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SMITHSONIAN 
MISCELLANEOUS COLLECTIONS 


VOR! 





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“EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES, 


AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN’’—JAMES SMITHSON 


(PusricaTIon 4310) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
1958 


THE LORD BALTIMORE PRESS, INC. 
BALTIMORE, MD., U.S. A. 





ADVERTISEMENT 


The Smithsonian Miscellaneous Collections series contains, since the 
suspension in 1916 of the Smithsonian Contributions to Knowledge, 
all the publications issued directly by the Institution except the An- 
nual Report and occasional publications of a special nature. As the 
name of the series implies, its scope is not limited, and the volumes 
thus far issued relate to nearly every branch of science. Papers in 
the fields of biology, geology, anthropology, and astrophysics have 
predominated. 

LEONARD CARMICHAEL, 
Secretary, Smithsonian Institution. 


(iii) 





IO. 


ET. 


CONTENTS 


. Aszot, C. G. Leading operations of the Smithsonian Astro- 


physical Observatory, 1895 to 1955. 8 pp. Sept. 22, 1955. 
(Publ. 4222.) 

Peterson, Menpet L. The last cruise of H.M.S. “Loo.” 55 
pp., 17 pls., 3. figs. Nov. 23, 1955. (Publ. 4224.) 

FaircHILp, G. B. Synonymical notes on neotropical flies of the 
family Tabanidae (Diptera). 38 pp. Jan. 11, 1956. (Publ. 
4225.) 

Coorer, G. ArtHur. New Cretaceous Brachiopoda from Ari- 
zona. 18 pp., 4 pls. Dec. 21, 1955. (Publ. 4227.) 

WetMorE, ALEXANDER. A check-list of the fossil and prehis- 
toric birds of North America and the West Indies. 105 pp. 
Jan. 25, 1956. (Publ. 4228.) 

Gazin, C. Lewis. Paleocene mammalian faunas of the Bison 
Basin in south-central Wyoming. 57 pp., 16 pls., 2 figs. Feb. 
28, 1956. (Publ. 4229.) 

Gazin, C. Lewis. The upper Paleocene Mammalia from the 
Almy formation in western Wyoming. 18 pp., 2 pls. July 31, 
1956. (Publ. 4252.) 

Gazin, C. Lewis. The geology and vertebrate paleontology of 
upper Eocene strata in the northeastern part of the Wind River 
Basin, Wyoming. Part 2. The mammalian fauna of the Bad- 
water area. 35 pp., 3 pls., 1 fig. Oct. 30, 1956. (Publ. 4257.) 

Kiruam, Lawrence. Breeding and other habits of casqued 
hornbills (Bycanistes subcylindricus). 45 pp., 6 pls., 2 figs. 
Nov. 8, 1956. (Publ. 4259.) 

Snoperass, R. E. Crustacean metamorphosis. 78 pp., 28 figs. 
Oct. 17, 1956. (Publ. 4260.) 

Cuapwick, L. E. The ventral intersegmental thoracic muscles of 
cockroaches. 30 pp., 18 figs. Jan. 15, 1957. (Publ. 4261.) 


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SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 1 


Roebling Fund 


LEADING OPERATIONS OF THE 
SMITHSONIAN ASTROPHYSICAL 
OBSERVATORY, 1895 to 1955 


By 
Cc. G. ABBOT 


Research Associate, Smithsonian Institution 





CO! Saar Pe 
iS OC Wows 2 


(PUBLICATION 4222) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
SEPTEMBER 22, 1955 


The Lord Baltimore Press 


BALTIMORE, MD., U. 8. A, 


Roebling Fund 


LEADING OPERATIONS OF THE SMITHSONIAN 
ASTROPHYSICAL OBSERVATORY, 
1895 TO 1955 


By C. G. Aszor 
Research Associate, Smithsonian Institution 


INTRODUCTION 


Having been associated with the Astrophysical Observatory almost 
from its inception in 1890, it seems good to me to print, in small com- 
pass in one place, references to the leading researches and instrumental 
developments carried out there. It may well be that there are now, 
and will be in future, those who, for one reason or another, may wish 
to refer to these events, and will appreciate having easy access to the 
original sources. 

The list is far from exhaustive, either as regards the work of the 
Astrophysical Observatory, or references to it. But I believe it is suffi- 
cient to present a fair picture of what has been accomplished. 


SECTION A 
Part 1.—lmproved and new instruments 


1. The bolometer rebuilt and equipped Annals of the Astrophysical Observa- 
with a balancing device close be- tory,) vol. I, pp. 47-56, 105-109, 
side it and at constant tempera- 1900; vol. 3, p. 42, 1913. 
ture. Result: The drift nearly 
eliminated and the wiggle 
greatly reduced. 

Galvanometer. Theory investi- Astrophys. Journ., vol. 18, No. 1, July 
gated and new galvanometer of 1903. 
tenfold sensitiveness built. Annals, vol. 1, pp. 244-252, 1900. 

3. Vacuum bolometer with self-con- Annals, vol. 4, pp. 45-64, 1922. 
tained Wheatstone bridge built. 

Result: Several-fold increase of 
sensitiveness, and increased 


i 


steadiness. 
4. Silver-disk pyrheliometer invented. Smithsonian Misc. Coll., vol. 56, No. 
About 100 copies have been 19, IQII. 


1 Hereafter referred to simply as “Annals.” 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL 131, NO. 1 


10. 


It. 


12, 


13. 


14. 


15. 


16, 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


built, standardized, and sold at 
cost to observers throughout the 
world. Repaired at cost and 
restandardized free when 
damaged. 

Water-flow and water-stir stand- 
ard pyrheliometers invented and 
used. The world’s scale of solar 
radiation measurements rests on 
them.? 

Pyranometer invented. Used daily 
on short-method solar-constant 
observations. Used by Moore in 
North Carolina and Chile. 
Copies made and sold at cost 
world-wide. 

Honeycomb pyranometer, or meli- 
keron, invented. Used by Ab- 
bot and Aldrich on human body, 
and by Sverdrup in polar re- 
gions. Copies made and sold 
at cost world-wide. 

Balloon recording pyrheliometer 
invented and used at high alti- 
tudes. 

Two-mirror coelostat invented. 

Slide-rule extrapolator invented. 
Constantly used in long-method 
solar observing. 

High-power lamp and other de- 
vices prepared by F. E. Fowle 
for researches on deep infrared 
spectrum. 

Highly sensitive radiometer in- 
vented for measuring energy 
spectra of stars. 


A prism of nearly normal wave- 
length dispersion invented. 


The kampometer invented, a 
highly sensitive instrument for 
measuring radiation. 

The periodometer invented, a me- 
chanical instrument for discov- 
ering periodic changes in data. 

A multiple rotating-sector dia- 


VOL. 131 


Annals, vol. 3, pp. 47-52, 1913; vol. 7, 
p. 105, 1954. 

Smithsonian Misc. Coll., vol. 95, No. 
23, 1937; vol. 111, No. 14, 1949. 


Annals, vol. 3, pp. 52-72, 1913; vol. 7, 
Pp. 99-101, 1954. 

Smithsonian Misc. Coll., vol. 87, No. 
15, 1932; vol. 110, No. 11, 1948. 


Smithsonian Misc. Coll., vol. 66, No. 7, 
1916. 

Annals, vol. 4, pp. 65-84, 1922; vol. 7, 
pp. 15-16, 21, 138, 1954. 


Smithsonian Misc. Coll., vol. 72, No. 
13, 1922. 

Annals, vol. 4, pp. 41, 300, 1922; vol. 
5, PP. 43-45, 1932. 


Smithsonian Misc. Coll., vol. 65, No. 4, 
1915. 

Annals, vol. 4, pp. 347-365, 1922. 

Annals, vol. 2, pp. 22-23, 211, 1908. 

Annals, vol. 4, pp. 84-86, 1922. 


Annals, vol. 4, pp. 23-25, 274-287, 1922. 


Astrophys. Journ., vol. 69, pp. 293-311, 
1920. 

Smithsonian Misc. Coll., vol. 104, No. 
14, 1945. 

Astrophys. Journ., vol. 11, No. 2, pp. 
135-139, March 1900, 

Smithsonian Misc. Coll., vol. 104, No. 
22, 1946; vol. 107, No. 19, 1948. 

Smithsonian Misc. Coll., vol. 89, No. 3, 
1933. 


Smithsonian Misc. Coll., vol. 87, No. 4, 
1932. 


Annals, vol. 5, p. 96, 1932. 


eee 


* A.P.O. modified form of Angstrém pyrheliometer is used in daily observa- 
tions. See Annals, vol. 6, pp. 50-55, 1942. 


NO. 


I 


ASTROPHYSICAL OBSERVATORY, 1895 TO I955—-ABBOT 


phragm combination invented, 
instantly exchangeable, for bo- 
lometer work. 


17. A continuously variable rotating 


sector invented, of accurate 
ratio, for photometry. 


18. A pair of telephoto cameras in- 


vented, electrically connected, 
for simultaneous exposure on 
flying objects. The invention 
comprises a_ belt-focal-plane 
shutter, surrounding film spools. 
Shutter and spools operated by 
a long spring and clockwork. 
The observer and assistant sepa- 
rated by a measured base line 
keep both cameras trained. Ob- 
server makes a series of expo- 
sures by a trigger, and second 
camera is simultaneously 
exposed. 


19. Apparatus invented for prevent- 


ing “personal equation” in ob- 
serving sudden phenomena. The 
observer notes the sector where, 
not the times when, the event 
occurs. 


20. Automatic recording radiation 


instruments invented. 


No published description. 


One camera on public exhibition in 
Langley case in the West Hall of 
the Arts and Industries Building, 


Smithsonian Institution. 


Apparatus 
Langley case (see above). 


Annals, vol. 7, pp. 144-146, 1954. 


on public exhibition 


in 


Part 2.—Various inventions, mainly for military use in World Wars I and II 


I. 


2. Variable-speed governor. 


Variable-speed 


power-transmis- 
sion mechanism, Claim 1, al- 
lowed “The combination of a 
driving element, a driven ele- 
ment, and means for establish- 
ing, and maintaining constantly, 
exactly and positively, a desired 
speed ratio between said ele- 
ments, or for continuously vary- 
ing said ratio.” 

For a 
clockwork to be of speed varied 
at will, without stopping, and 
continuously, through a several- 
fold range. Used for a Navy 
project. 


3. Self-propelled rotating projectile 


for smooth-bore guns, Combina- 
tion with smooth-bore ordnance. 


U. S. Patent No. 893416 of July 14, 


1908. 


U. S. Patent No. 2367254 of Janu- 


ary 16, 1945. 


U. S. Patent No. 1380172, and U. S. 
Patent No. 1380171, both of May 31, 


192I. 


4 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


4. Gyroscopic navigation instrument. 


R 


6. 


For measuring differences in 
longitude and latitude without 
sun or star observations. 

Compass and magnetic-dip indi- 
cator. Both this and No. 4 
used the principle of neutral 
flotation in liquid, and electric 
current therethrough for operat- 
ing. Germans independently dis- 
covered the mathematical prin- 
ciple of No. 4 and built such a 
machine but it failed. An Eng- 
lishman from National Labora- 
tory examined patent of No. 4, 
and said it carried superior fea- 
tures. Work on it stopped with 
the Armistice, November 1918. 

Instrument for navigating air- 
planes by daylight star observa- 
tions. Stars can be seen with a 
small telescope in daylight if 
the telescope field contains the 
star image. The instrument 
could be set to contain the star 
in its field before observing. 
Twelve stars and two planets 
were easily observed by W. H. 
Hoover in New Mexico. E. D. 
McAlister observed Altair from 
airplane at 21,000 feet. 

Instrument for automatic mapping 
of airplane course over ocean, 
to enable return to course of 
mother ship. The patent, No. 
2367254, above cited, was a part 
of this device. 

Solar distilling apparatus. 

Solar heater. 

Solar heat collector. 


General Electric Patent No. 1501886 
to C. G. Abbot July 15, 1924. 


General Electric Patent No. 1533683 
to C. G. Abbot April 14, 1925. 


Built and tested secretly. Never pub- 
lished. 


Built and tested secretly. Never pub- 
lished. 


Patent No. 2141330, December 27, 1938. 
Patent No. 2247830, July 1, 1941. 
Patent No. 2460482, February 1, 1949. 


SECTION B 


Part 1.—Researches 


. Bolometric map of infrared solar 


spectrum. 


Dispersion of rock-salt and fluo- 
rite. (Six-place decimals in re- 


Annals, vol. 1, pp. 5-204, 1900; vol. 5, 
P- 54, 1932. 

Smithsonian Misc. Coll., vol. 82, No. 1, 
1929. 

Annals, vol. 1, pp. 219-237, 253-262, 
1900. 


NO. 


10. 


Il. 


12, 


1g: 


14. 


15. 


16. 


17. 


18. 


10. 


I ASTROPHYSICAL OBSERVATORY, 1895 TO I1955—-ABBOT 5 


fractive index called ridiculous 
by Holland physicists. Identical 
in fifth place with Paschen 
work, however.) 

Structure of water-vapor bands 
Wy and Ws. 

Total solar eclipses, 1900, I9QoI, 
1908, 1918, 1910. 


Theory of sensitive galvanometer. 


“The cheapest form of light.” 

Solar-constant and solar-distribu- 
tion work, begun in 1902. 

Mount Wilson expeditions, begun 
1905. 

Theory of atmospheric transmis- 
sion. 

Methods for measuring the solar 
constant. 

Transmission of the spectrobo- 
lometer. 

Pyrheliometry. 


Details of solar-constant observ- 
ing. 

Sources of error in solar-constant 
work. 


Solar-contant results of stations 
compared. 

Normal solar-energy curves. Pre- 
ferred determination. 

Sun’s temperature. 

Atmospheric transmission, many 
stations, sea level up to 14,000 


feet altitude. 


Theory of vacuum bolometer, cor- 
rected later. 


Annals, vol. 1, pp. 263-264, 1900. 


Astrophysical Observatory special 
eclipse volume, 1900. 

Annals, vol. 2, p. 2, 1908; vol. 3, pp. 
3-6, 1913; vol. 4, PP. 20, 31, 34, 35, 
1922. 

Smithsonian Misc. Coll., vol. 69, No. 9, 
IQI9. 

Annals, vol. 1, pp. 244-252, 1900. 

Astrophys. Journ., vol. 18, No. 1, July 
1903. 

Annals, vol. 2, p. 5, 1908. 

Annals, vol. 2, pp. 2, 3, 21-82, 211-228, 
1908. 

Annals, vol. 2, pp. 7, 83-116, 1908. 


Annals, vol. 2, pp. 13-17, 1908. 
Annals, vol. 2, pp. 17, 57, 117-124, 1908. 
Annals, vol. 2, pp. 24, 51, 52, 1908. 


Annals, vol. 2, pp. 34-49, 1908; vol. 3, 
PP. 47-72, 1913; vol. 7, pp. 21-23, 1954. 

Annals, vol. 3, pp. 21-20, 1913; vol. 6, 
pp. 43-81, 1942. 

Annals, vol. 2, pp. 58-82, 1908; vol. 4, 
pp. 161-176, 1922; vol. 5, pp. 110-131, 
1932; vol. 6, pp. 33-42, 1942. 

Annals, vol. 2, pp. 85-98, 1908; vol. 3, 
p. 134, 1913; vol. 4, pp. 177-182, 
1922; vol. 5, pp. 244-245, 1932; vol. 
6, p. 163, 1942. 

Annals, vol. 2, pp. 104-106, 1908. 

Smithsonian Misc. Coll., vol. 74, No. 7, 
1923. 

Annals, vol. 2, pp. 106-107, 1908; vol. 
3, PP. 194-201, 1913. 

Annals, vol. 2, pp. 96-98, 110-112, 
1908; vol. 3, pp. 104-113, 1913; vol. 
4, Pp. 131-158, 1922; vol. 5, pp. 168- 
193, 1932; vol. 7, pp. 95-98, 1954. 

Annals, vol. 4, pp. 45-64, 1922; vol. 5, 
pp. 75-81, 1932. 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS 


20. Infrared and ultraviolet correc- 
tions for solar-constant work. 
21. Solar variation: 
a. First suspected. 


b. Clayton’s contributions. 


c. From solar-constant work 


1920-1930. 
d. Short up and down trends 
and (1) temperatures, 


(2) ionosphere. 

e. Accompanying (1) hurri- 
canes, (2) magnetic 
storms, 

f. Accompanying sunspots. 


g. Periodic—(1) 27-day, (2) 
6.6485-day. 


h. Long periodic and weather. 


22. A large family of periodic varia- 
tions: 
a. In the sun. 


b. In the weather. 


23. Defense of our solar-constant 
value (Abbot, Fowle, Aldrich). 
24. Brightness of the night sky. 


25. Direct and scattered radiation of 
sun and stars. 

26. Tower telescope on Mount Wilson 
and solar-drift curves. 


27. Nature of the sun’s sharp 
boundary. 


28. Volcanoes and climate. 


29. Summary of the work of the As- 
trophysical Observatory, 1890- 
1920. 

30. Radiometer measurements of stel- 
lar-energy spectra. 


VOL. 131 


Annals, vol. 5, pp. 103-110, 1932. 


Astrophys. Journ., vol. 19, p. 305, June 
1904. 

Annals, vol. 2, pp. 98-103, 117-179, 
1908. 

Annals, vol. 4, pp. 36, 185, 367-374, 
1922. 

Annals, vol. 5, pp. 246-269, 1932. 


(1) Smithsonian Misc. Coll., vol. 95, 
Nos. 12 and 15, 1936; (2) vol. 104, 
No. 13, 1945. 

(1) Smithsonian Misc. Coll., vol. 110, 
No. 1, and (2) No. 6, 1948. 


Smithsonian Misc. Coll., vol. 110, No. 
6, 1948. 

Annals, vol. 7, pp. 165-168, 1954. 

(1) Smithsonian Misc. Coll., vol. 104, 
No. 3, 1944; vol. 116, No. 4, 1951; 
(2) vol. 111, No. 13, 1949. 

Smithsonian Misc. Coll., vol. 122, No. 
4, 1953. 

Smithsonian Misc. Coll., vol. 122, No. 
4, 1953. 

Smithsonian Misc. Coll., vol. 128, No. 
4, 1955. 

Smithsonian Misc. Coll., vol. 128, No. 
3, 1955. 

Annals, vol. 4, pp. 323-366, 1922. 


Astron. Journ., vol. 27, No. 3, pp. 17- 
24, June 20, IQII. 

Astron. Journ. vol. 28, No. 16, pp. 129- 
135, March 1914. 

Annals, vol. 4, pp. 217-257, 1922. 

Smithsonian Misc. Coll., vol. 78, No. 5, 
1926. 

Scientia, vol. 19, pp. 171-181, March 
116. 

(See also Abbot, C. G., “The Sun,” 
IQOII.) 

Smithsonian Misc. Coll., vol. 60, No. 
29, 1913; vol. 65, No. 9, 1916. 

Annals, vol. 5, pp. 1-5, 1932. 


Astrophys. Journ., vol. 50, pp. 87-107, 
1924. 

Astrophys. Journ., vol. 69, pp. 203-311, 
1920. 


NO. I ASTROPHYSICAL OBSERVATORY, 1895 TO 1955—-ABBOT 7 


31. Campaign of observations of solar 
intensity on surfaces of different 
orientations and with various 
spectral regions, made at army 
camps for Quartermaster Corps, 
for a period of 8 years. 

32. Daily solar-constant values, 1920- 
1952, with 10-day and monthly 
means. 

33. Convenient table for solar-constant 
tabulations. 10-day and monthly 
mean excesses over 1.900 in 
hundredths percentages of 1.94. 
Thus 1.950 becomes 

1.950-1.900 
1.04 


s 100 = 2:58: 


Similarly 1.940 becomes 2.06. 

This difference, 0.52, is 0.53 per- 

cent of mean solar constant. 

Nore.—The tables in the two 
references cited above are 
printed without the decimal 
point for economy, and do not 
correspond with the descrip- 
tions above unless this fact 
is known. 


Annals, vol. 7, pp. 144-164, 1054. 


Annals, vol. 5, pp. 177-182, 1932; vol. 
6, pp. 85-162, 169-175, 1942; vol. 7, 
pp. 26-94, 1054. 

Smithsonian Misc. Coll., vol. 117, No. 
10, pp. 20-24, 1952; vol. 128, No. 4 
(table 1), 1955. 


Part 2.—Work of spectalists 


te 1b Be Aldrich: 

a. The melikeron, an approxi- 
mately black-body pyra- 
nometer, 

b. Reflecting power of clouds, 
and earth’s albedo. 

c. Eclipse expedition, June 
1918. 

d. A study of body radiation. 


e. Sunspots and the solar con- 
stant. 

f. Various researches on 
long-wave rays. 

g. Author (with W. H. 
Hoover) of volume 7 of 
Annals of the Astro- 
physical Observatory. 

2. F. E. Fowle: 

a. On atmospheric precipita- 

ble water. 


Smithsonian Misc. Coll., vol. 72, No. 
13, 1922. 

Annals, vol. 4, pp. 375-381, 1922. 

Smithsonian Misc. Coll., vol. 69, No. 9, 
1919. 

Smithsonian Misc. Coll., vol. 81, No. 6, 
1928. 

Annals, vol. 7, pp. 165-168, 1954. 

Annals, vol. 4, pp. 287-299, 1922. 


Annals, vol. 7, 1954. 


Astrophys. Journ., vol. 35, p. 149, 
1912, 


3: 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


b. On Avogadro’s number. 
c. On atmospheric ozone. 


d. On water-vapor absorption 
above 3 microns. 

e. On water-vapor absorption 
below 3 microns. 

f. Preparation of Physical 
Tables. 


W. H. Hoover: 

Besides his large part in vol- 
ume 7 of the Annals of 
the Astrophyical Observa- 
tory, as coauthor with L. B. 
Aldrich, he engaged in clas- 
sic researches on photosyn- 
thesis as a member of the 
staff of the Division of 
Radiation and Organisms, 
later a branch of the Astro- 
physical Observatory. 

a. Carbon-dioxide assimilation 
in a higher plant (with 
Earl S. Johnston and 
F. S. Brackett). 

b. The dependence of carbon- 
dioxide assimilation in a 
higher plant on wave- 
length of radiation. 

c. Improvements in use of 
standard water-flow pyr- 
heliometer, and in silver- 
disk pyrheliometer. 

d. Special studies of global 
sun and sky radiation 
(with L. B. Aldrich). 

e. Mechanical integrator for 
Brown recording poten- 
tiometer. 


Astrophys. Journ., vol. 40, p. 435, 1914. 

Smithsonian Misc. Coll., vol. 81, No. 
II, 1920. 

Annals, vol. 3, pp. 171-193, 1913. 


Annals, vol. 4, pp. 274-287, 1922. 
Smithsonian Physical Tables, 5th ed., 


1910; 6th ed., 1914; 7th ed., 1919; 
8th ed., 1934. 


Smithsonian Misc. Coll., vol. 87, No. 
16, pp. I-19, January 16, 1933. 


Smithsonian Misc. Coll., vol. 95, No. 
21, pp. I-13, February 27, 1937. 


Smithsonian Misc. Coll., vol. 122, No. 
5, pp. 1-10, August 14, 1953. 
Annals, vol. 7, pp. 99-104, 1954. 


Annals, vol. 7, pp. 144-164, 1954. 


Annals, vol. 7, pp. 138-139, 1954. 





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SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 2 


Mee As? CRUISE OF Ti Ms, “LOO” 


(WirtH 17 PLatEs) 


By 


MENDEL L. PETERSON 


Curator of Naval History 
U. S. National Museum 
Smithsonian Institution 












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(PuBLIcATION 4224) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
NOVEMBER 23, 1955 


THE LORD BALTIMORE PRESS, INC. 
BALTIMORE, MD., U.S. A. 


bhahebASY CRUISE: OF ELMS. shOO" 


By MENDEL L. PETERSON 
Curator of Naval History 
U. S. National Museum 
Smithsonian Institution 


(Wirt 17 PLateEs) 


In the spring of 1951 I was invited by Dr. and Mrs. George Crile, 
Jr., of Cleveland, Ohio, to accompany them on an expedition to ex- 
plore remains of a ship that had been wrecked about 5 miles off the 
main line of the Florida Keys over two centuries ago. The wreck had 
been shown to them by William Thompson, of Marathon, Fla. Only 
one thing was known about the ship—that it went down sometime 
after the year 1720. This was indicated by the fact that in 1950 
Dr. and Mrs. Crile and their party had recovered some copper coins 
from the site, one of which was a Swedish half-ore piece (pl. 2, fig. 1) 
dated 1720. The site was a reef named “Looe” on the charts and was 
located some 25 miles southwest of Marathon. At the time, the source 
of the name was unknown, and its presence on the charts was not 
considered significant. 

On Sunday evening, May 27, most of the members of the expedi- 
tion assembled in Miami, and the next day left for the Keys and 
Thompson’s yacht harbor, which was to be the base of operations. 
Here the entire party came together. It consisted of the sponsors, 
Dr. and Mrs. Crile; Mr. and Mrs. E. A. Link, of Binghamton, N. Y.,? 
Mr. and Mrs. James Rand, of Cleveland, Ohio; Mr. and Mrs. John 
Shaheen, of New York City ; William Thompson, of Marathon, Fla. ; 
Arthur McKee, of Homestead, Fla., an experienced diver on ship- 
wreck sites; and myself. Necessary supplies and equipment were as- 
sembled and tested, and the boats were made ready. We were to use 
a barge built on a Higgins boat hull, a small fishing launch, and later 
Mr. Link’s yawl, the Blue Heron. 

Early Wednesday morning, May 30, the party left for the reefs 


1 The participation of Mr. and Mrs. Link in the expedition was to prove very 
fortunate for the National Museum since it led to the establishment of the Link 
Fund through their generosity. This fund enables the Museum to participate in 
annual expeditions to explore historic wreck sites in the Florida Straits area. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 131, NO. 2 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


and by midmorning were over the wreck site (pl. 3). To locate the 
wreck exactly it was necessary to get into the water with face plates 
and carefully scan the sand bottom of a “valley” lying between two 
fingers of the reef which ran out to sea. Only the metal objects from 
the ship remained, and these were covered with a sand crust giving 
them the same color as the bottom—excellent camouflage, which made 
them almost invisible from the surface of the water. We detected the 
outlines of bars of metal, the ring of one of the ship’s anchors (pl. 4) 
jutting from the reef, two long cylindrical objects, encased in marine 
growths, which were recognized as guns from the ship, and, upon 
closer inspection, piles of round objects encrusted with sand—solid 
shot for the ship’s guns. 

After a preliminary survey from the surface, the boats were pulled 
over the wreck and the diving gear was prepared for operation. 
Tight-fitting rubber masks that completely covered the face were 
connected to the air compressors by long lengths of strong rubber 
hose. The divers strapped on their lead belts, put on their masks or 
helmets, and went over the side on the diving ladder and lifeline. 

The first object recovered was a large chunk of metal roughly tri- 
angular in cross section and stepped on the surface. It proved to be 
solid cast iron and was identified as permanent iron ballast cast to fit 
along the keelson of the ship. Clinging to it was a solid iron shot 33 
inches in diameter, a standard 6-pound shot of the eighteenth century. 
These finds immediately revealed two additional facts concerning the 
ship—she was most probably a warship, since merchantmen carried 
disposable ballast of stone, and she had 6-pounder guns in her 
batteries. 

The next day the attention of the divers was devoted to the smaller 
objects lying about in the sand “‘potholes” on the site. By the use of 
a powerful jet of water the sand was carefully washed away and the 
articles were uncovered (pl. 5). Soon basketfuls of sand-encrusted 
hull bolts, nails, solid iron shot, fragments of rum or brandy bottles, 
Chinese porcelain dishes, and earthenware, and many other objects 
were being emptied on the decks of the salvage boats. On the first 
of two brief dives that the author made on the site a basketful of 
solid iron shot was gathered (pl. 6). In this lot was found a 6-pound 
shot with an arrow on it, which was immediately identified as the 
broad arrow (pl. 7). This was the first indication of the nationality 
of the ship, as this symbol has been used for centuries by the kings 
of England and Great Britain to mark royal property. The occurrence 
of the broad arrow on the shot was not conclusive evidence that the 
ship had been British, since ordnance stores could have been captured 


NO. 2 LAST CRUISE OF H.M.S. “LOO”—PETERSON 3 


or stolen by the enemies of Britain. But until further evidence proved 
the contrary, we could consider the ship to have been British.? In the 
basket 12-pound, I-pound, and 4-pound shot were also found, giving 
additional information on the ship’s batteries. The broad arrow also 
appeared on the 12-pound shot. 

On Thursday, May 31, and Friday, June 1, numerous small objects 
were brought up, including more porcelain fragments, parts of clay 
pipes and rum bottles, the wooden knob of a walking stick, the eye- 
piece of a navigation instrument, pieces of stoneware decorated with 
blue flowers, and animal bones (later identified as pig and cow) from 
the pickled-meat stores of the ship. (See pls. 8, 9, and 10.) 

On Saturday, June 2, the party remained ashore to sort, clean, and 
begin the preservation process on the objects recovered. Fragments of 
wood were packed in fresh water for shipment to the National Mu- 
seum, the sand crust was cleaned from the cast-iron and other large 
iron objects by light hammering, and the objects were placed in baths 
of fresh water to leach out the sea salts. 

The cast iron was found to have been oxidized to a depth of one- 
half to three-fourths of an inch. The removal of the sand crust from 
all surfaces (those portions having been converted to crystalline mag- 
netite, which was very friable) had to be done with great care. The 
porous oxidized layer was saturated with chlorides, and to break these 
down the cleaned shot were placed in baths of sodium hydroxide. 
Most of the shot were packed with the sand crust on them, the crust 
protecting them from excessive drying while on the way to the 
Museum. 

On Sunday and Monday (June 3 and 4) a continual stream of ma- 
terial came up from the wreck and was added to the piles ashore at 
our base (pls. 11, 12). There was such a quantity of specimens that 
_it was decided that the author should devote a full day to identifying, 
sorting, cleaning, and preserving those that were to be retained. The 
boats went out as usual and that evening returned with one of the 
cannon barrels. Mr. Link had rigged the main boom of the Blue 
Heron and had lifted it to her side with block and tackle (pls. 13, 
14). Through skillful seamanship and favorable weather the 2,000- 
pound barrel was brought to Marathon, hanging beside the delicate 
mahogany hull of the yawl, which was protected with rope fenders. 
As soon as the barrel was on the ground at our base we began re- 
moving the sand crust with a hammer. As the crust fell away (pl. 15) 


2 Later a chain plate, which was originally bolted to the ship’s hull, was found 
by a salvage party from Miami. It also bore the broad arrow. 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


the beautifully molded contours of an eighteenth-century barrel were 
revealed (pl. 16). The appearance was deceptive, however, for while 
the physical form of the barrel was perfect it was evident that the 
surface of the iron had been oxidized deeply. 

Two more clues to the identity of the ship were furnished by the 
barrel. On the second reinforce over the trunnions was a crowned 
rose, and in the muzzle were the remains of a wooden tompion. The 
crowned rose was an insigne employed by the Tudor and Stuart 
monarchs of England and was not used as the principal mark on royal 
artillery after the death of Queen Anne in 1714. The fact that the 
tompion was in the barrel indicated that the ship had run aground 
through accident or storm and not as an aftermath to a naval engage- 
ment. The crowned rose enabled us to estimate the date before which 
the ship sank. Since the normal life of an iron barrel on shipboard 
was usually not over 35 or 40 years, the barrel had probably not been 
in active service after 1714 plus 35 or 40 years, or 1749-1754. It was 
therefore assumed that the ship must have sunk before the year 1750. 

The barrel was the last important object recovered from the wreck 
site in 1951, and we now had all the evidence that we could expect to 
recover that year. This evidence had told us that the ship was a 
British warship, had sunk between 1720 and 1750, had 6- and 12- 
pounders in her main battery, and had gone down as the result of an 
accident and not as a sequel to a naval action. We knew, of course, 
that the reef on which the ship had sunk was called “Looe Reef,” but 
we had not suspected a connection between the name and the wreck 
itself. 

After my return to Washington I began a search of the ship casualty 
lists for the eighteenth century published in Clowes “The Royal 
Navy,” and found the entry—“‘1743 Looe 44 guns, Capt. Ashby 
Utting, Lost in America.”’ Further research indicated that she carried 
6- and 12-pounders. The conclusion was obvious—the ship we had 
investigated was the Loo whose wreck had given her name to the reef. 
That night I phoned Dr. and Mrs. Crile in Cleveland and they immedi- 
ately called a friend in London. Within a week the Public Record 
Office had yielded a letter written by Captain Utting at Port Royal, 
S. C., February 15, 1743/44,° in which he described the wreck of his 
ship. A year later I was in London digging out all the documents in 
the Public Record Office relating to the ship. The account that fol- 
lows is based on letters, the Navy List, the Loo’s pay lists and muster 


8 The legal year began March 15. The calendar year was 1744. (See Appen- 
dix B for Utting’s letter.) 


NO. 2 LAST CRUISE OF H.M.S. ‘‘LOO’”—PETERSON 5 


rolls and other documents in the Public Record Office in London. All 
these documents are Admiralty papers. The references are given in 
the manner in which they are numbered in the collections of the Pub- 
lic Record Office, and bear the prefix ADM or AD. 

On June 14, 1743, Thomas Corbett, Secretary of the Admiralty, sat 
down in his London office and countersigned an order directing Capt. 
Ashby Utting to prepare his ship, the frigate Loo,’ for a cruise to 
North America: 


Having order’d His Majesty’s ship under your command to be refitted at 
Portsmouth, for a voyage to North America, cleaned, sheathed and graved,> and 
her provisions compleated to six months of all species, except beer, and of that 
as much as she can conveniently stow, and stored accordingly; you are hereby 
required and directed, to repair with her into Portsmouth Harbour, and strictly 
to observe the following instructions. 

You are to give constant attendance. 

Wie ie? Gis 
By TG: 


Thus began the last cruise of the Loo, the story of which might have 
been taken from a classic work of fiction. 

The Loo, a frigate of 40 to 44 guns, had seen long service in the 
Royal Navy. She had been built during the expansion of the British 
fleet incident to the War of the Spanish Succession. In this war 
England was fighting to prevent the seating of a Bourbon’ on the 
throne of Spain—a scheme of Louis XIV to strengthen the position 
of France in Europe. The Loo was to meet her end during another 
war in which Spain and Great Britain were enemies, a war that began 
as a result of the succession of Maria Teresa to the throne of Austria. 


4 Named for the old seaport town of Looe (also Loo), which lies on the rocky 
coast of Cornwall and which has supplied sturdy sailors to the Royal Navy 
since its beginning. 

5 The first Royal Navy vessel to be sheathed with lead was the Phoenix; this 
was done in 1670. The practice had been followed in the Spanish Navy since the 
middle of the sixteenth century and in some cases by English merchant ships 
(see Clowes, The Royal Navy, vol. 2, p. 240). Lead proved impractical, how- 
ever, and the practice of sheathing with thin fir boards was followed until the 
time of the American Revolution, when the British fleet was sheathed with 
copper. The thin fir sheathing was backed with pitch and horsehair, which dis- 
couraged worms from tunneling into the ship’s planking. Graving was the proc- 
ess of burning sea life from the bottom of a ship. 

® Tnitials of the Lords of the Admiralty, “W” for Daniel, Earl of Winchelsea, 
First Lord of the Admiralty, March 19, 1742, to December, 1744. “T.C.” for 
Thomas Corbett, Secretary of the Admiralty, 1742-1751. (Admiralty Out- 
Letters, ADM 2, vol. 60, p. 15, Public Record Office.) 

7 The grandson of Louis XIV, who ruled as Philip V of Spain (1700-1746). 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


She was most probably launched in 1706 since she first appears 
in the Navy List under the date April 1, 1707, when she was at “Long- 
reach taking in Guns.” ® Her complement at that time is given as 
190 men and her battery as 42 guns. 

The Navy List thereafter follows her career in terse monthly entries 
(see Appendix A). 

Her first two cruises carried her to Archangel and Newfoundland.?° 
By 1709 she was back at Sheerness refitting, and afterward was on 
patrol duty in the Channel.‘ Early in 1710 she was attached to the 
Dunkirke Squadron and in the fall of that year was on convoy duty to 
the eastern countries.’* In April 1711 she was in Holland “to bring the 
Queen’s wine to the Nore.’”’** She then sailed convoy to Russia and 
during the last two months of 1711 was refitted and again sent to duty 
in British waters, meeting ships from Virginia and convoying them 
to British ports.4* During the winter of 1712-13 she transported 
troops to Bayonne and returned with prisoners of war and then 
again served in the Channel, cruising against smugglers.1* In the fall 
of 1714 she was sent convoy to Port Mahon in the Mediterranean and 
on return was paid off and laid up."7 

Three years later the Loo was back in service as a hospital ship on 
duty with the Baltic Squadron.’* She was then laid up for the winter 
and the next spring again commissioned as a hospital ship and sent to 
the Mediterranean Squadron. A year later (April 1719) she appears 
in the Navy List with 30 guns and a crew of 125 men, which probably 
indicates that she had been reconverted to a frigate.1° Thus fitted out 
she served with the Mediterranean Squadron until the spring of 
1722.°° From that time until January 1728 she appears to have been 
laid up. On January 10, 1728, she was again in sea pay and until 


8 At least one earlier Loo is recorded. Like her successor, she was a frigate 
of 40 guns and was also lost through shipwreck, having run aground on the 
Irish coast near Baltimore, April 30, 1697 (Clowes, The Royal Navy, vol. 2, 
p. 536). 

® Navy List, Jan. 1, 1707-Dec. 31, 17090, ADM 8/10, Public Record Office. 

10 ADM 8/10. 

11 Thid. 

12 Thid. 

13 Thid. 

14 ADM 8/10 and 8/11. 

15 ADM 8/11. 

16 ADM 8/12. 

17 bid. 

18 ADM 8/13. 

19 Tbid. 

20 Tbid. 


NO. 2 LAST CRUISE OF H.M.S. “LOO’’—PETERSON 7 


July 1730 was on duty in British waters performing such tasks as 
transporting clerks and money to the pay at Plymouth and patrolling 
the Channel.?* In August she was ordered to the Mediterranean as 
convoy for transports going to Gibraltar and remained in the Medi- 
terranean cruising against the Barbary pirates “on the Coast of 
Sallee’ 

Coming home to Britain in August 1731, the Loo was again on 
Channel service until the next spring. For three years she was again 
laid up and on May 5, 1735, was commissioned and fitted out as a 
hospital ship for duty in the Channel service. In August of that year 
she joined the naval forces at Lisbon, still as a hospital ship, and 
served there until the spring of 1737.°* From that time until January 
1742 she was laid up and, war having broken out between Great 
Britain and Spain, was recommissioned as a frigate of 44 guns on 
January 5 and placed in the Channel service under the command of 
the Earl of Northesk. While on a cruise in the area of Cape Finnisterre 
(northwest Spain) the Loo, in company with the Dealcastle (24 guns) 
raided Vigo Bay, capturing four Spanish vessels in the harbor, an 
incident reported in the London Gazette for August 31, 1742.7 


21 Tbid. 

22 Thid. 

23 ADM 8/17 and 8/18. 

24 ADM 8/t19 and 8/20, 

25 Also mentioned in the Gentlemen’s Magazine for August 1742, p. 445, and 
September 1742, p. 494, giving an account of the Loo raiding in the Porto Nova 
and Santiago areas: “The Earl of Northesk, Capt. of his Majesty’s ship the 
Loo, being on a cruize off of Cape Finisterre, and the parts adjacent, received 
intelligence of a small Privateer being at Porto Nova, upon which he stood in 
there on the 30th of June, but the Privateer discovering him, got higher up the 
river than the Loo could venture, and it falling calm, Ld. Northesk was obliged 
to anchor close by the towns of Porto Nova, and St. Iago, into which he fired 
a few shot, then landed some men and dismounted 4 guns which were on a bat- 
tery at Porto Nova, and set fire to several houses at St. Iago. On July 7, Lord 
Northesk met with his Majesty’s Ship the Dealcastle, commanded by Capt. Elton, 
and receiving intelligence of some vessels being at Vigo, they run up the river 
and anchored before that town, where they made prizes of 4 vessels, 2 of which 
they set on fire, being light, and not having Sails on board to bring them out. 
They fired several shot into the Town to cover the boats while they cut away 
the vessels, there being a pretty smart fire at them with small arms from the 
shore. On July 19, upon intelligence that the privateer was still about the river 
of Porto Nova, the Loo run in and anchored under the Island of Blydones, where 
Lord Northesk put a Lieutenant and 60 men, with 2 of the ship’s 6 Pounders, 
into a Sloop taken at Vigo, and sent her up the river in quest of the privateer ; 
the Sloop could see nothing of her, but in her return chased a bark on shore, and 
set her on fire; and Lord Northesk landed some men, and burnt a village of 
about 40 houses.” 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I3I1 


Following this cruise, the Loo was again in the Channel service until 
May 1743, when she was ordered “To cruize between Bilbao and 
St. Jean de Luz,*® to intercept some Caracca ships *7 expected at 
St. Sebastian.” 78 At the conclusion of this cruise, which lasted some 
six weeks, the Loo returned to Portsmouth to refit for her ill-fated 
voyage to North America. Meanwhile Capt. Ashby Utting had as- 
sumed command under a commission dated 4 April, 1743.°° As the 
war between Great Britain and Spain had progressed, the people of 
the infant colonies of Georgia and South Carolina had felt increasing 
fears of an invasion by the Spanish from Florida and Cuba. In 1742 
the Spanish had attacked Fort Frederica in Georgia but had been re- 
pulsed by the troops of General Oglethorpe at the Battle of Bloody 
Marsh. After this attempt by the Spanish the colonists felt that the 
threat of devastation of their homes and farms was even greater. Con- 
sequently, the Lords of the Plantations in London had been petitioned 
by the Governor of South Carolina to send a large warship to the 
Carolinas for the protection of the coastline. The result was the order- 
ing of the Loo to the Charleston station. 

