much more lingually ex-
tended paraconid ridge on M,-..

Figure 1. External view of left mandible, Oligopithecus savagei, Type,
Y.C.B.E. 207. x 3 approx.

Discussion: In depth of jaw and canine morphology Oligo-
pithecus much more nearly resembles Propliopithecus than it
does Parapithecus or Apidium, three other primates from the
Fayum early Oligocene. On the other hand, the anteroposteri-
orly elongate P., a feature which typically characterizes post-
Oligocene cercopithecoids and pongids, is not to be seen in any
specimens of these three Fayum genera. Such a lower P, does
occur in a damaged mandible, A.M.N.H. 138389 from the
Fayum, lacking tooth crowns other than on P,;, described by
Simons (1961), but in spite of this resemblance P., of A.M.N.H.
13389 is three rooted, a feature of great rarity among Pri-
mates, while that of Oligopithecus is more normal in being two
rooted. Moreover, A.M.N.H. 13389 is, in comparable measure-
ments, over twice larger than the type of Oligopithecus. Pre-
sumably the former specimen belongs to yet another Egyptian
Oligocene primate species, diagnosis of which will have to await
recovery of more satisfactory material; see Simons (1961: 3).

4 Postilla Yale Peabody Museum No. 64

The relationship of Oligopithecus to Moeripithecus mark-
grafi, also from the Egyptian early Oligocene, is more obscure,
since comparison of M,. crown patterns in the two forms
fails to reveal any significant similarity. The overall mor-
phology of the tooth series preserved in Oligopithecus appears
to be more like that of certain Eocene prosimians than are these
patterns in other Fayum Primates, except that the P,. are
lost, the horizontal ramus is deep compared to tooth crown
all similarities to

height, and P, is anteroposteriorly elongate
Old World Anthropoidea. Among Eocene Primates perhaps the
greatest resemblances of Oligopithecus are to Omomyidae,
Anaptomorphidae and Necrolemurinae, which fact agrees well
with the supposition that Anthropoidea are more closely allied
to the so-called tarsioids of Eocene times than they are to other
known Kocene, and earlier, families of the order. The lingual po-
sition of the M,. hypoconulid in Oligopithecus creates a par-
tial posterior loph parallel to the anterior loph, between meta-
comid and protoconid, which might represent an early stage mn
the transition to the bilophodont lower molar pattern of Cer-
copithecoidea. This possibility will require further confirma-
tion before definite assignment of this primate to the latter
superfamily would be advisable. Placement of Oligopithecus
among the Hominoidea is also conceivable but M,-» crown
patterns are so primitive that such an association is equally
dubious. Clearly, however, the creature is not a ceboid or a

prosimian.

OLIGOPITHECUS SAV AGEI-

Figures 1 and 4.

Type: Left mandibular ramus, with C-M. inclusive, Y.C.P.E. 207.3

Horizon and locality: Yale Expedition quarry E, Fossil Wood Zone,
Fluviomarine Formation, early Oligocene age, Fayum Province, Egypt,
about 2.3 miles northeast of American Museum Quarry A (1907) and about

*Named for Dr. D. E. Savage of Berkeley University who discovered the
type and only specimen.

*By agreement with the Egyptian authorities concerned, types collected on
the Yale 1961-62 expedition will eventually be deposited in the paleontologi-
cal collections of the Cairo Museum of Geology. Pending such assignment
they are here identified by their field numbers.

August 7, 1962 Two New Primate Species 5

eight miles west-northwest of Quasr el Sagha Temple, thirty feet below the
top of the Fossil Wcod Zone, see Beadnell (1905).

Specific characters: Not distinguished from generic.

Discussion: The species, O. savage, is at present known
only from the type individual so that little can be said of varia-
bility in this primate. Although M, is missing in Y.C.P.E. 207
the fact that it had fully erupted is indicated by the preserva-
tion of the anterior root of this tooth in the type. A large wear
facet on the anterior crest descending from P., protoconid indi-
rates that this species must have possessed a sizeable upper
canine which sheared against this facet.

Measurements in millimeters of Oligopithecus savagei are
listed below in comparison with similar measurements on con-
temporary Fayum Primates taken from Kalin (1961).

Mandibular dimensions (Q. savagei).

Anteroposterior

Length of 6 Ehmowebs Mire a2 Berea 19.4
Depth of mandible beneath M, . 95
Depth of mandible beneath P, ........ 10.2

MEASUREMENTS OF TEETH

- Oligopithecus ; Parapithecus Propliopithecus Moeripithecus
savaget fraasi haeckeli markgrafi
left side left side left side right side

Canine
fenoth eo: 3.7 3:2) (Cor Be) 4.0 (right )
madth 2 s4...- 3.3 2.5 (C or P.) 4.3 (right)
P,
Ven othites every ar 4.2 3.3 3.5
Wwadth) 255.4. 3.1 2.5 3.8
P,
length) 2.4. 3.3 3.3 3.5
WAGth «2.5.5.7 3.0 2.5 4.0
M,
lengths ae -- 4.2 4.2 4.8 5.0
Willy "oper 34 3.2 4.5 4.8
M,
length ..... 4.2 4.3 4.8 5A
Width: © = 2... 3.5 3.4 4.5 54

6 Postilla Yale Peabody Museum No. 64

Genus APIDIUM Osborn, 1908

Type: dpidium phiomense Osborn, 1908.

Discussion: Although Osborn (1908) was reluctant to state
the ordinal position of this species, it subsequenlty came to be
regarded as a primate, and Gregory (1922) held that it might
be related to the basic cercopithecoid stock or to Oreopithecus;
see Simons (1959: 14). More recently a few authors have again
questioned the primate status of A. phiomense but its resem-
blance to Oreopithecus seems more than convergent, as was
discussed by the writer (1960). Moreover, the discovery of
several isolated upper teeth of a species of this genus, (de-
scribed below) in association in one quarry with several lower
jaws of this form, shows us that similarities to Oreopithecus
in the upper dentition are about equal to those seen in the
lower teeth. Nevertheless, A pidiuwm species are rather unlike
other known Oligocene Anthropoidea in that the cheek-teeth
are more polycuspidate, the mandibular ramus is shallow, and
the canine may have been small, or alternatively there may
have been three premolars—inadequate preservation of the
new materials rendering this point equivocal. Clearly the stock
of Apidium had differentiated from that of other Fayum Pri-
mates at a considerable remove in time from their common ocur-
rence in the Early Oligocene of Egypt, but it is in many ways
closest to Parapithecus insofar as the two forms can be com-
pared. The question, therefore, as to the relation of species
of these two genera to the earliest undoubted Old World
Anthropoidea is interrelated. Provisionally the study of A pi-
dium suggests that it, together with Parapithecus, may even-
tually be ranked with some certainty among the Anthropoidea.

APIDIUM MOUSTAFAIT,' new species
Figures 2, 3.
Type: Left mandibular ramus with P,-M,, Y.C.P.E. No. 269.

Hypodigm: ‘Type and Y.P.M. numbers 18007, left mandibular rumus
with damaged P,-M,; 18008, unassociated right P'-M*; 18009 right mandi-
bular ramus with P.-M,; 18018 left mandibular ramus with C?-M,; 18042

‘Named in honor of Dr. Y. Shawki Moustafa, whose earnest assistance
and advice were most helpful to our Egyptian expedition.

August 7, 1962 Two New Primate Species (

left mandibular fragment with P,-M,; 18081, unassociated left P*?, left
M?’, left M., left M;, all from Quarry G.

Horizon and locality: Yale Expedition Quarry G, below upper fos-
siliferous zone, Fluviomarine Formation, early Oligocene age, Fayum Pro-
vince, Egypt. One mile due north of American Museum Quarry A and
approximately 100 feet above top of lower Fossil Wood Zone.

Specific diagnosis: About one-eighth smaller than 4. phiomense, with
slighter development of lateral basal cingula and fewer accessory cuspules
on P, heel and on M,—., and as a whole, comparatively smaller and more
primitive than in 4. phiomense, with posterointernal angle of M, produced

more lingually.

Figure 2. A. Crown view and B. External view, 4 pidium moustafai
Type, Y.C.P.E. 260. x 5 approx.

Discussion: This species is both smaller and earlier occur-
ring than the type of 4. phiomense. That a significant lapse of
time exists between the populations from which the two species
are derived seems probable in view of studies of other faunal
elements from the upper part of the Egyptian Fluviomarine
Formation now in progress. Although the exact locality of the
type of A. phiomense is not known, it was reported by Osborn
(1908) as having been discovered upon the upper fossiliferous

8 Postilla Yale Peabody Museum No. 64

level, which begins about 100 feet stratigraphically above the
horizon of Quarry G. If the practice of some past students of
fossil Primates were followed here, a case for generic separa-
tion of these two species could probably be made, but this type
of splitting should not be attempted until these forms are much
better known.

MerAsSUREMENTS IN MILLIMETERS OF Apidium moustafai

Y.PM. Y.P.M. TYPE:
18018 18009 Y.C.P.E. 260
= D eas or ize
fenrtht. coc Soe eee 2.5
OV RCEEEI 9-025 4 56-© cre oF 2.0
P;
lenpth® 05.2565 2s seec<< 2.7 2.7 2.8
WARE ots Se ke 2.0 1.9 2.0
P,
length oso ae oe 2.7 2.7 2.8
tls La ie ate Bee, eee 2.3 22 2.3
M,
Jenrth?:< 3 se ee 3.5 6 3.7
MIG. 2! 2 Oe ee ee 2.8 2.7 3.0
M.
fenptlt 252. (eo 3.6 3.6
REN, ieee es i wis 5.2 3.1
M,;
lenp ahs. 22S 3.6 4.0
Width >. 23.2 Sncnee 3.0 2.9
Depth of mandible beneath
Bb Ae eer ee 6.0 71 6.8
Anteroposterior length
Pl ee eee 15.3 16.8

A-P length
PAM oe ee ee 8.6 9.0 8.8

August 7, 1962 Two New Primate Species 9

CONCLUSIONS

Recent collections of Primates from the Fayum early Olig-
ocene deposits of Africa indicate the existence of at least two
new forms of Primates. Study of the first of these, Oligopithe-
cus savagei, suggests that it is assignable to the Anthropoidea
and that it may represent an early stage in the differentiation
of Cercopithecoidea. Near agreement in expected size, and ap-
proximation in horizon and locality suggests that a catarrhine
frontal bone described by Simons (1959) may provisionally be
referred to this species. Placement of this frontal with A pidiwm
or Parapithecus, which are also of suitable size, seems less ap-
propriate in view of non-anthropoidean features such as are to
be seen in their anterior dentitions. Known Propliopithecus and
Moeripithecus are probably too large to have had a frontal this
size, but of course this skull fragment could well belong to a
species otherwise unkown. A second species, Apidium mousta-
fai, here described, appears to represent an earlier and ances-
tral population to Apidium phiomense which was recovered
from the upper part of the Fluviomarine Formation. Newly
discovered upper teeth of Apidium strengthen the view that
species of this genus, together with Oreopithecus bambolii, rep-
resent a fourth major group of Old World Higher Primates

distinct from cercopithecoids, pongids and hominids.

10 Postilla Yale Peabody Museum No. 64

Figure 3. Right mandibular ramus of A pidium moustafai, Y.P.M.
18009. x 6 approx.

August 7, 1962 Two New Primate Species 11

Figure 4. Crown view of dentition, Oligopithecus savagei, Type,
Y.C.P.E. 207. x 6 approx.

gfe Postilla Yale Peabody Museum No. 64

REFERENCES

Beadnell, H. J. L., 1905. The topology and geology of the Fayum provence
of Egypt. Publ. Sury. Dept., Ministry Public Works, Egypt, p. 1-101,
10 fig., 24 pl.

Gregory, W. K., 1922. The origin and evolution of the human dentition.
Baltimore, Williams and Wilkins Co., p. 1-548.

Kihlin, J., 1961. Sur les primates de POligocéne inférieur d’Kgypte. Annales
de Paléontologie, tome 47, p. 1-48, 18 fig., 7 pl.
Osborn, H. F. 1908. New fossil mammals from the Fayum Oligocene,
Egypt. Bull. Amer. Mus. Nat. Hist. v. 24, art. 16, p. 265-272 6 fig.
Simons, E. L., 1959. An anthropoid frontal bone from the Fayum Oligocene
of Egypt: the oldest skull fragment of a higher primate. Amer. Mus.
Nov., no. 1976, p. 1-16, 4 fig.

—_———§—,, 1960. Apidium and Oreopithecus. Nature. v. 186, no. 4727, p.
824-826, 1 fig.

——————, 1961. An anthropoid mandible from the Oligocene Fayum
beds of Egypt. Amer. Mus. Nov. no. 2051, p. 1-5, 1 fig.

SOG. 73
P&7

LAB a

YALE PEABODY MUSEUM

oF NaTurAu History

Number 65 August 24, 1962 New Haven, Conn.

STUDIES ON SPECIATION IN
MALDANID POLYCHAETES OF THE
NORTH AMERICAN ATLANTIC COAST

I. A taxonomic revision of three species of the subfamily
Euclymeninae.

CHARLOTTE Preston Mancum*

The three species of the polychaete family Maldanidae to be
treated below are among the most common marine annelids of
the Atlantic coast. Despite their abundance and familiarity,
persistent confusion has led to an undue multiplication of taxa
and the shuffling of species between them. Since Leidy (1885)
described Clymene torquatus, the first American member of the
subfamily Euclymeninae, these three species have been referred
to at least nine genera and probably twice as many species.

It will be noted that their morphology is indeed similar.
However, it is possible to distinguish them with certainty even
in the field. The present effort is intended to provide a means
of proper identification as well as a taxonomic grouping indica-
tive of evolutionary status.

*Department of Biology, Yale University, and Duke University Marine
Laboratory.

2 Postilla Yale Peabody Museum No. 65

METHODS AND LOCALITIES

The specimens considered were in all cases collected by the
author. These collections extend over the entire Atlantic range
of the various species. A preliminary effort was made to com-
pare them with holotypes and paratypes in the collections of
the Peabody Museum of Natural History, Yale University:
the American Museum of Natural History, New York; and the
U. S. National Museum, Washington, D. C. It soon became
apparent, however, that the conditions of the type specimens
would not permit analysis with the precision desired. As an
alternative, the author has reinvestigated many of the type
localities.

Collections were made at:

Passamaquoddy Bay, New Brunswick (45°N), intertidal to
about 1.5 meters below Mean Low Water.

Cape Cod Bay, Mass. (41°30'N ), intertidal.

Vineyard Sound, Mass. (41°30'N), intertidal to about 1
meter below M.L.W. Type locality for Clymenella torquata
(Verrill, 1873) and Clymene producta (Lewis, 1897).

Long Island Sound, Conn. (41 °15’N), 0 to about 1.5 meters
below M.L.W.

Isle of Wight Bay, Maryland (38°20’'N), 1.5 meters below
M.L.W.

Newport River, North Carolina (34°40'N), intertidal to
3 meters below M.L.W.

Summer River, Florida (29°50’N), intertidal to about 1
meter below M.L.W.

Bahia Parguera, Puerto Rico (18°N), 0.5 to about 1.5
meters below M.L.W.

Specimens were killed in 5% formalin in seawater, and later
transferred to 70% ethyl alcohol. Setae were dissected out
and mounted in Permount.

Specimens listed under the new combinations have been de-
posited in the Yale Peabody Museum of Natural History.

Aug. 24, 1962 Maldanid Polychaetes 3

Genus CLYMENELLA Verrill 1873

Clymenella torquata (Leidy, 1855)

Taxonomic summary:
Clymene torquatus Leidy, 1855. New Jersey.
Clymenella torquata (Leidy), Verrill, 1873.
Paraxiothea latens Webster, 1879.

The genus Clymenella was erected by Verrill (1873) for Clymene
5 Yi A y
torquatus Leidy, thus removing this species from the European
genus Clymene Savigny.

Description: Fully formed adult with 22 segments: single pros-
tomial achaetous segment, 18 setigerous and 3 preanal achaetous
segments. Cephalic plate present dorsally on prostomium, slanting
away from more ventral proboscis and forming acute angle in
lateral view of approximately 65° (Fig. 1A). Slightly elevated bor-
der of cephalic plate with four indentations: two dorsal and two
ventrolateral. Central depression of plate bisected by median keel.
Keel bounded on either side by deep furrow (nuchal organ), and
extending anteriorly beyond border as small papilla. No dark pig-
ment spots between prostomium and _ papilla.

First three setigerous segments with notopodial fascicles of long
slender setae and 6-7 neuropodial rostrate uncini (Fig. 2A). Fourth
setigerous segment extends for short distance over third as dis-
tinctly flanged collar. Four pairs of nephridia lying in coelom of
setigerous segments VI through X (Paterson and Krewson, 1960)
often visible as whitish ventrolateral masses. Mid-region translu-
cent. Setigerous segments IX through XVIII greatly elongate, with
increasing tendency for formation of highly vascularized ridge at
posterior end of each segment. Number of neuropodial rostrate
uncini increasing to about 30-40. Three preanal achaetous segments
reduced in length, giving appearance of longitudinal compression.

Conical papilla perforated by anal opening and surrounded by
caudal funnel with cirrated posterior margin. Relative lengths of
cirri size-dependent; smaller animals or recently regenerated tails
with irregularly long and short cirri; larger animals or older tails
with equally long cirri. No single cirrus outstanding in relative
length. Number of cirri variable, 10-30.

Color highly variable. Basic body color slightly iridescent vellow-
orange, interrupted longitudinally by bright red dorsal and ventral

4. Postilla Yale Peabody Museum No. 65

blood vessels, and transversely in posterior segments by capillary
beds concentrated in each segmental ridge.

Basic body color tinged by dull brown in certain localities, or
obscured by accumulation of green pigment in others (Mangum,
1962). Overall appearance of entire population may be green or
orange, but microgeographic population consistent with regard to
color phase.

Length and length-frequency variable. Size range of adults over
entire geographic population approximately 1-16 em, but range of

seteee

Imm

C

Figure 1. A. Prostomium of Clymenella torquata.
B. Prostomium of Clymenella mucosa.

C. Prostomium of Clymenella zonalis.

Aug. 24, 1962 Maldanid Polychaetes 5

local population much smaller. Mean length of Newport River,
N.C., summer population 4.2 em (+0.9 S.D.); Nantucket Harbor,
Mass., summer population 11.4 cm (+2.3 S.D.).

Straight, vertical tubes of sand and mucus, approximately 20 cm
in length. Material cohesive; tubes removable intact from sediment
No gelatinous egg masses.

Geographic range on North American Atlantic coast from New
Brunswick to northern Florida; recent immigrant to southwestern
England (Newell, 1949). Sand or mud, intertidal to about 110 m.

A distinct subspecies from the Gulf of Mexico has been described
by Hartman (1951) as C. torquata calida. Its deviation from the
Atlantic stem species may be somewhat exaggerated, for she be-
lieved C. torquata to possess 1) a cephalic plate with an entire,
uncrenulated border, and 2) alternate long and short caudal cirvyi.
However, the number (8-9) of crenulations or indentations that she
attributes to Gulf populations is twice that (4) consistently found
in Atlantic populations. If constant, this morphological difference
may indicate genetic divergence, and therefore a valid subspecific
distinction.

Material deposited at the Yale Peabody Museum: YPM No.
1482. W. Pivers Island, Newport River, N.C.

Clymenella mucosa (Andrews, 1891)

Taxonomic summary:
Aviothea mucosa Andrews, 1891. North Carolina.
Clymenella (Aviothella) mucosa (Andrews), Verrill, 1900

Axviothella mucosa (Andrews), Arwidsson, 1907.

The distinction between the original material and C. tor-
quata (Leidy) was made on the absence of a well-developed
collar on the fourth setigerous segment of the former (An-
drews, 1891). The species was placed in the genus Awiothea
Malmgren, which was later found to have been in prior use
for a genus of Coleoptera (Verrill, 1900). On re-examination
of the type species of Malmgren’s genus, A. catenata, Verrill
(1900) found that a fleshy fold was present on the third and
fourth setigerous segments, but that it was much less devel-
oped than C. torquata. He concluded (1900: 657) : “The collar

6 Postilla Yale Peabody Museum No. 65

is doubtless much narrower in life than the latter (C. tor-
quata), but it is of the same nature.”

He thus extended his genus Clymenella to include Awiothea
Malmgren which he renamed <Awiothella and regarded as a
subgenus. The occasional presence of a collar on the third
and fifth segments is also diagnostic. It was noted here that the
collar is not peculiar to Clymenella but is scattered throughout
the family.

The generic separation of Aviothella and Clymenella Vervill
was made by Arwidsson (1907), who simply reversed Verrill’s
decision that the two kinds of collars were worthy of only
subgeneric distinction. Although his classification has persisted
since that time, the extensive morphological similarities of the
two genera have caught the attention of a number of workers
(Hartman, 1945; Bookhout and Horn, 1949; Moment, 1951).
At least one (Moment, 1951) has questioned the validity of
separating the two as genera.

The present author now proposes a return to Verrill’s
(1900) original classification, on the premise that the differ-
ences that do exist do not warrant separation of C. torquata
and C. mucosa on the generic level.

Description: Mature adult with 22 segments: relative lengths of
segments and distribution of setae identical with C. torquata.

Flanged collar of C. torquata on fourth setigerous segment rep-
resented here by fleshy rim, only slightly more developed than
comparable ridges on contiguous segments.

Cephalic plate slanting more abruptly than that of C. torquata;
angle of prostomium in lateral view approximately 45° (Fig. 1B).
Border of cephalic plate with only two indentations, dorsal and
ventral; other features of plate identical with C. torquata. Ventral
area between plate and proboscis bulb speckled with dark pigment
spots.

Gonads visible through body wall of setigerous segments IX
through XIV as opaque whitish masses: present during summer
months. Nephridia obscured.

Caudal funnel cirrated, the longer processes alternating irreg-
ularly with the shorter. Median ventral cirrus always longer than
others (up to 1.5 mm in very large specimens).

Color variable. Basic body color yellow to white. Red pigment
diffusely localized in setigerous segments IV to IX, giving pink to

Aug. 24, 1962 Maldanid Polychaetes i

pale red appearance. Red coloration obscured in certain localities
by accumulation of green pigment in these segments and in seti-
gerous segments X to XIV as well (Mangum, 1962). Walls of
major blood vessels and parapodial capillaries also green in these
populations, resulting in transverse green crescents along posterior
surfaces of neuropodia in setigerous segments X to XVITI.

Length variable. Range approximately 1.5 to 7.5 em. Newport
River, N. C., population 5.4 em (+1.5 S. D.); Bahia Parguera, P.
R., population 2.7 em (+0.7 S. D.).

Tubes sandy, vertical, not sufficiently cohesive to permit removal
intact from sediment. ‘Tubes of females Y-shaped, with round gela-
tinous egg masses protruding from one arm during summer months.

Geographic range from North Carolina to West Indies and Gulf
of Mexico. Sand, intertidal and subtidal.

Material deposited at the Yale Peabody Museum: YPM No.
1483. W. Pivers Island, Newport River, N.C.

Clymenella zonalis (Verrill, 1874) new comb.

Taxonomic summary:
Pravilla zonalis Verrill, 1874. New England.
Pravilla elongata Webster, 1879. New Jersey.
Clymene producta Lewis, 1897. Massachusetts.
Euclymene (Euclymene) zonalis (Verrill), Verrill, 1900.

Euclymene (Macroclymene) producta (Lewis), Verrill 1900.
Mass.

Possibly ?Macroclymene elongata (Webster), Hartman, 1951.
Louisiana.

This species is perhaps the most problematic of the three.
In addition to the synonymies noted above, it has been identi-
fied by innumerable names in various ecological surveys of the
Atlantic coast. It is known as Euclymene collaris in the check-
list of the Marine Biological Laboratory, Woods Hole, Mass.
It was not included by Hartman (1945) in her survey of the
area surrounding Beaufort, N.C., where it is common on local
sandflats. Assignment to E. collaris Claparéde is incorrect,

8 Postilla Yale Peabody Museum No. 65

since it is a Mediterranean species with differing morphology
(Claparede, 1870).

In his original description, Verrill (1874) named the species
Prawilla zonalis. The genus Pravilla was placed in synonymy
with Clymene Savigny, subsequently found to have been pre-
occupied (Verrill, 1900). Verrill (1900) offered Huclymene
as a new name, and extended the generic description to include
Prawvilla, Although the description of the expanded genus ap-
pears in the same work as that of Clymenella, Verrill does not
emphasize the basis of generic distinction. Comparison of the
descriptive texts reveals that the character of the neuropodial
setae of the first three setigerous segments is the sole diagnostic
feature (1900: 654 and 658) :

Buclymene: “. .. especially by having on about three anterior
setigerous segments, one or two stout, bent spines, replacing the
rostrate uncini of the ventral parapodia (of Clymenella).”

ve

. rows of ventral rostrate uncinate anterior setae
having a series of apical hooks and a beard, on all the anterior
setigerous segment.”

Clymenella;

The present author is aware that categorical distinctions
in polychaete taxonomy are frequently made on the basis of
setal structure. However, it is felt that the differences in this
case are so slight that they do not override the importance
of other morphological similarities. The same difference, 7.e.,
rostrate uncini vs. a single bent spine, is used to separate
groups of species within the genus Clymenella (see discussion
by Munro, 1937). Hartman (1961) has recently recognized
?Clymenella cincta St. Joseph, 1899, as a valid species, although
it bears the anterior spines characteristic of Euclymene Verrill.

It may very well have been his pre-occupation with segment
numbers that led Verrill to seek generic differences, although
this is only implicit in the text. He proceeds to discuss **a very
aberrant species from near Vineyard Sound, Mass.,’’ described
by Lewis (1897) as Clymene producta. After examination of
the type he concludes that it differs from E. zonalis only in
the segment number. But the increase from 25 segments in his
E. zonalis type to ca. 70 segments in Lewis’ type was sufficient,
in Verrill’s judgment, to erect the sub-genus Macroclymene
with FE. (M.) producta (Lewis) as the type.

Aug. 24, 1962 Maldanid Polychaetes 8)

Hartman imphed the elevation of Macroclymene Verrill to
generic standing in her tentative identification of a Gulf of
Mexico fragment (Hartman, 1951) as ?Macroclymene elon-
gata (Webster), which had been previously synonymized by
Verrill (1900) with his E. zonalis. Macroclymene Verrill was
subsequently listed as a separate genus in her authoritative
Catalogue of the Polychaetous Annelids of the World (Hart-
man, 1959).

From the present author’s collections it is apparent that
the segment number of an individual and its range in a micro-
geographic population are simply size-dependent. The animals
continue to proliferate additional segments so long as growth
occurs, unlike the species whose adult growth consists only
of the enlargement of a definite number of segments. The
entire range described in the literature may be found within a
single microgeographic population, e.g., Vineyard Sound, Mass.
It hardly seems likely that intrapopulational reproductive iso-
lation exists concomitant with continuous differences in seg-
ment number. Therefore all of the Atlantic types should be
regarded as members of a single species.

The author was unable to find specimens on the Atlantic
coast south of North Carolina, and has not collected on the
Gulf coast. Judgment must again be withheld on the specimen
identified by Hartman (1951) as ?M. elongata (Webster).

Description: Segment number variable. Range: 18 to ca. 70.
Prostomial segment achaetous, followed by 15-ca. 65 setigerous and
3-4 preanal achaetous segments. Cephalic plate slanting dorsally
forming prostomial angle of approximately 65° (Fig. 1C). Border
of cephalic plate with five indentations: two dorsal, two ventro-
lateral and one ventral. Plate otherwise identical with C. torquata.
Dark pigment spots present ventrally, but fewer than C. mucosa.

First three setigerous segments with dorsolateral fascicles of long
slender setae, and 1-3 ventrolateral bent spines (Fig. 2B). 10-20
rostrate uncini beginning on setigerous segment IV and continuing
posteriorly.

Condition of collar identical with C. mucosa; fleshy rim on seti-
gerous segments III through V. Beginning with X and continuing
up to preanal achaetous segments, posterior segments greatly
elongated and swollen at terminals. Paired nephridia visible through
body walls of setigerous segments VII through XIV as elongate

10 Postilla Yale Peabody Museum No. 65

ventrolateral whitish masses. Preanal achaetous segments com-
pressed longitudinally.

Anal opening at center of conical papilla surrounded by cirrated
‘audal funnel. Number and relative length of cirri variable, resembl-
ing C. mucosa. Median ventral cirrus always longer than rest (up to
0.5 mm total length), but not so long as that of C. mucosa.

Color constant. No green populations found as yet. Basic body
color slightly iridescent vellow-orange, resembling C.
Distinct red
through IX,

crescents of

torquata.
bands surround portions of setigerous segments IV
and sometimes X in larger individuals. Transverse
red (lateral blood vessels) on posterior surfaces of

TABLE 1. COMPARISON OF CHARACTERS

Clymenella

Clymenella

Clymenella

Character torquata mucosa zonalis
Segment 22 22 18-70
number constant constant variable
Angle of 65° 45° 65°
prostomium constant constant constant
No. indentations 4, 8-9 2 5

in cephalic plate variable constant constant
border

Prostomial pigment absent present present
spots

Flanged collar on present absent absent
fourth setigerous

segment

Neuropodial uncini rostrate rostrate bent spine
on setigerous constant constant constant

segments I-IIT

Relative lengths
of caudal cirri

Basic body

all variable

yellow-orange

median ventral
longer
constant

yellow-white

median ventral
longer
constant

yellow-orange

color constant variable constant

Red bands in absent pale, diffuse bright,

mid-region constant distinct
constant

Green present but present but absent

dichromatism variable variable

Gelatinous absent present absent

egg mass

XY > SOpy

Figure 2. A. Rostrate uncini from anterior setigerous
segment of Clymenella torquata.

B. Neuropodial spines from anterior setigerous
segment of Clymenella zonalis.

Aug. 24, 1962 Maldanid Polychaetes iil

neuropodia from setigerous segment X to last setigerous segment.

Length variable; range 1.5 to ca. 20 cm.

Tubes sandy, sufficiently cohesive to permit removal from sedi-
ment intact. Straight, vertical for approximately 15-20 cm, then
frequently curved, becoming J-shaped. No gelatinous egg masses.

Geographic range from Maine to North Carolina. Intertidal to
50 m; in sand.

Material deposited at the Yale Peabody Museum: YPM No. 1484.
W. Pivers Island, Newport River, N.C.

ACKNOWLEDGMENTS

I am grateful for the sponsorship of Dr. L. M. Passano
throughout the course of this investigation. I would also like
to thank Dr. W. D. Hartman and Professor G. E. Hutchinson
for their criticisms of the manuscript. Dr. Meredith Jones has
been a most stimulating critic and helpful consultant, although
responsibility for the conclusions reached above is mine. The
facilities provided by the following marine laboratories enabled
me to cover the geographic area: Atlantic Biological Station,
St. Andrews, N. B.; Mt. Desert Island Biological Laboratory,
Salsbury Cove, Me.; Museum of Natural History, Nantucket,
Mass.; Marine Biological Laboratory, Woods Hole, Mass. :
Dept. of Research and Education, State of Maryland, Snow
Hill, Md.; Duke University Marine Laboratory, Beaufort,
N.C.; Bears Bluffs Laboratory, Wadmalaw Island, S.C.;
Marineland Research Laboratories, Marineland, Fla.: Insti-
tute of Marine Biology, University of Puerto Rico, La Par-
guera, P. R.

Part of this investigation was carried out while I was a
National Science Foundation Graduate Fellow, and part was
supported by a research grant from the National Science

Foundation (NSF G17862).

LireraTurRE Crrep

Andrews, E. A., 1891. Report on the Annelida Polychaeta of Beaufort,
North Carolina. Proc. U.S. Nat. Mus. 14:277-302.

Arwidsson, Ivar., 1907. Studien iiber die skandinavischen und arktischen
Maldaniden nebst Zusammenstellung der iibrigen bisher bekannten
Arten dieser Familie. Zool. Jahrb. Suppl. 9: 1-308.

Bookhout, C. G. and E. C. Horn, 1949. The development of Aviothella
mucosa (Andrews). J. Morph. 84: 145-183.

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Claparéde, Edouard, 1870. Les Annelides Chaetopodes du Golfe de Naples.
Soc. Phys. Geneve Mem. 20: 1-542.

Hartman, Olga, 1945. The marine Annelids of North Carolina. Duke Univ.
Marine Station Bull. No. 2. 51 p.

—————,, 1951. The littoral marine Annelids of the Gulf of Mexico. Publ.
Inst. Mar. Sci. 2: 7-124.

1959. Catalogue of the Polychaetous Annelids of the world.
Hancock Found. Publ. Occasional Paper No. 23. 628 p.

, 1961. Polychaetous Annelids from California. Hancock Pacific
Expedition Vol. 25. 226 p.

Leidy, Joseph, 1855. Contributions towards a knowledge of the mar-ne
invertebrates of the coasts of Rhode Island and New Jersey. Jour.
Acad. Nat. Sci. Philadelphia 3: 135-158.

Lewis, Margaret, 1897. Clymene producta sp. nov. Proc. Boston Soc. Nat.
Hist. 28: 111-115.

Mangum, C. P., 1962. The source of dichromatism in two Maldanid poly-
chaetes. Nature (in press).

Moment, G. B., 1951. Simultaneous anterior and posterior regeneration and
other growth phenomena in Maldanid polychaetes. J. Exp. Zool. 117:
1-14.

Munro, C. C. A. 1937. Polychaeta. John Murray Expedition. 1933-34. Sci-
entific Reports 4(8): 308-310.

Newell, G. E. 1949. Occurrence of a species of Clymenella Verrill (Poly-
chaeta fam. Maldanidae) on the North Kent coast. Nature 163: 648.

Paterson. M. C. and C. R. Krewson. 1960. Histological investigation of the
nephridia of Clymenella torquata, Biol. Bull. 119: 331-332.

Verrill, A. E. 1873. Report on the invertebrate animals of Vineyard Sound
and the adjacent waters, with an account of the physical characters of
the region. U. S. Comm. Fish. Rep. for 1871-1872:295-778.

. 1874. Explorations of Casco Bay by the U.S. Fish Commission,
in 1873. Am. Assoc. Adv. Sci. Proc. 22: 340-395.

. 1900. Additions to the Turbellaria, Nemertina and Annelida of
the Bermudas, with revisions of some New England genera and species.
Trans. Conn. Acad. Arts & Sci. 10: 595-671.

Webster, H. E. 1879. The Annelida Chaetopoda of New Jersey. N.Y.
State Mus. Nat. Hist. Ann. Rep. 32: 101-128.

DOS. 7S
2 $7

etitla

YALE PEABODY MUSEUM

oF Natura History

Number 66 September 10, 1962 New Haven, Conn.

SNAILS ON A PERSIAN HILLSIDE

Ecology—Prehistory—Gast ronomy

Cuarwes A. REED

The archeologist who is prehistorian may expect snail shells
in his excavations. Sometimes, even though the site may be far
from the nearest seas, such shells are marine and often were
used as decorations, indicating the continuity of human vanity
through the ages. Generally, the archeologist has had little
interest in such shells as snails, or even in the generic and
specific identifications furnished him by a malacologist; in-
stead, the archeologist is interested in any cultural uses of the
shells, and is intrigued by problems of their geographic origin
and the possibility of tracing prehistoric trade routes.

However, to the ecologically-oriented archeologists and the
various natural scientists with them working the past fifteen
years in Iraq and Iran, some of the local terrestrial snails
have become of prime interest, particularly as potential indi-
cators of past environmental conditions (including climate),
and as a source of food for past populations. Thus the con-
tinued presence in archeological sites of the same species of
snails in the same localities in northern Iraq, for periods some-
times measured in the tens of thousands of years, has been used,
with other evidence, to make a tentative reconstruction of the

2 Postilla Yale Peabcdy Museum No. 66

environments of that area over those periods (Reed and Braid-
wood, 1960).

This environmental reconstruction, based primarily on zo0o-
logical data, is partly in conflict for some time-periods with the
glaciological evidence of Wright (1961) and the palynological
evidence of Solecki and Leroi-Gourhan (1961). Wright’s stu-
dies indicate a colder climate for the period of the Wiirm max-
imum of the late Pleistocene than postulated by Reed and
Braidwood for those parts of this period for which they had
zoological remains, mostly of mammals and snails. Conclusions
derived from study of the pollen-grains recovered from Shani-
dar Cave in northern Iraq do not contradict Wright’s glacio-
logical evidence, but do indicate more fluctuating climatic phases
during the periods of the Baradostian and Mousterian cultures
than postulated by Reed and Braidwood.

Obviously if zoological remains, such as snail shells, are to be
useful in assessing past environments, the ecological conditions
of life

which each species can live—must be known. At present, such

and particularly the environmental limitations within

ecological data are not known precisely for any animal popula-
tion of southwestern Asia; while in general the botanical assess-
ment of past environments is probably capable of more exacti-
tude than is one based on zoological evidence, the latter should
not be ignored, and this present paper is a pveliminary effort
toward an understanding of the ecology of scme of the snails
excavated in various of the archeolegical sites in Iraq and
Iran.

To be useful as a climatic indicator, an animal population
should have narrow and definite environmental limitations, and
these should be known. For instance, a snail which ranges from
the Dead Sea to the Iranian Plateau, as does Helicella lang-
loisiana Bourguignat, is obviously useless as a climatic indi-
cator (Biggs, 1962).!

However, as our knowledge of the ecologic tolerances and
limitations of each animal and plant species increases, we can

|The same criticism might be brought against the use of //elix salomonica
as an environmental indicator, since it had been reported (Biggs, 1960)
trom Jericho in the lower Jordan valley. However, it is now believed that
this identification was an error (Biggs, personal communication).

Sept. 10,1962 Snails on a Persian Hillside 3

use the data derived from combinations of species, each with
varied requirements and with different present geographical
ranges, to clarify our concepts of the changing environments
of the past.

In all such attempts at environmental reconstruction we
begin with the assumption that a biologic population of a past
period, as represented by identified remains, had the same eco-
logical requirements as do members of a species with the same
morphology as studied today. In general, these assumptions of

TURKEY ‘ nia ON u
ios : w aS i} TURKEY 2 =
Co C ) a
SHANIDAR ¢, ie aes

Wis

=—-ae— International boundary
River

WARWAS|

SARAB
ee

CO: ASIAB
Kermanshah

Fig. 1. Map of the area discussed, showing the archeological sites mentioned.

4 Postilla Yale Peabody Museum No. 66

evolutionary stability and ecological uniformitarionism upon
which we build cur paleo-ecological reconstructions are prob-
ably more valid than is the loose framework of our present
knowledge, but we are warned by Johnson (1960), even if in a
different context, that these assumptions may be less valid for
the more precise details we hope to learn in the future.

In addition to the climatic problems outlined above, the
steady increase in numbers of one kind of snail, Helix
(Naegelea) salomonica Naegele through the late Pleistocene
and into the early Recent (post-Pleistocene) in all archeologi-
cal sites of northern Iraq, is considered to be evidence for the
increased use of these animals as food by a steadily-growing
population (Braidwood and Reed, 1957; Braidwood and Howe,
1960). The same general pattern is evident in western Iran,
and other species of Helix during this general period were sim-
ilarly being used in northern Africa.

Thus by the time the fourth” southwestern Asiatic prehis-
toric expedition of the Oriental Institute of the University of
Chicago was ready to go into the field in 1959, a preliminary
field study of the snails important to the archeologists had be-
come necessary.

Previous observations in northern Iraq and the problems
posed by them were as follows (see Braidwood and Howe, 1960,
as a general reference for geography, environment and chro-
nology):

1. Helix salomonica is not common in archeological sites
prior to the late cave-living period of the uppermost Pleistocene
(i.e., the Zarzian, ca. 15,000-12,000 years ago), then increases
in concentration to ca. 8,500 years ago, and was still numerous
at 7,000 years ago, but after that it disappeared almost entirely
from the archeological record of the area. It cannot be found
at present in some of the regions where it was previously so
plentiful, as for instance in the immediate area of Jarmo in
northeastern Iraq. The appearance and increase of these snails

2 The previous expeditions had been in 1947-1948, 1950-1951, and 1954-1955,
all in northeastern Iraq. The fourth, 1959-1960, went into western Iran. All
four expeditions have been under the direction of Dr. Robert J. Braidwood
of the Oriental Institute, University of Chicago. The present author was a
member of the two last expeditions.

Sept. 10,1962 Snails on a Persian Hillside 5

in archeolcgical sites are undoubtedly to be correlated solely
with changing human food-habits, but is their disappearance
from the archeological record due solely to changing human
food preferences? Possibly local destruction of the environment
(deforestation and brush-cutting, cultivation, over-grazing and
soil erosion) have destroyed the micro-habitat necessary for
the species so that its present distribution is discontinuous.
The species may thus be listed as “tabundant” over much of
northern Iraq at present (Harris, 1961), and have a number
of localities listed (Biggs, 1959) while yet being locally erad-
icated in an area (as at Jarmo) where once numerous. Harris
writes in general for the terrestrial gastropods which he has
listed as abundant, “As long as some perennial plants are
present, the supplement of annuals is quite sufficient to provide
food for the limited faunas common today. Where perennials
are absent, and only a poor annual flora is present, snails do
not occur, even though the rainfall is adequate.” However,
the situation is not so simple, as there may well be a sequence
of local eradication with the deterioration of the environment ;
for instance, as mentioned, H. salomonica now seems to be
absent entirely from the area around Jarmo, whereas Levan-
tina hurdistana (1..Pfr.) still flourishes there.

2. Different species of Levantina have been recorded from
different archeological sites in northern Iraq. L. mahanica Ko-
belt is found, in at least one long-occupied site, that of Shani-
dar Cave, for several tens of thousands of years, where the
shells of this species occurred earlier (ca. 60,000 or more years
ago) than did those of H. salomonica (somewhat more than
50,000 years ago) and in greater numbers than the latter in
the older deposits (prior to 27,000 B.C.). However, shells of
H. salomonica are more numerous in the later, post-Wiirm de-
posits (Solecki’s layers B and A) which coincide with the pe-
riod when this latter snail was being gathered for food in other
areas of southwestern Asia. In general, the concentrations of
the shells of Levantina never become high in archeological re-
mains in Iraq and Iran, and one assumes that it was rarely, if
ever, used for food. One wonders, too, why it was not so used,
since it is a large and meaty snail. However, the number of

6 Postilla Yale Peabody Museum No. 66

L. spiriplana (Olivier) found together at Jericho, in the lower
Jordan valley, suggested to Biggs (1960) that this species of
Levantina was being eaten at one time at that site.

Levantina has managed to survive, as at Jarmo, where Helix
salomonica cannot now be found. Moreover, at the three sites
(Shanidar Cave, Jarmo, and Warwasi), where recent collec-
tions have been made, the species of Levantina reported from
the prehistoric levels are the same (L. mahanica, L. kurdistana,
and L. diulfensis [| Mousson |, respectively) as are those found
in each of the areas now.

3. At the strictly taxonomic level, are the different species
of Levantina reported from northeastern Iraq (mahanica, kur-
distana, guttata | Olivier |) valid species, or are they—and the
forms since collected from western Iran (guttata and/or diul-
fensis)—merely geographical variants (1.e., subspecies) in a
wide-ranging and continuous population (one species), the
different parts of which exhibit considerable morphological
differentiation ?

It was with such questions in mind that the members of the
Iranian Prehistoric Project went into the field in west-central
Iran in 1959. We cannot claim to have solved any of these
problems, but we have added to our observations, and we feel
that a presentation is due of these, as well as of our present
level of understanding.

Unfortunately, late 1959 and early 1960 was a poor time for
“normal” environmental observations. The season was an ex-
ceptionally dry one over all of southwestern Asia. In the area
of Kermanshah, west-central Iran, where our group was _ lo-
cated, unusual cold and snow in November was followed by a
long period of winter dryness, during the greater part of what
is usually the rainy season. Then there was more snow in March
and finally some rain in April.

Observations to be reported were thus made in the area of
Kermanshah under the above-described circumstances. The
Kermanshah valley is a flat-bottomed alluvial valley, at a
general level of 4,000-4,400 ft. (ca. 1,230-1,3830 m), surrounded
by mountains, some of them high, steep and rugged. The aver-

Sept. 10,1962 Snails on a Persian Hillside re

age annual precipitation is variously reported as 18.1 in.
(Robison and Dodd, 1955) to nearly 17 in. (Bakker, 1956;
Ganji, 1960: the latter’s figures are for 15 consecutive years. )
This precipitation occurs entirely during the typical Mediter-
ranean “rainy season” (October to May, with March the month
of heaviest rain). The “average” is, however, not the “normal,”
as wide annual fluctuations occur, from 9 in. to 24 in. (22.5-
60.0 cm), although such extremes may not occur oftener than
once in 85 years (Bakker, 1956). Even two seasons recorded
as having the same precipitation could differ widely in the eco-
logical results of that snow and rain, depending upon the tem-
perature at the time, the intensity of the rain (or depth of
snow), and particularly upon the seasonal distribution.

The valley floor and much of the adjacent valley walls are
not now forested, and the botanists who have studied there
seem agreed that probably the areas now unforested have
mostly not been forested as long as present climatic conditions
have prevailed. Actually, the area seems to be one of an en-
vironmental transition, as measured by floral zones (Bobek,
1951; Pabot, 1961). The valley floor is now intensively cul-
tivated, and the valley walls heavily grazed and subjected to
continuous bush-cutting (I do not say brush-cutting because
the vegetation is too sparse to be called brush.) To what extent
the human activities, continuous for several thousand years,
have changed the original environment we cannot assess nor
can we at present definitely reconstruct the pre-agricultural
environment.

All of the above environmental factors, not available to
museum malacologists intent on taxonomic identifications, are
pertinent to the understanding of our problems. Actually, we
need to know additional types of ecological information not
yet gathered, such as the distribution, both geographically and
environmentally, of each species represented. Precise data of
this type would inform us concerning the limits within which
the total environment might vary and yet allow combinations
of certain species of snails to persist at one spot, as at War-
wasi. However, not only is such information not yet available,
but the nomenclatural confusions concerning certain of these
species are such that one finds difficulty in interpreting some of

8 Postilla Yale Peabody Museum No. 66

the identifications as published. It would be most desirable to
have the collecting, identifications, and ecological studies ac-
complished by one person or a coordinated team. Additionally,
we should know the fewest number of feeding periods per year
which will support each population of snail, and also the kinds
of variables (maximum and minimum daily temperatures and
maximum and minimum daily precipitations) which control the
emergence, feeding and breeding of each species. Eiseley
(1937) has considered in some detail other factors of the ecol-
ogy of terrestrial gastropods which have bearing on paleo-
environmental interpretations, and several other authors have
also considered different aspects of this general problem.

Some observations and collecting of empty shells had been
accomplished prior to the April rains, and by that time data
from several excavations could be added (Braidwood, Howe,
and Reed, 1961). Our observations were concentrated on Helix
and Levantina, for these alone (so far as we can see at present
for the area of our studies) are part of our more general pre-
historic archeological problems. These preliminary observa-
tions were as follows:

1. Shells of neither Helix nor Levantina were found any-
where on the open valley floor, on open rounded well-grazed
hills, in areas adjacent to streams, or in the typical oak-haw-
thorn-pistachio forest (Bobek, 1951) of some of the nearby
hilly areas.

2. Adjacent to the archeologic site of Warwasi (fig. 2) in
the T'ang-i-knisht valley, there is a southwesterly-facing, rock-
strewn slope covered with thornbushes and with a cliff above.
On the slope the most common shell was that of Helicella
langloisiana, occurring by the thousands. Some of these were
sharply-keeled, some almost rounded on the edges of the whorls,
with all gradations between these extremes. The second shell,
in frequency, was the high-spired Jaminia (Euchondrus) albula
(Mousson). Next most common was Levantina diulfensis; the
Levantina shells were accumulated at the base of the cliff and
some scattered down the slope. Fourth in frequency were shells

Sept. 10,1962 Snails on a Persian Hillside 9

of Helix salomonica, found only on the slope. Rarest were
shells of Buliminus (Buliminus) egregius Naegele and Zebrina
carducha (Mertens), which were found only in cracks in the
cliff and, presumably fallen from there, at the foot of the cliff.

3. Shells of Helix salomonica, but of no other snail, were
found in open fields on the top of the low divide between the
Kermanshah valley and the next valley to the south. The alti-
tude was around 5,500 feet (ca. 1,700 m) and the shells were
associated here only with a hardy ground-hugging perennial
too soft to be called a shrub but yet too resistant to be removed
by the primitive ploughs used in the area. This circumstance
agrees with the observations in Iraq of Harris (1961) on the
role of perennials in the survival of snails, but one wonders
what factor in this particular locality led to the survival of
H. salomonica and no other species.

4. Shells of Helix salomonica were found in great numbers
in an archeological site (Tepe Sarab) located in the open Ker-

Fig. 2. The hillside and cliff at Warwasi, where living snails were col-
lected in April of 1960. The Paleolithic archeological site of Warwasi is the
dark overhang at the base of the cliff, almost directly above the ear.

10 Postilla Yale Peabody Museum No. 66

manshah valley (Braidwood, Howe, and Reed, 1961); Tepe
Sarab is now dated at nearly 8,000 years ago and is thus prob-
ably somewhat more recent than is Jarmo. Large numbers of
H. salomonica in a site of this period agree with our findings
in northern Iraq. We are not suggesting that H. salomonica
was the major food source at Tepe Sarab (nor at any other
archeological site), as: a) in our experience, these snails can
only be gathered during or following a rain, and; b) at Sarab,

orains and

as at Jarmo, the people already had cultivated g

domestic animals.

5. In a site (Tepe Asiab), probably some 2,000 years older
than ‘Tepe Sarab and less than a mile distant, but near a per-
manent stream (the Kara Su) in the valley’s center, H.
salomonica is extremely rare; the major molluscan protein
source here seemingly was a clam, Unio tigridis Bgt. (So far as
we know, these earlier people at Tepe Asiab did not have cul-
tivated grains or domestic animals.) In northern Iraq at the
same time (ca. 10,000 years ago) we think H. salomonica was
being eaten in some quantities; perhaps the easy availability of
the fresh-water clams, still present in the river adjacent to
Tepe Asiab, made the gathering of snails unnecessary.”

6. In the rock shelter of Warwasi (Braidwood, Howe, and
Reed, 1961), adjacent to the slope and cliff mentioned in para-
graph 2 above, snail shells are found sparsely but continuously
through most of the Zarzian and through all of the deeper
Baradostian and Mousterian levels (these latter at least 40,000
years old and probably older). Shells of Helix salomonica be-
come numerous in the uppermost Zarzian layers (about 12,000

3 It is obvious, thus, that local biotic and/or cultural patterns change the
local archeological findings, making widespread archeological exploration
necessary before a total picture emerges. In the excavations at Tepe Sarab,
for instance, Unio tigridis is rare, although the clam-laden Kara Su is no
more than a half-mile away. Within some 2000 years a major shift in food-
habits had occurred and the people simply didn’t gather clams anymore.
Similarly today in the same region, we were told that the people will not
eat clams or snails, even under conditions of extreme starvation. There is
no truth to the assertion sometimes made by some prehistorians that primi-
tive people ate anything and everything they could gather. ‘There is now,
and seemingly has always been, the important factor of the “cultural filter”
in the collection of human food-stuffs, and consequently in the comparative
archeologie record.

Sept. 10,1962 Snails on a Persian Hillside mn

years ago), at which time these snails probably were being
gathered for food. Throughout these tens of thousands of
years of the later Pleistocene the species represented are the
same as those still present on the hillside and, except for the H.
salomonica of the more recent Zarzian levels, are considered to
be no more than random strays into the cultural deposits. In
levels below the upper Zarzian, Levantina is the most numerous,
and occurs earlier than any of the others; at its earliest occur-
rence it was exactly the same L. diulfensis as found on the hill-
side today.

Subsequent collections of living snails, on April 9 and April
18, 1960, were made on the slope and cliffs immediately ad-
jacent to the site of Warwasi in the Tang-i-knisht valley, which
is a lateral side-valley opening southerly into the main Ker-
manshah valley close to the town of Kermanshah. The mouth of
the secondary valley is bounded by high limestone cliffs, with
steep slopes of soil and talus rock at their bottoms. It was on
such a slope (fig. 2), southwesterly facing, that we did our
collecting. Toward the base of the slope there was relatively
little fallen rock, but higher and closer to the cliff the tumbled
rock was thick. Thorn-bushes, rarely over 10 inches high and
spaced some 10 or 15 feet apart, dotted the hillside, even grow-
ing among the fallen rocks but not up on the cliff. The ground
between the thorn-bushes is quite bare since the slope is heavily
overgrazed. This particular slope could be duplicated thou-
sands of times around the Kermanshah valley ; while we can say
the slope is typical of the area today, we must also assume that
the degenerate floral assemblage represents only a remnant of
the “natural” vegetation (whatever that may have been) before
intensive human use had removed most of the less hardy plants.
Thus we cannot now imagine the appearance of these steep hill-
sides some 8,000 years ago.

On the two nights mentioned, there were continued gentle
‘ains, followed by cool cloudy mornings (11°C-14°C ground
temperature) with occasional drizzle. Living snails were asur-
face both mornings. Since the conditions and the collections
were generally similar, the descriptions of the two events will be
combined.

12 Postilla Yale Peabody Museum No. 66

SPECIES ACCOUNTS
(Identifications were made by Rev. H. E. J. Biggs, from liv-
ing specimens air-mailed to England; see Biggs, 1962.)