Three days after Secretary Corbett signed the Loo’s orders Cap- 
tain Utting had received them and replied that he would “punctually 
comply” with them and use his “utmost endeavours” to get his ship 
ready for sea.*° The Loo sailed soon after Utting’s letter was posted 
and arrived at Portsmouth on the morning of June 18. Reporting his 
arrival there to the Secretary of the Admiralty, Utting complained 
that the 6-pounders ** on the upper deck of the Loo were “‘very indif- 
ferent and not fitt for a forrain voyage, being much honey combed.” *? 
a fact made known to him by his gunner, Samuel Kirk.** Utting 
recommended that a battery of 9-pounders, which had been mounted 
for the Hunnington, be substituted for the Loo’s worn-out 
6-pounders.** 


26 On the northern coast of Spain. 

27 That is, ships of the Caracas (Venezuela) Company, 

28 ADM 8/23. 

29 Commission and Warrant Book, 1743-1745, AD 6/16, p. 335. 

80 Admiralty In-Letters, ADM 1, vol. 2625, pt. 3, No. 146. 

81 Heavy guns of this period were rated by the weight of the solid shot they 
threw. The barrel of a long 6-pounder of this period weighed around 2,000 
pounds. 

82 That is, the barrels, which were cast iron, had small cracks in their bores. 

88 Kirk’s name is mentioned in the record of the Court Martial of Captain 
Utting held May 3, 1744 (Admiralty In-Letters, ADM 1, vol. 5283). 

84 ADM 1, No. 417. Utting’s recommendations were not followed. This is 
proved by the finding of the same 6-pounders on the wreck site of the Loo. They 


NO. 2 LAST CRUISE OF H.M.S. “LOO’’—PETERSON 9 


Preparations for the cruise proceeded swiftly. On June 20, the day 
after Utting wrote his letter concerning the guns, the Admiralty 
ordered the Captain to “make out” his pay books “to the 30 June, 
1742.” *° Five days later admiralty orders “about carrying candles up 
and down the ship and drawing off spiritous liquors and an order to 
cause the men’s allowance of rum to be diluted with water when in 
the West Indies” were issued.*® The order directed that “whenever 
the ship’s Company under your command are served with Rum, 
Brandy, or any other spirituous liquor, instead of Beer, the same be 
constantly issued out to them by the Purser upon the open Deck, and 
nowhere else ; and that you do order all officers and others under your 
command, never to draw off any arrack,*7 rum, brandy, or other 
spirituous liquors in places under deck, but always upon open deck.” *° 





bore the crowned rose, a device placed on royal guns during the reigns of the 
Tudors and the Stuarts. The Loo’s 6-pounders were therefore cast before the 
death of Queen Anne in 1714. Thus they would have been at least 30 years old 
at the time of the loss of the Loo—a fact borne out by Utting’s statement on their 
condition. 

35 ADM 2, vol. 60, p. 34. A year’s delay in paying the men was a common 
(even usual) occurrence at this time. 

86 Tbid., p. 41. These orders stemmed from the loss of the Tilbury, 60 guns, in 
the West Indies through fire on September 21, 1742. The incident was reported 
by Adm. Edward Vernon in a letter to Thomas Corbett written on the flagship 
Boyne in Port Royal harbor, Jamaica, October 3, 1742 (Admiralty In-Letters, 
ADM 1, vol. 233, extracts.) “I am heartily concerned for the melancolly 
account lately brought me by Captain Lawrence late of the Tilbury, who came 
in here the 24 September in the Island Sloop, with part of his officers and men, 
another part remaining on board the Defyance, in execution of my orders, and 
upwards of a hundred of them having perished in the sea or fyre, on her acci- 
dentally taking fyre, and burning, and sinking in the sea, amongst which are the 
Master, Boatswain, and Gunner, and a Marine Officer. But I cant proceed to 
enquire in it at a Court Martial, til the return of the Defyance, many evidences 
that saw the first of it, being absent in the Defyance, so all I can say of it at 
present is, that it took its rise from a Marine soldier’s snatching to get a bottle 
rum, out of the Purser’s boys hand, who had a candle in the other hand, declar- 
ing he would have a dram, and in the struggle with the boy, the bottle falling 
and breaking, and the candle with it the rum took fire, and communicating to 
more in the Pursers cabbin where the fyre first began, that could not be extin- 
guished by all their diligence afterwards, tho they say, they threw all their 
powder into the sea.” Admiral Vernon at the same time submitted a copy of a 
general order he had published to his forces two years before requiring that the 
rum ration be served to the men on deck, and that it be diluted with water. The 
new concoction became known as “grog” after Admiral Vernon who was called 
“Old Grog” from his habit of wearing a “grogram” cloak. “Grogram” was a 
coarse material of silk and mohair. The name is derived from “gros-grain.” 

87 A drink distilled from rum. 

88 Admiralty Out-Letters, ADM 2, vol. 59, p. 380. 


IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


Another order dated the same day directed Utting to have his ship 
“vichialled to four months only *® for a forreign voyage, and what beer 
she cannot take in to be made up with good brandy . . . and to cause 
half of one and half of the other to be issued.” It also instructed 
Utting to load the food as quickly as possible and “to take care, that 
the brandy supplied . . . be good and wholesome,” and to report to 
the Lords of the Admiralty “the usefullness of the allowance of half 
brandy and half beer and what effect it has upon the health of the 
men.” *° Utting silently conformed with these orders as they were 
received, but on July 2, still hoping to receive the battery of g-pounders 
to replace his wornout 6’s, he wrote: “The time for taken in my 
guns draws very near, and ye officers of ye ordinance here has no 
orders concerning ye 9 pounders, which I had wrote for and was in 
hopes I should have had them as ye ship would well bare them and 
make her a much better man of war.” ** He also requested that, if 
possible, he be told his ultimate destination since he knew only that 
he was to go to North America. 

On July 11 orders were issued to the commanding officers of the 
Rye, 20 guns; Flamborough, 20 guns, and the sloop Spy, 8 carriage 
guns and 12 swivels, all stationed in South Carolina, to place them- 
selves under the command of Captain Utting upon his arrival there. 
Utting was to carry these orders with him.*? The next day the Ad- 
miralty issued instructions to Utting concerning the impressment of 
seamen while in America, furnishing him with three press warrants. 
The instructions cautioned him that “it is not meant, that the trade of 
His Maj’s. subjects in America, or ships provided with Letters of 
Marque to cruise against the Enemy should be distressed thereby, 
but only that such prudent use be made of the said press warrants as 
may enable you to procure men to make up your complement, when 
proper opportunities offer it. You are to take great care, that no 
indiscreet or unreasonable use be made of them.” The instructions 
also directed that Utting was “never to molest the chief officers, such 
as the master, mate, boatswain or carpenter, or any seaman found on 
board with protections granted by us, pursuant to Act of Parlia- 
mient., 


89 Thus rescinding the order of June 14, which had directed the loading of a 
6-months’ supply of food. 

40 Admiralty Out-Letters, ADM 2, vol. 60, p. 42. Beer had been a standard 
beverage in the English Navy since earliest times. Easy to keep, it was superior 
to water, which grew putrid in the casks. 

41 ADM 1, vol. 2625, pt. 3, No. 418. 

42 ADM 2, vol. 60, p. 79. 

43 Thid., p. 80. 


NO. 2 LAST CRUISE OF H.M.S. “LOO’’—PETERSON II 


The same day detailed orders covering all phases of the cruise to 
North America were issued. They are an excellent example of the 
type of orders of that period given to senior officers destined for inde- 
pendent duty in remote parts of the Empire, and they are here quoted 
in their entirety.** 


TO CARRY GOVERNOR CLINTON TO NEW YORK AND THEN 
ATTEND ON SO. CAROLINA 


Whereas we have appointed His Maj’s. ship under your command to carry 
the Hon. Geo. Clinton, Esq. to his Government at New York, and then to attend 
on the Colony of South Carolina, you are hereby required and directd to make 
all possible dispatch in getting her compleated in all respects for the Sea, and 
you are to receive on board the said Mr. Clinton, with his Family and Equipage, 
and give them passage to New York, vichialling them as your Ship’s Company 
during their continuance on board, and allowing the Governor all such accom- 
modation as the Ship will afford. 

And whereas the ship under your command is only ordered to be vichialled to 
four months, and to have two months French Brandy instead of two months 
beer; and the Comrs. of the Vichialling having a large quantity of Brandy in 
store at Guernsey, in the Charge of Mr. Nich S. Dobree, a merchant in that Is- 
land, you are in your way down the Channel, to call off of Guernsey, without 
going into the Port, and send the enclosed letter with your Purser on Shore to 
the said Mr. Dobree; and receive from him such a quantity of Brandy as you 
think necessary for the use of your Ship’s Company and you can conveniently 
receive on board, which when you have done, you are to proceed directly to 
New York, without touching at the Madeiras, and there land the Governor, with 
his Family and Equipage; and having so done, you are to proceed on to South 
Carolina. 

And whereas His Maj’s Ships the Rye, Flamborough, and Spy Sloop, are 
stationed at South Carolina, You are to take them under your command, their 
Captains being directed to follow and observe your orders. 

When you arrive at South Carolina, you are to communicate these our instruc- 
tions to the Governor and Council of that Province, and to consult and advise 
with them from time to time, in what manner the ships under your command 
may be best employed in guarding the coasts, and securing the trade of that 
colony from any attempts of the Enemy, and to govern yourself according as 
shall be agreed on, using your best endeavors to take or destroy all such ships 
or vessels of the enemy, as shall come upon the coasts of the said Colony. 

And whereas it has been represented to us, that the Coast of North Carolina 
is very much infested with Spanish Privateers, who have even landed in the 
Country and carried off hogs and black cattle, to the great terror of the inhabi- 
tants of those parts, you are, when you see proper occasions, to extend your 
cruize as far as Cape Hatteras, or to order one of the ships under your Com- 
mand to do so, for the better protection of the trade of His Maj’s. Subjects in 
those parts; and you are to acquaint the Governor of North Carolina with this 
part of our instructions. 


44 Admiralty Out-Letters, ADM 2, vol. 60, pp. 81-84. 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


And whereas it has been represented to us, that the Town of St. Augustine 
depends much upon what comes by Sea for provisions, and would be greatly 
distressed, if His Maj’s. ships stationed at Carolina would sometimes cruize off 
that Port, to prevent provisions being carried to that place by Sea, you are to 
have a particular regard to that service, as far as may be consistent with the 
other necessary services on which you may be employed. 

And whereas we have directed the Captains of His Maj’s. ships attending on 
Virginia to hold a constant correspondence with you, you are, whenever you 
shall find the Enemy’s ships to be too strong for you, to send immediate advice 
thereof to the Captains of the said ships, whom we have directed to repair to your 
assistance, and you are to endeavour joyntly to take or destroy them. And if 
the Captains of the said Ships shall at any time send you notice of the Enemy 
being too strong for them, you are with all possible diligence to proceed to their 
assistance, communicating in the first place the intelligence you have received to 
the Governor and Council of South Carolina, and receiving their concurrence 
for your so doing, and when the service is performed, you are to return to your 
station. 

And whereas the Captains of His Maj’s. ships stationed in America, have of 
late years taken a very unwarrantable Liberty of lying in Port with their Ships, 
for the greatest part of the time they have remained abroad, to the dishonour of 
His Maj’s. service, and the disservice of the Colonies for whose protection they 
are appointed, and we being determined not to suffer any such neglect for the 
future, do hereby strictly charge and direct you to keep constantly at Sea, when 
the weather will permit, and cruize in proper stations for meeting with the 
Enemies ships or privateers, and for protecting the trade of His Maj’s. subjects, 
and guarding the said colony of Carolina from any attempts of the Enemy. 

You are not to fail to transmit to us, one in every two months an exact copy 
of the Journal, that it may be seen what care and diligence you have used in 
putting our instructions in execution and to order the Captains of His Maj’s 
ships under your command to do the same. 

And in order to enable you the better to keep the ships under your command 
in a good condition to cruize and protect the trade, as well as to annoy the 
Enemy, you are to cause them to be cleaned once in six months, at such times 
as it can be most conveniently done. 

When the ships you command are in want of provisions, you are to apply to 
the Contractors of the vichialling at Carolina, for the same, and never to leave 
the said Colony defenceless by going somewhere to vichial; and you are to 
take on board no more provisions at a time, than are necessary for the service 
on which you are employed. 

You are not to hoist the Union Flag on board the Ship you Command, on 
account of the Governor’s being on board, or on any other pretence whatever. 

In case of the death of any of the officers of the ships under your command, 
you are to appoint such other persons to act in their names, as by the quality of 
their Employments ought to succeed therein. 

When you shall receive our orders to return to Great Britain, you are to take 
in no more provisions than shall be sufficient to compleat what you may have on 
board to three months of all species at whole allowance, upon the penalty of 
making good what damage, His Majesty may otherwise receive thereby. 

You are, as you pass through the channel, to examine such ships and vessels 
as you shall meet with passing from Great Britain or Ireland to France, which 


NO. 2 LAST CRUISE OF H.M.S. ‘“LOO”—PETERSON 13 


you shall reasonably suspect to have Wool 4 on board, and upon discovering 
any with that comodity in them, to send them into the nearest Port, and deliver 
them into the care of the Collector of the Customs, in order to their being prose- 
cuted according to Law. 

You are by all opportunities to transmit to our Secretary for our information, 
an account of your proceedings, and of the condition of the ships under your 
command as to the number of men, and all other particulars and in case of in- 
ability by sickness or otherwise, to be careful to leave these our instructions 
with the next Commanding Officer. Given 12th July, 1743. 

Wo ifcCr0.Giv By 
By 
TG 
Capt. Utting, Loo, Spithead. 


Captain Utting must have received oral instructions that he was 
to carry Governor Clinton to New York several days before he re- 
ceived the above orders. In fact the Governor had either visited the 
ship or had otherwise instructed Captain Utting on the accommoda- 
tions that he desired aboard the Loo. Five days before the detailed 
orders on the cruise were written Utting had written Corbett “the 
carpenter will have compleated every conveniency Mr. Clinton desires 
by tomorrow night . . .” * 

On July 14 Utting acknowledged receipt of the orders of July 12 
and reported that his ship was “in all respects fitt for sea.” 47 Four 
days later the Admiralty instructed Utting, who was now at Spithead 
ready to sail, to convoy the storeship Pegasus “laden with naval stores 
for New York and South Carolina” to America, ordering that he 
“convoy her safely to New York, where you are to cause her to be 
unloaden as soon as possible, and then proceed with her to South 
Carolina.” 4 At the same time additional instructions on cruising 
while in America were issued.* 


TO CRUIZE BETWEEN CAPE FLORIDA, AND THE NORTH WEST 
PART OF THE GRAND BAHAMA WHEN THE SEASON OF THE 
YEAR WILL NOT PERMIT HIS CRUIZING OFF CAROLINA. 


In addition to our instructions to you dated the 12th instant, you are hereby 
required and directed, when the Season of the Year is not proper for your 
cruizing on the Coasts of South Carolina, and that neither the said Colony, nor 





45 The export of English wool was absolutely prohibited at this time in an 
effort to encourage the English woolen industry. The demand for English raw 
wool in the lowlands was great, and consequently the smuggling of it to the con- 
tinent was profitable. 

46 ADM 1, vol. 2625, pt. 3, No. 419. 

47 Thid., No. 420. 

48 ADM 2, vol. 60, p. 96. 

49 Thid., pp. 96-97. 


14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


that of Georgia is under any apprehension of being molested by the Enemy from 
Havanna or Augustin, to proceed with His Maj’s. Ship under your Command 
and Cruize between Cape Florida and the North West part of the Grand Ba- 
hama, ’til such time as the Season will permit your return to Carolina, taking 
care to have a sufficient quantity of provisions on board to last you on that 
service. 

You are diligently to look out for the Enemy’s ships passing through the Gulph 
of Florida for Europe, and use your utmost endeavours, to take, sink, burn or 
destroy them. 

But before you proceed on this Service, you are to communicate your design to 
the Governor of Carolina, and not to go thereupon, if you find any reasonable 
objections thereto. Given 18th July, 1743. 


Wak. sB. 
By 
TC 
Capt. Utting, Loo, Spithead. 


Utting had been thinking of the same operations plan as that of the 
Lords of the Admiralty, for on July 19, a day or so before he received 
the additional instructions, he had written: 


I was a little hurried to save post with my last yet dont know whether I ex- 
plained ye plans and time of cruising so plain as you could wish for fear of 
which beg you'll be pleased to indulge me with this to acquaint you. I propose 
(if ye service will allow me and you can git me orders) to saile from South 
Carolina ye 10 or 15 of October and cruise in and about ye Gulfe of Florida, as 
far as ye Cape °° if I can git there till ye middle of Jany. then return to Caro- 
lina. And as soon as I can water, victuall, and refitt, in all respects, then pro- 
pose to saile, and cruize on ye coast of Carolina of [off] St. Augustine or on 
such part of ye coast as I shall find the service require me most. Given ye 20 
gun ships proper stations as ye service shall require, on this coast I propose to 
keep all ye summer months 51; ye latter end of May shall go in for 6 or 8 days 
to victuall and water and then cruize till ye 20 or 25th. of July when as I shall be 
then about 12 months foull shall go in to heave down and about ye 2oth of 
Septr. shall saile to cruize on ye aforesaid station: yet I never propose to be in 
port above 2 months in ye year; after my first careening shall heave down every 
6 months. But as I am graved and tallowed 52 can go 12 months at first. There 
is an exceeding good careening place at Port Royall 5% which can be made ours 


50 Cape Florida. 

51 That is, keep to the sea during the summer months. 

52 See footnote 5, p. 5. In navy yards graving was usually done in a drydock. 
On remote stations it was necessary to careen the vessel by mooring her in a 
river, unloading her, and then “heaving her down” by pulling her over with 
tackles secured to trees on the bank. In this position half of her bottom was 
above water and could be cleaned. The process was repeated for the other side 
of the bottom. Hulls were coated with tallow as a protection against growths 
and water penetration of the ship’s planking. 

53 South Carolina. 


NO. 2 LAST CRUISE OF H.M.S. ““LOO’’—PETERSON 15 


conveniently to heave down without expense to the government. I have wrote 
to ye Navy Board for careening gear, but have not had an answer.®4 


On July 25 the Captain acknowledged receipt of the further instruc- 
tions on cruising and the orders to escort the Pegasus, and prepared 
to set sail.°° 

On August 6 Governor Clinton, his wife and her children, and suite 
of 15 persons came aboard the Loo.*® She probably sailed within a 
week, 

Six weeks later the Loo arrived safely in New York harbor with 
her charges, and the Pegasus. The Governor and his suite disem- 
barked on September 22.°7 Utting reported in a letter dated in New 
York Harbor September 29 that the voyage had been uneventful 
“with nothing worth their Lordships notice.” In the same letter the 
Captain made his first report on the trial ration of half brandy and 
half beer, stating that it agreed with men “extreamly well, and they 
are well pleased.” °* The ship, he reported, was unmooring as he 
wrote, and expected to sail that afternoon for South Carolina escorting 
the Pegasus. His departure was delayed until October 6, however, 
probably by adverse weather, but the bright lights of New York might 
have been the real reason, since Utting mentioned no cause for the 
delay. After a passage of five days the Loo arrived off Charleston 
Bar. In Charleston he found the sloop Spy ready for sea, the Rye 
“cleaned and almost fitt for sea,” the Flamborough “sheating.” °° He 
immediately delivered the Admiralty orders instructing the captains 
of these ships to place themselves under his command, and then issued 
orders giving each ship stations for cruising off the Carolina coast for 
the defense of the colonies and protection of English and colonial ship- 
ping. Captain Hardy of the Rye was directed to “cruize on the coast 
of South Carolina, between Charles Town Barr and the So.W most 
part of the same coast, keeping off St. Augustine, and as near into the 
shore as you shall judge proper when winds and weather will permitt 
to intercept any trade that may come from the Havanah to that 
place.” °° Hardy was also instructed to inform the Governor of 


54 ADM 1, vol. 2625, pt. 3, No. 421. 

55 Thid. 

56 ..o0’s General Muster Book, ADM 36, Ser. I, vol. 1823. 

57 Loo’s General Muster Book, ADM 30, Ser. I, vol. 1823. 

58 ADM 1, vol. 2625, pt. 3, No. 423. 

59 Tbid., No. 435. British ships were at this time sheathed with thin fir boards 
backed with horsehair and pitch. The sea worms ate through the thin board but 
were repulsed by the hair, and the ship’s hull planking was thus protected. 

60 ADM 1, vol. 60, No. 435. 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Georgia °*' of his activity off that coast, to remain at sea as long as his 
water and provisions would permit, and, after returning to Charles- 
ton to take on supplies, to return to his station and cruise as soon as 
his ship was ready for sea. Captain Hamar of the Flamborough was 
directed to cruise off the coast of North Carolina. “Whereas the 
Rt. Hono. the Lords Commiss. of the Admiralty has been informed 
that the coast of North Carolina has been much infested with priva- 
teers [Spanish] to the great determent of the inhabitants of that 
province, you,are to use your utmost endeavours to take or otherwise 
destroy them or any of the enemy’s ships as you may possible meet 
with in your cruise.’ °* Hamar was directed to inform the Governor 
of North Carolina ** of his cruising on that coast and told to keep to 
sea at all times possible.® Captain Newman (also spelled Newnham) 
of the sloop Spy was ordered to join the Loo and cruise with her until 
further orders.®® 

The execution of Utting’s plans was to be delayed, however, for the 
next day an “exstream hard gale of wind at ENE” struck the Loo 
as she lay at anchor off Charlestown Bar, obliging Utting to cut his 
“best bower cable’ ®*’ and go to sea “for fear of a hurricane.” For 
four days the Loo rode out the gale at sea, and when the storm was 
over Utting returned to his anchorage off Charleston, recovered his 
anchor and the next day (Saturday, October 25) “‘saild for Port 
Royall to refitt having received great damage in . . . masts and rig- 
ging.°> Upon examining the damage to the Loo, Utting and his offi- 
cers found the main yard sprung in three places and unserviceable. 
“The mainmast sprung in ye lower partners ® about 6 inches in tho 
not to bad but shall be able to fish *° him and make as serviceable as 
ever...” 7 Utting was mistaken, however, in his estimate of the 
damage, for closer examination revealed extensive damage to the mast 


61 Tbid. 

62 Tbid. 

63 ADM 1, vol. 60, No. 436. 

64 Tbid. 

65 Tbid. 

66 Tbid. 

67 The cable of the heaviest of the two anchors carried in the bow of a ship. 
The bower anchors were those used for anchoring under ordinary conditions of 
wind and sea. 

68 ADM 1, vol. 2625, No. 438. 

69 Planks fitted snugly around the base of a mast, a hatch, or a capstan cov- 
ering the opening in the decks. 

70 To splice a broken spar or mast by binding with splints and wedging firmly. 

71 ADM 1, vol. 2625, No. 438. 


NO. 2 LAST CRUISE OF H.M.S. “LOO’’—PETERSON 17 


below decks and the Loo was not to leave Port Royal until December 
30, when she began her last cruise. 

While lying at Port Royal Utting continued active direction of the 
vessels under his command from the Loo. On November 18 he issued 
two orders to Captain Newman of the sloop Spy. The first directed 
Newman to watch for a vessel expected from Havana with prisoners 
of war which were being exchanged and, should he meet with her, 
“to take out thirty of the best seamen on board for the service of his 
Majesty’s ship Loo.” * 

The second order directed the captain of the Spy to keep close touch 
with Charleston to obtain intelligence of the expected declaration of 
war against France, and if hearing of such declaration to rendezvous 
with the Loo.” 


By Captain Ashby Utting, Commander 
of his Majesty’s Ship Loo 
Whereas we are in dayly expectations to hear of the Declaration of a French 
War. 

You are hereby required and directed when on your cruise to call as often of 
Charles Town, as you shall think convenient to get the best information you can. 
And when you find any certain intelligence of a French War being declared 
either by Publick or private letters. You are immediately to proceed and joyn 
me of the N W part of the Grand Bahama, and if not find me there to proceed 
of the Isaack Rocks and the Bominies and if not at either of those places to 
proceed of Cape Florida and the Martiars [Fla Keys] 74 where you are to 
cruise for me Ten days and if not find me in that time you are to proceed to 
Hinds Bluff one of the Burry Islands where you are to fill up your water and 
then proceed and cruise between the N W part of the Grand Bahamas and Cape 
Florida till you meet me or as long as your provisions will last; and then return 
to Port Royall where you are to compleat your water and provisions to three 
months and to proceed to sea, and cruise between that Port and Georgia till 
further orders. Given under my hand on board the said ship in Port Royall 
Harbour the 18th day of November 1743. 

Ashby Utting 
To Captain Newnham of his 
Majesties Sloop Spy. 





72 ADM 1, vol. 2625, No. 426. 

73 ADM 2, vol. 2625, No. 426. 

74 Ponce de Leon named the Florida Keys “the Martyrs” because, he said, 
from the sea they bore a resemblance to the early Christian martyrs tied up on 
lines of stakes for execution. From 3 miles or so at sea the larger trees on the 
Keys indeed appear in long rows, the low-lying land of the Keys being out of 
sight over the horizon. In an age of Christian fervor, when religious signifi- 
cance was seen in every natural phenomenon, such an analogy would be the 
expected thing. The name “Martyrs” appeared on charts as late as the early 
1800's. 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


On November 25 Captain Hamer of the Flamborough was also 
ordered to be on the lookout for the prisoner exchange ship expected 
from Havana, and to remove seamen for the Loo.*® 

As work proceeded on repairing the storm damage to the rigging 
of the Loo, the carpenter discovered that the mainmast had been 
sprung in several places and reported to the Captain. Utting, on 
November 27, appointed the first and second lieutenants, the master,” 
the carpenter and the carpenter’s mate to “take a strict and careful 
survey” of the mast and report its “exact condition” to him.’” The 
survey board acted immediately and reported the same day that the 
mast had been severely sprung below decks and in their opinion was 
unfit for service.”® 

For over a month the crew and officers turned to getting a main- 
mast cut and rigged, during which operations Warren Bolitha, the 
First Lieutenant of the Loo, broke three ribs, and on December 29, 
the day before the Loo sailed, he requested the captain to “let him go 
home in order to get cured.” *° 

While Utting was struggling to refit the Loo’s damaged rigging, a 
letter arrived on December 14 from Capt. Charles Hardy of the Rye 
announcing that she, too, had sprung her mainmast. Utting immedi- 
ately ordered Hardy to replace the mast “as soon as possible” and to 
return to his station off the Carolina coast. As a precaution against 
confusion on the part of the commanding officer of any ship that might 
relieve the Rye while the Loo was away on its expected cruise toward 
Cuba, Utting instructed Hardy to pass on his orders to his relief.*° 

Meanwhile the Flamborough had been at sea and had fallen in with 
the ship that was bringing freed prisoners exchanged in Havana.** 
On December 15 Utting ordered Captain Hamer to search out the 
Spy, which was then cruising off Charlestown Bar, and transfer to 
her, for transportation to the Loo at Port Royal, 30 of the seamen 
whom he had impressed, and then to proceed to cruise off Georgia, 
sending a boat to the Governor of that colony “for any intelligence 
he may have of any of the enemy’s ships, or vessells being on that 
coast.” 8? 


75 ADM 1, vol. 2625, No. 432. 

78 Warships of this period had an officer in charge of the active sailing of the 
ship known as the Master. 

77 ADM 1, vol. 2625, No. 433. 

78 Thid. 

79 ADM ft, vol. 2625, No. 431. 

80 ADM 1, vol. 2625, No. 430. 

81 Among them, John Manley and Henry Spencer, who were to play a fateful 
part in the subsequent events. (ADM 1, vol. 2625, No. 446.) 

82 ADM 1, vol. 2625, No. 434. 


NO. 2 LAST CRUISE OF H.M.S. ‘‘LOO”—PETERSON 19 


At the same time he ordered the Spy to take aboard the 30 seamen 
from the Flamborough and then to cruise off Port Royal Bar and 
join the Loo when she came out.** 

On December 22, Utting ordered Captain Ward of the ship Tartar, 
which had arrived to relieve the Rye,** to cruise on the Carolina and 
Georgia coasts on the Rye’s old station.*® 

Finally, on December 30, work on the Loo having been completed 
and the winds and tide favorable, the ship crossed the bar at Port 
Royal and began her last cruise. In a final letter to the Admiralty 
before the ship weighed anchor, Utting explained the long delay occa- 
sioned by damage the Loo had received in the storm off Charleston 
October 16-20, which he had underestimated in his letter to the Ad- 
miralty dated November 12, 1743, at Port Royal Harbor. He re- 
ported that it had taken him more than a month to get a new mast 
cut, partially seasoned, and rigged ®* and took occasion to point out 
again to the Lords of the Admiralty the desirability of cutting several 
trees and seasoning them as a reserve to be used for the manufacture 
of masts or yards in the event of further damage to the ships under 
his command. 

At the same time Utting reported that he had relieved his first 
lieutenant, Mr. Bolitha, because of his injury, so that he could return 
home to England, and had promoted his second lieutenant and third 
lieutenant each one grade, then filling the vacancy left by the third 
lieutenant by the appointment of one William Lloyd whom he de- 
scribed as “a young gentm. well qualified for Preferement in his Maj’s. 
service.” 87 

After this last word from Utting, the Loo sailed to her station in 
the Florida Straits and began cruising against Spanish shipping. 

The morning of Saturday, February 4, 1744, found her cruising in 
the Straits off Havana. Around 8 o’clock in the morning a sail was 
sighted, and the Loo gave chase. As the stranger was neared, two 
seamen of the Loo, John Manley and Henry Spencer, who had been 
in the group of prisoners exchanged from Havana, informed Utting 
that they recognized the ship as the Billander Betty on which they had 
served. They told Captain Utting that while on a voyage in the 


88 ADM 1, vol. 2625, No. 427. 

84 Captain Newman of the Rye had been directed to convoy merchant ships to 
England from Charlestown in an order dated September 23, which was sent out 
by the Tartar. (ADM 2, vol. 60, p. 270.) 

85 ADM 1, vol. 2625, No. 429. 

86 ADM 1, vol. 2625, No. 424. 

87 ADM 1, vol. 2625, No. 424. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I3I1 


Betty (Capt. John Eades) from England to the Isle of May ** and 
South Carolina they had been captured by a Spanish vessel off the 
coast of South Carolina. The Spanish put aboard a prize crew and 
sent the Betty on to Havana, but the Spanish vessel herself was lost 
in returning to Havana. While prisoners in Havana, Manley and 
Spencer had heard that the Betty had been converted to a “snow” and 
was to make a voyage to Campeche. About noon, when the Loo came 
alongside her chase, Utting sent an officer to examine the stranger’s 
papers. Upon hearing that the master of the quarry could show only 
a common receipt, Utting decided to seize the ship for the proprietors 
of South Carolina and send her to Charleston. 

Before sending her off, however, Utting requested that an “Trish 
gentleman” on the snow be brought aboard the Loo for questioning. 
Before this gentleman left the snow, he was seen to throw a large oil- 
skin packet overboard. A boat from the Loo retrieved the packet and 
Utting discovered that it contained papers in French and Spanish. At 
this, he decided to take the prize in, with the Loo as escort. 

The examination of the prize had taken the entire afternoon, and 
when the Zoo set sail with her charge it was 6 p.m. and growing dark. 
The “Pan of Matanzas” (fig. 1), a flat-topped mountain behind 
Matanzas Bay on the coast of Cuba, bore south by east at a distance 
of 18 to 21 miles.*° Taking his fix on the mountain, Utting set sail 
and ordered a course northeast by north, the wind coming from the 
southeast. This course was kept until midnight, when Utting, be- 
lieving that he was clear of the Double Headed Shot Key in the west- 
ern end of the Salt Key Bank, instructed Randell, the first lieutenant 
and officer of the off-going watch, to alter the course to northeast and 
went below to his cabin to rest, having been continuously on deck since 
early morning.°° 

Shortly after, Randell was relieved of the watch by Robert Bishop, 
the master. Randell relayed these instructions to Bishop, reminding 
him to have the deep-sea lead line cast every half hour,®t and went 


88 “Maio” in the Cape Verde group occupied until the end of the eighteenth 
century by the English, who claimed a right to the island under the marriage 
treaty between Charles II and Catherine of Braganza of Portugal. The English 
occupation is recalled in the name “English Road,” which the port of Nossa 
Senhora de Luz is sometimes called. 

89 The bearing and the distance to the Pan of Matanzas were given by Lt. 
James Randell in his deposition to the court martial that tried Captain Utting. 
(ADM 1, vol. 5283.) 

90 Utting’s letter of February 15, 1744. (ADM 1, vol. 2625. (See Appendix 
B.)) 

91 Bishop’s deposition at the court martial. (ADM 1, vol. 5283.) 


NO, 2 LAST CRUISE OF H.M.S. ‘‘LOO’”—PETERSON 21 


below. Nothing to arouse the suspicion of Utting or Randell had been 
seen during the latter’s watch. The night was dark and cloudy, with 
visibility not over a quarter of a mile.*” 

At 12:30 a.m. and again at 1:00 the deep-sea lead line was cast 
according to orders, and no bottom was found at 300 feet. At about 
1:15 Bishop sent the lead-line crew to the side to clear the line for 
heaving and followed them to the gunwhale himself to see to this. To 
his great surprise he found the ship in “white water” and saw breakers 
ahead. He instantly “ordered the helm alee” and sent a message down 
to Captain Utting ** to call him on deck. As Utting rushed on deck 
he found the ship coming about into the wind and away from the reef 
on which the breakers were rolling. As the ship veered off the wind 
the head sails were caught across wind and the ship struck the reef 


Lh 


bast . Uw Pan de VMataneas 





Twat Matiigns 


Fic. 1.—The Pan of Matanzas from a vignette 
appearing on an English chart dated 1794. 


aft.°* At this the mainsail was set “in order to press her off,” and 
Utting ordered a boat out to sound around the ship.** The officers 
and men off watch and sleeping below were awakened by the shock 
of the ship striking the reef. John Vivian, the carpenter, rushed aft, 
whence the shock had come, and found the tiller broken off. He re- 
ported this to Utting just as another swell caught the ship and broke 
- off the rudder, at which she began shipping water in the hold. Utting 
ordered all pumps manned, and the water in the hold began to fall, 
but as the crew was getting out the boats “three or four severe seas” 
crushed the ship against the reef and she began sinking rapidly. 
When it became apparent that the ship could not be saved, Captain 
Utting ordered Mr. Bishop and Gunner Samuel Kirk to save as much 


92 Deposition of John Randolph, master’s mate, at the court martial of Cap- 
tain Utting. (ADM 1, vol. 5283.) 

93 Bishop’s deposition. (ADM 1, vol. 5283.) 

94 Utting’s letter of February 15, 1744. 

95 Bishop’s deposition. (ADM t. vol. 5283.) 

96 Utting’s letter of February 15, 1744. 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


of the bread and gunpowder as possible before the water rising in the 
hold ruined it. Bishop was able to save 20 bags of bread before the 
water forced him from the breadroom, and by the efforts of Gunner 
Kirk 6 barrels of gunpowder were saved. 

The prize, which had struck the reef just after the Loo, was rolling 
and pounding herself to pieces on the coast and, in order to save the 
men aboard her, Utting ordered her masts cut away and her guns and 


3 





—— 


HAMAS 


ABA 


Cb 


Fic. 2—Map of the Florida Straits, showing the course of the last cruise of 
the Loo. 1, Havana. 2, Pan of Matanzas. 3, Double Headed Shot Key. Dotted 
line, Captain Utting’s intended course. Solid line, actual course of the Loo. 


anchors thrown overboard. After this she lay more quietly in the swell 
and her men were saved. 

With the coming of daylight Utting and his officers saw with great 
surprise that they were ashore on a “small sandy Key about 13 
cables *? length long and 4 broad which lay on the edge of the Bank 
of the Martiers 3 leagues °* without them.” *° (See fig. 2.) 

At no time since the ship had struck had the pilots or officers 


97 A cable was 600 feet. 
98 A league was 3 nautical miles. 
89 Utting’s letter of February 15, 1744. 


NO. 2 LAST CRUISE OF H.M.S. “LOO’—PETERSON 23 


doubted that they were aground on Double Head Shot since, under 
normal circumstances, the course that they had steered could not have 
carried them to the Florida Keys. Utting sent Lieutenant Randell 
ashore to see if fresh water was to be had there, but he found none.*°° 
As full daylight came Utting landed all the men from the Loo and the 
prize with the exception of a few who were employed in cutting holes 
in the frigate’s deck to recover casks of water and such other supplies 
as could be saved. At about Io o’clock, to Utting’s great joy, a sloop 
was sighted offshore and a signal was made to her, but the sloop stood 
out to the northwestward.’ The captain immediately armed all the 
boats and with Lieutenant Randell and Mr. Bishop in command sent 
them in pursuit, instructing them to exert every effort to bring the 
sloop in, since it probably would be their only chance of succor. 

The desperate situation of the group was evident to all. Here were 
some 280 men stranded on a small sandy islet just off a hostile coast 
swarming with the savage Caloosa Indians who murdered Englishmen 
on sight.*°? 

To add to the insecurity was the evident fact that in a blow of any 
force the whole islet would be swept by waves. 

At night Utting posted watches, each consisting of 25 marines and 
25 sailors, around the island at the water’s edge as “centenells” to 
prevent a surprise night attack from the Caloosas “the Indians hav- 
ing numbers of canoes.” 1° 

The next morning (Monday) as daylight came, Utting and the men 
ashore were overjoyed to see the boats bringing in the sloop. As they 
came ashore Randell and Bishop reported that on the approach of the 
armed boats the Spanish crew had abandoned the sloop and were no 
doubt now headed for Havana in their boat. 

Meanwhile the men, frightened and confused, became “very rebel- 
lious and mutinous dividing into parties and growling amongst them- 
selves,’ *°* claiming that the officers no longer had authority over 
them, and clamored to leave the island immediately. Utting took no 
notice of them but, with the men who would work, continued efforts 
to recover water and other provisions from the wreck. 

All day Tuesday was spent in getting water casks from the Loo’s 
hold and in getting the sloop and boats ready for the escape. The 


100 Randell’s deposition. 

101 Tbid, 

102 Spaniards fared a little better, as the Caloosas knew they could be ran- 
somed. 

103 Utting’s letter of February 15, 1744. 

104 Tbid, 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


capacity and safety of the longboat were increased by adding planks 
to the gunwhales, giving her a higher freeboard and decreasing the 
chances of shipping water in a seaway while heavily loaded. Thus 
altered, the boat was able to carry 60 men.*” 

At about noon the next day, Wednesday, February 8, all the men 
were embarked—6o in the altered longboat, 10 in the yawl,'°* 184 “in 
the little Sloop not 30 tuns’’ *°7 and 20 in the captain’s barge. Utting 
sent the sloop, the longboat, and the yawl 3 or 4 miles offshore 
while he remained behind with the men detailed to the barge and laid 
most of the gunpowder which had been saved and some other fuel 
along the starboard gun deck of the Loo, the only deck remaining 
above water. By 2 o’clock all the preparations were completed and 
Utting fired the ship. As they rowed away the ship blazed to the top 
of her masts and blew up “in several places and was in flames fore 
and aft.” 1°° The burning ship was visible until sunset and while her 
hull was completely destroyed, Utting feared that the Spaniards would 
return and salvage her guns and anchors, since intelligence of the Loo’s 
end would be communicated to the Spanish in Havana by the escaped 
crew of the sloop. 

Utting’s plight was still grave, since the sloop was very much over- 
loaded and would have capsized in a blow. He placed First Lieutenant 
Randell and Third Lieutenant Lloyd in charge of the longboat, his 
Sailing Master Bishop was given command of the barge and “a mate” 
assigned to the yawl. Utting remained in command of the sloop. The 
motley fleet set a course for the Bahamas, the boats being ordered “in 
case of separation to make the best of their way over to the Bahama 
Bank for Providence.” *°° 

That night Utting carried a light to guide the boats, but they out- 
sailed the overloaded sloop and at midnight were lost from sight when 
Utting had to tack and stand to the northward after signaling his 
change of course with the light. At daybreak the boats were gone, and 
Utting, feeling that they were bound for Providence and being unable 
to set sail for an easterly course, set a course for South Carolina. In 
his report to the Admiralty he summed up the desperate situation in 
which he found himself with the overloaded sloop: “it blowing fresh 
and the sloop top heavy with men could not carry sail so obliged to 


105 Tbid. 

106 A heavy double-ended rowboat. 

107 Utting’s letter of February 15, 1744. 

108 Tbid. 

109 Thid. Providence had been settled in the seventeenth century. 


NO. 2 LAST CRUISE OF H.M.S. ‘“LOO’’—PETERSON 25 


bear away and take my fate through the Gulph of Florida **® for any 
port of Carolina even for St. Augustine (if I could fetch nowhere 
else) rather than all be drowned which Doe assure you had very 
little other prospect.” +14 

The fair weather continued and the overloaded sloop arrived in 
Port Royal harbor (pl. 17) on the night of February 13. Utting and 
the men were worn out from physical and mental strain, all realiz- 
ing that their escape from capture or drowning was just short of 
miraculous. 

Upon his arrival at Port Royal Utting began immediate steps to 
assemble evidence to protect himself in the court martial that he had 
to face for the loss of the Loo. His first step was to send one of his 
pilots, William Lyford, to the town of Beaufort 6 miles north of Port 
Royal to give a deposition before Robert Thorpe, justice of the peace. 
In the deposition Lyford stated that in his opinion the course the Loo 
had steered before she ran aground “was the best through the Gulph 
(and is generally allowed so to be) and was then of the opinion that 
such course would carry the said ship nearer the Bahama shore than 
the Florida; and this deponent further deposith and makes oath, that 
he is well acquainted with the Gulph of Florida having used it these 
thirty years past.” 12? 

Eight days later, on February 21, Utting was in Charleston start- 
ing proceedings to prove that the prize which he had taken was a legal 
one. John Manley and Henry Spencer, the two seamen who had 
recognized the prize as their former ship, appeared before James 
Grome, judge of the Court of Vice Admiralty of the Province of 
South Carolina, and swore under oath that the prize was the former 
Billander Betty, and that while on a voyage from England to the Isle 
of May and South Carolina, it had been captured off the coast of 
South Carolina on April 9, 1743— 
by a Spanish vessell bound from the Havannah to St. Augustine with about 
sixty or more soldiers on board, that the said vessell not being able to make St. 
Augustine return’d to the Havannah and in her passage was cast away, that the 
Billander so taken as aforesaid was carried to the Havannah and was there 
converted into a Snow and intended on a voyage to Campeachee but afterwards 
these Deponents hear’d that she was bound for the Mississippi. 

That these Deponents came to this province with the Flag of France and 
were press’d on Board his Majestys Ship the Loo under the command of Capt. 


Ashby Utting, that on a cruise in the said ship they met with a Snow which 
these Deponents very well knew to be the Billander Betty taken as aforesaid 


110 The Gulf Stream would carry him northward. 
111 Utting’s letter of February 15, 1744. 
112 Deposition accompanying Captain Utting’s letter of February 15, 1744. 


26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


by the Spaniards and converted into a Snow that the said Snow was taken by 
the Loo about a fourthnight ago, viz the 5th of this instant February and cast 
away with the said Man of War about nine leagues to the westward of Cape 
Florida.118 


On March 6, as the hearing proceeded, this deposition was intro- 
duced to the Court of Vice Admiralty as “Exhibit Marked I” and 
undoubtedly had a great influence in the outcome of the case in 
Utting’s favor. 

By March 12 the proceedings in the Vice Admiralty Court seem 
to have been concluded. Utting had only to wait for the return of 
his officers from the Bahamas, and then a warship for transportation 
to England if his court martial could not be held in America. 

The loss of the Loo had immediate repercussions in the colony of 
South Carolina, and on the same day the case in court was concluded, 
the Governor, James Glen, wrote a lengthy letter to the Lords of 
Trade in which he indicated the fears of invasion which had swept the 
southern frontier of the colony: 


After writing so lately by Captain Hardy,115 I had not so soon troubled your 
Lordships with another letter, but the loss of his Majestys Ship the Loo makes it 
my duty; this unlucky accident happened the fifth of February about one in the 
Morning, by her running on some rocks called the Martyres to the South West 
of Cape Florida. 

. my principal concern is to express to your Lordships how sensibly that 
loss affects this province. The long neglected Town of Beauford, upon the 
arrival of this Ship, and the assurances given that another would be sent out, 
began to revive, and many good houses were built, and many grants for Town 
Lotts were applyed for, so that I am persuaded that Town and the adjacent 
Country, would soon have been well settled, and consequently our Southern 
Frontier, where we are most vulnerable, would have been strengthened, but now 
I receive letters and petitions dayly from the best People in those parts, repre- 
senting their fears and the dangers to which they are exposed, and everything 
is at a stand, tho’ I have stationed one of our gallys (a very fine small vessel) 
there, I have likewise desired the Captains of the Man of War on this station, 
to keep a particular eye upon that Port, in their Cruizes along our coast.116 


118 Copies of papers relating to the proceedings of the Court of Vice Admiralty 
sent by Captain Utting to Thomas Corbett after his return to England (ADM 1, 
vol. 2625, No. 455.) 

114 Utting and his wife had been residents of South Carolina several years. 
The wait for transportation to England was probably not too burdensome to 
Utting. 

118 Of the Rye, which had sailed for England a short time before Utting 
reached Port Royal. 

116 Letter of James Glen dated March 12, 1743/44. Colonial Office original 
correspondence, CO5, vol. 370, pp. 141-142, Public Record Office, London. 


NO. 2 LAST CRUISE OF H.M.S. “LOO”—PETERSON 27 


He then went on to point out the suitability of Port Royal as a 
harbor and its strategic location in relation to the Florida Straits and 
the Spanish trade routes : 

And as most of the trade and treasure of France and Spain must come through 
the Gulf of Florida, where can it be so properly waited for as here, where a few 


great ships stationed, to cruize betwixt this and Cape Florida, a very easy navi- 
gation, must become masters of everything.117 


The Governor reported that Captain Dansant, captain of the Loo’s 
prize, would be sent off without being permitted to see the fortifica- 
tions of Charleston, and “The forty-four marines belonging to the 
Loo,” he stated, were being “lodged at the expense of this government 
and shall be well taken care of.” 11* The sailors were no doubt taken 
into the other ships present on the station, the Governor not mention- 
ing them. 

Governor Glen spoke a good word for Captain Utting with the 
Lords of Trade, describing him as ‘a Gentleman who by a long resi- 
dence in this Province, has established a character amongst all ranks 
of people here, for strict honors and veracity, as well as for his care, 
diligence, and knowledge, as an officer.” 7° 

Early in April Utting and his officers, who had arrived from the 
Bahamas, sailed for England and arrived there late on the night of 
May 24. The next day Utting reported his arrival to the Admiralty 
and requested an early court martial for the loss of the Loo. (See 
fig. 3.) 

Six days later, May 31, the court of 12 captains sat on board the 
ship-of-the-line Sandwich with Vice Admiral James Steuart presiding. 
After a consideration of the depositions and testimony of Utting and 
his officers the Court was “unanimously of the opinion that Cap". Utting 
and his several officers did in no wise contribute to her going ashore, 
but that it was owing to some unknown accident, it appearing to the 
Court, that the course the ship steered was a good one, and must have 
carried her thro the Gulph of Florida, with all safety had not some 
unusual current rendered the said course ineffectual.” 1*° 

On June 6 the Lords of the Admiralty ordered the Navy Board to 
procure funds from the Treasury and pay the officers and men of the 
Loo through the day she was lost. On August 10, 1744, the officers 
and men gathered on Broad Street in London and were paid the 1,510 


117 Thid, 

118 Thid, 

119 Tbid. 

120 Report of Court Martial dated June 1, 1744, Admiralty In-Letters, ADM 1, 
vol. 5283. 


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Fic. 3.— —Letter of Capt. Ashby Utting to the Secretary of the Admira Utting to athe Secretary of the Acti ed 
ing his arrival in England May 24, 1744, stand trial by court martial for the 
loss of the Loo. (Photostat feos oe Public R ecord Office, London.) 


28 










ae 


RK 


® 


NO. 2 LAST CRUISE OF H.M.S. “LOO’—PETERSON 29 


pounds 4 shillings 11 pence due them after deductions of 1,121 pounds 
4 shillings 11 pence for tobacco, clothing, hospital fund, pay advances, 
etc. James Compton, Captain, Royal Navy, a Navy Commissioner, 
kept an eye on the proceedings while navy clerks Stephen Mercer and 
Philip Stephens and treasury clerks John Wilson and Thomas Vaughn 
checked the pay list and disbursed the money.1** 

After his acquittal Utting attended the Secretary of the Admiralty 
frequently while waiting for an answer to his request for another com- 
mand. On June 12 he discussed with the secretary the possibility of 
getting command of the Mary Galley and the next day advised Cor- 
bett that several of the men and petty officers of the Loo desired to 
ship with him on his next cruise. He also reminded Corbett of the 
desirability of his return to the Carolina station since his wife was 
there:*#" 

On July 7, 1744, a commission was issued giving Utting command 
of the Gosport.1** While fitting out his new ship, Utting continued to 
hope that he would be able to get orders to return to the Carolinas. 
No one knew better than he the danger of invasion to which the 
colony was exposed through the loss of the Loo, and he was anxious 
for the safety of Mrs. Utting, who was at Port Royal. His fears were 
multiplied when, on August Io, he received a letter from his wife, 
dated July 5, in which she reported that the settlements south of 
Charleston had been evacuated because of fear of an invasion and that 
she was a refugee in the provincial capital. Utting’s patience reached 
the breaking point as he pleaded for orders to America: “This is a 
very shocking affair both to her and me and beg for God’s sake you'll 
be so good to use your interest with Lord Winchelsea *** to git me to 
some part of America.” 1”° 

The exigencies of war, however, outweighed the personal problems 
of Utting, and he was ordered to the Baltic to convoy a fleet of 
merchantmen to Elsinore, Denmark, and Bergen, Norway. 

On October 13 Utting was back in England with the convoy from 
Bergen. The next month he took a convoy to Ostend, leaving on the 
15th and returning to England on the 24th, assuming command of the 
Aldborough sometime between his return and November 29 under a 
commission dated November 7.1*° Utting’s wish to return to South 


121 [00's pay list dated Aug. 10, 1744, Admiralty Ships Pay Books, Treasurers 
Series I, ADM 33, No. 352. 

122 Admiralty In-Letters, ADM 1, vol. 2267. 

123 AD 6/16, Commission and Warrant Book, 1743-1745, p. 335. 

124 First Lord of the Admiralty. 

125 ADM 1, vol. 2625, No. 477. 

126 AD 6/16, Commission and Warrant Book, 1743-1745, p. 380. 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


Carolina was realized shortly after, when he was ordered to escort a 
convoy to America and assume his former command as senior officer 
present at Charleston. On March 26, 1745, he arrived at his old sta- 
tion and began the direction of naval operations off the Carolinas. 
The threat from the Spanish was still real, and naval patrols were 
necessary to prevent surprise attacks on the coastal settlements. 

Sickness plagued the Alborough and Utting was unable to keep the 
sea as he should have. Enemy privateers arrived off the coasts of 
Carolina and Georgia, and the captain was at his wit’s end to protect 
the coastal settlements from Fredrica, in Georgia, to Charleston with 
his little squadron. As a result some discontented merchants in 
Charleston complained to the Carolina proprietors that Utting was not 
doing his duty. These complaints seem to have been unjustified, for 
the Governor, Council, and several principal merchants refused to sign 
them. Thus misfortune harassed Utting during his last cruise until 
early in January 1746, when he died on board the Alborough in Rebel- 
lion Road, Charleston, just after returning from a patrol off the coast. 

On April 18, 1744, while Utting was on the high seas returning to 
face a court martial for the loss of the Loo, the Lords of the Ad- 
miralty had ordered the Navy Board to “cause a new ship of forty-four 
guns to be built in the room of the Looe lately lost near the Gulph 
of Florida.” 1**7 Today the name “Looe” is perpetuated by the sub- 
merged reef lying off the central Florida Keys, visited by occasional 
fishermen who must wonder at the strange name it bears, never 
dreaming that the quiet little reef was once the scene of as dramatic 
a story of shipwreck and rescue as can be found in the annals of the 
English colonies in America. 


127 Admiralty Out-Letters, ADM 2, vol. 205, p. 344. 


APPENDIX A 
EXTRACTS FROM (THE NAVY, LIST ‘RELATING TO THE. “LOO” 


Period 
April 1, 1707—May 31, 1707 
June 1, 1707—June 30, 1707 
July 1, 1707-September 30, 1707 
October 1, 1707—October 31, 1707 
November 1, 1707—November 30, 1707 


December 1, 1707-January 31, 1708 


February 1, 1708—March 31, 1708 
April 1, 1708—-April 30, 1708 

May 1, 1708-July 31, 1708 

August 1, 1708-February 28, 1709 
March 1, 17090-July 31, 1700 
August I, 1709—September 30, 1709 


October 1, 1709—October 31, 1709 
November 1, 1709—-November 30, 1709 


December 1, 1709—December 31, 1709 


January I, 1710-January 31, 1710 
February 1, 1710-February 30, 1710 


March 1, 1710—March 31, 1710 
April 1, 1710-April 30, 1710 
May I, 1710-May 31, 1710 


June I, 1710—June 30, 1710 


July 1, 1710-July 31, 1710 


August 1, 1710-September 30, 1710 
October 1, 1712—October 31, 1712 
November 1, 1710-December 31, 1710 
January 1, 1711—January 31, 1711 


Duty 


“Longreach taking in Guns” 

“Going to Archangell” 

“Gon to Archangell” 

“Arch-Angell” 

“Returned with the Russia ships to 
Grimsby” 

“Sheerness 
land)” 

“Downes for Newfoundland” 

“for Newfoundland” 

“Gon to Newfoundland” 

“at Newfoundland” 

“coming convoy from Newfoundland” 

“Coming convoy from Lisbon but last 
from Newfoundland” 

“Sheerness—refitting” 

“Sailed to Join the Tilbury Etc. at 
Goree and when she returns to join 
the Gosport and Strombolo between 
Dover and Beachy” 

“Downes ordered to cruise between 
Dover and Beachy head” 1 

“.. . Dover and Beachy head” 

“Cruizing between Dover and Beachy 
head” 

“Holland—ordered to cruize between 
Dover and Beachy Head” 

“Holland, to come to the Downs” 

“Dunkirke Squadron” “Sailed to 
Cruize on ye French Coast between 
Cape Barfleur and Harve de Grace” 

“Dunkirke Squadron” “Margate Roads 
ord. to Holland with the yachts and 
bring the Queen’s wine to the Nore” 

“Dunkirke Squadron” “Holland ord. to 
bring the Queen’s wine to the Nore” 

“Gone Convoy to the East Country” 

“At the Nore” 

“Sheerness Refitting” 

“Downes ordered to Scarboro to bring 
a ship to the Nore” 


(fitting for Newfound- 


1 Navy List, January 1, 1707—-December 31, 1709, ADM 8/10. 


31 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS 


Period 
February 1, 1711-February 28, 1711 
March 1, 1711—March 31, 1711 
April 1, 1711—April 30, 1711 
May 1, 1711-May 31, 1711 
June I, 171I-June 30, 1711 
July 1, 1711-September 30, 1711 
October 1, 1711—October 31, 1711 
November 1, 1711-November 30, 1711 
December 1, 1711-December 31, 1711 
January 1, 1712-March 4, 1712 
March 5, 1712-June 30, 1712 
July 1, 1712-July 31, 1712 
August I, 1712—-September 30, 1712 


October 1, 1712-October 31, 1712 
November 1, 1712-November 30, 1712 


December 1, 1712—December 31, 1712 
January I, 1713-January 27, 1713 
February 1, 1713-February 28, 1713 
March 1, 1713—March 31, 1713 
April 1, 1713—May 31, 1713 


June 1, 1713-June 30, 1713 


July 1, 1713-July 31, 1713 
August 1, 1713-September 30, 1713 


October 1, 1713-October 31, 1713 


November 1, 1713—December 31, 1713 


VOL. 131 


Duty 


“Spithead, ordered to the Downes” 

“In Holland to return to the Downes” 

“Tn Holland to bring the Queen’s wine 
to the Nore” 

“Sheerness, ordered to the Downes” 

“Ously Bay, for Russia” 

“Gone Convoy to Russia” 

“Coming from Russia. the Ist Convoy” 

“at the Nore” 

“Sheerness refitting” 2 

(No record found) 

“Convoy between Folmouth and Spit- 
head” 

“Cruizing for the homeward bound 
Virginia ships” 

“Cruising off the Orcades for the 
homeward bound Virginia ships” 

“Nore, ordered to the Downes” 

“Downs, ordered to Bayonne with the 
Transports” 

“Gone to Bayonne with transports for 
soldiers” 

“Gone to Bayonne for some prisoners 
of war” 

“Coming from Kinsale to Plyo. to fit 
and repair to Spithead” 

“Kinsale, ordered to Plymo., to fit and 
repair to Spithead” 

“Plymouth refitting and ordered to 
Spithead” 

“Spithead, ord." to Guernsey and Jer- 
sey for some disbanded men of Mor- 
daunts Regiment.” 

“Gone to Gurnsey and Jersey for some 
disbanded soldiers” 

“Spithead cruizing between Start and 
the Isle of Wight” 

“At Portsmouth to fit and cruiz be- 
tween ye Start and the Isle of 
Wight” 

“To intercept the traders cruizing be- 
tween Start and the Isle of Wight” ® 


2 Tbid., January 1, 1710—-December 31, 1711, ADM 8/11. 

8 Tbid., March 5, 1712—December 31, 1713, ADM 8/12. Constant patrol of these 
waters was maintained in an effort to prevent the smuggling of wool from 
Britain to the Continent, the export of raw wool being absolutely forbidden at 


this time. 


NO. 2 


Period 
January 1, 1714—March 31, 1714 


April 1, 1714-April 30, 1714 
May I, 1714-June 30, 1714 
July 1, 1714-September 30, 1714 


October 1, 1714—October 31, 1714 
November 1, 1714—December 31, 1714 
January I, 1715-January 31, 1715 
February 1, 1715-March 14, 1717 
April 1, 1717—October 31, 1717 


November 1, 1717—December 31, 1717 


January 1, 1718-March 25, 1718 
March 26, 1718—March 31, 1719 


April 1, 1719—April 30, 1719 


May 1, 1719-December 31, 1720 
January 1, 1721-April 30, 1721 
May 1, 1721-January 31, 1722 
February 1, 1722—-May 31, 1722 
June 1, 1722-August 31, 1722 
September 1, 1722-December 31, 1725 
January 1, 1726—-January 10, 1728 
January 11, 1728-March 31, 1728 
April 1, 1728-April 30, 1728 

May 1, 1728-July 31, 1728 

August 1, 1728-December 31, 1728 
January 1, 17290-January 31, 1729 
February 1, 1729-April 1, 1729 
April 2, 1729—-May 31, 1729 


June 1, 1729-July 31, 1729 

August 1, 1729-August 31, 1729 
September 1, 1729-September 30, 1729 
October 1, 1729-December 31, 1729 
January 1, 1730-March 31, 1730 
April 1, 1730-April 30, 1730 


LAST CRUISE OF H.M.S. “LOO’’—PETERSON 33 


Duty 


“To intercept the traders cruizing be- 
tween the Start and the Isle of 
Wight” 

“At Portsmouth fitting and then re- 
turns to her station” 

“In the Downes going to Port Mahon 
with a storeship.” 

“Gone to Port Mahon with a store- 
ship” 

“Coming from Port Mahon” 

“At Shearness refitting” 4 

“ordered to be laid up and paid off” 

(Not in service) 

“Baltick Squadron” 

“Hosp. Ship Looe” 

“To be paid off and laid up . . . Dep- 
ford” 

(Laid up) 

“Hosp. Ship Looe” 

“Mediterranean Squadron” 

“Looe .. 125 men 30 guns” (refitted 
as warship?) “Mediterranean Squad- 
ron” 

“Mediterranean Squadron” 

“Port Mahon” 5 

“In the Mediterranean” 

“Ordered home from Mediterranean” 

(Lists missing ) 

(Not in lists, laid up) & 

(Laid up) 

“At Woolwich” 

“At Longreach” 

“Nore” 

“Downes to examine ships” 7 

“Woolwich, not sheathed” 

(Laid up) 

“Woolwich fitting for Channel Serv- 
ice” 

“Downes” 

“Downes—Channel” 

“At Diep ordered to Spithead” 

“At Spithead” 

“Portsmouth Harbour” 

“Portsmouth Harbour, refitting for 
Channel service” 


4Ibid., January 1, 1714—December 1, 1714, ADM 8/13. 
5 [bid., January 1, 1715-April 30, 1721, ADM 8/14. 

6 Tbid., May 1, 1721—-December 31, 1725, ADM 8/15. 

7 Ibid., January 1, 1726-December 31, 1728, ADM 8/16. 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS 


Period 
May 1, 1730—May 31, 1730 
June 1, 1730—June 30, 1730 


July 1, 1730-July 31, 1730 
August 1, 1730-October 31, 1730 
November 1, 1730-January 31, 1731 


February 1, 1731-May 31, 1731 
June 1, 1731-July 31, 1731 


August 1, 1731-August 31, 1731 
September 1, 1731-November 30, 1731 


December 1, 1731—December 31, 1731 


January I, 1732—January 31, 1732 
February 1, 1732-February 28, 1732 
March 1, 1732-March 31, 1732 
April 1, 1732—-April 30, 1732 


May I, 1732-May 31, 1732 

June 1, 1732-December 31, 1734 
January 1, 1735-May 4, 1735 
May 5, 1735-June 30, 1735 


July 1, 1735-July 31, 1735 

August 1, 1735-August 31, 1735 
September 1, 1735-October 31, 1735 
November 1, 1735—April 30, 1737 
May 1, 1737—May 31, 1737 

June 1, 1737—December 31, 1739 
January 1, 1740-December 31, 1741 
January 1, 1742—January 3, 1742 
January 4, 1742-April 30, 1742 


May I, 1742-May 31, 1742 


June I, 1742-June 30, 1742 


VOL. 131 


Duty 


“Spithead—Channel” 

“Gone with the money and clerks to 
Plymouth” 

“Ordered to fitt for Gibraltar, Spit- 
head” 

“Gone to Gibraltar as convoy to the 
transports” 

“To remain in the Mediterranean” 

“Cruizing on the Coast of Barbary” 

“Cruizing on the Coast of Sallee” 

“Ordered Home” “From the Coast of 
Sallee” 

“Portsmouth 
Service” 

“Stationed between the Start and the 
Isle of Wight” 

“Stationed between the Start and the 
Isle of Wight” 

“Ordered to the Downes” & 

“Start and Isle of Wight” 

“Portsmouth, ordered to Plymouth” 

“At Spithead, Channel Service” 

“Gone to Plymouth with money and 
clerks” 

“Nore” 

(Not listed, laid up) ® 

(Not listed, laid up) 

“Sheerness, fitting for the Channel” 
(Again as a hospital ship) 

“At the Nore” 

“At Spithead” 

“Gone to Lisbon with Sir John Norris” 

“At Lisbon” 

“To be paid off” 

(Not listed, laid up) 1° 

(Not listed, laid up) 14 


refitting for Channel 


(Laid up) 
“Loo, 44 guns... Earl of Northesk 
Commander, Depford fitting for 


Channel Service” 

“At the Nore to convoy transports 
from the Downes to Spithead” 

“At Portsmouth refitting for channel 
service” 


8 Tbid., January 1, 1729-December 31, 1731, ADM 8/17. 

9 Ibid., January 1, 1732—December 31, 1734, ADM 8/18. 

10 Tbhid., January 1, 1735-December 31, 1739, ADM 8/109 and 8/20. 
11 Tbid., January 1, 1740—-December 31, 1741, ADM 8/21. 


NO. 2 
July 1, 1742-August 31, 1742 
September 1, 1742-September 30, 1742 
October 1, 1742-October 31, 1742 
November 1, 1742—November 30, 1742 
December 1, 1742-December 31, 1742 
January 1, 1743-March 31, 1743 
April 1, 1743-April 30, 1743 


May 1, 1743-May 31, 1743 


June 1, 1743-June 30, 1743 


July 1, 1743-July 31, 1743 


August I, 1743-August 31, 1743 
September 1, 1743-April 30, 1744 
May 1, 1744- 


LAST CRUISE OF H.M.S. ‘““LOO”—PETERSON 35 


“Cruizing 50 leagues off Capte Finis- 
terre” 

“Plymouth refitting for Channel Sery- 
ice” 

“In St. George’s Channel for 3 weeks” 

“Plymouth” 

“Spithead to clean at Portsmouth” 

“Portsmouth to cruize from 30 to 50 
Igs. W.S.W. of Cape Clear for 6 
weeks” 

“Plymouth, refitting for channel sery- 
ice” 

“To cruize between Bibao and St. Jean 
de Luz, to intercept some Caracca 
ships expected at St. Sebastian. . . 
to cruize 6 weeks on the Station” 

“Cruizing between Bilbao and St. Jean 
de Luz to intercept some Caracca 
ships expected at St. Sebastian . . .” 

“Portsmouth refitting for North 
America” 

“To attend on So. Carolina” 

“South Carolina” 

(No entry.12, The Loo had been lost 
February 5, 1744. Word apparently 
did not reach the Admiralty clerk 
keeping the navy lists until sometime 
in April. Entries of the location of 
ships were made on the first day of 
the month. ) 


12 Tbid., January 1, 1742-May 1, 1744, ADM 8/22, 8/23, and 8/24. 


APPENDIX B 


LETTER FROM CAPTAIN ASHBY UTTING TO THE ADMIRALTY 
REPORTING THE LOSS OF THE “LOO” 


Port Royall 
15th February, 1744. 


I am extremely sorry this should be the messenger of such disagreeable news 
as the loss of H.M.S. Loo. 

Will you please acquaint their Lordships that on the 4th day of February I 
was cruising on the station 8 leagues from the Cape of Florida when about 8 in 
the morning I saw a sail which I gave chase to and about noon spoke with her, 
she being an English “Snow”?! from Havannah and Missippy, but sailed by 
Frenchmen and two Spaniards, one that had been lately taken from the English 
and carried into Havannah. I having two men on board which was taken in 
her and the master having no copy of the condemnation and nothing to show for 
the sale but a common receipt. I seized her for the proprietors and was designed 
to send her into Charlestown but at the same time an Irish gentleman, a mer- 
chant that I had sent for on board, heaved a large packet overboard, which my 
boat took up and when opened found it full of French and Spanish papers, I 
then determined to see her in myself and also took her in tow. By the time I 
made sail it was 6 in the evening at which time the Pan of Mattances 2 bore 
S b E, the wind being SE. I steered NE b N till 12 at night by which time I 
was well assured I was got to the northward of the double Head Shott,? then 
hauled up NE. Till this time I was on deck myself and when thought I was 
passed all danger went and sat down in the cabin (as Doe assure you I did not 
go to bed one night in six of the time I was cruising here). 

At a } past one in the morning, the officer of the watch sent down to let me 
know he was in the middle of brakers and must Doe him the justice to say he 
behaved like an exceedingly good officer for before I was got upon deck which 
could not be ten moments, he had put the helm a Lee and the ship was at stays,* 
just as we hauled the main top sail the ship struck abaft but she pay’d off so 
far as to haul the head sails,5 when the Captain ® came and told me the tiller 


1A brig having a small trysail mast set astern of the mainmast. The trysail 
was a fore and aft sail with a gaff and, in some cases, a boom. 

2 A high, flat-topped hill lying inland from Matansas Bay on the northern coast 
of Cuba, a point on which mariners take bearings in setting a course up the 
Florida Straits (see fig. 1). 

8 A group of keys lying in the eastern end of the Salt Key Bank which ex- 
tends to the center of the southern end of the Florida Channel (New Bahama 
Channel). 

4 A vessel is said to be “ at stays” when heading into the wind in tacking. 

5 Swung off from the wind so far that the head sails were caught across the 
wind pushing the bow of the ship around toward the reef. 

6 The sailing master. 


36 


NO. 2 LAST CRUISE OF H.M.S. “LOO’’—PETERSON 37 


was broke short off the ship, continued striking, I ordered all the boats out as far 
as possible. Immediately after he came and told me the rudder was gone and 
that she made some water in the hold but not much, we set all the pumps to 
work as you must believe on this occasion, and rather gained on her. By this 
time we was getting the long boat out when there came three or four severe 
seas and bulged? her immediately and had 5 foot water in the hold; I ordered 
the master and gunner to come and save what bread and powder they could 
before the water was over all, which they did and saved 20 bags of bread and 
6 barrels of powder ® which was all we could save. 

By this time the “Snow” which shared the same fate, was on her broad side, 
the ship striking much and tareing all to pieces, and having no prospect of 
getting her off, ordered the masts to be cut away and all the upper deck guns 
and anchors to be thrown overboard, that she might lay quiet and by that means 
save the men which by good fortune she did, though all this time thought I was 
got on the double head Shott Bank when at daylight to my great surprise we 
was getting on a small sandy key about 1./1/2 cables® length long and ./1/2 
broad which lay on the edge of the Bank of the Martiers 3 leagues!® without 
them and lies from Cape Florida WSW 7 or 8 leagues is quite steep too, we hav- 
ing no ground at 50 fathom right up and down not 10 minutes before the ship 
was ashore and is the only dangerous place on the Florida shore and Doe assure 
you that from the day I got on that station, I always had the Drapsy 1! Line 
going every 1/2 hour from 6 at night till daylight in the morn, the only reason 
I can give for finding myself on the Florida shore when I expected I was on the 
double Shott Bank which lies from each other SE b E and NN b N at least 16 
leagues, must be occasioned by a very extraordinary and very uncommon new 
current; as soon as was daylight I landed all the men (but those that was em- 
ployed to scuttle the decks12 and get what water and what provisions we could, 
but could get but 2 butts out the whole day); at 10 o’clock this morning being 
Sunday we saw a small sloop when I immediately man’d and armed all the 
boats and sent them with orders to board her at all events and bring her here 
as she would be the means of carrying us off this dismal place, which I plainly 
saw that any common sea beat all over it and would certainly wash us all off, 
it being so low and dare not venture upon the main for the Indians which on 
this part of Florida are savages and innumerable, the next morning being Mon- 
day the boats to our great joy brought the sloop to us, the Spaniards having 
_all deserted her, she being about 25 or 30 tuns (at most) this day was employed 
in getting what provisions and water we could out of the ship with what men 
I could get to work which was but a few, though it was for all their good but 
all frightened and wanted to be gone for fear of the Indians and was very 
rebellious and mutinous dividing into parties and growling amongst themselves 


7 “Bilged”—stove in her planks at or below the waterline of the ship. 

8 A ship of 44 guns on foreign service normally carried 163 barrels of gun- 
powder in 1781 (Montaine, Will, The Practical Sea Gunners Companion, p. 73, 
London, 1781). 

® The cable was 200 yards or one-tenth of a nautical mile. 

10 The English and American marine league is equal to 3 nautical miles. 

11 Utting was speaking of the dipsey line, which is the deep-sea lead line. 

12 To cut openings in the decks. 


38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I3I1 


that they was all on a footing then; and they was as good as anybody and that 
everything was free. I thought it was best not to take any notice but prepared 
for our going as soon as possible. Here I found the service of the Marines 
whereof which was under command and did their duty extremely well and 
Centenells being obliged to mount 25 Marines and 25 seamen every night, though 
the place so small the Indians having numbers of canoes. 

Tuesday was employed getting what water we could out of the ship and put- 
ting the boats and sloop in order. Raised the long boat a strack 13 which enabled 
her to carry 60 men. On Wednesday being the 8th about noon I embarked all 
the men (which with the “Snow” included, was 274) viz 60 in the long boat, 
20 in the barge,!4 ro in the yaul!® and 184 in the little sloop not 30 tuns; sent 
the sloop, long boat and yaul to 3 or 4 miles from the shore. After they were 
gone I went on board the wreck with what barrels of powder we had saved 
(except a little we took in each boat) and laid in proper places on the Star- 
board side the gun deck, that side being out the water by her heelding off to 
Port and proper Fewell!® in every place we could when all being laid, about 2 
in the afternoon, I set her on fire and rowed off to the boats we kept in sight 
of her till sunset and she having blown up in several places and was in flames 
fore and aft but am much afraid the guns and anchors will fall into the hands 
of the Spaniards as they have frequent correspondence and trade with the In- 
dians, and it is my opinion the people which left the sloop we took, were over 
to Cuba in a launch directly to give them intelligence. This is the fatal end of 
H.M.S. Loo exactly as it happened. I sent in the long boat Mr. Randall and 
Mr. Lloyd, my first and third Lieuts., the Master in the barge and a mate in the 
yaul, myself and 2 Sevts. being in the sloop which when the hold was as full 
of men as could possibly be stowed, the deck was the same and much in danger 
of oversetting if any wind. I ordered them to follow me and in case of separa- 
tion to make the best of their way over to the Bahama Bank for Providence. 

I carried a light for them all night but as they all outsailed me they kept so 
far ahead that I lost sight of them by 12 at night when I was obliged to tack 
and stand to the northward, which did after making the proper signal but at 
daylight could see nothing of them, and it blowing fresh and the sloop top heavy 
with men could not carry sail, so obliged to bear away and take my fate through 
the Gulph of Florida for any part of Carolina even for St. Augustine (if I 
could fetch nowhere else) rather than all be drownded which Doe assure you 
had very little other prospect but thank God met with exceeding good weather 
and arrived at Port Royall on the 13th February at night and don’t in the least 
doubt but the boats are got safe to Providence long before this. 

I had two of the best pilots on board for the Gulph of Florida in all America 
who insist on it there could not be a better course steered and I have been 
numbers of times through the Gulph and am better acquainted with that and 
the coast of Carolina than any part of the world and had I had the honour to 
have command 20 sail and steering through for all our lives, should have steered 
the same course or rather more northerly, which as I observed before I cannot 





13 The addition of planks to the gunwhales of the boat increasing her capacity. 
14 The commanding officers boat. 

15 A heavy double-ended row boat. 

16 Fuel, combustibles. 


NO. 2 LAST CRUISE OF H.M.S. “LOO’’—PETERSON 39 


account for but by some uncommon and very extraordinary current; as fast as 
I can get my officers together shall send home their depositions. Some of them 
seem inclinable to go to the West Indies and some to the Merchant Service and 
some home. 

I have sent home the Mate of the Watch with the Log Book and my Second 
Lieut. was on board the Snow when cast away but I should have been very 
happy to have found Captain Hardy 17 not sailed that I might have come home 
directly but as I can’t be now and there being a great many chances against my 
coming home in a merchant ship, without being carried to Spain,18 has deter- 
mined me to stay here till some opportunity offers to come home in a Man-of- 
War or if I could possibly be indulged with a court martial in America, I shall 
think it the greatest favour and if found by the Court, which I hope I shall, that 
I have done my duty as an officer on this unforseen unhappy affair, to me as 
well as to his Majesty’s Service, beg their Lordships will be pleased to give me 
leave to rely on their goodness for my being employed again on His Majesty’s 
service. I have enclosed the deposition of Mr. Wm. Lyford, one of my pilots 
who has sailed the Gulph of Florida for many years and beg their Lordships 
will be pleased to let somebody enquire of General Oglethorpe for his corretor.19 

I am your most humble servant, 
Ashby Utting. 
Port Royal 
15th February 1743/4. 


17 Of the Rye, which ship had been ordered to England as convoy to merchant 
ships going from the Carolina colonies. 

18 The risk of capture of unprotected merchant ships was very great. 

19 Recommendation of Lyford. 


APPENDIX C 
MEMBERS OF THE CREW OF THE “LOO” ON HER LAST CRUISE 


The following is a list of names of all the men and officers appearing 
on the paybook of the Loo on her last cruise. Only those marked with 
an asterisk were on the ship when she was wrecked. 

In those days ships’ crews and officers were paid only at the end 
of a commission period, or at the end of a cruise. Men or officers who 
were transferred during a cruise were given a ticket by the purser 
showing the pay that was due them. These tickets were supposed to 
be held until the payday of the ship was announced in the newspapers, 
when they were presented at the designated pay office and the men 
received their pay. In actual practice, the interval of time between 
the issuance of the ticket and the actual payday was so great that many 
sailors suffered actual want, and to obtain funds, sold their tickets at 
enormous discounts to speculators. 


*Adam, Ervin 
*Adeane, W. 
Aiken, James 
Allen, Thomas 
Allman, John 
Anderson, James 
Angelo, Rogero 
*Arthur, James 
*Atkinson, Thomas 
Bagster, John 
*Baker, William 
Balderson, William 


Ball, William 
Balls, Benjamin 
Banke, John 
Baptista, John 


Barnes, William 
Barsey, Richard 


Barsey, Thomas 
*Bartlett, Joseph 
*Basham, Charles 


Bates, John 
40 


Quartermaster 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Landsman 

Quarters Servant 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Master 

Able Bodied Seaman 

Able Bodied Seaman 
and Quartermaster’s 
Mate 

Surgeon’s Mate 

Able Bodied Seaman 

Quarter Gunner 

Able Bodied Seaman 


Able Bodied Seaman 
Landsman 


2d Master’s Mate 
Able Bodied Seaman 
Gunner’s Mate and 
Quartermaster 
Able Bodied Seaman 


Deserted September 2, 1742 


Deserted October 14, 1743, in 
South Carolina 


Deserted November 10, 1743, 
at Port Royal, S. C. 


Deserted March 30, 1743, at 
Plymouth, England 


NO. 2 


Bates, William 
*Beckworth, Francis 

Belitha, Warren 

Bennet, John 


Bennett, Thomas 

Benson, Moses 
*Bent, John 
*Berry, John 


*Berry, Rowland 
Best, W. 
*Biggs, Thomas 
*Billonge, Jacob 
Birch, Robert 
*Bird, Richard 
*Bishop, Robert 
*Black, John 
Blackburn, John 
Blancher, Noah 
Bogue, Henry 
*Bond, Henry 


Boswell, David 


*Bould, William 
*Bousher, Walter 
Bradshaw, John 


Breamer, James 


*Briggs, William 
Briskingham, William 
*Bristoll, George 
Broughton, F. 
*Brown, James 
*Brown, Joseph 


*Brown, Nathaniel 

*Brown, Talbert 

*Brown, William 

*Buckley, John 

*Bugless, Ralph 

*Bugless, Stephen 
Bull, John 


Bullman, William 
Burdock, John 
Burns, Patrick 


Able Bodied Seaman 
Able Bodied Seaman 
Ist Lieutenant 

Able Bodied Seaman 


Landsman 

Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 


Able Bodied Seaman 
Master’s Mate 
Boatswain’s Servant 
Able Bodied Seaman 
Carpenter 
Able Bodied 
Master 
Able Bodied 
Able Bodied 
Able Bodied 
Able Bodied Seaman 
Able Bodied Seaman 
and Coxswain 
Able Bodied Seaman 


Seaman 


Seaman 
Seaman 
Seaman 


Able Bodied Seaman 


Able Bodied Seaman 


Able Bodied Seaman 
Able Bodied 
Servant 
Able Bodied 
Master’s Mate 
Able Bodied Seaman 
Able Bodied Seaman 
and Quarter Gunner 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Captain’s Servant 
Sailing Master’s Serv- 
ant 
Able Bodied Seaman 
Captain’s Servant 
Able Bodied Seaman 


Seaman 


Seaman 


LAST CRUISE OF H.M.S. “LOO”—PETERSON 41 


Deserted November 27, 1743, 
at Port, Royal;: SziC: 
Deserted September 2, 1742 


Deserted December 109, 1742, 
at Portsmouth, England 
(Returned) 


Deserted April 13, 1743 


Deserted April 6, 1743, at 
Plymouth, England 


Deserted September 16, 1742, 
at Plymouth, England 

Deserted October 15, 1743, in 
South Carolina 


Deserted April 13, 1743 


42 SMITHSONIAN MISCELLANEOUS COLLECTIONS 


Burrough, John 
Burroughs, John 


Burt, William 
Burthen, James 
*Burton, John 
*Bushnell, William 
Butchard, Samuel 
*Butcher, Richard 
Campbell, Allen 
Campbell, Edward 
*Campbell, John 
*Canton, William 
Carrol, Michael 


*Carroll, John 
Carter, James 
Carter, Samuel 
Cartwright, Benjamin 
Carty, John 


*Caunter, Henry 


*Chandler, Edward 
*Charming, Edward 
Charter, William 


Chippendall, Jona 
*Christopher, William 
*Churton, James 


Collins, John 
Compton, John 
*Conday, Richard 


Condray, Charles 


Conner, John 

Cook, John 

Cook, John 

Cook, John 

Cook, Thomas 
*Cormick, James 
*Cormick, John 

Cormick, Michael 

Cormick, William 
*Couch, James 


Courteney, F. 


Able Bodied Seaman 
Able Bodied Seaman 


Midshipman 

Able Bodied Seaman 
Able Bodied Seaman 
Carpenter’s Mate 
Able Bodied Seaman 
Able Bodied Seaman 
Captain’s Servant 
Corporal 

Able Bodied Seaman 
Ordinary Seaman 
Able Bodied Seaman 


Ordinary Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Captain’s Servant 
Able Bodied Seaman 


Steward and Ordinary 
Seaman 

Able Bodied Seaman 

Boatswain’s Mate 

Able Bodied Seaman 


Able Bodied Seaman 
Able Bodied Seaman 
Landsman and Able 
Bodied Seaman 
Quarter Gunner 
Captain’s Servant 
Able Bodied Seaman 
and Midshipman 
Ordinary Seaman 


Able Bodied Seaman 
Gunner’s Servant 
Ordinary Seaman 
Able Bodied Seaman 
Midshipman 
Able Bodied Seaman 
Surgeon’s Servant 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
and Ordinary Sea- 
man 


Clerk 


Deserted October 5, 1743 at 


New York 


Deserted October 4, 
New York 


Deserted October 13, 1743, in 


South Carolina 


Deserted September 16, 1742, 


at Plymouth, England 
Deserted April 13, 1743 


Deserted November 17, 1743, 


at Port Royal, S. C. 