1. Helix salomonica: Most of these snails were found a-
bove ground, but under the thorn-bushes on the lower three-
fourths of the slope. The snails were extended and moving, not
up in a bush, but were generally on the accumulation of dead
leaves and grass under a bush or at its edge. Only two individ-

A B

Fig. 3. A. Helix salomonica. B. Levantina diulfensis. Natural size.

uals were seen out between bushes, on practically bare ground.
A few were found under rocks (although none could be found
under rocks between rainy periods). The population of Helix
thinned out up the slope, and none were found in the upper
fourth of the slope, although thorn-bushes occur on that upper
fourth of the slope. On April 18 (although not on April 9) the
snails were observed copulating; of 840 picked up on the slope
on the latter date, 12 pairs were coupled, and many more of

the living snails did so in the jars after being collected.

Sept. 10,1962 Snails on a Persian Hillside 13

2. Levantina diulfensis: These snails were first found about
halfway up the slope, under identical habitat conditions (so far
as could be observed) as the Helix. They became much more
numerous on the upper parts of the talus slope and on the cliff
itself and continued on to the top of the cliff, a near-vertical
distance of at least 500 feet. On the upper parts of the talus
eround or

oS
rock, or on occasional cliff-side patches of moss or grass but

slope, and on the cliff, they usually occur on bare
unprotected by thorn-bush.

3. Jaminia (Euchondrus) albula (Mousson): Although
quite common as dead shells all over the hillside, living exam-
ples on these two mornings were rare. They were found coinci-
dent with the Helix, but also higher on the talus slope (not on
the cliff), in similarly protected spots where detritus had ac-
cumulated.

4. Buliminus (Buliminus) egregius Naegele: As dead shells,
these snails had been found on the surface of the highest part
of the talus slope, but more frequently on the cliff-side itself,
not only in protected crevices, but also out on small patches of
grass or moss where such occur on the rough cliff. They seem-
ingly did not continue to the top of the cliff, as the Levantina
did. Whereas the living Helix and Levantina could easily be
collected by the hundreds, Buliminus was rare, and only very
few living B. egregius were found. (Three of these are those
mentioned by Biggs (1962, p. 69) as being collected by Kent
Flannery on April 17; the correct date is April 18.)

5. Helicella langloisiana: When collecting empty shells, this
is by far the most frequent snail on the hillside, but the living
examples were few on the two mornings specified. The Helicella
were found on the slope under thorn-bushes and up the cliff
on and under small vegetational patches.

6. Zebrina carducha (Mertens): The few individuals of this
species were not distinguished by the collectors in the field from
specimens of Buliminus egregius; the two have been confused
even by experts, and the proper taxonomic position of the

14 Postilla Yale Peabody Museum No. 66

species carducha has only recently been established on the basis
of the internal anatomy (Forcart, 1962). As with B. egregius,
individuals of Zebrina carducha were limited to the cliff-side
niches. So far as can be determined in retrospect, the two
species were collected together.

It is obvious that the ecologic notes made to date on these
species of snails are not in themselves sufficient for valid paleo-
environmental conclusions. Still, a beginning has been made,
and one continues to wonder how much different the climate
could have been and yet have this same gastropod assemblage
represented—as it is at Warwasi—for periods of tens of thou-

sands of years of the late Pleistocene. During this time, a

1g
major period of glaciation occurred in the Zagros Mts.
(Wright, 1961), with considerable depression of the mean an-
nual temperature (possibly, but not necessarily as much as
12°C for the higher areas). The coincidental depression in alti-
tude of the permanent snow-line has not been studied for the
northeastern (interior) side of the Zagros Mts., as in the area
of Kermanshah, where the annual precipitation is much less
than on the outer (southwestern) aspect; in this latter area,
the snowline some 20,000 years ago was to be found at ap-
proximately 2,100 m to 1,500 m (ca. 6,750-4,850 ft.), if
Wright’s conclusions are correct.

The permanent snowline on the inner side of the mountains
would have been higher (even though the inner side is typically
colder), due to a lower annual precipitation (a precipitation
possibly no greater than that today [Bobek, 1954]). The
mean annual temperature, however, would have been lower
(possibly 5° C, possibly more) than that of today, so that
evaporation would have been less than it is now with a result-
ant more humid environment.

Whatever the details of the climatological factors, which
should be determined in major part by geological investiga-
tions, the snails at Warwasi remain the same. At Shanidar
Cave, on the “outer” side of the mountains, there is a human
occupation hiatus of some 17,500 years (between ca. 30,000
years ago and 12,500 years ago, as derived from C'™ deter-
minations), which period neatly coincides with that postulated

.

pul
7 |

Sept. 10, 1962 Snails on a Persian Hillside

for the Wiirm glacial maximum in North America and Europe.
The assumption is that man, because of cold, depressed snow-
line, and coincidentally depressed treeline, could not live in the
region of Shanidar Cave during this period (an assumption
which is perhaps questionable for the latter part of the period,
when conditions must have been warmer and both snowline and
treeline higher).

However, no such occupational gap has been detected for
Warwasi (although there are no C' determinations as yet on
any of the levels of this site). The inference is that man con-
tinued to live in the area of Warwasi, and thus of the Ker-
manshah valley as a whole, at an altitude of 1,300 m (4,200 ft.)
and higher, during a long period when he supposedly was ex-
cluded from the region of Shanidar Cave at 700 m (2,200 ft.).
One can only say from a study of the fauna (including snails )
that there was no noticeable faunal change at Warwasi
throughout this period of the last 40,000 years or so of the
Pleistocene, and at Shanidar Cave there was no noticeable
faunal difference between the last of the Baradostian cultural
layers (at ca. 30,000 years ago, prior to the occupational hia-
tus) and those of Solecki’s **Mesolithic” layer (following the
cultural gap, and beginning ca. 12,500 years ago) (Reed and
Braidwood, 1960).

Perhaps long-term experimental studies on the environ-
mental limitations of the snails of the Tang-i-knisht hillsides,
coupled with intensive field studies over varied environmental
areas where these snails may be found today, would throw some
light on these problems. There is, thus, much work for the
future.

GASTRONOMIC EXPERIMENTS

Several kinds of Helix, are the edible snails of southern
Europe, and the evidence of hundreds of thousands of similar
shells in archeological sites of 12,000 to 8,000 years ago in
northern Africa, as well as in Iraq and Iran, showed that snails
of this genus were being eaten then as well. Obviously, these
snails were good human food. Why, however, were the some-
what larger Levantina, occurring on the same slopes for at
least some tens of thousands of years, and as numerous or al-

16 Postilla Yale Peabody Museum No. 66

most as numerous as are the Helix on those slopes today—why
were these snails not eaten? (At least they do not occur in any
great numbers in any archeological site hitherto excavated by
the Oriental Institute and so we must assume they were not
often gathered in our area; an occasional Levantina shell at
Jarmo or Tepe Sarab may, we think, be due to a mistake by
some small child helping its mother in the gathering.)

It was with pleasurable anticipation, therefore, with respect
to the Helix salomonica but with some apprehension with re-
gard to the Levantina diulfensis that we took more than 300
of the former and more than 200 of the latter and prepared
them for eating in the best French tradition (Rombauer, 1951,
p. 257). All members of the expedition participated in the
experiment (April 19, 1960) ; indeed, we had a festive occasion,
with special ‘snail-picks” being provided, made from some of
the microlithic bladelets from Tepe Sarab (fig. 4). (We are
not claiming that these microliths were used originally as snail-
picks; we only showed that they could be.) This gourmet ex-
periment proved to our complete satisfaction that both the

vood.

Heliv and the Levantina are uniformly g

Fig. 4. Prehistoric microlith from Sarab, mounted to be used as a snail-pick
for gastronomic experiments.

However, it was assumed that 8,000 years ago the techni-
ques of French cooking were not available to the people of
west-central Iran (although we have no way of knowing, of
course, what variety of herbs they may have used to flavor
their food), so a few hardier spirits tried a second experiment.
Both H. salomonica and L. diulfensis were boiled 15 minutes,
and then eaten hot, without salt or any other flavoring. Sur-
prisingly, they are both acceptable food under these Spartan
conditions; the Helix comes out of the shell quite easily, the
Levantina perhaps a bit less so, and the latter retains maybe
a bit more mucus (tasteless), but it seems hardly possible that
such minor factors were those which restrained the prehistoric

Sept. 10,1962 Snails on a Persian Hillside ils

populations of the Zagros slopes from eating the Levantina.
We found these snails quite tasty, and cannot imagine why
they were not eaten in former times; the experiments, from
the point of view of such “action archeology,” were therefore
a failure, although gastronomically a success.

The mode of preparation of the snails for cating under
prehistoric conditions is a problem we have not solved. Since
most of the shells are intact as we find them, the animals must
have been killed prior to extraction, inasmuch as the living
animals cannot be extracted from their shells without breaking
these. Although the animals could be killed by drowning, we
presume that the mode of killing was by cooking, but have
no proof of this. We cooked them by boiling, for us a simple
and effective method; the boiling dees not affect the shells in
any way that we could see, although Matteson (1959) noted
that extended boiling of certain terrestrial snails from Tllinois
tends to cause the epidermis to flake away from the rest of
the shell. However, extended boiling—if the snails were boiled—
is not necessary for their preparation as food.

If the snails were cooked by boiling, as we first casually as-
sumed, the question arises as to what were the containers in
which they were boiled. Pottery is unknown prior to about
8,500 years ago, and for earlier times, we have no archeological
evidence of containers in which boiling water could have been
held. In answer to some who have suggested that this earliest
pottery was too coarse to have been used to hold boiling water,
Dr. Frederick Matson, who assisted in the excavations and has
studied the ceramics from Jarmo and Sarab, has written, ‘The
pottery from Sarab and Jarmo could easily withstand boiling
water. The vessels are made of fired clay, and, aside from their
porosity, would not react with the water. However, I doubt if
they were used to boil snails or prepare stews because most
of the larger flat-based vessels have very thin bases and there
might be a problem with respect to the weight of the water
unless the pots were nested in the fire with adequate support
beneath them. Also, I do not recall seeing smudge and burning
marks on the exteriors of the lower parts of the vessels that
would suggest such firing....A small amount of water in a pot
full of snails would not require a lot of basal support for

18 Postilla Yale Peabody Museum No. 66

weight. If the pot were covered with a flat slab of some sort
(skin, sherd, wood, or smaller jar that just fit the mouth) the
snails could be steamed without requiring the presence of much
water. ... I would not want to rule out the boiling or steaming
of the snails, because it would be physically possible, but I
wonder if it wouldn’t be easier to roast them in hot ashes”
(Matson, personal communication).

However, as mentioned, pottery was unknown for much of
the period for which we think eating of snails to have occurred,
and actually for neither the pre-pottery or pottery-making
cultures do we have any real knowledge of the mode of prepara-
tion of the snails.

In a similar situation in Alabama, where large numbers of
snail shells were found in pre-pottery cultural associations,
Morrison (1942, p. 381) thought that the snails were steamed
in pits beneath a fire. Again, we have no archeologic evidence
for or against such a hypothesis; we only know that, both for
our sites in southwestern Asia and for those in Alabama (as
well as those from prehistoric Jericho [ Biggs, 1960]), the
great majority of the shells show no signs of charring, and
thus we assume the animals were not roasted on hot rocks.

SUPERSTITION
Our cook and two house-boys were town bred; they were
familiar with snail-shells as shells, but were astounded to dis-
cover that each housed a living animal. They were of the opin-
ion that no one of all the people they knew had any idea that
these shells were anything other than what they were commonly
regarded to be: snake pillows!

ACKNOWLEDGMENTS

My zoo-archeological work in southwestern Asia in 1954-
1955 and 1960, upon which the present paper is based, was
financed in large part by grants from the National Science
Foundation to the Department of Anthropology and to the
Oriental Institute of the University of Chicago. While in the
field, all members of the expeditions assisted with the work in
one way or another, and to each of these numerous people I ex-
tend my thanks. I am particularly grateful to Kent Flannery,
Jim Knudstadt, and Gene Garthwaite for the gathering of

Sept. 10,1962 Snails on a Persian Hillside 19

more than 500 live snails in the vicinity of Warwasi on the
drizzling morning of April 18, 1960. I am most indebted to
the Rev. H. E. J. Biggs, of Bromley, Kent, for his identifica-
tions, not only of the living snails sent him, but also of hun-
dreds of dead shells collected from numerous recent surfaces
and ancient dwelling-places. Additionally, he has spent many
patient hours, initiating the present author (who is not a mala-
cologist) into the mysteries of terrestrial snails from the region
of the Zagros Mts. of Iraq and Iran.

SUMMARY

The finding of shells of certain terrestrial snails (particu-
larly of Helix salomonica and several species of Levantina) in
archeological context throughout the upper Quaternary of the
slopes of the Zagros Mts. of southeastern Iraq and western
Iran led to the speculations concerning: 1) the use of Heliw as
food by the prehistoric people involved, and: 2) the possible use
of the presence of these shells and others as ecologic indicators
of past environments.

Pleasurable gastronomic experiments indicated that both
Helix salomonica and Levantina diulfensis were equally accept-
eble as food to modern archeologists and their colleagues ;
however, during the late Pleistocene and early Recent when
snails were being gathered fer food, all evidence indicates that,
in the area studied by us, the Helix were eaten and the Levan-
tina were rejected. We have no explanation for this choice by
the prehistoric peoples involved.

The use (and possible mis-use) of terrestrial snails as paleo-
ecologic indicators are discussed, and some preliminary ecologic
notes are made on six species which have been found in late
Quaternary archeological sites and which presumably may have
importance to paleo-envircnmental studies. However, only a
bare beginning has been made in this type of study, particu-
larly for the area of the Zagros Mts., and much more intensive
study is necessary before any valid paleo-environmental deduc-
tions can be made on the basis of the terrestrial snails.

BiprioGRAPHyY
Bakker, A. J., 1956. Climate. Report to the Government of Iran on the
Development of land and water resources in Khuzistan. Food and Ag-
ricultural Organization of the United Nations (Rome), Rep. 553:44-57.

20 Postilla Yale Peabody Museum No. 66

Biggs, H. KE. J., 1959. Some land Mollusca from northern Iraq. Jour.
Conch., 24:524-3847.

—, 1960. Mollusca from prehistoric Jericho. Jour. Conch., 24:

379-387.

—, 1962. Mollusca of the Iranian plateau—II. Jour. Conch., 25:

65-72.

Bobek, H., 1951. Die natiirlichen Wiilder und Geholzfluren Trans. Bonn.
Gecg. Abh., 8:1-62.

——_—, 1954. Klima und Landschaft Irans in yor- und friigeschicht-
licher Zeit. Geog. Jhb. Osterreich, 25:1-42.

Braidwood, Robert J. and Bruce Howe, 1960. Prehistoric investigations in
Iraqi Kurdistan. Oriental Inst., Univ. Chicago, Stud. Ane. Orient.
Civiliz., no. 31: i-xxviii, 1-184.

Braidwood, Robert J., Bruce Howe and Charles A. Reed, 1961. The Iranian
Prehistoric Project. Science, 133:2008-2010.

Braidwood, Robert J. and Charles A. Reed, 1957. The achievement and
early consequences of food production: A consideration of the archeo-
logical and natural-h‘storical evidence. Cold Spring Harb. Symp. Quant.
Biol., 22:19-29.

Kiseley, Loren C., 1937. Index Mollusca and their bearing on certain prob-
lems of prehistory: A critique. Publ. Phila. Anthrop. Soc., 1:77-93.
Foreart, L., 1962. Revision of Buliminus carduchus von Martens. Jour.

Conch., 25:54.

Ganji, M. H., 1960. Iranian rainfall data. Uniy. Tehran Arid Zone Research
Centre, Publ. 3:1-191.

Harris, Stuart A., 1961. On the land snails of Iraq and their potential use
in determining past climatic conditions. Sumer, 17:107-113.

Johnson, Ralph Gordon, 1960. Environmental interpretation of Pleistocene
marine species. Jour. Geol., 68:575-576.

Matteson, Max R., 1959. Snails in archeological sites. Amer. Anthro.,
61:1094-1096.

Morrison, J. P. E., 1942. Preliminary report on mollusks found in the shell
mounds of the Pickwick Landing Basin in the Tennessee River Valley.
Smiths. Inst., Bur. Amer. Ethn., 129:337-392.

Pabot, H., 1961. The natural vegetation of the Khuzistan region and head-
waters. Food and Agriculture Organization of the United Nations
(Rome). 1-80.

Reed, Charles A. and Robert J. Braidwood, 1960. Toward the reconstruc-
tion of the environmental sequence of northeastern Iraq. Oriental Inst.,
Univ. Chicago, Stud. Anc. Orient. Civiliz., 31:163-173.

Robison, William C. and Arthur V. Dodd, 1955. Analogs of Yuma climate
in south central Asia (India-Pakistan-Afghanistan-Iran). Environ-
mental Protection Division, Headquarters Quartermaster Research and
Development Command, (Natick, Mass.). 1-24.

Rombauer, I. S., 1951. The Joy of Cooking. The Bobbs-Merrill Company,
Inc. (Indianapolis). i-vii, 1-1013.

Solecki, Ralph S. and Arlette Leroi-Gourhan, 1961. Palaeoclimatology and
archaeology in the Near East. Ann. New York Acad. Sci., 95:729-739.

Wright, H. E., Jr., 1961. Pleistocene glaciation in Kurdistan. Kisz. und
Gegenw., 12:151-164.

4

LAE

YALE PEABODY MUSEUM

oF NaTURAL History

Number 67 September 17, 1962 New Haven, Cenn.

NEW SUBSPECIES OF BIRDS
FROM LUZON, PHILIPPINES

KENNETH C. Parkers

CarRNEGIE Museum, Pirrspurcu, PENNSYLVANIA

During the course of my studies on Philippine birds, initi-
ated in 1956 following an expedition to central Luzon, taxo-
nomic revisions of several species have been undertaken, some
of which have already been published. In the present paper,
five additional subspecies from Luzon are described. Of these,
two represent forms from Luzon as a whole, as distinct from
those of other islands, while three reflect geographic variation
within Luzon itself. The latter may be added to Gilliard’s list
(1950:473) of bird species geographically variable on Luzon,
which I have already amended to some extent (Parkes, 1958: 2).

Gallicolumba luzonica griseolateralis, subsp. nov.

Type: Adult 2? (Y.P.M. No. 48108) collected at Mt. Si-
capo-o, Ilocos Norte Province, Luzon, Philippines, April 18,
1959, by D. S. Rabor (original no. 18557).

Diacnosis: Similar to G. 1. luzonica (Scopoli) of central
and southern Luzon (type locality, vicinity of Manila), but
crown darker gray, less clearly defined from back of head;

a Postilla Yale Peabody Museum No. 67

grays of wings somewhat darker; flanks and under tail coverts
of females darker, the rufous much mixed with gray, and lower
breast and abdomen purer white, less ‘‘stained” in appearance.

Rance: Northern Luzon; I know of no specimens presently
extant from areas where intergradation with luzonica might
take place.

Remarks: Sexual dimorphism in the Bleeding-heart Pigeon
does not appear to have been described in the standard ornitho-
logical literature, and has been barely mentioned by avicul-
turalists. Delacour (1959:63) states that the female is ‘ta little
smaller and duller, with a thinner bill and a smaller head. Her
underparts show more buff.” There are actually several color
differences between the sexes. Females have the edges of the
flight feathers of the wing more strongly rufescent and are
more heavily washed with rufous or pinkish-buff on the flanks
and under tail coverts. There is a marked tendency for the
iridescent color of the back to invade the area of the bend
of the wing, which is clear gray in males. Males have, on
the average, a somewhat larger red breast-spot, posterior to
which is, in most individuals, a strong pinkish wash. In females
of luzonica this wash is pinkish-buff mixed with gray, giving
the underparts a stained appearance; this effect is less well
marked in griseolateralis, as mentioned above in the diagnosis
of that race.

Delacour and Mayr (1946:95-96) treated all of the Philip-
pine representatives of Gallicolumba as subspecies of G. lu-
zonica. Later, however, Delacour (1959:63-64) recognized at
least two species (total number not clear from his discussion).
Peters (1937:133-134) admitted five species. Certainly the
level of differentiation represented by luzonica and griseo-
lateralis, or by criniger, leytensis and basilanica, more closely
corresponds to the usual Philippine (and continental) sub-
species than does that represented by luzonica versus criniger.
As is true of so many insular representative forms, the Bleed-
ing-heart Pigeons have attained several stages of differentia-
tion. To call all of these forms subspecies of luzonica is to
use our admittedly imperfect trinomial system to mask certain
obvious interrelationships. Within such a system it is impos-

Sept.17, 1962 Birds From Luzon, Philippines 3

sible to avoid compromise. In this case the most useful solution
seems to be the treatment of these pigeons as a single super-
species, divided into several species corresponding to what are
usually specific levels of differentiation in appearance among
continental forms. Some of these, in turn, exhibit geographical
variation at the subspecific level.

Genuinely wild Bleeding-heart Pigeons from Luzon, with ac-
curate locality data, are far outnumbered in museum collec-
tions by aviary specimens. All of the latter that I have seen
have been referable to the southern race, so that it appears
reasonably certain that the original material of luzonica, even
if obtained in a Manila market, came from this population.
A total of five northern and nine southern Luzon specimens
with accurate data formed the basis for these notes.

Copsychus saularis heterogynus, subsp. nov.

Tyrr: “6” [= adult 2 ] (Carnegie Mus. No. 139025), col-
lected at Pangil, Laguna Province, Luzon, Philippines, in
November, 1958, by N. A. Icarangal.

Diacnosis: Females differ from those of C. 5. mindanensis
(type locality, Mindanao) in being more heavily washed with
buff on the posterior underparts, and in having this area,
especially the flanks and under tail coverts, finely but distinctly
barred with white. Males are not separable from mindanensis.

Rance: Island of Luzon. No specimens from Polillo, Catan-
duanes or Marinduque were examined; these islands tend to be
inhabited by Luzon subspecies of polytypic Philippine species.

Remark: De Schauensee and du Pont (1959:3) have men-
tioned the fact that Dyals from the Sulus have somewhat larger
bills than those from the remainder of the Philippines. Actually
the Sulu birds represent the extreme of a slight cline in bill
size; although many specimens from both ends of the Philip-
pine archipelago can be matched with one another, the indi-
viduals with longest and heaviest bills are mostly from more
southern islands (Negros, Mindanao, Basilan, Sulus).

Of C. s. heterogynus, 14 males and 14 females from Luzon
were examined, and of C. s. mindanensis, the following: Sibuyan,

4 Postilla Yale Peabody Museum No. 67

3, 1.2 & Mindoro, id.e2 23) Samar, id. ee) Cebueaiac
(juv., identified by probability) ; Negros, 44, 42 ; Mindanao,
43,492; Basilan, 7¢, 12; Sulu archipelago, 46,192.

Dicaeum aeruginosum striatissimum, subsp. nov.

Tyre: adult 6 (Carnegie Mus. No. 95037), collected at
Solsona, Ilocos Norte Province, Luzon, Philippines, December
8, 1923, by R. C. McGregor e¢ al.

Diacnosis: Differs from D. a. aeruginosum (type locality,
Cebu) in being more heavily streaked below, with streaks ex-
tending farther back along flanks and on abdomen; dorsum
sootier, with less contrasting greenish wash on upper tail cov-
erts and edges of remiges and rectrices; white spot on inner
web of outer rectrix larger but less sharply defined. Differs
even more from D. a. affine of Palawan, which is markedly
greenish above and sparsely streaked below.

RANGE: Specimens examined from islands of Luzon and
Sibuyan.

Remarks: I follow Salomonsen (1960a:5-6) in considering
D. aeruginosum to be specifically separable from D. agile,
with which it has been associated by recent authors. Salomon-
sen believed that “D. a. aeruginosum shows no geographical
variation within its range,” but had examined only Luzon and
Mindoro specimens.

There is a cline in the Philippines from north to south in
decrease of ventral streaking and increase of greenish on back
and wings. Thus, although Luzon birds as a group are sufhi-
ciently distinct from those of the more southern islands to
be worthy of nomenclatorial recognition, the characters of
striatissimum are best developed in northern Luzon; the type
selected is therefore a northern bird. In some species (cf.
Dicrurus) the Mindoro population is most closely related to
that of Palawan, lying to the southwest. In Dicaeuwm aeru-
ginosum, on the other hand, Mindoro les on the main north-
south cline, and Mindoro birds are almost exactly intermediate
between striatissimum and aeruginosum. They are nearest the
former in back color and nearest the latter in amount of

~

Sept.17, 1962 Birds From Luzon, Philippines 5

ventral streaking (6 specimens examined). Of striatissimum,
16 Luzon specimens and 1 Sibuyan specimen have been exam-
ined. Of aeruginosum, 6 Cebu and + Negros specimens were
seen. The one known specimen from Mindanao (Salomonsen,
op. cit.:15-16) probably belongs here. Of affine, + Palawan
specimens were examined.

Dicaeum hypoleucum lagunae, subsp. nov.

Tyre: adult 2? (Carnegie Mus. No. 139066), collected at
Pangil, Laguna Province, Luzon, Philippines, in November,
1958, by N. A. Icarangal.

Diacnosis: Similar to D. h. obscurum from northern Luzon,
but underparts heavily washed with olivaceous green, brightest
and yellowest on mid-abdomen, rather than almost neutral
gray with relatively little greenish wash. Quite different from
the southern races pontifea, mindanense and hypoleucum, which
are progressively more bicolored, culminating in the sharply
black-and-white D. h. hypoleucum, the only race with pro-
nounced sexual dimorphism. In lagunae, as in obscurum, the
sexes are alike in color.

Rance: Presently known only from Laguna Province, south-
central Luzon, Philippines.

Remarks: Several birds of the foothills and highlands ap-
pear to have an interrupted range in Luzon, with distinctive
northern an dsouthern subspecies (cf. Ptilinopus merrilli, Cult-
cicapa helianthea). This hiatus may be an artifact based on
insufficient collecting, as there are still large areas of Luzon,
especially in the eastern Sierra Madre, which are all but un-
known ornithologically. In any case, Dicaeum hypoleucum is
here shown to be still another species known from areas in
northern and in southern Luzon, with a subspecies in either
area. I have examined 28 specimens from various northern
Luzon localities, in the collections of the American Museum
of Natural History and the Yale University Peabody Mu-
seum. Of lagunae I have seen only the type, in Carnegie
Museum, and three specimens in the American Museum (one
each from Mt. Makiling, Los Banos, and “Southern Luzon”).

6 Postilla Yale Peabody Museum No. 67

Salomonsen (1960b:3), when reviewing this species, stated
that he had seen specimens of obscurum from Mountain Proy-
ince only, although he supposedly had available the material
belonging to the American Museum of Natural History.

Dicaeum pygmaeum salomonseni, subsp. nov.

Tyre: adult ¢ (Y.P.M. No. 48608), collected at 1600’ on
Mt. Sicapo-o (Mt. Simminublan), Ilocos Norte Province, Lu-
zon, Philippines, April 4, 1959, by D. S. Rabor and R. B.
Genzales (original no. 21123).

Diacnosis: Similar to D. p. pygmaeum of central Luzon
> here
restricted to the vicinity of Manila), but males with dorsum

south to the Surigao Strait (type locality, ‘Luzon,’

blacker, less green, contrasting more with rump; flanks grayer,
less green; sides of head duller gray, less sharply defined from
the white of the throat. In both pygmaeum and salomonseni
the breast color is highly variable, but in the latter race the
orange is less often than in pygmaewm concentrated into a
suggestion of a definite breast spot. Females of salomonseni
are duller, grayer, less green above than those of pygmaeum,
with the yellow-green area of the rump duller and less exten-
sive; underparts with less buffy wash; flanks grayer, less

green.

Rance: Northern Luzon, known from Mountain and Ilocos

Norte provinces.

Remarks: Negros males are slightly paler on the sides of
head than males from southern Luzon, but are not subspecifi-
cally separable from D. p. pygmaeum. It is of interest to note
that Salomonsen (1960b:15-16) found that specimens from
the Babuyan Islands north of Luzon differed from “typical
pygmaeum” in certain characters, some of which (‘deeper
orange vinaceus [sic] tinge on the under parts, slightly darker
olive flanks, and the dark longitudinal patch on the center of
the lower breast and abdomen more well marked and distinct”’)
represent trends opposite to the differences observed between
the south Luzon pygmaeum and the north Luzon salomonseni.

Of. D. p. salomonsent, 6 specimens from Mountain Province,

Sept. 17, 1962 Birds From Luzon, Philippines i

16 from Ilocos Norte Province, and 1 from “North Luzon”
were examined. Of D. p. pygmaeum, the following Luzon speci-
mens were examined: Manila, 1; Bataan Prov., 5; Rizal Prov.,
5; Laguna Prov., 3; “Luzon,” 1. Also Marinduque, 1; Si-
quijor, 1; Mindoro, 5; Bohol, 2; Negros, 20.

This new flowerpecker is named for the eminent Danish or-
nithologist Dr. Finn Salomonsen, an authority both on Philip-
pine birds and on the family Dicaeidae.

ACKNOWLEDGMENTS

The 1956 expedition in which I participated was part of a
project of the Graduate School of Public Health, University
of Pittsburgh, under the sponsorship of the Commission on Viral
Infections, Armed Forces Epidemiological Board, and sup-
ported in part by the Office of the Surgeon General, United
States Department of the Army. Dr. Canuto G. Manuel, then
Chief Zoologist of the National Museum of the Philippines,
was helpful in many ways, including arrangements for two
visits to the Mount Makiling area in Laguna Province. Mr.
Reuben G. Almazan, Officer in Charge of Makiling National
Park, was kind enough to donate two specimens of Gallico-
lumba luzonica from a locally-trapped flock, and arranged for
permission to collect birds in the Park.

Specimens for comparison were borrowed through the cour-
tesy of the authorities of the American Museum of Natural
History, Yale University Peabody Museum of Natural His-
tory, Chicago Natural History Museum, and United States
National Museum. Specimens were also examined in sitw at the
first two museums named, and at the California Academy of
Sciences and the Museum of Vertebrate Zoology, University
of California. Study of Philippine specimens at the last two
named institutions took place during visits primarily con-
cerned with an unrelated project supported by a grant from
the National Science Foundation.

LarerRAtuRE Cirep

Delacour, J., 1959. Wild pigeons and doves. All-Pets Books, Fond du Lac,
Wisconsin. 116 p.

Delacour, J. and E. Mayr, 1946. Birds of the Philippines. Macmillan, New
York. 309 p.

8 Postilla Yale Peabody Museum No. 67

De Schauensee, R. M. and J. E. du Pont, 1959. Notes on Philippine birds.
Notulae Naturae, no. 322. 5 p.

Gilliard, E. 'T., 1950. Notes on a collection of birds from Bataan, I.uzon,
Philippine Islands. Bull. Am. Mus. Nat. Hist., 94; 457-504.

Parkes, K. C., 1958. A new race of the Blue-headed Fantail (Rhipidura

cyaniceps) from northern Luzon, Philippine Islands. Am. Mus. Novit.,
no. 1891. 5 p.

Peters, J. L., 1937. Check-list of birds of the world, 3. Harvard Univ.
Press, Cambridge. 311 p.

Salomonsen, F., 1960a. Notes on flowerpeckers (Aves, Dicaeidae). 2. The
primitive species of the genus Dicaeum. Am. Mus. Novit., no. 1991, 38 p.

Salomonsen, F., 1960b. Notes on flowerpeckers (Aves, Dicaeidae). 3. ‘The
species group Dicaeum concolor and the superspecies Dicaeum erythro-
thorax. Am. Mus. Noyit., no. 2016. 36 p.

bathe

YALE PEABODY MUSEUM

oF NatTurAL History

Number 68 September 24, 1962 New Haven, Conn.

STOMATOPOD CRUSTACEA COLLECTED BY THE
YALE SEYCHELLES EXPEDITION, 1957-1958

RaymMonp B. Mannine

Instirute oF Mariner Science, Universtry oF Miamt!

Although small in size and number of species, the collection
contains several rare species as well as several new records from
the Seychelles. Of particular interest is the discovery of an
undescribed species allied to Gonodactylus fimbriatus Lenz and
G. brevisquamatus Paul’son. In all, ten species in three genera
are represented, with all but two of the species in Gonodactylus.

In the following account, complete synonymies are given only
for the relatively rare species. In the case of common species,
reference is made to the synonymies compiled by Kemp, 1913,
and Holthuis, 1941, as well as more recent papers. The species
are arranged as in Kemp, 1913. In addition to the material
collected in the Seychelles proper, several specimens were col-
lected in the Maldives, at Ceylon, etc., during the expedition,
and these have been included in the account. Comparative
notes and sketches of G. fimbriatus and G. brevisquamatus have
been added in order to clarify their relationship to the new
Gonodactylus.

‘Contribution No. 404 from The Marine Laboratory, University of Miami.

2 Postilla Yale Peabody Museum No. 68

The measurement following the number of specimens is in
all cases total length, measured from the apices of the sub-
median spines of the telson to the anterior margin of the ros-
tral plate. All of the material has been deposited in the Yale

Peabody Museum (YPM).

ACKNOWLEDGMENTS

I would like to express my thanks to Dr. Willard D. Hart-
man, Curator, Division of Invertebrate Zoology, Yale Peabody
Museum for making this interesting collection available for
study. I would also like to thank Dr. EK. Deichmann, Museum
of Comparative Zoology at Harvard, who kindly loaned 4 speci-
mens of G. fimbriatus and Mr. R. W. Ingle, British Museum
(Natural History), who loaned a specimen of G. brevisqua-
matus for comparative purposes.

The illustrations of the new species were drawn by my wife,
Lilly Manning.

Thanks are due to the National Science Foundation for their
support of stomatopod systematics under Grant G-11235.

Pseudosquilla ciliata (Fabricius, 1787)
Pseudosquilla ciliata, Kemp, 1913:96; Holthuis, 1941:261.
MareriaL. 1¢, 51.3 mm; YSE Sta. 9; seined offshore, Point

Elizabeth, north of Trincomalee, Ceylon; N. Mahadeva; 22 August
1957; YPM 4326.
Lystosquilla maculata (Fabricius, 1793)

Lysiosquilla maculata, Kemp, 1918:111, pl. 8, figs. 86-91; Holt-
huis, 1941: 269, fig. 5.

MarerraL. Ibrk. ¢, 164.8 mm; Seychelles; collected by local
inhabitants; YPM 4342.
Gonodactylus chiragra (Fabricius, 1781)
Gonodactylus chiragra, Kemp, 1913:155, pl. 9, fig. 107; Holthuis,
1941:277, fig. 7.
Matrriat. 19, 51.2 mm; YSE Sta. 12; Foul Pt., Ceylon;
W. D. Hartman; 28 August 1957; YPM 4328.

Sept. 24, 1962 Stomatopod Crustacea 3

16, 46.7 mm; YSE Sta. 17; Funadu Is., N. Male Atoll, Mal-
dives; shoreward edge of reef, living in cavities in coral; W. D.
Hartman; 19 September 1957; YPM 4329.

16, 41.0 mm, 19, 23.1 mm; YSE Sta. 28; seaward reef, Ile du
Coin, Peros Banhos Atoll, Chagos Archipelago; A. J. Kohn, W. D.
Hartman; 23 October 1957; YPM 4333.

19, 15.2 mm: YSE Sta. 35B; northwest of Moyenne Is., Sey-
chelles; A. J. Kohn; 20 January 1958; YPM 4456.

12¢,37.7 mm; YSE Sta. 41; seaward reef, Menai Is., Cosmoledo
Atoll; A. J. Kohn; 10 December 1957; YPM 4338.

1 juv., 6.8 mm; YSE Sta. 53; Pasquére, Praslin Is., Seychelles ;
W. D. Hartman; 19 January 1958.

Gonodactylus platysoma Wood-Mason, 1895

Gonodactylus chiragra var. platysoma, Kemp, 1913:162, text-
fig. 1; Holthuis, 1941:281.

MarteriaL. 1 2, 86.1 mm; YSE Sta. 25C; Gan Is., Addu Atoll.
Maldives; A. J. Kohn: 18 October 1957; YPM 43381.

1 OP NAGE Sremmmys VO SIE es Sitanmee ze:

7; lagoon side, Iie du Coin, Peros
Banhos Atoll, Chagos Archipelago; W. D. Hartman; 22 October
1957; YPM 4332.

Gonodactylus spinosus Bigelow, 18938

Gonodactylus spinosus Bigelow, 1893:101; Bigelow, 1894:493 ;
Nobili, 1906:330; Borradaile, 1907:210; Bigelow, 1926:519, figs.
1, 2.

Gonodactylus chiragra spinosus, Lanchester, 1908:454, pl. 23,
fig. 14.

Gonodactylus demani spinosus, Kemp, 1918:165, pl. 9, fig. 112;
Tattersall, 1921:361; Kemp and Chopra, 1921:311;
1936:9; Dollfus, 1938:215, fig. 17; Chopra, 1939:172.

Ramadan.

Materia. 192, 17.2 mm; YSE Sta. 80A; Bird Is., Seychelles;
W. D. Hartman; 6 November 1957; YPM 4334.

Remarks. As first pointed out by Kemp and Chopra (1921)
and confirmed by Bigelow (1926), G. spinosus differs from G.

a 4\* ,

4. Postilla Yale Peabody Museum No. 68

demant in having the inner branch of the uropod setose on all
margins; in G. demani the inner margin is devoid of setae.

There is no reason to maintain spinosus as a variety or sub-
species of G. demani, Varietal status does not have nomencla-
tural significance, and as both spinosus and demani are sym-
patric in the Red Sea as well as in other areas, they cannot be
regarded as subspecies. No overlap or variation has been shown
in the main character used to separate the two, so they must
be regarded as distinct species.

Chopra (1939) records this species from Mauritius to the
Gulf of Suez, Red Sea, and Persian Gulf, and from Zanzibar to
the Sea of Java, and Borradaile (1907) reported it from the

Seychelles.

Gonodactylus falcatus (Forskal, 1775)
Cancer falcatus Forskal, 1775 :96.
Gonodactylus glabrous Brooks, 1886:62, pl. 14, fig. 5, pl. 15, figs.
7-9; Kemp, 1913 :167, pl. 9, fig: 118.
Gonodactylus falcatus, Holthuis, 1941:284, fig. 9a.

Mareriav. 14. 40.1 mm; YSE Sta. 9; seined offshore, Point
Elizabeth, north of Trincomalee, Ceylon; N. Mahadeva; YPM 4327.

1 6,28.3 mm; YSE Sta. 22; Hulule Is., N. Male Atoll, Maldives:
W. D. Hartman; 5 October 1957; YPM 4358.

3 2 9, 29.7-35.9 mm; YSE Sta. 36; in coral cavities, Anse a la
Mouche (northern reef), Mahé Is., Seychelles; W. ID. Hartman;
24 November 1957; YPM 4335.

2 2 2, 35.0-43.3 mm; YSE Sta. 36; Anse a la Mouche (northern
reef), Mahé Is., Seychelles; A. J. Kohn; 22 December 1957; YPM
1336, 4337.

Gonodactylus crinitus, new species
Figure 1]

Holotype. 12, 27.5 mm; YSE Sta. 38; Beau Vallon, Mahé Is.
Seychelles; A. J. Kohn; 1 February 1958: YPM 4459.

Paratypes. 1 2 , 26.4 mm; data as in holotype; YPM 5510.
9 2, 28.2-28.8 mm; YSE Sta. 55; La Passe, Silhouette Is., Sey-
chelles; A. J. Kohn; 6 February 1958; YPM 4340.

Sept. 24, 1962 Stomatopod Crustacea 5

Ye AONE >
Ss : Imm

Figure 1. Gonodactylus crinitus, n. sp. Female paratype, Mahé Is., Sey-
chelles. A. Sixth abdominal somite and telson. B. Dorsal view of right
uropod. C. Ventral view of right uropod. Telson length=2.9 mm.

6 Postilla Yale Peabody Museum No. 68

Description. Rostral plate with three sharp anterior spines:
median spine elongate, extending beyond base of eves; lateral spines
broader, shorter, extending to base of eyes; rostral width at antero-
lateral angles greater than median length.

Carapace with anterolateral margins distally truncate and extend-
ing well beyond base of rostral plate; lateral margins of carapace
sinuate; posterior breadth greater than anterior.

Mandibular palp two- or three-segmented; five epipods present;
antennal scale extending to or beyond middle of cornea.

First five abdominal somites wtih an impressed pit on upper
median portion of pleuron; second to fifth somites with “reversed-
L-shaped” groove anterolaterally: sixth abdominal somite with six
longitudinal carinae, submedians broader than intermediates and all
are swollen, with irregular margins.

Telson with eleven dorsal carinae and four posterior spines. sub-
medians with movable tips; anterior pair of tubercles present: me-
dian carina swollen, with irregular margins; posterior margin of
telson with a prominent median tubercle; above and anterior to this
tubercle lie a pair of submedian tubercles: three carinae between
median carina and carina of intermediate tocth; first pair not
extending to anterior margin, fattening and widening anteriorly,
converging posteriorly ; second pair much shorter than first. converg-
ing posteriorly, not extending anteriorly past middle of telson: first
and second pair (lateral to median carina) may fuse posteriorly in
a transverse carina that runs toward median distal tubercle; third
pair thinner, lving mesial to posterior portion of the carina of the
intermediate tooth; carinae of intermediate tooth long. well-defined,
extending almost to anterior margin of telson; marginal carinae
sharp, fusing posteriorly with carina of intermediate tooth; a trans-
verse, raised ridge, biconcave posteriorly, extends from base of
third pair of carinae to base of carina of submedian tooth and from
there to under the median tubercle; submedian teeth with movable
apices and with 13-17 minute denticles on inner margin and a spinule
on outer margin; intermediate teeth with a spinule on inner margin.

Uropods with penultimate segment of outer branch not setose, ex-
tending beyond distal segment; penultimate segment armed with
seven lateral spines, first four small, movable, last three fixed. large,
antepenultimate by far the largest; last three with tips corneous and
strongly recurved; distal segment of outer branch elongate, paddle-
shaped, dorsal margin setose on either side, median strip not setose,
ventral margin not setose; inner branch elongate, outer margin con-
vex, tapering to tip, inner half of dorsal margin setose, with proxi-

Sept. 24, 1962 Stomatopod Crustacea ve

mal third of outer half without setae, ventral margin without setae;
basal prolongation with proximal spine slender, almost as long as
distal spine; distal spine recurved dorsally, outer margin concave
distally.

Color. Largely faded in preservative; carapace and body mottled
green and white; eyestalks with scattered dark chromatophores ;
mottling very pronounced on sixth abdominal somite and _ telson,
carinae of telson appearing banded with white and green.

Measurements. Holotype: total length, 27.5 mm; carapace length
5.6 mm; telson length, 2.9 mm; telson width, 3.6 mm. Female para-
type, YSE Sta. 33: total length, 26.4 mm; carapace length, 5.0 mm;
telson length, 2.9 mm; telson width, 3.4 mm.

Discussion. G. crinitus is closely related to both G. fim-
briatus Lenz and G. brevisquamatus Paul’son, but differs from
both in having 11 dorsal carinae on the telson, lacking setation
on the dorsal surface of the proximal segment of the uropod,
and lacking ventral setation on the ventral surface of the
uropod. Differences between the three species are summarized

in Table 1.

G. fimbriatus has been synonymized with G. brevisquamatus
since the remarks of Kemp (1913) and Tattersall (1921)
based on an unpublished account by Patience in which the two
were synonymized. The two species have not been directly com-
pared since that time. Tattersall’s action was followed by Kemp
and Chopra (1921), Ramadan (1986), and Dollfus (1988).
Serene (1949) kept the two species separate in his discussion
of the position of G. strigatus Hansen. The two species are
very closely related and in my opinion they are distinct. An
adequate series might show that they deserve only subspecific
status.

Four specimens of G. fimbriatus (all 2? 2, total length 21.4-
30.7 mm; Zanzibar; Cooke, col.; MCZ 7817) and one of G.
brevisquamatus (2, 27.2 mm; Tella Tella Kebira, Sudanese
Red Sea; C. Crossland, col.; B. M. (N. H.) registry 1936.9.30.
1-2) were examined: the latter specimen was one of those
examined by Tattersall (1921). The following notes were taken
from this material.

8 Postilla Yale Peabody Museum

Tasie 1. Comparison of

y

Gonodactylus brevisquamatus Paulson, G. fimbriatus Lenz,

and G. crinitus n. sp.

G. brevisquamatus

GC. fimbriatus

G. crinitus

Telson

Dorsal Carinae

Submedian denticles

Intermediate denticles

Uropods
Exopod, prox. seg.

Exopod, distal seg.

Endopod

Basal prolongation

9

+ 14, in one
series

1 on submedian,
1 on intermediate
tooth

with a dorsal
patch of setae

broadly rounded
distally, outer
and inner margins
about equally
convex; setose
proximally on
dorsal margin,

all of dorsal

setae short

short, rounded
distally ; com-
pletely setose
ventrally

outer spine
much larger
than inner

9

10-12 in 2
series

6-10 in 2
series on
submedian,
6-8 on
intermediates

with a dorsal
patch of setae

tapering distal-
ly, outer margin
much more convex
than inner;

setose proximal-
ly on dorsal
margin, all of
dorsal setae

short

short, bluntly
truncate
distally ; com-
pletely

setose
ventrally

outer spine
much larger
than inner

11

13-17 in
one series

1 on submedian,
1 on intermediate

no dorsal patch
of setae

tapering distal-
ly outer and
inner margins
about equally
convex; non-
setose prox.

on dorsal
margin, all of
dorsal setae
long

elongate,
tapering
distally ;
no ventral
setae

inner
outer spine
slightly
larger than

G. fimbriatus and G. brevisquamatus are very similar in gen-

eral appearance. They differ primarily in the number and ar-

‘angement of submedian and intermediate spinules on the telson

Table 1) and in the shape of the uropod segments. A ventral
| | 2

view of the uropod of each species is shown in Figure 2; seta-

tion is omitted for clarity. In G. brevisquamatus, the distal

segment of the outer branch is broadly rounded and the inner

branch tapers distally. In G. fimbriatus, the distal segment

Sept. 24, 1962 Stomatopod Crustacea 9)

of the outer branch tapers and the inner branch is truncate
distally. Both are similar in lateral spination of the proximal
segment of the outer branch. The inner spine of the basal pro-
longation is comparatively longer in G. brevisquamatus. Both
species have a pair of short submedian carinae on the ventral
surface; these carinae converge anteriorly but do not meet.

Imm
EE)

Figure 2. A. G. brevisquamatus Paul’son, Red Sea, ventral view of right
uropod; B. G. fimbriatus Lenz, Zanzibar, ventral view of right uropod.
Smaller lateral spines present in both species; they are obscured by
the basal prolongation in 2A.

10 Postilla Yale Peabody Museum No. 68

The setation of the uropods is characteristic in both. The
proximal segment of the outer branch bears a prominent, circu-
lar patch of setae on the dorsal surface. The ventral surface
of the uropods is completely setose, with a thick, circular, prox-
imal patch of strong setae.

G. fimbriatus has been reported from Zanzibar (Lenz, 1905:
38, pl. 47, fig. 11; 1910:572) and Coétivy, Seychelles (Bor-
radaile, 1907 :211). Serene (1949 :231, 235) mentioned a speci-
men from the Mergui Archipelago, perhaps from Kemp’s
(1913:175) reference to Patience’s material.

Other than Patience’s questionable record of G. brevisqua-
matus from the Mergui Archipelago, that species is known only
from the Red Sea. Patience’s specimens may actually have
belonged to G. fimbriatus rather than G. brevisquamatus. The
following papers record G. brevisquamatus: Paulson, 1875:
127, pl. 21, fig. 3; Nobili, 1906 :331; Kemp, 1913:174, pl. 10,
figs. 115, 116; Tattersall, 1921 :362, pl. 27, figs. 5, 6; Kemp
and Chopra, 1921 :311; Ramadan, 1936:10; Dollfus, 1938 :222,
hie: 21:

Remarks. The curious arrangement of setae and spines on
the uropods of G. brevisquamatus and G. fimbriatus is perhaps
worthy of further mention. The large, recurved spines on the
uropod may be used by the animal to “lock” itself in its burrow.
The uropods are curved ventrally, so that the telson and uro-
pods together form half of a circle. The inner branches of the
uropods recurve dorsally. The setae of the uropods are so ar-
ranged that, when viewed from behind the animal, they appear
as a rounded “tbasket,” as if they were utilized as a filtering
apparatus. All of the specimens of G. fimbriatus had the setose
areas loaded with detritus and sand which had to be washed
away before the uropod structure could be observed.

The significance of these structures is not readily apparent.
The mouthparts and raptorial claws appear unmodified, and it
seems unlikely that these species are filter-feeders. Observations
on these species in the field would be most welcome.

Characters based on the relative length of the rostral plate
or antennal scale in relation to the eye must be used with some
caution in the stomatopods. Such characters vary according to

Sept. 24, 1962 Stomatopod Crustacea a fe

the state of contraction or expansion of the body at preserva-
tion. In future work, more attention should be given to the
shape of the uropods and their investment of setae. These
characters coupled with the configuration of the telson are ap-
parently diagnostic in this group of Gonodactylus (Group I
of Kemp, 1913).

Name. The specific name is from the Latin, “crinitus,”
fringed, and refers to the setation of the uropods.

Gonodactylus lenzi Holthuis, 1941

Pretosquilla glabra Lenz, 1905:388, pl. 47, fig. 18; Lenz. 1910:
572 [listed only].

Genodactylus glaber, Kemp, 1913:182, pl. 10, fig. 121; Kemp,
1915:186; Roxas and Estampador, 1930:124, pl. 3, fig. 5; Roxas,
1930:18; Chopra, 1934:42; Gravier, 1937:208: Seréne, 1947:385,
fio ll, pl. 2:

Gonodactylus lenzi Holthuis, 1911:288; Tiwari and Biswas, 1952:
362; Seréne, 1954:6 [larval stages, p. 34 et seq.].

MareriAL. 1 6, 25.7 mm; YSE Sta. 33; Beau Vallon, Mahé Is.,
Seychelles; inhabiting dead corals; A. J. Kohn; 1 February 1958;

YPM 4458.

Remarks. The specimen agrees with Kemp’s account in al-
most all respects. The posterior spines of the telson are of
equal size. The corrugations of the lateral margins of the tel-
son are not visible, but the three central bosses of the telson
are faintly corrugated.

The specimen figured by Seréne (1947) has the bosses of the
telson much more inflated than in the present specimen; both
Seréne and Gravier (1937) commented on the variability of the
median bosses of the telson.

The color pattern in preservative is striking. The antennular
flagella are banded purple and white. The rostral plate, oph-
thalmic somite and the two distal segments of the raptorial
claw are bluish with scattered dark chromatophores. The body
is cream colored and densely covered with dark chromato-
phores which are aggregated in dark patches on the posterior
portion of the carapace, sixth and seventh thoracic somites,

12 Postilla Yale Peabody Museum No. 68

and second to fourth abdominal somites. Kemp (1913) found
these patches on the first, fourth, and fifth abdominal somites.
The fifth and sixth abdominal somites, telson, and uropods are
outlined in blue.

This species has not previously been recorded from the Sey-
chelles. The records in the literature are: Zanzibar (I.enz,
1905) ; Ceylon (Lenz, 1910); Andamans (Kemp, 1913: Tiwari
and Biswas, 1952) ; Nicobars (Chopra, 1934) ; Bay of Batavia
(Holthuis, 1941); Poulo Condore Islands (Gravier, 1937) :
Nhatrang, Viet Nam (Gravier, 1937; Seréne, 1947, 1954) ;
Philippines (Kemp, 1913; Roxas, 1930; Roxas and Estam-
pador, 1930).

Gonodactylus gyrosus Odhner, 1928

Gonodactylus gyrosus Odhner, 1923:11, pl. i, figs. 4, 5; Ward,
1942:56; Tiwari and Biswas, 1952:362, fig. 5; Holthuis, 1953:61.

MarTeriaL. 19%, 41.5 mm; YSE Sta. 55; La Passe, Silhouette Is.,
Seychelles; A. J. Kohn; 6 February 1958; YPM 4341.

Remarks. The rostral plate of this specimen is somewhat
shorter than that illustrated by Tiwari and Biswas (1952), as
the median spine does not exceed the cornea. The cornea 1s
noticeably bilobed, and the eyes are comparatively shorter.

The color pattern is similar to that shown by Tiwari and
Biswas. The background color is cream, with many dark chro-
matophores in patterns on the anterior part of the body. The
fifth abdominal somite is lined anteriorly and posteriorly with
green. The sixth abdominal somite and telson are mottled green
and white.

The mandibular palp is present, apparently two-segmented.

This species has not previously been recorded from the Sey-
chelles. The records in the literature are: Arno Atoll, Marshall
Is. (Holthuis) ; Gilbert Islands (Odhner) ; Andamans (Tiwari

and Biswas); Diego Garcia, Chagos Archipelago (Ward).

Gonodactylus guerini White, 1861

Gonodactylus guerini White, 1861:48, pl. 6; Bigelow, 1931:139
[and synonymy |; Townsley, 1953:428, figs. 20, 21a-f; Serene, 1954:

52 [discussion only J.

Sept. 24, 1962 Stomatopod Crustacea te

MareriaLt. 19, 35.5 mm; YSE Sta. 414A; anchorage off Menai
Is., Cosmoledo Atoll; 25 fms.; W. D. Hartman; 12 December 1957;

YPM 4447,

Remarks. The specimen agrees well with the description
given by Bigelow (1931). The right submedian elevation of the
telson bears a curved row of six instead of five spines as in
Bigelow’s material.

The color is as noted by Bigelow, with two dark transverse
bands on the carapace; each of the thoracic and abdominal
somites are banded. The spines of the telson are flushed with
pink,

Bigelow gave a complete synonymy, which will not be re-
peated here, and recorded the following distribution: Hawan,
Marquesas, Fiji, and Mauritius, in depths ranging from a few
feet to 60 fms. Townsley (1953) reported on other Hawaiian
specimens in depths of 50 to 120 fms. The present specimen
extends the range 900 miles to the north of Mauritius.

Lirerature Crrep

Bigelow, R. P. 1893. Preliminary notes on the Stomatopoda of the Al-
batross collections and on other specimens in the National Museum.
Johns Hopkins Univ. Cire., 72(106) : 100-102.

—————,, 1894. Report on the Crustacea of the Order Stomatopoda col-
lected by the steamer Albatross between 1885 and 1891, and on other
specimens in the U. S. National Museum. Proc. U. S. Natl. Mus., /7:
489-550, pls. 20-22, text-figs. 1-28.

, 1926. On the type of Gonodactylus spinosus, a stomatopod
Crustacean. Amer. Nat., 60(671): 579-582, text-figs. 1-2.

, 1931. Stomatopoda of the southern and eastern Pacific Ocean
and the Hawaiian Islands. Bull. Mus. comp. Zool. Harvy., 72(4) :105-191,
pls. 1-2, text-figs. 1-10.

Borradaile, L. A. 1907. Stomatopoda from the Western Indian Ocean.
The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under
the leadership of Mr. J. Stanley Gardiner. Trans. Linn. Soc. London,
Zool., ser. 2, 72: 209-216, pl. 22.

Brooks, W. K. 1886. The Stomatopoda collected by H. M. S. Challenger
during the years 1873-76. Report on the Scientific Results of the
Exploring Voyage of H. M. S. Challenger, /6:1-116, pls. 1-16.

Chopra, B. 1934. On the stomatopod Crustacea collected by the Bengal
Pilot Service off the mouth of the River Hughli, together with some
notes on other forms. Rec. Ind. Mus., Calcutta, 36:17-43, text-figs. 1-5.

—— ——, 1939. Stomatopoda. The John Murray Expedition, Sci. Repts., 6
(3) :137-181, figs. 1-13.

14 Postilla Yale Peabody Museum No. 68

Dollfus, R. Ph. 1938. Stomatopoda (II). Catalogue synonymique des
especes jusqu’a présent recoltées dans la Mer Rouge, y compris la
partie sud du Canal de Suez et le Golfe d’Aden. Mission Robert Ph.
Dollfus en Egypte. X. Mém. Inst. Egypte, 37:185-236, text-figs. 1-23.

Forksal, P. 1775. Descriptiones animalium, avium...et insectorum, quae ‘n
itinere orientali observavit. Kd. Carston Niebuhr, Havniae (not seen).

Gravier, C. 1987. Stomatopodes des cotes d Indochine. Ann. Inst. Oceanogr.,
Paris, 7:175-211, text-figs. 1-23.

Holthuis, lL. B. 1941. Biological Results of the Snellius Expedition, XII.
The Stomatopoda of the Snellius Expedition. 'Temminckia, 6: 241-294,
text-figs. 1-9.

————.,, 1953. Enumeration of the decapod and stomatopod Crustacea
from Pacific coral islands. Atoll Research Bull., No. 24:1-66.

Kemp, S. 1913. An account of the Crustacea Stomatopoda of the Indo-
pacific region. Mem. Ind. Mus., Calcutta, 4:1-217, pls. 1-10, 10 text-
figs.

——, 1915. On a collection of stomatopod Crustacea from the Philip-

pine Islands. Philippine Journ. Sci., /0D:169-186, pl. i.

Kemp, S. and Chopra, B. 1921. Notes on Stomatopoda. Rec. Ind. Mus.,
Calcutta, 22: 297-311, text-figs. 1-4.

Lanchester, W. F. 1903. Marine crustaceans. VIII. Stomatopoda, with an
account of the varieties of Gonodactylus chiragra. In: Gardiner, J. S.,
The fauna and geography of the Maldive and Laccadive Archipelagoes.
Being an account of the work carried on and of the collections made
by an Expedition during the years 1899 and 1900. vol. 1: 444-459, pl. 23.

Lenz, H. 1905. Ostafrikanische Dekapoden und Stomatopoden gesammelt
von Herrn Prof. Dr. A. Voeltzkow. Jn: Voeltzkow, A., Wissenschaft-
liche Ergebnisse der Reisen in Madagaskar und Ostafrika in den Jahren
1889-95. Vol. III. Abh. Senckenb. naturf. Ges., 27: 341-392, pls. 47-48.

—__——_——_—_— , 1910. Crustaceen von Madagaskar, Ostafrika, und Ceylon. Jn:
Voeltzkow, A., Reise in Ostafrika in den Jahren 1903-1905 ..., 2: 539-
570.

Nobili, G. 1906. Faune carcinologique de la Mer Rouge. Decapodes et
Stomatopodes. Ann. sci. nat. Zool., Paris, ser. 9, 4:1-347, pls. 1-11, text-
figs. 1-12.

Odhner, 'T. 1923. Indopazifische Stomatopoden. Goteborgs Vet. Handl., Ser.
4, 27(4):1-16, pl. 1.

Paulson, O. 1875. Studies on Crustacea of the Red Sea with notes regard-
ing other seas. I. Podophthalmata and Edriophthalmata (Cumacea), pp.
xiv -- 145, pls. 1-21. Kiev.*

Ramadan, M. M. 1936. Report on a collection of Stomatopoda and De-
capoda from Ghardaqa, Red Sea. Bull. Fac. Sci. Egypt Univ., No. 6:1-
43, pls. 1-2.

* Paul’son’s rare 1875 paper on the Crustacea of the Red Sea originally
appeared in Russian; an English translation of this work was published
in 1961 for the National Science Foundation and the Smithsonian Institu-
tion by the Israel Program for Scientific ‘Translations. It is available in
the United States from the Office of Technical Services, U. S. Department
of Commerce, Washington 25, D. C.

Sept. 24, 1962 Stomatopod Crustacea 15

Roxas, H. A. 1930. The Puerto Galera Marine Biological Laboratory of
the University of the Philippines (A report to the President of the
University, together with a check-list of animals of the Puerto Galera
region), pp. 1-24, pls. 1-4.

Roxas, H. A. and EK. Estampador. 1930. Stomatopoda of the Philippines.
Natural app. Sci. Bull., Manila, /: 93-131, pls. 1-6.

Serene, R. 1947. Sur les stomatopodes rares trouvés en Indochine et n’ exist-
ant pas dans les collections du Museum. Bull. Mus. Hist. Nat., Paris,
ser. 2, 19(5): 381-389, 4 pls.

—————, 1949. Observations sur le Gonodactylus strigatus Hansen
(Crustacé Stomatopode). Bull. Soc. Zool. France, 74(4-5) : 225-231, 2
text-figs.

—, 1954. Observations biologiques sur les stomatopodes. Ann. Inst.
Oceanogr. Monaco, 29:1-91, pls. 1-10, text-figs. 1-15.

Tattersall, W. M. 1921. Report on the Stomatopoda and macrourus De-
capoda collected by Mr. Cyril Crossland in the Sudanese Sea. Journ.
Linn. Soc. London, Zool., 34: 345-398, pls. 27-28.

Tiwari, K. K. and S. Biswas. 1952. On two new species of the genus
Squilla Fabr.; with notes on other stomatopods in the collections
of the Zoological Survey of India. Rec. Ind. Mus., Calcutta, 49: 349-363,
text-figs. 1-5.

Townsley, Sidney J. 1953. Adult and larval stomatopod crustaceans oc-
curring in Hawaiian waters. Pacific Sci., 7 (4) : 399-437, text-figs. 1-28.

Ward, M. 1942. Notes on the Crustacea of the Desjardins Museum, Mauri-
tius Institute, with descriptions of new genera and species. Mauritius
Inst. Bull., 2: 49-113.

White, A. 1861. Descriptions of two species of Crustacea belonging to the
Families Callianassidae and Squillidae. Proc. zocl. Soc. Lond., 1861: 42-
44, pls. 6, 7.

ihe

YALE PEABODY MUSEUM

or Naturau History

Number 69 October 10, 1962 New Haven, Conn.

NOTES ON SOUTH AMERICAN FLAMINGOS

Luis EK. PeEwNa*

INTRODUCTION

During the Chilean winter of 1957 we had our first contact
with the Andean species of flamingos on an expedition organized
by the Yale University Peabody Museum of Natural History
to the high ranges of the Andes in the province of Antofagasta,
Chile. Later, in the summer of 1957-1958, we visited this re-
gion with Dr. Roger Tory Peterson and stayed there for a
month studying these birds. In the summer of 1960, together
with Dr. William G. Conway, Director of the New York Zoo-
logical Park and Mr. Bates Littlehale of the National Geo-
graphic Society, we spent another month on this work, and later,
during the month of March of 1960, we extended the expedition
for an additional few weeks traveling through the Patagonian
region of Argentina. Recently, between October 1960 and
March 1961, we made another expedition to the Patagonian
region, including Tierra del Fuego.

DESCRIPTION AND DISTRIBUTION

Three species of flamingos that live and nest in the moun-
tains in the province of Antofagasta, Chile and the neighboring

* Casilla 2974, Santiago, Chile.

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.

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No. 69

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Oct. 10, 1962 South American Flamingos 3

region of Bolivia have been identified and observed (a fourth
species called “Guajchatata” by the natives was not identified ) :
Phoenicopterus chilensis Molina is characterized by the lack
of black coloration on the sides and back of the body when it is
at rest. The reddest part is on its tail (figure 3 A and B).
Geographically it is the most widely spread, having been found
on the high Andean ranges of Peru and Bolivia, in the Pata-
gonian region of Chile and Argentina including Tierra del
Fuego, in Southern Brazil and in Uruguay. It is the only
species that during certain months lives close to the sea. The
natives call this bird the ‘“’Tococo,” named for its cry.

Phoenicoparrus andinus (Philippi) has a black fringe on
its tail and has been called the “Black Tail Flamingo.” The
winey coloration is very marked at the base of the neck and
part of the chest (figure 4 A and B). This species is character-
istic of the Andean regicn between 15° and 27° latitude south.
It permanently inhabits the marshes and lakes of brine or of
fresh water. It has been found throughout the year in the

‘igure 2. The camp at Lejia Lake (Antofagasta Andes Range) at 12,2
Figure 2. Th mp at Lejia Lal Antofagasta Andes Rang it 12,200
feet above sea level where Phoenicoparrus jamesi (Sclater) was found in

large numbers.

4 Postilla Yale Peabody Museum No. 69

Atacama lake in Antofagasta, Chile, in Champaja, Argentina
and in Lake Titicaca in Peru and Bolivia. The native name
for this species is “Jetete” or “Jititi.”

Phoenicoparrus jamest (Sclater) has two red stripes which
fall gracefully over its wings; they are almost plume-like and
arise from the upper part of the back. The black plumes are
seen Only as spots on the sides. The neck and chest are of a
very winey color (figure 5 A and B). The colors of the feet
and beak as distinguishing features are of less importance, as
from a distance one is all too likely to make a mistake, these
parts of the body being often covered with mud. This species is
typical of the high shallow Andean salt lakes, where they nest
in large colonies. It has been observed in the Laguna Colorada,
Bolivia, in the Laguna de Lejia, Antofagasta, Chile, and in
the Salar de Atacama, Antofagasta, Chile in July 1957. It has
also been collected in Lake Titicaca, Peru and Bolivia, and in
Abrapampa, Argentina, The native name for this bird is “Chu-
TUG.

MIGRATION

Nesting colonies, always homogeneous as to species, on La-
guna Colorada and in the Laguna Verde, both on the Bolivian-
Chilean border, were visited on different occasions in the months
of December, January and February. In general we were able
to determine that the resident species of the regions of high
altitude is Phoenicoparrus jamesi; of the regions of inter-
mediate altitude, Phoenicoparrus andinus; and of the lower
regions, Phoenicopterus chilensis, which wanders into the higher
altitudes at intervals.

Phoenicoparrus andinus (Philippi) and Phoenicoparrus
jamest (Sclater) are both extremely common, numbering in
the thousands while Phoenicopterus chilensis Molina did not
seem to form such large flocks, except perhaps in the Patagonia
of Argentina. At the end of spring (November and December)
both species of Phoenicoparrus are around the high marshes
and salt lakes which are over 4,000 meters high and frozen
during the winter. At the end of February, March or April
they return to the areas of milder climate, the Andean marshes
and lakes of lower altitude (2,300 to 3,500 meters). Although

Oct. 10, 1962 South American Flamingos 5

Ph. andinus prefers to nest in the marshes of lower altitudes,
we have not confirmed that personally.

Phoenicopterus chilensis Molina invades the Chilean and Ar-
gentine Patagonian region up to Tierra del Fuego, the cen-
tral region of Chile (coast of Santiago province to the island
of Chiloé), disappearing totally during the summer months,
taking shelter to nest, possibly in the hidden lakes of the cen-
tral Andean region and the south of Chile and Argentina, but
this has not been proved. Nesting colonies of these species are
only known in the high mountain range of Antofagasta and the
region of Natales, Magallanes (Chile). Formerly there existed
a large colony in Laguna del Maule (Chile), but this has disap-
peared owing to construction work on a dam.

Some strays or small flocks of immatures may be found in
the high lakes and marshes in the winter just as they may be
found in the low marshes of the Patagonian region in the sum-
mer. The few flocks that actually winter in the high region of
the Andes lakes and marshes make use of the moderate tem-
perature springs occasionally found there, but these examples
are relatively rare. The rest of these marshes are completely
frozen and uninhabitable by these birds.

RELATIVE ABUNDANCE OF THE THREE SPECIES

When we were with Dr. Peterson at the Laguna Colorada
(Bolivia) during the month of January 1958, we had been able
to observe that the number of flamingos that were inhabiting
this lake was in the vicinity of 6,000. Of these the following
are the approximate percentages :

Ph. jamest ..... 97 per cent
Ph. andinus |. . 2 per cent
Ph. chilensigs 4... -. 1 per cent

During the last expedition from January 25 to the first of
March 1960, the concentration of flamingos on this lake had
diminished to possibly not more than 4,500. Making a reason-
able estimate, based on the daily visits of flocks that were ar-

6 Postilla Yale Peabody Museum No. 69

riving in the vicinity of our camp, we arrived at the following
percentages of the species:

Ph. jamesi Seay se ee eee 86 per cent
Ph “ands. — 9 eae 2 per cent
Eh. chilensis = a. ee 12 per cent

As is seen the percentage of Ph. chilensis grew noticeably.
From our observations, we feel that this is owing to the fact
that while it was the egg-laying period for all the birds, it
was the final one for Ph. chilensis, before they banded together
to seek a more temperate climate. For the constancy of the
percentage of Ph. andinus, the logical conclusion is that this
species is only a visitor to these high lakes and that its nesting
is probably carried out elsewhere.

As the data that we gathered on our trip to Salar de Aguas
Cahentes (Chile) illustrated, we found only small groups of
flamingos, though this is possibly due to our inability to dis-
tinguish the species from the great distances that separated us
or to the short time at our disposal (end of February 1960).
In the Laguna de Lejia, around the said marsh, we found about
1,500 flamingos, the relative cccurrence of the different species

being as follows:

Ph. jamesi 1 per cent
Ph. andinus .. 98 per cent
Ph. chilensis 1 per cent

PLATE

Figure 3. A. Schematic drawing showing lateral view of the body of
Phoenicopterus chilensis Molina and the position of red coloration. B.
Dorsal view of Phoenicopterus chilensis Molina showing positions of red
coloration on its tail.

Figure 4. A. Schematic drawing showing lateral view of the body of
Phoenicoparrus andinus (Philippi) and the distribution of the red and
black coloration. B. Dorsal view of Phoenicoparrus andinus (Philippi),
“The Black Tail Flamingo.”

Figure 5. A. Schematic drawing showing lateral view of the body of
Phoenicoparrus jamesi (Sclater) and the distribution of the red and black
coloration. B. Dorsal view of Phoenicoparrus jamesi (Sclater) showing the
two red stripes of plumes that fall over its wings and the small areas of
black coloration.

~

Oct. 10, 1962 South American Flamingos

8 Postilla Yale Peabody Museum No. 69

It is possible that the non-use of this lake by Ph. jamesi is
due to the fact that the great depth of the Laguna de Lejia,
for example, is not favorable to them, as is true of other places.
This great depth does not affect Ph. andinus because, as we
had observed on many different occasions in the Laguna Verde
and the Laguna de Lejia, one of their customs is to swim and
look for food while submerging the head, neck and part of the
body, in a manner similar to ducks. This custom was not ob-
served in the other species of flamingos. Ph. chilensis when
looking for its food effects a type of dance, slowly kicking his
feet in the mud, turning around and keeping his head sub-
merged in the turbulent and muddy water.

MEASUREMENT OF EGGS

From the colonies of Ph. jamesi we obtained 13 eggs which
were measured. The data in millimeters are recorded below:

Oo) neat 9.0 x 5.8
9.2 x 5.6 8:8) x ord
O22 oul On Xx Dal
Orix oe S625
Oa 526 8.5 x 5.6
GOR 5e5 Seoeeoee

9.0 x 5.5

As ean be seen from the measurements, there is an enormous
variation in size of Ph. jamesi eggs, only two having the same

¢

dimensions.

Lirerature Crrep

Behn, F., A. W. Johnson and G. Millie, 1957. Exploraci¢n ornitologica a
las cordilleras del norte de Chile. Soc. Biol. Concep. Chile 32: 95-131.

Pena, Luis E., 1961. Results of Research in the Antofagasta ranges of
Chile and Bolivia. Yale Univ. Peabody Mus. Nat. Hist. Postilla no.
49 33-42.

Conway, W. G., 1961. To the High Andes for the Rarest Flamingo. Animal
Kingdom Mag. Zool. Soc. 63:34-50.

————, 1961. In Quest of the Rarest Flamingo. Nat. Geogr. Mag.
120:91-105.

LIFE Magazine, 1960. The Wonders of Life on Earth. p. 148-153.

SOS: 7 J
D8 7

Dile

YALE PEABODY MUSEUM

oF Naturau History

Number 70 November 5, 1962 New Haven, Conn.

A NEW CAVERNICOLOUS PSEUDOSCORPION
BELONGING TO THE GENUS MICROCREAGRIS

Witi1aM B. MucHMmMore

University oF Rocuester, RocuEster, New York

This remarkable new form was found among collections in
the Peabody Museum lent by Dr. C. L. Remington.

Microcreagris grandis, yn. sp.

Material: Holotype male | Holotype No. E101] (WM 403.
01001) found July 6, 1946 by T. O. Thatcher ‘on formations
in cave,” Lehman Caves National Monument, White Pine Co.,
Nevada. Paratype tritonymph (WM 402.01001) found June 8,
1937 by T. O. Thatcher and R. Loir “on stalagmite in cave,”
also in Lehman Caves National Mcenument. Types in the Pea-
body Museum of Natural History.

Diagnosis: Mair. A very large species for the genus, modified
for cavernicolous life by considerable attenuation of the ap-
pendages, but not by loss of pigment or reduction of eyes. Color
of carapace and palps a dark reddish-brown, the abdomen and
legs much lighter. Carapace about one-third longer than broad,
rather narrow at the anterior margin and greatest in breadth
near the posterior end; broad, smoothly rounded epistome pre-

2 Postilla Peabody Museum No. 70

sent; four well-developed eyes of nearly equal size present ; sur-
face smooth anteriorly, becoming rather scaly posteriorly and
reticulated on the sides; carapacal setae 24 (4-5).

Abdomen elongate and typical of the genus; tergal chaeto-
taxy 529027211282 10279: 9: 1229: ?omms stemaleichacko-

13 2 2
taxy ZS: (35-4 ) (5)21(4 ) 5 ( 4 ) 16 (5 ) 5 16 = FT 6 5 13 =o IY
?:?:?:mm; pleural membrane typically granulate; genital

area typical. Chelicera of typical facies, about twice as long as
broad; six setae on palm; fixed finger with 19 or 20 irregularly
large and small teeth; movable finger with 20-21 such teeth;
galea bifid just distad of the middle with each branch termi-
nally divided into two short, pointed projections, reaching
about to the end of the finger; galeal seta not reaching the tip
of the galea; flagellum of 10 setae nearly all of which are
deeply serrate on the anterior side; serrula interior with about
26 blades and serrula exterior with about 45 biades.

Palps very elongate for the genus; surfaces markedly granu-
late, except for the tibial pedicel, the pedicel and base of the
chelal hand, and the chelal fingers; setae long and acuminate.

Maxillae with + or 5 terminal setae. Proportions of the pod-
omeres as shown in figure 1. Tactile setae of chela as shown in
figure 2; fixed finger of chela with a marginal row of 127 low,
rounded or truncate teeth which are closely contiguous ; mov-
able finger with 118 similar teeth.

Trochanter 2.5, femur 5.9, entire tibia 5.0, tibial pedicel
alone 5.2, chela 4.6, and hand 2.6 times as long as broad; moy-
able finger 1.14 times as long as hand.

Lees. Legs generally similar to but more elongate and slender
than those of other members of the genus. Each coxa of legs I
and II heavily sclerotic but not produced into a definite pro-
cess. Fourth leg with tactile setae on metatarsus 0.15 and on
telotarsus 0.59 the length of the segment from the proximal
end. Subterminal setae of all tarsi subequally bifurcated and
each ramus with several spinose processes. Each claw with a
small denticle on the dorsal side about one third the length of
the claw from the proximal end.

MerasureMENts. Body length 5.49 (in mm); carapace 2.05

rreatest breadth 1.50; abdomen

long, anterior breadth 1.20, g1

Cavernicolous Pseudoscorpion

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4 Postilla Peabody Museum No. ‘70

3.43 long, 2.08 broad. Chelicera 1.04 long by 0.53 broad, and
with movable finger 0.70 long; galea about 0.075 long. Palpal
trochanter 1.31 long by 0.53 broad; femur 2.92 by 0.50; tibia
2.87 by 0.58 including pedicel, which is 1.24 by 0.28; chela
without pedicel 3.83 by 0.84; hand 1.93 by 0.75; movable fin-
ger 2.19 by 0.29. Leg I: basifemur 1.27 by 0.27; telofemur 0.91
by 0.24; tibia 1.27 by 0.18; metatarsus 0.58 by 0.18; telotarsus
0.85 by 0.12. Leg IV: entire femur 2.15 long; basifemur 0.99
long by 0.42 broad; telofemur 1.23 by 0.40; tibia 2.28 by 0.23;
metatarsus 0.75 by 0.17; telotarsus 0.96 by 0.15.

Femaue, Unknown.

Triroxympnu: Similar to the adult male except that the ap-
pendages are much less attenuated and all parts are lighter
in color. Carapace only one-sixth longer than broad; epistome
and eyes as in adult; carapacal setae 23 (4-6).

Chelicera as in adult; six setae on palm; flagellum of 9 setae;
galea well developed as in adult, but trifid in the distal third,
and without terminal subdivisions.

Palpal podomeres not so attenuated as in adult, especially
the tibial pedicel which is only 2.6 times as long as its least
breadth: other ratios are: trochanter 2.1, femur 4.4, entire
tibia 3.5, chela 4.1 and hand 2.1 times as long as broad; mov-
able finger 1.05 times as long as hand; fixed finger with seven,
and movable finger with three, tactile setae: fixed finger with
95 and movable finger with 93 marginal teeth.

Legs as in adult but stouter. Fourth leg with tactile setae
on metatarsus 0.19 and on telotarsus 0.49 the length of the
segment from the proximal! end.

MeasvremMents. Body length 3.15 (in mm); carapace 1.32
long, anterior breadth 0.87, greatest breadth 1.15; abdomen
1.82 long by 1.28 broad. Chelicera 0.77 long by 0.39 broad,
movable finger 0.51 long; galea 0.068 long. Palpal trochanter
0.78 long by 0.37 broad; femur 1.62 by 0.37: tibia 1.53. by
0.44 including pedicel, which is 0.57 by 0.25; chela without
pedicel 2.46 by 0.61; hand 1.24 by 0.61; movable finger 1.3]
by 0.22. Leg I: basifemur 0.72 long by 0.19 broad; telofemur
0.54 by 0.17; tibia 0.69 by 0.12; metatarsus 0.37 by OA105

Nov. 5, 1962 = Cavernicolous Pseudoscorpion 5

telotarsus 0.43 by 0.10. Leg IV: entire femur 1.35 long; basi-
femur 0.63 long by 0.29 broad; telofemur 0.71 by 0.29; tibia
1.28 by 0.16; metatarsus 0.45 by 0.13; telotarsus 0.59 by 0.13.

Remarks. Although the eyes are not reduced and the derm
is heavily sclerotized and pigmented, M. grandis is certainly
specially modified for life in caves by its large size and great
attenuation of the appendages. In support of this contention
is the fact that the specimens were taken ‘ton formations” and
“on stalagmite” within the cave where they obviously were at
home. In proportions of the body and appendages it is rather
similar to M. cavernicola Vachon (from caves in Portugal)
which is undoubtedly a true troglobite. While reduction of the
eyes and pigment usually accompanies increase in body size and
attenuation of the appendages in adaptation of a pseudoscor-
pion to cavernicolous life, there is no reason to believe that all
these modifications must necessarily proceed together. It is rea-
sonable to suppose that M. grandis is exclusively troglobitic in
spite of the retention of eyes and pigmented derm, features
which have not proved disadvantageous and which have not
yet been lost by random mutation. Indeed, it is difficult to im-
agine that such a form as M, grandis could live successfully in
the epigean environment of a typical Microcreagris, where its
size and attenuation would put it at a definite disadvantage.

It is impossible to fit WM. grandis into the key given by Cham-
berlin (1962, p. 333) inasmuch as it has four eyes and an ex-
tremely slender tibial pedicel, a combination of characters
which is not provided for in couplet 1. There is no difficulty,
however, in separating it from other American species on the
basis of its size and proportions.

It seems appropriate to note here that Obisium cavicola
Packard from New Market Cave, Virginia, which Beier (1932)
and Hoff (1958) tentatively place in the genus Microcreagris,
does not in fact belong here. As will be shown in another place,
it is a member of the genus Chitrella.

6 Postilla Peabody Museum No. 70

REFERENCES

Beier, M. 1932. Pseudoscorpionidea I. Chthoniinea et Neobisiinea in Das
Tierreich, Berlin, vol. 57, p. 1-258.

Chamberlin, J. C. 1962. New and little-known false scorpions, principally
from caves, belonging to the families Chthoniidae and Neobisiidae
(Arachnida, Chelonethida). Bull. Amer. Mus. Nat. Hist., vol. 123, art.

6, p. 299-352.

Hoff, C. C. 1958. List of the pseudoscorpions of North America north of
Mexico. Amer. Mus. Novitates, no. 1875, p. 1-50.

Vachon, M. 1946. Description dune nouvelle espece de Pseudoscorpion
(Arachnide) habitant les grottes Portugaises; Microcreagris caver-
nicola. Bull. Mus. Nat. Hist. Nat., Paris, vol. 18, p. 333-336.

ee

SUSE ZE

[L§7

Pine

YALE PEABODY MUSEUM

or NatTurAL History

Number 71 November 14, 1962 New Haven, Conn.

SCOPELOGADUS (?) CAPISTRANENSIS,
A NEW FOSSIL MELAMPHAID (PISCES:
TELEOSTEI) FROM CAPISTRANO BEACH,

CALIFORNIA

ALFRED W. EBELING

Melamphaidae, a family of bathypelagic fishes that hitherto
has been known only from living material, comprises 5 genera
and about 33 species. Regan (1911), followed by Ebeling
(1962), referred it to the Stephanoberyciformes, an order of
peculiar spiny-rayed deep-sea fishes allied with the Berycifor-
mes, but having a hypertrophied open cephalic sensory canal
system lined with delicate bony ridges, usually a single trian-
gular supramayillary bone, often a regressed lateral line, no
orbitosphenoid bone, and possibly various other adaptations
to life in the deep sea. Ebeling (1962) presented a key to the
5 melamphaid genera: Melamphaes Giinther 1864, Sio Moss
1962, Scopelogadus Vaillant 1888, Poromitra Goode and Bean
1883, and Scopeloberyx Zugmayer, 1911. Ebeling and Weed
(in press) revised Scopelogadus, which contains three living
species, including two subspecies.

Scopelogadus, like other Melamphaidae, is mostly circum-
tropical at depths between 100 meters and the bottom. S.
beanti, however, is antitropical and inhabits both the temperate

2 Postilla Peabody Museum No. 71

North Atlantic and the region of the Subtropical Convergence,
which is an area of sinking of water masses at about 40°S lat.
(Ebeling and Weed, in press).

Very few fossils of bathypelagic fishes have been reported.
Only the Gonostomatidae, Paralepididae, and Myctophidae are
listed in Berg (1940) as other than Recent (“Miocene to Re-
cent’). To my knowledge, the only fossil melamphaids were
collected by Dr. Andreas B. Rechnitzer on May 2, 1956, from
Miocene shales along the sea cliff south of Capistrano Beach
in Orange County, California. Because counts and measure-
ments of these two small specimens generally agree with those
of Scopelogadus, they are provisionally placed in this genus
(Table 1).

The Capistrano Miocene locality consists mainly of finely
laminated diatomaceous shale and mudstone. From the included
Foraminifera fauna, Dr. M. N. Bramlette of the University of
California, Scripps Institution of Oceanography inferred that
the shale-mudstone deposits probably accumulated on the sca
floor below 100 fathoms during the Upper Miocene Age (Mil-
ler, 1951). Dr. Carl L. Hubbs (pers. comm.), also of Scripps,
added that although the lower parts contain algae (20 species
thus far discovered), a few fish (herring), and many fish scales,
the upper parts are bathypelagic deposits, in which have been
found, besides the melamphaids, fossils of the bathypelagic fish
Cyclothone and of the pelagic crustacean Pleuroncodes. Miller
(1951) described a new fossil species of petrel. Oceanodroma
hubbsi, which was also found near the bathypelagic site. More
recently, a grunion-like shallow water fish (Atherinidae) was
uncovered. As suggested by Dr. Hubbs, it would appear from
this stratification of faunas that the deposits originated in a
shallow basin, which gradually deepened as a result of either a
rise in sea level or a depression of the bottom.

Scopelogadus (7) capistranensis, new species

Fig. 1
The following description is mainly of the holotype. In-
formation from an impression of a second, smaller specimen
compliments data on numbers of pectoral and pelvic rays and

tranensl

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4 Postilla Peabody Museum No. 71

the position of the pelvic fin relative to that of the pectoral.
Additional characters are given in Table 1.

Body with greatest depth, at pelvic insertion, about 0.33
standard length. Caudal peduncle with length 2.5 times its
least depth. Distance from tip of snout to base of first anal ray
almost 0.7 standard length. Head large, 0.45 standard length ;
its depth about 0.7 its length; 3 or more weak spines faintly
visible at posteroventral angle of preopercle. Branchiostega!
rays at least 6.

Fig. 1. Scopelogadus (?) capistranensis, holotype YPM No. 3965, 38.7 mm
standard length.

Dorsal fin with about 13 rays (total); origin at middle of
body: distance from tip of snout to base of first ray equals
distance from this ray to base of caudal fin, which also equals
distance from tip of snout to base of pectoral fin. Anal fin
with 9 or 10 rays (total); origimates under fourth from last
dorsal ray. Pectoral fin with more than 12 or 13 rays, possibly
15. Pelvic fin with about 8 rays; inserts directly under pectoral.
Caudal fin with 19 principal rays.

Vertebrae on holotype 10 precaudal plus 14 caudal (the first
caudal vertebra overlies the first distinct haemal spine, the
urostyle is counted as one element) ; in smaller specimen about
24 to 26. The arch of the first haemal spine apparently lacks
the anteroventrally projecting spurs characteristic of some
species in Melamphaes and Scopeloberyz.

MATERIAL

ae ‘ : : 4
The types are two impressions of whole specimens, cata-
logued Yale University, Peabody Museum Paleontological Col-

Nov. 14, 1962 Scopelogadus (7) capistranensis 5

lections No. 3965. The larger, standard length 38.7 mn, is
selected as holotype. The vertebral column and most of the
fin rays are easily discernible on the holotype (Fig. 1). The
impression of the smaller specimen is much fainter and there-
fore difficult to interpret. On the holotype are impressions of
various head bones, including the line of fusion of the hyo-
mandibular with the front of the preopercle, sections of the
opercular series, parts of the jaws, branchiostegal rays, fin
supports, and the caudal skeleton.

DERIVATION OF NAME

The species name capistranensis refers to the locality of dis-
covery of the fossils.

IDENTIFICATION WITH Scopelogadus

The fact that Scopelogadus (?) capistranensis has 19 prin-
cipal caudal rays and thoracic pelvic fins places it with the
berycoid-like fishes. Its general shape, positioning of fins, num-
bers of fin rays, etc. further refer it to the Melamphaidae. A
definite identification of the fossils with Scopelogadus was im-
possible, although the specimens are provisionally referred to
this genus by virtue of comparisons with each melamphaid
genus. In Table 1 are listed the only characters measureable on
the fossils, along with ranges of values for all five melamphaid
genera. At the bottom of each “genus column” are: first, the
total number of characters whose ranges exclude values for
S. capistranensis and second, this number of disagreements for
meristic characters only. Both Scopelogadus and Poromitra
had only two disagreements. Even though the next lowest pro-
portion of disagreements, 4/18, of Scopeloberyax is not signifi-
cantly different from 2/18 X?=0.20 with one d.f., 0.75>p>
0.45), the fossils agree with Scopelogadus in both meristic
counts and general overall shape, which associate them most
strongly with this genus.

ACKNOWLEDGMENTS

I am indebted to Dr. Andreas B. Rechnitzer, formerly of the
Scripps Institution of Oceanography, La Jolla and the Naval

6 Postilla Peabody Museum No. 71

Electronics Laboratory, San Diego for donating the fossils
and to Dr. Carl L. Hubbs for much useful information on the
constitution of the fossil beds.

Lrrerature Crrep

Berg, L. S., 1940. Classification of fishes, both recent and fossil. Trav.
Instit. Zool. Acad. Sci. U.S.S.R., 5: 1-517.

Ebeling, A. W., 1962. Melamphaidae I, Systematics and zoogeography of
the species in the bathypelagic fish genus Melamphaes Giinther. Dana-
Rep. No. 58. 164 p.

Ebeling, A. W. and W. H. Weed. Melamphaidae III. Sytematics and
distribution of the species in the bathypelagic fish genus Scopelogadus
Vaillant. Dana-Rep. (in press).

Goode, G. B. and T. H. Bean, 1895. Oceanic ichthyology.... (Smithsonian
Contrib. Know. nos. 981, 982). Washington. Smithsonian Institution. 555
p. Atlas of plates. .

Giinther, A., 1864. Addenda. Jn Catalogue of the Physostomi, containing
the families Siluridae, Characinidae, Haplochitonidae, Sternoptychidae,
Scopelidae, Stomiatidae in the collection of the British Museum. Cat.
Fish. British Mus., 5: 1-455. London.

Miller, L., 1951. A Miocene petrel from California. The Condor, 53: 78-80.

Moss, S. A., 1962. Melamphaidae II. A new melamphaid genus, Sio, with a
redescription of Sio nordenskjolditi (Loénnberg). Dana-Rep, no. 56,
10 p.

Regan, C. 'T., 1911. The anatomy and classification of the teleostean fishes
of the orders Berycomorphi and Xenoberyces. Ann. Mag. Nat. Hist.
@Sersne) sieeel—o-

Vaillant, L., 1888. Expéditions scientifiques du Travailleur et due 'alis-
man pendant les années 1880, 1881, 1882, 1883....Poissons. Paris, G.
Masson. 406 p.

Zugmayer, E., 1911. Diagnoses de poissons nouveaus provenant des cam-
pagnes du yacht *Princesse-Alice” (1901 a 1910). Bull. Instit. Oceanogr.
Monaco, (193): 1-14.

GOS, 73

LE)

pila

YALE PEABODY MUSEUM

oF Natura History

Number 72 December 20, 1962 New Haven, Conn.

A MEROITIC TOMB INSCRIPTION
FROM TOSHKA WEST

Bruce G. TRIGGER

The Meroitic inscription presented herewith is the only funer-
ary inscription to have been discovered by the University Mu-
seum of the University of Pennsylvania—Peabody Museum of
Yale University Expedition to Egypt in the large cemetery
(called TW-A) just north of the winter Post Boat station at
Toshka West. The cemetery is in the north part of Toshka op-
posite Gebel Agg, and lies between the river and the hamlet of
Duki Dawur. It was found to contain several hundred graves
of the late Meroitic, X-group, and Nubian Christian periods.
No reference to this cemetery is found in Emery’s and Kirwan’s
(1935) report on their archaeological survey of this region. A
description of the cemetery and of the work done there in 1961
and 1962 can be found in two preliminary reports on the work
of the expedition written by its director, Professor William K.
Simpson of Yale University, These have appeared in the II-
lustrated London News (1961) and in Eapedition, the bulletin
of the University Museum of the University of Pennsylvania
(1962). The inscription, which was found during the 1962 field
season, is illustrated on page 38 of the second article and on the
cover of the issue.

2 Postilla Peabody Museum No. 72

The stela, on which the inscription was carved, was found
face up and reused among the blocks in the largely denuded
superstructure of a Christian grave numbered TW-A 198. The
superstructure was of the type found by Monneret de Villard
(1985: 132-141) at er-Rammal. Without much question the
stela had been plundered from one of the Meroitic graves
nearby for use as a building block. Since none of the Meroitic
superstructures were preserved, however, it is impossible to dis-
cover from which grave it came. Funerary inscriptions were
generally written either on offering tables or on stelae, more
commonly the former. The stelae have a variety of shapes and
apparently were set up in little shrines which projected from
the front walls of the Meroitic superstructures (Griffith, 1911:
29). Sometimes representations of the deceased were painted or
carved on these stelae, but more often there was only a text.

The Toshka West inscription is engraved on a block of grey,
fine to medium grained Nubian sandstone. It has been given the
Pennsylvania—Yale expedition number TW-A 198.2, and was
assigned to the expedition for a subsequent division of finds be-
tween the University Museum and the Peabody Museum. The
block is roughly rectangular, though the left side tapers slightly
toward the base. Its measurements are: height 52 cm, width
40-45 cm, thickness 6.5-11 cm. The inscribed surface undulates
slightly but is smooth except for some ridges in the lower right
hand corner. These antedate the inscription. There is no evi-
dence that this face was artificially worked, and it seems to be
merely the natural, sand-scoured surface of a local rock expo-
sure. This contrasts sharply with the carefully leveled surfaces
of the fragments of two funerary stelae found by the expedi-
tion at Arminna West. The block was apparently quarried by
scoring it along the sides and top edge and then prying it
horizontally along the lines of natural bedding. The back is
domed where large flakes were knocked marginally from the top
and sides in order to lessen its weight. The sides were then
smoothed near the upper face though traces of the scoring and
prying still remain visible underneath. The upper edge, which
suffered accidental chipping at a later time, was probably
meant to be convex. Below the inscription the face of the block
was quite rough, and no attempt was made to square this sec-

Dec. 20, 1962 A Meroitic Tomb Inscription 3

tion. Presumably when the stela had been set up this part was
covered over with mud plaster.

The rows of letters, which read from right to left, are sep-
arated by incised lines 2 mm wide and 2.5-4.5 cm apart. The
ninth line from the top appears to have been engraved from
both sides. The letters, which average about 1.5 cm in height
(omitting tails) are engraved to a depth of 1 to 1.5 mm, about
the same depth as the lines. The lines and the letters still have
traces of the red paint used to accent the text. The whole sur-
face of the stone, especially towards the bottom, is reddened,
the coloration being clearly artificial and distinct from the
natural pink color found in some of the Nubian sandstones.
This no doubt represents the working out of pigment from the
letters rather than a deliberate coloring of the whole surface
of the block. A pronounced red line under the final row of let-
ters, where presumably the stone was embedded in the super-
structure, marks the termination of this reddening. This sug-
gests that the pigment had been weathered while the stone was
still in site.

The similarity of some Meroitic letters, particulary e and 1,
and m, 8, and h creates a danger of error in transliteration,
although a comparison of the form of letters found in recog-
nizable phrases and words clears up most cases of uncertainty.
The greatest margin of doubt must remain in the translitera-
tion of personal names. Meroitic tombstones do not carry
dates. Nevertheless, changes in the style of the letters may offer
clues as to the general date of the inscription. Griffith (1911:
17-21) was able to distinguish three stages in the development
of cursive Meroitic, and Hintze (1959), working on materials
from Meroé, has recently published a more detailed paleo-
graphic table which gives the forms of letters associated with
various reign dates. Although the study of Meroitic pale-
ography is not yet sufficiently advanced to provide a real basis
for dating inscriptions, the conclusions resulting from a com-
parison of this inscription with the forms given by Griffith and
Hintze are of some interest. The form of the letters showing
the most marked changes through time (Griffith 1911: 11) all
belong to the later stages of the writing. In general, the letters
most closely resemble the forms found in Griffith’s columns 14

4 Postilla Peabody Museum No. 72

and 16. Column 14 belongs to the Transitional phase and is
dated 25 B.C. to A.D. 250; 16 is Late and is dated A.D. 250 to
400. The rather equal distribution of resemblances suggests a
date close to A.D. 250. Hintze’s system provides a greater
variety of forms, so much so that different forms of the same
letter on our stone can often be placed in several columns. The
temporal distributions of these placements are not, however,
such as to be damaging to Hintze’s system. The majority of

forms are to be found in the columns dated A.D. 170-350 and
A.D. 246-266.This dating seems in accord with the conclusions
reached using Griffith’s system. Two of Griffith’s (1911: 33;
1912: 57) paleographically dated inscriptions which also bear
the same introductory formula as does this one are dated to

Dec. 20, 1962 A Meroitic Tomb Inscription 5

the late period. Although a detailed report on the material
from the Toshka cemetery has not yet been prepared, a date in
the third century A.D. would not seem out of keeping for many
of the Meroitic graves there.

—
ee

eras NS :
Fr AF FBECI3 2 YE SII RYVKS $323/3
SSIPYS UB BSI CLBII ATE EI AS

re SSS
MBAS) BESTE PASS 77
153 BSS BX S280 Hs be YS USBI S72 E_
IDS-IPY Rik SE LS Ald: 02 15) His
EEE ner aa ps
O° ULE GSE ae
= LEE EE MY" YS SP,
as WH UES oy UTS UF /CSUYE TS
‘Sy = “
OO LEE SE Tg
SSUES s IS hitcig =
horas See SABES LGB EIR.
ZL Sees ee
A/S PUR Foo

TWA 198.2
occu wus-aT 1962

TRANSLITERATION AND COMMENTARY

The following transliteration of the text follows Hintze’s
system. The arrangement of lines is that of the original. Words
which are carried over from one line to the next are indicated
by a hyphen. The colon is used to represent the Meroitic word
divider. The text is divided into sections, each of which is de-

6 Postilla Peabody Museum No. 72

noted by a letter of the alphabet. These sections will be dis-
cussed individually in the commentary.

(a) wés : wethyimgh : séri : wetrri

(b) gé : mli : wés qéwi : (c) pélmés : adbli-
s atkitnideye : kditeléwi :

(d) pélmés : adblis : adileméme :
kditeléwi : (e) qntkleb : kditeleé -

EE ee

wi : (f) hrphiphrste : pteremé

=<

tiye : kditeléwi : (g) apéte :a-
remelis : aqéhléye : yetmde -

: léwi : (h) apéte arémelis : htpiye
10. kditeléwt : (1) apétekdiyt : tki-
vi teléwi : (3) pelmésleb : apétele -
12. b : hrphleb : kditebetéwi :

13. (k) atémsb : h/S-i-h/s- es

Like all funerary texts this one consists of three sections,
1) an invocation usually addressed to Isis and Osiris, 2) the
name and description of the person commemorated, and 3) a
terminal formula or formulae, sometimes called a benediction.

Section (a) constitutes the introductory invocation. It is
longer than the usual wé3t : séreyi (Isis, Osiris), since each of
these god’s names introduces another phrase terminating in lt
or 7, which are believed to be vocative particles. The interven-
ing phrases are apparently adjectives describing Isis and
Osiris but their meaning is unknown (Griffith 1911: 34). There
is a similar invocation on a stone of unknown provenience now
in the Cairo Museum (Griffith 1912: 57). Another example oc-
curs on funerary inscription 76 from Karanog, although here
the second # is written né and one stroke has been left out of
the fifth letter of the second word. The first substitution is quite
common, however, (Griffith 1911: 14) and in writing yi one
stroke is commonly omitted (ibid., 33). The alternative read-

Dec. 20, 1962 A Meroitic Tomb Inscription 7
ing wetnééinégeli is impossible (tbid., 13, section 3). A variant
also occurs in inscription 4 from Faras (Griffith 1922: 570)
in the second half of which mklh [Great God? (Griffith 1912:
57) ] replaces the name of Osiris.

Section (b). This section gives the name of the deceased,
which is referred to in the literature as the A name. As in some
other inscriptions the name is introduced by the expression
gé, which Griffith renders tentatively as ‘the honorable.” The
expression géwi regularly follows the name of the deceased and
it is not often separated from it by a word divider. Hence the
name is Mli-wés, the prefix mli indicating that it is a woman’s
name (Griffith 1911: 55). This prefix is often separated from
the rest of the name by a word divider.

Following this we find eight parallel phrases listing people
who stand in some sort of relationship to the deceased. In their
fullest form these phrases consist of a title, the person’s name,
and a word describing the relationship between him and the
dead person. Griffith (1911: 88) called this last word the
descriptive phrase. Grammatically it stands in apposition to
the name of the deceased. The terminal particle léwi (which
sometimes replaces géwi in the preceding phrase may either
represent a copula or be for emphasis (tbid., 35). Two descrip-
tive phrases which very frequently occur in funerary inscrip-
tions are absent here. These are tedheléwi and terikeléwi. The
former follows the name of the mother (the B word) of the de-
ceased, and the latter that of his father (the C word) and they
seem to give a reading: A borne by the woman B, begotten of
the man C. On the other hand six of the sections in this inserip-
tion contain the descriptive phrase kditeléwi which is unrecorded
in Griffith’s word lists. Griffith (1911:38, 39) lists two forms
beginning with kdi (which is believed to mean “woman’’), kdis-
betéwi and kditéwi, both of which are associated with feminine
A names. Taking the forms which are known, it would seem to
be composed of kdi (woman?) and teléwi (the locative particle
plus the copula? léwi). Teléwi, however, seems to be added
only to place names (Griffith 1911: 23, 40). If, as conceivably
might be the case, te was written here as a variant for té, the
genitive suffix, then this word might be closely related to kditéwi
and be read “a woman? of A.” But this sort of comparison,

8 Postilla Peabody Museum No. 72

which has constituted the main approach to the study of
Meroitic grammar so far, tends to provide little in the way of
satisfactory solutions to such problems in the absence of a
more fundamental break-through in the study of the language.

Sections (c) and (d). Despite the lack of a divider, the ini-
tial s in line 3 undoubtedly belongs with the final word on the
preceding line. Pélmés, a variant of plmeés or pelmés, is derived
from the Egyptian p» imy-r_ ms meaning strategos or com-
mander. The form pélmés adblis does not appear in the word
lists but seems to be a variant of pelmés adblité, with a genitive
particle s substituted for the alternative (?) genitive particle
té. The expression is translated “‘commander of the land” or as
Zyhlarz (1956: 33) has suggested “commander of the deserts.”
The closely related term “commander of the water” is written
pelmés atélis and pelmés atélit (Griffith 1912:64). The last
portion of the name Adileméme is the same as that of the name
Arleméme occurring in inscription 24 from Faras (Griffith
1922 :583).

Section (e). gntkleb may be a person’s name or it may con-
sist of the root qntk plus the plural particle leb. Neither form
could be located elsewhere. The leb ending would seem to make
a personal name less likely.

Section (f). Arphw and variants, a civil title believed by
Griffith (1922: 567) to refer to the governor or leading person
of a community, in rank inferior to a pagar and a pesaté. phrs
(Pachoris) is the old name of Faras; te is the locative particle.
Hence ‘‘a civil official in Pachoris.”

Sections (g) and (h). apéte, from the Egyptian ipwty, mes-
senger or envoy; arémelis, “of (the) Rome.” Presumably the
so-called s-genitive here has the force of a dative, as Griffith
(1911: 72) himself has suggested. The considerable number of
people holding this title would suggest that it is applied to
individuals having dealings with Roman Egypt, perhaps to
government sponsored traders. Hence the title may mean some-
thing like “agent to the Romans.” yetmdeléwi is a common des-
cripive phrase rendered as “cousin of” or “kinsman of” (Grif-
fith 1912: 65).

Section (i). The first word seems to consist of apéte plus kzt
which is sometimes added to a noun to give it a feminine form.

Dec. 20, 1962 A Meroitic Tomb Inscription 3

Hence it would appear to read “a female ambassador.” Here
b]
however, it may well be used as a person’s name as Griffith
’ J
(1911:59) shows a variant to be used elsewhere. The terminal
yi is unaccounted for. thiteléwt seems to be another unlisted
descriptive phrase.

Section (j). The first three words have been made plural by
the addition of the suffix leb. This list of titles may either be
resumptive or refer to Mli-wés’s relatives (?) in general. The
descriptive phrase used here would seem to be a plural form,
since it contains a b-infix. But such a form has occurred only
once before in an inscription commemorating a single person.
This is in inscription 99 from Karanog, and Griffith (1911: 70)
believed it to have been written inadvertently. Moreover the
téwi-ending normally becomes tebkwi in the plural. Hence a
completely different word or form may be involved. The ending
betéwi occurs in inscriptions 89 and 125 from Karanog (See
Griffith 1911: 40).

Section (k). The form até introduces the terminal formula
Type A, the commonest and invariably the first of a number of
such titles. até is rarely separated by a divider from the follow-
ing letters, which in almost all the variants of this formula are
ms. msb is found in Type Ae (Griffith 1911: 46) but here any
resemblance between our inscription and any of the varieties
listed by Griffith ceases. The second word is very carelessly
written and may be read several ways. It does not bear resem-
blance to any other of the types of terminal formulae, except
perhaps a crude and unconvincing one to type F (Griffith 1911:
52), and hence it would appear that we have here only one for-
mula. Griffith (ibid., 46) suggests that the general meaning of
this phrase is “abundant water may you drink,” this wish being
made on behalf of the deceased for his afterlife. The formation
of the various forms is exceedingly speculative and no purpose
could be served in discussing them here

CONCLUSIONS

As can easily be seen in the preceding discussion, there exists
at present only a rudimentary understanding of the Meroitic
language. The sound values of the letters have been generally

10 Postilla Peabody Museum Nowra

established, names can be read, and certain loan words, mostly
from Egyptian, have been recognized. The meaning of a small
number of native words and some of the basic elements of gram-
mar have also been established. Griffith’s lexicons, which sys-
tematically arrange the known forms, should someday provide
a substantial base for further work. But until either bilingual
texts are discovered or the linguistic affiiations of Meroitic
are worked out so that known cognate languages can provide
a sound basis for systematic investigation, the hope of much
progress in the understanding of Meroitic appears dim indeed.
Until such a time the “translation” of even relatively simple
and well-studied formulae such as appear on tomb inscriptions
must remain an identification of known words and grammatical
forms eked out with many others of vague or uncertain mean-
ing as well as with guesses and blanks. With these limitations in
mind we offer the following translation of the Toshka inscrip-
tion:

(a) O (unknown adjective) Isis! O (unknown adjective)
Osiris!
(b) (the honorable) Mli-wés (is here commemorated)

(c) a kinswoman of the commander of the deserts
(land?) whose name is Atkitnideye

(d) a kinswoman of the commander of the deserts
Adileméme

(e) a kinswoman of Qntkleb [or of qntks |
(f) a kinswoman of the governor [or civil
official] of Faras Pteremétiye

(g) a kin [or a cousin] of the envoy to the
Romans Aqéhléye

(h) a kinswoman of the envoy to the Romans

Ht piye

(1) a relation (?) of Apétekdiyi [or of a female
ambassador |,

(j) a relation (?) of commanders, envoys, and gover-
mOrs: ((723)

(k) (Abundant water may you drink in the afterlife.)

Dec. 20, 1962 A Meroitic Tomb Inscription 11

Despite plundering and reuse, this Meroitic funerary in-
scription from Toshka West has been preserved virtually in-
tact. It appears to date from the third century and commemo-
rates a woman whose name was Mli-wés. She has no specific ti-
tles, although this is often the case when women are commemo-
rated. Her name is followed by a list of the names and titles of
a number of individuals whom we presume to be her relatives. In
spite of orthographic peculiarities all the titles which are listed
are known, with the possible exception of qntk, if it is a title.
Although close parallels exist for the name apétekdiyi, we are
not sure that it is here being used as a personal name. A search
of the literature has failed to turn up occurrences of the other
personal names in the inscriptions from the Meroitic cemeteries
at Karanog, Shablul (Griffith 1911), and Faras (abid., 1922).
The inscription is somewhat unusual in that the formulae giv-
ing the names of the parents of the deceased are lacking. Also
several new descriptive phrases are found here.

This text is of interest since it one of the very few Meroitic
funerary texts to have come from the Toshka-Arminna region,
and the only one preserved intact. Junker (1925: 104) found a
few fragments of broken funerary inscriptions in the Meroitic
cemetery at Arminna East, and a few more have been found by
the Pennsylvania-Yale expedition at the village site at Ar-
minna West. These are as yet unpublished. Apart from a few
graffiti scratched on rocks and a number of inscribed potsherds
from both Toshka and Arminna, these constitute the entire cor-
pus of Meroitic writing from this section of Lower Nubia.