Deserted December 14, 1742 


VOL. 131 


NO. 2 


Coverley, William 

Cowe, Peter 
*Cowen, Philip 

Cowen, William 


*Cowey, Robert 
Cox, Anthony 
Creese, John 


*Crilly, Thomas 
Cross, Samuel 
*Crow, Philip 
*Crowley, Bryan 
Cunnan, John 
*Curry, John 
Davidson, Alexander 
Davies, Griffith 


Davies, Matthew 
*Davies, Thomas 

Davies, William 

Dawson, William 


Day, Joseph 
Deacon, J. 


*Dean, James 
Delancy, Lawrence 
*Demount, J. 
Dent, Digby 
*Dickson, William 
Dixon, David 
Donnaly, Sam 
*Donnovan, John 
Douglas, David 
-Douglass, Robert 
Dove, Benjamin 
Dover, Saunders 
Dowes, William 


Downing, Robert 
Downing, Thomas 


*Dowsing, Samuel 
Driscoll, John 

*Driscoll, William 
Drisdall, Alexander 


Duncan, John 


Ordinary Seaman 

Ist Lieutenant 

Able Bodied Seaman 
Able Bodied Seaman 


Midshipman 
Able Bodied Seaman 
Able Bodied Seaman 


Ordinary Seaman 
Captain’s Servant 
Ordinary Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Captain’s Servant 
Able Bodied Seaman 
Ordinary Seaman 


Able Bodied Seaman 
Captain’s Servant 
Cook 

Able Bodied Seaman 


Able Bodied Seaman 
Able Bodied Seaman 


Able Bodied Seaman 
Able Bodied Seaman 
Ordinary Seaman 
Captain 

Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Carpenter’s Servant 
Corporal 

Captain’s Servant 
Captain’s Servant 
Trumpeter 

Able Bodied Seaman 


Able Bodied Seaman 
Able Bodied Seaman 
and Master’s Mate 
Able Bodied Seaman 

Ordinary Seaman 
Able Bodied Seaman 
Able Bodied Seaman 


Carpenter’s Mate 


LAST CRUISE OF H.M.S. “LOO’’—PETERSON 43 


Deserted September 2, 1742 


Deserted October 13, 1743, in 
South Carolina 


Deserted August II, 1743, at 
Plymouth, England 


Deserted October 28, 1742, at 
Plymouth, England 


Deserted October 4, 1743, at 
New York 

Deserted December 12, 1742, 
at Portsmouth, England 

Deserted March 30, 1743, 
Plymouth, England 


Deserted April 13, 1743 


Deserted December 12, 1742, 
at Portsmouth, England 


Deserted April 6, 1743, at 
Plymouth, England 
Deserted September 10, 1742 


44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


Able Bodied Seaman 
Able Bodied Seaman 


Surgeon’s Mate 
Able Bodied Seaman 


*Duncan, William 
Dunn, George 

*Dunn, Matthew 
Dunstar, James Deserted October 5, 1743, at 
New York 

Able Bodied Seaman 

Able Bodied Seaman 


Landsman 


Dyar, Anthony 

*Dyer, Darby 
Eades, Thomas Deserted September 2, 1742, 
at Plymouth, England 

Able Bodied Seaman 

Able Bodied Seaman 

Master’s Servant 

Landsman 


*Easton, Thomas 
*Edgecombe, William 
Ellis, Benjamin 
*Ellory, Robert 
Ervin, Adam 
Evans, George 
Fanson, Andrew 
*Farmer, William 


Able Bodied Seaman 

Captain’s Servant 

3d Lieutenant and 2d 
Lieutenant 

Able Bodied Seaman 

Midshipman and Mas- 
ter’s Mate 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 


Farrel, James 
Ferguson, Robert 


Field, John 
*Fisher, William 
Fitzgerald, Morris 
Fitzpatrick, Henry Deserted December 12, 1742, 
at Portsmouth, England 


Fletcher, John Ordinary Seaman 


*F ling, Timothy Able Bodied Seaman 

Forrest, Richard Able Bodied Seaman 

Forsith, William Able Bodied Seaman 

*Forster, Randal Able Bodied Seaman 

Fortiene, Joseph Able Bodied Seaman Deserted November 17, 1743, 
at Port Royal, S. C. 

Fraser, Daniel Able Bodied Seaman Deserted November 27, 1743, 
at Port Royal, S. C. 

Frost, George Sailmaker and Mid- 

shipman 
*Fullmore, Henry Boatswain 


Gally, Thomas 
Gibson, George 
Gilbert, Thomas 


*Gilmore, Arthur 
Gilmore, John 

*Gold, William 
Goldsmith, John 

*Good, John 


Master’s Servant 
Able Bodied Seaman 
Able Bodied Seaman 


Quarter Gunner 

Able Bodied Seaman 
Able Bodied Seaman 
Ordinary Seaman 
Surgeon’s 2d Mate 


Deserted April 16, 1743, at 


Plymouth, England 


Deserted April 13, 1743 


Goodsides, Abraham 
Gordon, James 
Gordon, Robert 


Midshipman Deserted October 8, 1742 


Midshipman 


NO. 2 


Gorman, David 

Graham, Matthew 
*Graham, Samuel 

Grant, Henry 


*Green, Henry 
Gregory, Jo. 
Gresham, Charles 
Grossier, John 
Grun, Nicholas 

*Hallet, John 
Hamilton, James 
Hampshire, William 
Hancock, George 
Hancock, John 


*Harman, William 
Harris, John 
Harris, Richard 
Harris, Thomas 
Harrison, Theodore 
Hartie, John 


*Hartman, Christian 
*Harwood, Lewis 
*Hatch, John 


*Hatfield, Willey 
Hawkins, John 
Hawkins, John 
Hay, David 
Hayes, Samuel 
Hays, James 
Headley, Christian 
Heaver, James 
Hemins, John 

*Henderson, James 
Henry, William 
Henton, John 

*Hickey, Thomas 

*Higgenson, James 

*Higginson, John 

*Highmas, Thomas 

*Hinds, Michael 

*Hogg, Peter 
Holliday, Richard 
Holmes, John 

*Hope, John 
Horlock, Joseph 


Able Bodied Seaman 

Coxswain 

Able Bodied Seaman 

3d Lieutenant’s Serv- 
ant 

Ordinary Seaman 

Master’s Servant 

Landsman 

Landsman 

Able Bodied Seaman 

Second Gunner 

Midshipman 

Able Bodied Seaman 

Able Bodied Seaman 

3d Lieutenant’s Serv- 
ant 

Able Bodied Seaman 

Boatswain’s Servant 

Pilot 

Master’s Servant 

Able Bodied Seaman 

Ordinary Seaman 


Able Bodied Seaman 


2d Gunner and Master 
at arms 
Able Bodied Seaman 
Captain’s Servant 
Ordinary Seaman 
Master’s Mate 
Able Bodied Seaman 
Able Bodied Seaman 
Trumpeter (?) 
Able Bodied Seaman 
Midshipman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Ordinary Seaman 
Boatswain’s Servant 
Quartermaster’s Mate 
Master’s Servant 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Midshipman 
Quartermaster 
Surgeon’s 2d Mate 


LAST CRUISE OF H.M.S. “LOO’—PETERSON 


Deserted September 2, 1742 
Deserted September 10, 1742 


Deserted June 30, 1743, at 


Portsmouth England 


Deserted October 13, 1743 


Deserted December 26, 1742 


Deserted April 13, 1743 


46 SMITHSONIAN 


Horseman, Arthur 
Hughes, John 
*Hull, Lancaster 


Hulsal, Arthur 
Hunt, John 
Hunt, John 
Hussey, I. 
*Hyslop, Thomas 
Ingerton, Dennis 


Jackman, William 
Johnson, Luke 


Jolly, George 
*Jolly, Thomas 


Jones, Anthony 
Jones, David 
Joynes, James 
*Juba, Luke 
Julian, Charles 


Keeler, Robert 
*Keighley, William 
*Kelley, Morgan 

Kelsey, William 


*Kennedy, John 
Kent, Ambrose 


Keys, Robert 
Killrick, Isham 


*Kilsey, William 
King, Daniel 


*King, Nathaniel 
*Kingsbury, William 
*Kirk, James 
*Kirk, Samuel 
*Kivey, John 
*Knowles, Edward 
*Knowling, James 
Lake, Mark 


Lamb, William 


Able Bodied Seaman 
Able Bodied Seaman 


Able Bodied Seaman 
and Midshipman 
Able Bodied Seaman 
Sailing Master 
Captain’s Servant 


Armorer 
Able Bodied Seaman 


Ordinary Seaman 
Able Bodied Seaman 


Ordinary Seaman 
3d Lieutenant’s Serv- 
ant 
Able Bodied Seaman 
Able Bodied Seaman 
Quarter Gunner 
Purser’s Servant 
Able Bodied Seaman 


Master at Arms 

Able Bodied Seaman 

Able Bodied Seaman 

Surgeon’s Servant and 
Able Bodied Sea- 
man 

Able Bodied Seaman 


Quartermaster 

Able Bodied Seaman 
and Midshipman 

Captain’s Servant 

Landsman 


Able Bodied Seaman 
Captain’s Servant 
Gunner’s Servant 
Gunner 

Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Ist Lieutenant’s Sery- 

ant 
Quarter Gunner 


MISCELLANEOUS COLLECTIONS 


VOL. 131 


Died March 24, 1742, at 
Plymouth, England 

Deserted December 12, 1742, 
at Portsmouth, England 


Deserted August II, 1743, at 
Plymouth, England 

Deserted April 13, 1743 

Deserted April 6, 1743, at 
Plymouth, England 


Deserted December 19, 1742, 
at Portsmouth, England 


Deserted November 17, 1743, 
at Port Royal, S. C. 


Deserted September 2, 1742, 
at Plymouth, England 


NO. 2 


*Lander, Robert 
*Langston, Richard 
*Lather, Samuel 
Lavermore, Joseph 
Lawler, John 
*Ledgerwood, James 
*Lemarr, Stephen 
*Leslie, George 
Lewis, Christopher 
*Lewis, Edward 
Lewis, Theodore 
*Lewiswentz, David 
*Limb, Giles 
Linch, William 
Liston, William 
*Lloyd, William 
Lobb, Thomas 
Lodge, John 
Logan, Charles 


Long, Samuel 


Long, William 

Lookert, Patrick 

Luch, John 
*Lyford, William 


*Maby, John 
Malt, Issac 


Manley, George 
Mannon, John 
*Marriot, William 
Martin, John 
Maxwell, James 
May, William 


*McCann, John 
McCarty, Daniel 
McChownley, Lott 
McCleland, Robert 


*McClockland, William 
McCowley, Charles 
McDugal, James 
McKensie, Samuel 
McKnight, James 

*McNeal, William 
Meachem, James 


Able Bodied Seaman 
Quartermaster 
Able Bodied 
Able Bodied 
Able Bodied 
Able Bodied 
Able Bodied 
Purser 

Able Bodied Seaman 
Ordinary Seaman 
Able Bodied Seaman 
Captain’s Servant 
Able Bodied Seaman 
Able Bodied Seaman 
Ordinary Seaman 
Midshipman 
Surgeon’s Servant 
Boatswain’s Servant 
Ordinary Seaman 


Seaman 
Seaman 
Seaman 
Seaman 
Seaman 


Able Bodied Seaman 


Bodied 
Bodied 


Seaman 
Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
and Midshipman 
Ordinary Seaman 
Able Bodied Seaman 


Able 
Able 


Ordinary Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Quartermaster 
Ist Lieutenant’s Serv- 
ant and Able Bodied 
Seaman 
Able Bodied 
Able Bodied 
Able Bodied 
Able Bodied 


Seaman 
Seaman 
Seaman 
Seaman 


Able Bodied Seaman 
Ordinary Seaman 
Able Bodied Seaman 
Quarter Gunner 
Ordinary Seaman 
Quartermaster 

Able Bodied Seaman 


LAST CRUISE OF H.M.S. ‘“‘LOO’—PETERSON 


Deserted April 16, 1743 


Deserted April 13, 1743 


Deserted November 5, 1742, 


at Plymouth England 


Deserted March 30, 1743, at 


Plymouth, England 


Deserted October 2, 1742 


Deserted October 3, 1743, at 


New York 
Deserted April 13, 1743 


Deserted October 4, 1743, at 


New York 


48 SMITHSONIAN MISCELLANEOUS COLLECTIONS 


Mellan, John 
Meridith, John 
Mignam, George 
*Miller, James 
*Miller, Nicholas 
Miller, Stephen 
Mills, Nathan 
*Millsom, James 
Mitchell, R. 
*Molineaux, John 


Moore, Samuel 


More, Thomas 
Morran, William 
Morrison, John 
Morrow, Hugh 
*Mortimer, Robert 
Murphy, Francis 


*Murphy, James 
*Murphy, John 
Murphy, Patrick 
Murphy, Samuel 
*Nagan, James 
*Nellson, Robert 
*Newgent, Patrick 
*Newson, Richard 
Nobbs, Stephen 
Norman, Edward 


*Norman, William 
Northesk, Earl of 
*Nott, Ebinezer 


*Nuikle, Robert 
Ogburne, John 
Oliver, Richard 

*Orr, Archibald 

*Osbourne, William 
Parker, Peter 
Parr, William 


*Parrott, Samuel 
Patrick, Richard 
Patten, Hugh 


Pearse, Samuel 


Boatswain’s Mate 
Ordinary Seaman 
Surgeon 

Able Bodied Seaman 
Able Bodied Seaman 
Boatswain’s Servant 


Able Bodied Seaman 

Ordinary Seaman 

Ordinary Seaman and 
Able Bodied Sea- 
man 

Yeoman 


Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Captain’s Servant 

Able Bodied Seaman 
Able Bodied Seaman 


Able Bodied Seaman 


Able Bodied Seaman 
Able Bodied Seaman 
Ordinary Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Landsman 

Ordinary Seaman 


Gunner’s Servant 

Captain 

Ordinary Seaman and 
Able Bodied Sea- 
man 

Able Bodied Seaman 

Captain’s Servant 

Landsman 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Quartermaster 


Captain’s Servant 
Cook’s Servant 


Able Bodied Seaman 


Able Bodied Seaman 


Deserted April 13, 1743 


Deserted September 2, 1742 


Deserted September 10, 1742, 


at Plymouth, England 


Deserted August II, 1743, at 


Plymouth, England 


Deserted December 29, 1743, 


at Port Royal, S. C. 


Deserted September 2, 1742 


Deserted December 12, 1742, 
at Portsmouth, England 


Deserted March 30, 1743, at 


Plymouth, England 
Deserted April 16, 1743 


VOL. 131 


NO. 2 


*Pearson, Thomas 


Pegan, Roger 


Pegan, Thomas 
Pelican, John 


Pemell, Thomas 
*Peters, William 


Phallem, Edmund 
*Phillips, Thomas 
*Phonix, Philip 
*Pickering, Michael 
*Plantain, Charles 

Poole, Thomas 

Potterfield, George 

Powell, Philip 


Pownswell, Edward 


*Pretty, Thomas 
*Price, James 
*Priest, Lewis 
Puttick, William 
*Quaco, John 


*Quin, John 
Ramsey, Patrick 
*Randall, James 


*Randall, James 
*Randall, John 


Randell, James 
*Randolph, J. 


*Ratsey, George 
Reed, Thomas 


*Rhode, John 


*Richards, Nicholas 
*Richardson, William 
Richey, David 
*Richmond, Andrew 
*Rider, Charles 


Ordinary Seaman and 
Able Bodied Sea- 
man 

Able Bodied Seaman 


Able Bodied Seaman 
Able Bodied Seaman 


Able Bodied Seaman 
Able Bodied Seaman 
and Midshipman 
Able Bodied Seaman 

Cook 
Able Bodied Seaman 
Clerk 
Able Bodied Seaman 
Pilot 
Able Bodied Seaman 
Able Bodied Seaman 


Able Bodied Seaman 


Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Captain’s Servant and 
Ordinary Seaman 

Able Bodied Seaman 

2d Gunner 

2d Lieutenant and Ist 
Lieutenant 

Able Bodied Seaman 

Able Bodied Seaman 
and Midshipman 

Ist Lieutenant 

Able Bodied Seaman 
and Master’s Mate 

Cooper and Able Bod- 
ied Seaman 

Able Bodied Seaman 


Able Bodied Seaman 
and Ordinary Sea- 
man 

Able Bodied Seaman 

Able Bodied Seaman 

Quartermaster’s Mate 

Able Bodied Seaman 

Able Bodied Seaman 


LAST CRUISE OF H.M.S. “LOO’—PETERSON 


Deserted November 27, 1743, 


at, Port Royal, S/C: 


Deserted March 17, 1742, at 


Plymouth, England 


Deserted March 30, 1743, at 


Plymouth, England 


Deserted December 12, 1742, 


at Portsmouth, England 


Deserted October 6, 1743, at 


New York 


49 


50 SMITHSONIAN MISCELLANEOUS COLLECTIONS 


Risden, Joseph 
Roach, Henry 


Roberts, David 
Roberts, Edward 


Roberts, John 
Roberts, John 
Roberts, Joseph 

*Roberts, William 
Robinson, Alexander 
Robinson, John 
Rogers, Robert 
Rowe, Edward 
Rowe, George 


Rowe, Peter 


Royall, James 
*Russell, Richard 
Ryan, John 


St. Lawrence, Samuel 
Salisbury, Edward 
*Salmon, Joseph 
*Salter, James 
*Saunders, Joseph 
Scannel, John 
Scott, George 
Scott, Richard 
*Shaw, Patrick 
*Shearing (John or 
Joseph) 


*Shearing, Thomas 
Sherwood, William 
Shirley, Washington 

*Shoart, Oliver 
Simmonds, Richard 


*Simms, James 
Singleton, William 

*Skinner, Philip 
Sluman, John 
Smith, Benjamin 

*Smith, James 
Smith, Paul 
Smith, Theodore 


Able Bodied Seaman 
Able Bodied Seaman 


Ordinary Seaman 


Sailmaker’s Mate 
Captain’s Servant 
Captain’s Servant 
Captain’s Servant 


Able Bodied Seaman 
Landsman 

Able Bodied Seaman 
Quarter Gunner 


Able Bodied Seaman 


Able Bodied Seaman 
Boatswain’s Servant 
Able Bodied Seaman 


Ordinary Seaman 

Able Bodied Seaman 

Ist Captain’s Servant 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Gunner 

Landsman 

Able Bodied Seaman 

Ist Lieutenant’s Servy- 
ant and 3d Lieuten- 
ant’s Servant 

Able Bodied Seaman 

Able Bodied Seaman 

3d Lieutenant 

Able Bodied Seaman 

Quarter Gunner 


Ordinary Seaman 
Ordinary Seaman 
Ordinary Seaman 
Gunner 
Able Bodied Seaman 
Able Bodied Seaman 
Servant 
Boatswain’s 
and Able 
Seaman 


Servant 
Bodied 


VOL. 131 


Deserted December 12, 1742, 
at Portsmouth, England 
Deserted October 28, 1742, at 

Plymouth, England 
Deserted December 19, 1742, 
at Portsmouth, England 


Deserted September 2, 1742 


Deserted December 12, 1742, 
at Plymouth, England 

Deserted October 4, 1743, at 
New York 


Deserted September 10, 1742, 
at Plymouth, England 
Deserted April 13, 1743 


Deserted April 6, 1743, at 
Plymouth, England 


Deserted September 2, 1742 


Deserted October 4, 1743, at 
New York 


NO. 2 


Softley, Robert 


Somerwel, Joseph 
Southard, Thomas 


Spare, Thomas 
Sparks, Joseph 
*Spinks, Stephen 
Spry, Nicholas 


Stanford, Richard 
Stapleton, A. 
Stephenson, James 


*Steuart, Neil 
Stevenson, Henry 


Stewart, Alexander 
*Stiveash, Stiven 
*Stoneham, John 

Stradder, Forbel 

Stroud, John 
*Sullivan, Thomas 

Swain, William 


*Sweeny, Daniel 
*Swickman, Thomas 
Swift, Theodore 


*Tabler, Thomas 


*Taylor, John 


*Taylor, Jonathan 
*Taylor, William 


Thatcher, John 
*Thompson, Richard 

Thoyer, Peter 
*Tipper, John 

Tobyn, George 


Todd, Alexander 


*Torginton, Joseph 
Treacey, William 


Able Bodied Seaman 


Able Bodied Seaman 
Able Bodied Seaman 


Able Bodied Seaman 
Able Bodied Seaman 


Able Bodied Seaman 


Quartermaster’s Mate 
Landsman 


Surgeon 
Able Bodied Seaman 


Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Landsman 

Boatswain 

Ordinary Seaman 

3d Lieutenant’s Serv- 
ant and Captain’s 
Servant 

Able Bodied Seaman 

Landsman 

Able Bodied Seaman 


Captain’s Servant and 
Able Bodied Sea- 
man 


Carpenter’s Servant 
and Able Bodied 
Seaman 


Able Bodied Seaman 
Captain’s Servant and 
Ordinary Seaman 

Quartermaster 

Pilot 

Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 


Able Bodied Seaman 


Carpenter’s Servant 
Able Bodied Seaman 


LAST CRUISE OF H.M.S. “LOO’’—PETERSON 


Deserted March 30, 1743, at 


Plymouth, England 
Deserted April 13, 1743 


Deserted September 10, 1742, 


at Plymouth, England 


Deserted March 17, 1742, at 


Plymouth, England 


Deserted September 2, 1742, 


at Plymouth, England 


Deserted October 15, 1743, in 


South Carolina 


Deserted April 13, 1743 


Deserted August 5, 1743, at 


Portsmouth, England 


Deserted October 6, 1743, at 


New York 


Destered March 30, 1743, at 


Plymouth, England 


Deserted December 26, 1742, 
at Portsmouth, England 


52 SMITHSONIAN MISCELLANEOUS COLLECTIONS 


*Triming, Thomas 

*Trist, Nathaniel 
Trott, Thomas 
Trotter, Benjamin 
Trouve, Paul 


*Trunker, William 
Tunis, Michael 
Turford, Thomas 


*Utting, Ashby 
Vincent, Aaron 
*Vivian, John 
*Wadlin, Richard 
*Walker, James 
Walker, John 


Walker, Peter 
Wallis, Benjamin 


Warceys, Thomas 
*Ward, Samuel 
*Weatherill, John 

Wedlock, John 


Welch, Michael 
Welsh, Peter 
*Wemuss, Robert 
Wemy, James 
Wheeler, Robert 
*White, Richard 
*White, Robert 
Whitver, Thomas 
Wilkinson, Edward 
*Williams, Richard 
*W illmot, Theodore 
Wills, J. 
Wilson, George 


Wilson, Lawrence 
Wise, John 


Woodgate, William 
*Wotton, William 
Wright, William 


*Yeates, James 
*Young, John 
Younger, John 


Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
3d Lieutenant’s Sery- 
ant 

Able Bodied Seaman 
Able Bodied Seaman 
Carpenter 


Captain 
Able Bodied Seaman 
Carpenter 


Ordinary Seaman 
Able Bodied Seaman 


Able Bodied Seaman 


Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 


Able Bodied Seaman 
Ordinary Seaman 
Cook’s Servant 

Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 


Ist Lieutenant’s Sery- 
ant 

Able Bodied Seaman 

Able Bodied Seaman 
and Midshipman 


Able Bodied Seaman 
Able Bodied Seaman 


Captain’s Servant 
Ordinary Seaman 
Captain’s Servant 


Deserted September 10, 1742, 


at Plymouth, England 


Deserted October 2, 1742, at 


Cork, Ireland 


Deserted December 12, 1742, 


at Portsmouth, England 


Deserte@ September 16, 1742, 


at Portsmouth, England 


Deserted December 12, 1742, 


at Portsmouth, England 


Deserted September 23, 1742, 


at Plymouth, England 


VOL. 131 


MARINE LIST 


Colonel Cotterel’s Regiment : 
*Allen, Robert 
Baker, John 


*Ball, William 
*Bond, George 
*Brooke, Joshua 
*Brooks, George 
Brooks, John 
Brown, George 
Brown, John 
Cant, Thomas 
Clint, Richard 
*Cole, Joseph 
*Cook, Samuel 
Corbett, Thomas 
Diamond, Robert 
*Dight, Edward - 
*Douglass, Edmund, Sergeant 
Drake, Richard 
*Farries, William, Sergeant 
Finch, Samuel 
*Fitzsimmons, Thomas 
*Gaddish, Lazarus 
*Gleddon, Richard 
*Gould, William 
Grovenor, Francis 


Hall, William 
*Hardeman, William 
*Harding, Matthew 
*Haydon, Timothy 

Hodge, Hugh 


*Hold, Thomas 
~ Hughes, Edward 
*Hyatt, Samuel 
Isaac, Titus 
Johnson, Adam 
*Jones, Philip 
*King, Joshua 
Lee, John 
Martin, Thomas 
*Matthews, Joseph 
*McCraw, Daniel 
Miller, Richard 


Mitchell, John 


LAST CRUISE OF H.M.S. “LOO’’—PETERSON 53 


Deserted December 26, 1742, 
at Portsmouth, England 


Deserted June 30, 1743, at 
Portsmouth, England 


Deserted December 12, 1742, 
at Portsmouth, England 


Deserted December 2, 1742, 
at Portsmouth, England 


54 SMITHSONIAN MISCELLANEOUS COLLECTIONS 


*Morgan, John 
*Morgan, William 
*Mould, Thomas 
*Murray, Peter 
*Overshott, John 
Pearce, George 
Phillips, Edward 
Phillips, William 
Prest, Timothy 
*Ridghewothh, Thomas 
*Risden, Samuel 
Roberts, John 
Rowls, John 
Searle, John 
Short, George 
Spraeg, Nicholas 
*Stevens, Samuel 
*Stone, John 
Thrasher, John 
*Toll, John 
Trovana, William 
*Turner, Jonas 
*Turpin, John 
*Vaughan, Hector, Lieutenant 
*Walker, William 
Whiteker, Thomas 
*Williams, Thomas 
Wills, Samuel 
*W oodeson, John 
Woodley, Nicholas 
Colonel Wynyard’s Regiment 
Aldridge, William 
Chappel, Samuel 
Clayton, Joseph 
Hancock, Justinian 
Hope, Richard 
*Kent, Benjamin 
*Korgett, John 
Parker, Daniel 
*Stokes, William 
*Trowes, Richard 
*Wright, Samuel 


VOL. 131 


APPENDIX D 


ROSTER OF THE FAMILY AND SERVANTS OF GEORGE CLIN- 
TON, GOVERNOR OF NEW YORK FROM SEPTEMBER 1743 TO 
OCTOBER 1753, PASSENGERS ON THE “LOO” DURING HER LAST 
VOYAGE TO AMERICA, AUGUST-SEPTEMBER 17431 


Clinton, Ann (Governor’s wife) Cunningham, Archibald 
Clinton, Ann Davies, Ann 

Clinton, George (Governor ) Ellis, Thomas 

Clinton, Henry Fenton, Sarah 

Clinton, Lucy Harvey, Phillip 
Clinton, Mary Ryves, James 

Ascough, John Vanham, John 

Aurneo, Ann White, Margaret 
Blundell, Christopher Williams, William 
Catherwood, John Willson, Ann 


_ 


SOON ANDwW DH 


1 Based on the Muster Roll of the Loo, ADM 36, vol. 1823. 


APPENDIX E 
COMMANDING OFFICERS OF THE “LOO” 

MP arland MiNOUEEE: cae cbichasisioie cae cieines ore April 4, 1707—October 1, 1709 
PETOR DEER A OMAN ojnis, oe ctesaiecte e w/tnavouse aoaierere October I, 1709- ? 
PRAVV AGUA ES EAtt: cya /scrorsiciaeielererais, a eranie chorale 1700- 1715 
PREM IAME ori Miad wacats's slo wcee 3. ar oierd 5 vera esd eainisie April 1717(?)—December 1717 
POP EOEHEHE, GEGERE Kies diaisiae wed onercvess April 1718- 1720 

SCOte—(C CAntAA! oalay.c nae ove stnsteg sci October 1720- 1722 

Waterhouse, Tho. (Commodore)....... February 1727/28—-October 1728 
Me Mather laces J cle dmis tee cate e October 22, 1728-April 2, 1729 
PBetmeley, VV MISARD ws in coicnles von e baa es April 2, 1729- ? 
BUESCRE GNOME 2rd Bers Bietercls aeileln'ecsieainm ne May 5, 1735-May 1737 
pe Obeasic, BAG) Ol eisio ic ca acted aveie's cieccle es January 4, 1741/42-September 1742 
MPSPCRIE MOPESISY 8 tc ghee caioes hiaiaionorvera-a arate hoch September 17, 1742-April 11, 1743 
oN ECMO CEM OMLIND igs eh nO ccls! prceistioieinl veh eleyecw des April 11, 1743-February 5, 1743/44 


1 Based on the Navy List, 1707-1744. 


55 





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ye 


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a 





SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 2131, NO: 2, PL. 2 





1. Copper coins recovered from the wreck site of the Loo in the summer of 
1950. Left to right: Spanish 8 maravedi circa 1600, Spanish 8 maravedi circa 
1640, and Swedish 4 ore dated 1720. This last coin was important evidence 


in dating the wreck. Lent to the U. S. National Museum by Dr. and Mrs. 
George Crile, Jr. 





2. Small Queen Anne pewter teapot from the wreck of the Loo. Giit 
to the National Museum from E. A. Link. The pot was damaged by fire 
when the Loo was burned by her captain after being wrecked. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


VOL. 2315, NO. 2; 





PES 


The remains of the ship lay between two coral reefs in 


(Photograph by Dr. George Crile, Jr.) 


he wreck site. 


35 feet of water. 


The salvage boats of the expedition at anchor over t 


2, Pees 


NO. 


id, 


ONIAN MISCELLANEOUS COLLECTIONS 


S 


SMITH 





oY} STOAOI 


(‘Af ‘apg 95 


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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131-"NO.. 2, PL. 6 





A basket of coral-sand-encrusted shot coming up from the bottom. 
(Photograph by Dr. George Crile, Jr.) 


2, Pla 


131, NO. 


VOL. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


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SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 3 


SYNONYMICAL NOTES ON NEOTROPICAL 
FLIES OF THE FAMILY TABANIDAE 
(DIPTERA) 


By 
G. B. FAIRCHILD 


Gorgas Memorial Laboratory, Panama 





(PUBLICATION 4225) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
JANUARY 11, 1956 


THE LORD BALTIMORE PRESS, INC. 
BALTIMORE, MD., U. S. A. 





SYNONYMICAL NOTES ON NEOTROPICAL 
FLIES OF THE FAMILY TABANIDAE 
(DIPTERA)! 


By G. B. FAIRCHILD 
Gorgas Memorial Laboratory, Panama 


The nomenclature of the Neotropical flies of the family Tabanidae 
has long been in a state of great confusion, in spite of the efforts of a 
number of students to bring it into some sort of order. The main dif- 
ficulties seem to have been the lack of adequate collections in any one 
place and the very numerous inadequate descriptions by several of the 
older authors. The existing catalogs of Kertész (1900, 1908) and 
Surcouf (1921) for the Tabanidae of the World are quite uncritical 
and are chiefly lists of names. The catalog of the Neotropical Tabani- 
dae prepared by Krober (1934) was a great step forward, but subse- 
quent work has modified greatly the understanding of generic and 
higher categories, and he failed in many cases to appreciate the value 
of a study of the type specimens of the older descriptions. His cata- 
log, therefore, although extremely useful, has often proved unreliable. 

During the fall of 1953 I was enabled, through the aid of a generous 
travel grant from the Marsh Fund of the National Academy of Sci- 
ences, to visit the British Museum in London and the Muséum 
d’ Histoire Naturelle in Paris. The trip was undertaken for the purpose 
of studying and comparing specimens with the types of Neotropical 
Tabanidae (horse flies and allies) contained in the collections of the 
British Museum in London and the Muséum d’Histoire Naturelle in 
Paris. The Neotropical species of Tabanidae described by Francis 
Walker between 1848 and 1860, by M. J. Macquart between 1834 and 
1855, and by J. M. F. Bigot in 1892 have been a serious stumbling 
block to students for many years. Not only did these three authors 
among them describe some 300 species, but their descriptions were, for 
the most part, so superficial and inadequate that a large proportion of 
their names have remained unrecognized or misinterpreted. Further- 
more, some 27 generic names have been based on these species, often 
without adequate knowledge of their characters. Although the primary 


1 Published through a grant from the Gorgas Memorial Institute of Tropical 
and Preventive Medicine, Incorporated. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 131, NO. 3 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


purpose of the trip was to study the type specimens of these three 
authors, most of which are deposited in either London or Paris, it 
was felt important to examine all other types of Neotropical Tabanidae 
available in these two institutions. 

Since time would not permit the careful description and drawing 
of all the species likely to be found, it was felt of utmost importance 
to take over for comparison specimens of as many species as possible. 
Through the courtesy of the authorities of the Museum of Compara- 
tive Zoology at Cambridge, Mass., and of the U. S. National Museum, 
a collection of nearly 600 species of Neotropical Tabanidae was 
secured and taken to London and Paris. 

Although not a few of the types I had hoped to see have been lost 
or destroyed in the course of the nearly 100 years since they were 
described, I was fortunate in being able to see a good many additional 
species of more recent date which I had not expected to find. These in- 
cluded a number of the types of species described by Osten Sacken, 
Williston, Townsend, Surcouf, Ricardo, Summers, and Krober. I was 
able to bring back specimens matched with the types of about 220 
Neotropical species, fairly complete notes, in some cases with drawings, 
on a further 107 types, and miscellaneous notes on an additional 60 to 
70 species, not types, of which I had not previously seen specimens. 

It is a pleasure to acknowledge my indebtedness to Dr. Alexander 
Wetmore of the National Academy of Sciences, who facilitated the 
procuring of travel funds, and to Dr. Joseph Bequaert and Dr. Alan 
Stone for their generosity in lending material in the collections under 
their care. Capt. N. D. Riley, C. B. E., Keeper of Insects at the British 
Museum, most generously placed the facilities of that institution at 
my disposal; and I am most especially grateful for the invaluable help 
and cordial hospitality of H. Oldroyd and Paul Freeman of the Diptera 
section of the Museum. At Paris, M. E. Seguy, custodian of the 
Diptera section in the Muséum d’Histoire Naturelle, put the collections 
at my complete disposal and did everything possible to make my short 
stay pleasant and profitable. The drudgery of taking dictation and 
typing the extensive notes fell to my wife, without whose invaluable 
assistance the work could not have been completed. 

The Tabanidae at the British Museum are arranged primarily on 
a taxonomic basis, the various groups following one another irrespec- 
tive of locality. Each drawer is marked with the genera it contains and 
a colored slip indicating the geographical regions represented. There 
is also a card catalog of the species in the collection. All types are in- 
corporated in the general collection but are marked with small circular 


No. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 3 


labels, usually red for primary types, green for cotypes, and yellow 
for paratypes. 

In its present state, the arrangement and labeling are largely the 
work of E. E. Austen, the late curator, and H. Oldroyd, the present 
curator. Austen is largely responsible for verifying Walker’s types, 
which, until his time, were not marked as such. The Bigot collection, 
containing Macquart and Bigot types, was remounted on double 
mounts after receipt by the British Museum. In most cases only one 
specimen of a series bore a label—those of Macquart which Bigot had 
pasted onto larger labels, or his own folded and often much defaced 
labels. In repinning this material great care seems to have been taken 
to put the labels back on the same specimens, though in one or two 
cases there appears to have been an exchange of labels. Mr. Oldroyd 
has done the great service of marking all the types with distinctively 
colored labels, a procedure that greatly facilitated their recognition. 

At Paris, the collections are housed in large glass-topped cardboard 
boxes. There has been no effort to rearrange the Tabanidae, and the 
collection is really a series of separate collections. Although most of 
the Tabanidae are together, the Macquart collection is in its own series 
of boxes, not mixed with subsequent additions. Surcouf’s material 
is also separate. This policy seems the only sound one under the pre- 
vailing conditions, as M. Seguy is in charge of several other orders 
besides Diptera, and has but one assistant. The Meigen collection of 
Diptera, as well as several other largely European collections, is thus 
preserved. For the most part, the Macquart types are not labeled as 
such and bear only their original labels, so that reference to the orig- 
inal descriptions is often necessary. The box labels under which the 
species stand are, I believe, a later addition, and are not very helpful 
or consistent. Since most of the specimens are types or easily recog- 
nized species, determination as to which specimen is a type is usually 
not difficult. The Macquart collection is also divided geographically, 
the Neotropical, Nearctic, etc., species placed together. Owing to lack 
of realization that Mexican material may have been considered Nearc- 
tic, I quite likely missed seeing the types of several of Macquart’s 
species, as I lacked time to go through other than the Neotropical 
boxes. 

Most of Walker’s Neotropical species were described in the “List 
of the Specimens of Dipterous Insects in the Collection of the British 
Museum,” which is here abbreviated to “List” with volume, page, and 
year. His other publications are more fully cited. Macquart’s species 
appeared mostly in a series of articles entitled “Dipteres Exotiques 
Nouveaux ou Peu Connus,” here abbreviated to “Dipt. Exot.’ This 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


series appeared more or less simultaneously in the Mémoires de la 
Société Royale des Sciences, de l’Agriculture et des Arts de Lille, and 
in a separately published form put out by Librairie Encyclopédique de 
Roret, Paris. There is no difference in the text, but the pagination is 
different, and in some cases the reprint is of an earlier or later date. 
I have not been able in all cases to check as to which edition a given 
page number refers, but since the work is adequately indexed in both 
editions, this is not of great importance. The possible conflict between 
Supplement 4 of Macquart’s work and the Diptera Saundersiana of 
Walker, both dated 1850, does not seem to concern any names for 
Neotropical Tabanidae. Mr. Oldroyd felt that since Macquart’s paper 
was read in June 1849, though not published until sometime in 1850, 
while Walker’s paper appeared after September 12, 1850, it is best 
to assume priority for Macquart’s names where conflict occurs. Nearly 
all of Bigot’s species were described in an article entitled “Descrip- 
tions de Diptéres Nouveaux” published in 1892 in Mém. Soc. Zool. 
France, vol. 5, and will be cited here merely by the date and page. 
Secondary references will be cited only by author and date, the full 
reference cited only in the bibliography. 

Although fairly detailed notes, and in some cases camera lucida 
sketches, of nearly all the types examined were made, it has seemed 
better to present the results in the more condensed form of an anno- 
tated list of the types examined. Much information that might aid 
in the determination of specimens has thus had to be omitted, and the 
list is primarily of nomenclatorial interest. It is planned, as time and 
opportunity permit, to make the more interesting and perhaps more 
valuable descriptive matter of the notes available, together with figures 
of homotypes, in connection with planned revisionary studies now 
being undertaken in collaboration with Dr. C. B. Philip. In the mean- 
time, copies of the full notes will be deposited in the U. S. National 
Museum and the British Museum for reference purposes. 

I have refrained from specifically selecting lectotypes of species with 
more than one specimen in the type series. To do so at the time of 
examination would have used more time in writing labels than I could 
spare; to do so now would lead me into explanations and justifica- 
tions for my action in each case and would unduly lengthen this paper. 
In cases where more than one species is obviously involved in the type 
series, I have indicated which one I believe should represent the name. 

The species discussed here are nearly all listed by Krober (1934) 
in his catalog of the Neotropical Tabanidae. A few species listed by 
him have been omitted here, such as T. pruinosus Bigot from Mexico, 
fully treated by Philip (1950), and T. parvidentatus Macquart, dis- 





NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 5 


cussed by Bequaert (1940). Several other names are Nearctic and 
cataloged by Philip (1947). Names omitted by Krober are so indicated 
here, except in the case of those originally published with no locality 
or a non-Neotropical locality. 

The type material of the following 64 species was not found either 
in London or Paris. The species are listed below alphabetically, with 
place and date of publication, locality, and collection as originally given 
where this information is available to me. Several names are homo- 
nyms and so indicated. Species for which no locality was originally 
given, or where the author states his ignorance of the provenance 
of the specimen, are indicated by “Loc. ?.” In many cases these species 
may not have been Neotropical, and in the case of the collection at 
Paris, no search of the Old World or Nearctic collections was made 
for them. In London, the card catalog of types was checked, and 
so it is reasonably certain that Walker’s species now missing from the 
collections are truly lost. In some cases additional notes on status have 


been added. 


Dichelacera abiens Wlk. 1848, List, 1: 191. West Indies, B. M. Seen by Ricardo 
(1904) but subsequently lost (Bequaert, 1940). 

Tabanus advena Wlk. 1850, Newman’s Zoologist, 8, App., p. Ixix. Loc.? B. M. 

Tabanus albiscutellatus Macq. 1850, Dipt. Exot., Suppl. 4:34. Mexico = Jeu- 
caspis Wied. (Osten Sacken, 1878; Krober, 1934; Fairchild, 1941). Perhaps 
in Nearctic coll. 

Tabanus albivittatus Macq. 1834, Hist. Nat. Dipt., 1: 206. Loc.? Coll. Percheron. 
Probably not Neotropical. T. albivittatus Schuurm. Stekh. 1926 is a homonym. 

Pangonia ardens Macq. 1838, Dipt. Exot., 1(1):197. Saint Leopold. Coll. 
Serville. 

Tabanus argentifrons Wik. 1848, List, 1: 186. Loc.? B. M. 

Pangonia bicolor Macq. 1850, Dipt. Exot., Suppl. 4:27. Mexico = Esenbeckia 
semiflava Wied. (Bellardi, 1859; Krober, 1934). Homonym of P. bicolor 
Macq. 1846. New Holland. 