ACKNOWLEDGMENTS

The 1962 excavations of the Peabody Museum of Yale Uni-
versity and the University Museum of the University of Penn-
sylvania were directed by Professor William K. Simpson of
Yale University and were supported by the E. B. Coxe, Jr.
Fund of the University Museum, a grant from the Bollingen
Foundation, established by Mr. Paul Mellon, to Yale Univer-
sity, and by a grant of the Department of State through Pub-
lic Law 480. The author’s participation in this year’s work
was facilitated by a grant of the Augusta Hazard Fund of

12 Postilla Peabody Museum No. 72

Yale University, and the major part of the work on this in-
scription was accomplished while he was the recipient of a Pre-
doctoral Fellowship of the Canada Council.

BrsiioGRAPHY

Emery, W.B. and L.P. Kirwan, 1935. The Excavations and Survey between
Wadi es-Sebua and Adindan. Cairo, Government Press.

Griffith, F. Ll., 1911. Karanog. The Meroitic Inscriptions of Shablul and
Karanog. Philadelphia, University Museum.

, 1912. Meroitic Inscriptions Part II. Napata to Philae and
Miscellaneous. London, Egypt Exploration Fund.

, 1922. Meroitic Funerary Inscriptions from Faras, Nubia. Re-
cueil d’études égyptologiques dédiées 4 la mémoire de Jean-Francois
Champollion. Paris, Librairie Champion.

Hintze, F., 1959. Studien zur Meroitischen Chronologie und zu den Opfer-

tafeln aus den Pyramiden von Meroe. Abhandlungen der Deutschen
Akademie der Wissenschaften zu Berlin. Berlin.

Junker, H., 1925. Bericht ueber die Grabungen der Akademie der Wis-
senschaften in Wien auf den Friedhoefen von Ermenne (Nubien).
Akademie der Wissenschaften in Wien, Philosophisch-historische
Klasse, Denkschr. 67, Band 1.

Monneret de Villard, U., 1935. Nubia Medioevale, vol. I, Cairo, Imprimerie
de l'Institut Frangais.

Simpson, W. K., 1961. Discoveries, From Old Kingdom to Coptic Times, at
Toshka West. The Illustrated London News, vol, 239 no. 6363, p.
94-95.

, 1962. Nubia, 1962 Excavations at Toshka and Arminna. Ex-
pedition, 4. (4): 36-46. Philadelphia.

Vycichl, W., 1958. The Present State of Meroitic Studies. Kush, 6:74-81.
Zyhlarz, E., 1956. Die Fiktion der Kuschitischen Voelker. Kush, 4:19-33,

A ae

+4 Gs

Josie

YALE PEABODY MUSEUM

oF Naturau History

Number 73 December 20, 1962 New Haven, Conn.

NEW BIRDS FROM PALAWAN
AND
CULION ISLANDS, PHILIPPINES

S. Ditton RIPLEY sanp D. S. Rasor

INTRODUCTION

BY D. S. Raror

The Palawan Expedition of 1962 was sponsored jointly by
the Yale Peabody Museum, the Entomology Section of the
U. S. Army Medical Research and Development Command of
the Office of the Surgeon General, under the auspices of the
Bernice P. Bishop Museum, and by Silliman University at
Dumaguete City, P. I. Collections were made on Palawan
Island March 26 until May 18, 1962, working in southern
Palawan from March 26 until April 21, 1962, and in northern
Palawan from April 25 until May 18, 1962.

The main collecting localities in southern Palawan included
the forested areas in the vicinity of the municipality of Brookes
Point, from the sea coast up to about 6,700 feet to the top of
the main peak of Mount Mantalingajan. Collecting camps
were established at Macagua (sea level to 500 feet), Pinigisan
(2,100-2,500 feet), Magtaguimbong (3,600-4,350 feet), and

os Postilla Yale Peabody Museum No. Za

on the ridge and top of the peak of Mount Mantalingajan
itself (5,000-6,700 feet).

The collecting localities in northern Palawan included Sitio
Malabusog (sea level to 500 feet) of Barrio 'Tinitian, munici-
pality of Roxas, and Sitio Tarabanan (sea level to 1,000 feet)
of Barrio Concepcion, municipality of Puerto Princesa.

A small collecting party worked on Balabac Island from
April 17 until May 2, 1962, and rejoined the main body of
the expedition party in northern Palawan on May 5, 1962.

Mr. Max Thompson, Research Fellow of the Bernice P.
Bishop Museum, who was a member of the expedition party,
headed a small group that carried on additional collecting for
the Bishop Museum, on Busuanga and Culion Islands of the Cal-
amianes Group, north of Palawan, from May 20 until June
20, 1962, after the main party had already left for home. His
collections of 141 birds from both islands were included in our
study.

It is interesting to note that six months earlier, in August
and September, 1961, a small collecting party of the Danish
**Noona Dan” Expedition, together with two Filipino collectors
from the Philippine National Museum, worked in the same
collecting localities where we collected later on, with the ex-
ception of the ridge and top of the main peak of Mount Man-
talingajan itself, which they failed to reach. Our collecting
party made camps on the very camping sites of the Danish
Expedition collectors in the lower elevations of the Mount
Mantalingajan Range and even employed the same head moun-
tain guide for the work in the highlands of Mount Mantaling-

ajan.

Salomonsen (1961, Dansk. Ornith. Foren. Tiddsskr., vol.
55, p. 219-221; 1962, vol. 56, p. 129-134) reported and de-
scribed one new species of tree-babbler, Stachyris hypogram-
mica, and two new subspecies, a mountain tailor-bird, Ortho-
tomus cucullatus viridicollis and a mountain” leaf-warbler,
Phylloscopus trivirgatus peterseni, from the collections made

by the “Noona Dan” collectors in southern Palawan.

Dec. 20, 1962 New Birds from Palawan, Culion Islands 33

Salomonsen reported six forms of true mountain birds which
were taken by the “Noona Dan” collectors, which included

the following:

1. Zosterops montana

2. Muscicapa westermanni
3. Setcercus montis

4. Phylloscopus trivirgatus
5. Orthotomus cucullatus

6. Stachyris hypogrammica

Our Palawan Expedition Party secured very good series of
most of the above mentioned mountain birds, except Setcercus
montis, of which only 5 specimens were taken by us. In
addition, two more true mountain birds were collected by our
party, increasing the list of true mountain birds from Palawan
to 8. These forms are Brachypteryx montana and Cettia Mon-
tana.

We are reporting six new subspecies of birds from our re-
cent collections on Palawan, most of them from the highlands
of southern Palawan in the Mount Mantalingajan_ local-
ities, and from the Calamians. It is very possible that the
highlands of northern Palawan, notably in the localities of
Cleopatra’s Needle, the high peak which dominates this area
may still yield additional forms distinct from those that we
have collected in the Mount Mantalingajan localities of south-
ern Palawan. The highlands of northern Palawan remain as
still another largely unworked locality.

We are indebted to the authorities of the American Museum
of Natural History and of the Chicago Natural History
Museum for the use of some of the comparative materials in
our present study.

D. S. Rabor acknowledges his indebtedness to the Frank M.
Chapman Memorial Fund of the American Museum of Nat-
ural History, to the Entomology Section of the U. 5. Army
Medical Research and Development command of the Office of
the Surgeon General, and to the Yale Peabody Museum, for

the research grants, which enabled him to come to the United

4 Postilla Yale Peabody Museum No. 73

States to study and prepare the report on the present bird
collections from Palawan, Balabac, and the Calamian Islands
of Culion and Busuanga, in collaboration with Dr. Ripley.

NrEw SUBSPECIES FROM PALAWAN
BY

S. Ditton RieLey anp D. S. Razor

Ninox seutulata palawanensis, subsp. nov.

Tyre: Yale Peabody Museum No. 73202 from Tarabanan,
Concepcion, Puerto Princesa, Palawan Island, Philippines, sea
level to 1,000 feet elevation. Adult male collected May 8, 1962
by D. S. Rabor.

Diacnosis: Similar to N. sc. randi of most of the larger
Philippine islands in the reddish chocolate-brown upper- and
underparts, but decidedly smaller. Slightly smaller than N.
sc. totogo of Botel Tabago but lighter brown on upperparts
with decidedly more reddish wash; light parts on the streaked
individual feathers of underparts purer white with much less
buff, and brown streaks darker: blackish tail bands similar in
width, number and location on tail feathers, but brown bands
in palawanensis tinged more ashy. Closest to Ninow scutulata
borneensis in size, but upperparts slightly darker brown with
more red wash; blackish tail bands much narrower, more in

number and darker, and the brown bands tinged more ashy.

MrasurEMENts: N. sc totogo from A’Oshima, Loo Choo
Islands, and Okinawa: Wing 8 (2) 214, 210, 2 (5) 204-
212.5 (av. 208.5); tail ¢ (2) 119, 115, 2 (6) 1115-118" Can:
115.6); culmen from base 6 (2) 24.5, 23.5, 2 (4) 238.5-25
(av. 24.5):;tarsus ¢ (2) 26, 24.5, 2 (5) 25-26.5 (av. 26 mm):
N. sc. borneensis from Borneo: Wing ¢ (4) 176-184.5 (av.
179.6), 2 (2) 182.5, 176, sex ? (4) 183-187.5 (av. 185.6)
tail ¢ (4) 98-102 (av. 99.5), 2 (2) 96.5, 96, sex ? (4) 94-
104 (av. 100.1); culmen from base 6 (4) 21.5-22.5 (av. 22.1),
@ (2) 22, 22.5, sex ? (4) 215-24 (av. 22:2); tarsus® 6° (49)

Dec. 20,1962 New Birds from Palawan, Culion Islands ix

23-25 (av. 24.4), 9 (2) 24.5, 23.5, sex ? (4) 22.5-24 (av.
23.6 mm). N. sc. palawanensis from Palawan Island. Wing 4
195.5; tail 108; culmen from base 23.5, tarsus 25 mm.

Rance: Palawan Island.

Remarks: The three races totogo, borneensis and palwanen-
sis are lightly differentiated from one another, based on plum-
age coloration. In size, however, palawanensis is intermediate
between totogo and borneensis.

Pitta erythrogaster thompsoni, subsp. nov.

Tyre: Yale Peabody Museum No. 73203 from 6.5 km south-
west Culion, Culion Island, Philippines. Adult male collected
June 7, 1962 by Max C. Thompson. Orginial field no. BBM-
PI 5290, Bernice P. Bishop Museum.

Diacnosis: Closest to P. e. propinqua of Palawan and Bal-
abac Islands but back, rump, tail and scapulars pale blue, not
cobalt. Differs from P. e. erythrogaster in the much paler blue
of back, rump, tail and scapulars, and in the much narrower
dull green band on mantle, as in propinqua.

MeEaAsurEMENTs: P, e. propinqua from Palawan: Wing 4
(3) 96-100 (av. 98); tail ¢ (3) 34-86.5 (av. 34.8); culmen
from base 6 (3) 13.5-14.5 (av. 14); tarsus 6 (8) 32-33.5
(av. 32.8 mm); weight 6 (8) 50.7-53.4 (av. 51.2 grams).

P. e. erythrogaster from Luzon: Wing ¢ (4) 96.5-99 (av.
97.5), 2 99.5; tail 6 (4) 32-37.5 (av. 35.1), 2 34.5; culmen
from base 6 (2) 23.5, 23:5, 2 23; tarsus ¢ (4) 33.5-35.5
(av. 34.2), 2 35 mm; weight 6 (4) 56.2-69 (av. 63.3), °
95.2 g.

P. e. erythrogaster from Mindanao: Wing ¢6 (2) 98.5,
HOOD. 2 (2) 95.5, 98; tail ¢ (2) 3615, 33.5, 2 (2) 32, 34;
culmen from base 6 (2) 23.5, 24, 2 22; tarsus 6 (2) 35.5,
35.5, 2 (2) 34, 34.5 mm.

6 Postilla Yale Peabody Museum No. 73

P, e. erythrogaster from Negros: Wing 46 (3) 98-100.5
(av. 99.5), 2 99;tail ¢ (3) 38-85.5 (av. 34.8), 2 32; culmen
from base 6 (3) 22-23.5 (22.6), 2 23; tarsus ¢ (3) 33-33.5
(av. 33.3), 2 33.5 mm.

P. e. erythrogaster from Samar: Wing (sex?) 94.5; tail
35.5; tarsus 33 mm.

P. e. erythrogaster from Bongao: Wing ¢ 95; tail 36.5;
e e te) S
culmen from base 22; tarsus 35 mm.

P.e. thompsoni from Culion: Wing é (2) 94.5, 94; tail 35,

36; culmen from base 22.5, 23.5; tarsus 33, 34.5 mm.
Rance: Culion Island.

Remarks: The Negros population of P. e. erythrogaster
comes closest to P. e. thompsoni in the intensity of the blue
color on the back, rump, tail and scapulars, but the Culion
form is still paler on these parts. Also, the dull green band on
the mantle in the Negros birds is much wider as is character-
istic of erythrogaster.

We take great pleasure in naming this form after Mr. Max
C. Thompson, Research Fellow of the Bernice P. Bishop
Museum, the young collector who was a great help in the field
work in Palawan, Balabac, and the Calamians.

Brachypteryx montana sillimani, subsp. nov.

Tyre: Yale Peabody Museum No. 73204, from Magtaguim-
bong, Mount Mantalingajan, 3,600-4,350 feet, Palawan Is-
land, Philippines. Adult male collected April 11, 1962, by
D. S. Rabor.

Diagnosis: Closest to B. m. poliogyno of northern (north-
western) Luzon and Mindoro, and B. m. andersoni of southern
Luzon, but differs in the following characteristics: a) Male
averages much brighter and richer slate-blue upper- and under-
parts, with much less wash of gray on abdomen; b) female
with much duller rusty brown undertail coverts: c) longer

culmen.

Dec. 20,1962 New Birds from Palawan, Culicn Islands 74

MerasvureMENts: B. m. poliogyna from northern Luzon:
Wing, ¢ (8), 65-68.5 (av. 66.6); 2 (2) 68.5, 62.5; tail, ¢
(3) 45.5-47 (av. 46.3), 2 (2) 46, 40; culmen from base, ¢
(3) 15.5; 2 (2) 16.5, 15.5; culmen from anterior margin of
nostril d (3) S.9, 2 (2))9,8-o5 tarsus, 6 (3). 26-27.5 (26.5),
272) 28, 27.

B.m. poliogyna from Mindoro: Wing, 6 (5) 65.5-67.5 (av.
66.7), 2 (8) 60.5-64 (av. 62.3); tail, 6 (5) 45-46.5 (av.
46), 2 (3) 41-43 (av. 42); culmen from base, 6 (5) 15.5-
17 (av. 16.2), 2 (3) 15.5-17 (av. 16.8) ; culmen from anterior
marci of nostril, 6 (5) 8.5-9 (av. 8.9), 2 (8) 8.5-9.5 (9);
tarsus, 6 (5) 28-29.5 (av. 28:5), 2 (3) 26-29 (av. 27-6).

B. m. andersoni from southern Luzon: Wing, ¢ (2) 71, 66,
O(2)63:55 tail, ¢ (2) 45.5,°47, 2 (2), 42.5, 43; culmen
irom base, ¢ (2) 16.5, 16; 2 (2) 16:5, 16; culmen from
anterior margin of nostril, 6 (2) 9, 8.5, 2 (2) 9; tarsus, 4
(2) 28.5, 9 (2) 28, 27.

B. m. sillimant from southern Palawan: Wing, ¢ (3) 68-
70.5 (av. 69), 2 (3) 63-68 (av. 66); tail, 8 (3) 46.5-48
(av. 47.3), 2 (8) 42-45 (av. 42.3); culmen from base, ¢
(3) 17-17.5 (av. 17.8), 2 (8) 17-18 (av. 17.5); culmen from
anterior margin of nostril, ¢ (3) 9-9.5 (av. 9.3), 2 (3)
Seay. 9-5 )r; tarsus, 6 (3) 27-29 Gav. 28.1); 2 (3) 27
28 (av. 27.6).

Rance: The highlands of southern Palawan as far as known.

Remarks: In the male, the Palawan form is the brightest
and richest in coloration of plumage, with the least powdery
gray wash on the slate-blue color, both on the upper- and
underparts, when compared to males of other races of the
species in the Philippines, Borneo, and Sumatra. The female of
this race also averages brighter and richer slate-blue on the
upperparts than those of the other races.

The Palawan birds resemble most closely those of northern
Luzon

The species is recorded for the first time on Palawan Island

8 Postilla Yale Peabody Museum No. 73

and is a new addition to the recently known mountain bird
fauna of this island.

The various races of the species within the geographic unit
of the Philippines include:

B. m. poliogyna—northern (northwestern) Luzon,

B. m. andersoni—southern Luzon,

B. m. brunneiceps—Negros,

B. m. mindanensis—Mt. Apo, Mindanao,

B. m. malindangensis—Mt. Malindang, Mindanao,

B. m. sillimani—southern Palawan.

Some of the larger islands with highlands attaining ele-
vations of 3,000 feet or more may have representatives of this
species that remain to be discovered. In southern Luzon, in the
Mount Isarog localities, the species ranges as low as 2,200 feet
along shaded banks of mountain streams.

The new race is named after Dr. Robert B. Silliman, Vice-
President of Silliman University, Dumaguete City, Negros
Oriental, Philippines, for his interest in and active stimulation

of biological research in the university.

Muscicapa westermanni palawanensis, subsp. nov.

Tyre: Yale Peabody Museum No. 73205 from Mount Man-
talingaian Peak, 5,500-6,000 feet, Palawan Island, Philippines.
Adult female, collected April 14, 1962 by D. S. Rabor.

Dracnosts: Like M. w. rabori and M. w. westermanni but dif-
fers in: a) bill decidedly more robust than that of either, as
shown by the comparative index derived from the ratio between
length of culmen from base and width of maxilla at level
of frontal feathering: b) in the female, the upperparts being a
purer, darker slate gray, and with the tawny-olive wash on the

back and rump still further reduced; c) longer wing and tail.

MerasurEMENts: M. w. palawanensis: wing 38 (5) 58.5-61
(av. 59.5), 2 (1) 55.5; tail ¢ (5) 42:5-44 (av. 43.3)5 2
(1) 40: culmen from base ¢ (5) 13-13.5 (av. 18:3) , 2 (1)
13; width of bill at level of frontal feathering ¢ (5) 4.6-
5.5 (av. 4.9), 2 (1) 5.1; tarsus ¢ (5))13:5-14.5 Cave 141mm).

EE

Dec. 20, 1962 New Birds from Palawan, Culion Islands 9

M. w. rabori: wing $ (9) D-O9 » (ave O19), 2 (3) o5-
57 (av. 56); tail ¢ (9) oe ae 41.2), 2 (8) 39-41, (av.
39.8); culmen from base 6 (9) 12-13.5 (av. 12.8), 2 (38)
12-13 (av. 12.8); width of bill at level of frontal feathering
6 (9) 3.8-4.9 (av. 4.4), 2 (3) 4-44 (av. 4.2)53 tarsus ¢ (9)
14-15.5 (av. 14.6), 2? (8) 13.5-15 (av. 14.3 mm).

M. w. westermanni: wing ¢ (16) 55-57.5 (av. 56.2), 2
GIO) 85225 ages Se 54.7); tail 6 (16) 39-41.5 (av. 40),
(10). 37-89.5 (38.5); culmen from base ¢ (16) 12-13.3 (av.
P25), 2 ene 12-18 (av. 12.5); width of bill at level of
frontal feathering ¢ (16) 4-4.6 (av. 4.3), 2 (10) 4-4.6)
(av. 4.4); tarsus ¢ (16) 13.5-15 (av. 18.9), 2 (10) 14-14.5
(av. 14.2 mm).

Rance: Mountains of southern Palawan Island, Philippines,
as far as known.

Remarks: The Palawan males are indistinguishable from
the males of the various Philippine races of the species, as is
characteristic of this particular species if based on plumage
coloration and color pattern.

The main bases for naming the Palawan population are:
a) the decidedly more robust bill in both sexes compared to
that of any other race of this species; b) the distinctive plum-
age coloration of the female; and c) the longer wing and tail
lengths as compared to that of the other races in the Philip-
pines. In order to arrive at comparative figures which deter-
mine the comparative degrees of heaviness of the bill in the
various Philippine races, we are using the ratio of the length
of culmen from base to the width of maxilla at level of frontal
feathering. We call this figure the “index of bill heaviness.”
The lower this figure is, the more heavy or robust the bill. We
included the figure for both sexes to arrive at indices which
will give a general idea as to the average heaviness of the bill
in either sex of a particular race.

The indices for bill heaviness of the races in the Philippines
are:

M. w. palawanensis—2.70 (53,12).

M. w. westermanni—2.87 (163,102 ).

M. w. rabori—2.91 (94,32).

10 Postilla Yale Peabody Museum No. 73

Cettia montana palawana, subsp. nov.

Type: 6 ad (Y.P.M. No. 54238), collected April 12, 1962,
by D. S. Rabor on Mount Mantalingajan, 6,700 feet, Palawan
Island, Philippines.

Diacnosis: This form differs strikingly in color from the
other subspecies of Cettia montana from India east to Laos
and south to Sumatra and Borneo. In olivaceous brownish tone
this population is paler than sepiaria and more olive, less
warm brown than oreophila, but altogether darker, more suf-
fused than mainland birds. Olive yellow is on the throat and
abdomen with warm olive brown on the flanks and invading the
breast in a light wash in some specimens forming an incomplete
breast band. These yellowish throat feathers tend to be
streaked along the shaft and adjacent areas of the vanes
with dull ashy, difficult to see on account of the make-up of
the skins. This yellowish tone of the underparts is strikingly
different from all other forms of Cettia montana. Like other
Palawan animals, however, this warbler points up the South-
east Asian affinities of this island.

MeEAsvrREMENTs: Type: wing, 54.5; tail, 56; culmen from
base, 15.5; culmen from anterior margin of nostril, 8.5; tarsus,
22 mm. Seven other males measure: wing, 54.5-57.5 (av. 55.8) ;
tail, 55.5-60 (av. 57.6); culmen from base, 14.5-15.5 (av.
15.1); culmen from anterior margin of nostril, 8-8.5 (av.
8.4); tarsus, 20-22.5 (av. 21.6). Six females measure: wing,
51-55.5 (av. 52.4) 3 tail, 47.5-54 (av. 51-5) ‘culmen from
base, 13.5-16 (av. 14.9); culmen from anterior margin of
nostril, 8-8.5 (av. 8.4); tarsus, 21-22 (av. 21.3).

Rance: Mountains of Palawan Island, Philippines.

Remarks: Stays singly or in pairs among the dense tangles
and low stunted growth on the mountain ridges, always actively
hopping from branch to branch, all the while giving out its
very characteristic short note. Frequently the course of the
bird may be followed by the sound of these notes, even though

the bird itself may not be visible among the dense growth.

Dec. 20, 1962 New Birds from Palawan, Culion Islands 11

The species is recorded for the first time on Palawan, and
in the Philippines, as a geographical unit and is a new addition

to the recently known mountain birds of this island.

Lonchura leucogastra palawana, subsp. nov.

Tyre: Yale Peabody Museum No. 73206 from Macagua,
Brooke’s Point, sea level to 250 feet, Palawan eee Philip-
pines. Adult male, collected April 3, 1962 by D. S. Rabor.

Diacnosis: Closest to L. 1. manueli in the deep chocolate
brown on chest becoming blackish brown to almost black on
throat and chin, but differs in: a) having the deep chocolate
brown on the chest, and the blackish brown or black on the
neck, throat and chin occupying a decidedly much larger area
up the sides of the neck and face; b) having a smaller bill.

MerasvureMENts: L. 1. palawanas: Wing é (14) 48.5-51.5
(av. 50.8), 2 (14) 49-52.5 (av. 49.9); tail 6 (14) 33-36.5,
@ (14) 31.5-87.5 (av. 35.4); length of culmen ¢ (14) 11.5
13 (av. 12.4), 2 (14) 11-13 (av. 12); greatest width of cul-
memed (140) 7-5-8. (av. 725), 2 (1a) 727-5 (av. 7-4) > bill
index 6 (14) 19-20.5 (av. 19.2), 2 (14) 18.5-20.5 (av.
19.5); tarsus 6 (14) 12.5-18.5 (av. 18), 2 (14) 12-18.5 (av.
15 mm).

E. l bE A Eas Wing ¢ (7) 47.5-50 (av. 49.2), 2 50.5;
tail do C7) 27-35.5 (av. 33), 2 33; HON of culmen ¢ (7)
10.5-12 (av. 11.6), 2 12; greatest width of culmen ¢ (7)

2
G:5-( (av. 6:7), 2 6:52 bill) imdex ¢ (7) 17-19 (av. 18.3),
ON UanSsmS |i) Leo oo Gave ls), 0o Lo.o mm:

L. 1. castanonota: Wing é (2) 49.5, 50.5; 2 (2) 49; tail
gz) aa-o,0046 9 (2) 35.5, 36.5; length of culmen ¢ (2)
11.5, 12, @ (2) 12, 12.5; greatest width of culmen 6 (2) 7,
ONC) Weni-o-, bill index 6 (2), 195, 20; 2 (2) 20; 20.5;
tarsus ¢ (2) 12.5, 13, 2 (2) 13 mm.

L. 1. everettt: Wing ¢ (9) 50-54 (av. 51.3), 2 (7% Me 52
(ay. 50-6)|;-tail ¢ ( eo

j
9) 33-37.5 (av. 85.3), 2 (7);

1 Postilla Yale Peabody Museum Nous

(av. 36.4); length of culmen ¢ (9) 12-12.5 (av. 12.3), 2
(7) 11.5-13 (av. 12.3); greatest width of culmen ¢ (7) 7.5-
8.5 (8), 2 (6) 7.5-8-5: (av. 7.9); bill mdex 3 .(7) 19:5-21
(av. 20.3), 2 (6) 19.5-20.5 (av. 20.3); tarsus 6 (9) 12-138.5
(av. 13.1), 2 (7) 12-13-5 (av. 13 mm).

L. l. manuelt: Wing 6 (7) 50.5-55 (av. 51.9), 2 (4) 48-
54 (av. 51.6); tail ¢ (7) 33-37.5 (av. 36.2), 2 (4) 32.5-35
(av. 33.8); length of culmen ¢ (7) 12.5-13 (av. 12.8), 2?
(4) 12.5-13.5 (av. oe oreatest coe of culmen 38 (7) 8-

t=)
Sp (ave 9), 2 (4) 5-8-5 (aye ; bill mdex 4 (7) 2025-
21> (av. ZO), 2 (4 ) 20-22 sie Bak ; tarsus ¢ (7) 12.5-

21.5
13.5 (av. 13), @ (4) 13-13.5 (av. 13.4 mm).

EL. l. smythiest: Wing id 51.5, 2 50.55 tall 36 3ihy Saar
length of culmen ¢ 12, 2? 13; greatest width of culmen ¢ 7.5,
S's bill index ¢. 19.5, 2 2i> tarsus 13:2 13:5 mm:

We follow Parkes’ measurement of the bill index, the index
derived from the sum of the greatest length of culmen plus its

greatest width.

Rance: Palawan, Busuanga and Culion as far as known in
the Philippines; the highlands of eastern and northern Borneo.
P} g

Remarks: Parkes (1958, Proc. U. S. Nat. Museum, vol. 108,
no. 3402, p. 279-284) reviewed the taxonomy and nomen-
clature of this species and described two new races, bringing
the total number of races that he recognized for the species to
five. He gave the ranges of these various races as:

L. 1. leucogastra—Thailand, Malay Peninsula, and Sumatra.
L. 1. castanonota—Southern Borneo.

L. l. everetti—Luzon and the adjacent islands of Mindoro,

Catanduanes, and Polillo, in the Philippines.

L. 1. manueli—Southern half of the Philippine Archipelago
and the highlands of northern Borneo, eastern Borneo,

and Sarawak.

en
~o

. smythiesi—Known only from the vicinity of Kuching,

Sarawak, Borneo.

Dec. 20, 1962 New Birds from Palawan, Culion Islands 13

Parkes had some Palawan specimens in his studies of this
species and he included them with the variable manweli. In this
connection he commented that “Although, as mentioned above,
manuelt is quite constant in its characters for a bird whose
range encompasses so many islands, there is a certain amount
of intraracial variation present. The most noticeable of these
variations is a tendency for Palawan specimens to have smaller
bills than those of the other islands within the range of
manueli as here defined.”

Our material, considering only the fully adult and properly
sexed specimens, consists of 8 (7¢, 12) L. lL. leucogastra, 4
(26, 22) castanonota, 16 (9 ¢, 72) everetiz, VL (7s,
42) manueli, 2 (16,12) smythiesi, and 28 (146, 142)
palawana. Our material of palawana includes 23 birds from
Palawan, two from Busuanga, one from Culion, and two from
northern Borneo. In addition, we also examined but did not
include the measurements, of two adult Palawan specimens of
undetermined sex.

In studying the various races of L. leucogastra, especially
those which are found in the Philippines, we made the following
observations:

a) Age of birds and their plumage. Immature and subadult
birds of the three Philippine races are indistinguishable from
one another. Only fully adult birds show to the best advantage
the characters which are of any value in differentiating the
various races.

b) Color of upperparts. L. l. castanonota differs distinet-
ively from the other races in having the deep rufous chestnut
upperparts. The other races differ very slightly from one
another in the general colors of the upperparts in being brown
streaked with white on the back and crown. There is a tendency,
however, toward a very gradual deepening in the intensity of
the brown upperparts in the various races (excluding casta-
nonota), starting from everetti as the lightest and ending with
leucogastra, in the order everetti > smythiesi > manueli >
palawana — leucogastra.

The degree of development of the white shaft streaks on the
feathers of the back up to the crown is a very variable charac-

14 Postilla Yale Peabody Museum No. 73

ter, even within the same race, and depends much on the age
of the individual bird and the freshness of its plumage. Within
the same population, in any one race, there are individuals
where these white shaft streaks have totally disappeared from
the crown, giving this part a uniform color. These white shaft
streaks are not as well pronounced on the back in some birds
and yet are very distinct in others of the same race and from

the same population of a particular locality.

c) Upper tail coverts. There is a gradual increase in the
degree of intensification from the original plain brown upper
tail coverts, which hardly contrast from the plain brown
color on the rump and lower back in smythiest, to deeper brown,
blackish brown, blackish, and finally to black in castanonota.
Among the races there is also a tendency for this color on the
upper tail coverts gradually to invade the rump, thus in-
creasing the area that it occupies. We summarize below the
condition of the upper tail coverts in the various races:

L. l. smythiesi—plain brown, almost with no contrast to

the rest of the rump and back.

L. l. everetti—plain brown with tendency to be slightly
more intense than the rump and lower back, thus
beginning to show a contrast.

L. 1. manueli—darker brown, contrasting distinctly with the
rump and the rest of back, but covers only a small
area of the upper tail coverts.

L. 1. palawana—blackish brown contrasting distinctly with
the rump and rest of back, occupying a larger area
on the upper tail coverts than in manuweli, and in some
specimens already beginning to show the tendency to
invade the rump.

L. 1. leucogastra—deeper blackish brown to almost black,
this color having invaded the greater part of the rump
or all of it, the whole area contrasting distinctly with
the rest of back.

lL. 1. castanonota—very intense blackish brown to black,
occupying the upper tail coverts and the rump, and

contrasting distinctly with the rest of back.

Dec. 20, 1962 New Birds from Palawan, Culicn Islands 15

d) Anterior underparts.

L. l. smythiesi—chocolate brown, becoming deeper and
richer on throat and chin.

L. l. everetti—chocolate brown, becoming deeper and richer

on throat and chin, as in smythiesi.

L, 1. manueli—deep chocolate brown on the chest, becoming
blackish brown on throat and chin, these colored areas
being separated by a wide band of light brown on the

sides of the chest and neck from the upperparts.

L. 1. palawana—as in manueli, deep chocolate brown on the
chest becoming blackish brown to black on throat and
chin, but these colored areas occupying a decidedly
much larger portion of the chest, neck, throat and chin,
so that the plain brown band separating them from

the upperparts is very much reduced.

L. l. leucogastra—the entire anterior underparts very
intense blackish brown to black, this color extending up
the sides of the chest, neck, throat and chin, coming in
direct contact with the much lighter brown of the upper-
parts.

L. l. castanonota—as in leucogastra, with the tendency to

be black instead of intense blackish brown.

The newly-described race is intermediate between manuwelt
and leucogastra in color pattern. When the various races
(excluding castanonota, because it is easily differentiated
from the others) are arranged in the order of increasing inten-
sity of the colors of the upperparts and underparts, the fol-
lowing arrangement results:

smythiesi > everetti > manueli > palawana — leuco-

gastra,

We did not have the opportunity to examine specimens
from the Sulu Archipelago but Parkes found the two birds

that he examined to be small-billed. a condition similar to the

16 Postilla Yale Peabody Museum No. 73

Palawan race. From this character and from geographic con-
sideration we are inclined to include the Sulu Archipelago
birds with the race palawana. The Philippine races of L.
leucogastra have the following ranges:

L. 1. everetti—Luzon and the adjacent islands of Mindoro,
Catanduanes, and Polillo.
L. 1. manueli—Central and southern Philippines.

L. 1. palawana—Palawan, Busuanga, Culion, Sulu Archi-

pelago.

OOS. 73

Ege:
Pe

YALE PEABODY MUSEUM

oF NaTurRAL History

Number 74 March 31, 1963 New Haven, Conn.

REVISION OF THE TYPE SPECIES OF THE
ORDOVICIAN NUCULOID PELECYPOD GENUS
TANCREDIOPSIS

A. Lert McALEsTER

ABSTRACT

The type species of the early Paleozoic nuculoid pelecypod genus
Tancrediopsis Beushausen, 1895, is shown to be Ctenodonta contracta
Salter, 1859, from Middle Ordovician rocks of southern Quebec, Canada.
This species is redescribed from a series of 189 well-preserved silicified
specimens, about half of which were collected at the original type locality.
This large sample permits the definition of limits of variation in the species
and reveals previously unknown morphologic features, among which are
strong pedal muscle scars and external escutcheonal perforations. The
species is easily confused with closely related sympatric species of Tan-
crediopsis, and criteria are discussed for distinguishing it from these
similar forms. The correct name for “Ctenodonta”’ contracta Salter is shown
to be Tancrediopsis cuneata (Hall).

INTRODUCTION

This paper is the first in a projected series to be devoted
to generic-level revisions of the systematics and phylogeny of
Paleozoic nuculoid pelecypods. As a vital first step toward
clarifying the early history of this common and long-ranging
group, a restudy is being made of the type species of each ge-
neric name that has been proposed for Paleozoic nuculoids.

2 Postilla Yale Peabody Museum No. 74

This redescription of the generic type species is being patterned
directly on a study of Paleozoic gastropod type species which
has been compiled by Knight (1941). As in Knight’s work, it
is planned to make these redescriptions as objective as possible
by basing them only on known original specimens of the species
involved (see Knight, 1941, p. 1, for an excellent discussion of
the value of objectivity in such work). An exhaustive search
of the literature has indicated that about 60 names have been
proposed for Paleozoic pelecypod genera which have at some-
time been considered to show taxodont dentition or other ev-
idence of nuculoid affinities.’ The redescription of the primary
types of the type species of these genera is now about half
completed, and it is expected that these revisions will be sub-
mitted for publication as one unit within a year.

As a further step toward understanding the morphology,
adaptations, and phylogeny of Paleozoic nuculoid genera, a
much longer range program is planned for assembling and
studying additional non-type material of the type species of
many of these genera. In this program, particular emphasis
will be placed on genera that cannot be adequately understood
from the surviving original type specimens. Every effort will
be made to assemble enough material of each species to permit
the application of modern concepts of population systematics
for determining the true nature and variability of specific and
generic characters. It is planned to publish these more compre-
hensive revisions intermittently, one genus at a time, as ad-
equate material of the type species can be assembled. This
paper is the first of this series of type species revisions based
on additional material that was not available to the original
author of the species. It treats the common Ordovician nu-

culoid genus T'ancrediopsis Beushausen (1895), whose type
tan) b Py

1The term “nuculoid” is used here in the broadest sense to include all
Nucula-like forms from Paleozoic rocks. In practice, this means all
Paleozoic peleeypods with taxodont dentition, because the convergent
development of such dentition in other unrelated groups, such as arcids
and the fresh-water genus /ridina, did not take place until after the
close of the Paleozoic. As thus defined, “nuculoid” is approximately
equivalent to the Subclass Protobranchiata (other than the Solemyidae)
which has recently been suggested to include all peleeypods with proto-
branch ctenidia (see Cox, 1959).

Mar. 31, 1963 Revision of T'ancrediopsis 3

species, Ctenodonta contracta Salter (1859), has never been
adequately understood.

This revision of Ctendonta contracta is based on Salter’s
15 original specimens from the Geological Survey of Canada
collections, supplemented by an excellent series of 174 si-
licified specimens from the collections of the Yale Peabody
Museum. About half of these additional specimens are from
the original type locality of the species. The Yale material
was collected in the early years of this century by C. EK. Beecher
and P. EK. Raymond as a part of an ambitious program of
restudy of important silicified early Paleozoic faunas of eastern
North America. Many hundreds of pounds of rock from several
localities were collected and etched, but the program was cut
short by Beecher’s untimely death in 1904, with the result
that this great wealth of material has never been studied. The
etched Middle Ordovician collections have now been sorted for
pelecypods and have yielded about 1500 identifiable specimens.
These collections are particularly rich in well-preserved nu-
culoid species, and it is expected that this material will provide
a basis for future studies on the earliest evolutionary radiation
of this important group of pelecypods.

I am most grateful to the National Science Foundation for
support of these studies of Paleozoic nuculoid pelecypods under
its Program for Systematic Biology, Division of Biological
and Medical Sciences (Grant No. G19961). I am also greatly
indebted to D. J. McLaren, T. E. Bolton, and G. W. Sinclair
of the Geological Survey of Canada for the loan of Salter’s
types and for generous cooperation on this project. Finally,
I wish to thank D. W. Harvey and Martha Erickson for their
skillful preparation of the photographs and drawings, and
C. J. Durden, who made the preliminary sorting of the silicified
Peabody Museum material.

Genus TANCREDIOPSIS
Author. Beushausen, 1895, pe wO:

Type Species. Ctenodonta contracta Salter (118595 p37)
[=T'ellinomya cuneata Hall, 1856, p. 892] by subsequent des-
ignation of Cossmann, 1897, p. 94.

4 Postilla Yale Peabody Museum No. 74

Discussion. Most Ordovician nuculoid pelecypods have been
described under the generic name Ctenodonta Salter, 1852,
which has as its type the large and distinctive, but rather
uncommon Ordovician species Ctenodonta nasuta (Hall), 1847
(see Wilson, 1956, pl. 2 for illustrations of this species). In
an attempt to subdivide further the complex of species tradi-
tionally assigned to “Ctenodonta,” Beushausen proposed the
subgeneric name T'ancrediopsis for more common, smaller, early
Paleozoic nuculoids typified by the Ordovician species Cteno-
donta contracta Salter (1859) and the Silurian species Nucwa
sulcata Hisinger (1841), both of which were originally desig-
nated by Beushausen as the “types” of his new subgenus.
The first subsequent designation of one cf these species as the
type of the subgenus was made by Cossmann two years after
the original description (1897, p. 94), when he chose Cteno-
donta contracta Salter as the type species. It will be shown
later that the correct name for this species is T’ancrediopsis

cuneata (Hall).

Beushausen’s name T'ancrediopsis seems to have been ignored
by all later workers on Ordovician pelecypeds, but it has gained
some usage as a generic name with workers describing Silurian
and Devonian nuculcids (Prosser and Kindle, 1913; McLearn,
1924; Reed, 1931; Northrop, 1989; Sherrard, 1960). In this
regard it should be noted that even though Beushausen origi-
nally named an Ordovician and a Silurian species as the “types”
of the subgenus, the name was proposed to facilitate the deserip-
tion of Devonian species which were the subject of his mono-
graph. While it is still too early to determine the final usefulness
of the generic name, it appears likely that many of the post-
Ordovician species to which it has been apphed cannot be con-
sidered as congeneric with the type species, “Ctenodonta con-
tracta.”’ The name T'ancrediopsis will, however, prcbably prove
useful in the future as a generic subdivision of the heteroge-
neous assemblage of Ordovician forms now included in “Cteno-
donta.”” Here again, a final determination of the value of the
name must await further study of other genera and species of
Ordovician nuculoids. In anticipation of such studies it appears

that many Ordovician nuculoid species show closer affinities to

Mar. 31, 1963 Revision of T'ancredio psis 5

“Ctenodonta contracta,” the type of T'ancrediopsis, than to
Ctenodonta nasuta, the distinctive and uncommon type of the
genus Ctenodonta. For this reason, the transfer of many Or-
dovician species from Ctenodonta to T'ancrediopsis may prove

desirable in future revisions.

Tancrediopsis cuneata (Hall)

Figures 1-80

Tellinomya cuneata Hall, 1856, p. 392, figs. 6, 7. Hall, 1857a,
p. 188, figs. 6, 7. Hall, 1857b, p. 143, figs. 6, 7. [not | Hall,
1862.9; 38; figs. 1, 2.

Ctenodonta contracta Salter, 1859, p. 37, pl. 8, figs. 4, 5.
Logan, 1863, p. 175, figs. 160a, 160b. Wilson, 1956, p. 28,
plaZzoties. 4-9.

Ctenodonta (Tancrediopsis) contracta (Salter). Beushausen,
1895; p: 70.

[?] Tellinomya contracta? (Salter). Walcott, 1884, p. 76,
pl lio figs. 15, 15a.

Revised description. Shel] of small size (median height of 136
measurable specimens 8 mm), equivalved, strongly convex, thick
and massive, constricted posteriorly.” Shape variable, height
ranging from 62 to 86 per cent of length (median of 86 meas-
urable specimens 71 per cent). Surface sculpture of very fine,
widely spaced concentric ridges which are commonly divided
into rod-like pustules, especially near the outer margin (figs.
4-7, 18, 19, 24, 26, 28, 35, 37). Sculpture usually obscure or
absent, probably because of difficulty of preservation rather
than absence on original shell. The dorsal margin shows a
preminent oval lunule anterior to the umbones and a more
elongate and obscure posterior escutcheon (figs. 65-73). Lu-
nule and escutcheon variable in size and shape. Strong. chev-
ron-shaped, taxodont dentition with approximately equal num-

2 By analogy with living nuculoids, the smaller, contracted end of T.
cuneata is considered to be posterior and the larger end to be anterior
as shown in figs. I and 2.

6 Postilla Yale Peabody Museum No. 74

bers of teeth on both sides of umbo, teeth decreasing abruptly
in size but apparently continuous in umbonal region (figs. 46-
64, 73-80). One well-preserved specimen shows several tiny
perforations along the margins of the escutcheon, probably
representing an extreme elongation of the dental sockets (fig.
71; see Sorgenfrei, 1937, and Trueman, 1952, for discussions
of similar structures in Cenozoic nuculoids). Resilifer
absent. Several specimens show a strong, external ligament
structure preserved as a silicified replica along the anterior
third of the escutcheon posterior to the umbones (figs. 65-67,
69, 70, 72, 80). Strong, subequal adductor scars deeply im-
pressed into the thick shell material, bounded on interior side
by thickened shell material making raised ridge which is most
prominent behind the anterior scar (figs. 75-80). Two small
but deeply impressed subequal pedal muscle scars cecur just
below the hinge plate at the dorsal end of these adductor
ridges (figs. 74-80). A few well-preserved specimens also show
faint impressions just below the posterior hinge plate which
may represent cther pedal or visceral muscle scars (figs. 75,
77). Pallial line not preserved, probably very faint or absent
on original shell material. Original calcareous shell material

unknown, replaced by amorphous silica in all specimens.

Types. Lectotype ef Tellinomya cuneata Hall, here des-
ignated, the specimen shown by Hall, 1856, as fig. 7 (and
fig. 6 if both figures represent the same specimen), p. 392;
whereabouts unknown. Type locality: “Pauquette’s Rapids on
the Ottawa River” [between Allumette Island, Quebec and
Ontario mainland, about three miles south of Waltham, Que-
bec, Canada (see Kay, 1942, pl. 6) |. Stratigraphic position:
“Beds lying at the junction of the Trenton and Black River
limestones” [Rockland beds cf the Ottawa formation, Middle
Ordovician (lower Trenton stage of Twenhofel, 1954) ]. Lecto-
type of Ctenodonta contracta Salter, by designation of Wil-
son, 1956, p. 23, No. 1171b in the collections of the Geological
Survey of Canada, Ottawa, Ontario, Canada. This specimen
is one of Salter’s original syntypes which was figured by him
(1859 )as pl. 8, figs. 5, 5a. Type locality: *‘Allumette Islands,”

Mar. 31, 1968 Revision of T'ancrediopsis vi
Quebec [probably Paquette Rapids between Allumette Island,

Quebec and Ontario mainland, about three miles south of Wal-
tham, Quebec, Canada (see Kay, 1942, pl. 6) |]. Stratigraphic

lun

anterior posterior

Figure 1. Vanerediopsis cuneata (Hall). Generalized dorsal view of artic-
ulated valves showing lunule (lun), ligament (lig), escutcheon (es) and
escutcheonal perforations (ep).

apm

aam

anterior posterior

=
Re el

Figure 2. Tancrediopsis cuneata (Hall). Generalized interior view of
right valve showing anterior adductor muscle scar (aam), anterior pedal
muscle scar (apm), posterior pedal muscle scar (ppm), posterior adductor
muscle scar (pam), and additionai possible pedal or visceral muscle scars
along posterior dorsal margin (pm?).

position: “Allumette limestenes” [probably Rockland beds of
the Ottawa formation, Middle Ordovician (lowest Trenton
stage of T'wenhofel, 195+) |.

8 Postilla Yale Peabody Museum No. 74

The location of Hall’s original figured specimen is unknown,
for it is not listed in the type catalogues of either the Ameri-
can Museum of Natural History or the New York State Mu-
seum (Whitfield and Hovey, 1898-1901; Clarke and Ruede-
mann, 1908), the two institutions which contain most of Hall’s
original material. The specimen may be as yet unrecognized
among the non-type material at these or other institutions.
Hall’s original figure leaves little room for doubt, however, as
to the identity of the species in question, and the designation
of a neotype therefore seems unnecessary.

Material. This revised description is based on 189 silicified
valves. Fifteen of these are Salter’s original types of Cteno-
donta contracta from the collections of the Geological Survey
of Canada, Ottawa, Ontario, Canada (catalogue No. 1171).
The remaining 174 specimens are from the collections of the
Peabody Museum, Yale University, New Haven, Connecticut,
U.S. A. All of Salter’s specimens and almost half of the Yale
material (75 specimens) were collected at Paquette Rapids
(between Allumette Island, Quebec and Ontario mainland,
about three miles south of Waltham, Quebec, Canada; see Kay,
1942, pl. 6) which is the original type locality of both T'el-
linomya cuneata Hall and Ctenodonta contracta Salter. The
remaining Yale specimens (99) were collected from Middle
Ordovician limestones of approximately the same age exposed
at Pointe Bleue on the shores of Lake St. John, about five
miles north of Roberval, Quebec, Canada (see Dresser, 1916,
map 184A). Both Yale collections were made in 1903 by C. E.
Beecher and P. E. Raymond. The original specimens studied
by Salter were collected by W. E. Logan in 1845. The distribu-
tion of specimens of T'ancrediopsis cuneata at these institu-
tions and localities is shown in more detail in Table 1.

The material shows considerable variation in the quality of
the silicified preservation. In some specimens the whole valve is
preserved, but many valves were incompletely replaced by silica
and do not show the entire outline. The fragile posterior ex-
tremity is commonly missing, but the heavy hinge plate and

dentition are almcst always preserved. The specimens from

Mar. 31, 1963 Revision of T'ancrediopsis 9

TABLE 1]

Distribution of specimens of Tancrediopsis cuneata (Hall) used in this
study. For consistency, each valve of articulated individuals is counted as
one “specimen.”

Locality

and Right Left Articulated Total
Source valves valves valves — specimens

Paquette Rapids

Yale Peabody Museum... 44. 29 2 75
Geological Survey
OhmCanadaneeeeeeer. 6 7 2 15
Pointe Bleue, Lake St. John
(All Yale Peabody
IMIS GUM) ever e eieiee 19 20 60 99

RG Gaull Simi sete sta srites hate, onset 69 56 64. 189

Lake St. John are generally more coarsely silicified and show
fewer morphologic details than do those from Paquette Rapids.

Occurrence. The species is known with certainty only from
the type locality and the Lake St. John Iccality mentioned
above, both of which probably represent horizons in the mid-
dle part of the Middle Ordovician (lowest Trenton stage of
the American standard, approximately lower Caradoc of the
European standard, see Twenhofel, 1954). Several additional
Ontario and Quebec occurrences are listed by Wilson (1956,
p- 23), and the species may be represented in the Ordovician
of Nevada (Walcott, 1884). Restudy cf other important
Ordovician pelecypod faunas may prove the species to be more
abundant and widespread than is apparent from the evidence
now available. In most such faunas nuculoid forms are not
silicified, but are preserved as internal or composite molds (see
McAlester, 1962). For this reason, an artifical internal mold
of a well-preserved, silicified specimen of 7’. cuneata has been
figured here to facilitate comparison with other faunas (fig.

47).

10 Postilla Yale Peabcdy Museum No. 74

Discussion. There has been considerable confusion in the
identification of Salter’s species “Ctenodonta contracta” and
other closely related species from the Paquette Rapids locality.
The Yale collections from this locality contain about 700 sili-
cified nuculoid specimens, and an analysis of these specimens
has revealed that at least seven species of nuculoids occur in
appreciable numbers at Paquette Rapids. Several additional
nuculoid species probably also occur in the fauna but are quite
rare and are represented by only a few specimens from the
Yale collections. Five of the seven common species are distinc-
tive and cause little confusion in identification. Wilson (1956,
pl. 2) illustrates four of these distinctive species, which are
identified by her as: Ctenodonta astartaeformis Salter, Cteno-
donta levata (Hall), Ctenodonta nasuta Hall, and Ctenodonta
logant Salter. A fifth common species, a small Palaeoneilo-like
form, is not mentioned by Wilson and may be as yet un-
described.

It is the final two common nuculoid species in the Paquette
Rapids fauna which are easily confused. Both are medium-sized
(for nuculoids), thick-shelled forms with contracted posterior
extremities, strong adductor impressions, and similar patterns
of dentition. Analysis of the large Yale collections shows that
the two forms do, however, show consistent and distinct dif-
ferences in shape, which are most cbvious from comparing the
shell exteriors. To facilitate comparison the two species will
be referred to here as “Form A” and “Form B.” Form A
has more central umbones, a more elongate and_ gently-
sloping posterior constriction, and a differently shaped an-
tericr-ventral and anterior margin. Internally, the differences
are less distinctive, but the more central umbones, less abrupt
posterior constriction, and longer posterior dentition of Form
A can usually be recognized. Form B also seems to have lacked
the very fine, pustulose, concentric sculpture seen on well-pre-
served specimens of Form A. Form B also appears to have had
weaker and somewhat differently oriented pedal muscle scars.
In Form A the chevron-shaped teeth point toward the umbo,
whereas they tend to point away from the umbo in Form B.

These differences in shape and sculpture are shown in fig. 3.

Mar. 31, 1963 Revision of T'ancrediopsis aE

There are no morphological intermediates between these two
common forms in the Paquette Rapids fauna, and they cer-
tainly represent two closely related sympatric nuculoid species.
Among approximately 700 identifiable nuculoid specimens from
the locality in the Yale collections, about 250 (86 per cent)
are form B, 75 (or about 11 per cent) are Form A, and the
bulk of the remainder (about 375 specimens or 53 per cent) rep-
resent the five additional species mentioned above. The lectotype
of Hall ’s species T'ellinomya cuneata is readily recognizable

T. cuneata T. “abrupta”’
(‘Form A") (''Form B")

INTERIOR
(right valves)

EXTERIOR
(left valves)

Figure 3. Internal and external views showing morphologic differences be-
tween the closely related sympatric species Tancrediopsis cuneata (Hall)
and Tancrediopsis “abrupta”’ (Billings).

from the original figure as belonging to Form A (sub-central
umbones, elongate posterior, etc.) as are also the lectotype
and other original specimens of Ctenodonta contracta Salter.
The lectotypes of both species are from Paquette Rapids, and
there is little doubt that the two names are synonyms.

Hall’s name “T'ellinomya cuneata” was not formally pro-
posed as a new species, but was merely first listed in explana-
tion of figures of ‘“T'ellinomya” [—=Ctenodonta| in a discus-
sion of the genus (1856) which was later reprinted in two
forms (1857a, 1857b). Characters of the species are, however,
briefly mentioned in Hall’s text (p. 892 in the original report).
The figures and discussion of the species in Hall’s report cer-

12 Postilla Yale Peabody Museum No. 74

tainly constitute a valid description or “indication” of a new
species as prescribed in the rules of nomenclature (see Stoll,
1961, p. 15), and Hall’s name, which has three-years priority
over Ctenodonta contracta Salter, is therefore accepted here
as the correct name for the species. This strict usage of prior-
ity may violate the new “nomen oblitum’’ provision of the
nomenclatural cede which provides that ‘ta name that has re-
mained unused as a senior synonym in the primary zoological
literature for more than fifty years is to be considered a for-
gotten name (nomen oblitum). . . [and] is not to be used un-
less the Commission so directs” (Stoll, 1961, p. 23). However,
the application of the rule is ambiguous in this case because
T'ellinomya cuneata has appeared in what is presumably the
“primary zoological literature” as an incorrectly suppressed
senior synonym of Ctenodonta contracta within the past fifty
years (Bassler, 1915, p. 302). In addition, the prescribed pro-
cedure for applying the rule is extremely cumbersome and in
my opinion will do more to create nomenclatural instability
than to correct it because every “tnomen oblitum’? must be
laboriously referred to the Commission for action. For these
reasons I prefer a strict interpretation of the Law of Priority
in this case (for additional objections and comment on the
“nomen oblitum’” provision see the Bulletin of Zoological No-
menclature, Volume 19, Part 6, 28 December, 1962).

The name of the second common species (‘Form B”) is not
yet certain, but it appears that the lectotype of Ctenodonta
abrupta Billings (1862; see also Wilsen, 1956) is a represent-
ative of “Form B,” and this may be the correct name for the
species. These two closely related species (T'ancrediopsis cu-
neata and “Ctenodonta abrupta’) are certainly congeneric
and thus C. abrupta would become T'ancrediopsis “abrupta”

(Billings).

In addition to these two commen species of T'ancrediopsis,
a few other specimens from the Paquette Rapids fauna may
represent closely related but very rare species. Several such
specimens in the Yale collections do not appear to fall within
the range of shape variation of either 7’. cuneata or T. “ab-

rupta,” but at present these specimens are too rare either to

Mar. 31, 196: Revision of T'ancrediopsis 1033

warrant specific names or to cause confusion in the identifica-
tion of JT. cuneata. The original figured specimen of Cteno-
donta gibberula Salter (1859, pl. 8, fig. 6) may represent one
such species but, regrettably, the specimen appears to have
been lost (see Wilson, 1956, p. 24). Pending the discovery of
the original specimen or similar additional material, it does
not seem prudent to recognize Salter’s name. The two names
C. abrupta Billings and C, gibberula Salter appear to be the
only names proposed for Paquette Rapids specimens which are
closely related to 7’. cuneata.

In comparison to the Paquette Rapids fauna, nuculoids are
somewhat less common and mere poorly preserved in the silici-
fied fauna from Lake St. John, but the fauna does contain
many articulated specimens which are rare at Paquette Rapids
(see Table 1). The nuculoid species at Lake St. John all ap-
pear to be indistinguishable from those at Paquette Rapids,
although the relative abundances differ at the two localities.
At Lake St. John, 7’. cuneata is over three times as abundant
as T', “abrupta” (99 vs. 30 specimens), whereas this situation
is reversed at Paquette Rapids where 7’. “abrupta” strongly
dominates (approximately 250 specimens vs. 75 specimens of
T’. cuneata).

LITERATURE CITED

Beushausen, Ludwig, 1895. Die Lamellibranchiaten des rheinischen Devon
mit Ausschluss der Aviculiden: Kgl. Preussischen Geol. Landesanstalt
Abh., Neue Folge, no. 17, 514 p., 28 pls.

Billings, Elkanah, 1862. New species of Lower Silurian fossils [part 2]:
Montreal, Geol. Survey Canada, p. 25-56 [reprinted in 1865 as part of
“Palaeozoic Fossils,” v. 1]

Clarke, J. M., and Ruedemann, Rudolph, 1903. Catalogue of type specimens
ot Paleozoic fossils in New York State Museum: New York State Mus.
Bull. 65, 847 p.

Cossmann, Maurice, 1897. Revue critique de paléozoologie, premiére année
Nos. 1-4): Paris, Société d’Editions Scientifiques, 186 p.

Cox, L. R., 1959. The geological history of the Protobranchia and the dual
origin of taxodont Lamellibranchia: Malacological Soc. London Proc.,
v. 33, p. 200-209.

Dresser, J. A., 1916. Part of the district of Lake St. John, Quebec: Canada
Geol. Survey Mem. 92, geol. ser. no. 74, 88 p., 5 pls.

Hall, James, 1856. On the genus Tellinomya and allied genera: Canadian
Naturalist and Geologist, v. 1, p. 390-395.

14 Postilla Yale Peabody Museum No. 74

———, 1857a. On the genus Tellinomya and allied genera: Regents
of the Uniy. of the State of New York, State Cabinet of Nat. History,
10th Ann. Rept., App. C, p. 181-186.

, 1857b. On the genus Tellinomya and allied genera, in Descrip-
tions of new species of Palaeozoic fossils, ete., extracted from 10th
Ann. Rept. Regents of the Univ. of the State of New York: p. 141-146.

——-, 1862. Physical geography and general geology; Remarks upon
the condition of the fossils, etc., in Hall, James, and Whitney, J. D.,
Rept. Geol. Survey Wisconsin, v. 1: p. 1-72 (chap. 1), 425-448 (chap. 9).

Hisinger, Wilhelm, 1841. Lethaea Svecica sue petrificata Sveciae, iconibus et
characteribus illustrata [2nd supplement, part 2]: Holmiae, 6 p., pls.
40-42. a

Kay, G. M., 1942. Ottawa-Bonnechere graben and Lake Ontario homo-
cline: Geol. Soc. America Bull., v. 53, p. 585-646.

Knight, J. B., 1941. Paleozoic gastropod genotypes: Geol. Soc. America
Spec. Paper 32, 510 p., 96 pls.

Logan, W. E., and others, 1863. Geology of Canada; Geological Survey of
Canada, Report of progress from its commencement to 1863, ete.:
Montreal, Dawson Brothers, 983 p.

McAlester, A. L., 1962. Mode of preservation in early Paleozoic pelecy-
pods and its morphologic and ecologic significance: Jour. Paleontology,
v. 36, p. 69-73.

McLearn, F. H., 1924. Paleontology of the Silurian rocks of Arisaig, Nova
Scotia: Canada Geol. Survey Mem. 137, geol. ser. no. 118, 180 p., 30
pls.

Northrop, S. A., 1939. Paleontology and stratigraphy of the Silurian rocks
of the Port Daniel-Black Cape region, Gaspé: Geol. Soc. America
Spec. Paper 21, 302 p., 28 pls.

Prosser, C. S., and Kindle, E. M., 1913. Pelecypoda, in Systematic pale-
ontology of the Middle Devonian deposits of Maryland: Maryland
Geol. Survey Middle and Upper Devonian, p. 214-279, 14 pls. [in
separate volume].

Reed, F. R. C., 1931. Some new lamellibranchs from the Silurian of the
Ludlow district: Annals and Mag. Nat. History, ser. 10, v. 8, p. 289-
304.

Salter, J. W., 1852. Note on the fossils above mentioned, from the Ottawa
River: British Assoc. Adv. Sci. Rept., 21st Mtg., 1851, Notices and
Abs., ete., p. 63-65.

—___———,, 1859. Fossils from the base of the Trenton group: Geol
Survey Canada, Figs. and Descriptions of Canadian Organic Re-
mains, decade 1, 47 p., 10 pls.

Sherrard, Kathleen, 1960. Some Silurian lamellibranchs from New South
Wales: Linnean Soc. New South Wales Proc., v. 84, p. 356-372.

Sorgenfrei, Theodor, 1937. Some remarks on the hinge of nuculids and
ledids: Vidensk. Medd. Dansk naturh. Foren. Kgébenhavn, vy. 100, p.
369-375.

Stoll, N. R., chairman, 1961. International code of zoological nomencla-
ture adopted by the XV International Congress of Zoology: London,
Internat. Trust for Zool. Nomenclature, 176 p.

Trueman, E. R., 1952. Observations on the ligament of Nucula: Mala-
cological Soc. London Proc., v. 29, p. 201-205.

Mar. 31, 1963 Revision of T'ancrediopsis 15

Twenhofel, W. H., chairman, 1954. Correlation of the Ordovician forma-
tions of North America: Geol. Soc. America Bull., v. 65, p. 247-298.

Walcott, C. D., 1884. Paleontology of the Eureka district [Nevada]: U.S.
Geol. Survey Mon. 8, 298 p., 24 pls.

Whitfield, R. P., and Hovey, E. O., 1898-1901. Catalogue of the types and
figured specimens in the paleontological collection of the geological
department, American Museum of Natural History: Am. Mus. Nat.
History Bull. v. 11, 500 p.

Wilson, A. E., 1956. Pelecypoda of the Ottawa formation of the Ottawa-
St. Lawrence Lowland: Geol. Survey Canada Bull. 28, 102 p., 9 pls.

16 Postilla Yale Peabody Museum No. 74

Figures 4-21. Tancrediopsis cuneata (Hall). A series of exterior views of
silicified right valves showing variation in shape and sculpture. All figures
are twice natural size. Precise locality information for the “Paquette Ra-
pids” and “Lake St. John” localities is given in the text. YPM=/Yale
University Peabody Museum collections, New Haven, Connecticut, U.S.A.
GSC = Geological Survey of Canada collections, Ottawa, Ontario, Canada.

Figure 4. YPM 22966, Lake St. John. Figure 5. YPM 22967, Lake St.
John. Figure 6. YPM 22968, Lake St. John. Figure 7. YPM 22969,
Lake St. John. Figure 8. YPM 22970, Lake St. John. Figure 9. YPM
22971, Paquette Rapids. The specimen lacks the anterior and pos-
terior extremities. See also fig. 48. Figure 10. YPM 22972, Paquette
Rapids. The specimen lacks the posterior extremity. Figure 11. YPM
22973, Lake St. John. Figure 12. YPM 22974, Lake St. John. Figure
13. YPM 22975, Paquette Rapids. The specimen lacks the anterior
extremity. See also fig. 50. Figure 14. YPM 22976, Paquette Rapids.
The specimen lacks the anterior and posterior extremities. Figure 15.
YPM 22977, Paquette Rapids. See also figs. 52, 68. Figure 16. YPM
22978, Lake St. John. Figure 17. GSC 1171c, Paquette Rapids. One of
Salter’s original specimens of Ctenodonta contracta, figured by Wil-
son, 1956, as figs. 7 and 8 of pl. 2. See also fig. 70. Figure 18. YPM
22979, Paquette Rapids. The anterior part of the specimen is missing.
Figure 19. GSC 1171k, Paquette Rapids. One of Salter’s original
specimens of Clenodonta contracta, figured by Wilson, 1956, as fig.
9 of pl. 2. The specimen lacks the anterior extremity. Figure 20.
GSC ll7le, Paquette Rapids. One of Salter’s original specimens of
Ctenodonta contracta. See also fig. 66. Figure 21. YPM 22980,
Paquette Rapids. See also figs. 47, 55, 77, 79.

Mar. 31, 19638 Revision of T'ancrediopsis 7

Figures 22-47. Tancrediopsis cuneata (Hall). Figures 22 through 42
are a series of exterior views of silicified right valves showing variation
in shape and sculpture. Figures 43 through 45 are dorsal views of three
articulated silicified specimens (anterior end to left). Figure 46 is an
interior view of the largest known specimen, a silicified left valve. Figure
47 is a latex internal mold of a silicified right valve showing the appear-
ance of the species as normally preserved in non-silicified faunas. All
figures are twice natural size. Precise locality information for the
“Paquette Rapids” and “Lake St. John” localities is given in the text.
YPM= Yale University Peabody Museum collections, New Haven, Con-
necticut, U.S.A. GSC = Geological Survey of Canada collections, Ottawa,
Ontario, Canada.

Figure 22. YPM 22981, Lake St. John. Figure 23. YPM 22982, Lake
St. John. Figure 24. YPM 22983, Lake St. John. Figure 25. YPM
22984, Lake St. John. Figure 26. YPM 22985, Lake St. John. Figure
27. YPM 22986, Lake St. John. Figure 28. YPM 22987, Lake St. John.
Figure 29. YPM 22988, Lake St. John. Figure 30. YPM 22989, Lake
St. John. Figure 31. YPM 22990, Lake St. John. Figure 32. YPM
22991, Paquette Rapids. The specimen lacks the anterior extremity.
See also fig. 58. Figure 33. YPM 22992, Lake St. John. Figure 34.
GSC 1171L, Paquette Rapids. One of Salter’s original specimens of
Ctenodonta contracta. See also fig. 43. Figure 35. GSC 1171j, Paquette
Rapids. Figure 36. YPM 22993, Lake St. John. See also fig. 44.
Figure 37. YPM 22994, Lake St. John. Figure 38. YPM 22995, Lake
St. John. See also fig. 45. Figure 39. Lectotype of Ctenodonta con-
tracta Salter, GSC 1171b, Paquette Rapids, figured by Salter, 1859,
as figs. 5 and 5a of pl. 8. The specimen lacks the posterior extremity.
See also figs. 61, 67, 75, 80. Figure 40. GSC 1171m, Paquette Rapids.
One of Salter’s original specimens of Ctenodonta contracta. The
specimen lacks the anterior extremity. See also figs. 60, 73, 76. Figure
41. GSC 1171i, Paquette Rapids. One of Salter’s original specimens of
Ctenodonta contracta. See also fig. 69. Figure 42. GSC 1171d,
Paquette Rapids. One of Salter’s original specimens of Ctenodonta
contracta, Figure 43. GSC 1171L, Paquette Rapids. One of Salter’s
original specimens of Ctenodonta contracta. See also fig. 34. Figure
44. YPM 22993, Lake St. John. See also fig. 36. Figure 45. YPM
22995, Lake St. John. See also fig. 38. Figure 46. GSC 117la, Paquette
Rapids. One of Salter’s original specimens of Ctenodonta contracta.
Figure 47. Latex cast of YPM 22980, Paquette Rapids. See also figs.

Ze OO Mills D>

18 Postilla Yale Peabcdy Museum No. 74

Figures 48-64. Tancrediopsis cuneata (Hall). A series of internal views of
silicified valves showing dentition and adductor musculature. Figures 48
through 56 are right valves; figures 57 through 64 are left valves. All
figures are twice natural size. Precise locality information for the
“Paquette Rapids” locality is given in the text. YPM= Yale University
Peabody Museum collections, New Haven, Connecticut, U.S.A. GSC =
Geological Survey of Canada collections, Ottawa, Ontario, Canada.

Figure 48. YPM 22971, Paquette Rapids. See also fig. 9. Figure 49.
YPM 22996, Paquette Rapids. Figure 50. YPM 22975, Paquette Rap-
ids. See also fig. 13. Figure 51. YPM 22997, Paquette Rapids. Figure
52. YPM 22977, Paquette Rapids. See also figs. 15, 68. Figure 53.
YPM 22998, Paquette Rapids. Figure 54. GSC 1171g, Paquette Rap-
ids. One of Salter’s original specimens of Ctenodonta contracta. See
also fig. 72. Figure 55. YPM 22980, Paquette Rapids. See also figs.
21, 47, 77, 79. Figure 56. YPM 22999, Paquette Rapids. See also fig.
74. Figure 57. YPM 23000, Paquette Rapids. Figure 58. YPM 22991,
Paquette Rapids. See also fig. 32. Figure 59. YPM 23001, Paquette
Rapids. See also fig. 78. Figure 60. GSC 1171m, Paquette Rapids.
One of Salter’s original specimens of Ctenodonta contracta, See also
figs. 40, 73, 76. Figure 61. Lectotype of Ctenodonta contracta Salter,
GSC 1171b, Paquette Rapids, figured by Salter, 1859, as figs. 5 and
5a of pl. 8. See also figs. 39, 67, 75, 80. Figure 62. YPM 23002,
Paquette Rapids. Figure 63. YPM 23003, Paquette Rapids. Figure
64. YPM 23004, Paquette Rapids. See also fig. 71.

Mar. 31, 19638 Revision of T'ancrediopsis 19

Figures 65-80. Tancrediopsis cuneata (Hall). A series of enlarged views of silici-
fied valves showing details of ligament, dentition, and musculature. All figures are
three times natural size except Figure 65 (twice natural size) and Figure 71 (six
times natural size). Precise locality information for the “Paquette Rapids” locality
is given in the text. YPM=Yale University Peabody Museum collections, New
Haven, Connecticut, U.S.A. GSC=Geological Survey of Canada collections, Ot-
tawa, Ontario, Canada.

Figure 65. GSC 1171f, Paquette Rapids. One of Salter’s original figured speci-
mens of Ctenodonta contracta, illustrated by Salter, 1859, as figs. 4 and 4a of
pl. 8. Dorsal view of articulated valves (anterior end to left) showing lunule,
escutcheon, and silicified replica of ligament. Figure 66. GSC 117le, Paquette
Rapids. One of Salter’s original specimens of Ctenodonta contracta. Dorsal
view of right valve showing lunule, escutcheon, and silicified replica of liga-
ment. See also fig. 20. Figure 67. Lectotype of Ctenodonta contracta Salter,
GSC 1171b, Paquette Rapids, figured by Salter, 1859, as figs. 5 and 5a of pl. 8.
Oblique dorsal view of left valve showing lunule, escutcheon, and_ silicified
replica of ligament. See also figs. 39, 61, 75, 80. Figure 68. YPM 22977, Paquette
Rapids. Oblique dorsal view of right valve showing lunule. See also figs. 15,
52. Figure 69. GSC 1171li, Paquette Rapids. One of Salter’s original specimens
of Ctenodonta contracta. Oblique dorsal view of left valve showing escutcheon
and silicified replica of ligament. See also fig. 41. Figure 70. GSC 1171c, Paquette
Rapids. One of Salter’s original specimens of Ctenodonta contracta, figured by
Wilson, 1956, as figs. 7 and 8 of pl. 2. Oblique dorsal view of right valve interior
showing lunule, escutcheon, and silicified replica of ligament. See also fig. 17.
Figure 71. YPM 23004, Paquette Rapids. Enlarged dorsal view of left valve
showing lunule, escutcheon, and tiny perforations along posterior margin of
escutcheon. See also fig. 64. Figure 72. GSC 1171g, Paquette Rapids. One of
Salter’s original specimens of Ctenodonta contracta. Oblique dorsal view of
right valve showing lunule, escutcheon, and silicified replica of ligament. See
also fig. 54. Figure 73. GSC 1171m, Paquette Rapids. One of Salter’s original
specimens of Ctenodonta contracta, Oblique dorsal view of left valve showing
lunule, escutcheon, and dentition. See also figs. 40, 60, 76. Figure 74. YPM 22999,
Paquette Rapids. Oblique ventral view of interior hinge region of right valve
showing anterior and posterior pedal muscle scars and dentition. See also fig. 56.
Figure 75. Lectotype of Ctenodonta contracta Salter, GSC 1171b, Paquette
Rapids, figured by Salter, 1859, as figs. 5 and 5a of pl. 8. Ventral view of left
valve interior showing dentition, adductor muscle scars, strong rounded anterior
and posterior pedal muscle scars, and faint additional pedal? muscle scars
below posterior dentition. See also figs. 39, 61, 67, 80. Figure 76. GSC 1171m,
Paquette Rapids. One of Salter’s original specimens of Ctenodonta contracta.
Oblique ventral view of left valve interior showing dentition and posterior ad-
ductor and pedal muscle scars. See also figs. 40, 60, 73. Figure 77. YPM 22980,
Paquette Rapids. Ventral view of right valve interior showing adductor muscle
scars, strong rounded anterior and posterior pedal muscle scars, and faint addi-
tional pedal? muscle scars below posterior dentition. See also figs. 21, 47, 55, 79.
Figure 78. YPM 23001, Paquette Rapids. Oblique ventral view of left valve
interior showing dentition, anterior adductor muscle scar, anterior and posterior
pedal muscle scars. See also fig. 59. Figure 79. YPM 22980, Paquette Rapids.
Interior view of right valve showing dentition, adductor muscle scars, and pedal
muscle scars. See also figs. 21, 47, 55, 77. Figure 80. Lectotype of Ctenodonta
contracta Salter, GSC 1171b, Paquette Rapids, figured by Salter, 1859, as figs.
5 and 5a of pl. 8. Interior view of left valve showing dentition, adductor muscle
scars, pedal muscle scars, and silicified replica of ligament. See also figs. 39, 61,
67, 75.

50S. 73
Tab

LAID

YALE PEABODY MUSEUM

oF NaTuraAL History

Number 75 May 27, 1963 New Haven, Conn.

DAVID BALDWIN, O. C. MARSH AND THE
DISCOVERY OF THE FIRST CONTINENTAL
PALEOCENE FAUNAS OF THE NEW WORLD

Ewuwyn L. Simmons

Recent reorganization of the collection of fossil vertebrates
at the Yale Peabody Museum has brought to light material
for establishing a hitherto unreported chapter in the history
of western paleontological exploration—the time, place and
circumstances of discovery of the first known land mammals
of the New World Paleocene.’

David Baldwin, who was to make this outstanding discovery
while in the employ of Professor O. C. Marsh of the Yale College
Museum [Yale Peabody Museum], was first recommended to
Marsh in 1876 by Lieutenant W. L. Carpenter. In the previous
year both Carpenter and Baldwin, who held a position as
packer, had taken part in paleontological activities of the
Wheeler Survey of New Mexico on which expedition Professor
E. D. Cope of Philadelphia was also present. According to
Schuchert and LeVene (1940:179), Baldwin ‘‘began to collect
for Marsh in New Mexico in May, 1876, and continued inter-
mittently until 1880. Marsh was then concentrating on Eocene

‘Research reorganization of these collections was supported by National
Science Foundation grants 14255 (1960) and GB-247 (1962).

2 Postilla Yale Peabody Museum No. 75

O4

did not please him; moreover, with

mammals and the material of that age sent in by Baldwin
boxes and 20 packages

his customary absorption in the work at hand he failed to
appreciate the value of the bones (14 boxes) collected by
Baldwin from red beds, supposed to be Triassic age but later
determined to be Permian. The terms under which Baldwin
was to work were evidently not clearly defined by Marsh at
the start, and a long wrangle resulted. The two finally agreed
to submit their differences to Lieutenant Carpenter.”

Eventually Carpenter arranged a settlement which was satis-
factory to both, but Baldwin clearly had been irritated by
the erratic nature in which he received payment from Marsh
in support of his field activities and in 1880 shifted his employ-
ment and allegiance to Professor E. D. Cope. Baldwin then
sent his collections made in the region in 1880-1881 to Phila-
delphia which formed the basis of the first report of the Paleo-
cene (Lower Eocene) faunas of New Mexico, published by
Cope 1881la and b). Between 1880 and 1888 Baldwin collected
for Cope, and his work resulted in the Cope Collection of
Paleocene mammals now located at the American Museum of
Natural History, New York. Serving as it did as the basis
for Cope’s fundamental and extensive contributions on the
nature and initial differentiation of Cenozoic Mammalia, this
remains one of the most important collections of fossil Mam-
malia ever assembled, and it stands as a fitting memorial to
Baldwin’s diligence in the field.

While still working for Marsh, from July 1879 through
the remainder of that summer, Baldwin began to find and
send in to Marsh teeth of middle Paleocene mammals from
the area which he had previously mapped and recovered mam-
malian bone fragments. His correspondence with O. C. Marsh,
now preserved at the Peabody Museum as part of the extensive
archive of Marsh’s correspondence, makes it possible to learn
much about his activities in opening up this great fossil field.

Although its importance was not recognized at that time,
this discovery came as the culmination of three years’ paleon-
tological exploration of the San Juan and Gallinas River
drainages by Baldwin. During 1876, 1877 and 1878 Baldwin

May 27, 1963 Discovery of New World Paleocene 3

was collecting almost continuously for Marsh, working par-
ticularly in Wasatch Eocene deposits of the Almagre and
Largo formations in the drainage of Gallinas River, New
Mexico. From these beds he sent in to New Haven a consid-
erable collection of early Eocene Mammalia including much
Coryphodon material. Nevertheless many of the specimens were
fragmentary and incomplete and Marsh apparently was not
particularly impressed by their quality. Very little was done
with these shipments, many of which have only recently been
sorted and identified.

An innovation relating to the history of recovery of fossil
vertebrates was Baldwin’s adoption in 1876 of the methed of
enclosing fragile fossil specimens in a clay paste prepared
on the site of collection. A specimen of Coryphodon molestus,
YPM 15239,° collected by Baldwin on Nov. 29/30, 1877 shows
a further elaboration in that the clay-coated jaw was tied up
between two cradle-lhke, hollowed and hand-carved boards
(prepared by him) which effectively bandaged it for shipment.
A note with the specimen by Baldwin states “N.B. Lower jaw
done up in mud and boards in bottom of sack be careful very
rotten.” As far as can be determined from the Yale Peabody
Museum records this is the first fossil vertebrate to have been
housed for shipment in a manner foreshadowing modern meth-
ods. A letter from another of Marsh’s collectors dated in the
same year (1877) suggested that a coating of plaster of Paris
would be useful in holding bones together during removal from
a hard matrix in the field (see Schuchert and LeVene, 1940:
175). With these two suggestions in mind it only remained
for Marsh to instruct his collectors to bandage fossil speci-

mens in plaster jackets in the field—an invention for which
he took (and is often given) credit (Schuchert, 1939:15).
By the spring of 1879 Baldwin had prepared a map of the
reck types and river drainages of the San Juan Basin, New
Mexico (Figure 1) and had begun collecting in horizons, which
he regarded as Cretaceous, underlying the “Coryphodon beds.”
Actually these deposits were cf Paleocene age but this could

2 Abbreviations used in this paper: YPM, Yale Peabody Museum, New
Haven; AMNH, American Museum of Natural History, New York.

4 Postilla Yale Peabody Museum No. 75

hardly have been realized at that time by either Marsh or
Baldwin, inasmuch as continental deposits and faunas of this
epoch were then entirely unknown in North America. During
this period of his explorations David Baldwin shipped mate-
rials to Yale both from Animas City, Colorado and from
Abiquiu a small town on the Rio Chama about twenty-five
miles due north of Los Alamos, New Mexico. In June, 1879
Baldwin sent the following geological observations to his em-
ployer in New Haven.

Animas City Colorado

June 16th, 1879
Prot. ©: C. Marsh

Dear Sir:

Inclosed send you a list of the bones I send you by
mail today. They are mostly weathered specimens. The
mammals entirely so. In April I sent you the horizons of
this basin cut in paper [figure 1]. The beds in which I
have found these mammalian remains extend no farther
west than about haff way between the Animas and La
Plata rivers and their northern limit is about ten miles
South of this place. They show themselves about twenty
miles down the Rio Puerco from the head on the west side
of Naceimento Mountain and from that point westward
form a nearly continuous wall of bluffs nearly to the East-
ern line of the Navajo Reservation. They do not extend
to the road from Santa Fe to Fort Wingate. That road
for the first forty miles west of the Puerco Crossing is on
the Marine Cretaceous in part though mainly upon the
Lignite and the Strata containing the reptillian remains on
the San Juan river. These mammalian beds do not extend
as far west as the Eastern line of the Navajo Reservation
on the south side of the San Juan river but they extend
far beyond what I have (untill this winter) considered as
the limit of the Eocene basin. On the Gallina creek side
to my mind everything is in confusion. I cannot believe
that I have ever sent you a fossil from those beds from
there. In your address at Nashville you spoke of the
Coryphodon beds as being the oldest known Eocene and it
was that statement that showed me that I had probably

May 27, 1963 Discovery of New World Paleocene 5

the best field in the world in which to look for Cretaceous
Mammals it being a point where the oldest Eocene rested
upon the newest fresh water Cretaceous. But I do not
know but what you have had some other one sending you
fossils from the New Mexican Eocene and probably these
beds have been known to you for three or four years past.

Please tell me if I have ever sent you anything from
the Coryphcdon beds before. Are these last fossils from
the Coryphodon beds?

I have not found any small reptiles above the coal
yet though fragments of large bones are abundant. Please
write to me at this place. I will work on these mammalian
beds two or three weeks or perhaps a month before going
back to Abiquiu.

Very truly yours,

D. Baldwin”

During the following month Baldwin continued to collect
along the San Juan river and its tributaries the Animas and
Canon Largo in the new extension of the ‘‘Eccene basin”
which he had found, still supposing that these exposures were
of Cretaceous age. Marsh’s replies to Baldwin’s letters dur-
ing this pericd are not preserved, but it seems likely from the
questions asked by Baldwin in the above letter of June 16, 1879
that he had not troubled to provide his collector with much
information as to the nature of the fauna he was securing.

After searching in the New Mexican Paleocene for several
months Baldwin at last succeeded in recovering mammal teeth
which he sent to Marsh. In his letter of July 12 he discussed
the occurrence of other fossils in these beds; his washing of
sediments in an attempt to recover mammalian teeth may be
the first time this technique, now widely applied, was used in
America.

Animas City Colorado

July 12, 1879
“Prof. O. C. Marsh:

Enclosed you will find list of fossils sent to-day by
mail.

6 Postilla Yale Peabody Museum No. 75

I have found fragments of large Reptillian bones in
the lower whitish beds resting upon the Reptilian Strata
above the lignite and within 150 ft. perpendicular of the
hard sandstone capping these lower whitish beds (See cut
paper horizons sent you April 22nd, 79). This fact I sup-
pose brings the Cretaceous to the hard sandstones capping
the lower whitish beds.

A man living on the San Juan River has shown me
where he picked up two years ago a piece of a small jaw
having grinding teeth. He says it looked like the jaw of a
man or monkey but says the teeth were smaller than A &
B which I sent you today. The place where he found it
was in the Reptillian horizon below the lower whitish
beds the same horizon as the bones send in Box 2 1879
[YPM accession no. 1297] and about 400 yds. distant.
I have packed dirt from the place where he said he found
it 8 or + miles to the San Juan river and panned it out
carefully to see if I could find a speck of a tooth or any
fragment of small bone but could find nothing. It was a
Cretaceous mammal but he has lost it. I have hunted for
days and days in that vicinity and have found large rep-
tiles in abundance but no small ones and no mammals.

I shall work my way from here down the Animas and
up Canon Largo to Abiquiu where I will try the Red
rocks for a while.

Please write to me at Abiquiu and also please send
me a check for $200 as I shall greatly need it.

Very truly yours,
With the exception of the 2nd part of Description of
Jurassic fossils I have received nothing from you since
yours of Nov. 2, 1878.

D. Baldwin

I shall order my mail to be sent from here to Abiquiu.”

To this letter and a succeeding one of August 4th are
appended the following field numbers, locality data and de-
scriptions sufficient to confirm the identity of the Paleocene

May 27, 1963 Discovery of New World Paleocene 7

mammal teeth concerned which also still retain their original
field designation slips (A and A, etc.).

List of Registered Package No. 6
Animas City Colo. July 12—1879
Package contains AX A,A&B,A&X&C,AXD, A&E [YPM
accession no. 1247 |
A. & A [YPM 11887] found west side Animas River, Taos
Co., New Mexico, June 21st, 1879, near Cox’s Ranch.

A & B [YPM 14459] one molar in piece lower jaw. Found
scuth side San Juan River, Taos Co., New Mexico, June
28th, 1879, First Canon west of Canon Largo.

A & C [YPM 14476] found on La Plata side of divide
between Animas and La Plata rivers Ute Reservation
Colcrado July 9th, 1879—two molars in cement.

A & D Ute Reservation near A and C, July 9th, 1879.

A & E Ute Reservation La Plata side of divide July 10th,
1879 Piece of lower jaw teeth broken off—unmarked.

D. Baldwin

List of bones sent by mail from Abiquiu August 4th,
1879, [YPM accession no. 1259 ].

A & F [YPM 14461] from south side San Juan River, Taos
Co., New Mexico four miles west of mouth of Canon
Largo—weathered fragments of skull mammal July 21st,
1879.

A & G [YPM 14477] south side San Juan River west of

and small bones, July

Canon Largo. One back molar
22nd, 1879.
D. Baldwin

Although the above listed localities are somewhat imexact,
any locality data at all is rather unusual for the nineteenth
century and is an evidence of the type of precision which Marsh

8 Postilla Yale Peabody Museum No. 75

consistently expected of his collectors. It must be remembered
that some specimens secured in the 1870’s (and even much
later) by other institutions cannot now be located stratigraph-
ically within the particular intermontane basin of their provy-
enance. Taking the evidence cf Baldwin’s correspondence and
the collection labels reproduced here, these initial finds of
Paleocene Mammalia appear to have come from two different
areas (Figure 1) about fifteen to twenty miles apart. The
first being approximately between the towns cf La Plata and
Aztec, perhaps close to localities worked later by Granger
in 1916 for the AMNH, while the second set of material appar-
ently came from a spot six or seven miles north of Angel Peak
on the south side of the San Juan River.

Although much of the material sent in at this time by
Baldwin is insufficient for identification, three of the specimens
of the July 12, 1879 shipment and two of the shipment of
August 4 are adequate for generic and probable specific
assignment. Although these scanty fragments provide little
of morpholegic or taxonomic interest, when taken together
they do show that these initial discoveries are of species of

ge. The tax-

Torrejonian rather than Puercan provincial a
onomic position of these specimens is as follows: 1.) YPM
11887, Periptychus rhabdodon, fragmentary horizontal rami of
beth mandibles, with damaged left P, and right M.; 2.) YPM
14459, Periptychus rhabdodon, mandibular fragment with left
M,; 3.) YPM 14461, Periptychus cf. P. carinidens maxillary
fragment with a worn right M-; 4.) YPM 14476, Tetra-
claenodon cf. T. puercensis, left maxilla with M>° (M°® larger
than most JT’. puercensis, but close to T’. puercensis AMNH
3937); 5.) YPM, 14477, Neoclaenodon cf. N. procyonoides M”

ard associated bone fragments.

The first of the above listed specimens, YPM 11887. was
described by Marsh (1894: 260) as the type of “Eohyus robus-
tus.” "This was the only specimen from Baldwin’s Paleocene
collection at the Peabody Museum to which Marsh gave tax-
cnomic attention. The latter, by 1894, had recognized that
YPM 11887 and associated materials were from ‘“‘the so-
called Puerco deposits” (actually Torrejon horizons) thus

May 27, 1963 Discovery of New World Paleocene 9

lying considerably lower stratigraphically than the Wasatch-
ian age beds to the east in which Coryphodon occurs as the
commonest guide fossil. Marsh (1877:362) first mentioned
the genus “Eohyus” from the “Coryphodon beds” but did not
designate a species name and did not describe his material
except to observe that ‘“‘these remains are clearly Suilline in

39

character, ... In 1894 he designated a species for this
Wasatchian, Eocene specimen, “Kohyus distans,”’ the type
of which consisted of a single upper right third molar figured
by him (1894: 261). In the opinion of Professor Marsh (1894:
260) Cope’s genus Periptychus was a junior synonym of
Eohyus. It is clear, however, that this is not the case. What-
ever “Eohyus distans” is (the specimen cannot now be located),
if from the “Coryphodon beds” as stated by Marsh, it cannot
take priority over Periptychus, for the latter genus does not
range into the Eocene. Moreover, Gazin (1955:10) regarded
“Kohyus” as probably indeterminate. There can be little doubt
that the “type” of “Kohyus distans” was from an Eocene
horizon, for at the time of its first mention (1877) Baldwin
had not yet begun to collect in areas which could have yielded
Paleocene materials. Consequently, both Marsh’s generic and
species designations of “Hohyus robustus’ are invalid. The
“type” (YPM 11887) is a member of Periptychus, as was
first noted by Sinclair (1914:267). The latter genus has clear
priority over Eohyus through the description of P. carinidens
by E. D. Cope (1881a:337). Moreover, YPM 11887 may
be assigned with some confidence to Periptychus rhabdodon, a
Torrejonian Paleocene species.

CONCLUSIONS

A small series of fossil mammals and other vertebrates, five
of which can be taxonomically identified with some certainty,
collected by David Baldwin for O. C. Marsh in the spring and
summer of 1879, comprise the first continental Paleocene fauna
to be discovered in the New World. Baldwin’s pioneer collect-
ing activities in Northwestern New Mexico are traced as they
relate to this major discovery from his field label records,
maps and correspondence preserved at the Yale Peabody Mu-

10 Postilla Yale Peabody Museum No. 75

seum of Natural History. The species “Hohyus robustus”
Marsh (1894) based on one of the specimens discovered in
1879 by Baldwin is a junior synonym of Periptychus rhab-
dodon, a Torrejonian, Paleocene condylarth.

REFERENCES

Cope, E. D., 188la. Mammalia of the Lower Eocene beds. Am. Nat., p. 337-
338.

Cope, E. D., 1881. On some Mammalia of the Lowest Eocene Beds of New
Mexico. Proc. Am. Philos. Soc. 19: 485-495.

Gazin, C. L., 1955. A review of the upper Eocene Artiodactyla of North
America. Smiths. Mise. Coll., 128(8): 1-96.

Marsh, O. C., 1877. Introduction and succession of vertebrate life in
America. Am. Jour. Sci. ser. 3, 14: 338-378

Marsh, O. C., 1894. Description of Tertiary artiodactyles. Am. Jour. Sci.
AT: 259-274.

Schuchert, C, 1939. Othniel Charles Marsh 1831-1899. Biog. Mem. Nat.
Acad. Sci. 20 (1): 1-78.

Schuchert, C. and LeVene, Clara M., 1940. O. C. Marsh, Pioneer in Paleon-
tology, Yale Press, New Haven, ix + 541 p.

Simpson, G. G., 1951. Hayden, Cope, and the Eocene of New Mexico. Proc.
Acad. Nat. Sci. of Phila. Vol. c III. p. 1-21, April 25, 1951.

Sinclair, W. J., 1914. A revision of the bunodont Artiodactyla of the middle
and lower Eocene of North America. Bull. Amer. Mus. Nat. Hist. 33:
267-295.

Figure 1.

The first geological map of the North American Continental Paleo-
cene, prepared by David Baldwin in April, 1879. The areas of two dis-
coveries of Paleocene Mammalia made by him during that year are indi-
cated by stipple.

YPM 1447]

YPM 14476

YPM 11887

May 27, 1963 Discovery of New World Paleocene

11

Figure 2. The initial five Paleocene mammals discovered in North

America. Damaged areas stippled.

YPM 14477. Neoclaenodon ct. N. procyonoides M® and associated
bone fragments.

YPM 14461. Periptychus ct. P. carinidens maxillary fragment
with a worn right M?’.

YPM 14476. Vetraciaenodon ct. T. puercensis, left maxilla with
M** (M® larger than most 7. puercensis, but close to T. puer-
censis AMNH 3937).

YPM 11887. Periptychus rhabdodon, fragmentary horizontal rami
of both mandibles, with damaged left P, and right M,.

YPM 14459. Periptychus rhabdodon, mandibular fragment with
left M,.

oe

YALE PEABODY MUSEUM

oF Naturau History

Number 76 October 30, 1963 New Haven, Conn.

MOLT AND BREEDING IN POPULATIONS

OF THE SOOTY TERN STERNA FUSCATA

N. Puitie ASHMOLE

Epwarp Grey [Nsrirute, Oxrorp, ENGLAND*

I have recently discussed (Ashmole, 1963) the breeding cy-
cles of the Sooty Tern Sterna fuscata in all parts of its range.
Although there are many areas for which adequate informa-
tion is lacking, it appears that this species in different locali-
ties shows three different types of breeding cycle: namely, every
twelve months, every nine and one half months, and every six
months. In the places where breeding occurs every six months
it has not yet been shown whether the same individuals breed
in successive breeding periods, but I am at present carrying
out work on Christmas Island (Pacific Ocean) designed to
determine this.

During study of a large number of Sooty ern skins in
United States and British museums I cbserved an unexpected
difference in the state of the primaries between samples of
birds from populations where breeding is annual and from
populations where breeding occurs every six months. This dif-
ference, described in the present paper, suggests that Sooty
Terns in populations where six-monthly breeding occurs have

* Present address: B. P. Bishop Museum, Honolulu 17, Hawaii.

2 Postilla Yale Peabody Museum No. 76

evolved special modifications of the species’? normal molt pro-
gram, which enable them to breed at approximately six-month
intervals and yet to replace their remiges often enough to
maintain reasonable flying efficiency.

Sooty Terns in most populations appear to have a pattern
of molt similar to that in a number of other tern species: that
is, they undergo a postnuptial or “basic”! molt after breeding
in which all feathers are replaced, and a prenuptial or “alter-

”

nate’ molt shortly before the onset of breeding which does not

involve the primaries or secondaries. Although Dwight (1901)

39

says “The Terns undergo two complete moults in a year. ..,

I know of no tern species for which there is adequate evidence
that all the remiges are replaced twice each year. In some
species none of the remiges are replaced more than once, while
in others the inner primaries are replaced twice, the outer ones
only once (Ashmole, in prep.). In the Sooty Tern I have found
no indication that any of the primaries are replaced more than
once between one breeding period and the next.

As in other terns, molt and breeding in the Sooty Tern are
more or less mutually exclusive. (However, Brown Noddies
Anous stolidus on Ascension Island and perhaps elsewhere
sometimes breed and molt at the same time (Dorward and Ash-
mole, 1963).) Few museum specimens are accompanied by infor-
mation as to whether the individuals were involved in breeding
activities when collected, but Table 1 shows that most Sooty
Terns collected on breeding grounds in all parts of the world
have complete sets of primaries and rectrices. From some locali-
ties there are a few birds just completing the replacement of
their primaries (primary molt scores 98 and 99), while from
some breeding stations there are birds which have recently
started a molt (primary molt scores nearly all below 30).°
I have already shown (Ashmole, 1963) that on Ascension Is-
land individuals complete a molt before starting to breed, but
some at least start their postnuptial molt before their chicks

',° These terms are those advocated by Humphrey and Parkes (1959).

*“Primary molt scores” are stages on the scale from 1 (= molt of pri-
maries just started) to 99 (—molt of primaries almost completed); for
details of the method of scoring see Ashmole (1962).

Oct. 30, 1963 Sooty Tern Sterna Fuscata 3

Taste 1. Molt of primaries and rectrices of Sooty Terns collected
on breeding grounds in different areas.?

7——- Primaries ——, —Rectrices—

Pro- Range in Pro-
Number portion — scores of portion
of birds — of birds molting of birds
Locality examined molting birds? molting
GuLF oF Mexico
AND W. INpIEs
Gorpus Christi; Lexas) <7... 17 0 _ 0
Windaiinl IISS Sogcansis oeoudaa 12 0 — 0
EavuaroriAL ATLANTIC
Fernando Noronha ........ 33 :2i1 2-28, 98 1 li55
PASCENSIONMs 2 oaie sie ete eielc cies 107 20 1-20 13
SoutH ATLANTIC
Trinidade /Martin Vaz .... 11 0 a= 0
INpIAN OcEAN
Waccadiveplssm sae eee 11 18 Dink, .09
Nortu Paciric
ILEN GEM Goocopeauoooeacbeoc 39 03 98 05
iepisabelas Mexico’ oo... 5-4. 11 18 Qala 09
TERS OCORLOMM ET eee coi 15 .20 4, 24, 48 .20
Clipperton lero 24 04 99 13
EavuatoriIaAL Paciric
@hristmasels terre ee ee 16 19 2, 98, 98 06
Culpepper /Wenman,
Galapacosiancrcrr ceric 10 30 All 98 10
Soutn Pactric
WGorderlowewlen cme clases + cer 13 0 — “20
INorfolkilivss: sccn Socio: stele 10 10 4 10
Kermadecslisaercticne sneer 18 0 a 0
INGE ~ SooobbdosonDoUgmaUG 11 0 a 0
SivonOvalegeerer cet cmccniceos 40 30 1-24 15
Kauehi, Tuamotu Arch. ... 26 04 98 .O4
Mar GguesasS vanities 19 21 All 98 05
@enOe He anton ccs tremors 13 0 5 0

Notes. 1. Only breeding localities from which I have examined at least ten
birds are included in this table.

2. See footnote in text for explanation of “primary molt scores.”

4 Postilla Yale Peabody Museum No. 76

become independent. Both on Ascension and elsewhere body
molt of Sooty Terns occurs almost entirely while the birds are
absent from the breeding grounds; I have so little information
about it that I shall not consider it further in this paper.

PRIMARIES

Examination of molting birds on Ascension, and of skins of
molting individuals, shows that in the Sooty Tern molt of the
primaries normally starts with the first (innermost) feather
and progresses outwards to the tenth (outermost long pri-
mary). Sooty Terns when breeding should thus have the inner-
most primaries oldest and the outer ones progressively newer,
the whole series forming a smooth sequence. This was found to be
the case in 86 per cent of all skins of birds collected on the breed-
ing grounds and not in process of molt. However, in some birds
there are striking differences in the condition of adjacent pri-
maries ; one finds a sudden break in the normal age-sequence
part way through the series. This I have called a ‘‘discon-
tinuity.””* It should be emphasized that the discontinuities were
not caused merely by molt in progress when the bird was col-
lected ; most individuals were not molting at all, and in the few
which had recently started a molt the arrangement of old and
new feathers at the discontinuity could not be explained as a
result of the molt then in progress. It was evidently the re-
sult of an unusual molt sequence in the past, followed by a ces-

sation of molt prior to breeding.

* Like many other species (Dwight, 1901) Sooty Terns have a pale “frost-
ing” or “silvering” on the dark primaries and secondaries, which gradually
wears off, thus making it easy to detect large differences in the age of
adjacent feathers. I have recorded discontinuities only when the difference
in the condition of adjacent feathers was sufficiently striking for there to
be no doubt that they were of very different age. In badly set specimens
it is difficult to assess the relative ages of the small inner primaries,
especially as they tend to be protected from wear by the overlying sec-
ondaries. I may therefore have overlooked relatively new innermost pri-
maries in some birds, and the figures for the occurrence of discontinuities
between primaries 1 and 2 must be considered as minimum ones. Discon-
tinuities further out in the series are not likely to have been overlooked,
and there were few birds in which I was doubtful whether the feathers
had been molted in regular sequence.

Oct. 30, 1963 Sooty Tern Sterna Fuscata 5

Discontinuities are found at all points in the primary series,
but Table 2 (from which molting birds are excluded) shows
that they are not distributed at random. Nearly all popula-
tions (see Table 3 for details) contain a small proportion of
birds with first primaries much newer (occasionally much
older) than the second, but in most populations (grouped in
the bottom row of Table 2) discontinuities at other points in
the primaries are rare. The sample from Ascension is separated
since not only does it contain an especially large proportion of
birds with discontinuities between primaries 1 and 2, but it also
has a number with discontinuities between primaries 2 and 3;

collected on Ascension, on the Phoenix and Line Islands, on Bedout Island,

Taste 2. Distribution of discontinuities in the primaries of Sooty ‘Terns

and in the other localities mentioned in Table 3.

No. and %
Number (above) and percentages — of wings

Number (below) of wings with discontinu- with no

of wings ities at each point in the primaries— _ discon-
Locality examined 46% % 4% % & BW %% %o_ tinuities
Ascension 164 No. 20 8 1 2 0 0 0 0 0 133
Island (82

birds) To 125, 0:6 110) 0" 10) <0) 0 81
Phoenix 64 Nox Zin ON 2562) o GS) ie 12 34.
and Line (=32
Islands birds ) % 3 @ 8 8) co © 2B} il iy) 53
Bedout 10 INOw 27210) 100) OO A 2 2
Island (=5

birds)
Other 946 No. 59 7 8) pp 45 Me ayo) 854.
areas (=473

birds) % 6 0.7 0.6 05 05 04 0 0.3 0.5 90

Notes. 1. Molting birds are not included.

2. For each locality, the upper figures are the numbers of wings which
show discontinuities at each point in the series of primaries. Wings
are used rather than birds since the two wings on a single bird
sometimes have discontinuities in different places.

3. The lower figures show the number of discontinuities at each posi-
tion as percentages of the number of wings examined. Since there
are sometimes two discontinuities in one wing, and both are in-
cluded in the figures given, the percentages total more than 100
in some cases.

6 Postilla Yale Peabody Museum No. 76

these are rare in other populations. Samples from two areas
only contain an appreciable number of birds with discontinui-
ties further out in the series than the second primary. One of
these areas includes the Phoenix Islands and Line Islands in
the central equatorial Pacific, while the other is represented by
Bedout Island off the northwest coast of Australia. The distri-
bution of discontinuities in the wings of birds from these areas
(Table 2) seems certainly to indicate that many birds in these
populations have a very different molt program from the birds
breeding on Ascension and in the other localities from which
specimens were examined.

Table 3 shows that, from population to population, there is
no correlation between the frequency of occurrence of disconti-
nuities between primaries 1 and 2 and that of discontinuities at
other points in the series. Thus while 21 out of 38 birds from
the Phoenix and Line Islands have discontinuities among pri-

Taste 3. Occurrence of discontinuities among the primaries in adult Sooty Terns
from different breeding areas.

Occurrence of Occurrence of
discontinuities only discontinuities
between primaries among

———-I and 2 =~ Z primaries 2-10—
Number Number
Number (and pro- Number (and pro-
of portion ) of portion )
specimens — with dis- specimens — with dis-
Geographical area available continuities available continuities
GuLF oF MExIco AND W. INDIES ...... 74 13 (.18) 80 4 (.05)
(incl. Corpus Christi (Texas),
Virgin Is.)
FQuatrortaL ATLANTIC
Fernando Noronha, Rocas Reef ... 34 2 (.06) 38 4 (.11)
FNRADSOOD. Ils SaoooooacbesoocpaudDe 82 10 (.12) 106 6 (.06)
SOUMED PAGE AINIDIC ees perlite) srreucKeiens 19 ORG) 19 OMG)
(Trinidade/ Martin Vaz,
St. Helena)
INDIAN TOGEAN screeners ete ores 21 2 (.10) 24 1 (.04)
(incl. Gloriosa, Seychelles,
Rodriguez, Laccadive Is., Chagos)
NorRTHWEST AUSTRALIA .........0000- 5 1 (.20) 5 3 (.60)

(Bedout I.)

Oct. 30, 1963 Sooty Tern Sterna Fuscata

Taste 3 (Continued)

Occurrence of
discontinuities only
between primaries

————1 and 2

Geographical area

available

Number
Number — (and pro-
of portion )
specimens — with dis-

continuities

INORIENWAES TE EPA CTBEG oinlstal-ds acu sen=ie=) alate 13 3 (.23)
Paracel Is., Philippines, Ryu
Kyu Is., Bonin Is., Marianas)
NorteH CENTRAL PACIFIC) ...........: 79 2 (.03)
(Wake, Hawaiian chain incl.
Laysan, Johnston)
INJORIMENEAS TN yA CIFIC i. .jeleleioie)- 1 shel = evel 61 1 (.02
(Revilla Gigedo Is. inel. Socorro,
Clipperton, Lower California,
west coast of Mexico)
EQuaToriIaAL Paciric
NVIGAR SERIA TPS Se Foye ies Vel oueiuetelete sass 5 1 (.20)
Puoentrx Is. (Howland, Baker, 9 (i)
Canton, Enderbury, Phoenix)
Line Is. (Palmyra, Christmas, 23 0 (—)
Jarvis, Malden, Starbuck)
Gaxaracos Is. (Culpepper/Wenman) — 10 E10)
SOULEWESI IVACTNIGH ss. ons. see +s 2 ve 53 4 (.07)
(Lord Howe, Norfolk, Kermadec Is.,
Fiji, Tonga, Samoa)
SouTHeAst Pactric
Cook Is., Society Is., Tubuai Is., 72 3 (.04)
Tuamotu Is. incl. Kauehi,
Marquesas Is., Oeno, Henderson,
Ducie, Easter, San Felix
SWOOTOD, VONGATEVA, .\e1se.215 21536 902 34 0 (==)

Occurrence of

discontinuities
among
-———primaries 2-10
Number
Number (and pro-
of portion )
specimens — with dis-
available continuities
13 1 (.08)
82 0 (—)
63 2 (.03)
5 1 (.20)
11 9 (.82
27 12 (44)
10 1 (C10)
55 1 (.02
73 1 (.01)

42 By (0)

Notes. 1. Only adult birds collected on the breeding grounds or within a few miles of
them are included in this table.

tho

Birds which were in process of primary molt are excluded, with two exceptions:

(a) birds whose tenth primaries only were growing have been included, and
(b) birds whose first and/or second primaries only were growing, have been
used in the right-hand section but not in the left-hand section of the table: this
accounts for the differences between the columns showing “Number of speci-

mens available” in the two sections.

3. Where ten or more birds were available for examination of primaries 2-10,
from one island or from a group of islands within a circle of radius 25 miles,
the name of the island or group is italicized.

8 Postilla Yale Peabody Museum No. 76

maries 2 to 10, only 1 out of 382—less than the average propor-
tion—have discontinuities between primaries 1 and 2. Only on
Ascension, as already mentioned, are there an appreciable num-
ber of birds in which primary 2, together with 1, is strikingly
different in age from the rest. Probably in this case birds with
the first and second primaries very different in age from the
next outwards should be classed with those in which only the
first primary is affected.

I suspect that discontinuities far out in the series are nor-
mally produced under quite different circumstances from those
between primaries 1 and 2. I have already suggested with re-
spect to the Ascension population (Ashmole, 1963) that birds
with first (or first and second) primaries newer than the next
outwards may be young birds breeding for the first time; in im-
mature Sooty Terns successive sequences of primary replace-
ment often overlap, so that as one sequence is nearing comple-
tion with the growth of the outermost primaries, another se-
quence is starting with the innermost ones. If molt stops for
breeding at the completion of cne sequence, the outermost
feathers will be new, but so may be the innermost ones, with
a discontinuity outside them.

It is not possible to explain in this way the extremely high
incidence of discontinuities (Table 3) among the outer pri-
maries in the samples from the Phoenix and Line Islands, while
the small sample from Bedout Island may also require a dif-
ferent explanation. It can be no coincidence that it is on cer-
tain of the Phoenix Islands and Line Islands, alone of the
places from which I have examined an appreciable number of
specimens, that Sooty Terns are known to have two breeding
periods each year (Ashmole, 1963).