Haematopota bivittata Macq. 1834, Hist. Nat. Dipt., 1: 212. Amer. Merid.= 
-Diachlorus bivittatus Wied. 1828 (Krober, 1934). From description, a Dia- 
chlorus, though not credited to Wiedemann in original description. 

Tabanus bivittatus Macq. 1845, Dipt. Exot., Suppl. 1: 35. Brazil. Coll. Spinola. 

Tabanus bonariensis Macq. 1838, Dipt. Exot., 1 (1):142. Buenos Aires. Coll. 
Serville. Kroéber (1934) places in Agelanius with acupunctatus Rond. 1868, 
as synonym. iw 

Tabanus brevivitta Wik. 1850, Newman’s Zoologist, 8, App., p. xcvi. Loc.? 
B. M. Homonym of T. brevivitta Wlk. 1848. Australia. 

Tabanus castaneus Macq. 1834, Hist. Nat. Dipt., 1: 198. Cayenne. The descrip- 
tion indicates probable synonymy with T. unicolor Wied. 

Tabanus castaneoventris Macq. 1838, Dipt. Exot., 1 (1) : 152. Loc.? Mus. Paris. 

Tabanus chrysoleucus Wlk. 1854, List, 5, Suppl. 1: 327. Brazil. B. M. 

Tabanus despectus Krober 1930, Dipt. Pat. S. Chile, 5 (2):158. Chile. B. M. 
(Paratype.) This and several other species described in same paper appear 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


not to have been returned to B. M. by Kroéber. T. despectus Fchld. 1942 is 
a homonym, but proposal of a replacement name now seems unnecessary, as 
the species is uncertainly distinct. 

Tabanus dorsivitta Wlk. 1850, Dipt. Saund., 1:39. S. America? B. M. The 
African T. dorsivitta Wlk. 1854 is a synonym of T. taeniola P. de B. 

Tabanus flammans Wlk. 1848, List, 1: 153. Loc.? B. M. 

Tabanus flavibarbis Macq. 1845, Dipt. Exot., Suppl. 1: 169. Cayenne. Coll. 
Spinola. Kréber (1930h) claims to have seen the type and places guyanensis 
Macq. as a synonym. The latter has, however, page priority. 

Tabanus flavifascies Macq. 1845, Dipt. Exot., Suppl. 1: 36. Colombia. Coll. de 
Breme. 

Tabanus formosus Wik. 1848, List, 1: 148. Loc.? B. M. 

Chrysops frontalis Macq. 1838, Dipt. Exot., 1 (1): 160. Saint Domingue. Col- 
lection not stated. 

Tabanus fullo Wik. 1850, Newman’s Zoologist, 8, App., p. Ixvii. Loc.? B. M. 

Chrysops fulviceps Wlk. 1854, List, 5: 285. Para. B. M. The description agrees 
best with C. incisa Macq. Ricardo (1901) says type could not be found at 
B. M. 

Tabanus fulvifasciatus Macq. 1834, Hist. Nat. Dipt., 1: 206. lLoc.? Coll. 
Percheron. 

Tabanus fulviger Wik. 1850, Dipt. Saund., 1: 65. Loc.? B. M. 

Chrysops geminata Macq. 1850, Dipt. Exot., Suppl. 4:39. Mexico. The name 
is a homonym of geminata Wied. and a presumed synonym of virgulatus Bell. 
1859. 

Tabanus gigas Macq. 1834, Hist. Nat. Dipt., 1: 200. Loc.? A homonym of 
T. gigas Herbst 1787. 

Pangonia hebes Wik. 1848, List, 1: 137. Loc.? B. M. Ricardo (1901) says type 
lost, but see Oldroyd (1954). Probably Australian. 

Tabanus hispidus Wlk. 1850, Dipt. Saund., 1: 63. Loc.? B. M. 

Tabanus honestus Wik. 1850, Dipt. Saund., 1:64. Loc.? B. M. Lutz (1907) 
lists as possible synonym of fuscofasciatus Macq. 

Pangonia inconspicua Wlk. 1848, List, 1: 137. Loc.? B. M. Ricardo (1901) 
records as missing from B. M. Coll. 

Esenbeckia insignis Krober 1931, Zool. Anz., 94: 255. Brazil. B. M. 

Tabanus lagenaferus Macq. 1838, Dipt. Exot., 1 (1): 148. Loc.? Mus. Paris. 

Tabanus longifrons Krober 1930, Dipt. Pat. S. Chile, 5 (2):152. Chile. B. M. 
Apparently not returned by Krober. 

Tabanus longipennis Macq. 1834, Hist. Nat. Dipt., 1: 201. Loc.? Coll. Percheron. 

Chrysops lugubris Macq. 1846, Dipt. Exot., Suppl. 1: 44. Brazil. Coll. Robyns, 
Bruxelles. The description suggests possibility that this was not a ChArysops. 

Tabanus maculipennis Macq. 1834, Hist. Nat. Dipt., 1: 198. Brazil. Coll. Serville. 
A homonym of T. maculipennis Wied. 1828, and Brullé 1832. Not same as 
maculipennis Macq. 1846. Not listed by Krober (1934). 

Tabanus maculipennis Macq. 1846, Dipt. Exot., Suppl. 1:34. Brazil. Coll. 
Spinola. A homonym of maculipennis Wied. 1828, and Macq. 1834, but a 
different species, probably a synonym of Tabanus (Hybomitra) quadripunctata 
and the genotype of Dasyphyrta End. (Krober 1934). 

Tabanus marginenevris Macq. 1855, Dipt. Exot., Suppl. 5:29. Amer. Merid. 

Tabanus microcerus Wlk. 1848, List, 1: 150. Loc.? B. M. 


e 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 74 


Tabanus nigripalpis Macq. 1845, Dipt. Exot., Suppl. 1: 40, pl. 4, figs. 7-8. New 
Grenada. Coll. Bigot. Placed by Kroéber (1931i, 1934) in Catachlorops, near 
rufescens Fab. A specimen, headless, in Mus. Paris labeled as rufescens by 
Macquart does not disagree with the description, which appears to be based 
on two different species. 

Pangonia nigronotata Macq. 1850, Dipt. Exot., Suppl. 4:27. Mexico. Seen 
in Mus. Paris by Philip and discussed by him (1954b). 

Tabanus olivaceiventris Macq. 1847, Dipt. Exot., Suppl. 2:18. Brazil, Para. Coll. 
de Villiers. 

Tabanus opulentus Wk. 1848, List, 1: 148. Loc.? B. M. 

Tabanus ornatifrons Krober 1930, Dipt. Pat. S. Chile, 5 (2): 153. Chile. B. M. 
Apparently not returned by Krober. 

Pangonia planiventris Macq. 1850, Dipt. Exot., Suppl. 4: 23. Mexico. Placed by 
Kréber (1934) as a possible synonym of saussurei Bell. In Nearctic Coll. at 
Mus. Paris, where seen by Philip (1954b). 

Tabanus planus Wik. 1850, Dipt. Saund., 1:61. Loc.? B. M. 

Pangonia prasiniventris Macq. 1846, Dipt. Exot., Suppl. 1: 29. Colombia. Coll. 
Fairmaire. A specimen ex Coll. Bigot in B. M. is determined by Macquart 
and can be made neotype if the original fails to turn up. 

Tabanus pubescens Wlk. 1854, List, 5: 220. Brazil. B. M. A homonym of T. pu- 
bescens Strom 1768, and Macquart 1847. 

Tabanus pudens Wik. 1850, Dipt. Saund., 1:36. Brazil, Rio de Janeiro. Lutz 
(1907) = occidentalis Wied. Listed as (Neotabanus) by Kroéber (1934), but 
he saw no specimens. 

Tabanus redactus Wlk. 1850, Dipt. Saund., 1: 66. Loc.? B. M. 

Tabanus ruber Macq. 1845, Dipt. Exot., Suppl. 1:42. Mexico. Coll. Guerin. 
A homonym of 7. ruber Thunb. 1827. Renamed subruber by Bellardi, 1850. 
May be among Nearctic species in Mus. Paris. 

Pangomia rufa Macq. 1838, Dipt. Exot., 1 (1): 110. Lima (Peru). Coll. Serville. 
There are specimens determined by Bigot and Krober in B. M. 

Tabanus scutellatus Macq. 1839, Dipt. Exot., 1 (2): 186. Loc.? Mus. Paris. 
Kroéber (1934) claims to have seen the type and says it = macula Macq., but 
I was unable to find it and doubt the synonymy. In any case scutellatus has six 
years’ priority. 

Tabanus secundus Wik. 1848, List, 1: 180. Loc.? B. M. 

Tabanus subsenex Wk. 1850, Dipt. Saund., 1: 38. S. America. B. M. Lutz 

- (1907) = triangulum Wied. 

Tabanus surinamensis Macq. 1838, Dipt. Exot., 1 (1) : 132. Surinam. Coll. Ser- 
ville. The description suggests T. nebulosus de Geer. Kréber (1934) placed in 
Tabanus with a query. 

Pangonia translucens Macq. 1845, Dipt. Exot., Suppl. 1: 26. Brazil. Coll. Guerin. 
The species has long been placed in Esenbeckia, (Krober, 1934; Fairchild, 
1942d). 

Tabanus trifarius Macq. 1838, Dipt. Exot., 1 (1):144. Chile. Coll. Serville. 
Genotype of Archiplatius End. Krober (1934) as Agelanius. 

Tabanus trifasciatus Macq. 1834, Hist. Nat. Dipt., 1: 204. Loc.? Coll. Percheron. 
T. austeni var. trifasciatus Szil. 1915, appears to be a homonym. 

Tabanus variventris Macq. 1847, Dipt. Exot., Suppl. 2: 18. Brazil. Coll. Spinola. 
There is a specimen in Mus. Paris labeled by Macquart as “T. erythrogaster 
Colomb.” and also as “7. variventris J. Macq. 2e. Suppl. Colombie.” The 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


former appears to be a MS. name. This specimen is a color variant of hirti- 
tibia Wlk., but is not the true type of variventris, which was from Brazil and 
probably different. 

Tabanus venustulus Kréber 1930, Dipt. Pat. S. Chile, 5 (2) :155. Chile. B. M. 
Apparently not returned to B. M. after description. 

Tabanus viridiflavus Wik. 1850, Newman’s Zoologist, 5, App., p. [xvi. Brazil. 
B. M. Lutz (1907) = mexicanus L. The description indicates Chlorotabanus 
inanis Fab. in the synonymy of which the name has long been placed. 

Pangonia xanthopogon Macq. 1838, Dipt. Exot., 1 (2): 179. Brazil, Goyaz. Mus. 
Paris. Lutz (1909) regarded a specimen in Mus. Paris labeled fulvibarbis, 
the Latin equivalent of the Greek xanthopogon, as probably the type. I did not 
know of this at the time and did not see the specimen. 


It is to be noted that most of the types of Macquart which could not 
be found in the Paris Museum are species he received from other col- 
lectors. Thus, of four species from the Marquis de Spinola, none was 
found; of eight from the Serville collection, none is in Paris, and one 
was found in the British Museum. Species described from the collec- 
tions of de Villiers, Robyns, Guerin, and de Breme, six in all, are all 
missing from the Paris Museum. The Bigot material and most of the 
Fairmaire species are in the British Museum save for a few appar- 
ently lost. Of the 50 species I have been able to check, described from 
the Paris Museum collections, only 6 are apparently missing, and these 
were mainly species without locality which may have been placed 
elsewhere in the collections. 

Dr. C. B. Philip, who has examined a large proportion of the types 
discussed here, as well as many of Wiedemann’s types, has very gen- 
erously gone over the manuscript of this paper and made numerous 
suggestions and corrections. He was also fortunate in discovering 
certain types of Macquart’s species in Paris which I did not see, and 
I append here the information on them he has furnished, as his obser- 
vations cannot properly be placed with my own. 

Pangonia bicolor Macq. 1850. 2 ¢ cotypes in Paris confirm synonymy with 

P. semiflava Wied. 

Chrysops frontalis Macq. 1838. 1 ¢ type in Paris with dichoptic eyes and pecul- 
iar wing pattern. 

Tabanus fulvilateralis Macq. 1838 is the same as the Nearctic T. (Hybomitra) 
haemaphorus Marten 1882, and agrees with a specimen from Alaska in Dr. 

Philip’s collection. 


Tabanus scutellatus Macq. 1839. 2 2 cotypes in Paris are not the same as 
T. macula Macq. 1845. 


In the following list the names are arranged alphabetically. Each 
name is followed by a condensed citation to its original proposal with 
the generic name under which it was proposed in parentheses, and the 
location of the material studied indicated by “B. M.” for the British 





NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 9 


Museum, “M. P.” for the Muséum d’Histoire Naturelle in Paris. 
Names of which I have seen the types are preceded by an asterisk (*). 
Where I was able to match the type with a specimen in my possession 
I have placed an (H) after the location of the type. Some of these 
specimens are the property of the U. S. National Museum or the 
Museum of Comparative Zoology at Harvard University, and will be 
returned to those collections. The remainder are in my collection and 
will be retained for the time being. 

Names which appear to be valid are in boldface; all others in 
italics. Synonymy believed to be new is indicated by (N. S.). In the 
case of confirmation of older synonymy, an attempt has been made to 
indicate the earliest authority for it, although in some cases this has 
not been possible. Since I have not examined the types of Wiede- 
mann’s species, cases where his names appear to be the earliest valid 
ones are accepted from the literature. 

The supraspecific categories of Neotropical Tabanidae are still in a 
chaotic condition, though Dr. I. M. Mackerras has in preparation a re- 
vision of the whole family, and I have been privileged to see his manu- 
script. Most of the names here used are in the sense of Krober’s 
(1934) catalog, the exceptions being the following: Fidena includes 
Fidena and Melpia of the catalog. Scaptia s.s. includes Osca and Cal- 
hosca. Scaptia subgenus Pseudoscione includes Listriosca, Listrapha, 
Parosca, Listraphella, and probably Lilaena of the catalog. Stenotabanus 
includes all the small Tabanus-like species with bare subepaulets and at 
least some of the species placed in Stypommuia and Stypommisa in the 
catalog. Aegialomyia is treated as a subgenus of Stenotabanus. Dasy- 
basis is used for the species placed in Agelanius in the catalog, follow- 
ing Stone (1944), the criterion being bare subepaulets, generally 
broad frons, and often pilose eyes, the species being mainly Chilean. 
Dasychela End. is used for those species with generally hairy eyes, 
long antennal tooth, bare subepaulet, and fleshy labella placed in 
Dicladocera in the catalog. The bulk of the species are from Colombia, 
Ecuador, and Peru. Dicladocera is retained for the mainly southern 
Brazilian forms, which differ in having usually bare eyes and more or 
less sclerotized labella. Dichelacera includes Catachlorops, Amphichlo- 
rops, and Psalidia as subgenera. Tabanus includes Allioma and Che- 
lommia of the catalog, and, in fact, all Tabaninae with setose subepau- 
lets except Leucotabanus. Lophotabanus, Hybomitra, Philipotabanus, 
and Hemuchrysops are retained as subgenera. Future careful work 
will no doubt modify many of the above categories, but this is not the 
place for detailed discussion of generic concepts. If the species ap- 
pears to have been correctly placed generically by the original de- 


Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


scriber, I have not added any generic placement. If not, I have added 
what appears to me the correct generic name in boldface type. In 
the cases where a good modern description or discussion clearly refer- 
ring to the species exists, I have added the appropriate reference to it. 


*adustus Wk. 1850, Dipt. Saund., 1:34. (Tabanus) B. M.(H) = *rubigini- 
pennis Macq. 1845, Kréber (1934, 1940) as Dicladocera. 

*albicans Macq. 1845, Dipt. Exot., Suppl. 1:37. (Tabanus) B. M.(H). Not 
Macq. 1833 or 1834. Not *L. canithorax Fchld. 1941. = Leucotabanus 
exaestuans L. 1767 (N. S.). 

*albidicollis Macq. 1850, Dipt. Exot., Suppl. 4:32. (Tabanus) B. M.(H) = 
T. importunus Wied., Kréber (1934). Fairchild (10942f). 

*albidocinctus Big. 1892, 5:686. (Tabanus) B. M.=Stenotabanus. Krober 
(1934) as Leucotabanus. 

*albifasciata Macq. 1845, Dipt. Exot., Suppl. 1: 28. (Pangonia) B. M. = Scione. 
Kréber (1934) has it albofasciata in error. 

*albifrons Macq. 1838, Dipt. Exot., 1(1):108. (Pangonia) M. P.(H)= 
(Pseudoscione). Kroéber (1934) as Lilaeina Borg. Genotype. 

*albipectus Big. 1892, 5: 611. (Mycteromyia) B. M. Kroéber (1933b) = Fidena 
lingens Wied. 

*albitarsis Macq. 1850, Dipt. Exot., Suppl. 4:36. (Lepiselaga) M. P. Krober 
(1920). 

*albithorax Macq. 1838, Dipt. Exot., 1 (1): 107. (Pangonia) M. P.= Scaptia. 
Kroéber (1930k, 1934) as Osca. 

*alboater Wik. 1850, Zoologist, 8, App., p. Ixvii. (Tabanus) B. M.(H) = 
*T. atricornis Big. 1892. = T. albibarbis Wied., Krober (1932b). *7. angusti- 
frons Macq. 1847, *T. senior Wlk. 1850, and Chelommia amazonensis Barr. 
1949, are probably all variants. Alboater and atricornis agree precisely with 
each other but differ from albibarbis in open first posterior cell and less 
fumose wings. Senior agrees best with albibarbis det. Krob. Angustifrons 
has clear wings and slightly different antennae and palps. 

*albohirtus Wlk. 1857, Trans. Linn. Soc. London, 17: 338. (Tabanus) B. M.= 
Dasybasis. Krober (1930i, 1934) as Agelanius. 

*albomaculatus Wik. 1854, List, 5: 207. (Tabanus) B. M.(H) =*T. discifer 
WIlk. 1850. Krober (1931c) as Gymnochela discifer; (1934) as Chelommia 
discifer. 

*albomarginata Krob. 1930, Zool. Anz., 90 (3-4):76. (Spheciogaster) B. M. 
Fairchild (1939) as Acanthocera. 

*albopicta Big. 1892, 5: 633. (Dichelacera) B. M.(H) =—Catachlorops potator 
Wied., Lutz (1907) = *Dichelacera marmorata Big., Lutz (1907). 

*albovarius Wlk. 1854, List, 5: 206. (Tabanus) B. M.(H) =*T. unicinctus 
WIk. 1857 = *Leucotabanus Jeuconotum Fchld. 1941 (N.S.). 

*albovittatus Krob. 1930, Dipt. Pat. S. Chile, 5, fasc. 2:146. (Therioplectes) 
B. M.=*Dasybasis scutulatus Kréb. 1930. (N.S.) Krober (1934) as 

siladynus. 

*alcis Will. 1806, Trans. Ent. Soc. London, pt. 3, p. 302. (Tabanus) B. M.= 
Dichelacera, Bequaert (1940). 

*alene Towns, 1895, Trans. Amer. Ent. Soc., 22: 59. (Tabanus) B. M. = *Steno- 
tabanus parallelus Wlk., Bequaert (1940). 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD II 


*alteripennis Wik. 1860, Trans. Ent. Soc. London, 5: 275. (Tabanus) B. M.(H) 
= T. caliginosus Bell. 1859, Hine (1925). Fairchild (1953) as (Philipo- 
tabanus). Philip (1954). 

*altivagus O. S. 1886, Biol. Cent.-Amer., Dipt., 1:45. (Chrysops) B. M. 

*amabilis Wik. 1848, List, 1: 154. (Tabanus) B. M.=T. (Hybomitra) quad- 
ripunctatus Fab. 1805. Krober (1934). 

*angustifrons Krob. 1930, Zool. Anz., 90(3-4):74. (Diachlorus) B. M.= 
? D. ochraceus Krob. 1928. Not Macquart 1850. Type appears teneral and 
may be composite. (N.S.) 

*angustifrons Macq. 1847, Dipt. Exot., Suppl. 3:12. (Tabanus) B. M.(H) = 
? T. albibarbis Wied. See remarks under alboater Wlk. (N.S.) 

*angustifrons Towns. 1895, Trans. Amer. Ent. Soc., 22: 59. (Tabanus) B. M.= 
T. townsendi Johns., nom. nov. = T. lucidulus Beq. 1940 in part.=T. lu- 
ctdulus Fchld. 1951. 

*angustus Macq. 1838, Dipt. Exot. 1 (1):136. (Tabanus) M. P.(H) =*T. 
polytaenia Big. 1892= ? T. duplovittatus Rond. 1868, Brethes (1910). 
*apicalis Macq. 1847, Dipt. Exot., Suppl. 2:20. (Tabanus) B. M.(H). Two 
specimens labeled type and paratype, the first bearing a Macquart label. The 
species was described as headless, but the present type has its own head. The 
paratype has head glued on. It is possible that Bigot or someone else glued 
a head of this group onto the true type and switched the label to the more 
perfect specimen. The type above agrees with “bigoti var. B,” the paratype 

with “bigoti var. A” of Fairchild (1942). —=T. bigoti Bell., nom. nov. 

*approximans Wlk. 1848, List, 1: 108. (Chrysops) B. M.(H) = Diachlorus 
ferrugatus Fab. 1805. 

*ataenia Macq. 1838, Dipt. Exot., 1 (1): 156. (Diabasis) M. P. This specimen 
is probably the one from Para mentioned in original description. = Diachlorus 
curvipes Fab. 1805, Lutz (1907). Type headless. 

*atricornis Big. 1892, 5: 683. (Tabanus) B. M.(H) =*T. alboater Wik. (q. v.) 
= ? T. albibarbis Wied. Not T. atricornis Meig. 1838. 

*atrifera Wlk. 1860, Trans. Ent. Soc. London, 5: 272. (Pangonia) B. M.= 
Pangonia haustellata (Fab.) 1781. Palearctic. Philip (1954). 

attenuatus Wlk. 1848, List, 1: 159. (Tabanus) B. M. Probably Oriental; does 
not appear to be Neotropical, and may not be the true type. 

*aureopygia Krob. 1931, Zool. Anz., 95:24. (Fidena) B. M. 

*aquribarbis Macq. 1847, Dipt. Exot., Suppl. 3:12. (Tabanus) B. M.(H) = 
*Dasychela macula Macq. 1845. Krober (10934, 1940) as Dicladocera macula. 

*aurimaculata Macq. 1838, Dipt. Exot., 1(1): 109. (Pangonia) M. P.(H) = 
Fidena, Krober (1934). 

*auripes Ric. 1900, Ann. Mag. Nat. Hist., ser. 7, 5: 176. (Erephrosis) B. M.(H) 
= Fidena, Kréber (1934). 

*qurisquammatus Big. 1892, 5: 665. (Atylotus) B. M. = Atylotus fulvus Meig. 
Palearctic. (N.S.) Lutz (1907) =T. unicolor Wied. 

*auroguttata Krob. 1930, Zool. Anz., 90(3-4):71. (Chrysops) B. M. Bequaert 
(1944). A distinct species from C. auroguttata pallidifemorata Krob. 

*aurora Macq. 1838, Dipt. Exot., 1(1):142. (Tabanus) M. P. Not T. aurora 
of Lutz (1914, 1918), Bequaert (1926), or Kréber (19209c). Near T. ferreus 
WIk. 1848 and T. impressus Wied. 1828. 

*austem Krob. 1930, Zool. Anz., 86(11-12) : 294. (Tabanus) B. M. Not Tabanus 
(Ochrops) austeni Szil. (1915, Ent. Mitt. Berlin, 4: 100). For Tabanus 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


(Phaeotabanus) austeni Krob. I hereby propose the name Tabanus sannio, 
nom. nov. 

*badia Wlk. 1848, List, 1: 132. (Pangonia) B. M. = Fidena venosa Wied. 1828, 
Krober (1930g) as Sackenimyia venosa; (1934) as Melpia. 

*badia Krob. 1931, Rev. Ent., 1 (4): 402. (Dicladocera) B. M.(H) = Dasy- 
chela. Fairchild (1940b). 

*bahiana Big. 1892, 5: 612. (Pangonia) B. M. = *Pangonia flavescens Ric. 1900a. 
= Esenbeckia vulpes Wied. 1828, Krober (1932a). 

*basalis Wik. 1848, List, 1: 133. (Pangonia) B. M.(H) = Fidena basilaris Wied. 
1828. Lutz (1909) as Phaeoneura basilaris Wied. Krober (1933b) as F. basi- 
laris Wied. Not Pangonia basalis Macq. 1847. Palearctic. Not listed by 
Kroéber (1934). 

*basi-rufus Wlk. 1850, Dipt. Saund., 1: 32. (Tabanus) B. M.(H) = Dasychela, 
Bequaert and Renjifo (1946). Krober (1940) as Dicladocera. 

*basi-vitta Wlk. 1850, Zoologist, 8, App., p. Ixviii. (Tabanus) B. M.(H) = 
*T. viduus Wik. 1850. = *T. bitinctus Wilk. 1857. =? T. marginenevris 
Macq. 1855. Type not found, but specimens det. Bigot in B. M. agree with 
basivitta, (N.S.) 

*bicolor Big. 1892, Wien. Ent. Zeitg., 11: 162. (Bolbodimyia) B. M.(H). Stone 
(1954). 

*bicolor Big. 1892, 5: 636. (Stibasoma) B. M. = Stibasoma triste (Wied.) 1828. 
Lutz (1915). Kroéber (1934). Not Tabanus tristis Fab. 1708. 

*bifascies Wlk. 1848, List, 1: 191. (Dichelacera) B. M.(H). Barretto (1949b) 
as Rhamphis End. 

*bifenestratus O. S. 1886, Biol. Centr.-Amer., Dipt., 1:52. (Tabanus) B. M. 
Philip (1952). 

*bipartitus Wlk. 1848, List, 1:158. (Tabanus) B. M.(H) =*T. (Lophota- 
banus) oculus Wlk. 1848, p. 157. Kroéber (1934). Fairchild (1942b) as 
Bellardia. 

*bitinctus Wlk. 1857, Trans. Ent. Soc. London, 4 (5) : 123. (Tabanus) B. M.(H) 
= *T, basivitta Wlk. 1850. (N.S.) 

*brasiliensis Ric. 1901, Ann. Mag. Nat. Hist., ser. 7, 8:314. (Chrysops) 
B. M.(H) =*Chrysops incisa Macq. (q. v.). (N.S.) 

*brunnipes Krob. 1929, Encycl. Ent., Dipt. 5: 116. (Stenotabanus) B. M.= 
Stenotabanus, with bare subepaulets. The specimen is a paratype. 

*callicera Big. 1892, 5: 686. (Tabanus) B. M. Appears related to Stenotabanus 
pequeniensis Fchld. 1942a. Subepaulets with sparse macrotrichiae. Lutz 
(1907) = ? rubrithorax Macq. 1838. 

*callosus Macq. 1848, Dipt. Exot., Suppl. 3:11. (Tabanus) B. M.(H) = 
? T. trivittatus Fab. Not T. callosus Fairchild 1942c. Type lacks antennae 
and palps and further series may show intergrading with other named forms, 
as in /ineola complex. Philip (1954b). 

*campechianus Towns. 1897, Canadian Ent., 29: 197. (Tabanus) B. M. A 
Tabanus nearest yucatanus Towns. but quite distinct. 

*carbo Macq. 1850, Dipt. Exot., Suppl. 4:33. (Tabanus) B. M. and M. P.(H) 
= Veprius presbiter Rond. 1863. Kréber (1929a) as Stypommia. The types 
have hind tibial spurs, bare subepaulets and subcosta and well-developed 
ocelli, so that carbo will replace Rondani’s specific name. (N.S.) 

*castanea Big. 1892, 5: 633. (Dichelacera) B. M.(H) —Dicladocera. Krober 
(193Ic) as var. of Gymnochela satanica Big.; (1934) as syn. of Chelommia 





NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 13 


satanica Big. Not Chelommia castanea Barr. 1949a. Subepaulets bare. Type 
agrees quite well with a specimen det. Fairchild as Dicladocera unicolor Lutz 
from the description. Barretto (1948) as syn. of Amphichlorops satanica Big. 
= ? Amphichlorops ferruginea Barr. 1948. 

*castanea Surc. 1919, Mes. Arc. Mérid. Equat. Amér. du Sud, 10: 222. (Ere- 
phopsis) M. P. = Fidena castaneiventris Krob. 1934, nom. nov. Not Fidena 
castanea (Perty) 1830. 

*caustica O. S. 1886, Biol. Centr.-Amer., Dipt., 1:44. (Pangonia) B. M.= 
Esenbeckia wiedemanni Bell., Krober (1934). Philip (1954a) as a distinct 
species. 

*chilensis Macq. 1838, Dipt. Exot., 1(1):145. (Tabanus) M. P.(H) = Dasy- 
basis. Krober (1934) as Agelanius. 

*chionostigma O. S. 1886, Biol. Centr.-Amer., Dipt., 1:54. (Tabanus) B. M.= 
Stibasoma, Fairchild (1o4ob). Labels on type do not agree with original 
description. 

*cinerascens Big. 1892, 5: 610. (Mycteromyia) B. M. Krober (1930f). 

*cingulifer Wik. 1857, Trans. Ent. Soc. London, 4(5) : 123. (Tabanus) B. M.= 
Leucotabanus exaestuans Linn. 1767. 

*claripennis Big. 1892, 5:675. (Atylotus) B. M.(H) = Tabanus hookeri Knab 
1915 (N.S.). Original locality given as Australia. Bequaert (1940). 

*clausus Macq. 1847, Dipt. Exot., Suppl. 2:17. (Tabanus) B. M.(H) = Tabanus 
fuscus Wied. 1828. Lutz (1907); Krober (1934) as syn. of Chelotabanus 
fuscus Wied. 

*colombensis Macq. 1846, Dipt. Exot., Suppl. 1:37. (Tabanus) B. M.(H) = 
T. amplifrons Krob., Fairchild (1942c). =*T. fur Will. 1901. Krober 
(1933a) as syn. of T. occidentalis Linn. (N.S.) 

*columbiensis Krob. 1930, Mitt. Mus. Hamburg, 44:177. (Melpia) B. M.(H)= 
Fidena, Krober (1934); Fairchild (1951a). 

*commixtus Wlk. 1860, Trans. Ent. Soc. London, 5: 273. (Tabanus) B. M.(H) 
= *Tabanus maya Bequaert 1932. Fairchild (1942c). Krober (1934) as syn. 
of T. lineola Fab. (N.S.) 

*communis Krob. 1931, Stett. Ent. Zeitg., 92: 282. (Agelanius) B. M.=*T. fre- 
quens Krob. 1934, nom. nov. Not 7. communis Krob. 1930. = Dasybasis, 
with bare subepaulets. 

*compactus Wik. 1854, List, 5, Suppl. 1: 222. (Tabanus) B. M.(H) = Stibasoma 
fulvohirtum Wied. 1828, Osten Sacken (1886). 

*comprehensa Wik. 1850, Dipt. Saund., 1: 11. (Pangonia) B. M.(H) = Elaphella 
cervus Wied. 1828, Ricardo (1904). 

*confinis Wik. 1848, List, 1: 160. (Tabanus) B. M.=? T. aurilineatus Sch.- 
Stekh. 1926, Oriental. Not T. confinis Zett. 1840. Lutz (1907) = T. taeniotes 
Wied. Kréber (1933a) as (Neotabanus). Not Neotropical, in my opinion. 

*confligens Wik. 1854, List, 5, Suppl. 1: 326. (Tabanus) B. M.(H). Nom. nov. 
pro *T. tenens Wlk. 1850, Zoologist, 8, App., p. Ixv, and 1854, List, 5, Suppl. 
1: 123. Not tenens Wlk. 1850, Dipt. Saund., 1: 49. E. India. = T. cinerarius 
Wied. 1828, Krober (1931¢, 1934) as Chelommia or Gymnochela. 

*conica Big. 1857, Ann. Ent. Soc. France, ser. 3, 5: 278. (Pangonia) B. M.(H) 
= Mycteromyia Philippi, genotype. Krober (1930f). 

*consequa Wlk. 1850, Zoologist, 8, App., p. cxxi. (Tabanus) B. M.(H) = 
? T. lineola var. carneus Bell., Fairchild (1942c). = *7. globulicallosus 
Krob. 1931. = ? T. dorsiger var. pallidefemorata Krob. 1929. = ? T. ochro- 


14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


philus Lutz 1914. This is the small pale form of lineola var. carneus (N.S.). 
Until the relationships in this group can be thoroughly worked out it seems 
better to retain dorsovittatus Macq. (q.v.) as separate. 

*convergens Wik. 1848, List, 1: 198. (Chrysops) B. M. = Diachlorus ferrugatus 
Fab., Ricardo (1904). 

*cornuta Wilk. 1857, Trans. Linn. Soc. London, 17(3):337. (Pangonia) 
B. M.(H) = Rhabdotylus planiventris (Wied.) 1828, Lutz (1907). Krober 
(1934) as Amphichlorops. 

*corone O. S. 1886, Biol. Centr.-Amer., Dipt., 1: 51. (Tabanus) B. M.(H). 

*cribellum O. S. 1886, Biol. Centr.-Amer., Dipt., 1:52. (Tabanus) B. M. = 
Stenotabanus, Stone (1938); Philip (1950). 

*cyaneum Wk. 1848, List, 1: 208. (Hadrus) B. M.=Selasoma tibiale Fab., 
Ricardo (1904). 

*depressa Macq. 1837, Ann. Soc. Ent. France, 6: 429; 1838, Dipt. Exot., 1: 111. 
(Pangonia) B. M. and M. P.(H) =Scaptia lata Guerin 1830. Walker 
(1850), genotype of Osca. Rondani (1863), genotype of Diatomineura. 

*derivatus Wlk. 1848, List, 1: 151. (Tabanus) B. M. = ? (Lophotabanus). 
Type headless, unrecognizable, a male. Name should not be used for speci- 
mens, in my opinion. Described from North America. Not listed by Krober 
(1934). 

*desertus Wlk. 1850, Zoologist, 8, App., p. Ixix. (Tabanus) B. M.(H) = 
*T. nuntins W1k. 1854. = *T. univittatus Macq. 1855. = T. sallei Bell. 1859. 
= T. angustivitta, Fchld. 1942. =*T. (Agelanius) ruficolor Krob. 1934. = 
*T. discifer Big. 1892.—= ? T. dorsiger var. angustivitta Krob. 1929. Bodkin 
and Cleare (1916) (N.S.). 

*detersus Wlk. 1850, Dipt. Saund., 1:31. (Tabanus) B. M. = Stenotabanus 
near pompholyx Fchld. Krober (1930c) as Macrocormus. 

*discifer Wlk. 1850, Dipt. Saund., 1:35. (Tabanus) B. M.(H) =*T. albo- 
maculatus Wik. 1854. Krober (1931c) as Gymnochela; (1934) as Chelommia. 

*discifer Big. 1892, 5:684. (Tabanus) B. M.(H) =*T. desertus Wlk. 1850. 
Not *T. discifer Wlk. 1850. (N.S.) 

*diversipennis Wlk. 1848, List, 1: 165. (Tabanus) B. M.(H) = *Esenbeckia 
fascipennis Macq. 1838, Kréber (1934). 

*diversipes Macq. 1848, Dipt. Exot., Suppl. 3:13. (Diabasis) B. M.=Dia- 
chlorus bicinctus Fab. 1805. Lutz (1913) ; Krober (1928b). 

*dives Wik. 1848, List, 1:166. (Tabanus) B. M.(H) =*Stibasoma flavi- 
ventre Macq. 1847, Lutz (1915). Krober (1934) as var. of fulvohirtum 
Wied. 

*dominicanus Krob. 10931, Stett. Ent. Zeitg., 92: 301. (Tabanus) B. M.= 
(Lophotabanus) Bequaert (1940). 

*dorsoguttata Macq. 1850, Dipt. Exot., Suppl. 4:24. (Pangonia) M. P. = 
Scaptia (Pseudoscione). Krober (1930k) as Parosca; (1934) as Listrapha. 

*dorsovittatus Macq. 1855, Dipt. Exot., Suppl. 5:30. (Tabanus) B. M.(H). 
= T. lineola var. carneus Bell., Fairchild 1942. Lutz (1907) = ? trilineatus 
Latr. Krober (1933a) suggests a var. of carneus. Type in poor shape; agrees 
with pale specimens of var. carneus from Pernambuco, Brazil. (N.S.) See 
under consequa Wlk. 

*ebrius O. S. 1886, Biol. Centr.-Amer., Dipt., 1:49. (Tabanus) B. M.(H) = 
(Philipotabanus) subgenotype. Fairchild (1942f). 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 15 


*edwardsi Krob. 1930, Dipt. Pat. S. Chile, 5(2): 131. (Mycteromyia) B. M.= 
*Mycteromyia hirtipalpis (Big.) 1892. (N.S.) Hack (1953). 

*elongatus Macq. 1845, Dipt. Exot., Suppl. 1:38. (Tabanus) B. M.(H) = 
Dichelacera (Psalidia) vespertina Bequaert and Renjifo 1946, nom. nov. 
Not T. elongatus Wied. 1828. Krober (1934) as Rhamphidommia. Barretto 
(1948) as Amphichlorops. 

enderlein Kréb. 1931, Zool. Anz., 94(9-10):252. (Esenbeckia) B. M. The 
holotype ¢ is in Berlin. The specimens in B. M. det. Krober, 1 ¢ and 2 2, do 
not agree with KroOber’s statements as to sex and locality, are probably not 
the same species (see Lutz and Castro 1935) and are not true types. 

*equatoriensis Sure. 1920, Mes. Arc. Mérid. Equat. Amér. du Sud, 10: 219. 
(Scione) M. P.(H). Kréber (1930j). 

*erebus O. S. 1886, Biol. Centr.-Amer., Dipt., 1:50. (Tabanus) B. M.(H) = 
Astigmatophthalmus satanus Krob. 1931, Stone (1938). Kroéber (1934) as 
syn. of T. alteripennis Wilk. Fairchild (1942f) as Tabanus. 

*eriomera Macq. 1838, Dipt. Exot., 1(1):109. (Pangonia) M. P. = Fidena. 
Lutz (1909) as Epipsila. Structurally close to F. rhinophora Bell. 

*erythraeus Big. 1892, 5:687. (Tabanus) B. M.(H) = T. impressus Wied. 
1828. (N.S.) Not Afylotus erythraeus Big. 1892, 5: 661. Krober (1934) as 
syn. of T. monochroma Wied. 

*erythrocephalus Big. 1892, 5:668. (Atylotus) B. M.(H) = Bolbodimyia, 
Kréber (1930h). Stone (1954). 

*erythronotata Big. 1892, 5: 612. (Mycteromyia) B. M.(H) = Fidena. Lutz 
(1909) as Bombylopsis. Krober (1934) as Melpia. 

*eutaeniatus Big. 1892, 5: 664. (Atylotus) B. M.(H) = ? Tabanus triangulum 
Wied., Fairchild (1942c). Type pale and denuded. Does not wholly agree 
with description. (N.S.) Lutz (1907) suggests syn. of T. ditaenia Wied., a 
species from unknown locality. 

*excelsus Surc. 1919, Mes. Arc. Mérid. Equat. Amér. du Sud, 10: 228. 
(Tabanus) M. P. Not T. excelsus Ricardo (1913, Ann. Mag. Nat. Hist. 
(8)11: 543, India). The type bears the MS. name excelsior Surc., indicating 
an intention on Surcouf’s part to change his homonym; excelsior is hereby 
proposed for excelsus Surc. 1919, not Ricardo 1913. Kroéber (1934) as 
? Stypommia, Bequaert and Renjifo (1946) as Agelanius. = Dasybasis. 

*exeuns Wik. 1850, Dipt. Saund., 1:12. (Pangonia) B. M. = ? (Pseudoscione) 
molesta Wied. 1828. Near seminigra Ric. Enderlein (1925) as Melpia geno- 
type. Not Melpia Wik. Kroéber (1930k) as syn. of Parosca molesta Wied.; 
(1934) as Listrapha molesta Wied. 

*fallax Macq. 1847, Dipt. Exot., Suppl. 2:20. (Tabanus) B. M. Not T. fallax 
Macq. 1845, Africa. Kréber (1932b, 1934) as Chelotabanus fallax. Type 
very dirty. Apparently related to bigoti, but I could not match among my 
material. A new name may be needed when this group is straightened out. 

*fasciata Macq. 1834, Hist. Nat. Ins. Dipt., 1: 194; 1838, Dipt. Exot., 1(1) : 107. 
(Pangonia) M. P. Not P. fasciata Latr. 1811, Egypt. Lutz (1907) as syn. 
of Esenbeckia esenbeckii Wied. 1830. There are 10 2 in Mus. Paris, appar- 
ently conspecific. Related to insignis Krob., filipalpis Will. clari Lutz, and 
nigricorpus Lutz. 

*fasciata Wlk. 1850, Dipt. Saund., 1:68. (Dichelacera) B. M.(H) = *Di- 
chelacera analis Hine 1920. = D. costaricensis Krob. 1931, Fairchild (1940b). 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Osten Sacken (1886), Ricardo (1904), Kréber (1934) as syn. of D. cervi- 
cornis Fab. (N.S.) 

*fascipennis Macq. 1845, Dipt. Exot., Suppl. 1:35. (Tabanus) B. M.(H) = 
(Philipotabanus) Fairchild (1942f). 

*fascipennis Macq. 1838, Dipt. Exot., 1(1): 110. (Pangonia) M. P.(H) = 
*Esenbeckia diversipennis Wlk. 1848, Kroéber (1934). 

*fascipennis Krob. 1930, Zool. Anz., 88(9-10) : 237. (Hemichrysops). The type 
lacks hind legs, but another specimen (Carillo, Costa Rica) in B. M. retains 
them and shows no spurs. Subepaulet sparsely setose. The genus is tabanine 
not pangoniine and close to Philipotabanus, from which it may be separated 
by more protuberant face and sunken frons. The specific name will fall as a 
homonym of Tabanus fascipennis Macq. 1845. The new specific name of 
vecordis is hereby proposed and the species may be known as Tabanus 
(Hemichrysops) vecordis, nom. nov. 

*fenestratus Macq. 1838, Dipt. Exot. 1(1):139. (Tabanus) M. P.(H) = 
Pachyschelomyia notopleuralis Barr. 1950. Brethes (1910) and Krober 
(1934) as Stibasoma. The species is an aberrant one, resembling the African 
Ancala africana in inflated tibiae and general fascies, but with bare subepau- 
lets and sclerotized labella. It seems closest to Phaeotabanus Lutz among 
Neotropical species. Barretto’s name seems tenable. Not T. fenestratus Fab. 
1794. (N.S.) 

*fenestrata Macq. 1845, Dipt. Exot., Suppl. 1:26. (Pangoma) B. M. = 
(Pseudoscione). Lutz (1909) as Diatomineura. Kroéber (1930k) as Parosca; 
(1934) as Listrapha. Very close to seminigra Ric. and longipennis Ric. 

*ferreus Wlk. 1848, List, 1:151. (Tabanus) B. M.(H). Lutz (1907) = ? ? 
D. rufipennis Macq. Krober (1932b) as Chelotabanus ferreus. Very close to 
T. impressus Wied. 1828, but darker. 

*ferrifer Wik. 1850, Dipt. Saund., 1:30. (Tabanus) B. M. (H) = Tabanus 
nebulosus de Geer, Philip (1952). Fairchild (1942f). 

*ferruginea Macq. 1839, Dipt. Exot. 1(2):2095. (Pangonia) M. P.(H) = 
*Esenbeckia illota Will., Fairchild (1942d). Not Tanyglossa ferruginea 
Latr. 1809. The type is like the form treated by Fairchild (1942d) as E. 1. 
enderleint Krob. but with slightly more slender palpi. 

*ferruginosus Wlk. 1850, Dipt. Saund., 1:40. (Tabanus) B. M. = Steno- 
tabanus. Krober (1930c) as Macrocormus. Subepaulet bare. Near *jacu- 
lator Fchld. and fulviventris Macq. 

*filiolus Will. 1901, Biol. Centr.-Amer., Dipt., 1, Suppl. : 261. (Tabanus) 
B. M.(H) = *Tabanus haemagogus Will., Hine (1925). Bequaert (1931). 

*flavescens Ric. 1900, Ann. Mag. Nat. Hist., ser. 7, 5: 174. (Pangonia) B. M.= 
Esenbeckia vulpes Wied., Lutz (1907); Krober (19322). 

*flavinotum Krob. 1934. Rev. Ent., 4(3):300. (Tabanus) B. M.(H). Nom. 
nov. pro T. nigriflavus Krob. (1931, Konowia, 10(4) : 292) = Leucotabanus, 
Fairchild (1941). Not T. nigriflavus Krob. 1930. 

*flavipennis Macq. 1850, Dipt. Exot., Suppl. 4:35. (Diabasis) B. M. The 
type is in poor shape, subepaulets with a few macrotrichiae, antennae 
Tabanus-like. Said to be from Philippine Islands. The species is a Tabanus, 
in my opinion, and the name should not be added to the Neotropical fauna 
without further evidence. It will preoccupy 7. flavipennis Ric. 1914, from the 
Moluccas. 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 17. 


*flaviventris Macq. 1847, Dipt. Exot., Suppl. 3:90. (Tabanus) B. M.(H) = 
Stibasoma, Lutz (1915) with dives Wlk. 1848, as syn. = *T. dives Wk. 
1848; = St. euglossa Lutz 1915 (fig. only, no description) ; = *S¢. stilbium 
Fchld. 1953; = St. mallophoroides, J. Beg. 1944, not Walker 1857; = ? St. 
sulfurotaeniatum Krob. 1921 (N.S.). Trinidad specimens lack complete yel- 
low hind marginal bands on all tergites except the second, but differ in no 
other way from the type and specimens from South and Central America. 

*flavohirta Ric. 1902, Ann. Mag. Nat. Hist., ser. 7, 9: 437. (Scione) B. M. The 
specimen in B. M. is an allotype. The type ¢ was returned to Budapest. 

*fulva Ric. 1902, Ann. Mag. Nat. Hist., ser. 7, 9:435. (Scione) B. M. The 
specimen is a paratype; others returned to Budapest. Krober (1930j). 

*fulvilateralis Macq. 1838, Dipt. Exot., 1(1):137. (Tabanus) M. P. = (Hy- 
bomitra). Eyes pilose, vertical tubercle and setose subepaulets. Possibly 
Palearctic or Nearctic; probably not from Cayenne, as stated. 

*fulvitibialis Ric. 1900, Ann. Mag. Nat. Hist., ser. 7, 5:177. (Erephrosis) 
B. M. = Fidena, Krober (1934). Near rhinophora Bell. but with longer 
palpi and face, more slender antennae and wholly dark abdomen. 

*fulviventris Macq. 1845, Dipt. Exot., Suppl. 1:36. (Tabanus) B. M.(H). 
Kroéber (1930a) as Stypommisa. Subepaulets setose. 

*fulvosericea Kroéb. 1931, Zool. Anz., 95(1-2):26. (Fidena) B. M.(H) = 
*Scione rufescens (Ric.) 1900; = *Scione aureopygia Fchld. 1942. (N.S.) 

*fumifera Wlk. 1854, List, 5, Suppl. 1:323. (Pangonia) B. M. = Fidena, 
Krober (1933b). *F. loricornis Krob. 1931 and Erephopsis pseudoaurimacu- 
lata Lutz 1909 are separable with difficulty, and all three species are from the 
Amazon basin. 

*fur Will. 1901, Biol. Centr.-Amer., Dipt., 1, Suppl. : 261. (Tabanus) B. M.(H) 
= *Tabanus colombensis Macq. Not T. fur Will. 1887 (N.S.). 

*furcata Big. 1892, 5:631. (Bellardia) B. M.(H) = *Dichelacera (Psalidia) 
fulminea Hine. Not D. (Ps.) furcata (Wied.) 1828. This is the light form 
of the species named ocellata by Enderlein and festiva by Hine. See Fair- 
child (1951a) (N.S.). Not listed by Krober (1934). 

*furunculus Will. 1901, Biol. Centr.-Amer., Dipt., 1, Suppl. : 260. (Tabanus) 
Bs MiB). 

*fuscicrura Big. 1892, 5:662. (Atylotus) B. M.(H) = Tabanus subruber 
Bell., Philip (1952). 

*fuscinevris Macq. 1839, Dipt. Exot., 1(2):184. (Tabanus) M. P.(H) = 
*Catachlorops intereuns (Wlk.) 1856 (N.S.). Lutz (1907) noted it was 
probably Neotropical. Kréber (1934) not listed. Oldroyd (1954). 

*fuscipennis Macq. 1847, Dipt. Exot., Suppl. 2:14. (Dichelacera) B. M.(H) = 
Catachlorops psoloptera (Wied.) 1828. Bequaert (1924) genotype of 
Catachlorops. Krober (1934, 1939); Barretto (1946). Type agrees with 
specimen of psoloptera det. Barretto. 

*fuscipennis Macq. 1838, Dipt. Exot., 1(1):156. (Diabasis) M. P. = Leptapha 
fumata (Wied.) 1821 (N.S.). 

*fuscofasciatus Macq. 1838, Dipt. Exot., 1(1): 140. (Tabanus) M. P.(H) = 
*Tabanus hilarii Macq. 1839. 

*fuscus Ric. 1902, Ann. Mag. Nat. Hist., ser. 7, 9: 431. (Erephrosis) B. M. 
Lutz (1907) and Kréber (1933a) as syn. of Fidena winthemi Wied. Differs 
from winthemi in B. M. in broader frons, stouter antennae, wholly pollinose 
clypeus, darker legs and white-haired pleura. Close also to obscuripes Krob. 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


but differs in broader palps, frons, and antennae, and in abdominal coloring, 
which is as in winthemi. Specimen in B. M. a cotype; the other was in 
Budapest. 

*fusiformis Wlk. 1850, Dipt. Saund., 1:19. (Pangonia) B. M.(H) = Esen- 
beckia transluscens Macq., Hine (1920, 1925); Krober (1934). - 

*globulicallosus Krob. 1931, Stett. Ent. Zeitg., 92: 302. (Tabanus) B. M.(H) = 
*Tabanus lineola var. consequa Wlk. 1850. = T. lineola var. carneus Bell. 
small form (N.S.). 

*srandis Ric. 1904, Ann. Mag. Nat. Hist., ser. 7, 14:371. (Dichelacera) 
B. M.(H). Krober (1934). 

*guiterasi Brunetti 1922, Bull. Ent. Res., 13: 401. (Chrysops) B. M.(H) = 
Chrysops flavida Wied. 1821, Bequaert (1940). The original description 
states a d in B. M., a 2 in Berlin, but the specimen now at B. M. is a 2 and 
there is now no d in B. M. 

*guyanensis Macq. 1845, Dipt. Exot., Suppl. 1:41. (Tabanus) B. M.(H) = 
? Tabanus flavibarbis Macq., Dipt. Exot., Suppl. 1:41. = T. flavibarbis of 
Krober (1929c, 1930h) and Bequaert (1926). The type of flavibarbis could 
not be found. The description of guyanensis precedes on the same page 
CRESS). 

*haemagogus Will. 1901, Biol. Centr.-Amer., Dipt., 1, Suppl. : 261. (Tabanus) 
B. M.(H). Bequaert (1931) with *filiolus Will. as syn. 

*halteratus Krob. 1931, Ann. Mus. Hung., 27:344. (Catachlorops) B. M. 
Very close structurally to C. d’almeidai Pech. but darker, more brownish 
facial pollinosity, brown callus and not strongly bicolored fore tibiae. Speci- 
mens in B. M. from Br. Guiana det. luctuosa by Krober are the same, but 
*luctuosa Macq. is a different species. See Barretto (1946). 

*hemiptera Surc. 1912, Bull. Mus. Hist. Nat. Paris, pp. 61-63. (Stibasoma) 
M. P. The specimen is ex coll. Bigot labeled “nov. Holl.” The head is glued 
on and may not belong. Labella fleshy, hence not a Stibasoma. I believe 
near riveti Surc., macula Macq., and minos Schin., and will go in Dasychela. 
I doubt its being Australian. Not listed by Kréber (1934). 

*hilariti Macq. 1839, Dipt. Exot., 1(2): 185. (Tabanus) M. P.(H) = *T. fusco- 
fasciatus Macq. 1838f = ? T. acer Brethes 1910. (N.S.) 

*hinnulus Wk. 1850, Zoologist, 8, App., p. exxii. (Dichelacera) B. M.(H). 
=*Dichelacera marginata Macq. 1847, Ricardo (1904). Kroéber (10934) 
as syn. of marginata Macq. 

*hirtipalpus Big. 1892, 5:619. (Diatomineura) B. M.(H) = Mycteromyia 
= *Mycteromyia edwardsi Krob, 1930. Krober (1930f) erected the genus 
Caenopangonia for this species on the basis of supposedly hairy eyes. The 
type, however, has bare eyes. (N.S.) 

*hirtitibia Wlk. 1850, Dipt. Saund., 1:33. (Tabanus) B. M.(H) = ? Ta- 
banus cinnamomeus Schin. 1868. —= Chelommia fibulata End. 1924. Alli- 
omma Borgm. is based on another species of the same group. Bequaert and 
Renjifo (1946). (N.S.) 

*illota Will. 1901, Biol. Centr.-Amer., Dipt., 1, Suppl. :254. (Pangonia) 
B. M.(H) = *Esenbeckia ferruginea (Macq.) Fairchild 1942. Philip 
(1954a). 

*immaculata Macq. 1838, Dipt. Exot., 1(1):115. (Dichelacera) M. P.(H). 
= Amphichlorops angustifrons Kroéb. 1932 = ? Amphichlorops ferruginea 
Barr. 1948. Lutz and Neiva (1914), Kroéber (1934), and Barretto (1946) 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD I9 


treat as Catachlorops. Specimen in B. M. det. immaculata by Krober agrees 
with type of *Catachlorops fuscipennis Macq. The type of tmmaculata is very 
close to flavus Wied. and vespertina Beq. and will go into Amphichlorops. 
(N.S.) 

*immaculata Krob. 1930, Stett. Ent. Zeitg., 91(2) : 148. (2Rhinotriclista) B. M. 
= Scione, Kroéber (1934). 

*imponens Wlk. 1857, Trans. Ent. Soc. London, 4: 122. (Tabanus) B. M.(H) 
= Tabanus olivaceiventris Macq. 1847 (N.S.) = *Atylotus pulverulentus 
Big. 1892. Kroéber (1929a) as (Lophotabanus) with pulverulentus as syn. 
Bequaert (1926). 

*importunus Macq. 1847, Dipt. Exot., Suppl. 2:18. (Tabanus) B. M. = (Neo- 
tabanus), Krober 1933. Not T. importunus Wied. 1828. Type very dirty and 
denuded, apparently a member of lineola complex. 

*incertus Macq. 1838, Dipt. Exot., 1(1):151. (Tabanus) M. P.(H) = Tabanus 
nebulosus de Geer 1776 = *T. ferrifer Wlk. 1850. Not T. palpalis var. 
incertus Szil. 1926, East Indies. Blanchard’s (1852) reference of specimens 
from Chile repeated by Kroéber (1934) is certainly an error. (N.S.) 

*incipiens Wlk. 1860, Trans. Ent. Soc. London, 5:275. (Tabanus) B. M. = 
Stenotabanus, Krober (1930h, 1934). Type headless, subepaulets bare. Near 
St. maculifrons Hine, but probably now indeterminable with any certainty. 

*incisa Macq. 1845, Dipt. Exot., Suppl. 1:177. (Chrysops) B. M.(H) = 
*Chrysops brasiliensis Ric. 1901 = C. fulviceps, Krober 1925, Bequaert 1940. 
=? C. fulviceps Wik. 1845. =C. aurofasciata Krob. 1926. (N.S.) Not 
C. fulviceps Lutz 1909. Not C. incisa Fairchild 1942. There are 3 2 cotypes; 
the one bearing Macquart’s hand label has been selected and labeled as lecto- 
type. The others are different species. 

*incisuralis Macq. 1847, Dipt. Exot., Suppl. 2:12. (Pangonia) B. M. = ? Fi- 
dena opaca (Brethes) 1910 = Fidena albibarba End. 1925, p. 203, not 
Melpia auribarba var. albibarba End. 1925, p. 276 = Fidena abominata Philip 
1941 = ?? Tanyglossa hirsuta Thunberg (1827, Nova Acta R. Soc. Sci. 
Upsala, 9:67, Brasilia). Not Pangonia incisuralis Say 1823. Enderlein 
(1925) as syn. of hirsuta Thunb. and says incisuralis Lutz (1909) = albi- 
barba End. Krober (1933b, 1934) lists as a valid species of Fidena and saw 
type. Castro (1945) identifies incisuralis of Lutz (1909) with longipalpis 
End. 1925. All specimens seen by Lutz, Enderlein, Brethes, Castro, and my- 
self are from southern Brazil and Argentina. The eyes are practically bare 
and frons with a pair of low bosses at base. The correct name for this 
species must await examination of Brethes and Thunberg’s types, if still in 
existence, 

incompleta Macq. 1845, Dipt. Exot. Suppl. 1:27. o 9; 1850, op. cit. Suppl. 
4:25. (Pangonia) M. P. Only a female remains in Paris and its labeling 
indicates it may not be the studied in 1845. The description was mainly 
based on the male, which seems to have been a different species, as noted by 
Schiner (1868) and Szilady (1926). Krober (1930j) as Scione, but his 
description indicates a different species. Schiner (1868) as genotype of 
Diclisa. The specimen in Mus. Paris is very close to *Sc, minor Macq. (q.v.), 
but with frons a little wider, palps, subcallus, and legs uniformly brown, no 
median black patches on abdomen. It would seem that the validity of the 
name must rest on the description of the ¢ rather than the specimen in Paris. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


*inconspicuus Wik. 1848, List, 1:171. (Tabanus) B. M. = Chlorotabanus 
inanis Fab., Kroéber (1930c). 

*indecisus Big. 1892, 5: 666. (Atylotus) B. M.(H) = Tabanus = *Atylotus 
simplex Big. 1892, p. 667. Not *Tabanus simplex Wik. 1850. Kréber (1934) 
as Tabanus. (N.S.) 

infuscatipennis Macq. MS., Sure. 1919, Mes. Arc Mérid. Equat. Amér. du Sud, 
10(2): 230. (Tabanus) M. P. 1 @ specimen under this name does not agree 
with description of T. ruber Macq., for which it is supposed to be a substitute 
name, or with specimens det. as infuscatipennis by Bequaert. The specimen 
in Paris has no labels on it, but stands under a box label reading “T. infus- 
catipennis Cat. Mus.” with “Colombie” added in pencil. Contrary to Bequaert 
and Renjifo (1946) no description appears to have been based on this speci- 
men. The specimens upon which Surcouf based his description of T. ruber 
Macq. were not found. Kréber (1934) does not list. 

*inornatus Wik. 1848, List, 1: 199. (Chrysops) B. M.(H) = Diachlorus 
bivittatus (Wied.) 1828, Ricardo (1901). Kréber (1934). 

*innotescens Wlk. 1854, List, 5, Suppl. : 327. (Tabanus) B. M.(H) = Cryp- 
totylus pallidipalpis Stone (1944) = Tabanus aurora, Bequaert (1926) and 
Kroéber (1929c). Not *7. aurora Macq. (N.S.) 

*intereuns Wlk. 1856, Dipt. Saund., 1, pt. 5: 450. (Tabanus) B. M.(H) = 
*Catachlorops fuscinevris (Macq.) 1839. Krober (1934). Barretto (1946). 
(N.S.) 

*interruptus Macq. 1838, Dipt. Exot., 1: 156. (Diabasis) M. P. = Diachlorus 
immaculatus (Wied.) 1828, Lutz (1913). Kroéber (1928b). 

*jamaicensis Newst. 1900, Ann. Trop. Med. Parasit., 3: 465. (Atylotus) 
B. M.(H) = Stenotabanus (Aegialomyia), Fairchild (1951a). Bequaert 
(1940) as (Stenotabanus). 

*latipalpis Macq. 1850, Dipt. Exot., Suppl. 4:25. (Pangonia) M. P.(H) = 
(Pseudoscione). Ricardo (1900) as Diatomineura. Enderlein (1922, 1925, 
1929) as Listrapha genotype. Kréber (1930f) as Parosca; (1934) as 
Listrapha. 

*lativentris Macq. 1838, Dipt. Exot. 1(1):153. (Tabanus) M. P.(H) = 
Rhabdotylus planiventris (Wied.) 1828. Blanchard, in Gay (1852) records 
from Chile, probably in error. Kréber (1934) species incert. sedis. (N.S.) 

*Jativitta Wik. 1848, List, 1: 184. (Tabanus) B. M. = Tabanus obsoletus 
Wied. 1828, Lutz (1907); Krober (1934). Type headless. 

*leucothorax Ric. 1900, Ann. Mag. Nat. Hist., ser. 7, 5: 179. (Diatomineura) 
B. M.(H) = (Pseudoscione). Krober (1930f) as Parosca; (1934) as 
Listrapha. 

*limbatus Big. 1892, 5:642. (Therioplectes) B. M.(H) = *Dichelacera uni- 
fasciata Macq. 1838, Brethes (1910); Kroéber (1934); Barretto (1949b). 
*limbithorax Macq. 1855, Dipt. Exot., Suppl. 5:22. (Pangonia) B. M. = 
Scaptia. Krober (1930f) as Parosca; (1934) as Listrapha. Ferguson (1924) 
concluded on the basis of comparison by Austen with type of limbithorax 
that niveovittata Ferg. and Henry was a synonym and hence the species 
Australian. Although Krober (1930f, 1934) treated the species as Neotropi- 
cal, he saw no material other than the type. It should be excluded from the 

Neotropical fauna. 

*limonus Towns. 1897, Ann. Mag. Nat. Hist., ser. 6, 20:21. (Tabanus me-xi- 

canus var.) B. M.(H) = *Cryptotylus longiappendiculatus Macq. 1855. 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 21 


Knab (1916) as syn. of Iuteoflavus Bell. Not Cryptotylus limonus Fairchild 
(1940a). (N.S.) 

*litigiosus Wlk. 1853, Dipt. Saund., 1:37. (Tabanus) B. M.(H) = Phaeo- 
tabanus Lutz and Neiva (1914); Bequaert (1924) genotype of Phaeota- 
banus. The @ in B. M. is now headless but agrees with current interpretations 
of the species and with Krober’s (1930b) description and figs. It should be 
taken as lectotype. The ¢ is a different species, unknown to me. 

*lividus Wik. 1848, List, 1: 162. (Tabanus) B. M.(H) = Tabanus importunus 
Wied. 1828. Kréber (1934) as syn. of T. viduus Wlk. (N.S.) 

*longiappendiculatus Macq. 1855, Dipt. Exot. Suppl. 5:32. (Tabanus) 
B. M.(H) = Cryptotylus. = Tabanus luteoflavus Bell. 1850. = *T. purus 
Wilk. 1860. = *T. mexicanus var. limonus Towns. 1897. = T. pallidus Krob. 
1930. = T. pallidulus Kroéb. 1934. Krober (1934) as (Macrocormus). 
(N.S.) 

*longipalpis Macq. 1848, Dipt. Exot., Suppl. 3:9. (Pangonia) B. M. = 
Histriosilvius Krober 1930d genotype; redescribes and figures type. Lutz 
(1909) as Esenbeckia; Ricardo (1900a) as Diatomineura; Enderlein (1925) 
as Protosilvius. 

*longipennis Ric. 1902, Ann. Mag. Nat. Hist., ser. 7, 11: 433. (Diatomineura) 
B. M. = (Pseudoscione) Lutz et al., 1918; Fairchild (1951a) genotype of 
Pseudoscione; Enderlein (1922) ; Krober (1930k) as Listriosca. 

*longirostris Macq. 1847, Dipt. Exot., Suppl. 2: 12. (Pangonia) B. M.(H) = 
Fidena nigripes (V. Roder) 1892, nom. nov. Not Pangonia longirostris 
Hardwicke 1825. Kroéber (1933b) as longirostris. = ? Erephopsis brevistria 
Lutz 1909. The type also agrees with specimens in B. M. det. awrifasciata 
End. by Krober. 

*loricornis Krob. 1931, Zool. Anz., 95(1-2):32. (Fidena) B. M.(H) = *Pan- 
gonia basalis var. Wlk. 1854, List, 5, Suppl. 1: 322, not *basalis Wlk. 1848. 
Ricardo (1900a) says Walker’s second basalis 1854 not same as his first. 
Kroéber’s type of loricornis is the second specimen discussed by Ricardo, not 
the type of Walker’s 1854 description, though I believe the two are con- 
specific. 

*lucidulus Wlk. 1848, List, 1: 188. (Tabanus) B. M. = *Tabanus obliquus 
Wilk. 1850. Not T. lucidulus, Fchld. (1951a) and not T. lucidulus Austen in 
litt., Bequaert (1940), the latter = T. obumbratus Beq. 1940. The synonymy 
of the three Jamaican species of this group appears to stand as follows: (1) 
*T. lucidulus Wlk. 1848 = *T. obliquus Wik. 1850. = T. lucidulus Krob. 
1930. (2) T. townsendi Johns. = *7. angustifrons Towns. not Macq. = 
*T. Iucidulus Bequaert in part 1940. = *T. lucidulus Fchld. 1951. (3) 
T. obumbratus Beq. 1940 = *T. Iucidulus Austen in litt. The true lucidulus 
does not appear to have been seen by Bequaert. It has a narrower frons, 
small oval callus less than half width of frons and unconnected with the 
median ridge, as figured by Krober. Wings quite heavily fumose. 

*luctuosus Macq. 1838, Dipt. Exot., 1(1):319. (Tabanus) B. M. = Cata- 
chlorops, Kroéber (1934). Barretto (1946) with nigripennis Krob. 1931 as 
synonym. The type from Brazil has wings wholly black, with all cells fenes- 
trate; the specimen from Surinam is different, with apex of wing hyaline. 
Krober (1939) seems to have used a form similar to the Surinam species in his 
redescription of /uctuosa. His nigripennis, from description and figures, is 
composite, the description agreeing fairly well with /uctuosa, the figures not. 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


*lugubris Macq. 1838, Dipt. Exot. 1(1):108. (Pangonia) M. P. = Esen- 
beckia, Lutz (1909). Krober (1934). 

*lutzi Surc. 1921, Gen. Insect., Taban. p. 54; 1923, Ann. Ent. Soc. France, 
91(3) : 242. (Stigmatophthalmus) M. P. = Dasychela. Close to *D. riveti 
Surc. Kroéber (1940) as Dicladocera (Stigmatophthalmus). St. altivagus 
Lutz, the genotype, is quite different. Not listed by Krober (1934). 

*macroceratus Big. 1892, 5: 687. (Tabanus) B. M.(H) = *Pseudacanthocera 
sylveirii (Macq.) 1838. Krober (1934). 

*macrodonta Macq. 1839, Dipt. Exot. 1(2):183. (Tabanus) M. P.(H) = 
Psalidia furcata (Wied.) 1828, Lutz (1907). 

*macula Macq. 1845, Dipt. Exot. Suppl. 1:43. (Tabanus) B. M.(H) = 
*Dasychela auribarbis Macq. 1847. Kréber (1940) as Dicladocera, with 
auribarbis, argyrophorus Schin, 1868 and scutellatus Macq. 1839 as syno- 
nyms. Bequaert and Renjifo (1946) as Dasychela with auribarbis Macq., 
argyrophorus Schin. and submacula Wlk. 1850. The type of scutellatus Macq. 
could not be found, but I doubt its identity with macula. The type of *sub- 
macula Wlk. is somewhat different though closely related. 

*maculifrons Krob. 1931, Stett. Ent. Zeitg., 92: 277. (Agelanius (Archiplatius) ) 
B. M.(H) = *Dasybasis maculiceps (Kréb.) 1934, nom. nov. Not Tabanus 
maculifrons Hine 1907. 

*maculinevris Macq. 1855, Dipt. Exot., Suppl. 5:31. (Tabanus) B. M. Krober 
(1930a, 1934) as Stypommisa. Near *fulviventris Macq. (q.v.). 

*maculipennis Krob. 1929, Zool. Anz., 83(1-4):52. (Stypommia); op. cit., 
(5-8): 117. (Stypommisa) B. M.(H) = *Stenotabanus venosus Big. 
1892. Not Tabanus maculipennis Wied. 1828, Brullé 1832, Macquart 1834, or 
Macquart 1846. 

*maculiventris Macq. 1850, Dipt. Exot. Suppl. 4:33. (Tabanus) M. P.(H) 
= Dasybasis. Kréber (1934) as Tabanus, with rubromaculatus Blanch. 1852 
as synonym. The type is labeled “rubromarginatus Gay Chili,” probably a 
lapsus for rubromaculatus, as well as with its published name, so that all three 
names refer to same specimens. 

*maletectus Big. 1892, 5:664. (Atylotus) B. M. Kréber (1934) as Tabanus. 
The subepaulets are bare, labella fleshy, eyes bare, antentiae missing. Frons 
broad with large black callus filling lower third of frons. Perhaps best in 
Dasybasis, but I do not recognize the species. 

*mallophoroides Wlk. 1857, Trans. Ent. Soc. London, IV, 5: 123. (Tabanus) 
B. M. = *Stibasoma dyridophorum Knab 1913, Bequaert (1944). Lutz 
(1915) as Stibasoma. Krober (1934) as Stibasoma, Dyridophorwm has less 
yellow on dorsum of abdomen but otherwise same. It is possible that festivwm 
Wied. and panamense Curr. are but races or color forms of the same species. 

*manifestus Wik. 1850, Dipt. Saund., 1: 41. (Tabanus) B. M. = Tabanus quin- 
quevittatus Wied. 1821, fide Philip in litt. 

*marginata Macq. 1847, Dipt. Exot., Suppl. 2:14. (Dichelacera) B. M.(H) = 
*Dichelacera hinnulus Wlk. 1850, Ricardo (1904). Lutz (1915). 

*marginatus Macq. 1848, Dipt. Exot. Suppl. 3:172. (Tabanus) B. M. = 
Tabanus. Not Tabanus marginatus Wlk. 1848 (November) Australia. 
Ricardo’s (1901) transference of Walker’s species to Silvius makes it also 
a homonym of marginatus Macq. 1838. Kréber (1934) confused with Silvius 
marginatus Macqy 1838. The type is fragmentary and probably unidentifiable 
with certainty. 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 23 


*marginatus Macq. 1838, Dipt. Exot., 1(1), pl. 19, fig. 1. (Silvius) M. P. = 
*Pseudacanthocera slyveirii (Macq.) 1838, 1, cp. 155. Not Silvius mar- 
ginatus (Wlk.) Ricardo 1901. Both Macquart names almost surely were 
based on the same specimens; the types show only “marginatus,” the name 
appearing on the figure, so it seems probable that the name was changed to 
honor the collector Sylveira shortly before publication but after the plates 
were made. Lutz (1907) as syn. of Acanthocera coarctata Wied. 

*marmorata Big. 1892, 5:634. (Dichelacera) B. M.(H) = Catachlorops 
potator Wied. 1828; Krober (1934). 

*minor Macq. 1847, Dipt. Exot., Suppl. 2:29. (Pangonia) B. M.(H) = 
Scione. Krober (1930j) with aurea Szil. and incompleta Macq. as synonyms, 
but description drawn from other material than the type. His treatment con- 
fusing, and I surmise he intended to synonymize incompleta Schiner, not 
incompleta Macq. with minor Macq. 

*minor Macq. 1850, Dipt. Exot., Suppl. 4: 33. (Tabanus) M. P. = Dasybasis. 
Krober (19301, 1934) as (Agelanius). 

*misera O. S. 1886, Biol. Centr.-Amer., Dipt. 1:47. (Diclisa) B. M.(H) = 
Scione aurulans (Wied.) 1828, Hine (1925). Fairchild (1942d). 

*missionum Macq. 1839, Dipt. Exot., 1(2):186. (Tabanus) M. P. = Dasy- 
basis. Lutz et al. (1918) as Neotabanus. 

*modestus Krob. 1931, Stett. Ent. Zeitg., 92(1-2) : 203. (Tabanus (Agelanius) ) 
B. M. = Dasybasis modestinus Krob. 1934, nom. nov. Not JT. modestus 
Wied. 1828. 

*montium Surc. 1919, Mes. Arc Mérid. Equat. Amér. du Sud, 10: 229. (Ta- 
banus) M. P. = Dasybasis. Bequaert and Renjifo (1946) as Agelanius. 
Near osornoi Beq. and excelsus Surc. 

*multifascia Wlk. 1850, Dipt. Saund., 1:68. (Dichelacera) B. M.(H) = Di- 
chelacera cervicornis (Fab.) 1805; Ricardo (1904). 

*nana Wilk. 1850, Dipt. Saund., 1:11. (Pangonia) B. M. = Pseudelaphella. 
Genotype, Krober (1930e). 

*neglectus Will. 1901, Biol. Centr.-Amer., Dipt., 1, Suppl. : 256. (Chrysops) 
B. M.(H) = Chrysops latifasciata Bell., Hine (1925). Krober (1934) as 
synonym of C. incisa Macq. 

neo-submacula Krob. 1931, Rev. Ent., 1(4):4-9. (Dasyrhamphis) B. M. 
Specimens so det. by Kréber in B. M., not types, which are in Berlin, agree 
with a specimen labeled “Tabanus macula var. n. sp.” by Macquart in B. M. 
This specimen, though labeled as a type, appears not to have formed the 
basis of any published description. *Submacula Wlk. and *macula Macq. are 
conspecific and distinct from neo-submacula det. Krober. 

*nigra Krob. 1931, Rev. Ent., 1(3): 200. (ihamphidommia) B. M. = Amphi- 
chlorops Barretto (1948). 

*nigrifascies Big. 1892, 5:607. (Mycteromyia) B. M. = ? Fidena. The type 
is in execrable condition, but is not Mycteromyia and is probably not from 
India as described. 

*nigripennis Guerin Meneville 1835, Icon. Regne Animal, Insectes, pl. XCVII, 
fig. 2; 1838, Voy. Coquille, Zool., II: 288. (Pangonia) M. P. The specimen 
is labeled “Pangonia nigripennis nob. nov. sp.” in Macquart’s hand and bears 
a Guerin-Meneville label, so may be the type. It is congeneric and possibly 
conspecific with Fidena aureosericea Krob., but is not the same as piceohirta 
Wlk. Nigripennis and piceohirta are placed as synonyms of Sackenimyia 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


venosa (Wied.) 1821 by Kroéber (1930g, 1934). The palpi are very small and 
deeply grooved. = Fidena. 

*nigrithorax Krob. 1930, Zool. Anz., 90(3-4):75. (Diachlorus) B. M. A 9 in 
B. M. from Br. Guiana is close but lacks median black on third tergite. It 
stands over the box label podagricus Fab. 

*nigriventris Krob. 1931, Zool. Anz., 94(9-10) : 254. (Esenbeckia) B. M.(H). , 

*nigrivittata Macq. 1850, Dipt. Exot., Suppl. 4:23. (Pangonia) M. P. = 
Fidena marginalis (Wied.) 1830, Lutz (1907, 1909) ; Kroéber (1933b, 1934). 
The synonymy is probable but not certain as there are several similar species 
here. 

*nigrohirta Wik. 1848, List, 1: 132. (Pangonia) B. M. = Fidena venosa 
(Wied.). Krober (1930¢g). 

*nitens Big. 1892, 5:609. (Mycteromyia) B. M.(H) =Fidena. Krober 
(1933b) redescribes, but his description and figures inconsistent and probably 
composite. Fairchild (1951a) genotype of Bombylopsis Lutz and Ionopsis 
Lutz. 

*nivalis Wlk. 1850, Dipt. Saund., 1:71. (Scepsis) B. M.(H) genotype. Sur- 
couf (1921); Krober (1928a). 

*notabilis Wlk. 1850, Dipt. Saund., 1:18. (Pangonia) B. M. = Esenbeckia, 
Kroéber (1932a) with inframaculata Lutz as synonym. If notabilis and in- 
framaculata are really synonymous, the species will go in Proboscoides Philip, 
but since the type of notabilis lacks the proboscis, this is uncertain. See Fair- 
child (1951a, p. 445). 

*nuntius Wik. 1854, List, 5, Suppl. 1: 207. (Tabanus) B. M.(H) = *Tabanus 
desertus Wlk. 1850. (N.S.) Kréber (1934) says type lost. Philip (1952) 
as synonym of obsoletus Wied., fide Hine MS. 

*obesus Big. 1892, 5: 660. (Atylotus) B. M. The type is headless. The bare 
subepaulets, large size, 14 mm., and general fascies suggest Dasybasis, but I 
know of nothing resembling it from Mexico or Central America. 

*obliquus Wlk. 1850, Dipt. Saund., 1: 28. (Tabanus) B. M. = *Tabanus lu- 
cidulus Wlk. 1848. 

*obscurehirta Krob. 1930, Zool. Anz., 86 (11-12) :284. (Tabanus (Phaeota- 
banus) aphanoptera var.) B. M. Krober (1934) as var. obscuripilis, nom. nov. 
Not T. obscurchirta Ric. 1908. — Phaeotabanus obscuripilis Krab. 

*obscuriventris Krob. 1930, Zool. Anz., 87(1-2):6. (Tabanus (Maérocormus) ) 
B. M.=*Tabanus (Lophotabanus) albocirculus Hine 1907. Krober (1934) 
as obscurigaster, nom. nov. Not T. obscuriventris Krober 1929. (N.S.) 

*ocellus Wlk. 1848, List, 1: 143. (Pangonia) B. M. = Dasychela. Ricardo 
(1900a) noted it was a tabanine. Kroéber (1930h) as Tabanus. Says an arti- 
fact. The species is close to Triceratomyia Bequaert, to Dasychela lim- 
bativena End., and Dicladocera badia Krob., and the detached head now with 
the specimen clearly belongs to it. The antennae are now lost. 

*ochraceus Macq. 1850, Dipt. Exot., Suppl. 4:36. (Diabasis) M. P.(H) = 
Diachlorus bimaculatus Wied. (N.S.) Not D. ochraceus Kréber 1928 
which = ? *D. angustifrons Krob. 

*ochraceus Macq. 1838, Dipt. Exot., 1(1): 149; 1846, op. cit. Suppl. 1: 42. 
(Tabanus) M. P. 2 @ types. One = Cryptotylus unicolor Wied., the 
other = Amphichlorops flavus Wied. (N.S.) 

*oculatus Big. 1892, 5: 606. (Chrysops) B. M. = Chrysops molesta Wied. 1828, 
Ricardo 1901; Krober (1926, 10934). 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 25 


*oculus Wlk. 1848, List, 1:157. (Tabanus) B. M.(H) =Tabanus (Lopho- 
tabanus). Krober (1934) as Bellardia. Fairchild (1942b, 1953). Only the 
Honduras specimen is now in B. M. 

*ornativentris Krob. 1929, Konowia, 8 (2):182. (Hybostraba) B. M.(H) = 
Tabanus nebulosus subsp. Kroéber (1929, I.c.) lists as synonym of Lopho- 
tabanus druyvestetjni Szil., a synonym of nebulosus de Geer. Fairchild 
(1942f) as synonym of ferrifer Wlk. The species is, in my opinion, a darker 
and smaller race of nebulosus. 

*pachycephalum Big. 1892, 5:636. (Stibasoma) B. M. = *Stibasoma chio- 
nostigma O. S. 1886. Fairchild (1940b). 

*pachypalpus Big. 1892, 5:631. (Dichelacera) B. M.(H) = Tabanus. Near 
bigoti Bell. and validus Hine. Same as rufipennis Macq. 1846 (Dipt. Exot., 
Suppl. 1: 41), specimen in B. M., but different from *rufipennis Macq. 1838, 
types in Paris. 

*pallidefemorata Kroéb. 1930, Zool. Anz., 90:72. (Chrysops auroguttata var.) 
B. M. = Chrysops pallidefemorata, Bequaert (1944) ; Pechuman (1037). 
*pallidetinctus Krob. 1930, Zool. Anz., 86(11-12):207. (Tabanus (Phaeota- 
banus)) B. M.(H) = Tabanus (Philipotabanus) caliginosus Bell., Fair- 

child (1953). 

*parallelus Wlk. 1848, List, 1: 187. (Tabanus) B. M. = Stenotabanus, 
Bequaert (1940) with *7. alene Towns. as synonym. 

*patellicornis Krob. 1930, Zool. Anz., 88(11-12):307. (Pseudelaphella) 
BoM. (H): 

*pavida Will. 1901, Biol. Centr.-Amer., Dipt., 1, Suppl. :253. (Pangonia) B. M. 
=Esenbeckia. Enderlein (1925) and Krober (1934) as Ricardoa. Palpi 
long, curved, and deeply grooved outwardly. Philip (1954a). 

*penicillata Big. 1892, 5: 610. (Mycteromyia) B. M. = Fidena. Lutz (1909) 
as Erephopsis, but probably not same species. Krober (1930k) as Melpia; 
(1934) as Fidena. The only black species with reddish legs I have seen. 

*perplexus Wlk. 1850, Dipt. Saund., 1:32. (Tabanus) B. M.(H). Krober 
(1940) as Dicladocera. The subepaulets are setose and the species belongs 
with hirtitibia Wik. in the group treated as Chelommia by Barretto (1949a). 

*peruviana Big. 1892, 5:635. (Dichelacera) B. M.(H) = ? Dasychela lim- 
bativena End. 1925. The subepaulets are bare and the specimen resembles 
Dasychela badia Krober. 

*peruvianus Macq. 1848, Dipt. Exot., Suppl. 3:173. (Tabanus) B. M.(H). 
The specimen has the first posterior cell closed, a fact not mentioned in the 
original description, and may not be the true type. The subepaulets are setose. 
Krober (1931c) as Gymnochela; (1934) as Chelommia. Barretto (19494) 
as Chelommia. Bequaert and Renjifo (1946) as Dichelacera (Psalidia). 

*piceo-hirta Wlk. 1848, List, 1: 132. (Pangonia) B. M. = Fidena venosa Wied. 

*bictipennis Macq. 1834, Hist. Nat., Dipt., 1: 199. (Tabanus). 2 2 ex coll. 
Bigot in B. M. under Acanthocera longicornis Fab. bearing labels “Brazil 
ex coll. Serville” which may be types of pictipennis. As noted by Fairchild 
(1939) the description agrees well with longicornis. (N.S.) Not listed by 
Kroéber (1934). 

*pictipennis Macq. 1850, Dipt. Exot., Suppl. 4: 32. (Tabanus) M. P. = ? Cata- 
chlorops. May be a teneral specimen of a well-known species. Krober 
(1934) as synonym of Tabanus uruguayensis Lynch-Arrib. 1882. Not 
*Tabanus pictipennis Macq. 1834. Not Catachlorops pictipennis Krob. 1931. 