It has been argued (Ashmole, 1963) that the Sooty Terns on
Ascension are breeding as often as they can—that breeding,
followed by a complete molt, occupies about nine and a half
months. But if this is the minimum time needed by the Ascen-
sion birds, it is difficult to see how the birds in the Phoenix and
Line Islands (where there are two breeding periods each year)
could breed and undergo a complete molt in a period of only
six months. It is therefore not surprising that it has been
tentatively suggested in the past (Richardson and Fisher,

Oct. 30, 1963 Sooty Tern Sterna Fuscata 9

1950; Hutchinson, 1950; Chapin, 1954) that in the areas
where breeding occurs every six months, different populations
cf birds might be involved in successive breeding periods, so
that each individual would breed only once a year. I also felt
that this must be the explanation, until I examined specimens of
the Sooty Terns from the islands concerned, and found that
many of the breeding birds had some old and some new pri-
maries, and had evidently not undergone a complete molt be-
tween breeding periods.

As it has been pointed out, Sooty Terns in other parts of
their range, where breeding occurs only once each year, have a
straightforward complete replacement of all their remiges after
breeding. If the individuals on the Phoenix and Line Islands
were also breeding only once a year, how could one explain the
fact that many of them do not have a complete set of new re-
miges when they start breeding? The most reasonable hypoth-
esis seems to be that at least some individuals breed in succes-
sive breeding periods and replace only some of their wing
feathers in the short interval in between. It is likely, however,
that no one individual breeds in every breeding period.

The sample from Bedout Island (N.W. Australia) is very
small, but three cut of the five birds show discontinuities among
primaries 2-10. Sooty Terns have so far only been recorded as
breeding in autumn on Bedcut. However, it would not be sur-
prising if they were found to have two breeding periods there
each year, since Serventy (1952) has shown that they breed in
autumn on islands to the north of Bedout, but in spring on
islands further south (see Appendix in Ashmole, 1963, for
details). Bedout is at about the latitude at which several other
species of sea birds change from autumn to spring breeding,
and some of them are already known to breed in both seasons,
on Bedout and certain other islands (Serventy, 1952

Of the other localities from which there are reasonably large
samples, Fernando Noronha (where the interval between suc-
cessive breeding periods has not yet been determined) has most
birds with discontinuities among primaries 2-10, but the pro-
portion is far lower than in the samples from the Line Islands
and the Phoenix Group. Of the latter, the Line Islands have the
lower proportion, but even this is significantly higher than

10 Postilla Yale Peabody Museum No. 76

among the Fernando birds (P <.01). It is evident, therefore,
that the populations from the Line and Phoenix Islands show
peculiarities which are almost entirely absent in the other popu-
tions sampled. This is expected on my hypothesis that peculiari-
ties in the molt normally occur only in individuals which breed
in successive breeding periods about six months apart.

Table 3 shows that although the proportion of birds with
discontinuities among primaries 2-10 is far higher in both the
Phoenix and Line Islands than in any other area apart from
Bedout Island, it is appreciably higher in the small sample
from the Phoenix Group than in that from the Line Islands.
This difference as it stands is statistically significant (at the
5 per cent level), but both samples are heterogeneous (birds col-
lected from several different islands, in different years and at
different stages in the breeding periods), so I doubt whether :t
is meaningful.

Examination of the precise arrangement of new and old
feathers in the primaries of birds from the Phoenix and Line
Islands and from Bedout Island, may help towards an under-
standing of the molt program in these populations and of the
way in which breeding and molt are interrelated; some actual
examples are therefore given below. In the examples, N repre-
sents an apparently brand new feather, (N) a newish one, (O)
an oldish feather, and O an old, worn feather. Where a series
of adjacent feathers have been replaced in a regular sequence,
they may grade from one category to another; in such cases
the two terminal members of the series are joined by an arrow
headed towards the newest feather. The figures 1, 2, 3 and 4
represent successive stages in the growth of a feather (see Ash-
mole, 1962 :255).

1. Suvoroyv Ne EN N Oo. OF oO O O O O
Rie IN| EN N OF O03 0 Oe JOO) O

2. Fernando Noronha L. 1 (N) (N) (N) (N) (N) (N) (N) (N) O
Be GND GNOL GN) CN) G0, GO CO GORY

3. Jarvis I. Ear GN) NON) (CN LOO” HOF Oro
(Line Is.) R: GN) (NY (N) CN) O(N) Ol OS TOF TOMO

Oct. 30, 19638 Sooty Tern Sterna Fuscata Re

4. Enderbury I. IGN) Sz IN
(Phoenix Is.) REN) INO

5. Fernando Noronha L. O >1(O) EN) END CNY) END)
EO) eee ea (((0))) GND) (END) ND) QING)

6. Baker I. L.. (0). (0) (0), NL NO), (0) ©) CO) (O)
(Phoenix Is.) Re (0) (0). (O)) NIN (0), (0) "(0)).(O), CO)

7. Jarvis I. L. 4 ENS) GN) CO) CO) CONE GN) FCN EGN)
(Line Is.) Rese OND ND) ENO) (O) ECO) ACO) TENDER GND REND

8. Palmyra I. Ea Ne IN EN, Ne NG (O)k(O) Ne Ne SN
(Line Is.) FUNG Ni ee N oe NG ae Ni ING NIG SN) DSN

9. Christmas I. (0) aN One Oe O SSO EN) GN))
(Line Is.) ER (O)) fe (GN) tO) ©) ©) IND) ee)
10. Enderbury I. TONG) IN) (CO) CO) Re Ni CO)

(Phoenix Is.) R. (N) NN (O) (©) ©) N (CO)

In most birds with discontinuities among primaries 2-10, both
from areas where breeding occurs annually and from those
where it occurs every six months, the feathers inside the dis-
continuity are newer than those outside it (see examples 1-4).
This condition is the one which would arise if a normal sequence
of primary replacement stopped part way through the series.
Patterns of the type shown in example 5, in which there is a
definite discontinuity between old feathers on the inside and
newer ones on the outside, would result if, after primary re-
placement had stopped part way through the series for breed-
ing, it later started again where it had left off, and then con-
tinued outwards.’ However, the condition shown in example 5
is uncommon in all Sooty Tern populations. This suggests that
normally, when a bird with the inner feathers newest (as in
example 1) starts to molt again, the innermost primaries,
rather than those immediately outside the discontinuity, are
replaced first; this is in fact what is occurring in examples 2
and 3.

° Something equivalent to this certainly occurred in the replacement of the

secondaries of some Black Noddies Anous tenuirostris on Ascension Is-
land (Ashmole, 1962).

12 Postilla Yale Peabody Museum No. 76

The other examples shown are of the more complex situation
in which there is more than one discontinuity among the pri-
maries in one or both wings. Patterns of this type, which are
found in a significant proportion of the birds from areas where
breeding occurs every six months, have not yet been found
among birds from annual-breeding populations. They could not
be produced during a molt program in which each primary was
always replaced in regular sequence outwards through the
series, but they could arise if an incomplete primary replace-
ment was succeeded, after the breeding period, by another
incomplete molt, and if feathers replaced late in the first of
these molts tended to be skipped during the next. In this case
the bird in example 1, if it underwent another incomplete molt,
might in the next breeding period be in a condition similar to
example 6, for instance:

1 2 3 4 5 6 7 8 9 10

LaCO)(@) CO), Ne ONE NPN Om Ono
Re (O)¥(@O) 16@) 2 ON) OND UNE Ol aOn nO

Another partial molt could lead to conditions comparable to
those in examples 7-10, for instance:

Nee N: Ni (©) CO) (O).2(O) Ne Nea

BeoN. Ni oN (O)(O) (ONO), No EN, EN
or

LE: NN. NiO) (0), ORO)IN ON TO

RUN. Ni, N _(OjnCO), (O).CON ON, CoN a ©

It will be clear from what has been said that it is not neces-
sary to postulate a random molt sequence to account for the
complex patterns of old and new primaries found in the wings
of some Sooty Terns. The indications are that in these birds
as in all other terns molt in the primaries proceeds from the
inside outwards, but that many individuals in populations
where breeding occurs every six months fail to replace all their
primaries between successive breeding periods, and that subse-
quent molts are modified by the, presence in their wings of a
mixture of old and new feathers.

Oct. 30, 1963 Sooty Tern Sterna Fuscata 13

SECONDARIES

As in other tern species, replacement of the secondaries in
Sooty Terns starts much later than that of the primaries, but
is completed at the same time or only a little later. Replace-
ment normally starts at the two ends of the series of sec-
ondaries, and it is some of the middle feathers (often numbers
12 and 13 counting from the carpal joint inwards) which are
the last to be replaced. After the complete replacement of the
secondaries in this manner there should be no appreciable dis-
continuities within the series, but the feathers at the two ends
of the series will be oldest, and the middle ones newest. This is
in fact the situation found in nearly all specimens from most
Sooty Tern breeding colonies, including Ascension Island (Ta-
ble 4). This must imply that all the secondaries are replaced
once between breeding periods. However, in several populations

Taste 4. Numbers of Sooty Terns with discontinuities among the secondaries, in relation
to the incidence of discontinuities among primaries 2-10, in different populations.

Birds without Birds with
discontinuities discontinuities
-—among primaries 2-10— —among primaries 2-10
Number with Number with
discontinuities discontinuities
Geographical Number among Number among
area examined secondaries examined secondaries
Gur or Mexico anv W. INpres 72 0 (+2?) 2 1
PRSCENSION Ms i deseo o onesie ere TA. 1 6 1
HEN TAIN OCEAN: iy enc, shee c2)ereus @\ eis 22 Om (Gotealle)) 1 I
LPR NOS IE Sacred oes DOSE 2 2 & 6 (+2?)
‘LABS rtd Ii SR Barge acer ee ae enh ee ae 1153 Ay (=f s1/2)) 11 11
SOUMED MEA CTRIC) acisie epee aie ciaeierere 116 4 (+1?) 1 0
(incl. Marquesas,
but not Suvoroy)

SSENV OR O Voaera ny eh yor Meichoxe sspears, she loveis 26 SS G62) 3 20 (=?)

Notes. 1. Birds undergoing molt of primaries or secondaries are excluded, except for
those in which only the first primaries have so far been shed.

ho

. Additional figures in brackets, followed by question-marks, represent birds
whose secondaries show probable, but not striking, discontinuities.

14 Postilla Yale Peabody Museum No. 76

birds are found which have some secondaries much older than
the rest. I did not examine the secondaries of all specimens, and
there were a good many doubtful cases, so I cannot give reli-
able figures for the frequency of this peculiarity in all popula-
tions. However, Table 4 gives the information for those popula-
ticns from which the secondaries of a fair proportion of the
available specimens were examined.

As might be expected, the populations in which many birds
show discontinuities in the primaries (Phoenix and Line Is-
lands) also contain many birds with discontinuities among the
secondaries. However, in these populations some of the birds
without discontinuities in the primaries, nevertheless have sec-
ondaries which do not seem to have been replaced in a smooth
sequence: evidently the molt cycle is not entirely normal even
in these birds.

Among localities where discontinuities in the primaries are
rare, the island of Suvoroy, south of the equator in the central
Pacific, is the only one from which I have a fairly large sample,
in which many birds have discontinuities among the secondaries
(Table 4). In these birds some of the middle secondaries tend
to be much older than the rest, suggesting that the secondary
molt has stopped before completion. This situation invites
comparison with the Black Noddies on Ascension Island (Ash-
mole, 1962), where the primary molt was apparently never cut
short at the start of breeding, but some of the old middle sec-
ondaries, which are normally molted slightly later than the
last primaries, were sometimes retained through the breeding
period and replaced immediately after it. The occurrence of a
similar phenomenon among the Sooty Terns on Suvoroy sug-
gests that the breeding cycle there may be abnormal in some
respect, but there is very little information on the times of
breeding (Ashmole, 1963).

RECTRICES

Replacement of the tail feathers of Sooty Terns normally
starts with the outermost feathers (number 6 on each side),
the central pair (number 1) being molted next; molt probably
then continues in the sequence 2, 3, 5 and 4. It is possible that

Oct. 30, 1963 Sooty Tern Sterna Fuscata 15

in the annual-breeding populations all the rectrices are then
replaced again before the next breeding season, but in the
Ascension population there was evidence that the outer pair
alone are replaced twice (Ashmole, 1963). In the Ascension
birds the outer webs of the outermost feathers are normally
white in the breeding period, but are more often, if not always,
dark in the non-breeding period. In other populations there ts
much variation in the color of these feathers, and samples from
different populations sometimes also differ markedly. In some
areas nearly all the birds taken on the breeding grounds have
entirely white outer webs to the outer rectrices (69 out of 82
birds from the North Central Pacific), but in other places (e.g.
Fernando Noronha, the Southeast Pacific, the Phoenix and
Line Islands, and Suvoroy) the proportion is much lower. It is
likely that in the populations in which breeding occurs every
six months not even the outermost rectrices are always replaced
twice between successive breeding periods. I cannot suggest any
explanation of the different frequencies with which dark color
is present in the outer webs of these feathers in other popula-
tions.

DISCUSSION

The data presented in this paper, together with the informa-
tion on the times of breeding of various Sooty Tern popula-
tions given by Ashmole (1963), show that the schedule of
breeding and molt evolved among the Sooty Terns of the Phoe-
nix and Line Islands is remarkable both in that breeding occurs
every six months, and in that the program of molt is flexible to
a unique degree. It appears that an individual sometimes under-
goes a complete molt without interruption but at other times
replaces only some of its primaries and secondaries between one
breeding cycle and the next.

I have already mentioned that in the populations where
breeding occurs every six months, individuals which have under-
gone only a partial molt before breeding generally have the
outer primaries older than the inner ones. Since the outer pri-
maries are also more subject to wear, it is not surprising to
find some individuals with outer primaries in extremely poor

16 Postilla Yale Peabody Museum No. 76

condition while breeding. These birds are doubtless below their
maximum flying efficiency, but if the curtailed molt has enabled
them to breed in a breeding period which they would otherwise
miss, the disadvantage may on balance be outweighed. However,
it is clear that the molt program in the Phoenix and Line Island
populations, in which the inner primaries are on an average
replaced more often than the outer ones, although the latter
get more wear, is not the most efficient that might be evolved.
More birds would be close to maximum flying efficiency for
more of the time if molt always started from where it had left
off, so that the primaries were always replaced in order of age.
This evidently happens sometimes but cannot be common.

Although I have argued that the presence of discontinuities
among the primaries of breeding individuals from the Phoenix
and Line Islands implies that these birds were involved also in
the previous breeding period only six months before, I am not
suggesting that individuals breed every six months. It seems
unlikely that a pair could raise a chick successfully in one
breeding period and yet be ready to breed again in the next
breeding period only six months later. With the time required
for courtship plus incubation for a month and feeding the
young for two to three months (longer if the juveniles are not
independent as soon as they leave the colony), very little time
would be left before the next breeding period. I would guess
that birds which raise a chick in one breeding period may then
undergo a complete molt, missing the next breeding period;
this would account for the proportion of birds from the Phoe-
mix and Line Islands which appear to have undergone a per-
fectly normal and complete molt. Many of the birds, however,
losing their eggs or their chicks while fairly young, could be
ready to try again in the next breeding period, after only a
partial molt.

This reasoning is of course highly speculative, and further
discussion of the factors controlling the schedule of breeding
and molt in the Sooty Terns of the Phoenix and Line Islands
will be profitable only when we have more information on the
sequence of events in individual birds from one of these popula-
tions.

~

Oct. 30, 19638 Sooty Tern Sterna Fuscata 1
ACKNOWLEDGMENTS

This account of molt in populations of Sterna fuscata forms
a complement to my study of the species on Ascension Island
during the course of the British Ornithologists’ Union Cente-
nary Expedition (Ashmole, 1963). The present paper is a
product of my examination of a large number of skins of
Sterna fuscata during the summer of 1960, while I was the
holder of a Seessel fellowship at Yale University. I am most
grateful to Professor G. E. Hutchinson who suggested a study
of specimens of the species and who with Dr. S. Dillon Ripley
enabled me to visit Yale. I must also thank Dr. Philip S.
Humphrey, who gave me much of his time during my work in
the Yale Peabody Museum, and whose wide knowledge of molt
was a constant stimulant. My visit to Yale was made while I
was a member of the Edward Grey Institute of Field Orni-
thology, Oxford; it is a pleasure to thank the Director, Dr.
David Lack, for encouraging me to go to Yale, for helpful
discussion of my work and for reading a draft of this paper.
My wife has also made many helpful criticisms of the manu-
script.

Tern specimens were studied in the collections of the follow-
ing institutions: Peabody Museum of Natural History, New
Haven (which contains the bulk of the collections made by the
“Blossom” South Atlantic expedition, sponsored by the Cleve-
land Museum of Natural History); American Museum of Nat-
ural History, New York (which has fine series of Sooty Tern
skins collected by the Whitney South Sea Expedition) ; United
States National Museum, Washington, D.C.; Museum of Com-
parative Zoology, Cambridge, Mass.; Museum of Zoology, Ann
Arbor, Michigan; Department of Conservation, Cornell Univer-
sity; British Museum (Natural History), London. I am grate-
ful to the curators of all these collections for the facilities they
offered me. In addition, skins were generously loaned by the
authorities of the California Academy of Sciences, the Los
Angeles County Museum, the Carnegie Institution, Pittsburgh,
the Academy of Natural Sciences of Philadelphia, the Museum
of Zoology, Ann Arbor, Michigan, the Kansas University Mu-

18 Postilla Yale Peabody Museum No. 76

seum of Natural History, the Chicago Natural History Mu-
seum, the United States National Museum and the Bernice P.
Bishop Museum, Hawaii.

SUMMARY

The Sooty Tern Sterna fuscata in most parts of its range
breeds at the same season in each year, and study of museum
specimens shows that the individuals replace all their remiges
and rectrices between breeding seasons. On Ascension Island,
where breeding occurs every nine and one half months, there
is also a complete molt between successive breeding periods.
However, among birds from the Phoenix Islands and Line
Islands in the central equatorial Pacific, where breeding occurs
every six months, many individuals have “discontinuities”
among the primaries and secondaries, indicating that they
have not undergone a complete molt between successive breeding
periods. These populations have apparently evolved a uniquely
flexible molt program, such that under certain circumstances
(perhaps the successful rearing of a chick) breeding is followed
by a complete molt, but often molt stops and the bird breeds
again before all of the primaries and secondaries have been
replaced. It is suggested that because of this flexibility in the
molt, individuals are sometimes able to take part in successive
breeding periods only six months apart.

LireraAtTurRE Crrep

Ashmole, N. P., 1962. The Black Noddy Anous tenuirostris on Ascension
Island. Part 1. General biology. Ibis, 103b: 235-273.

Ashmole, N. P., 1963. The biology of the Wideawake or Sooty Tern Sterna
fuscata on Ascension Island. Ibis, 103b: 297-364.

Chapin, J. P., 1954. The calendar of Wideawake Fair. Auk, 71: 1-15.

Dorward, D. F., and N. P. Ashmole, 1963. Notes on the biology of the
Brown Noddy Anous stolidus on Ascension Island. Ibis, 103b: 447-457.

Dwight, J. 1901. The sequence of moults and plumages of the Laridae
(gulls and terns). Auk, 18: 49-63.

Humphrey, P. S., and K. C. Parkes, 1959. An approach to the study of
molts and plumages. Auk, 76: 1-31.

Hutchinson, G. E., 1950. Marginalia: Wideawake Fair. Amer. Scientist,
38: 613-616.

Richardson, F., and H. I. Fisher, 1950. Birds of Moku Manu and Manana
Islands off Oahu, Hawaii. Auk, 67: 285-306.

Serventy, D. L., 1952. The bird islands of the Sahul Shelf. Emu, 52: 33-59.

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YALE PEABODY MUSEUM

oF Natura History

Number 77 November 11, 1963 New Haven, Conn.

A NEW PEREGRINE FALCON
FROM THE CAPE VERDE | eaten

poee

EASTERN ATLANTIC OMEAN

,

S. Ditton RipLtey anp GEorGE E. Watson* f

In the course of preparing a survey paper on the peregrine
falcons, Falco peregrinus, of Eurasia and North Africa, we
had occasion to examine three falcons taken in Cape Verde
Islands a number of years ago by the “Blossom” Expedition
of the Cleveland Museum of Natural History. On the basis of
these same specimens, the peregrine was recorded as a migrant
on the Cape Verde Islands by Bourne (1955) who also saw
“large hawks ... around the rocks of S. Thiego” and sug-
gested that “the species may breed.” More recently, Abbé
Réné de Naurois saw peregrines nesting on Cima, one of the
Rhombos Islets between Raza and Fogo (pers. comm. 1963).
We thank both these authorities for their assistance.

The Cape Verde specimens are as large as F.. p. peregrinus
and F. p. calidus of the northern Palearctic, but they differ
radically from these two subspecies in showing a strong rufous
wash on the head, mantle and underparts as in some specimens
of the smaller Mediterranean race, broohkei, and all specimens

* Smithsonian Institution, Washington 25, D.C.

2 Postilla Yale Peabody Museum No: 737

of the North African desert subspecies, pelegrinoides. We,
therefore, describe the Cape Verde breeding population of the
peregrine falcon as:

Falco peregrinus madens, subsp. nov.

Tyre: Adult 2 (Y.P.M. 44551), collected at Provocao,
Brava, Cape Verde Islands, in 1924, by John da Lomba and
Robert H. Rockwell on the ‘Blossom’ Expedition.

Diacnosis: The adult female differs from typical European
Falco p. peregrinus in being much more brownish and saturated,
with the crown and nape feathers strongly tinged with brown and
rufous producing an irregular rufous nuchal collar. The brown-
ish shading extends down onto the mantle but disappears on
the lower back and rump which are grey as in peregrinus.
The black “moustaches” are edged rufous and the light cheek
patches are suffused with the same color. The general appear-
ance is of a tawny wash or suffusion throughout the underparts
including the flanks and thighs but not the lower abdomen and
vent which are isabelline.

The adult male also differs from peregrinus in having the
crown, nape and mantle washed with brown and rufous to
produce a distinct rufous nuchal collar. Below, there are traces
of rufous wash on the black edges of the “*moustaches” and the
underparts are suffused throughout with dull pinkish buff.
The immature female has a pronounced rufous nuchal patch
and rufous edging to the feathers of the anterior part of the
crown. It is heavily streaked below and roughly similar in tone
of color to immature specimens of brookei although the rufous
wash on the brownish black ‘tmoustaches” is striking.

This population differs from the migrant tundra form, cali-
dus, in being darker, blackish brown, not grey or slaty on the
upperparts, and in being washed with rufous on the head and
cheeks and isabelline or pinkish buff on the remainder of the
underparts.

Compared with the darkest specimens of the smaller Medi-
terranean subspecies, brookei, the two adult birds are as dark or
even darker on the upperparts, less slaty, more blackish, with
the dark shade extending further down the back. There is a

Nov. 11, 1963 A New Peregrine Falcon 3

much more pronounced rufous wash over the head, nape, cheeks
and underparts, but the dark spots and barring are somewhat
reduced below.

From pelegrinoides, which it resembles in having a rufous
wash about the head, madens differs by being larger and al-
together darker, more blackish above, and more heavily washed
with darker buff below. The spotting on the underparts 1s
heavier, especially in the male.

Wing Tail Culmen

MEASUREMENTS: (mm ) (mm ) (mm )
YPM 44551 (Type) ad. 2 340 156 545
YPM 44553 (paratype) ad. 6 320 151 60.0
YPM 44552 (paratype) juv. 2 315 152 54.5

Rance: Cape Verde Islands; recorded so far on Brava, Sao
Thiago, and Cima islands.

Remarks: The adult female is in freshly molted definitive
basic plumage; the male collected on Sao Thiago April 22,
1924, was undergoing prebasic molt (primary + completing re-
growth) ; and the young bird, collected on Brava, date unknown,
was just beginning first prebasic molt (primary 5 lost, scattered
first basic feathers on throat, upper breast and mantle).

The Cape Verde peregrine population continues a north to
south cline of increasingly darker coloration from calidus
through peregrinus to brooket. It shows in the rufous wash of
the head and underparts an approach to the paler Canary
Island and North African desert peregrine, F. p. pelegrinoides.
A male identified as pelegrinoides (No. 537300 in the Ameri-
can Museum of Natural History) which had been kept at Tring,
England for five years, but which was said to have come from
Morocco (Hartert, 1915), is as dark as the male from the
Cape Verde Islands, although the black on the mantle does not
extend so far down the back. Below, this captive bird is even
darker, providing a good example of the well-known tendency
of individual peregrines to molt into darker or lighter plumage
in response to altered diet and environment.

F. p. pelegrinoides and the desert peregrine of Asia, F’. p.
babylonicus, have recently been separated from the forest and

4 Postilla Yale Peabody Museum No. 77

tundra peregrines as a distinct species by Vaurie (1961) and
Dementiew and I]jitschew (1961). The existence of popula-
tions intermediate in color, such as this from the Cape Verde
Islands and another in North Africa (‘“‘punicus” see Klein-
schmidt, 1912-1927, p. 95-96), and records of cross-mated
pars producing young in India (Dodsworth, 1915), however,
induce us to retain all the peregrines in one cosmopolitan
species. We shall discuss further the significance of intermediate
specimens in another paper.

The long wing measurement of the Cape Verde Island popu-
lation is unexpected, since there is a distinct north to south
cline of decreasing wing length in the continental populations
from northern Europe to Africa. It is possible, however, that a
large predator nesting in the cliffs of a small oceanic island
would need to forage at a distance so that a mutation for long
wings would be at a selective advantage in a small population.
Furthermore, the cold Canary Current from the north and
local upwelling result in a climate remarkably temperate for
the tropics, especially in summer (Murphy, 1924). If Berg-
mann’s rule which appears to be valid in this species is related
to temperature, the low summer temperature of the Cape
Verde Islands may help account for the longer-winged pere-
grine.

The subspecific name is derived from the present participle
of the Latin verb madeo and refers to the “saturated” plumage
of the Cape Verde Island peregrine.

Lirerature Crrep

Bourne, W. R. P., 1955. Birds of the Cape Verde Islands. Ibis 97: 508-556.

Dementiew, G. P. and V. D. Iljitschew, 1961. Bemerkungen iiber die
Morphologie der Wiisten-Wanderfalken. Der Falke 8: 147-154.

Dodsworth, P. T. L., 1913. Some notes on the nesting and plumage of the
shahin falcon (Falco peregrinator) and the black-cap faleon (Falco
atriceps) ... Some further notes on the nesting . . . Journ. Bombay
Nat. Hist. Soc. 22: 197-198, 629-630.

Hartert, E., 1915. Notes on falcons. Novit. Zool. 22: 168-176.

Kleinschmidt, O., 1912-1927. Falco Peregrinus. Berajah.

Murphy, R. C., 1924. The marine ornithology of the Cape Verde Islands,
with a list of all the birds of the archipelago. Bull. Amer. Mus. Nat.
Hist. 50: 211-278.

Vaurie, C., 1961. Systematic notes on Palearctic birds. No. 44 Falconidae:
The genus Faleo (Part 1, Paleo peregrinus and Falco pelegrinoides).
Amer. Mus. Novit. 2035: 1-19.

ys/

patella

YALE PEABODY MUSEUM

oF Natura History

Number 78 April 15, 1964 New Haven, Conn.

ONTOGENY AND EVOLUTION IN THE MEGAPODES
(AVES: GALLIFORMES)’*

Grorce A. CiLark, JR.

DeEPARTMENT OF BioLocy, YALE UNIvERsITy2
3
a}

INTRODUCTION

Unlike all other birds, megapodes of Australia and the
Pacific Islands incubate their eggs in mounds or holes by heat
from fermentation, sun, or volcanic activity. Furthermore, meg-
apodes are unique among birds in being able to fly weakly
on the day of hatching and in having no parental care for
young.

These and other reptile-like aspects of megapode reproduc-
tion have been interpreted in two contradictory ways. Some
authorities (e. g. Portmann, 1938, 1950, 1955) have main-
tained that megapodes are the most primitive of living birds,
while others (e. g. Pycraft, 1910) have stated that the similari-
ties of megapodes and certain reptiles are due to convergent
evolution. A related and also unresolved problem has been the
primitiveness of mound-building megapodes relative to those
laying their eggs in holes (cf. Frith, 1962).

1 This study is based on a dissertation presented for the degree of Doctor
of Philosophy at Yale University.

2 Present address: Department of Zoology, University of Washington,
Seattle, Washington.

2 Postilla Yale Peabody Museum No. 78

Despite their anomalous nidification and precocity of young,
megapodes as adults are structurally similar to other members
of the Order Galliformes (e. g. pheasants) as exemplified by the
overlap in adult osteological proportions (cf. data of Verheyen,
1956). Morever, the family Megapodiidae and the New World
gallinaceous family Cracidae (chachalacas, guans, curassows)
are particularly difficult to separate at the family level on a
morphological basis (cf. Miller, 1924). Megapodes and cracids
have been classified as the two most primitive gallinaceous fami-
lies (Huxley, 1868; Peters, 1934).

Unlike the megapodes, most other species of Galliformes have
a simple nest on the ground, but some pheasants, cracids, and
the highly aberrant hoatzin (Opisthocomus) nest in trees.
Since avian development often varies in accord with nidification,
it was anticipated that the study of megapode embryos and
juveniles would reveal clues pertinent to the analysis of mega-
pode phylogeny.

Prior to this study, the only detailed accounts of structure
of embryonic or juvenile megapodes were based on the genus
Megapodius (cf. Pycraft, 1900; Friedmann, 1931; Becker,
1959). These previous investigations had led to contradictory
conclusions on the homologies of the early plumages (cf. Nice,
1962) and on the phylogenetic origins of the family (cf.
Frith, 1962). The object of the present study was to attempt
to resolve the controversy over the phylogeny of megapodes
through examining the morphology of embryos and juveniles
representing several genera of megapodes.

MATERIALS AND METHODS

Specimens. Thirty embryos of the megapodes Talegalla
jobiensis and Leipoa ocellata were studied (Tables 2, 3).
The 11 Talegalla embryos were collected for this investigation
in New Guinea during 1959-60 by E. T. Gilliard and S. D.
Ripley in separate expeditions. The 19 Leipoa embryos were
collected by me during 1960 in the mallee about 25 miles north
of Griffith, New South Wales, Australia; the collecting area
was favorable in having an unusually high density (Frith,
1959) of active Leipoa mounds which were as frequent as one
per 50 acres in the limited suitable terrain. The eggs of Leipoa

April 15, 1964 Ontogeny and Evolution in Megapodes 3

were marked as found and allowed to incubate in the mounds.
Subsequent collections provided accurate ages for three embryos
and minimal ages for certain others (Table 2). The tempera-
ture is ordinarily relatively uniform for Leipoa eggs together
in a mound (Frith, 1959), and the time between egg layings
by a hen is usually four or more days (Frith, 1959, and a
few cases in this study). Thus when actual or minimal age of
one embryo was known, minimal ages of progressively larger
embryos in that mound were estimated by adding four more
days for each. Since the first eggs were probably laid on
September 4 or later, as judged from previous years (Frith,
1959), some specimens (Nos. 12, 16, 19, 1195, of Table 2)
could be assigned presumed maximal ages; smaller embryos
from the same mounds could also be assigned maximal ages,
again using the hypothesis of four or more days between egg
layings in a mound.

Eighty-two juvenile specimens (including 79 study skins)
of megapodes were examined at the American Museum of Nat-
ural History and Yale Peabody Museum (YPM). Among these
were the following species (with numbers of each) : Megapodius
freycinet (59), M. laperouse (5), M. pritchardu (1), Macro-
cephalon maleo (1), Aepypodius arfakianus (2), Talegalla
cuviert (2), T. fuscirostris (4), T. jobiensis (4), Alectura
lathami (3), and Leipoa ocellata (1). More than 140 embry-
onic and juvenile specimens representing 22 genera of non-
megapode Galliformes were used for comparison.

Methods. Characters were chosen for interspecific morpho-
logical comparisons according to 1) potential accuracy of
description or measurement, as determined by reproducibility
in repeated examinations, and 2) potential phylogenetic signifi-
cance demonstrated by the extent of intergeneric variation and
its possible phylogenetic interpretations.

Measurements. Measurements, selected for their applica-
bility over a wide range of sizes, were:

Wine: folded and flattened, with a rule from the anterior
edge of the wrist to the end of the manus, or, in feathered speci-

4 Postilla Yale Peabody Museum No. 78

mens, to the most distant tip of a remex. Due to the distal
shriveling of the ensheathed remiges of embryonic megapodes,
wing lengths over 20 mm (Tables 2, 3, 5) were rounded to the
nearest 5 mm.

Tarsus: with Vernier calipers from the posterodorsal sur-
face of the ankle along the tarsometatarsus to the level of the
proximal surface of the base of the hallux.

CuLMEN: with calipers from the tip to the most posterior
unfeathered point on the dorsal midline.

HuMERUs; RADIUS: respective maximal lengths with calipers.

Tuirp (mippLe) picir: straightened, with a rule from the
tip to the most distal point of webbing connecting with an
adjacent toe.

Megapode embryos Nos. 1, 20, and 21 (Tables 2, 3) were
too immature to measure by these criteria.

Values in the Tables (2, 3, 5) are means of two measure-
ments, each of which, unless otherwise noted, was rounded to
the nearest millimeter. Estimated maximal ranges of variation
in measuring were + 1 mm for dimensions of 2 to 10 mm and
up to + 3 mm for dimensions of 150 mm; these maximal esti-
mates were derived from the ranges in duplications of more than
500 measurements. Among the factors possibly affecting the
accuracy in measuring were 1) unavoidable errors in aligning
and reading calipers and rule, 2) structures changing in shape
as well as length, 3) variations in the positions of parts of
specimens at fixation, and 4) (for anatomical specimens) rate
of fixation with 10 per cent formalin.

Weights (Tables 3, 5), recorded by collectors in the field,
are given only for fresh specimens, as weights of preserved
specimens would be unreliable. The weights and their cube
roots were plotted on arithmetic and double logarithmic graphs
against the various linear dimensions; if any one of the weights
for Talegalla were grossly in error, this would have been seen
as a point lying relatively far from the plot for the other
points. Factors possibly influencing accuracy in weighing in-
clude uneven removal of the yolk sac of embryos before weigh-

April 15, 1964 Ontogeny and Evolution in Megapodes 5

ing, uneven drying of surface moisture on the feathers of
embryos, and variations in the contents of the digestive tract
of juveniles.

MORPHOLOGY OF EMBRYOS AND JUVENILES

Time in embryonic development. Young embryos of the
megapode Leipoa developed slowly compared with embryos of
phasianids (e. g. Gallus, Phasianus, Coturnix), as shown by
the much later occurrence of the first gross appearance of
egg tooth, feathers, labial groove, etc., in Leipoa (Table 1).
Through the first 20 days, these Leipoa embryos attained a
much smaller absolute size than did embryonic chickens (domes-
tic G. gallus) as illustrated by comparing linear dimensions of
Letpoa and chickens (Fig. 1; Tables 2 and 3). As an example,
after 20 days of incubation the wing of an embryonic Leipoa
was less than 50 per cent as long as that of a chicken (Fig. 1).

The normal incubation period of Leipoa is generally at least
twice as long as that of known phasianids or turkeys (cf.
Table 1; see also Frith, 1959, on Leipoa, and Romanoff, 1960,
on phasianids). This lengthy incubation period of Letpoa ts

Tasre 1. Time of certain gross morphological changes in embryos of the
megapode Leipoa and of phasianids. Age in days after laying of the

egg.
Leipoa Phasianus — Coturnix Gallus
age age age age
Egg tooth formed ..... 21-22 9 5-6 614-7
Labial groove formed .. 21-22 ? ? 10
Feathers appear ....... 11-21 9 5-6 61/-7
Toes are first separated 11-21 10-12 7-8 8-9
Scales appear on legs .. 29-54 13 8?-9 11-12
Eyelids come together .. 29-61 15 10-11 13
Labial groove lost ..... 29-61 ? ? 19
Flatehin gem 3) 3..0ic 6 eos e- 60-73 23-24 16 20-21

Sources of data: Leipoa ocellata, specimens of this study; Phasianus
colchicus, Fant, 1957, and Westerskov, 1957; C. coturnix japonica,
Padgett and Ivey, 1960; domestic G. gallus, Hamilton, 1952.

6 Postilla Yale Peabody Museum No. 78

related to both the slow early development and the large size
at hatching (see p. 27 for discussion of the effects of incuba-
tion temperatures).

18
4

140 wv,
ec Gallus
E =~
eo !00
£
=

60

fe Leipoa

Days

Figure 1. Chronological growth of the wing in Leipoa ocellata and do-
mestic G. gallus (data from Tables 2 and 3). Curves showing length against
time were fitted by inspection and should not be considered as quantita-
tively accurate.

Relative proportions and growth. At hatching T'alegalla
and Leipoa are about two to 15 tinfes heavier than other newly
hatched Galliformes of the genera Coturnia, Colinus, Phasianus,
Gallus, and Meleagris (Lyon, 1962; Westerskov, 1957; Roman-
off, 1960; see also Table 6). It is of interest that Letpoa and
Talegalla at hatching have proportions and size like those of
adult C. coturnix japonica (Table 5). The genus Megapodius
is intermediate in hatching weight (Table 5) between Talegalla
and phasianids or turkeys.

As a means of comparing changes in proportions during the
growth of different species of the Order Galliformes, arithmetic
and double logarithmic plots (e. g. Figs. 3, 4, 5) were prepared
using the linear measurements of embryos and juveniles (data

of Tables 2, 3, and 5). Such proportional growth was described

April 15, 1964 Ontogeny and Evolution in Megapodes 7

approximately in certain cases by using the conventional allo-
metric equation, Y = AX, or the equivalent form, log Y = log
A+B log X, where X and Y are the values of two dimensions.
A and B (Table 4) were calculated using Bartlett’s method as
described by Simpson, Roe, and Lewontin (1960). B values for
different species were compared using a modified t-test (Simp-
son et al., 1960). Correlation coefficients for the sets of data
expressed as B values in Table 4 were all significant at the
0.001 level.

To compare growth of linear dimensions relative to total
body size in different species, the cube root of weight was used
as one criterion for body size (see Amadon, 1943, for the

Taste 2. Data for specimens of Leipoa ocellata. All are embryos except
1195. For procedures of measuring, see text. All lengths in milli-
meters. Estimated ages in days, Symbols: S, specimen number; W,
wing length; T, tarsal length; C, culmen length; H, humerus length;
R, radius length; Td, length of third digit; m, male; f, female;—,
observation could not be made.

S Ww T Cc H R Td Sex Age
1 a -= a: — 11-?
2 if 3 3 5 3 3} — 21-22
3 9 6 5 6 6 4. — 22
4 12 8 af 10 10 6 _- 29
5 13 8 8 10 9 7 + P-54
6 20 12 9 13 13 11 m 19-58
7 20 13 9 13 12 10 m ?-55
8 25 14 10 16 14 12 f ?-59
9 25 15 10 llie¢ 18 13 m ?
10 40 19 12 20 20 15 m ?
11 45 20 iat 23 20 15 m ?-62
12 45 PMA 12 22 21 15 m 45-70
13 55 22 12 22 23 16 f 48-61
14 70 24 14 28 26 19 m 2-63
15 7 23 13 27 25 IY —— 52-64
16 (3) 25 14 28 27 18 m 49-74
17 80 26 13 28 29 20 f 56-69
18 80 24 14 31 28 20 f ?-67
19 85 26 14 29 30 20 £ 60-73
1195* 115 28 — 38 37 24 — 2-73

* This specimen, found dead in the field, was lacking its head.

8 Postilla Yale Peabody Museum No. 78

LEGEND
8 ae
B® Talegalia °
13 @ Megapodius res S a
O Aepypodius
6 i Vv
V Alectoris v
5 © Phasianus ee 7

Cube Root of Weight

Culmen (mm)

Figure 2. Relationships of the cube root of body weight in grams to the
culmen length in the megapodes Talegalla jobiensis, Aepypodius arfa-
kianus, Megapodius freycinet and the phasianids Alectoris chukar and
Phasianus colchicus. All data from this study except that for Phasianus,
for which mean values for males were taken from Westerskov, 1957. See
text for discussion,

explanation of this procedure). Since weights were unknown
for most specimens, a linear criterion for body size was also
chosen. As the culmen length had a relatively direct relation-
ship to the cube root of body weight over a fifty fold range of
weights for eight specimens of embryonic and juvenile T'alegalla
jobiensis and for six juvenile specimens of the phasianid Alec-
toris chukar (Fig. 2), culmen was selected as a convenient
linear measure for body size in these specimens. Moreover, simi-
lar analyses revealed that culmen is a relatively good measure
for body size in embryonic chickens (10-21 days; matched
lengths from figures of Hamilton, 1952, with weights from
Romanoff, 1960) and in juvenile Phasianus colchicus from
zero to nine weeks posthatching (Westerskov, 1957; see also
Fig. 2 of this study). Since the culmen is a less sensitive and
less accurate indicator of body size than is the cube root of
body weight, certain interspecific differences have possibly gone
undetected due to the use of culmen as a major standard for
body size.

April 15, 1964.

Ontogeny and Evolution in Megapodes

9

The scales for the cube root of weight in Figs. 3, 4, 5, are
calculated from a mean value of 3.24 for the ratio of culmen
length to the cube root of body weight in grams for the eight
weighed specimens of Talegalla jobiensis. Due to the relative

imprecision of culmen measurements (compared with weights )

Taste 3, Data on specimens of Talegalla jobiensis and domestic G. gallus.
Nos. 1196, A, B, C, S, T, U, V, W are posthatching specimens. For
procedures of measuring, see text. All lengths are in millimeters.
Weights in grams. Ages in parentheses are estimated from stages in
Hamilton (1952). Symbols: S, specimen designation; W, wing length;
T, tarsal length; C, culmen length; H, humerus length; R, radius
length; Td, length of third digit; m, male; f, female;—, observation

could not be made.

Talegalla:
Ss WwW
21 —
22 9
23 11
24 12
25 18
26 20
27 45
28 80
29 100
30 100
1196 115
A 115
B 160
C 164
Gallus:
N 8
O 10
P 11
Q 25
R 27
Ss 85
ab 110
U 135
Vv 170
WwW 160

Den

10
10
15
18
21
24
24

aor oO

ao |?

—_
bo bo ©

_
©

Age (days)

(10-11)
(11-12)
(12)
(19)
(19-20)

Note: Specimens A, B, C are study skins.

10 Postilla Yale Peabody Museum No. 78

50 50
LEGEND LEGEND
OQ Leipoa ie) Q Leipoa
B Jolega/la 12) @ Jolegalla
O Ggallus O Ggoallus 8
40 40
| . - °
¢ Pe Z
E E
iy ei) Ke E 30 Oo
3 | og r "s
9° o
@ g 3 oa
a A Z
=
20 fs :
a
a
Vr
10 a
a>
we
A O
5 10 15 20 25 5 10 15 20 25
Culmen (mm) Culmen (mm)
aaa See —
LS 3.1 4.6 6.2 Usk! Sel 4.6 6.2
AY Weight AY Welght

Figure 3A, (Left) Growth of the radius relative to the culmen in
Leipoa ocellata, Talegalla jobiensis and domestic G. gallus (see alsa
Fig. 4). B. (Right) Growth of the third digit relative to the culmen in
these three species (see also Fig. 4).

Cube roots of weights in grams calculated by the method indicated in
Texte Gpemo))

and probable interspecific variations in the mean ratio of cul-
men to the cube root of body weight, the cube root values in
Figs. 3, 4, and 5, are probably not precise for individual speci-
mens shown on the graphs3 nevertheless, these cube roots of
weights help to indicate, in an approximate way, the relative
growth of the different species.

As shown by either arithmetic (e. g. Figs. 3, 5A) or loga-
rithmic plots (Fig. 4), growth of linear dimensions relative to
culmen in the two species of megapodes is generally similar to
that of Gallus (see also Table 4). It should be emphasized,
however, in view of the necessarily small sample sizes and
inherent limits of accuracy in measurement, that these analy-
ses tend to mask certain differences in relative growth. For
example, in embryonic chickens the radius (Fig. 4A) and
humerus temporarily have lower rates of relative growth fol-
lowed again by higher rates (this study) ; the data of Roman-

April 15, 1964 Ontogeny and Evolution in Megapodes 11

off (1960: 1146) show that the slow growth of these structures
in chickens occurs about 14-17 days of incubation. As a con-
sequence, the radius and humerus of chickens near hatching
are a few millimeters shorter than those of similar-sized embryos
of Talegalla or Leipoa (data in Tables 2, 3). In addition,
measurements of three juveniles of the phasianid C. coturniar
japonica revealed for this form also a slow mean rate of embry-
onic growth of radius and humerus relative to other dimensions
followed by increased relative rates after hatching. The rela-
tively short radius and humerus of Gallus and Coturnia in
older embryos and at hatching are possibly adaptive in prevent-
ing premature flying of the young birds; such an adaptation
would be analogous to the retarded development of remiges in
juveniles of forms such as petrels and hawks. No trace of a
relatively slow embryonic growth of radius and humerus was
found in the megapodes.

Culmen measurement in the utilized samples covers a rela-
tively small range (less than 20 mm), but this handicap is offset
somewhat by the utility of this measurement for study skins.
The culmen is measured linearly over a curved surface but
nevertheless is empirically useful. In measuring the culmen of

Tasie 4. Interspecific comparison of allometric growth of dimensions rela-
tive to culmen. None of the interspecific differences in exponent is sta-
tistically significant. See text for details.

Exponent (B) with Size
95 per cent Coefficient of

Dimension Species confidence interval (A) sample
Tarsus Leipoa 1.6 + 0.2 0.24 18
4 Talegalla 1.5 + 0.3 0.20 13
WY Gallus 1.4 4+ 0.2 0.18 10
Humerus Leipoa 1.5 + 0.4 0.19 18
» Talegalla 1.4 + 0.2 0.15 10
is Gallus 1.4 + 0.2 0.19 10
Radius Leipoa 1.6 + 0.2 0.26 18
” Talegalla 1.4 + 0.2 0.17 10
ue Gallus 1.5 = 0.2 0.27 10
Third digit Leipoa 1.5 + 0.3 0.24 18
es mh Talegalla 1.5 + 0.2 0.23 10

ey ae Gallus 1.5 + 0.2 0.20 10

12 Postilla Yale Peabody Museum No. 78

late embryonic and juvenile chickens (Gallus; Table 3), the
presence of the comb necessitated estimating culmen lengths
in eight specimens through the projection of lines from the
postero-lateral margins of the horny bill dorsally to the mid-
line; however, this approximation did not alter the interpreta-
tions as shown by using other combinations of dimensions. At
hatching in Gallus, Leipoa, and T'alegalla, the culmen may lose
up to 1 mm in length through loss of periderm, but this small
change does not affect the interpretations of relative growth.

Analogous to the shorter culmen after hatching are reduc-
tions (about 5 mm) in wing length of juveniles of these species
through loss of natal downs and also the decrease (less than
1 mm) in length of the third digit through loss of the claw
pad at hatching. Here again the interpretations of relative
growth were not affected.

Relative and proportional growth of gallinaceous wings was
too complex to permit adequate representation in a simple
equation, but, as shown by graphs (e. g. Fig. 5), relative
growth of the wing in Talegalla, Leipoa, and other Galli-
formes was similar within the size range considered. The propor-
tional growth illustrated in Fig. 5B suggests possible interspe-
cific differences which, however, are not especially striking. Data
for the Jungle Fowl (G. gallus) were used in Fig. 5 to provide
a larger sample, but data for chickens (domestic G. gallus;
Table 3) gave similar results.

Juvenile Megapodius have an unusually short culmen con-
trasted with those of juveniles of other megapodes or other
Galliformes; the mean ratio of culmen length to the cube root
of body weight for three Megapodius freycinet (Fig. 2; Table
5) was 2.1, compared with 3.24 for eight T'alegalla jobiensis.
Young juvenile Megapodius (Table 5) also differ from young
juveniles of Talegalla in having a longer wing relative to the
cube root of body weight.

Measurements of wing, tarsus and culmen of more than 110
other juvenile specimens representing 22 genera of non-mega-
pode Galliformes (cf. Table 5) were plotted on graphs and
compared. These species generally appear to have proportional
growths similar to those of T'alegalla, Leipoa, and Gallus.
However, a juvenile Craz rubra of the cracids (Table 5) was

Radius (mm)

April 15, 1964 Ontogeny and Evolution in Megapodes_ 13

exceptional in having a relatively short wing (shown also by
figures of young Craw globicera (= rubra) in Heinroth, 1931).
The shorter wing at hatching in Craw is apparently associated
with the generally less well-developed feathers (p. 24). Forms
such as ducks (e. g. Anas) which have delayed formation of
juvenal remiges show plots of alar growth quite unlike those of
Galliformes.

These analyses, although necessarily based on small samples,
indicate that embryonic megapodes undergo proportional and
relative growth analogous to that occurring up to several weeks
posthatching in phasianids. Certain forms such as Megapodius
and Crax show interesting deviations from the general gal-
linaceous conditions. Larger samples might reveal additional
interspecific differences and possibly intraspecific variations
according to individuals, sex or locality.

Some qualitative comparisons of embryos and juveniles.
Embryos of Leipoa (e. g. Nos. 2 and 19) and of chickens
shortly prehatching behaved similarly when taken from the
shell, i. e. the embryos gaped and kicked. Even Leipoa embryos

5 LEGEND 5

aes 8 LEGEND
= mained 40 4 Le/lpoa 8
alegalla Bo ° 2 Talegalia °
3 ©°G. gallus — 30 oG. gallus °
E 's
€
2 5. 20:
2
a
he
10
7
5
3
3 5 7 10 20 30
3 5 Tf ake) 20 30 Culmen (mm)
Culmen (mm)
SF
— 1.5 rary. & \.
ES) 12:2; 75.1 6.2 3 ns
“YWeight” V Weight

Figure 4A. (Left) Double logarithmic plot of growth of the radius rela-
tive to the culmen in Leipoa ocellata, Talegalla jobiensis and domestic
G. gallus. B, (Right) Double logarithmic plot of growth of the third digit
relative to the culmen in these three species.

Cube roots of weights in grams calculated by the method indicated in
LEXE ((219)))-

14 Postilla Yale Peabody Museum No. 78

Taste 5. Comparison of dimensions of some juvenile Galliformes. Speci-

mens arranged by increasing tarsal length. For procedures of measuring,

see text. Lengths in millimeters. Symbols: W, wing; T, tarsus; C, culmen;
m, male; f, female; g, grams.

Woh iG WwW T ic

Numida meleagris (f) 20 17 10 Penelope purpurascens 105 27 12
Chrysolophus pictus 20 18 7 Megapodius pritchardii 85 28 6
Phasianus colchicus 30 19 8 Gennaeusleucomelanos 105 28 15
Opisthocomus hoazin 40 19 12 Chrysolophus pictus 115 30 14
Gennaeus leucomelanos 50 19 10 Alectoris chukar

Syrmaticus mikado 30 20:48 (f; 154 g) 120 30 16
Chrysolophus pictus 26 20 7% Tragopan temmincki 130 30 14
Phasianus colchicus 35 21 10 Ortalis wagleri 110 31 15
C. coturnia japonica 90 22 12 Alectoris chukar
Alectoris chukar (5195-2) a0 ese
(m; 73 g) 95 22 14 £Mitu tomentosa 85 32 14
Phasianus colchicus 85 23 13 #&£z%Talegalla fuscirostris 120 32) 15
Meleagris gallopavo 45 24 10 Alectura lathami 105 33 15
Ortalis wagleri 45 24 10 #£Phasianus colchicus (f) 110 33 18
Ortalis vetula 70 24 12 Francolinus gularis 120 33 15
Megapodius freycinet Megapodius freycinet
(f; 63.6 g¢) 100 24 8 (f; 117 g) 125 33 10
Numida meleagris (m) 110 24 14 Megapodius freycinet
Megapodius laperouse 95 25 8 (123.5 g) 130 33 11
Ulectorimchubar Opisthocomus hoazin 165 34 19
(m; 121 g) 115 26 15 Macrocephalon maleo 140 35 15
Meleagris gallopavo 55 27 9 Crax rubra (m) 75 36 15
C. coturnix japonica Dendragapus obscurus
(adult) 100 2% 13 (f) 175 36 18

considerably larger than chickens at hatching show this charac-
teristic embryonic behavior.

Meyer (in Meyer and Stresemann, 1928) noted the large fat
deposits in late embryonic Megapodius; both Talegalla and
Leipoa embryos (this study) also have subcutaneous fat bodies
distributed similarly to those of chicken embryos but covering
a wider area in embryos near hatching. These deposits in older
Talegalla and Leipoa embryos are especially well developed
laterally along the neck and beneath portions of the ventral
feather tract.

The genus Megapodius (Miller, 1924; confirmed in this
study) is unusual among Galliformes in having a small web

April 15, 1964

180 Oo
170
B
160 |
oO
1504
@)
140 fe)
130 4
120 4
a
_ lod fe)
iS
E @)
100 | a
oO
2 o
= 904
AO
80 { as
w
704 a
604 (@)
@®
504
ao] a
6)
304
(@.ay
204 é
ol
a
10 20 30
Culmen (mm)
a eed
SHI 6.2
WYWeight

Figure 5A. (Left) Growth of the wing relative to the culmen in Leipoa
ocellata, Talegalla jobiensis and G. gallus. Cube roots of weights in grams
calculated by the method indicated in the text (p. 9). B. (Right) Propor-

Ontogeny and Evolution in Megapodes 15

LEGEND
Leipoa &
Tolega/ia @
G. gallus O

(mm)

WING

180

170

150 4

120 4

0 4

100 4

90 4

80

50 4

40 4

LEGEND
XQ Leipoa
B Jolega/lia
O Ggollus

Tarsus

40 50 60
(mm)

tional growth of the wing versus the tarsus in these three species.

between the second and third toes but, unlike forms such as
Leipoa, Talegalla and Gallus, none between the third and

fourth toes.