26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


*pictus Krob. 1930, Zool. Anz., 86(11-12) : 264. (Eutabanus) B. M. Subepaulets 
bare. Nearest Stenotabanus but distinct. Bequaert (1939). 

*plangens Wlk. 1854, List, 5, Suppl. 1: 199. (Tabanus) B. M.(H) = Tabanus 
lineola var. Fairchild (1942c). 

*polytaenia Big. 1892, 5: 667. (Atylotus) B. M.(H) = *Tabanus angustus 
Macq. 1838, Krober (1934). 

*praetereuns Wlk. 1850, Dipt. Saund., 1:69. (Dichelacera) B. M. = Cata- 
chlorops. Closest to halteratus Krob. and d’almeidai Pech. Lutz et al. 
(1918). Barretto (1946) in key only. 

*primitivus Wlk. 1848, List, 1:177. (Tabanus) B. M. Lutz (1907) = trivit- 
tatus Latr. Krober (1930c) as Macrocormus, redescribes. Near strigimacu- 
lus Fchld. 1942. 

*pruinosus Krob. 1931, Stett. Ent. Zeitg., 92(2): 276. (Tabanus (Agelanius) ) 
B. M.(H) = Dasybasis. Kroéber (1934) as pruinivitta, nom. nov. Not 
T. pruinosus Big. 1892, or Hine 1900, or Surcouf 1906, or Krober 1929. 

*pubescens Macq. 1847, Dipt. Exot., Suppl. 2:20. (Tabanus) B. M. The type 
is in very bad condition, probably composite and not certainly Neotropical. 
Not 7. pubescens Str6m 1768 or Walker 1854. 

*pulchra Will. 1901, Biol. Centr.-Amer., Dipt., 1, Suppl. : 263. (Dichelacera) 
B. M.(H) = Dichelacera salvadorensis Lutz 1915. (N.S.) 

*pulverulentus Big. 1892, 5: 665. (Atylotus) B. M.(H) = Tabanus olivacei- 
ventris Macq. 1847. (N.S.) = *Tabanus imponens Wlk., Krober (1929). 

*pumiloides Will. 1901, Biol. Centr.-Amer., Dipt., 1, Suppl. : 260. (Tabanus) 
B. M. = Stenotabanus. Fairchild (1953). 

*bunctipennis Macq. 1838, Dipt. Exot., 1(2):185. (Tabanus) M. P.(H) = 
Tabanus (Hybomitra) quadripunctatus Fab. Fairchild (1942f). Not 
T. punctipennis Macq. 1847 (Nearctic), or Stypommisa punctipennis End. 
1925. 

*purus Wlk. 1860, Trans. Ent. Soc. London, 5: 274. (Tabanus) B. M.(H) = 
*Cryptotylus longiappendiculatus Macq. 1855. Hine (1925) as synonym of 
luteoflavus Bell. (N.S.) 

*pyrausta O. S. 1886, Biol. Centr.-Amer., Dipt., 1: 43. (Pangonia) B. M.(H) 
= Fidena rhinophora Bell. 1850, Fairchild (1953). 

*quadrimaculatus Macq. 1845, Dipt. Exot., Suppl. 1:39. (Tabanus) B. M. = 
Dichelacera. Krober (1932a) as Gymnochela; says poeciloptera Schiner 
same; (1934) as Amphichlorops. Close to *testacea Macq., and *alcis Will. 

*reinburgi Surc. 1919, Mes. Arc Mérid. Equat. Amér. du Sud, 10: 233. (Esen- 
beckia) M. P. 

*repanda Wk. 1848, List, 1: 190. (Dichelacera) B. M.(H) = *Dichelacera 
testacea Macq. 1846, Krober (1934). 

*reticulatus Kréb. 1930, Zool. Anz., 86(11-12):298. (Tabanus (Phaeota- 
banus)) B. M.(H) = (Philipotabanus), Fairchild (1942f). 

*riveti Surc. 1919, Mes. Arc. Mérid, Equat. Amér. du Sud, 10: 226. (Tabanus) 
M. P.=Dasychela. Kroéber (1940) as Dicladocera. 

*rubescens Macq. 1838, Dipt. Exot., 1(1): 143. (Tabanus) M. P. = Dasybasis. 
Type headless and denuded, but not same as specimens ex coll. Bigot in B. M. 
so labeled. Near *trigonophorus Macq. but probably now indeterminable. 

*rubescens Big. 1892, 5: 663. (Atylotus) B. M. = Tabanus campestris Brethes 
IQI1, nom. noy. Kréber (1933a) redescribes. Subepaulets setose. Not rubes- 
cens Macq. 1838, Bellardi 1859. 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 27 


*rubidus Macq. 1847, Dipt. Exot., Suppl. 2:19. (Tabanus) B. M. Not T. ru- 
bidus Wied. 1821. Kroéber (1934) as T. rubricosa, nom. nov. Not T. rubri- 
cosa Wulp. 1881. Type headless and excessively dirty, probably now indé- 
terminable. Near *fallax Macq. and albibarbis Wied. 

*rubiginipennis Macq. 1845, Dipt. Exot., Suppl. 1:39. (Tabanus) B. M.(H) 
= *Tabanus adustus Wlk. 1850. Krober (1940) as Dicladocera. Bequaert 
and Renjifo (1946) as Hybomitra. Eyes bare, subepaulet setose. Near 
bigoti Bell. 

*rubiginosa Summers 1911, Ann. Mag. Nat. Hist., ser. 8, 7: 213. (Dichelacera) 
B. M. = (Psalidia). Krober (1934) as Dichelacera. Close to fulminea 
Hine, but with bicolored fore tibiae and widely open cell Rs5. 

*rubribarbis Big. 1892, 5:630. (Atvlotus) B. M.(H) = Dichelacera (Psa- 
lidia) furcata (Wied.) 1828, Krober (1932b). 

*rubrinotatus Big. 1892, 5: 676. (Atylotus) B. M.(H) = *Tabanus guyanen- 
sis Macq. 1845. Kroéber (1934) does not mention. (N.S.) 

*rubripes Macq. 1838, Dipt. Exot., 1(1): 138. (Tabanus) M. P.(H). The 
type is labeled “Sylveira Bresil 1832” though the original description says 
“Cayenne, Sylveira.” All other Sylveira material was from Brazil, so I be- 
lieve Macquart erred here in transcribing the locality. Krober (1930c, 1934) 
as (Macrocormus). The species is close to sorbillans, but distinct. Speci- 
mens in B. M. det. Bigot are sorbillans Wied. 

*rubrithorax Macq. 1838, Dipt. Exot., 1(1):143. (Tabanus) M. P.(H) = 
Stenotabanus. Type headless, but appears to belong in group of pequeniensis 
Fchld. with a few setae on subepaulet. 

*rubriventris Krob. 1930, Mitt. Mus. Hamburg, 44:165. (Osca) B. M. = 
Scaptia. 

*rufa Macq. 1847, Dipt. Exot., Suppl. 2:13. (Dichelacera) B. M. = ? Dichela- 
cera submarginata Lutz 1915. Rondani (1850) transfers to Tabanus and 
changes name to brasiliensis, nom. nov. Lutz (1907) = ? D. januarii var. 
Krober (1934) = januarit Wied. A pale form with reduced wing markings 
and broad frons. 

*rufescens Ric. 1900, Ann. Mag. Nat. Hist., ser. 7, 6:204. (Erephrosis) 
B. M.(H). The holotype is in Budapest; this is a paratype. — *Scione 
aureopygia Fchld. 1942. = *Fidena fulvosericea Krob. 

*ruficornis Krob. 1931, Stett. Ent. Zeitg., 92:287. (Tabanus (Agelanius) ) 
B. M. = Tabanus erythrocerus Krob. 1934, nom. nov. Not T. ruficornis Fab. 
Subepaulet setose. Close to *desertus Wlk. and *johannesi Fchld. 

*rufipennis Macq. 1838, Dipt. Exot., 1(1):138. (Tabanus) M. P. = Diclado- 
cera. Subepaulet bare, labella with sclerotized plate. Very close to *castanea 
Big. and unicolor Lutz, and all may be variants of same species. *Satanica 
Big. also close, but distinct. 

*rufipes Macq. 1850, Dipt. Exot., Suppl. 4:37. (Silvius) M. P. = ? Veprius. 
Type very dirty, 9-10 mm. Hind tibial spurs and ocelli present. Proboscis 
short and fleshy. Frons wider than high, divergent, callus transverse, barlike. 
Third antennal segment with a basal plate and probably four annuli. 

*rufithorax Wlk. 1848, List, 1:165. (Tabanus) B. M. = Catachlorops, 
Krober (1934). Barretto (1946) in key only. 

*rufiventris Macq. 1838, Dipt. Exot., 1(1):145. (Tabanus) B. M. = *Tabanus 
bifloccus Hine 1925. Not T. rufiventris Fab. 1805 or Macquart 1845. Kréber 
(1934) does not list. Bequaert (1940) = ? T. hookeri Knab. 


28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


*rufiventris Macq. 1845, Dipt. Exot., Suppl. 1: 38. (Tabanus) B. M. = *Ta- 
banus simplex Wlk. 1850. Not T. rufiventris Fab. 1805 or Macq. 1838. = 
Tabanus (Hybomitra) indiorum Beg. and Renjifo 1946, nom. nov. pro rufi- 
ventris Macq. 1845. (N.S.) 

*rufohirta Wik. 1848, List, 1: 131. (Pangonia) B. M.(H) = Fidena venosa 
(Wied.), Krober (1930g, 1934). 

*rufopilosus Big. 1892, 5:620. (Veprius) B. M. Ricardo (1901) as Silvius. 
Kroéber (1930d, 1934) lists under Veprius with a query. Hind tibial spurs, 
bare subepaulet and subcosta, fleshy labella, and pilose holoptic eyes. 

*yufus Krob. 1931, Stett. Ent. Zeitg., 92: 287. (Archiplatius) B. M.(H) = 
*Tabanus desertus Wlk. Not 7. rufus Scop. 1763 or Palisot de Beauvois 
1809. Kroéber (1934) as T. (Agelanius) ruficolor, nom. nov. (N.S.) 

*satanica Big. 1802, 5:632. (Dichelacera) B. M.=Dicladocera. Krober 
(1931c) as Gymnochela with castanea Big. as syn.; (1934) as Chelommia. 
The subepaulets are bare and the species congeneric with wnicolor Lutz and 
*rufipennis Macq. It also agrees well with the description of 7. scutellatus 
Macq., but the type of the latter could not be found at B. M. or Paris. Bar- 
retto (1948) as Amphichlorops with castanea Big. as synonym. 

*scapularis Macq. 1847, Dipt. Exot., Suppl. 2:15. (Dichelacera) B. M.(H). 
Hine (1917). Krober (1934). Nearest to *marginata Macq. 

*scutellata Macq. 1838, Dipt. Exot., 1(1):155. (Diabasis) M. P.(H) = 
Diachlorus. Krober’s (1928b) redescription and figure not accurate. Frons 
parallel-sided, callus weakly trifid above. 

*scutellatus Krob. 1931, Ann. Mus. Hung., 27: 348. (Catachlorops) B. M. 
Barretto (1946) keys out with rufescens Fab. 

*scutulatus Kr6éb. 1930, Dipt. Pat. S. Chile, 5(2):143. (Therioplectes) 
B. M.(H) = Dasybasis. = *Tabanus (Therioplectes) albovittatus Krob. 
1930. (N.S.) Krober (1934) as Sziladynus. 

*seminigra Ric. 1902, Ann. Mag. Nat. Hist., ser. 7, 9: 432. (Diatomineura) 
B. M. = (Pseudoscione). Lutz (1907) and Kroéber (1934) as synonym of 
Listrapha tabanipennis Macq. The type of *tabanipennis in Paris is a Fidena. 
Specimens det. tabanipennis by Bigot in B. M. are seminigra Ric. Close to 
*fenestrata Macq. and *longipennis Ric. 

*semisordidus Wlk. 1854, List, 5: 208. (Tabanus) B. M.(H) = Tabanus 
importunus Wied., Krober (1920c, 1034). 

*semiviridis Ric. 1900, Ann. Mag. Nat. Hist., ser. 7, 8:181. (Pangonia) 
B. M.(H) = Esenbeckia prasiniventris Macq., Bequaert and Renjifo 
(1946). Kroéber (1932a, 1934). Described as from Barengo, Old Castile, 
Spain, but the original label is indecipherable and might have been “Vene- 
zuela.” 

*senior Wik. 1850, Dipt. Saund., 1:67. (Tabanus) B. M.(H) = Tabanus albi- 
barbis Wied. 1824. (N.S.) Bodkin and Cleare (1916). Krober (1934) not 
listed. 

*shannoni Kroéb. 1930, Dipt. Pat. S. Chile, 5(2):144. (Therioplectes) B. M. 
= Dasybasis. 

*simplex Wlk. 1850, Dipt. Saund., 1:34. (Tabanus) B. M. = *Tabanus rufi- 
ventris Macq. 1845. Not T. rufiventris Fab. 1805. = Tabanus (Hybomitra) 
indiorum Bequaert and Renjifo 1946. (N.S.) Krober (1934) suggests = 
bifloccus Hine, but confusion here with rufiventris Macq. 1838. Fairchild 
(1942a) under umbraticolus, which is entirely distinct. 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 29 


*simplex Big. 1892, 5:667. (Tabanus) B. M.(H) = *Tabanus indecisus 
(Big.) 1892. Not *T. simplex Wik. (N.S.) Krober (1934) = ? T. (Neo- 
tabanus) signativentris Brethes. There are three cotypes, the lectotype being 
the one with the Bigot name label. The other two are different species. 

*sparsa Wik. 1850, Dipt. Saund., 1:71. (Dichelacera) B. M.(H) = Diclado- 
cera guttipennis (Wied.) 1828, Lutz (1907). Eyes sparsely pilose. 

*stigmaticalis Krob. 1931, Stett. Ent. Zeitg., 92: 2909. (Tabanus) B. M.(H) = 
*Tabanus (Philipotabanus) grassator Fchld. 1953. Type headless. My 
specimen becomes allotype. (N.S.) 

*subfascipennis Macq. 1855, Dipt. Exot., Suppl. 5:35. (Chrysops) B. M. = 
Chrysops variegata de Geer. Krober (1934) as var. of variegata. A large 
dark form, wing apex unusually dark and outer border of crossband concave. 

*submacula Wlk. 1850, Dipt. Saund., 1:30. (Tabanus) B. M.(H) = Dasy- 
chela, Bequaert and Renjifo (1946) who place as synonym of macula Macq. 
Krober (1931b) as synonym of neo-submacula Krob. 1931; (1934) places in 
synonymy of both macula Macq. and neo-submacula Krob. Both the latter 
are distinct species, in my opinion. 

*subvaria Wlk. 1848, List, 1:150. (Tabanus) B. M.—=Esenbeckia, Krober 
(1932a, 1934). Walker (1849) = Pangonia fuscipennis Wied. var. Ricardo 
(1900) = Pangonia, a distinct species. Close to *Esenbeckia notabilis Wlk., 
the proboscis rather heavy, incompletely sclerotized. 

*sulphureus Macq. 1847, Dipt. Exot., Suppl. 2:19. (Tabanus) B. M.(H) = 
Chlorotabanus inanis (Fab.), Krober (1934). Lutz (1907) as a pale form 
of mexicanus L, 

*tabanipennis Macq. 1848, Dipt. Exot., 1(1):108. (Pangonia) M. P. The 
lectotype = Fidena castanea Perty 1833 of Kréber (1930k). The specimen 
from de la Mana Leschen, a paratype, = ? *Pseudoscione seminigra Ric. 
Specimens in B. M. det. tabanipennis by Bigot = seminigra Ric. The lecto- 
type of tabanipennis also = *Fidena unicolor Macq. 1845. (N.S.) 

*tanycerus O. S. 1886, Biol. Centr.-Amer., Dipt., 1: 46. (Chrysops) B. M.(H) 
= Assipala Philip 1941 genotype. 

*tenens Wik. 1850, Newman’s Zoologist, 8, App., p. Ixv. (Tabanus) B. M.(H) 
= Tabanus cinerarius Wied. 1828. Not T. tenens Wik. 1850, Dipt. Saund., 
1:49. Walker (1854) changes name to confligens. Krober (1934) as 
Chelommia, T. cinerarius was proposed as a new name for T. glaucus Wied. 

- 1819 (Zool. Mag., 1(3) : 42) thought to be preoccupied by T. glaucus Meig. 
1820 (Syst. Beschreib. Europ. Zweifl. Ins., 2:51) but this appears to be an 
error, and the species should be known as Tabanus glaucus Wied. 18109. 

*tenuirostris Wik. 1860, Trans. Ent. Soc. London, 5: 272. (Pangonia) B. M. = 
Esenbeckia flavohirta Bell. 1859. Krober (1934) as Iicardoa flavohirta. 

*tenuistria Wlk. 1848, List, 1:143. (Pangonia) B. M.=Fidena. Krober 
(1930g) as Sackenimyia redescribes type; (1934) as Melpia. 

*tepicana Towns. 1912, Can. Ent., 44(1):287. (Pangonia) B. M. = Esen- 
beckia. Philip (1954a). 

*terminalis Macq. 1855, Dipt. Exot., Suppl. 5:36. (Chrysops) B. M.(H) = 
Diachlorus curvipes Fab. 1805. (N.S.) 

*terminus Wik. 1848, List, 1: 160. (Tabanus) B. M. = ? Tabanus sorbillans 
Wied. Type is a d and seems to match sorbillans fairly well. Kréber (1933a) 
as a valid species of (Neotabanus). (N.S.) 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


*testacea Macq. 1846, Dipt. Exot. Suppl. 1:29. (Dichelacera) B. M.(H). 
Kroéber (1934) with repanda as queried synonym = *Dichelacera repanda 
Wilk. 1848. Bequaert and Renjifo (1946) as (Catachlorops). Close to 
*quadrimaculatus Macq. and *alcis Will. 

*testaceiventris Macq. 1848, Dipt. Exot., Suppl. 3:9. (Pangonia) B. M.(H) 
=Esenbeckia, Krober (1932a) with uwmbra Wlk. 1850 as synonym = 
*E. umbra Wik. 

*testaceomaculatus Macq. 1838, Dipt. Exot., 1(1):144. (Tabanus) M. P.(H) 
=Dasybasis, Kroéber (19301) as (Agelanius). = ? *D. trigonophorus 
Macq. 1838, p. 185. 

*testaceus Macq. 1838, Dipt. Exot., 1(1):137. (Tabanus) M. P. Closest to 
*T. indecisus Big. but differs. Krober (1930c) claims he saw type in London. 
Not T. testaceus Forskal 1775. 

*tinctipennis Krob. 1931, Zool. Anz., 94(9-10) : 256. (Esenbeckia) B. M. 

*tinctus Wlk. 1850, Dipt. Saund., 1:29. (Tabanus) B. M. = Tabanus eggeri 
Schin. 1868 (Palearctic), nom. nov. pro 7. intermedius Egger 1859, not 
Walker 1848. Walker’s name appears to be the oldest for this European 
species. Bequaert (1940) suggests may not be Neotropical. (N.S.) 

*transposita Wlk. 1854, List, 5:151. (Dichelacera) B. M.(H) = Catachlo- 
rops, Kréber (1931i, 1934). Fairchild (1940b). 

*trifaria Macq. 1838, Dipt. Exot., 1(1) : 163. (Chrysops) M. P.(H). Krober 
(1926, 1934). 

*trifascia Wlk. 1850, Dipt. Saund., 1:37. (Tabanus) B. M. Type in very 
poor condition, a ¢ Stenotabanus, but not further identifiable in present 
state of knowledge of this group. It is not the ¢ of *callosus Macq. as sug- 
gested by Krober (10934). 

*trigonophorus Macq. 1838, Dipt. Exot., 1(2) : 185. (Tabanus) M. P. = Dasy- 
basis. Very close to *testaceomaculatus Macq. 1838, p. 144, structurally, but 
both types very denuded and certainty impossible. 

*tristis Big. 1892, 5:621. (Dasybasis) B. M.= ? Protodasyapha, Krober 
(1930k). A do. Eyes densely pubescent, holoptic, facets demarcated. Ocelli 
present. Subepaulet bare, hind tibiae spurred, labella fleshy, antennae subu- 
late, Pangonia-like. 

*tritus Wlk. 1857, Trans. Linn. Soc. London, 17(3) : 338. (Tabanus) B. M. = 
Dasybasis. Kréber (1930i, 1934) as Stypommia, the genotype of which St. 
patagonica End., he considers a synonym of tritus Wlk., but his figures and 
descriptions of patagonica and tritus do not agree with each other or with 
Walker’s type. 

*umbra Wlk. 1850, Dipt. Saund., 1:19. (Pangonia) B. M.(H) = *Esenbeckia 
testaceiventris Macq. 1848, Krober (1934). 

*unicinctus Wlk. 1857, Trans. Ent. Soc. London, 4: 122. (Tabanus) B. M. = 
*Leucotabanus albovarius (Wlk.) 1854. (N.S.) = *Leucotabanus leu- 
conotum Fchld. 1941. Not T. unicinctus Loew 1856. The type lacks antennae 
and is poorly preserved. It is either albovarius Wlk. or canithorax Fchld. 
with the former more likely. *Leuconotum Fchld. is the same as *albovarius 
WIk. 

*unicolor Macq. 1845, Dipt. Exot., Suppl. 1:27. (Pangonia) B. M.(H) = 
Fidena castanea Perty 1830, Kréber (1930k, 1934). = *Pangonia tabani- 
pennis Macq. 1848. Lutz (1909) makes xanthopogon Macq. 1838 also a syno- 
nym of castanea Perty. Lutz (1907) says unicolor Will. 1895 is different. 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 31 


*unifasciata Macq. 1838, Dipt. Exot., 1(1):119. (Dichelacera) M. P.(H). 
Barretto (1949b) with *limbata Big., trigonotaenia Lutz and soror Krob. as 
syns. 

*ynipunctatus Big. 1892, 5: 663. (Atylotus) B. M.(H) = *Tabanus (Lopho- 
tabanus) piraticus Fchld. 1942. (N.S.) Philip (1952) = T. jilamensis 
Hine. Not T. (L.) unipunctatus, Kroéb. 1929. Jilamensis Hine appears to be 
distinct. See Fairchild 19422. 

*univittatus Macq. 1855, Dipt. Exot., Suppl. 5:30. (Tabanus) B. M.(H) = 
*Tabanus desertus Wlk. 1850. (N.S.) 

*yacillans Wik. 1850, Dipt. Saund., 1:70. (Dichelacera) B. M.(H) = ? Cata- 
chlorops capreolus Wied. 1828. (N.S.) Kréber (1934) as synonym of potator 
Wied. 1828. Not vacillans, Barr. 1946. The type keys out to capreolus in 
Barretto’s key (1946). Lutz (1907) = potator Wied. 

*valterii Macq. 1838, Dipt. Exot., 1(2):184. (Tabanus) M. P. Related to 
cinerarius Wied. and importunus Wied. but apparently distinct. 

*varius Wlk. 1848, List, 1: 209. (Diabasis) B. M.(H) = Scaptia. Ricardo 
(1904) as ? Diatomineura. Krober (1930k) = Calliosca schoenemani End. 
genotype. 

*yaripes Wik. 1854, List, 5, Suppl. 1: 298. (Chrysops) B. M.(H) = Dia- 
chlorus curvipes Fab. 1805, Ricardo (1901). 

*varipes Wlk. 1857, Trans. Linn. Soc. London, 17(3) : 337. (Tabanus) B. M. 
Krober (1934) = ? ? Tabanus. Subepaulet setose, a small vertical tubercle, 
eyes sparsely short pilose under high magnification. Looks Nearctic or 
Palearctic to me. 

*venenatus O. S. 1886, Biol. Centr.-Amer., Dipt., 1:53. (Tabanus) B. M.(H) 
= Rhabdotylus. Kroéber (1934) = Amphiclorops. Fairchild (1942b) = 
Stibasoma. Very close to *Rhab. viridiventris Macq. 

*venosus Big. 1892, 5:685. (Tabanus) B. M.(H) = *Stenotabanus maculi- 
pennis (Kroéb.) 1929. (N.S.) Kroéber (19302) as Stypommia, but his 
venosus not same species as the type. St. pequeniensis Fchld. 1942 also close. 

*viduus Wl1k. 1850, Newman’s Zoologist, 8, App., p. lxviii. (Tabanus) B. M.(H) 
= *Tabanus basivitta Wlk. 1850. (N.S.) Krober (1930h) =T. (Lopho- 
tabanus) lividus Wik. 1848, but this latter = importunus Wied., in my 
opinion. 

*viridiventris Macq. 1838, Dipt. Exot. 1(1): 112. (Pangonia) M. P.(H) = 

_ (Pseudoscione). Enderlein (1922, 1925) as Parosca genotype. Krober 
(1930k) as Parosca; (1934) as Listrapha. 

*viridiventris Macq. 1838, Dipt. Exot., 1(1):141. (Tabanus) M. P.(H) = 
Rhabdotylus. Close to venenatus O. S. and planiventris Wied. Carrera and 
Lane (1945). 


SUMMARY 


A study of the type specimens of Neotropical Tabanidae preserved 
in the British Museum of Natural History in London and the Muséum 
d’Histoire Naturelle in Paris is reported. The types or type material 
of 335 species were seen and an attempt made to place them generi- 
cally. New synonymy is proposed in about 70 cases and new generic 
or subgeneric placement in 76 cases. Three new names are proposed 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


for homonyms and four species previously considered Neotropical 
are shown to be probably or surely from other regions. Specimens 
compared and agreeing with the types of 181 names (homotypes) 
were brought back and will eventually be deposited in American 
museums. Of the names treated, 189 appear to be valid, pending com- 
plete information on the status of the earlier names of Wiedemann, 
Thunberg, and some other authors; 110 are synonyms, and 40 are 
homonyms—in a fair number of cases a name may be both. In a few 
cases it was impossible to fix the status of a name owing to the condi- 
tion of the type or uncertainty as to the material’s being a true type. 
For the sake of completeness a list is also given of the Walker and 
Macquart species and a few others whose types could not be found— 
64 in all. The Walker types are presumed lost, while some of the Mac- 
quart types not seen by me are in Paris and others may turn up else- 
where. 


REFERENCES 
Barretto, M. P. 
1946. Estudos sobre Tabanidas Brasileiros II. Sobre 0 genero Catachlorops 
Lutz com descricdes de sete especies novas. An. Fac. Med. Univ. 
Sao Paulo, vol. 22, pp. 151-183, 29 figs. 
1948. Estudos sobre Tabanidas Brasileiros V. Sobre o genero Amphichlo- 
rops Lutz, com as descricdes de quatro especies novas. Rev. Ent., 
vol. 19, No. 3, pp. 401-417. 
1949a. Estudos sobre Tabanidas Brasileiros VII. O genero Chelommia 
End., com as descricdes de tres novas especies. Rev. Brasileiro 
Biol., vol. 9, No. 1, pp. 39-48, 13 figs. 
1949b. Estudos sobre Tabanidas Brasileiros VIII. Sobre os generos Dichela- 
cera Macq., e Rhamphis End. com a descricao de un novo genero. 
An. Fac. Med. Univ. Sao Paulo, vol. 24, pp. 63-80. 
1950. Estudos sobre Tabanidas Brasileiros XIII. O genero Phaeotabanus ~ 
Lutz e descricao de dois novos generos. An. Fac. Med. Univ. Sao 
Paulo, vol. 25, pp. 89-100, 6 figs. 
BeELuArpI, L. 
1859-62. Saggio di Ditterologia Messicana. Mem. Reale Accad. Sci. 
Torino, ser. 2, vol. 19, pt. I, pp. 1-80, 3 pls. (1859) ; vol. 21, pt. 2, 
pp. I-99, 2 pls. (1861), and Appendix, pp. 1-28, 1 pl. (1862). 
BEQUAERT, J. 
1924. Notes upon Surcouf’s treatment of the Tabanidae in the Genera In- 
sectorum and upon Enderlein’s proposed new classification of this 
family. Psyche, vol. 31, No. 1, pp. 24-40. 
1926. Jn Medical Report Hamilton Rice Seventh Expedition to the Amazon, 
etc., chap. XV, Insecta (Tabanidae), pp. 214-235. Cambridge. 
1931. Tabanidae of the peninsula of Yucatan, Mexico, with descriptions of 
new species. Journ. New York Ent. Soc., vol. 39, No. 4, pp. 533- 
553, 3 figs. 
1937. Triceratomyia, a new South American genus of Pangoniinae (Dipt. 
Tabanidae). Rev. Ent., vol. 7, No. 4, pp. 350-353, 1 fig. 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 33 


1939. Résultats scientifiques des croisiéres du navire-école Belge Mercator, 
vol. 2, XV, Tabanidae, pp. 169-170. 
1940. The Tabanidae of the Antilles. Rev. Ent., vol. 11, Nos. 1-2, pp. 253- 
360, 32 figs. 
1944. Further studies on the Tabanidae of Trinidad, B.W.I. Psyche, vol. 
51, Nos. 1-2, pp. 12-21. 
BEQuAERT, J., and Renjiro S., SANTIAGO. 
1946. The Tabanidae of Colombia. Psyche, vol. 53, Nos. 3-4, pp. 52-88. 
BIGOTS) 2M. EF, 
1892a. Descriptions de diptéres nouveaux. Mém. Soc. Zool. France, vol. 5, 
pp. 602-601. 
1892b. Nova genera dipterorum. Wiener Ent. Zeitg., vol. 11, No. 6, pp. 
161-162. 
BLANCHARD, E. 
1852. In Gay, C., Historia fisica y politica de Chile. Zoologia, vol. 7, Ta- 
banidae, pp. 386-397. 
Bopk1n, G. E., and Creare, L. D., Jr. 
1916. Notes on some animal parasites in British Guiana. Bull. Ent. Res., 
vol. 7, pp. 179-190, 3 figs. 
BRETHES, J. 
1910-1911. Dipteros nuevos o poco conocidos de Sud-America. An. Mus. 
Nac. Hist. Nat. Buenos Aires, ser. 3, vol. 13, pp. 472-480 (1910), 
481-484 (1911). 
CarrERA, M., and LANE, J. 
1945. Diptera de Caioba (Estado do Parana). Arq. Mus. Paranaense, 
vol. 4, art. 5, pp. 127-136, 7 figs. 
Castro, G. M. pe OLIveErRA. 
1945. Notas sobre Tabanidas Brasileiros I. Fidena longipalpis End. 1925. 
Mem. Inst. Oswaldo Cruz, vol. 42, No. 2, pp. 281-286. 
ENDERLEIN, G. 
1922. Ein neues Tabanidensystem. Mitt. Zool. Mus. Berlin, vol. 10, No. 2, 
PP. 333-351. 
1925. Studien an blutsaugenden Insekten, I. Grundlagen eines neuen Systems 
der Tabaniden. Mitt. Zool. Mus. Berlin, vol. 11, No. 2, pp. 255-409, 
5 figs. 
1929. Zur Kenntnis einiger von Herrn Oskar Schoenemann gesammelten 
chilenischen Melpiinae. Wiener Ent. Zeitg., vol. H6, No. 2, pp. 66- 
71, 1 fig. 
FAIRCHILD, G. B. 
1939. Notes on the genus Acanthocera Macquart. Rev. Ent., vol. 10, No. 1, 
Pp. 14-27, 23 figs. 
1940a. Notes on Tabanidae from Panama I. The genera CH Potabanus and 
Cryptotylus. Rev. Ent., vol. 11, No. 3, pp. 713-722, 5 figs. 
1940b. Notes on Tabanidae from Panama II. The genus Dichelacera Macq. 
and related genera. Ann. Ent. Soc. Amer., vol. 33, No. 4, pp. 683- 
700, 2 pls. 
1941. Notes on Tabanidae from Panama IV. The genus Leucotabanus Lutz. 
Ann. Ent. Soc. Amer., vol. 34, No. 3, pp. 629-638, 8 figs. 
1942a. Notes on Tabanidae from Panama III. The genus Chrysops Meigen. 
Proc. Ent. Soc. Washington, vol. 44, No. 1, pp. 1-8, 1 pl. 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


1942b. Notes on Tabanidae from Panama V. The genus Tabanus, subgenus 
Bellardia Rondani. Psyche, vol. 49, Nos. 1-2, pp. 8-17, 1 pl. 
1942c. Notes on Tabanidae from Panama VII. The subgenus Neotabanus 
Lutz. Ann. Ent. Soc. Amer., vol. 35, No. 2, pp. 153-182, 2 pls. 
1942d. Notes on Tabanidae from Panama VIII. The genera Pityocera, 
Scione and Esenbeckia. Ann. Ent. Soc. Amer., vol. 35, No. 2, pp. 
183-199, 2 pls. 
1942e. Notes on Tabanidae from Panama IX. The genera Stenotabanus, 
Lepiselaga and related genera. Ann. Ent. Soc. Amer., vol. 35, No. 
3, pp. 289-300, 1 pl. 
1942f. Notes on Tabanidae from Panama X. The genus Tabanus and 
résumé of the Tabanidae of Panama. Ann. Ent. Soc. Amer., vol. 
35, No. 4, pp. 441-474, 2 pls. 
1951a. Descriptions and notes on Neotropical Tabanidae. Ann. Ent. Soc. 
Amer., vol. 44, No. 3, pp. 441-462, 1 pl. 
1951b. The generic names for Tabanidae proposed by Adolpho Lutz. 
Psyche, vol. 57, No. 4, pp. 117-127. 
1953. Notes on Neotropical Tabanidae with descriptions of new species. 
Ann. Ent. Soc. Amer., vol. 46, No. 2, pp. 259-280, 2 pls. 
Fercuson, E. W. 
1924. Notes on the nomenclature of Australian Tabanidae: Subfamily 
Pangoniinae. Bull. Ent. Res., vol. 14, No. 3, pp. 251-263. 
1926. Additional notes on the nomenclature of Australian Tabanidae. Bull. 
Ent. Res., vol. 16, No. 4, pp. 293-308, 12 figs. 
Hack, W. H. 
1953. Segunda contribucion el estudio de los Tabanidos Argentinos. An. 
Inst. Med. Reg. Univ. Nac. Tucuman, vol. 3, No. 3, pp. 339-347, 
10 figs. 
Hine, J. S. 
1917. Costa Rican Diptera collected by Philip P. Calvert, 1909-1910. Paper 
2, Tabanidae and Asilidae. Trans. Amer. Ent. Soc., vol. 43, pp. 
291-299. 
1920. Description of horseflies from Middle America II. Ohio Journ. Sci., 
vol. 20, No. 8, pp. 311-3109. 
1925. Tabanidae of Mexico, Central America and the West Indies, Occ. 
Pap. Mus. Zool. Univ. Michigan, No. 162, pp. 1-35. 
KERTESZ, COLOMANNUS. 
1900. Catalogus tabanidarum orbis terrarun universi. 78 pp. Mus. Nat. 
Hungarico Ed. Budapest. 
1908. Catalogus dipterorum. Vol. 3, 348 pp. Mus. Nat. Hungaricum. 
Budapest. 
Knap, F. 
1916. What is Tabanus mexicanus? Ins. Ins. Mens., vol. 4, pp. 95-100. 
Kroser, O, 
1926. Die Chrysops—arten Sud—und Mittelamerikas, nebst den Arten der 
Inselwelt und Mexikos. Konowia, vol. 4, Nos. 3-4, pp. 210-256, 
319-375, 5 pls. 
1928a. Neue Dipteren des Deutsches Entomologisches Museum in Dahlem. 
Ent. Mitt., vol. 17, No. 1, pp. 31-41, 9 figs. 


NO. 


3 


1928b. 


19292. 
1929b. 


1920¢. 


1920d. 
1930a. 


1930b. 


1930C¢. 


1930d. 


1930e. 


1930f. 


1930¢. 


1930h. 


19301. 


1930}. 


1930k. 
10931a. 
193Ib. 
193ICc. 
1931d. 


1931e. 


1931f. 


1931g. 


NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 35 


Die amerikanischen Arten der Tabaniden—subfamilie Diachlorinae 

End. Arch. Schiffs-u. Tropen-Hygiene, etc., vol. 32, No. 2, pp. 
73-123. 

Uber einige kleinere Gattungen der sud amerikanischen Tabanini. 
Zool. Anz., vol. 83, Nos. 1-4, pp. 47-63; Nos. 5-8, pp. 115-137. 

Die Ausbeute der deutschen Chaco—Expedition 1925—26. XIII. 
Tabanidae. Konowia, vol. 8, No. 2, pp.174-193, 2 pls. 

Ergebnisse einer zoologischen Sammelreise nack Brasilien, insbeson- 
dere das Amazonasgebiet, ausgefuhrt von Dr. H. Zerny. Ann. 
Naturhist. Mus. Wien. vol. 43, pp. 243-255, 13 figs. 

Die Stenotabaninae und die Lepiselaginae Sudamerikas. Encycl. 
Entom., ser. B, 11, Diptera, vol. 5, No. 3, pp. 101-154, 07 figs. 

Nachtrage zu den kleinen Gattungen der sudamerikanischen Ta- 
banini. Zool. Anz., vol. 86, Nos, 9-10, pp. 248-265, 8 figs. 

Die Tabanidenuntergattung Phaeotabanus Lutz. Zool. Anz., vol. 86, 
Nos. II-I2, pp. 273-300. 

Die Untergattungen Macrocornus Lutz und Chlorotabanus Lutz. 
Zool. Anz., vol. 87, Nos. 1-2, pp. 1-18, 13 figs. 

Die Tabaniden subfamilie Silviinae der neotropischen Region. Zool. 
Anz., vol. 88, Nos. 9-10, pp. 225-239, I1 figs. 

Die Pityocerini (Tabanidae) der neotropischen Region. Zool. Anz., 
vol. 88, Nos. II-12, pp. 305-312. 

Die Tribus Pangoniini der neotropischen Region. Zool. Anz., vol. 89, 
Nos. 7-10, pp. 211-228. 

Die Tabanidengattung Sackenimyia Big. Zool. Anz., vol. 90, Nos. 
I-2, pp. I-12, 6 figs. 

Neue Tabaniden und Zusatze zu bereits beschreibenen. Zool. Anz., 
vol. 90, Nos. 3-4, pp. 69-86, 21 figs. 

Tabanidae. Diptera of Patagonia and South Chile, pt. 5, fase. 2, 
pp. 106-161, 2 pls. British Museum. 

Die sudamerikanischen Arten der Gattung Scione Wlk. (= Rhino- 
tricliista End.). Stett. Ent. Zeitg., vol. 91, No. 2, pp. 140-174. 

Die Pelecorhynchinae und Melpiinae Sudamerikas. Mitt. Zool. 
Staatinst. Zool. Mus. Hamburg, vol. 44, pp. 149-196, 33 figs. 

Neue sud- und mittelamerikanische Arten der Dipterengattung 
Tabanus L. Stett. Ent. Zeitg., vol. 92, No. 2, pp. 275-305. 

Neue neotropische Tabaniden aus den Unterfamilien Bellardiinae 
und Tabaninae. Rev. Ent., vol. 1, No. 4, pp. 400-417, 18 figs. 

Die Tabanus—Untergattung Gymnochela End. Zool. Anz., vol. 96, 
Nos. 1-2, pp. 49-61, 9 figs. 

Neue Arten der Gattung Fidena Wlk. Zool. Anz., vol. 95, Nos. 1-2, 
Pp. 17-37, 19 figs. 

Neue Arten aus dem Genus Esenbeckia Rond. Zool. Anz., vol. 94, 
Nos. 9-10, pp. 245-257, 7 figs. 

Die Tabanus—Gruppen Straba End. und Poecilosoma Lutz (= 
Hybostraba und Hybopelma End.) der neotropischen Region. Zool. 
Anz., vol. 94, Nos. 3-4, pp. 67-80, 20 figs. 

Die kleinen Gattungen der Dichelacerinae End. aus der sud- 
amerikanischen Region. Rev. Ent., vol. 1, No. 3, pp. 282-208, 
11 figs. 


36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


193th. Dreizehn neue neotropische Tabanusarten. Konowia, vol. 10, No. 4, 
pp. 291-300. 

1931i. Neue sudamerikanische Tabaniden des ungarischen National-Mu- 
seums und einiger anderer Institute. Ann. Mus. Nat. Hungarici, 
vol. 27, pp. 329-350, 19 figs. 

1932a. Das Genus Esenbeckia Rondani und die Gymnochela—Untergattung 
Amphichlorops Lutz. Rev. Ent., vol. 2, No. 1, pp. 52-90, 32 figs. 

1932b. Die Tabaniden—Subfamilie Bellardiinae End. der neotropischen 
Region. Rey. Ent., vol. 2, No. 2, pp. 289-302, 6 figs. 

1933a. Das Subgenus Neotabanus der Tabanidengattung Tabanus s. lat. 
Rev. Ent., vol. 3, No. 3, pp. 337-367. 

1933b. Die neotropischen Arten der Tabanidengattung Fidena Wlk. Arch. 
Naturg., N. F., vol. 2, No. 2, pp. 231-284. 