A few qualitative gross morphological changes appear at
a greater absolute weight, and, for larger embryos, at a detec-

16 Postilla Yale Peabody Museum No. 78

tably greater linear size, in the megapodes (Leipoa and Tale-
galla; this study) than in Phasianus (Westerskov, 1957) or
Gallus (structures from Hamilton, 1952, matched with weights
from Romanoff, 1960). Examples of these phenomena in T'ale-
galla versus phasianids (Table 6) include first appearance of
feathers, egg tooth, labial groove, and coming together of the
eyelids.

Taste 6. Comparison of weights at times of certain qualitative morpholog-
ical changes in Talegalla jobiensis, Phasianus colchicus, and domestic
G. gallus. Weights in grams. Talegalla weights in parentheses were esti-
mated from culmen lengths using the relationship reported in the text

Gre Ye

Macroscopic : Talegalla Phasianus Gallus
character weight weight weight
First appearance, feathers ........ 3.5 - 4.7 0.7-1.7 0.4- 1.2
First appearance, egg tooth ........ 4.7 - 5.0 0.7-1.7 0.4- 1.2
Formation of separate toes ......... 3.5 - 4.7 1.4-4.8 0.7- 2.3
Formation of scales on legs ........ (5.5)-14.3 3.2-5.8 2.3- 7.3
Kyelids coming together ........... 22- (40) 4.7-8.5 §.2-11.0
eV evtchinn oie ere reeis says aieectarchetrer aie eotens 110+ 23 33

Sources of data: Talegalla from this study; Phasianus from Westerskov
(1957) ; Gallus morphology from Hamilton (1952) combined with Gallus
weights from Romanoff (1960: 1147).

Tarsal seutellation. My observations on the tarsal scutella-
tion of megapodes support the findings of Ogilvie-Grant (1893).
Megapodius, Aepypodius, and Talegalla are alike in having a
single row of large scutes down most of the foresurface of the
tarsus (tarsometatarsus), but Aepypodius has two rows dis-
tally. Alectura and Leipoa have two rows of large scutes down
the foresurface, while Macrocephalon has many small scutes.
Tarsal scutellation is similar in juveniles and adults within a
species of megapode.

Turkeys, many phasianids and some cracids have two rows
of large scutes on the foresurface, while many cracids possess
only one row; Opisthocomus has many small scutes.

Feathering of the oil gland. T'alegalla jobiensis has a naked
oil gland (no feathers on the tip; Fig. 6, this study) and thus

April 15, 1964 Ontogeny and Evolution in Megapodes 17

Figure 6. Oil glands of domestic G@. gallus (19 day embryo; ca. 5.5),
Magapodius laperouse (YPM 89; juvenile; ca. 3X), and Talegalla
jobiensis (No. 29, embryo; ca. 1.5), from left to right. Dorsal view.

is like Alectura and Leipoa (Miller, 1924; confirmed in this
study). In contrast, Megapodius laperouse (Fig. 6, this study)
has a tufted oil gland as was reported by Miller (1924) for
other species of Megapodius and for Macrocephalon. Most
(ralliformes, excluding megapodes, have tufted oil glands (Fig.
6 of this study; Miller, 1924; see also Table 7 for a summary
of this character in other birds).

Eutaxy. Unlike other gallinaceous families, megapodes have
variation in eutaxy (presence of the fifth secondary; Steiner,
1918; Miller, 1924). As anticipated from reports on allied
species (1. e. Alectura and Leipoa; Miller, 1924), T'alegalla
jobiensis is eutaxic (this study). Both Talegalla and Leipoa
are eutaxic at the first embryonic appearance of the second-
aries. Megapodius laperouse (YPM 89) is also eutaxic, but
M. pritchardii (Pycraft, 1900) and some (but not all) mem-
bers of M. freycinet (Steiner, 1918; Miller, 1924) are diasta-
taxic (lacking the fifth secondary). Macrocephalon is also
diastataxic (Miller, 1924). In contrast, all other Galliformes,
including chickens, are eutaxic (Miller, 1924; see also Table 7
for a summary of diastataxy and eutaxy in other birds).

Carotid arteries. In agreement with the data reviewed by
Glenny (1955) for Megapodius freycinet, M. pritchardu,
Macrocephalon, and Alectura, the megapodes dissected in this
study (e. g. Leipoa No. 17, Talegalla No. 29, Megapodius
laperouse YPM 89) had a left dorsal carotid artery but none

18 Postilla Yale Peabody Museum No. 78

on the right side; in contrast, chicken embryos possessed both
right and left dorsal carotids. Gleniry (1955) has reported that
all Galliformes except megapodes are bicarotid (see Table 7
for a summary of this feature in other birds).

Early plumages. Studer (1878) and Pycraft (1900) be-
lieved that megapodes molt natal downs before hatching,
but Portmann (1955) and Becker (1959) have contended

Taste 7. Status of dorsal carotid arteries, disastataxy versus eutaxy, and
oil gland feathering in nongallinaceous birds. Symbols: 2, bicarotid; 1, uni-
carotid; EK, eutaxy; D, diastataxy; T, tufted oil gland; N, naked oil gland;
O, no oil gland.

Taxonomic Carotid Fifth Oil

group arteries secondary gland
SINITVAIING BON oteetsrerapercieerersi neko evoletee 2 E 4u
SOT SUULUCSS ae ere tr eucy craie eet iaee ier omatets 1,2 D,E O,?
Gavi aes Waeccrecce scx eusseesistaeerepers 2 D T
POGICIPEMITONMES! Ciscralerelee selects 1 D Al
Procellariitonmesi yen itr 2 (1) D AY
CHOI MWINGNCIS BoobnasdoouoaocoKS 2 D it
Pelecanifonmesiy ceccc ccele ces 1,2 D, E Mt
Wiconiiformesmnseeie. eecleeiaci 1,2 D oN
AMATI aA el eects etevs eiettous so eharereusuors 2 D at
ATV CIMA? Pn ytee nie sisciatnn siecle ene 2 D 48
Halcomitormesesareace mecca 2 D ts N
GrUiON MeSH whence ehorsieionceeatene ier: 1,2 DH TNO
Charadniitonmesmarrtrirrert sia 2 (1) D, E at
@olumbitormesh a sao cn ose eee 2 D, EB N,O
Psittacikonmes eine ceo corre NY D fr @)
Miusophacidae sas) -seleriierorels cis 2 E At
G@uculidae: Ba wcecivaeecnerocnes tes 2 E N
Sitriciornmesy seciece a aceerrtercr 2 D TN
Caprinauleiformes! ye -iite ie. D N,O
FNyYorshhito ales yogononobeonodnor In DE N
@olliormMes? 250 aos see omer 1 E N
MromonilOEMmess reese ee eta 1 E N
C@oraciiformes! Gf ceeeeeeeeeeee 1,2 D, E ei
PictfOrMeS 1 oAcite ache ce eho 1,2 E TNO
Passeritormesere cei tierce 1 E N

Sources of data: arteries, Glenny, 1955; eutaxy and diastataxy, Steiner,
1956; oil gland, Beddard, 1898, and Miller, 1924.

April 15, 1964 | Ontogeny and Evolution in Megapodes 19

that megapodes lack natal downs and that their first feathers
represent the phylogenetic precursors of natal downs. In con-
trast, Friedmann (1931) stated that megapodes at hatching
bear juvenal feathers in opposition to several authors (e. g.
Ogilvie-Grant, 1893), who referred to the downy young. In
order to determine which, if any, of these conflicting views is
correct, it was necessary to analyze many features of pterylo-
sis, feather growth, and molt.

In the embryonic early growth of the megapode feathers,
those of the tail are longest. For example, on one Leipoa
(No. 5) the caudal sheaths (10 mm long) were 5 mm longer
than the next longest ones on the cervical region and femoral
tract. Similarly, a T'alegalla embryo (No. 24) with tail feath-
ers of 10 mm had the next longest sheaths (8 mm) on the
cervical region. Precocious embryonic early growth of caudal
natal downs occurs in chickens (Hamilton, 1952) and Coturnix
Quail (Padgett and Ivey, 1960) and is apparently a gallina-
ceous trait.

Although a row of 9 or 10 relatively large papillae initially
were formed on the posterior surface of the manus (e. g. on
Nos. 3, 22, 23), of these only primaries 1 through 8 were large
on older embryos and newly hatched T'alegalla and Leipoa
(see also Pycraft, 1900, for Megapodius). Such embryonic
repression of the juvenal outer primaries (9 and 10) is charac-
teristic for many Galliformes.

Embryonic megapodes do not molt, contrary to the report
of Studer (1878), who was misled partly by the ease with which
immature sheaths are dislodged from the skin. Indeed, feather
maturation, manifested by hardening, does not occur on the
body in T'alegalla and Leipoa until the last quarter of incuba-
tion as determined by dissection of sheaths from eight tracts.
At hatching, as in other Galliformes, the feathers on the body
are fully grown or nearly so, but the vanes of the remiges
continue growing.

Feather sheaths at hatching are longer on T'alegalla and
Leipoa than on chickens. To illustrate this condition, the mean
lengths (M) and coefficients of variation (CV) were calculated
for six sheaths from each of three embryos near hatching. The
six sheaths were taken from corresponding positions on six

Postilla Yale Peabody Museum No. 78

Si)

i

(i=

i BS IWS \\N

Z

Soot

Figure 7. Comparison of the tip of secondary No. 9 of the right wing
(top; ca. 4X) with a natal down from the body (bottom; ca. 3X).

Leipoa ocellata No. 19; 60-73 days of incubation.

April 15, 1964 Ontogeny and Evolution in Megapodes_ 21

tracts on the body of each of the embryos. The values were:
Gallus (19 day) M 13.8 mm (CV 37.6); Letpoa (No. 19) M
28.6 (CV 39.9); and Talegalla (No. 30) M 36.5 mm (CV
38.8). In view of the great variation in lengths of sheaths
within a tract, these values are useful only to indicate the
great difference between megapodes and chickens.

Sheaths on the body of Talegalla and Leipoa embryos
appeared conventional, having opaque and unshriveled tips,
but sheaths of remiges, alula quills, and certain alar upper
coverts of the older T'alegalla and Leipoa embryos had unusual
translucent and shriveled tips as noted by Pycraft (1900) for
remiges of embryonic Megapodius. Pycraft (1900) figured a
constriction of the sheath of the Megapodius remex in the
region of transition from opaque to translucent portions. This
constriction does not occur in Leipoa and Talegalla (this
study); due to lack of a suitable specimen of Megapodius,
it was not possible to check Pycraft’s report of a constriction
in that genus.

Within the translucent tips of the sheaths of remiges on
older Talegalla and Leipoa embryos are weak filaments which
are distal portions of the central barbs of the tip of the remex
(Fig. 7). These distal filaments are easily dislodged in removing
remiges from the sheaths so that some or all filaments are
missing from the expanded remiges of embryos (as in Fig. 7)
and juveniles. Unlike the correspondingly placed natal downs
on the tips of juvenal remiges of phasianids or cracids, these
filaments on the tips of remiges of embryonic megapodes are

weakly developed and lack barbules.

On juveniles of six megapode genera (this study), the feath-
ers at hatching have 1) barbule-free distal ends of central barbs
of body feathers (Fig. 7); 2) a central rhachis; 3) a large
aftershaft on the body feathers (Fig. 7) ; 4) a well-formed vane
in the remiges; these features in common demonstrate that
megapodes had common ancestors possessing such features at
hatching. In contrast, the feathers of chickens at hatching have
1) barbule-free distal ends of central barbs; 2) a distinct
rhachis only in the short and growing juvenal remiges; 3) no
aftershaft ; 4) a well formed vane only in the growing remiges.

22 Postilla Yale Peabody Museum No. 78

Hall (1901), Blasyzk (1935), and Frith (1962) have
reported for juvenile Leipoa and Alectura that the feathers
on the body at hatching are later carried out on the tips of
the growing second feathers. The finding of these connections
(this study) on Letpoa ocellata (Fig. 8), Alectwra lathami,
Talegalla jobiensis, and Megapodius freycinet, demonstrates
that this is another general feature of megapodes. As the first
feathers are easily dislodged from the tips of the second ones,
the rarity of observations of these junctions on preserved speci-
mens is to be expected. These connections resemble those be-
tween natal downs and juvenal feathers in other Galliformes.

Figure 8. A natal down attached to the tip of a juvenal rectrix from
juvenile Leipoa ocellata. (YPM 1195) ca. 3X.

April 15, 1964 Ontogeny and Evolution in Megapodes 23

However, since similar connections occur between other genera-
tions of feathers in Galliformes (Watson, 1963), these attach-
ments, considered alone, do not demonstrate conclusively that
the first feathers on the body of megapodes are natal downs.

Nevertheless, the homology of megapode feathers on the
body at hatching with the natal downs of other Galliformes
is shown by the following features in common: 1) the preco-
cious early growth of embryonic tail feathers; 2) the plumula-
ceous structure of the feathers on the body at hatching relative
to the more pennaceous structure of later generations of feath-
ers and of the first remiges; 3) attachment of the first feathers
to the tips of growing feathers of the second generation; 4)
barbule-free distal ends of central barbs; 5) start of the first
body molt within two weeks posthatching (data on Leipoa
timing from Hall, 1901, and Frith, cited in Nice, 1962).

The following group of characters demonstrates that the
megapode first remiges are juvenal like those of other Galli-
formes: 1) only eight primaries at hatching but ten on older
juveniles and adults; 2) similar lengths of growing primaries
Nos. 1( first basic = postjuvenal) and 10 (juvenal) on juve-
nile Megapodius (YPM 89) as in certain juvenile phasianids
(cf. Heinroths, 1928) ; 3) remiges more pennaceous than other
feathers at hatching; 4) similar location of the distal filaments
on the embryonic remiges of megapodes and of the correspond-
ing natal downs on other Galliformes ; 5) time of initial loss of a
first remex (two weeks posthatching in Leipoa; Hall, 1901) ;
synchrony of molt of natal downs on the body and juvenal
remiges is characteristic for Galliformes.

The lengths of rhachises in the natal downs of Galliformes
can be partly correlated with the size of the newly hatched
birds. For example, the young of small phasianids, e. g. Cotur-
nix, lack rhachises in their natal downs, while turkeys (Melea-
gris; Pycraft, 1900, and confirmed in this study; and Agrio-
charis; this study) and tragopan pheasants (this study), both
of which are larger at hatching than are the small phasianids,
have short rhachises in their natal downs. Megapodes, still
larger at hatching, have longer rhachises (Fig.7). Certain cra-
cids, e. g. Crax, are exceptional in being large at hatching

24 Postilla Yale Peabody Museum No. 78

(over 100 grams; Heinroth, 1931), while lacking or having
only short rhachises in their natal downs (this study).

As might be expected from the data thus far presented,
many phasianids molt the last of their natal downs at a body
size smaller than that of juvenile megapodes at the time of loss
of the last natal downs. For example, Phasianus colchicus at
160 grams has lost nearly all the natal downs (Westerskov,
1957), while T'alegalla (e. g. B of Table 3) at this weight
retains many natal downs on the breast, back and head.

Thus the hatching plumages of megapodes and other Galli-
formes are homologous but differ structurally.

Structures associated with hatching. Several authors (e. g.
Frith, 1959) have reported megapodes at hatching kicking
their way out of the shell, and some observers (e. g. Elvery in
Campbell, 1901) have emphasized the difference from hatching
m chickens. A relatively detailed description of megapodes at
hatching is that of Bergmann (1961), who observed that, in
Talegalla cuvieri, at the time of breaking open of the shell, the
only parts of the body to break through the shell membrane
were the legs and feet. Thus T'alegalla is unlike both chickens
(Hamilton, 1952) and Coturnix Quail (Clark, 1960) which
use the egg tooth of the beak conspicuously in breaking open
the shell.

Although Friedmann (1931) could not find an egg tooth on
one Megapodius pritchardii embryo, and Bergmann (1961)
could not find an egg tooth on Talegalla cuvieri at hatching, I
(1960, 1961) have found egg teeth on both Talegalla jobiensis
and Leipoa ocellata embryos (latter observation made indepen-
dently by Frith, 1962). Frith has kindly shown me one speci-
men of prematurely hatched Leipoa bearing an egg tooth,
which, together with my finding that many other specimens of
newly hatched megapodes lack egg teeth, suggests that egg
teeth are usually lost about the time of hatching in megapodes.
The egg teeth of chickens near hatching are approximately
two times larger in linear dimensions than the fully grown egg
tooth of Leipoa (Fig. 9) or Talegalla. Especially when con-
sidered relative to body size at hatching, the megapode egg
tooth is quite small. I (1961) have reviewed the occurrence

April 15, 1964 Ontogeny and Evolution in Megapodes 25

of egg teeth in birds as a whole; egg teeth probably occur on
most, if not all, birds. Megapodes are the only birds for which
egg teeth are thought to be nonfunctional at hatching.

In T'alegalla and Leipoa the Musculus complexus or “hatch-
ing muscle” is located dorsally on the neck immediately under
the skin (and under fat deposits in larger embryos), attached
anteriorly to the parietal of the skull, and posteriorly con-
nected to the third, fourth, and fifth cervical vertebrae and the
muscular complex overlying these vertebrae. The two complexus
muscles were separated in the dorsal midline in the 20 examined
anatomical specimens of megapodes: in Leipoa by minimal

Figure 9. Egg tooth of an embryonic Leipoa ocellata. (No. 9) Overlying
periderm removed. Ca. 7X.

distances of 1.5 (No. 4) to 3 mm (No. 19) and in Talegalla
by 2.5 (No. 26) to 5 mm (No. 30). In contrast, in chicken
embryos near hatching, the two complexus muscles met in the
dorsal midline (Fig. 10). The anterior insertions meet in the
dorsal midline long before hatching and after hatching move
laterally, separating in the dorsal midline (Fisher, 1958; this
study). The M. complexus of megapodes and chickens also
differed in the apparent lack of a temporary enlargement about
the time of hatching in megapodes. In chickens near hatching
this muscle appears swollen, protruding above the level of
adjacent cervical muscles and reaching a thickness of at least
2.5 mm, whereas in megapodes no swelling was observed and
maximal thickness was always less than 1 mm. Similarly,
although maximal width of the complexus muscle in each of
four chickens near hatching was 7 mm, in none of the mega-
podes did this width exceed 5-7 mm, which was reached only
in the largest specimens (e. g. Nos. 19, 30).

26 Postilla Yale Peabody Museum No. 78

Length measurements of the M. complexus were unreliable
due to the lack of a clear posterior boundary of the muscle.
When measurements of width and midline separation were ana-
lyzed relative to body size by plotting on arithmetic and double
logarithmic graphs, no indications of prehatching variations
other than growth and individual variations were detected for
the megapodes, but the precision of these measurements (about
+ 0.5 mm) is not very great relative to the dimensions meas-
ured. These observations do not eliminate the possibility of a

~

Figure 10. The Musculus complexus of domestic G@. gallus (19 day
embryo; ca. 12) and of Talegalla jobiensis (No. 30; ca. 14%).
Talegalla on the right.

transient enlargement of the M. complexus at hatching in meg-
apodes, but they provide no support for such a view. The
separation in the dorsal midline and apparent lack of special
enlargement of the complexus muscle at hatching in megapodes
are very likely correlated with the larger size of megapodes
at hatching.

The small egg tooth and unusual features of development
of the M. complexus of megapodes appear to be associated with
the different methods of hatching in megapodes and phasianids.

DISCUSSION AND CONCLUSIONS

Gallinaceous growth and maturation. The embryonic mega-
podes Leipoa after the first 20 days were relatively immature
compared with chickens of similar age. Although slow early
embryonic development is a_ reptile-like character, not too
much phylogenetic significance can be attributed to this con-

April 15, 1964 Ontogeny and Evolution in Megapodes 27

dition in Letpoa, since the slow developmental rate is asso-
ciated with the methods of incubation including relatively
low incubating temperatures. It is possibly phylogenetically
significant that Letpoa can hatch successfully (Frith, 1959)
at incubating temperatures so low (below 95°F) as to be lethal
for chicken embryos (Romanoff, 1960) ; however, data on the
normal range of egg temperatures of wild birds in general
(Huggins, 1941) about that megapodes are perhaps not
unusual among birds with respect to tolerated incubating tem-
peratures.

Interpretation of the chronology of embryonic megapodes
is complicated by great individual variation. For example, nor-
mal prehatching periods in Leipoa from different mounds range
from 50 to 90 days in association with intermound variations
from 96° down to 80°F in incubating temperatures (Frith,
1959). Since incubating temperatures of the megapode T'ale-
galla jobiensis (Ripley, 1964) are within the range for
Leipoa (Frith, 1959), it is possible, though unproven, that
Talegalla has an embryonic chronology similar to that of
Leipoa. Analysis of differences in embryonic chronology be-
tween megapodes and phasianids is further complicated by the
great interspecific variation among phasianids incubated at
100°F. For example, Colinus weighing 6 grams (egg weight,
9 @) and Phasianus weighing 18 grams (egg weight, 32 g)
are both hatched in 24 days, while chickens of 31 grams (egg
weight, 60 @) are hatched in only 21 days (Romanoff, 1960:
1143). Data are not available for a quantitative comparison
of the effects of varied incubation temperatures on the devel-
opment of chickens versus megapodes.

Both the phasianid Phasianus colchicus (Westerskov, 1957)
with an adult (male) weight of 1400 grams and the megapode
Alectura lathami (Coles, 1937) with a slightly higher adult
weight (Heinroth, 1922) reach full size about 25-30 weeks
after laying of the egg, indicating that the posthatching growth
of Alectwra is neither unusually fast nor slow compared with
that of phasianids.

The data of this study show that Leipoa and Tclegalla
before hatching undergo proportional and relative growth
analogous to that occurring up to several weeks posthatch-

28 Postilla Yale Peabody Museum No. 78

ing in other Galliformes. The similarity of relative growth in
young Galliformes is in agreement with the morphological
homogeneity of adults (cf. data of Verheyen, 1956). The rel-
ative growth appears, in this case, to be phylogenetically
generally more conservative than chronological growth. The
differences in relative growth of radius and humerus between
megapodes and phasianids do not indicate that either group
is more primitive than the other.

The noted interspecific variations in the size of embryos at
the first macroscopic appearance of certain structures may
represent interspecific differences in the growth of anlage of
these structures, for, as Schmalhausen (1926) and others have
pointed out, relative growth itself can produce qualitative
changes in form.

Although the weight of a bird at hatching is relatively
directly correlated with the weight of the egg (Heinroth,
1922), the ratio of the size of the egg relative to that of
adults often shows considerable intergeneric variation (Hein-
roth, 1922). Megapodes and certain small phasianids (e. g.
Coturnix) have eggs generally in the range from 8 to 18 per
cent of adult body weight in contrast to other phasianids and
turkeys with eggs weighing less than 5 per cent of adult body
weight (Heinroth, 1922).

The precocity of megapodes at hatching is associated with 1)
the large absolute egg size and correspondingly large size of
young at hatching together with 2) an embryonic relative
growth of the wing analogous to that occurring up to several
weeks posthatching in phasianids. No birds other than mega-
podes have large eggs plus extensive embryonic growth of the
wings.

Megapodes and reptiles. Portmann (1938) listed the fol-
lowing as primitive (reptile-like) traits of megapodes: lack of
natal downs, possible lack of an egg tooth at hatching, absence
of parental care for young, eggs incubated in sand by solar
heat, long incubation period, large clutch size, slow growth to
adult size, and precocity of young at hatching. However, as
shown by my study, megapodes do have natal downs, and at
least some species have egg teeth. Furthermore, there is no

April 15, 1964 Ontogeny and Evolution in Megapodes 29

good evidence for an especially slow posthatching growth of
megapodes.

Moreover, the many adaptive interrelationships (coadapta-
tions) of the reptile-like characters of megapodes should be
considered. For example, the long incubation period is cor-
related with the methods of incubation and the large size and
precocity of young at hatching. The precocity of young is also
correlated with the lack of parental care which in turn is asso-
ciated with the incubating methods and clutch size. The reptile-
like traits of megapodes all belong to one, or perhaps two,
group(s) of coadapted characters. Considered in this way, the
evidence for special affinities of megapodes and reptiles is uncon-
vincing, since the points of similarity are all related to com-
mon reproductive adaptations.

The case for special reptilian affinities of megapodes would
be greatly strengthened if there were reptile-hke characters
relatively independent of the central adaptation in megapodes :
however, no such characters have yet been found. As one exam-
ple, there is reported to be a significant difference in the caloric
values of reptilian and avian egg yolks (Slobodkin, 1962), yet
samples of yolk collected during this study from relatively
fresh eggs of Letpoa and Gallus had values agreeing with
those of other avian species (Slobodkin, 1962).

Furthermore, advocates of the primitiveness of the mega-
podes among birds as a whole have generally failed to analyze
the possibility of convergent evolution. In short, evidence for
the primitiveness of megapodes among birds as a whole is
unacceptable.

Evolution of the megapode family. Megapodes are basi-
cally similar in morphological development to phasianids. Dif-
ferences in the structure of natal downs, in absolute and rela-
tive sizes of eggs, in sizes of subcutaneous fat bodies, in develop-
ment of the hatching apparatus, etc., are all directly or
indirectly correlated with the sizes of the young at hatching.

Huxley (1868) emphasized that, in contrast to other Galli-
formes, megapodes and cracids are alike in depth of the sternal
notches and in position of the hallux. From this anatomical
basis, he postulated that these forms, isolated respectively in

30 Postilla Yale Peabody Museum No. 78

the Australian and Neotropical regions, are remnants of an
ancestral gallinaceous stock which has been replaced through
most of the Old World and Nearctic region by more modern
Galliformes.

However, the differences at hatching in feather structure be-
tween cracids and megapodes support the generalization that
megapodes and cracids are not especially closely related in
evolution, contrary to some current classifications (e. g. Peters,
1934).

The contemporary megapodes are characterized by 1) rha-
chidial natal downs on the body, 2) long juvenal remiges and
large body size at hatching, 3) a relatively high ratio of egy
to adult weights compared with other Galliformes, and 4) the
unicarotid condition; it is likely that these distinctive traits
were present in a population ancestral to all living megapodes.
Megapodes are apparently unique among birds in having such
long and weak natal downs preceding the embryonic juvenal
remiges. These weak natal downs are clearly vestiges rather
than preadaptations and indicate the evolution of megapodes
from unknown gallinaceous ancestors possessing a natal plum-
age and less precocious chicks resembling those of extant phasi-
anids.

This phylogenetic interpretation is also supported by the
finding of a vestigial egg tooth and the apparent lack of special
enlargement of the complexus muscle at hatching; these fea-
tures strongly indicate an evolutionary origin of megapodes
from forms less precocious at hatching. One aspect of the evolu-
tion of megapodes has been the transition from the use of the
egg tooth in hatching to kicking open the shell.

The variation in the number of carotid arteries in birds as
a whole (Table 7) appears to be due to much convergent evolu-
tion. The most readily conceived sequence is a loss of one
carotid artery (Glenny, 1955), but a possible evolutionary
increase cannot be excluded. The occurrence of only one carotid
in megapodes in contrast to two in all other known Galliformes
suggests that megapodes are specialized in this respect.

My conclusions, based on morphology, are compatible with
the concept of Mainardi and Taibel (1962: Fig. 4), based
largely on erythrocyte antigens, that megapodes, cracids, and

April 15, 1964 Ontogeny and Evolution in Megapodes 31

phasianids have evolved as three separate lines from unknown
gallinaceous ancestors.

It is pertinent that there are living forms intermediate in
structure of feathers at hatching and in precocity of young
between megapodes and phasianids such as Phasianus or Gallus.
For example, the phasianid genus T'ragopan has natal downs
with short rhachises (this study), relatively long juvenal remi-
ges at hatching (Beebe, 1918), and initial flight on the third
day posthatching (Nice, 1962; after the Heinroths). Although
Tragopan probably does not represent the phylogenetic ances-
tors of megapodes, certain aspects of its structure and behavior
of young aid in visualizing the evolutionary origin of the mega-

podes.

Evolution within the megapodes. Megapodius and Macro-
cephalon lay their eggs in holes (Megapodius also uses mounds )
and are known to lay their eggs communally, while the four
other genera use mounds exclusively as far as known. (In
accord with the study of Ripley (1964) the form Ewlipoa
wallace is here included in the genus Megapodius.)

The specialized Macrocephalon is somewhat intermediate in
adult proportions of wing, tarsus, and tail between other large
megapodes (4 genera) and the smaller Megapodius (data in
Ogilvie-Grant, 1893). The relatively uniform color of Megapo-
dius and its relative simplicity of nesting habits have led some
authors (e. g. Becker, 1959) to consider Megapodius primitive
among the megapodes. The uniform color pattern of Mega-
podius resembles that of Aepypodius or Talegalla and may
indeed be a primitive trait among living megapodes. But sim-
plicity of nesting site (e. g. the incubation of eggs in holes in
the ground) does not necessarily imply primitiveness as illus-
trated by the specialized brood-parasitic avian species which
also build no nests.

Since one trait of the megapodes is the relatively high ratio
of egg weight to adult weight, and since megapodes have evolved
from apparently more conventional gallinaceous ancestors, it 1s
likely that, during megapode evolution, sizes of eggs increased
relative to adult size. Although megapode evolution has very
likely also involved an increase in the absolute size of eggs and

32 Postilla Yale Peabody Museum No. 78

chicks at hatching, the absolute sizes of newly hatched young
do not necessarily indicate the relative primitiveness of the
contemporary megapodes. Indeed, if, as seems likely, the evolu-
tion of megapodes has involved an increase in the absolute size
of eggs and hence of young at hatching, then a large ancestral
adult would have been better preadapted, in terms of size, than
a small ancestral adult for the evolution of larger absolute sizes

of eggs.

More critical features suggesting the direction of evolution
within the megapodes are the proportions at hatching. In this
respect Megapodius is more remote than T'alegalla or Leipoa
from the conditions in non-megapode Galliformes. In view of
the relatively shorter bill and longer wing at hatching and the
unusual webbing of the toes in Megapodius, the simplest hypo-
thesis is that Megapodius has secondarily evolved from a form
like T'alegalla or Aepypodius. Thus Megapodius, perhaps most
reptile-like of the megapodes in certain respects, is structurally
specialized.

The small size (and relatively short culmen) of adult Mega-
podius appear to be adaptive in reducing potential ecological
competition where Megapodius and other megapode genera
occur sympatrically (Ripley, 1960). From the present study
it is apparent that a shorter culmen and smaller body size at
hatching also characterize Megapodius when compared with
other megapodes.

Megapodius and Macrocephalon have possibly primitive char-
acters in the occurrence of diastataxy (variable in Megapo-
dius) and the tufted oil gland. Distribution of these characters
in birds as a whole (Table 7) indicates that there is no neces-
sary correlation in the presence of these features and that they
have been subject to considerable convergent evolution. Despite
the contention of Steiner (1918, 1956) that diastataxy is prim-
itive because it occurs in “primitive” birds, there is no con-
vincing evidence against the possibility that diastataxy might
evolve from eutaxy (see Humphrey and Clark, 1961, for a
review of the various hypotheses on the origin of diastataxy).
Similarly, there is no reason to assume that a tufted oil gland
is necessarily primitive.

April 15, 1964 Ontogeny and Evolution in Megapodes 33

In view of the intraspecific constancy of tarsal scutellation
and its intergeneric variation in the megapodes, it appears use-
ful in dividing the megapodes into subgroups ; however, in view
of the range of variation within the megapode family, it would
probably be unwise to emphasize this feature in attempting to
determine the affinity of megapodes with other gallinaceous
families.

\lll

Megapodius Leipoa

Alectura

=
Talegalla =

Aepypodius =

L
Macrocephalon

Stem megapode
population

Pheasant-like
gallinaceous
ancestors

Figure 11. Provisional phylogeny of the family Megapodiidae. The
smaller branches leading from the genera represent speciation.

From these considerations, the first phylogeny to cover inter-
generic relationships within the megapodes has been developed
(Fig. 11). The ancestral stem population (Fig. 11) would
have possessed large adult and chick sizes, like Talegalla,
rhachidial natal downs, a relatively long culmen at hatching,
and egg laying in mounds. If this phylogeny is correct, then
current classifications (e. g. Peters, 1934) are misleading in
placing Megapodius first in the sequence of megapode genera.

In examining megapode development, I have found no charac-
ters indicating that megapodes are especially primitive birds ;
indeed, the evidence demonstrates the specialized nature of
megapode ontogeny which has probably evolved from a phasi-
anid-like condition.

34 Postilla Yale Peabody Museum No. 78

ACKNOWLEDGMENTS

Valuable suggestions and constructive criticism were given
by Professors S. D. Ripley, G. E. Hutchinson, J. L. Brooks,
and J. P. Trinkaus. Dr. E. J. Boell gave much excellent coun-
sel. Dr. P. S. Humphrey initially suggested the topic and
provided many helpful suggestions. Dr. H. J. Frith, of the
Wildlife Survey Section, Australian Commonwealth Scientific
and Industrial Research Organization, made arrangements
which greatly facilitated my field collecting. I am also much
indebted to Dr. D. Amadon, the American Museum of Nat-
ural History, Mr. B. K. Brown, Dr. E. T. Gilliard, Mr. D.
Heath, Mr. E. A. Heath, Dr. H. Levene, and many others
who have helped in a variety of ways. Mrs. Shirley Hartman,
Mr. G. di Palma, and my wife prepared the figures. I am
especially grateful to my wife for her encouragement and aid.

Financial support was provided by the National Science

Foundation Grant G-10735, awarded to Dr. S. D. Ripley.

SUMMARY

Many differences found in development between megapodes
and phasianids are associated with megapodes having before
hatching proportional and relative growth equivalent to that
occurring up to several weeks posthatching in phasianids.

Contrary to published reports, megapodes at hatching bear
juvenal remiges and natal downs on the body and are thus like
other Galliformes, although there are structural differences in
the natal downs. Vestigial natal downs preceding the embry-
onic juvenal remiges indicate that megapodes evolved from
forms with more conventional gallinaceous feathering at hatch-
ing and less precocious young.

This interpretation of megapodes as evolutionarily special-
ized is also upheld by their vestigial egg teeth and apparent
lack of a special enlargement of the complexus muscle which -
aids in the hatching of other Galliformes.

Compared with other megapode genera and other Gallifor-
mes, young juvenile Megapodius have a long wing and unus-
ually short bill. It is therefore concluded, contrary to published

April 15, 1964 Ontogeny and Evolution in Megapodes 35

reports, that, despite its apparent simplicity in color pattern
and egg laying habits, Wegapodius is specialized among mega-
podes.

A phylogeny of the megapode genera is proposed on the
basis of proportions at hatching, tarsal scutellation, foot web-
bing, eutaxy, oil gland feathering, and other characters.

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164-177.

Becker, R., 1959. Die Strukturanalyse der Gefiederfolgen von Megapodius
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Beebe, W., 1918. A monograph of the pheasants. Vol. 1. Witherby, London.
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Bergmann, S., 1961. My father is a cannibal. Hale, London. 192 p.

Blaszyk, P., 1935. Untersuchungen iiber die Stammesgeschichte der
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Campbell, A. J., 1901. Nests and eggs of Australian birds. Pt. 2: 525-1102.
Sheffield, England.

Clark, G. A., Jr., 1960. Notes on the embryology and evolution of the
megapodes (Aves: Galliformes). Yale Peabody Mus. Postilla no. 45:
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Clark, G. A., Jr., 1961. Occurrence and timing of egg teeth in birds. Wilson
Bull. 73: 268-278.

Coles, C., 1937. Some observations on the habits of the Brush Turkey
(Alectura lathami). Proc. Zool. Soc. Lond, 107A: 261-273.

Fant, R. J., 1957. Criteria for aging pheasant embryos. J. Wildl. Mgt. 21:
324-328.

Fisher, H. I., 1958. The “hatching muscle” in the chick. Auk 75: 391-399.

Friedmann, H., 1931. Observations on the growth rate of the foot in the
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[ie 14

Frith, H. J., 1959. Breeding of the Mallee Fowl, Leipoa ocellata Gould
(Megapodiidae). CSIRO Wildl. Res. 4: 31-60,

Frith, H. J., 1962. The Mallee-fowl. Angus & Robertson, Sydney. p. xii +
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Glenny, F. H., 1955. Modifications of pattern in the aortic arch system of
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Hall, R., 1901. Descriptions of the young of some Australian birds. Vict.
Nat. 18: 19-25.

Hamilton, H. L., 1952. Lillie’s development of the chick. 3rd ed. Holt,
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Heinroth, O., 1922. Die Beziehungen zwischen Vogelgewicht, Eigewicht,
Gelegegewicht und Brutdauer. J. f. Orn. 70: 172-285.

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Heinroth, O., 1931. Beobachtungen bei der Aufzucht eines Knopfschnable-
Hokko’s (Crax globicera) und eines Mitu’s (Mitua mitu). J. f. Orn.
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Heinroth, O. and M. Heinroth, 1928. Die Vogel Mitteleuropas. Bd. 3.
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Huggins, R. A., 1941. Egg temperatures of wild birds under natural condi-
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Lyon, D. L., 1962. Comparative growth and plumage development in
Coturnix and Bobwhite. Wilson Bull. 74: 5-27.

Mainardi, D., and A. M. Taibel, 1962. Studio immunogenetico sulle
parentele filogenetiche nell’ordine dei Galliformi. Inst. Lombardo, Rend.
Se., B 96: 131-140.

Meyer, O., and EK. Stresemann, 1928. Zur Kenntnis der Entwicklung von
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Ogilvie-Grant, W. R., 1893. Catalogue of the game birds .. . of the
British Museum. p. 445-472.

Padgett, C. S., and W. D. Ivey, 1960. The normal embryology of the
Coturnix Quail. Anat. Record 137: 1-11.

Peters, J. L., 1934. Check-list of the birds of world. Vol. 2. Harvard Univ.
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LHe

YALE PEABODY MUSEUM

oF NaTurAL History

Number 79 April 20, 1964 New Haven, Conn.

NOTEWORTHY AMPHIPODA (CRUSTACEA) IN THE
COLLECTION OF THE YALE PEABODY MUSEUM

if
Eric L. Mitts (

DEPARTMENT OF BiIoLocy UEEN’s Universtry, Kineston, ONTARIO
’ 4 S)

During the last half of the nineteenth century the Peabody
Museum of Natural History, Yale University, was the focus of
much marine biological research. A. E. Verrill and 8. I. Smith
amassed a considerable collection of North American inverte-
brates and exchanged specimens with European biologists. In
the course of my curatorial work on the amphipod crustaceans
in this collection a number of interesting locality records and
type specimens have come to light.

The type specimens are primarily those of S. I. Smith and
B. W. Kunkel. Smith apparently based his descriptions on sev-
eral specimens of each species and designated no types, so I
have chosen lectotypes where this was warranted by the condi-
tion and locality data of the collections. The type terminology
used follows the recommendations of the International Code of
Zoological Nomenclature (Stoll et al., 1961). Four of Smith’s
species are redescribed here from lectotype specimens. They are
Melita nitida, Ampithoe longimana, A. valida and Cymadusa
compta. These are common New England or Middle Atlantic
Coast species, so these redescriptions should be helpful for com-

2 Postilla Yale Peabody Museum No. 79

parative purposes. A generic diagnosis has been included for
each redescribed species.

I have examined some probable type specimens of Bermuda
amphipods described by Kunkel (1910), including those of
Pariphinotus tuckeri, Ceradocus colei, C. parkeri, Elasmopus
magnispinatus, Eusiroides verrilli and Ampithoe pollex. The
poor condition of the specimens makes designation of lecto-
types inadvisable for all species except Husiroides verrillt.

Paratypes of the recently described species Ampelisca vado-
rum and specimens of Listriella clymenellae from the type local-
ity are also present in the Peabody Museum collections and are
listed here. Interesting locality records for a number of other
species are also included in this paper.

Professor W. D. Hartman encouraged the curatorial and
systematic work on these collections. I am grateful for his sup-
port and for the stimulus of conversations on systematic zool-
ogy. Dr. Fenner A. Chace, Jr., of the United States National
Museum, helped me with problems of type nomenclature. This
work was supported financially under NSF grants G-10772 and
GB-583 to Yale University.

Abbreviations used in the species discussions are: YPM-Yale
Peabody Museum catalogue number; USFC—United States
Fish Commission; USNM—United States National Museum
accession number.

Family AMPELISCIDAE G.O. Sars
Ampelisca vadorum Mills

Ampelisca vadorum Mills, 1963, p. 971-989, figs. 1-3.

Material: YPM 5642-Sta. 1, Long Island Sound, Conn.,
41°11.1’N, 73°08.8’W. Coarse sand, depth 10-12 meters.
Dredge with stramin liner. 24 May 1962. S. W. Richards,
collector. 2 adult ¢ 6 10 adult 2 2, paratypes. YPM 5643—
same data 10 juv. 6 ¢, 10 juv. 2 2, paratypes.

Ampelisca spinimana Chevreux

Ampelisca spinimana Chevreux, Chevreux and Fage, p. 81-82,

we

fig. (o.

April 20, 1964 Peabody Museum Amphipoda 3

Material: YPM 4698—USNM 37152. Sta. 36B, (Steamer
“Bache”’) 42°18’N, 69°49’W, 23 14 mi. ENE 14 N from Race
Pt., 142 fm. (260 meters). Soft blue mud. Sept. 1873. 1
specimen.

This species has been recorded only recently from North
America and appears to be widespread on the continental slope

(Mills, 1963).

Ampelisca eschrichti Kr¢yer

Ampelisca eschrichti Krgyer, Gurjanova, 1951, p. 307-308,
fig. 170.

Material: YPM 500845 fm. (82 meters). Georges Bank,
41°25'N, 66°45’W, 16 Sept. 1872. S. I. Smith and Oscar
Harger. 1 juv. ¢.

Seven lots of this species are present in the Yale collections.
Six are from boreal and subarctic waters of Labrador and the
Gulf of St. Lawrence. However, one collection, described above,
is a southern record for the species on the East Coast of North
America. Smith and Harger (1874) record ‘‘Ampelisca sp.”
in this haul on the east side of Georges Bank. Ampelisca macro-
cephala is known to be common on Georges Bank (Roland L.
Wigley, personal communication), but the present species must
be rare or absent now, with the recent warming of New England
waters. (Taylor, Bigelow and Graham, 1957).

Family PHLIANTIDAE Stebbing
Pariphinotus tuckeri Kunkel

Pariphinotus tuckeri Kunkel, 1910, p. 19-21, fig. 6.

Material: YPM 5613—Bermuda, 1901, W. G. Van Name.
broken ¢;1 9.5.5 mm.

Kunkel’s (1910) description was undoubtedly based on these
two specimens. Since the specimens were dessicated and the

male now lacks head and urosome Kunkel’s description must
stand.

A Postilla Yale Peabody Museum No. 79

Family LILJEBORGIIDAE Stebbing
Listriella clymenellae Mills
Listriella clymenellae Mills, 1962, p. 158-162, figs. 1, 2.

Material: YPM 4492—Sta. F (Sanders), Barnstable Har-
bor, Mass., sand flats in Clymenella tubes, August, 1960, H. L.
Sanders. 5 specimens. YPM 4493—Sta. A (Sanders), Barn-
stable Harbor, Mass., in tube of Clymenella torquata, 5 August
1959, Eric L. Mills. 1 specimen.

These specimens were collected at the type locality (Mills,
1962.)

Family GAMMARIDAE Leach
Crangonyx pseudogracilis Bousfield
Crangonyx pseudogracilis Bousfield, 1958, p. 102-105, fig. 16.

Material: YPM 5473—Axelshop Pond, Mount Carmel, Conn.,
weedy area, 15 Oct. 1961, Eric L. Mills. 9 juveniles.

This is apparently the first record under this name for Con-
necticut. Bousfield (1958) suggested that the species would be
present in Atlantic state watersheds. Kunkel’s (1918) records
of Crangonyx gracilis Smith from New Haven, Conn., and
Providence, R. I. (as Eucrangonye gracilis), are partially ref-
erable to C. pseudogracilis (Bousfield, 1958). Bousfield has
shown that Crangonyx gracilis has been authentically recorded
only from the Great Lakes basin, while C. pseudogracilis is a
widespread species of warm, shallow fresh waters.

Genus Melita Leach 1813/1814 (cited by Stebbing, 1906.)

Stebbing (1906, p. 421) defines the genus as follows:

“Body slender, peraeon smooth, pleon with one or more of the
segments dorsally dentate and armed with bristles. Head not ros-
trate, lateral corners rounded. Side-plate 4 the largest, emarginate
behind. Eyes usually distinct, rather small. Antenna 1 slender,
longer than antenna 2, Ist and 2nd joints rather long, 3rd not
short, with accessory flagellum. Mouth-parts, so far as known, nor-
mal. Upper lip with small central emargination. Lower lip, inner

April 20, 1964 Peabody Museum Amphipoda 5

lobes tolerably distinct. Mandibular palp rather slender. Maxilla
1, inner plate with several setae, outer with 11 spines. Maxilla 2,
inner plate sometimes with setae on inner margin. Maxillipeds, outer
plates with teeth on inner margin, passing into slender spines on
apex. Gnathopod 2 larger, often unequal, and one in the male some-
times larger than the other, sometimes approximately chelate. Perae-
pods 3-5, 2nd joint well-expanded. Peraeopods 4 and 5 subequal,
longer than others. Branchial vesicles simple. Marsupial plates nar-
row. Uropod 2 the shortest. Uropod 3 projecting much beyond the
others, outer ramus long, 2nd joint wanting or rudimentary, inner
ramus very short. Telson small, deeply cleft. Some characters sub-
ject to much variation within the species.”

Melita nitida Smith
(Fig. 1)

Melita nitida Smith, (in: Verrill and Smith, 1873, p. 560-561).
Della Valle, 1893, p. 716. Holmes, 1905, p. 505-506, figs.
Kunkel, 1918, p. 99-100, fig. 22. Shoemaker, 1935a, p. 70-
al, tig, 2.

Diagnosis: 1) Gnathopod 1 propodus oblong, palm trans-
verse, with projection defining lower corner. 2) Gnathopod 2
propodus, palm and lower margin forming a single even curve.
3) Hind margins of peraeopod 3-5 bases finely serrated and
setose. 4) Pleon and urosome not dentate, with only a few
spines on each side of the midline of urosome segment 2.

Description: Mave (8.7 mm) LECTOTYPE. Fig. 1 (a-p). Head
only as long as first two body segments combined; interantennal lobe
large, rounded above, rather square below. First antenna stout, a
little longer than second; peduncle about equal in length to flagel-
lum; flagellum with 20 articles; accessory flagellum of 3 articles.
Second antenna stout; 4th peduncle segment with a few small ven-
tral setae; 5th peduncle segment with several groups of long curved
setae on ventral margin; flagellum of 12 articles, about half again
as long as 5th peduncle segment.

Epistome finely setose and slightly angular below, but not medially
notched or emarginate. Mandible, incisor process with 3 teeth, lacinia
mobilis finely dentate, molar process strong, projecting outwards ;
palp, second segment with 3 groups of marginal setae, 3rd segment
inner margin lined with long setae. Lower lip inner plates not fully

6 Postilla Yale Peabody Museum No. 79

distinct from outer plates; both plates finely setose on the medio-
distal margins. Maxilla 1, inner plate distal margin oblique, with
several setae; outer plate with 9 pectinate spines; palp, second seg-
ment with 5 spines and several setae. Maxilla 2, inner and outer
plates equal in length, inner with distal margin oblique and heavily
setose, outer with distal margin transverse and setose. Maxilliped,
inner plate slightly emarginate distally with an oblique row of setae;
outer plate inner margin evenly set with 11 small spines which
grade distally into 3 long stout spine setae; palp curved distally,
second segment setose medially, 3rd segment sparingly setose.

Coxal plates rather shallow, first 3 deeper than wide, oblong,
rounded distally; fourth with a posterodistal lobe rounding evenly
into the distal margin; fifth and sixth small, with rounded anterior
and posterior lobes; seventh small, rounded below.

Gnathopod 1, basis flattened and slightly expanded; carpus longer
than propodus, lower margin with several groups of long, strong
setae; propodus oblong, slightly curved, dactyl arising in middle of
almost transverse palmar margin; propodus palmar margin defined
by finely setose prominence at corner; dactyl very short, stout. Gna-
thopod 2, propodus greatly expanded, thick, palmar margin set with
spinules, rounding evenly into an almost straight lower margin;
dactyl long, stout, when closed fitting into a long oblique groove
lined with setae on inner side of propodus.

Peraeopods 1 and 2 almost equal in size, short, rather slender.
Peraeopods 3, 4 and 5 very similar, highly spinose, bases with groups
of spines on anterior margin, posterior margins finely serrated and
setose; peraeopod 3 the shortest; peraeopods 4 and 5 about equal in
length.

Epimeral plate posterior margins setose, first two slightly rounded
posteriorly, with a small tooth at the posterodistal corner ; third with
square or slightly toothed posterodistal corner. Pleopod peduncles
and rami slender; coupling spines 2, hook-shaped. Dorsal pleonal
and urosomal teeth absent.

Urosome short, second segment with 3 to 5 articulated spines
on either side of the dorsal mid-line. Uropod 1 extending slightly
beyond uropod 2, peduncle margins with strong spines, rami with
apical spine groups; outer ramus outer margin with 4 spines, inner
margin 2 spines; inner ramus outer margin with 3 spines, inner mar-
gin with 5 spines. Uropod 2, peduncle margins spinose, rami with
apical spine groups, outer ramus outer margin with 4 spines, inner
margin with 2 spines; inner ramus outer margin with one spine,
inner margin with 3 spines. Uropod 8 peduncle stout; outer ramus

April 20, 1964 Peabody Museum Amphipoda u

very long, margin and apex with groups of strong setae; inner ramus
very small, with 2 apical spines, set in a sinus of the peduncle.
Telson short, deeply cleft and spread, lobes with 2 to several spines
apically.

Femae (9.3 mm) PARALECTOTYPE. Fig. 1 (u-z, aa). Very
similar to male in most features. Antennae slightly shorter, but fla-
gellum articles same in number as in male. Gnathopod 1 smaller
than in male, propodus shape similar. Gnathopod 2 smaller than in
male, carpus lower margin longer and with more groups of stout
setae; propodus similar in shape to that of male, but palmar margin
minutely serrated and with a stout spine at the lower corner; propo-
dus lacking setose groove into which dactyl of male fits; dactyl with
a tooth near the tip. Peraeopod 4 coxa projected into a curved,
finger-like lobe anteriorly, rather than the short, rounded lobe of the
male.

Type locality: Great Egg Harbor, New Jersey.

Material: YPM 1247-Great Egg Harbor, New Jersey.
A. E. Verrill and S. I. Smith. April 1871. “Melita polita.” 1
lectotype ¢, 8.7 mm, 8 paralectotype ¢ ¢ (one dissected by
Kunkel, 1918), 5 paralectotype ? 2. YPM 4897—New Haven,
Connecticut, S. I. Smith. 11 6 6,8 2 2,5 juv. YPM 4899—
probably New Jersey or Long Island Sound. S. I. Smith. No
data. “Melita polita.” 1 3, 4 juv.

Discussion: Shoemaker (1935a) figured specimens of this
graceful species from Sinaloa, Mexico. The type specimens
described here have been refigured for comparative purposes,
although Shoemaker’s figures characterize the species well. At
the time of Shoemaker’s paper the species was known from
Cocos Island, Ecuador, Panama, the west coast of Mexico.
and from Louisiana to Cape Cod (see Pearse, 1913, and Kunkel,
1918). E. L. Bousfield (personal communication) has collected
the species as far north as Pugwash, Nova Scotia.

Ceradocus colei Kunkel

Ceradocus colei Kunkel, 1910, p. 41-48, fig. 15

8 Postilla Yale Peabody Museum No. 79

Material: YPM 4532—-no data. Specimen dry, examined
1961. Probably Bermuda (no locality label with specimen.)
1 2, about 5 mm.

This was probably the “single female specimen” mentioned
in Kunkel’s description. Over the course of the years the speci-
men dried out and has broken into three pieces. New figures of
the species are essential to replace Kunkel’s meagre drawings,
but they could not possibly be made from this material.

‘6

Ceradocus parkeri Kunkel

Ceradocus parkeri Kunkel, 1910, p. 39-41, fig. 14.

Material: YPM 4534—no data. Specimen dried out. Ber-
muda? (No locality label with specimen.) 1 ¢, 6 mm.

This specimen lacks most of the head appendages [ appar-
ently lost since Kunkel’s (1910) description] and is not satis-
factory for a redescription.

Elasmopus magnispinatus Kunkel

Elasmopus magnispinatus Kunkel, 1910, p. 54-56, fig. 20.

Material: YPM 4543-no data. Probably Bermuda, 1901,
ASE. Verrill 292): 8 juvend on

Kunkel’s description and figures must stand, since his original
specimens are in poor condition.

Gammarus (Gammarus ) fasciatus Say

Gammarus (Gammarus) fasciatus Say, Bousfield, 1958, p. 69-
72, fig. 4.

Material: YPM 4506—Mashpee River, near Route 28, Cape
Cod, Mass. Weed, cool stream. 16 July 1961. Eric L. Mills,
collector. 1 ovig. 2, 12 juvs. YPM 4529—Mill River below
Axelshop Pond, Mount Carmel, Hamden, Conn. Roots and
weeds, edge of river. 15 Oct. 1961. Eric. L. Mills, collector. 28
juvs. YPM 4628—Woodbridge, Conn. A. E. Verrill, 5 specimens.

April 20, 1964 Peabody Museum Amphipoda 9

These three collections help to fill in the range and occur-
rences listed by Bousfield (1958). The Woodbridge, Conn. col-
lection is undoubtedly that mentioned by Kunkel (1918, p.
107). Smith (1874a) first documented the occurrence of the
species in Connecticut.

Gammarus (Rivulogammarus ) lacustris lacustris G. O. Sars.

Gammarus robustus Smith, 1874b, p. 610, pl. 2, fig. 7-12; Bar-
nard, 1958, p. 55.

Gammarus limnaeus Smith (in part), Shoemaker, 1920, p. 16.

Gammarus lacustris lacustris, Bousfield, 1958, p. 80-81, fig. 8.

Material: YPM 4876—“Gammarus robustus,’” Wasatch
Mountains (Utah). L. E. Ricksecker, collector. 3 broken 2 2.

The status of Smith’s Gammarus robustus has remained a
mystery since its description, although Shoemaker (1920)
placed it with G. limnaeus (= G. lacustris) and Bousfield
(1958) suspected that this move was correct. Smith’s original
specimens have now come to light in the Yale collections, and,
after examining them, Dr. Bousfield has informed me that they
are females of G. 1. lacustris. The above synonymy establishes
the status of Smith’s species with certainty.

Family PLEUSTIDAE Stebbing
Neopleustes pulchellus (Kr¢yer )

Neopleustes pulchellus (Kréyer), Gurjanova, 1951, p. 643-645,
figs. 439, 440.

Material: YPM 1301—USFC, (Bl), 50 fm. (92 meters),
1872. USNM 35646. Georges Bank, 41°25’N, 66°25’W. S. I.
Smith and O. Harger. 1 ?.

This is apparently the southernmost record in New England.
Holmes (1905) recorded the species (as Paramphithoe pul-
chella) from Grand Manan, New Brunswick, and Shoemaker
(1930) includes Casco Bay, Maine, in his distribution records.
As mentioned in the case of Ampelisca eschrichtt Krgyer, warm-

10 Postilla Yale Peabody Museum No. 79

ing trends in New England waters may make Georges Bank
unsuitable at present for some arctic and subarctic species,

perhaps including Neopleustes pulchellus.

Family ATYLIDAE G. O. Sars
Atylus swammerdami (H. Milne-Edwards )

Paratylus swammerdami (H. Milne-Edwards), G. O. Sars,
1895, p. 463-465, pl. 163.
Atylus swammerdami, Barnard, 1958, p. 31.

Material: YPM 5632—-Sta. 1231. “Fish Hawk.” Vineyard
Sound Lightship, W by N 14 N; Cuttyhunk N by W 3% W;
Gay Head SE % E. 16 fm. (29 meters). Sand. 29 Aug. 1887.
1 juv., 6 mm.

There are apparently no other records of Atylus swam-
merdami from North America. This specimen agrees well with
specimens from the west of Ireland collected by Canon A. M.
Norman about 1866 (YPM 5620), but differs slightly from
Sars’ (1895) figures in having a more convex hind margin of
the peraeopod 5 basis and a setose hind margin of the peraeo-
pod 4 basis which is not projected at the posterodistal corner.

Other species of the genus are apparently not common on the
East Coast of North America, although A. carinatus (Fabri-
cius) occurs as far south as the St. Lawrence estuary (Brunel,
1961b), and Shoemaker (1932) summarizes occurrences of
Nototropis (now Atylus) minikot Walker from Chesapeake
Bay to Brazil.

Family EUSIRIDAE Stebbing
Eusiroides verrilli (Kunkel)
Pontogeneia verrilli, Kunkel, 1910, p. 29-31 fig. 10.

Eusiroides verrilli, Schellenberg, 1929, p. 273-282.

Material: YPM 5338—#11. Castle Harbor. Bermuda? A. E.
Verrill. 1 lectotype 2, 9 mm, (figured by Kunkel) ; 2 paralecto-

April 20, 1964 Peabody Museum Amphipoda 11

type adult ¢ ¢,8 and 10 mm; 32 paralectotype ? 2 ; 3 para-
lectotype juveniles.

A dissected adult female, 9 mm, is apparently the specimen
used by Kunkel (1910, fig. 10) for his description of the species
(as Pontogeneia verrilli). It is designated a lectotype. Kunkel’s
description and figure seem adequate, so the species has not been
redescribed.

Family PODOCERIDAE Stebbing
Dulichia spinosissima Kr¢gyer

Dulichia spinosissima Kr¢éyer, Gurjanova, 1951, p. 990-991,
fig. 688.

Material: YPM 5618—Sta. 81. West Harbor, Fisher’s Is.,
N.Y. Fisher’s Is. Sound. Sand and shells. 314 fm. (6.4 meters).
10 Aug. 1874. USFC. 1 broken juv. ¢.

Brunel (196la) records this species from the Gulf of St.
Lawrence. The present specimen is apparently the southern-
most record of the species.

Family AMPITHOIDAE Stebbing

Genus Ampithoe Leach 1813/1814 (cited by Stebbing, 1906.)

Stebbing (1906, p. 631) defines the genus as follows:

“Head without rostrum. Side-plates 1-5 well developed, 5th as
wide as 4th, with a very small hind lobe. Antenna 1 without acces-
sory flagellum, usually longer than antenna 2, though with shorter
peduncle. Mouthparts prominent below the head. Upper lip distally
rounded. Lower lip ... inner lobes well developed; outer lobes
bifid; mandibular processes prominent. Mandible normal, principle
and secondary plate multidentate, spines in spine row numerous,
molar of moderate size; 3rd joint of palp sometimes widened dis-
tally and crowded with setae, at others not widened and slightly
armed. Maxilla 1, inner plate very small, usually with 1-3 setae,
outer plate with 10 spines; 2nd joint of palp with several spical
spines. Maxilla 2, outer plate the larger, inner distally-narrowed,
inner margin not very elongate. Gnathopod 1 subchelate, usually
the smaller. Gnathopod 2 usually subchelate, stronger in ¢ than in

12 Postilla Yale Peabody Museum No. 79

? and generally of a different shape. Peraeopods 1 and 2, 2nd joint
expanded, sometimes greatly, for the cement glands, the secretion
from which issues through the apex of the finger to supply fibres
for constructive purposes. Peraeopods 4 and 5 longer than the
others. Uropod 3, outer ramus carrying 2 reverted spines. Telson
short, usually or always having the angles of the apex minutely

hooked.”

Ampithoe longimana Smith
(Figs. 2, 3)

Amphithoe longimana Smith, (in: Verrill and Smith, 1873, p.
563). Holmes, 1905, p. 509, figs. Kunkel, 1910, p. 87, fig.
34. Kunkel, 1918, p. 147-149, fig. 43.

Ampithoe longimana Smith, Stebbing, 1906, p. 634, 738.
Pearse, 1913, p. 376. Barnard and Reish, 1959, p. 36-37,
pl. 12.

Diagnosis: 1) Antenna 2 peduncle segments 4 and 5 of ¢
long. 2) Gnathopod 1 propodus of é long, even in width. 3)
Uropod 3 rami very short, rounded. 4) Peraeopod 5 basis con-
vex, spine at posterodistal angle.

Description: Mare (10.1 mm) LECTOTYPE Fig. 2 (at), 3
(u). Head short, length less than first two body segments combined ;
interantennal lobe with rounded margins, rather prominent; inferior
antennal sinus with oblique, gently-rounded posterior margin; eye
yellowish-brown in alcohol, with about 50 facets.

Antenna | as long as body; peduncle extending a little beyond
4th peduncle segment of antenna 2, Ist peduncle segment with a
few setae and one short spine distally, 2nd segment longer than
Ist, lightly setose; 3rd segment about 1% length of second; flagel-
lum 114, to 2 times length of peduncle, of 29 articles. Antenna 2
stout; peduncle long, 4th segment about equal to first two segments
of antenna 1 peduncle in length, 5th segment slightly longer than
4th ; flagellum equal in length to 5th peduncle segment, of 23 articles.

Mouthparts projecting below head. Upper lip rounded and finely
setose below. Mandible, palp slender, third segment lightly expanded
distally, armed with several long plumose setae; incisor process
strongly toothed; lacinia mobilis large, associated with a few slender
spines; molar process with ridged triturating surface. First maxilla,

April 20, 1964 Peabody Museum Amphipoda 13

palp apical segment curved, outer plate armed with 10 strong den-
tate spines. Second maxilla inner plate short, outer margin short,
oblique, inner margin setose along its entire length; outer plate
projecting over inner distally. Lower lip outer plates with marked
lateral projections; medial margins with deep rounded incisions.
Maxilliped, outer plate armed with rather small spines; inner plate
inner margin partially oblique, set with plumose setae.

Gnathopod 1, coxa projected anterodistally ; basis with an antero-
distal rounded lobe armed with a few spines; carpus elongated;
propodus long, even in width, palmar margin short and transverse,
armed with a blunt spine near the dactyl and several setae, lower
margin of segment slightly concave and heavily set with groups of
setae; dactyl long, inner margin dentate. Gnathopod 2 basis with
an anterodistal rounded lobe; carpus wide, about 3/4 length of
propodus; propodus only slightly shorter than that of gnathopod 1
but much wider, lower margin slightly concave, palm oblique, con-
cavity defined by a ventral projection, with a quadrate setose lobe
near base of dactyl, palm and lower margin set with groups of long
setae; dactyl dentate on inner margin, extending length of palm.

Peraeopods 1 and 2 short, very similar; basis expanded medially ;
propodus narrow, with a few setae ventrally; dactyl short, 1/3
length of propodus.

Peraeopod 3, coxa with a large anterior lobe and small posterior
lobe; basis expanded, rounded anteriorly and posteriorly; dactyl
stout, curved.

Peraeopods 4 and 5 very similar in form, but 4 shorter than 5,
extending at most to the middle of peraeopod 5 propodus. Peraeo-
pod 4, coxa slightly lobed anteriorly; basis rounded posteriorly,
with a small posterodistal concavity marked by a single spine;
merus, carpus and propodus about equal in length. Peraeopod 5 coxa
rather narrow, slightly concave distally; propodus longer than car-
pus, about equal in length to merus.

Gills all simple, short and broad.

Epimeral plates; first very short, rounded anteriorly, slightly
concave posteriorly; second strongly rounded anteriorly, posterior
margin convex proximally, becoming slightly concave near postero-
distal corner; third strongly rounded anteriorly, posterior margin
convex above posterodistal corner. Pleopod rami about half again
as long as peduncles; coupling spines short, apex with one hook on
one side, 2 on the other.

Uropods all extending to same point. Uropod 1 stout, peduncle
with 7 spines on outer margin, 10 on inner margin; outer ramus

14 Postilla Yale Peabody Museum No. 79

shorter than inner, outer margin evenly set with 8 spines, inner mar-
gin with 2 spines only; inner ramus outer margin with two spines,
inner margin with 3 spines. Uropod 2 stout; peduncle margins each
with 4 to 5 spines; outer ramus slightly shorter than inner, outer
margin with 6 spines, inner margin with 4 spines; inner ramus outer
margin with 8 spines, inner margin with 5 spines. Uropod 3 short;
rami very short, rounded, outer ramus with 2 lateral hooks, inner
ramus with an apical spine, 2 lateral spines and a tuft of apical
setae; distal margin of peduncle with 6 spines.
Telson short, narrowed, quadrate, and with 4 setae distally.

Frema.e (9.7 mm) PARALECTOTYPE Fig. 3 (v-y). Antenna I
peduncle more slender than in male but proportions about the same;
flagellum of 25 segments, extending 3/4 length of body. Antenna
2 less robust than that of male, peduncle segments 4 and 5 slender,
almost equal in length; flagellum of 17 articles.

Gnathopod 1 carpus shorter and smaller than male, almost straight
ventrally ; propodus about equal to carpus in length, palmar margin
rounded, oblique, dactyl long, dentate, half its length beyond palm.
Gnathopod 2 propodus shorter and smaller than in male, palmar
margin convex near dactyl, gradually becoming concave ventrally,
palm with quadrate lower corner.

Oostegites present on gnathopod 2 and peraeopods 1-3, lobate,
broad, fringed on most margins with long curved setae. Uropod 3
with 1-3 spines on inner ramus inner margin.

Other features as in male.

Type locality: Vineyard Sound, Mass.

Material: YPM 5214—Vineyard Sound, Mass. U. S. Fish
Commission, 1871. 1 lectotype ¢, 8 paralectotype ¢ 6. YPM
5215—Vineyard Sound, Mass. U. S. Fish Commission, 1871. 8
paralectotype 2 2 (1 figured). YPM 4631—Great Egg Har-
bor, N.J. (2) A. E. Verrill and S. I. Smith, 1872, 1 juv. oe
YPM 5238—no data. Several dessicated specimens. YPM 5239-
Provincetown, Cape Cod, low water, USFC. Aug. 22, 1879.
USNM 35662. 2 adult 6 6,1 juv. 3,1 ovig. 2 YPM 5240—-
Long Island Sound, USFC, 1874. USNM 35619. 1 adult ¢.
YPM 5241—Long Island Sound, USFC, 1874. USNM 35618.

Discussion. The material here described and figured was
almost certainly that used by Smith for his original descrip-

April 20, 1964 Peabody Museum Amphipoda 15

tion of the species (in: Verrill and Smith, 1873). Smith did
not mention the numbers of animals in his original collection.
Among the specimens in this series is one which Kunkel (1918)
apparently used for his figure 43. However, since this specimen
was incomplete and no trace could be found of missing append-
ages, the lectotype chosen was a male in better condition.

One difference was noted between Smith’s description and
the material at hand. All specimens examined had yellow-brown
eyes, while Smith stated their color was black. Fresh specimens
of the species from Sengekontacket Pond, Martha’s Vineyard,
Mass., in my personal collection, have dark brown eyes in alco-
hol. Thus it seems that eye pigment gradually leaches out after
years in preservative.

Barnard (in: Barnard and Reish, 1959) outlines the range
of A. longimana as including the United States East Coast,
Bermuda (see Kunkel, 1910), and parts of southern and lower
California. Barnard’s figures show that the California popula-
tions are virtually identical with those on the East Coast. Table
1 outlines the main morphological differences between females
of the present species and females of Ampithoe rubricata
(Montagu).

Ampithoe pollex Kunkel

Amphithoe pollex Kunkel, 1910, p. 93, fig. 36.
Grubia indentata Stout, 1913, p. 656-657. Shoemaker, 1941,
p- 188.

Ampithoe pollex, J. L. Barnard, 1954, p. 29-31, figs. 27-28.

Material: YPM 5291—probably Bermuda, A. E. Verrill, 2
6 6, 5.5 mm and 3.8 mm.

Kunkel’s type material consists of two damaged males which
agree well with his figures (1910) and with the description and
figures of Barnard (1954). Since the species is so well charac-
terized no attempt has been made to figure the poor specimens
at hand.

16

TABLE lI,

Postilla Yale Peabody Museum

longimana and Ampithoe rubricata.

No. 79

Main morphological differences between females of Ampithoe

A, longimana 9

A.rubricata 9

Eye
Antenna 1

Antenna 2

Mandible

Gnathopod 1

Gnathopod 2

Peraeopod 5

Uropod 1

Uropod 2

Uropod 3

Telson

Round, large.
Peduncle slender,

Peduncle segment 4 and 5
slender,

Palp segment 3 almost
linear.

Propodus narrow, palm
short; dactyl longer
than palm.

Carpus lower margin long,
shallowly rounded.

Slightly oval, small.
Peduncle stout.

Peduncle segment 4 and 5
more stout.

Palp segment 3 flattened,
inflated distally.

Palm longer, dactyl equals
palm length.

Carpus lower margin short,
more sharply rounded.

Very similar in both

Basis hind margin widest
proximally, convex.
Spine at posterior angle.

Outer ramus inner margin
with 2 spines.

Outer ramus inner margin
and inner ramus outer
margin with spines.

Rami very short, rounded.

Inner ramus with a few
spines and few apical
setae,

Outer ramus with no
lateral setae,

Few apical setae.

Basis even width throughout,
hind margin linear. No
spine at posterior angle.

Outer ramus inner margin
spineless.

Outer ramus inner margin
and inner ramus outer
margin lacking spines.

Rami % length of peduncle
or more, less rounded,

Inner ramus with several
spines, large clump of

apical setae,

Outer ramus with clumps of
lateral setae.

Several apical setae (3/side)

April 20, 1964 Peabody Museum Amphipoda 17

Ampithoe valida Smith
(Fig. 4)

Amphithoe valida Smith (in: Verrill and Smith, 1873, p. 563).

Amphithoe rubricata (in part), Della Valle, 1893, p. 459.
Holmes, 1905, p. 510. Kunkel, 1918, p. 149-151.

Ampithoe valida, Stebbing, 1906, p. 635. J. L. Barnard, 1954,
p. 34-35, pl. 31. Alderman, 1936, p. 68.

Amphithoe shimizuensis Stephensen, 1944, p. 77-80, figs. 28, 29.

Diagnosis: 1) Gnathopod 1 carpus and propodus broadly
expanded, carpus with broad lobe on lower margin. 2) Gnatho-
pod 2 propodus massive, palm almost transverse, with a blunt,
square tooth medially. 3) Uropod 3 rami half length of pedun-
cle, outer ramus with 2 stout out-curved spines. 4) Antenna
2 only slightly shorter than antenna 1.

Description: Mave (12.7 mm) LECTOTYPE. Fig. 4 (a-q). Head
slightly shorter than first two body segments combined ; interanten-
nal lobe large, slightly truncated, corners rounded.

Antenna 1 about half as long as body, peduncle segments 1 and 2
equal in length; flagellum of 35 articles. Antenna 2 slightly shorter
than antenna 1, peduncle segment 5 slimmer than 4, about equal to
it in length; flagellum with 20 articles.

Epistome slightly rounded and finely setose below.

Mandible, incisor process with 6 sharp teeth; lacinia mobilis large
and toothed; 6 pectinate spines between incisor and molar processes ;
molar process well-developed, ridged and sclerotized; palp 3rd seg-
ment broad, with several setae on the curved distal margin. Maxilla
1, last segment of palp rounded distally, bearing 8 spines and a
seta; outer plate with 10 stout spines; inner plate small, unarmed.
Maxilla 2, inner plate setose along entire inner margin; outer plate
ptojecting over inner plate distally. Maxilliped, palp segments short ;
outer plate with 11 small short spines on inner margin, grading into
longer spines distally; inner plate broadly rounded distally and set
with marginal setae.

Gnathopod 1, coxa projected anterodistally ; basis with an antero-
distal rounded lobe laterally ; carpus broad, with a lateral fringe of
long setae on the upper margin, lower margin broadly lobed, extend-
ing partly over the base of the propodus; propodus almost as broad
as carpus, palmar margin rounding evenly into lower margin, defined

18 Postilla Yale Peabody Museum No. 79

by a spine at the lower corner; dactyl stout and short. Gnathopod 2,
basis stout, with a large anterodistal lateral lobe; merus projecting
slightly below carpus; carpus short, with a few setae on a lower
lobe; propodus massive, slightly expanded distally, palm almost
transverse, with a small square median tooth and a rounded prom-
inence at the lower corner; dactyl stout, tip fitting behind promi-
nence at lower corner of palm.

Peraeopods 1 and 2 very similar; basis expanded, setose on the
hind margin; carpus and propodus slender. Peraeopod 3. basis
rounded, with a few spines on the anterior margin; propodus with
5 spines on anterior margin; dactyl pointed outward or forward.
Peraeopods 4 and 5 quite similar, peraeopod 4 short, extending to
about middle of peraeopod 5 propodus, its basis with a sharp notch
posterodistally.

Epimeral plate 1 rounded anteriorly, with a slight posterodistal
tooth and a squarish projection posteriorly ; epimeral plate 2 rounded
posteriorly, with a slight tooth at the posterodistal corner; epimeral
plate 3 larger than 2, quite similar in shape. Pleopod peduncles
stout, coupling spines 9, mushroom-shaped; rami about half again
as long as peduncles, of 16-18 segments.

Urosome short, flattened. Uropods all extending to same point.
Uropod 1 long; outer ramus slightly over half as long as peduncle,
shorter than inner ramus, outer margin with 6 spines, inner margin
spineless; inner ramus outer margin spineless, inner margin with 4
spines. Uropod 2, outer ramus outer margin with 5 spines, inner
margin spineless, inner ramus outer margin spineless, inner margin
with 3 spines. Uropod 3 short, peduncle about twice length of rami,
dorsal surface with 3 spines at base of outer ramus and 2 at base
of inner ramus; outer ramus with 2 out-curved spines distally; inner
ramus with 4 short spines and a tuft of setae distally and a small
spine on the outer margin.

Telson short, distal corners slightly notched, apex slightly acute
(rounded in some specimens), dorsal surface with setae near mar-
gins and at distal corners.

Femate. The collections contain a number of juvenile females,
all previously identified as A. valida. All, however, are indistin-
guishable from females of A. longimana. It appears that Smith’s
original collection contained both species and that he or a subse-
quent viewer of the material mistook the female of 4. longimana for
that of 4. valida. Smith (in: Verrill and Smith, 1873, p. 5638) says
of the female of A. valida: “The female differs in having the hands
of the first pair of legs slightly more elongated, and those of the

April 20, 1964 Peabody Museum Amphipoda 19

second pair more elongated than in the male, and the palmar margin
slightly oblique.”

This description fits the gnathopod condition of a paralectotype
juvenile male exactly, and presumably the female of the species is
very similar. It is possible that Smith’s description was based on
this paralectotype male.

Alderman (1936) undoubtedly saw a female of A. valida, since
his description is based on a mating pair. Of the female gnathopod
characters he says only (p. 68): “First gnathopod somewhat more
slender in female than in male.... Fifth joint of second gnathopod
in female produced as in male, shorter than sixth joint. Palm
smooth, convex. Second joint as in male.”

Further collections are necessary to clarify the morphology of
the female.

Type locality: Beesley’s Point, New Jersey.

Material: YPM 1230—Beesley’s Point, New Jersey. A. E.
Verrill and S. I. Smith, April 1871. 1 lectotype ¢ (12.7 mm),
5 paralectotype adult ¢ 6,1 paralectotype juv. ¢.

Discussion. Uncertainties about the identification and
description of females have been discussed previously.

Smith’s description (in: Verrill and Smith, 1878) fits the
specimens at hand very closely, except that the specimens have
pale brown eyes, rather than black. This is certainly attrib-
utable to leaching of color in alcohol, as discussed for A.
longimana.

On the United States East Coast A. valida is apparently
known only from the type locality, although Smith (in: Ver-
rill and Smith, 1873) noted its presence in Long Island Sound.
Alderman (1936) and Barnard (1954) both record the species
from the United States West Coast (California and Oregon,
respectively). Alderman’s description differs from Smith’s speci-
mens in a few points, all probably attributable to size difference.

20 Postilla Yale Peabody Museum No. 79

These differences are:

Alderman’s description Smith’s specimens

Size 9-11 mm 10-13 mm
Antenna 1

flagellum 50 articles 35 articles
Antenna 2

flagellum 25 articles 20 articles
Antenna 2 peduncle Half again as Equal to 5 in

segment 4 long as 5 length
Maxilla 1 inner plate 9 spines 10 spines

Barnard’s figures indicate that his specimen may have been
slightly subadult, since the limbs are less spinose than the New
Jersey material figured here and the male gnathopod 1 is
slightly less developed. However, agreement is very close.

Holmes (1905) first placed A. valida in the synonymy of
A. rubricata, and his action was followed by Kunkel (1918).
Holmes stated (p. 510): “Specimens sent to the U. 8S. National
Museum by Professor Smith under the name Amphithoe valida,
and which I have examined, agree well with Professor Smith’s
description of that species, which is certainly identical with A.
rubricata.”’ Holmes’ mistake could have been due to the examina-
tion of immature specimens of A. valida. Mature males are
quite distinct in the character of both pairs of gnathopods.

Genus Cymadusa Savigny 1816.

Ruffo (1947, p. 168) defines this genus as follows: “Charac-
ters of the genus Amphithoe Leach. .., but first pair of anten-
nae provided with an accessory flagellum of 1-6 articles. The
genus is particularly distinguished by the following combina-
tion of characters; mandible furnished with a well-developed
palp, first pair of antennae provided with an accessory flagel-
lum.”

April 20, 1964 Peabody Museum Amphipoda 21

Cymadusa compta (Smith) new combination

(Figs. 5, 6)

Amphithoe compta Smith, 1873 (in: Verrill and Smith, 1878,
p-. 564.)

Grubia compta, Holmes, 1905, p. 510-511, figs. Kunkel, 1918,
p. 151-153, fig. 45. PShoemaker, 1921, p. 102.

Amphithoides comptus, Stebbing, 1906, p. 645.

PGrubia sp., Pearse, 1913, p. 376.

Not Grubia compta, Pearse, 1913, p. 376, fig. 6.

Grubia hirsuta Chevreux (in part), Schellenberg, 1925, p.
186-187.

Cymadusa filosa Savigny (in part), Pirlot, 1939, p. 64-67.

Diagnosis: 1) Coxae of gnathopods 1 and 2 sparingly setose.
2) Gnathopod 1 propodus palmar margin slightly concave. 3)
Gnathopod 2 carpus of é long, without distinctly projecting
ventral lobe. In 2, gnathopod 2 carpus with a large, round-
margined ventral lobe. 4) Gnathopod 2 propodus equal in width
to carpus, dactyl longer than palmar margin. 5) Antenna 1
flagellum with about 38 articles; antenna 2 flagellum with 20-
28 articles.

Description: Mare (8.6 mm). LECTOTYPE. Fig. 5 (a-p). Head
slightly shorter than the first two body segments combined; inter-
antennal lobe square and projecting; hind margin oblique; inferior
margin slightly concave. Eye slightly ovate, straw-colored in speci-
mens in alcohol.

Antenna 1, first segment of peduncle as long as head; second seg-
ment equal in length to first, third segment about 1/3 length of
second; accessory flagellum of one main article and a small setose
apical article; flagellum slender, long, extending almost to end of
body, of about 33 articles.

Antenna 2 slightly shorter than antenna 1, peduncle stout, seg-
ments 4 and 5 about equal, each armed ventrally with 8-9 groups
of long setae; flagellum of 20-28 articles.

Epistome rounded ventrally and finely setose.

Mandible, incisor process large, curved, with 6 sharp teeth; lacinia
mobilis stout, armed with several small teeth; 8 setae between incisor
and molar processes; molar process ridged and heavily sclerotized;

22 Postilla Yale Peabody Museum No. 79

palp slightly curved, 8rd segment expanded distally and set with
several long pectinate setae.

Hypopharynx with well-developed, finely setose, inner lobes, outer
lobes bilobed medially.

Maxilla 1, inner plate small, acute apically, with 7 setae on the
medial margin; outer plate with 10 stout toothed spines; palp seg-
ment 3 expanded distally, bearing 7 short stout spines and one or
two setae. Maxilla 2, outer plate large, distal margin oblique, set
with long setae; inner plate small, narrowed distally, with an oblique
row of setae joining setae found along the entire medial margin.

Maxilliped, inner plate narrow, with 2 or 3 stout spines and
several long setae distally and a row of long plumose setae along the
medial margin; outer plate set with several long, curved setae dis-
tally and with a series of 13 stout spines along the medial margin;
palp segments 3 and 4 with groups of long setae medially.

Gnathopod 1, coxal plate small, projecting anteriorly ; basis stout,
with several groups of long plumose setae near the hind margin;
merus with a long anteroventral projection extending nearly half-
way along the carpus, set with groups of long plumose setae; carpus
long, broad, anterior margin almost straight and square, ventral mar-
gin set with thick groups of long plumose setae; propodus
shorter than carpus, with thick groups of plumose setae on upper
margin, palmar margin oblique, set off by a stout spine at the lower
angle, and a slight hump near the attachment of the dactyl, lower
margin rounded, receding into a narrow connection with the propo-
dus, with groups of long setae; dactyl stout, curved, inner margin
serrated.

Gnathopod 2, coxal plate deeper than wide, corners rounded; basis
heavily set with long plumose setae; merus small, with a rounded
anteroventral projection armed with groups of long setae; carpus
long and broad, upper and lower margins heavily set with long
plumose setae; propodus almost as long as carpus and about as wide,
with groups of long plumose setae on the upper margin, palmar mar-
gin oblique and slightly concave, with a small hump near the base
of the dactyl and a small acute projection at the ventral corner;
dactyl stout, inner margin serrated.

Peraeopod 1, coxal plate deeper than wide, lower anterior corner
rounded, posterior corner almost square; basis stout, glandular
(glandular material also present in ischium and merus); carpus
and propodus about equal in length, both with several groups of
setae on ventral margin: dactyl short, stout, curved. Peraeopod 2
very similar.

April 20, 1964 Peabody Museum Amphipoda 23

Peraeopod 3, coxa with a posterior rounded lobe, lower corners
rounded; basis ovate, armed with 6 spines on the anterior margin;
merus and carpus about equal, short, stout, with one and two spines
on the posterior margin respectively; propodus about half again as
long as carpus, with 4 stout spines on the posterior margin and 5
groups of setae on the anterior margin; dactyl short, stout, turned
posteriorly.

Peraeopod 4 shorter than peraeopod 5; coxa small, with a rounded
anterior lobe; basis narrow, posterior margin concave distally and
with 1-2 short spines; merus with 2 spines on posterior margin;
carpus slightly shorter than merus, with one spine on the posterior
margin; propodus slightly expanded distally, with 5 spines on ante-
rior margin; dactyl stout, turned anteriorly. Peraeopod 5 similar to
peraeopod 4, but more stout and longer; coxa not lobed, slightly
narrowed distally; basis with a posterior proximal lobe, 3 spines
on the posterior margin; propodus more heavily setose than that of
peraeopod 4.

Gills short, slightly longer than wide, simple.

Pleon slightly compressed dorsoventrally. Pleopod rami about
half again as long as peduncles, with about 13-14 segments. Pleopod
coupling hooks anchor-shaped. Epimeral plate 1 small, posterior
margin convex, lower posterior corner acute, with a small spine.
Epimeral plate 2 with a rounded projection anteriorly, posterior
angle acute, posterior margin concave distally, convex proximally.
Epimeral plate 3 larger than 2, acute posteriorly, posterior margin
gently convex.

Urosome slightly flattened dorsoventrally. Uropods all extending
to same point. Uropod 1 stout, peduncle with 5 spines on inner mar-
gin, 6 spines on outer, produced into a long ventral spine-like proc-
ess which extends between the rami; both rami with distal spine
groups, inner ramus longer than outer; outer ramus outer margin
with 4 spines, inner margin with two spines; inner ramus outer mar-
gin with one spine, inner margin with 4 spines. Uropod 2 shorter
than 1, peduncle inner margin with 3 spines, outer margin with 4
spines; both rami with distal spine groups; inner ramus longer than
outer, inner margin with 4 spines, outer margin with 2; outer ramus,
inner margin with 8 spines, outer margin with 4. Uropod 3 short,
peduncle with 4 spines on inner margin, 2 spines and 2 groups of
ventral setae on outer margin, and 3 short spines at the base of the
outer ramus; inner ramus straight, armed with 3 spines on the inner
margin, an apical spine and tuft of setae, and one spine on the outer
margin; outer ramus curved, with 2 distal spines which curve outward.

24 Postilla Yale Peabody Museum No. 79

Telson short, broad, corners acute, with setae set in notches; 3
groups of lateral setae on each side; dorsal surface with 4 groups
of setae.

FemaLe (14.4 mm) PARALECTOTYPE. Fig. 6 (v-z). Quite
similar to the male, but antennae a little shorter and more slender
and gnathopods considerably smaller.

Antenna 1 flagellum with 37 articles; antenna 2 flagellum with
27 articles.

Gnathopod 1 with only a few long simple setae proximally on the
hind margin of the basis; merus with a short, acute anteroventral
projection set with a few simple setae extending about 1/3 the
length of the carpus; carpus broad, with a broad ventral oblique
lobe set with simple setae ventrally ; propodus about as long as car-
pus, with only a few long setae on upper margin, palmar margin
oblique, but not as much as in male and shorter, defined by a stout
spine at the lower corner and a slight hump near the attachment of
the dactyl, lower margin slightly rounded, set with groups of setae,
receding into narrow connection with the propodus; dactyl stout,
inner edge serrated.

Gnathopod 2, basis with several groups of long setae, merus small,
with a rounded anteroventral projection armed with groups of long
setae; carpus short and broad, with a ventral lobe thickly set with
groups of long setae, upper margin almost bare of setae; propodus
as long as carpus and about as wide, with a few groups of setae
on the upper margin, palmar margin oblique and almost straight,
defined by a stout spine at the lower corner; dactyl stout and ser-
rated on the inner margin.

Oostegites long, ovoid, margins with many close-set long setae.

Type locality: Vineyard Sound, Mass.

Material: YPM 5209—Vineyard Sound, Mass., USFC, 1871.
1 Lectotype ¢ (8.6 mm); 3 paralectotype adult ¢ ¢ (1 dis-
sected by Kunkel, 1918); 3 paralectotype juv. ¢ ¢: YPM
5210—Vineyard Sound, Mass., USFC, 1871. 8 ovig. 2 2, 4 juv.
2 2, paralectotypes. YPM 5211—Long Island Sound, USFC,
1874. 1 juv. $, 1 ovig. 9. YPM 5212—Long Island Sound,
USFC, 1 ovig. ?, 5 juvs. YPM 5242—Long Island Sound,
USFCYUSi4l Tt jaye o.-

Discussion. This species, described by Smith (in: Verrill and
Smith, 1873) as Amphithoe compta, has a complicated nomen-

April 20, 1964 Peabody Museum Amphipoda 25

clatural history. Stebbing (1906) transferred the species to the
genus Amphitoides Kossman, believing it to have only one
hook on the outer ramus of uroped 3. Smith’s description makes
no mention of uropod 38. Holmes (1905) placed the species in
the genus Grubia. Kunkel (1918) pointed out Stebbing’s mis-
take and retained the genus Grubia. Schellenberg (1925) stated
“Tch halte nach allem die Arten von Chevreux, Kunkel und
Pearse fiir identisch” and placed Grubia compta in G. hirsuta
Chevreux. With Pirlot’s (1938, 1939) resurrection of the genus
Cymadusa Savigny to replace Grubia, G. compta and its several
synonyms were listed with Cymadusa filosa Savigny, as was
Schellenberg’s “Grubia hirsuta.” Ruffo (1947) suggested that
Smith’s species was erroneously included in Pirlot’s list. This
has proved to be the case, since C. compta differs greatly from
C. filosa and other Cymadusa species in its gnathopod charac-
ters and seems to have a disparate range.

The known range includes New England (present material),
south to North Carolina (Kunkel, 1918, on Smith’s authority).
Pearse’s (1913) record of the species from Florida is C. filosa,
as Barnard (1955) pointed out, judging by Pearse’s figures,
in which gnathopods 1 and 2 have been reversed. Pearse also
records “Grubia sp.?” from Key West, Florida; this may be
C. compta. Shoemaker’s (1921) record of “Grubia compta”
from Barbados may be C. filosa. The latter species seems never
to have been taken north of Florida and is undoubtedly tropical.

Kunkel describes the preference of C. compta for eelgrass.
The species was very common in eelgrass with Ampithoe longi-
mana Smith at Sengekontacket Pond, Martha’s Vineyard, Mass.,
in August, 1962.

Cymadusa filosa Savigny
(For synonymy see Barnard, 1955, and Ruffo, 1947.)

Material: YPM 5208—no data. Bermuda (?).2 ¢ 4,2 2 2,
2 juv. Probably Kunkel’s types of Grubia coei. YPM 5213—
Bermuda. “‘Grubia crassicornis.” 2 2 2,2 juv.

Kunkel (1910, p. 97 and fig. 38) dissected and figured a sub-
adult male of YPM 5208 as Grubia coei. These figures and the

26 Postilla Yale Peabody Museum No. 79

specimens are in close agreement with Shoemaker’s (1985b) and
Ruffo’s (1947) figures of Cymadusa filosa Savigny and support
Pirlot’s (19389) and Ruffo’s views of the status of Grubia coei
as synonymous with C. filosa.

Although YPM 52138 was labelled “Grubia crassicornis,” the
specimens do not agree with Kunkel’s figures (1910, fig. 87) of
G. crassicornis Costa and so cannot be the specimens on which
the figure was based. They are clearly specimens of Cymadusa
filosa,

The confusion of C. compta (Smith) with C. filosa Savigny
has been discussed earlier in this paper.

C. filosa has been recorded from widely separated localities,
including Bermuda (Kunkel, 1910), Florida and Puerto Rico
(Shoemaker, 1935b), Mediterranean Sea, Red Sea, Canary
Islands (Ruffo, 1947), Hawaii, Indian Ocean, Australia and
West Africa (summary in Barnard, 1955).

Family COROPHIIDAE Stebbing
Erichthonius difformis H. Milne-Edwards

Erichthonius difformis H. Milne-Edwards, Gurjanova, 1951, p.
950-951, fig. 661.

Material: YPM 5633—USFC. No other data. 2 adult ¢ 6,2
juv. YPM 5665—Bay of Fundy, USFC, 1872. 20 adults, 4 juv.

Eleven collections of species of the genus Erichthonius occur
in the Yale collection. Based on male gnathopod 2 characters
primarily, nine are FE. rubricornis (Stimpson), and all were
collected in the region from Georges Bank to Halifax Harbour,
Nova Scotia. Two collections, listed above, contain E. difformis.
Most specimens in these two collections agree quite well with
Sars’ (1895) figures. However, some variability in the form of
the male second gnathopod occasionally makes specific identi-
fication difficult. Taxonomic problems have been noted in other
combinations of species in this genus. Dahl (1946) maintained
that E. difformis and E. hunteri were specifically distinct, while
Enequist (1950) suspected that the two forms could be the
same species, the variability in form being caused by the effect
of environment on growth rates.

April 20, 1964 Peabody Museum Amphipoda 27

It is noteworthy that the form of the male second gnathopod
in E. rubricornis is closer to that of E. hunteri (as illustrated
by Sars) than Holmes’ (1905) figures would indicate. For this
reason, as well as to clarify the problems outlined above, a
critical morphological study of the common North Atlantic
species would be of great value.

The distribution of FE. difformis on the East Coast of North
America is not clear. Brunel (1961a) records only E. rubri-
cornis and E, tolli from the Gulf of St. Lawrence region. Some
of S. I. Smith’s references to E. difformis in New England refer
to E. rubricornis, since some of his specimens in the Yale collec-
tion, labelled EF. difformis, are actually E. rubricornis. Holmes
(1905) seems to have recognized this fact. The present speci-
mens indicate only that E. difformis occurs south at least to

the Bay of Fundy.

LITERATURE CITED

Alderman, A. L. 1936. Some new and little known amphipods of
California. Univ. Calif. Publ. Zool. 41 (7): 53-74.

Barnard, J. L. 1954. Marine Amphipoda of Oregon. Ore. St. Monogr. Zool.
8: 1-103.

1955. Gammaridean Amphipoda in the collections of Bishop
Museum. Bull. Bishop Mus., Honolulu. 215: 1-46.

1958, Index to the families, genera and species of the
Gammaridean Amphipoda (Crustacea). Occ. Pap. Allan Hancock
Fdn. 19: 1-145.

Barnard, J. lL. and D. J. Reish. 1959. Ecology of Amphipoda and
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Bousfield, E. L. 1958. Fresh-water amphipod crustaceans of glaciated
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Brunel, P. 196la. Liste taxonomique des invertébrés marins des parages de
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1961b. Inventaire taxonomique des invertébrés marins du golfe
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Chevreux, E. and L. Fage. 1925. Faune de France 9. Amphipodes. Paris,
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Dahl, E. 1946. Notes on some Amphipoda from the Gullmar Fiord. Ark.
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Della Valle, A. 1893. Fauna und Flora des Golfes von Neapel und der
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Enequist, P. 1950. Studies on the soft-bottom amphipods of the Skagerrak.
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Frizzel, D. L. 1933. Terminology of types. Amer. Mid]. Nat. 14 (6) : 637-668.

Gurjanova, E. F. 1951. Amphipoda-Gammaridea of the seas of the
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its biology. Crustaceana 4(2): 158-162.

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Pearse, A. S. 1913. Notes on certain amphipods from the Gulf of Mexico,
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Pirlot, J. M. 1938. Les amphipodes de Vexpédition du Siboga. Siboga
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—1939. Amphipoda. In: Résultats scientifiques des croisi¢res
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Ruffo, S. 1947. Studi sui Crostacei anfipodi, XIII, Sulle specie mediterranee
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1929. Revision der Amphipoden Familie Pontogeniidae. Zool.
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1913-18, 7, pt. E: 1-30.

1921. 5. The Amphipods. In: Reports on the Crinoids,
Ophiuroids, Brachyura, Tanaidacea and Isopoda, Amphipods and
Kchinoidea of the Barbabos-Antigua Expedition of 1918. Iowa Studies
in Nat. Hist., 9: 99-102.

——————. 1932. The amphipod Nototropis minikoi on the East Coast of
the United States, Proc, biol. Soe. Wash. 45: 199-200,

1935a. A new species of amphipod of the genus Grandidierella
and a new record for Melita nitida from Sinaloa, Mexico. J: Wash.
Acad. Sci. 25: 65-71.

1935b, The amphipods of Porto Rico and the Virgin Islands.
In: Scientific Survey of Porto Rico and the Virgin Islands, N. Y.
Acad, Sci., 15 (2): 229-262.

Smith, S. I. 1874a. Crustacea of the fresh waters of the United States. A.

Synopsis of the higher fresh-water Crustacea of the northern United
States. Rep. U. S. Comm. Fish., 1872-73 (1874): 637-660.

1874b, Report on the amphipod crustaceans, In: Annual Rept.
of U. S. geol. geogr. Sury. of the Territories embracing Colorado for

1873 (F. F. Hayden). p. 608-611.

April 20, 1964 Peabody Museum Amphipoda 29

Smith, S. I, and O, Harger. 1874. Report on the dredgings in the region
of St. Georges Banks in 1872. Trans. Conn. Acad. Arts Sci. 3: 1-57.

Stebbing, T. R. R. 1906. Amphipoda I, Gammaridea. Das Tierreich, 21:
806 p.

Stephensen, K. 1935. The Amphipoda of N. Norway and Spitsbergen with
adjacent waters. Troms¢@ Mus. Skr. 3 (1): 1-140.

——— 1944, Some Japanese amphipods, Vidensk. Medd. dansk
naturh. Foren, Kbn. 108: 25-88, 33 figs.

Stoll, N. R., et al., eds. 1961. International Code of Zoological Nomencla-
ture adopted by the XV International Congress of Zoology, Int. Trust
for Zool, Nomenclature, London, 176 p.

Stout, V. R, 1913. Studies in Laguna Amphipoda II. Zool. Jb. (Syst.) 34:
633-659.

Taylor, C. C., H. B. Bigelow and H, W. Graham. 1957. Climatic trends
and the distribution of marine animals in the Northeast. Fish. Bull.
U. S. 115: 293-345.

Verrill, A. E. and S. I. Smith. 1873. Report upon the invertebrate animals
of Vineyard Sound and adjacent waters. Rep. U. S. Comm. Fish.,
1871-2: 295-778.

30 Postilla Yale Peabody Museum No. 79

Figure 1.
Melita nitida Smith. Great Egg Harbor, New Jersey.

LECTOTYPE ¢ (8.7 mm). a, head and antennae. b, upper lip. c, man-
dible. d, lower lip. e, maxilla 1. f, maxilla 2. g, maxilliped (outer plate
separated). h, gnathopod 1. i, gnathopod 1 propodus and dactyl, inner.
j, gnathopod 2. k, gnathopod 2 propodus and dactyl, inner. 1, peraeopod
1. m, peraeopod 2. n, uropod 1. 0, uropod 2. p, epimeral plates 1-3 and
3rd _ pleopod.

PARALECTOTYPE ¢ (8.9 mm). q, uropod 3.

PARALECTOTYPE ¢ (7.1 mm), r, peraeopod 3. s, peraeopod 4. t,
peraeopod 5.

PARALECTOTYPE @ (9.3 mm). u, antenna 1. v, antenna 2. w, telson.
x, gnathopod 1, y, gnathopod 1 propodus and dactyl, inner. z, gnathopod
2. aa, gnathopod 2 propodus and dactyl, inner.

April 20, 1964 Peabody Museum Amphipoda 31

Postilla Yale Peabody Museum No. 7

Figure 2.

Ampithoe longimana Smith. Vineyard Sound, Massachusetts.

LECTOTYPE ¢ (10.1 mm). a, head. b, antenna 1. c, antenna 2. d, upper
lip. e, mandible (with lateral view of palp terminal segment). f, lower

lip. g, maxilla 1. h, maxilla 2. i, maxilliped (outer plate separated). j,

gnathopod 1, inner. k, gnathopod 2, inner. 1, peraeopod 1. m, peraeopod 2.

n, peraeopod 3, 0, peraeopod 4, p, peraeopod 5. q, uropod 1. r, uropod 2.
s, uropod 3. t, telson.

April 20, 1964 Peabody Museum Amphipoda 33

Od Postilla Yale Peabody Museum No: 79

Figure 3.

Ampithoe longimana Smith, Vineyard Sound, Massachusetts.

LECTOTYPE ¢ (10.1 mm). u, epimeral plates 1-3 with pleopods.

PARALECTOTYPE 9 (9.7 mm). vy, antenna 1. w, antenna 2. x, gnatho-
pod 1, inner, y, gnathopod 2, inner.

April 20, 1964

Peabody Museum Amphipoda

35

36 Postilla Yale Peabody Museum No. 79

Figure 4.
Ampithoe valida Smith. Beesley’s Point, New Jersey.

LECTOTYPE ¢ (12.7 mm). a, antenna 1. b, antenna 2. c, upper lip.
d, mandible (lacinia mobilis and incisor in detail). e, lower lip. f, maxilla
1. g, maxilla 2. h, maxilliped (outer plate separated). i, gnathopod 1. j,
gnathopod 2. k, epimeral plate 1. 1, epimeral plate 2. m, epimeral plate 3
and pleopod 3. n, uropod 1. 0, uropod 2, p, uropod 3. q, telson.

PARALECTOTYPE ¢ (7.9 mm). r, peraeopod 1. s, peraeopod

ets
peraeopod 3. u, peraeopod 4, v, peraeopod 5. w, head.

April 20, 1964 Peabody Museum Amphipoda 37

38 Postilla Yale Peabody Museum Norris

Figure 5.
Cymadusa compta (Smith). Vineyard Sound, Massachusetts.

LECTOTYPE ¢ (8.6 mm). a, head, b, upper lip. c, mandible. d, lower
lip. e, maxilla 1. f, maxilla 2. g, maxilliped (outer plate separate). h, gnath-
opod 1 (inner). i, gnathopod 2 (inner). j, peraeopod 2, k, epimeral plates
1-3 (right to left), with pleopod 3. 1, pleopod 1 coupling hooks. m, uropod
1 (side view of ventral spine below). n, uropod 2. 0, uropod 3. p, telson.

Peabody Museum Amphipoda 39

April 20, 1964

40 Postilla Yale Peabody Museum No. 79

Figure 6.
Cymadusa compta (Smith). Vineyard Sound, Massachusetts.
PARALECTOTYPE ¢ (7.5 mm). q, peraeopod 3, r, peraeopod 5.

PARALECTOTYPE ¢ (12 mm). s, antenna 1, t, antenna 2. u, peraeo-
pod 4.

PARALECTOTYPE 9 (14.4 mm). v, antenna 1, w, antenna 2. x, gnatho-
pod 1 (inner). y, gnathopod 2 (inner). z, peraeopod 1.

April 20, 1964 |= Peabody Museum Amphipoda 41

na

Prasopy Museum or Natrurat History

YALE UNIVERSITY
Number 80 May 27, 1964 New Haven, Conn.

CYPRIA PETENENSIS, A NEW NAME FOR THE
OSTRACOD CYPRIA PELAGICA BREHM 1932

Epwarp Frrcuson Jr.’

Lincotn Universiry or Missourt, JEFFERSON Ciry

G. Evetyn HutTcuHinson,

Yate Universiry, New Haven, C

CrypE E. GouLpEnN, \
Yate Universiry, New Haven, Conn.
~,

Recently during the preparation of a comprehensive account
of the lake plankton (Hutchinson, im press) a rather unfor-
tunate case of homonymy was discovered in the genus Cypria.

In 1932 Brehm brought forward the name Cypria pelagica
for a planktonic species from Laguna de Petén, Guatemala.
Brehm mentioned several characters of the species and com-
mented that they suggest ‘*‘Beziehungen zu der siidamerikanis-
chen Cypria pellucida Sars.” Since his remarks merely indi-
cate apparent affinities with pellucida but unhappily contain no
“statement that purports to give characters differentiating the
taxon,” the name pelagica Brehm (1932) appears to be, under
Article 13a of the International Code, invalid and must be
treated as a nomen nudum.

‘Research supported by National Science Foundation Grant GB-1534.

SAT ifsGs se

2 Postilla Yale Peabody Museum No. 80

In the next year Klie (1933) published a description of
Cypria javana subspec. pelagica from lakes in Java and Bali,
which is apparently identical with a pelagic species from the
Lake of Colombo erroneously identified by Apstein (1907) as
Cypris purpurascens Brady. Whatever the status of Klie’s form
may be, the name pelagica was validly proposed as subspecific,
and in view of the invalidity of Brehm’s use of pelagica in the
previous year, cannot be used for any other subsequently des-
cribed Cypria.

Brehm (1939) returned to the problem and gave an illus-
trated description of Cypria pelagica, presumably based on the
material to which he had referred in 1932. Recently we have had
the opportunity to examine specimens of what is doubtless the
same species, collected in the Laguna de Petén by Drs. Georgi-
ana B. and Edward S. Deevey. In view of the history of the
species a new specific name is clearly required.

Famity CYPRIDAE Baird 1845
Genus CYPRIA Zenker 1845

Cypria petenensis n. sp.
Figs. 1-6

Specific characters: Frmate—Eye prominent. Valves translu-
cent, surface with scattered hairs and without other ornamentation.
anterior and posterior margins with sparsely placed hairs; shell
seen from the side boldly arched, greatest height near middle, ante-
rior and posterior margins broadly rounded, ventral margin of right
valve convex, that of left slightly concave; length of valves 0.52
mm—0.53 mm, height 0.31 mm—0.32 mm. Natatory setae of second
antennae reaching beyond tips of end-claws by more than twice the
length of the claws; distal end of antepenultimate podomere bearing

a short seta with an elongate plumose terminal portion; olfactory
club apparently with four segments. Mandibular palp is a very
striking structure, respiratory plate with five prominent setae; ante-
penultimate and penultimate podomeres with long, plumose setae
on ventral margins; ultimate podomere with two smooth spines and
a longer two-segmented spine bearing two lateral rows of short
setae at tip. Proximal podomere of maxillary palp very wide and
bearing four long setae on laterodistal margin; width of distal
podomere equal to length with three pectinated spines; spines of

50m

50yu

AMY
WSK
SSE

6

Cypria petenensis n. sp—Drawings of figures 1, 3, 4, 5, and 6 were made
from the female holotype stained with lignin pink and mounted in polyviny!
lactophenol. The drawing of figure 2 was made from the androtype stained
in a 1% alcoholic solution of eosin Y and mounted in Canada balsam.
1. Second antenna of female holotype. 2. Lateral view of right valve of
androtype. 3. Maxilla and maxillary palp of female holotype. 4. Furcal
ramus of female holotype. 5. Mandibular palp of female holotype. 6. Third
thoracic appendage of female holotype.

4 Postilla Yale Peabody Museum No. 80

all three masticatory processes two-segmented and smooth. Ultimate
podomere of third thoracic appendage with two short setae of
unequal lengths and a long, reflexed seta equal to the combined
lengths of the antepenultimate, penultimate and ultimate podo-
meres; terminal margin of penultimate podomere with a patch of
setae. Furcal ramus strongly developed, stout, length approximately
10 least width; dorsal seta short, occasionally inconspicuous and
removed from the subterminal spine by the length of spine; ter-
minal and subterminal spines pectinated; length of subterminal
spine about eight-tenths that of terminal; terminal seta one-third
length of subterminal spine.

Matr—Valves similar to those of female. Testes extend as cir-
cuitous bands along margins of valve. Prehensile palps of first
thoracic appendages dissimilar; penis triangular. Males appear in
numbers equal to females.

Type locality: The type specimens were collected on March 10,
1961 by Drs. Georgiana and Edward S. Deevey from the Laguna
de Petén, Guatemala.

Type specimens: Microscopic mounts of the holotype and andro-
type are deposited in the Yale Peabody Museum (YPM 5670, 5671).
Paratype male and females have been deposited in the British
Museum (Natural History) and the Zoologisches Museum, Ham-
burg, Germany.

REFERENCES

Apstein, C., 1907. Das Plankton im Colombo-See auf Ceylon. Zool. Jahrb.
(Abt. Syst.) 25:201-244.

Brehm, Vinzenz, 1932. Notizen zur Stsswasserfauna Guatemalas und
Mexikos. Zool. Anz. 99 (3-4) :63-66.

——————, 1939. La Fauna Microscopica del Lago Petén, Guatemala.
Anales de La Escuela Nacional de Ciencias Biologicas 1:173-204.
Hutchinson, G. E. (in press) Treatise on Limnology, vol. Il. John Wiley,

New York and London.
Klie, Walter, 1933. Die Ostracoden der Deutschen Limnologischen Sunda-
Expedition. Archiv fiir Hydrobiol. Suppl. 11:447-502.

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