1934. Catologo dos Tabanidae da America do Sul e Central, incluindo a 
Mexico e as Antilhas. Rev. Ent., vol. 4, Nos. 2-3, pp. 222-276, 291- 
333- 

1939. Das Tabanidengenus Catachlorops Lutz. Verhoff. Deutsch. Kol. Mus. 
Bremen, vol. 2, pp. 211-232, 4 pls. 

1940. Das Tabanidengenus Dicladocera Lutz. Verhoff. Deutsch. Kol. Mus. 
Bremen, vol. 3, No. 1, pp. 58-92, 3 pls. 

Lutz, A. 

1907. Bemerkungen iiber die Nomenclatur und Bestimmung der Brasilian- 
ischen Tabaniden. Zentralbl. Bakteriol., vol. 44, No. 1, pp. 137-144. 

1909. Tabaniden Brasiliens und einiger Nachbarstaaten. Zool. Jahrb., 
Suppl., vol. 10, No. 4, pp. 619-602, 3 pls. 

1913. Tabanidas do Brasil e alguns estados visinhos. Mem. Inst. Oswaldo 
Cruz, vol. 5, No. 2, pp. 142-191, 2 pls. 

1915. Tabanidas do Brasil e de alguns estados visinhos. Segunda memoria. 
Mem. Inst. Oswaldo Cruz, vol. 7, No. 1, pp. 51-119, 3 pls. 

Lutz, A., and Nerva, A. 

1914. As Tabanidae do Estado do Rio de Janeiro. Mem. Inst. Oswaldo 
Cruz, vol. 6, No. 2, pp. 69-80. 

Lutz, A., Araujo, H. C. pe Souza, and Fonseca FiLuo, O. pa. 

1918. Viagem scientifico no Rio Parana e a Asuncion, com volta por Buenos 
Aires, Montevideo e Rio Grande. Mem. Inst. Oswaldo Cruz, vol. 
10, pp. 104-199. 

Lutz, A., and Castro, G. M. pE OLIVEIRA. 

1935. Sobre algumas novas especies de motucas do genero Esenbeckia Rond. 

Mem. Inst. Oswaldo Cruz, vol. 30, No. 3, pp. 543-562. 
MACcKERRAS, I. M. 

1954. The classification and distribution of Tabanidae I. General review. 

Australian Journ. Zool., vol. 2, No. 3, pp. 431-454, 10 figs. 
Macoguankt, J. 

1834. Histoire naturelle des insectes diptéres. Jn Suites 4 Buffon, vol. 1, 
pp. 186-216. 

1838-1855. Diptéres exotiques nouveaux ou peu connus. Mém. Soc. Roy. 
Sci., Agr., Arts, Lille. Contemporaneously published as a separate 
work by Librairie Encylopédique de Roret, Paris, with same title but 
separate pagination and in numbered volumes and supplements. The 
original parts, with pages, plates, and dates of issue are here listed 


NO. 3 NEOTROPICAL FLIES, TABANIDAE—FAIRCHILD 37 


with the corresponding issues of the Roret edition in parentheses. 
1838, pp. 9-225, pls. I-XXV (1838, vol. 1, pt. 1, pp. 1-221, pls. I- 
XXV) ; 1839, pp. 121-323, pls. I-XIV (1838, vol. 1, pt. 2, pp. 1-207, 
pls. I-XIV); 1841, pp. 283-413, pls. I-X XI (1840, vol. 2, pt. 1, 
pp. I-135, pls. I-X XI) ; 1842, pp. 63-200, pls. I-X XII (1842, vol. 2, 
pt. 2, pp. 1-140, pls. I-X XII); 1843, pp. 162-460, pls. I-XXXVI 
(1843, vol. 2, pt. 3, pp. 1-304, pls. I-XXXVI); 1845, pp. 133-364, 
pls. I-XX (1846, Suppl., pp. 1-238, pls. I-XX); 1847, pp. 21-210, 
pls. I-VI (1847, 2° Suppl., pp. 1-104, pls. I-VI) ; 1848, pp. 161-237, 
pls. I-VII (1848, Suite de 2° Suppl. [or third Supplement], pp. 
1-77, pls. I-VII) ; 1850, pp. 309-379, pls. I-XIV, and 1851, pp. 134- 
209, pls. XV-X XVIII (1851, 4° Suppl., pp. 1-336, pls. I-X XVII) ; 
1855, pp. 25-156, pls. I-VII (1855, 5° Suppl., pp. 1-136, pls. I-VII). 
Otproyp, H. 
1954. The horse-flies (Diptera: Tabanidae) of the Ethiopian region. Vol. 
II. Tabanus and related genera. x + 341 pp. British Museum. 
OsTEN SACKEN, C. R. 
1878. Catalogue of the described Diptera of North America (2d ed.). 
Smithsonian Mise. Coll. vol. 16, No. 270, xlvi ++ 276 pp. 
1886. Biologia Centrali-Americana, Insecta, Diptera, 1 (Tabanidae), pp. 
43-60. 
PEcHUMAN, L. L. 
1937. Notes on some neotropical species of the genus Chrysops. Rev. Ent., 
vol. 7, Nos: 2-3, pp. 135-141, 2 figs. 
Purp, C. B. 
1941. A new genus of Neotropical deer flies (Dipt. Tabanidae). Rev. Ent., 
vol. 12, No. 3, pp. 470-474. 
1947. A catalog of the blood-sucking fly family Tabanidae (horse flies and 
deer flies) of the Nearctic region north of Mexico. Amer. Midl. 
Nat., vol. 37, No. 2, pp. 257-324. 
1950. New North American Tabanidae II. Ann. Ent. Soc. Amer., vol. 43, 
No. I, pp. 115-122. 
1952. The Linnaean and Degeerian species of American Tabanidae. Ann. 
Ent. Soc. Amer., vol. 45, No. 2, pp. 310-314. 
1954a. New North American Tabanidae VIII. Notes on and keys to the 
genera and species of Pangoniinae exclusive of Chrysops. Rev. 
Brasileira Ent., Sao Paulo, vol. 2, pp. 13-60. 
1954b. New North American Tabanidae VII. Descriptions of Tabaninae 
from Mexico. Amer. Mus. Nov. No. 1695, pp. 1-26, 15 figs. 
Ricarpo, G. 
1900a. Notes on the Pangoninae of the family Tabanidae in the British 
Museum collection. Ann. Mag. Nat. Hist., ser. 7, vol. 5, pp. 97- 
121, 167-182, 1 pl. 
1900b. Description of five new species of Pangoninae from South America. 
Ann. Mag. Nat. Hist., ser. 7, vol. 6, pp. 201-204. 
1901-1902. Further notes on the Pangoninae of the family Tabanidae in the 
British Museum collection. Ann. Mag. Nat. Hist., ser. 7, vol. 8, 
pp. 286-315; vol. 9, pp. 424-438. 
1904. Notes on the smaller genera of the Tabaninae of the family Tabanidae 
in the British Museum collection. Ann. Mag. Nat. Hist., ser. 7, 
vol. 14, pp. 349-373. 


38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


RonpDaANI, C. 
1850. Osservazioni sopra alquante specie di esapodi ditteri del Museo Tori- 
nense. Nuoy. Ann. Bologna, ser. 3, vol. 2, pp. 165-197. 
1863. Dipterorum species et genera aliqua exotica revisa et annotata, novis 
nonnullis descriptis. Arch. Zool. Anat. Canestrini, vol. 3, pp. 1-99. 
ScHINER, J. R. 
1868. Reise der Osterreichen Fregatte Novara um die Erde, etc. Zoologische 
Teil, Diptera, pp. i-vi, 1-388, 4 pls. 
STONE, A. 
1938. The horseflies of the subfamily Tabaninae of the Nearctic region. 
U. S. Dept. Agr. Misc. Publ., 305, pp. 1-171, 79 figs. 
1944. Some Tabanidae from Venezuela. Bol. Ent. Venezolana, vol. 3, No. 3, 
pp. 125-138. 
1954. The genus Bolbodimyia Bigot. Ann. Ent. Soc. Amer., vol. 47, No.2, 
pp. 248-254. 
Surcour, J. M. R. 
1919. Diptéres. Brachycéres piqueurs (Tabanidae). Mes. Arc de Meérid- 
ian Equatorial en Amérique du Sud, vol. 10, No. 2, pp. 217-233. 
1921. Genera insectorum, fasc. 175, Diptera, Tabanidae, pp. 1-182, 5 pls. 


Brussels. 
Sziuapy, Z. 
1926. New and Old World horseflies. Biol. Hungarica, vol. 1, No. 7, pp. 
1-30, I plate. 


THUNBERG, C. P. 
1827. Tabani septendecim novae species descriptae. Nova Acta Reg. Soc. 
Sci. Upsal., vol. 9, pp. 53-62; ibid., Tanyglossae septendecim novae 
species descriptae, pp. 63-75. 
WALKER, F. 
1848-1854. List of the specimens of dipterous insects in the collection of the 
British Museum. Pt. 1, pp. 1-299 (1848); pt. 5, pp. 1-330, I pl. 
(1854) ; pt. 5, suppl. I, pp. 1-350. 
1850a. Insecta Saundersiana: or characters of undescribed insects in the 
collection of W. W. Saunders. I. Diptera, pt. 1, pp. 1-75, 2 pls. 
1850b. Characters of undescribed Diptera in the British Museum. New- 
man’s Zoologist, vol. 8, Appendix, art. 1, pp. Ixv-lxxii, art. 5, 
pp. xcv-xcix, art. 25, pp. CXxXi-Cxxii. 
WIEDEMANN, C. R. W. 
1821. Diptera exotica. 
1828-1830. Aussereuropaische Zweiflugelige Insekten, vol. I, pp. i-xxxii, 
1-608, 7 pls. (1828) ; vol. 2, pp. i-xii, 1-684, 5 pls. (1830). 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 4 


Charles DB. and Mary Waux CHalcott 
Research Fund 


NEW CRETACEOUS BRACHIOPODA 
FROM ARIZONA 


(WitTH 4 PLatEs) 


By 
G. ARTHUR COOPER 


United States National Museum 
Smithsonian Institution 









ae le 
4st HONOR Uy 
7, Te 
NAINGTO wily 












(PuBLIcaTION 4227) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
DECEMBER 21, 1955 


THE LORD BALTIMORE PRESS, INC. 
BALTIMORE, MD., U. S. A. 





Charles D. and Mary Waux Walcott Research Fund 


NEW CRETACEOUS BRACHIOPODA 
FROM ARIZONA 


By G. ARTHUR COOPER 


U. S. National Museum 
Smithsonian Institution 


(WirH Four Prates) 


Cretaceous brachiopods, except for the occurrences of Kingena in 
Texas, are a great rarity in the United States, and it is also unusual 
to find here more than one kind of brachiopod of that age in abun- 
dance. However, in 1947, Dr. J. B. Reeside, Jr., of the U. S. Geologi- 
cal Survey, called my attention to several species of brachiopods from 
the Cretaceous Mural limestone of Arizona. These were insufficient 
for study ; therefore, in the summer of that year I went to the Bisbee 
area of Arizona with Dr. Ellis Yochelson, now of the U. S. Geological 
Survey, to obtain more material. 

Occurrence of Cretaceous brachiopods in Arizona is mentioned by 
Ransome (1904, p. 6) in his description of the Bisbee Quadrangle. 
The single occurrence cited is said to be the only one on the quadrangle 
and is a small hill on the east side of the quadrangle opposite the mouth 
of Glance Canyon and about 3 miles east of Glance. This hill is in the 
NW4iSWiNE? sec. 36, T. 23 S., R. 25 E. and lies about 0.2 
mile east of U. S. Highway 80 about 12.7 miles west-northwest of 
Douglas, Cochise County. The location is thus easily accessible be- 
cause of its proximity to an excellent road, and it is from this locality 
that Dr. Yochelson and I collected the specimens described below. 

The low hill from which the brachiopods were taken consists of 
massive limestone, through which the brachiopods are scattered. They 
are not concentrated in bands, although some pieces were found in 
which they were fairly common. A large part of the collection con- 
sisted of small lumps showing one or two specimens. Although some 
large pieces were taken, they were not rich and it was best, therefore, 
to collect individual specimens or small groups in small pieces. The 
limestone lumps taken produced few specimens aside from brachio- 
pods. These included small oysters, small rudistids, and a few poorly 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 131, NO. 4 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 13% 


preserved small echinoids. The brachiopods are fairly well silicified. 
The silicification is not coarse or crude in the large specimens, but it is 
in some of the immature ones. In general the specimens are brittle 
and fragile and must be handled with great care. 

These brachiopods occur in the Mural limestone but, because of the 
isolated character of the hill, the exact stratigraphic position of the 
specimens is uncertain. Ransome, who defined the Mural, states that 
“The little hills near the eastern edge of the quadrangle north of Hay 
Flat are composed mainly of the hard limestones of the upper member 
of the formation. Some of the beds here contain abundant corals 
(Astrocenia and another form not collected). Caprina, and a number 
of little brachiopods (Rhynchonella, Terebratella and Terebratula) 
not seen at any other locality in the quadrangle.” 

Stoyanow (1949, p. 20) divides the Mural limestone into three 
units: (a) Basal thinner-bedded limestone with Orbitolina texana; 
(b) massive “rudistid” limestone, and (c) thinner-bedded limestone 
with Orbitolina texana. He says: “In the basal beds of the Mural 
limestone, small brachiopods, corals, specimens of Lima muralensis, 
and large forms of Lunatia? sp. often occur. The massive limestone 
is usually replete with FRadiolites? sp., whereas the specimens of 
Caprina sp. are comparatively rare and come from the thinner-bedded 
layers below the reef.’’ These remarks suggest that Stoyanow identi- 
fied the brachiopod beds as low in the Mural. Perhaps brachiopods 
occur in more than one level and were not seen by Ransome. At any 
rate the Mural limestone is now placed (Cobban and Reeside, 1952) 
at about the middle of the Albian stage in the Lower Cretaceous. 


CRANISCUS HESPERIUS Cooper, new species 
Plate 3A, figures I-3 


Pedicle valve unknown. 

Brachial valve a low cone about medium size for the genus, sub- 
rectangular in outline; length about two-thirds the width; maximum 
width in anterior third; sides slightly oblique and gently rounded; 
anterior margin broadly rounded; anterolateral extremities narrowly 
rounded; posterior margin nearly straight. Apex approximately 
central, blunt ; anterior slope steep ; lateral slopes about as steep as an- 
terior slope, but posterior slope gentle. Surface irregular. 

Interior with low median ridge rising to a sharp point at the valve 
middle; anterior adductor scars narrowly elliptical, obliquely placed 
and forming low ridges which, with the median ridge, divide the valve 
into three parts; posterior adductor scars large, but lightly impressed. 
Anterior half with strong pallial ridges. 


NO. 4 CRETACEOUS BRACHIOPODA, ARIZONA—COOPER 3 


Measurements in mm.—Length, 10.6; maximum width, 14.0; height, 
3.4. 

Types.—Holotype U.S.N.M. No. 124192. 

Horizon and locality —Mural limestone, from a small hill 300 yards 
east of U. S. Highway 80, NW4SW4iNE4 sec. 36, T. 23 S., R. 25 E., 
Bisbee Quadrangle, Arizona. 

Discussion—This genus has not hitherto been identified among 
North American fossils. The laterally elongated muscle scars and short 
median septum are characteristic. The species is like Craniscus suessi 
(Bosquet) from the Maastrichtian of Holland in having the apex near 
the middle and a long, flat posterior slope, but the Dutch species is 
more swollen anteriorly and is a much deeper shell. 


CYCLOTHYRIS AMERICANA Cooper, new species 
Plate 1A, figures I-17 


Shell of about medium size for the genus, subtriangular in outline; 
maximum width at or near the middle; valves subequal in depth, the 
brachial valve having a slightly greater depth; anterior commissure 
gently uniplicate; surface costellate, costellae numbering about 32 to 
42 along the anterior margin. 

Pedicle valve moderately convex in lateral profile, with the maxi- 
mum convexity near the middle; beak apicate, nearly straight to sub- 
erect, making an angle of 60° to 80°. Umbo swollen; beak ridges 
moderately strong and defining a fairly broad, gently concave inter- 
area; sulcus originating in the anterior third to half, shallow and oc- 
cupying about one-third the width. Flanks gently convex, descending 
steeply to the sides. Deltidial plates conjunct, auriculate; foramen 
submesothyrid, oval in outline. 

Pedicle valve interior with strong but small teeth; dental plates 
‘stout and fairly long, not surrounding the muscle field which is large 
and broadly elliptical ; diductor scars subreniform in outline ; adductor 
scars posterior to diductors ; no pedicle collar. 

Brachial valve strongly convex in lateral and anterior profiles ; umbo 
swollen ; fold low or defined only as a wave of the commissure; flanks 
convex ; posterolateral slopes steep. Brachial valve interior with long, 
slitlike sockets bounded by moderately strong socket ridges; hinge 
plate divided ; crura curved, short, of radulifer type. 


MEASUREMENTS IN MILLIMETERS 


Length Width Thickness 


Holotype U.S.N.M. No. 124193a........... 14.0 13.6 9.0 
Paratype U.S.N.M. No. 124193b........... 11.8 12.0 9.0 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Types—Holotype U.S.N.M. No. 124193a; figured paratypes 
U.S.N.M. Nos. 124186a-e ; measured paratype U.S.N.M. No. 124193b. 

Horizon and locality —Mural limestone, from a small hill 300 yards 
east of U. S. Highway 80, NW4SW4iNE43 sec. 36, T. 23 S., R. 25 E., 
Bisbee Quadrangle, Arizona. 

Discussion.—This species is characterized by its moderate size, con- 
vex valves and costellate surface. One of the significant features of 
the Arizona species is the poor development of the fold and sulcus, 
which is strongly developed only at the front. A few specimens of 
thynchonellids have been found in the American Lower Cretaceous 
which are probably referable to Cyclothyris. None of these resembles 
the present species. The National Museum has a specimen of Cyclo- 
thyris from the Main Street formation, Fort Worth, Tex., but it is 
transversely elliptical rather than strongly triangular as the Mural 
species is. Of three specimens of Cyclothyris from the Edwards lime- 
stone, Presidio County, Texas, two are strongly transverse, but the 
third is suggestive of C. americana. 

The British species most suggestive of C. americana is C. nuciformis 
(Sowerby), but this differs in having a pronounced fold and sulcus 
for at least half the valve length. Some young specimens from the 
Cretaceous of France called Cyclothyris compressa (Lamarck) or 
C. depressa (Sowerby) suggest the American species but they are 
usually more transverse or are demonstrable aberrations of a variable 
species. 


CYCLOTHYRIS species 


A single specimen (U.S.N.M. No. 124216) differing importantly 
from C. americana was found with the other specimens described 
herein. Although somewhat crushed, it differs from C. americana in 
being much wider and in having a fairly prominent fold and sulcus 
that originate a short distance anterior to the middle. 


RECTITHYRIS VESPERTINA Cooper, new species 
Plate 1B, figures 18-37 


Shell small for the genus, elongate oval in outline and with the 
maximum width at the middle; sides gently rounded; anterior margin 
narrowly rounded. Valves unequal in depth, the pedicle valve having 
the greater depth. Posterior margin narrowly rounded to subcarinate ; 
anterior commissure rectimarginate to faintly uniplicate; lateral com- 
missure nearly straight. Surface smooth except for concentric lines 
and varices of growth. 


NO. 4 CRETACEOUS BRACHIOPODA, ARIZONA—COOPER 5 


Pedicle valve moderately convex in lateral profile and with the 
maximum convexity at about the middle; anterior profile fairly 
strongly convex ; umbonal region narrowly convex to subcarinate, the 
narrow swelling continued nearly to the median region where it dies 
out. Anterior third flattened to faintly sulcate. Beak erect, obliquely 
truncated ; foramen broadly elongate, moderately large, oval to circu- 
lar, submesothyrid to mesothyrid. Deltidial plates conjunct, not 
covered by beak, suture visible. Beak ridges strong. 

Interior of pedicle valve with large and thick teeth; dental plates 
obsolete ; pedicle collar small. Muscle marks too indistinct to discern 
individual scars or pattern of field. 

Brachial valve shallow, gently convex in lateral profile and broadly 
convex in anterior profile; umbonal region gently swollen; beak ob- 
_ scured by the overlapping of the deltidial plates ; median region gently 
swollen and forming a barely perceptible fold which appears at the 
front margin as a gentle wave of the commissure in the direction of 
the brachial valve; flanks gently inflated and with short, steep sides. 

Interior with short stout loop having short crura and short blunt 
crural processes; descending lamellae short; transverse ribbon broad 
in adults, fairly strongly elevated and with a flattened crest at its 
middle; outer socket plate moderately broad, moderately concave; 
inner socket plates nonexistant to small; inner socket ridge strong, 
overlapping the teeth. Cardinal process small, wide and short. Muscu- 
lature and pallial marks poorly impressed, elongate, somewhat tear- 
shaped. 


MEASUREMENTS IN MILLIMETERS 


Brachial 
Length length Width Thickness 
Holotype U.S.N.M. No. 124194b...... 17.6 14.0 14.0 0.4 
Paratype U.S.N.M. No. 1241094a...... 16.9 13.9 12.5 8.8 
, " es ET 2ATOAC As cues 20.3 17.4 17.4 11.6 
aR , ee VIEZATO“G 28H. t 5.4 4.7 3.9 ay 
5 ig MEILZATOACHSAAk 4 11.4 9.2 8.9 5.2 


Types.—Holotype U.S.N.M. No. 124194b; figured paratypes 
U.S.N.M. Nos. 124187, 124188, 124194¢, d, 124195, 124195a, 124196c; 
unfigured paratypes U.S.N.M. Nos. 1241948, e, 124196a, b, 124205, 
124218. 

Discussion.—This species is characterized by the unequal convexity 
of the valves, the suberect to erect beak (Thomson classification, 
1927), rectimarginate to faintly plicate anterior commissure, and short, 
stout loop. I am not completely happy about the assignment of this 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


species to Rectithyris but this appears to be the only genus at present 
to which it can be assigned. Points of agreement with Rectithyris as 
defined by Sahni (1929) appear: the mesothyrid foramen, the easily 
visible deltidial plate, rudimentary cardinal process, the inner hinge 
plates, even though they are not strongly developed, and the constric- 
tion of the broad transverse ribbon to form a median, flattened crest. 
Deviations from Rectithyris are the curvature of the beak, less strongly 
triangular loop, and smaller foramen. 

Some points of similarity exist between the Arizona species and 
Neoliothyrina. The loop of the latter is like that of R. vespertina and 
according to Sahni’s figures (1929, pl. 9, figs. 19, 20) shows the same 
type of variation. The loop shown in figure 20 has nearly parallel sides 
and the crural processes are well anterior to the crural bases, whereas 
the loop shown in figure 19 has the crural processes located almost at 
the junction of the crura and the crural bases. The beak characters and 
other details of Neoliothyrina, however, are too different to permit 
use of that name. 

No known species of Rectithyris is like this American species ; con- 
sequently, no direct comparison is possible. 

Variation of the loop.—Variations in parts of the loop are evident 
in many of the specimens, but these variations could not be correlated 
with shape or shell differences. In some specimens the brachial valve 
is distinctly flatter than in others, but this difference did not accord 
with loop differences. In the young, loop variation is evident in the 
length of the descending branch anterior to the crural base. One 
specimen, paratype U.S.N.M. No. 124196a, has the crural process 
given off almost at the junction of the descending branch with the 
crural base. In another, plate 1B, figure 33, the crural process is lo- 
cated a short distance anterior to the crural base. In this specimen the 
crural base appears as a ridge bounding the inner socket plate. This 
is true of a somewhat larger and more-elongate specimen shown in 
figures 29-31 on the same plate. This is not true, however, of the 
largest and oldest loop figured, same plate, figures 34 and 35, in which 
the descending process is short and the posterior extension of the 
crural base is buried in the formation of modest inner hinge plates. 
The presence of inner hinge plates appears to be an age character, at 
least in this case. 

Abnormal specimen.—This species shows considerable variation in 
exterior as well as interior features. Such variability is to be expected, 
but the occurrence of a freak specimen having the crural processes 
united is unusual. This specimen is paratype U.S.N.M. No. 1241952. 
The beak is broken and most of the brachial valve broke from the 


NO. 4 CRETACEOUS BRACHIOPODA, ARIZONA—COOPER 7 


specimen during the etching and was not recovered. Unfortunately 
the loop is thus revealed from the dorsal side which is not the most 
advantageous view for appreciation of the structure. 

The loop is of the normal size as shown by other specimens. The 
descending branch is very stout and the transverse band is strong and 
thick. The crural processes appear to have been normal but the points 
grew inward and united to form a transverse band, thinner than the 
anterior one but with the band convex toward the pedicle valve and 
having a form like that of the normal ribbon. 


GEMMARCULA ARIZONENSIS Cooper, new species 
Plate 2A, figures 1-28; plate 4B, figures 3-6 


Shell small, attaining a width of slightly more than one-half inch, 
transversely elliptical in outline; wider than long and with a narrow 
hinge. Widest at about the middle. Sides rounded; anterior margin 
subnasute to broadly rounded; anterior commissure rectimarginate to 
faintly uniplicate, the uniplication clearly visible only in old specimens. 
Valves unequal in depth, the pedicle valve having the greater depth. 
Surface multicostate, the costae appearing in three generations. Costae 
numbering 20 to 24 on the front margin of an average adult. 

Pedicle valve moderately to strongly convex in lateral profile and 
broadly to strongly convex in anterior profile, the convexity in both 
profiles depending upon age. Umbo somewhat narrowly convex, the 
convexity continued anteriorly as an indistinct fold which is bounded 
somewhat indistinctly by two costae stronger than those surrounding 
them; median region swollen; flanks and anterior slope steep. Beak 
irregular from pedicle pressure against rough surface; interarea wide 
and long; foramen large and circular; deltidial plates disjunct in the 
young, conjunct in old specimens and forming a symphytium. 

Interior of the pedicle valve with short but stout dental plates, 
strong transverse teeth; callosity of pedicle collar on floor of delthy- 
rial cavity thick; median septum low, extending anteriorly to beyond 
the valve middle. 

Brachial valve gently to moderately convex in Jateral profile, broadly 
but gently convex in anterior profile; umbo gently convex, often 
abraded by pedicle pressure against the substratum. Fold barely per- 
ceptible except in old specimens, and usually defined by a median 
crowding and smaller size of the costellae. Flanks gently swollen and 
with long, gentle slopes to the margins. 

Interior of the brachial valve with a thick concave notothyrial cal- 
losity buttressed by a strong median septum that extends to about the 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


valve middle; plates bounding sockets broad; outer socket ridges 
broad ; sockets long and deep. Crura short; crural processes short and 
pointed ; loop having form of early dallinid development, braced at its 
junction with the septum by a wide plate concave toward the anterior ; 
descending branches slender; ascending branches broad; transverse 
ribbon broad and with two prongs on each side which are directed 
toward the pedicle valve. Hinge plate usually obscured by callus which 
smooths the notothyrial region ; cardinal process a wide, bilobed thick- 
ening on the posterior margin of the notothyrial platform formed by 
the callus covering the hinge plate. 

Development of the loop.—The smallest specimens showing interior 
details are 2.0 mm. wide (paratype U.S.N.M. No. 1242004) and 2.7 
mm. (paratype U.S.N.M. 124198a). In these specimens the septum 
is continuous from the hinge platform to the valve middle where it is 
expanded ventrally toward the pedicle valve to form the pillar. Viewed 
from the anterior the expanded pillar is divided by a groove and the 
ventral and dorsal sides diverge slightly from each other, the begin- 
nings of the loop ring. Crural processes and descending branches are 
either not preserved or not yet formed, probably the former. The 
notothyrial region is filled with solid callus. 

The next larger specimen is 3.6 mm. in width (paratype U.S.N.M. 
No. 124199). Both valves are preserved. The pedicle valve shows a 
wide and completely open delthyrium with no trace of the deltidial 
plates. Thickening on the floor of the delthyrial cavity is strong and 
the median ridge anterior to this thickening is well developed. Inside 
the brachial valve the notothyrial region is solid as in the preceding 
specimen, but the septum is short and high. The crural processes are 
well developed and the crura are short and thick. The descending 
lamellae join the lower or dorsad diverging branches of the anterior 
end of the pillar, the anterodorsal side of which is now more deeply 
cleft. The ventral portion of the septum is elongated and the two 
lamellae diverging widely from the pillar are roofed by a flat plate, 
the pre-campagiform hood of Elliott. 

A third specimen of 5 mm. width (paratype U.S.N.M. No. 1242014) 
also shows the median septum and its anterior expansion. This speci- 
men differs from the preceding one in having a definite concavity in 
the notothyrial callus, bounded by the crural bases. Other details of 
the loop can only be inferred. 

A fourth specimen 5.7 mm. in width (paratype U.S.N.M. No. 
124198b) shows additional details of the loop, but part must be in- 
ferred from ridges and remnants. The notothyrial callosity is thick 
and is buttressed by a strong median septum. The crura are very short 


NO. 4 CRETACEOUS BRACHIOPODA, ARIZONA—COOPER 9 


and the descending lamellae are moderately broad and extend from the 
crural processes anteriorly to join the median part of the curved lateral 
plates at the distal end of the septum. The incision at the anterior end 
is much deepened and the anterior ends of the broken loop ring are 
beginning to diverge widely. 

No specimens between 6 and 12 mm. preserving good details of the 
loop were taken from the acid residues. Specimens 12 mm. wide or 
wider evidently have adult loops, but none of them are complete. The 
lateral branches connecting the septum to the loop are broad, stout, and 
long, frequently being strengthened by a median triangular plate. The 
loop of an adult specimen 16.5 mm. wide (paratype U.S.N.M. No. 
124220) has a broad transverse ribbon with long ears. 


MEASUREMENTS IN MILLIMETERS 


Brachial Mid- Hinge Thick- 
Length length width width ness 


Holotype U.S.N.M. No. 124197a.. 10.1 8.8 12.3 8.7 5.2 
Paratype U.S.N.M. No. 124197b.. 13.1 II.1 13.5 10.4 8.7 
e ot WE2A2008'... (12:2 9.6 14.0 10.8 6.5 
is - “Fh *124200D 13-4 10.6 14.0 10.7 VAS 
bi “ ry) SEAA206G ETS 9.0 13.0 10.9 6.4 
“ if = AT 242000) 722 5.3 7.5 ? 4.2 
c as “+ B242060.0:3.. 525 4.3 5.8 5.9 2.6 


Types——Holotype U.S.N.M. No. 124197a; figured paratypes 
U.S.N.M. Nos. 124197b, d-j, 124198a, b, 124199, 1242014, b, 124220; 
measured paratypes U.S.N.M. Nos. 124206 a-e; unfigured paratypes 
U.S.N.M. Nos. 124197c, 124200<, b. 

Horizon and locality.—Mural limestone, from a small hill 300 yards 
east of U. S. Highway 80, NWiSW4NE} sec. 36, T. 23 S., 
R. 25 E., Bisbee Quadrangle, Arizona; Rancho Nuevo, 3 miles east of 
Santa Rosalia, Sonora, Mexico. 

Discusston.—This species is characterized by having a low and in- 
distinct fold and sulcus and differs from all other described species of 
Gemmarcula in this respect. This is the first report of the genus in 
North America. Gemmarcula arizonensis is about the same size as 
G. aurea, type species of the genus, and has a cardinal process like it, 
but the exterior is different as noted above. Gemmarcula menardi 
(Lamarck), a well-known species in France and Great Britain, is 
larger than the American species and has a much more pronounced 
fold and sulcus. It also differs from the Arizona shell in having a 
more elaborate cardinal process. 


Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


GEMMARCULA MENARDI (Lamarck) 
Plate 2B, figures 29-34 


Figures of the interior and exterior of this fine species are intro- 
duced for comparison with G. arizonensis. Note elaborate develop- 
ment of the cardinal process in this European species. 

Horizon and locality.—Cretaceous (Green Sand), LeMans, Sarthe, 
France. 


PSILOTHYRIS Cooper, new genus 


Generally small to medium size, oval to subpentagonal in outline; 
anterior commissure rectimarginate to uniplicate; valves unequally 
convex, the pedicle having the greater depth and convexity. 

Pedicle valve with strong beak ridges; beak erect; foramen round, 
small to large, submesothyrid to mesothyrid; deltidial plates disjunct 
te conjunct, often worn away by pedicle pressure. Interior with stout, 
thick teeth, buttressed by stout dental plates; no pedicle collar; muscle 
scars not discernible in available material. 

Cardinalia small; hinge plate undivided, short, often upturned on its 
anterior edge; inner socket plate concave, stout; crura short, stout; 
crural process long and slender ; loop simple, long dalliniform, forming 
a broad ribbon and having a broad transverse band in the adult; 
median septum of the adult short, slender, and may or may not but- 
tress the hinge plate, and reduced to a faint myophragm at the valve 
middle. Young stages with loop metamorphosis like that of the Dal- 
linidae. 

Type species.—Psilothyris occidentalis Cooper, new species. 

Discussion—This species is characterized by its smooth exterior, 
simple uniplicate folding, short inner hinge plate and long dalliniform 
loop. Details of the formation of the hinge plate and the development 
of the loop are discussed under the specific description. The combina- 
tion of characters exhibited by Psilothyris is different from any de- 
scribed Cretaceous dallinoid and is also unlike any described smooth 
Jurassic zeilleroid and dallinoid. 

Of smooth Jurassic zeilleroids that resemble Psilothyris more or less 
closely, Zeilleria, Microthyridina, Rugitela, and Ornithella are pro- 
vided with a strong median septum in the brachial valve and the fold- 
ing of all of them is different from that of Psilothyris. Aulacothyris 
and Antiptychina are differently folded, these two genera having 
a strongly sulcate brachial valve. The dallinoids Plesiothyris and 
Obovothyris have long septa and different folding. Epicyrta has a 
carinate brachial valve and is thus quite different externally. 





NO. 4 CRETACEOUS BRACHIOPODA, ARIZONA—COOPER II 


This genus, although not named until now, was recognized by 
Deslongchamps (1884, p. 189) in his discussion of the genus Zeilleria. 
He characterizes the division as having a relatively short, thick beak 
having a very large foramen. The shell is globular, short, and compact. 
It is unique in the Cretaceous. 


PSILOTHYRIS OCCIDENTALIS Cooper, new species 
Plate 3B, figures 4-24; plate 4A, figures 1, 2 


Shell small, attaining a length of five-eighths inch; outline sub- 
pentagonal with the length slightly greater than the width; greatest 
width located slightly posterior to the middle; sides sloping medially ; 
anterior margin subtruncate; posterior margin forming an obtuse 
angle. Anterior commissure uniplicate; lateral commissure straight. 
Valves unequal in depth, the pedicle valve deeper; surface smooth 
except for concentric lines and varices of growth. 

Pedicle valve strongly convex in lateral profile, with the maximum 
convexity slightly posterior to the middle; anterior profile strongly 
convex ; umbonal region inflated ; beak small, erect ; beak ridges strong. 
Median region swollen; anterior slope flattened; flanks swollen and 
steep. Foramen small, round, mesothyrid, slightly labiate. Deltidial 
plates conjunct, suture visible. Interior of pedicle valve with large 
teeth supported by stout dental plates. No pedicle collar. Muscle 
marks lightly impressed. 

Brachial valve in lateral profile flattened in the median region but 
convex at the posterior and anterior; anterior profile broadly and 
gently convex. Umbonal region swollen but median area flattened ; 
flanks narrowly convex. 

Interior of the brachial valve with short, undivided hinge plate 
deeply excavated anteriorly and thickened, elevated or puckered on the 
anterior edge. Socket ridge short, stout. Crura short; crural processes 
long and slender in the adult loop but short and blunt in the young. 
Loop long and free in the adult; loop short, broad, and attached to a 
short septum on the floor of the valve in the young. Septum in the 
adult short and confined to a position at the beak and under the hinge 
plate and may or may not support the hinge plate. 

Development of the loop.—In the smallest specimen available, 
measuring 3 mm. in length and probably the same in width (paratype 
U.S.N.M. No. 124190k), the notothyrial region is deeply concave 
and without a hinge plate. The crura are slender and arise from ridges 
bordering the notothyrial cavity. The descending lamellae are short 
and their anterior ends converge to unite with a septal blade or pillar 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


that springs from the floor of the valve near the middle. This pillar 
is much expanded longitudinally with free edges extending a short 
distance posteriorly, but also anteriorly to a point about three-fourths 
the length of the valve from the beak. The anterior extension appears 
to be a long, broad-ribboned ring. The details cannot be ascertained 
because this part of the structure is obscured by silicious material. 
This is the pre-campagiform stage. 

Details of the cardinalia are clear in a specimen without loop meas- 
uring 4.2 mm. in length (paratype (U.S.N.M. No. 124202d). The 
floor of the notothyrium is thickened by ridges joining the crural bases 
and the septum is now extended posteriorly to meet the center of these 
lateral ridges (pl. 3B, fig. 13). The expanded anterior end of the 
septum is free, but no other details are available. 

A specimen measuring 4.5 mm. in diameter (paratype U.S.N.M. 
No. 124202g) is somewhat more advanced than the previous one. The 
notothyrial cavity is now deeply concave, the lateral extensions thick- 
ened and anteriorly excavated to simulate a hinge plate supported by 
the median septum. The crural processes on the loop are well devel- 
oped and are located just anterior to the hinge plate. The descending 
branches of the loop attach to the distal expanded end of the septum 
near its dorsal extremity. The greatest change has taken place at the 
free part of the septum, the anterior end of which is distinctly divided 
by an incision in its anterior end and the remains of a ring mounting 
the ventral edge is clearly visible. This is probably the frenuliniform 
stage of development. 

A still more advanced stage, the terebrataliform stage, is shown by 
a specimen 6.6 mm. in diameter (paratype U.S.N.M. No. 1242021). 
The hinge plate is still deeply concave and the crural processes moder- 
ately long. The septum is well developed and high but does not now 
reach to the middle. The descending branches of the loop are broad 
and are now extended far anterior to the end of the septum. The loop 
is nevertheless still attached to the distal end of the septum by two 
short branches, but the septum does not extend anterior to its point 
of contact (plate 3B, figure 17). The specimen does not preserve a 
ring or ascending branch, but remnants of it are visible. 

The next specimen of the series is 7.6 mm. in length and slightly 
less in width (paratype U.S.N.M. No. 124190h). The hinge plate has 
become considerably shallower by anterad growth of a transverse 
plate at its anterior end; the crural processes are large and the branch 
between them and the hinge plate is now nearly obsolete. The descend- 
ing branches are free of the septum, but projections, which face in- 
ward and represent the remnant of the septal attachments, appear at 





NO. 4 CRETACEOUS BRACHIOPODA, ARIZONA—COOPER 13 


about their middle. The septum has been absorbed to a mere remnant 
which extends for a short distance only anterior to the hinge plate. 
Except for the incompletely developed hinge plate and the remnants 
of the septal attachments, the loop is essentially adult in character. 
This stage is the dalliniform stage. 

By summarizing the evidence from these few specimens it is possible 
to give a fairly complete account of the loop development. Prior to 
3 mm. the median septum must show as a small projection from the 
floor. At 3 mm. the septum has become elongated and the descending 
branches have grown anteriorly to meet the sides of the elongated free 
distal expanded part of the septum and the ring bud starts to develop. 
By 44 mm. the expanded end of the septum splits laterally and the 
ring enlarges, the septum having a deep reéntrant anteriorly and the 
descending branches extended a considerable distance anteriorly. At 
6.6 mm. the loop is strong, with broad descending branches extended 
beyond the anterior end of the septum, and the loop attachment to the 
septum is a small process. At 7.6 mm. the loop is now free of the 
septum which has become nearly completely resorbed except for the 
short remnant supporting the hinge plate. Remnants of the process 
attaching the loop to the septum can be seen in specimens having at- 
tained a length of 12.5 millimeters. 

The hinge plate is deeply concave in the young, but in the 7.6 mm. 
stage the transverse plate forming the flat and undivided hinge plate 
forms, and this lengthens with advancing age. In old age it becomes 
puckered or upturned on its free edge. 


MEASUREMENTS IN MILLIMETERS 


Brachial 
Length length Width Thickness 
Holotype U.S.N.M. No. 124191....... 15.1 12.4 13.9 9.7 
Paratype U.S.N.M. No. 124189....... 13.3 II.0 12.2 8.1 
° f of eEZATOOD 55 -10)512 11.4 9.6 10.7 6.0 
: . oh DPA TOO Cs tis.e 12.4 10.4 11.7 7.5 


Types.—Holotype U.S.N.M. No. 124191; figured paratypes 
U.S.N.M. Nos. 1241902, c, e, f, h-j, 124202a, d-i; measured paratypes 
U.S.N.M. Nos. 124189, 124190, b ; described but unfigured paratypes 
U.S.N.M. No. 124190k; unfigured paratypes U.S.N.M. Nos. 124190d, 
g, 124202b. 

Horizon and locality —Mural limestone, from a small hill 300 yards 
east of U. S. Highway 80, NW4SW4iNE# sec. 36, T. 23 S., 


14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


R. 25 E., Bisbee Quadrangle, Cochise County, Arizona; Rancho 
Nuevo, 3 miles east of Santa Rosalia, Sonora, Mexico. 

Discussion.—This species is characterized by its compact form, 
small foramen, moderately strongly uniplicate commissure, flatly con- 
vex brachial valve and deep pedicle valve. No other species of this 
genus is now known in North America to which this one can be com- 
pared. The species most like P. occidentalis outside of North America 
is “Waldheimia”’ tamarindus (Sowerby) from the British Isles. A 
variety of forms now appears under this name in the British Isles, | 
but specimens of P. tamarinda from Faringdon, England, are most 
like the American species. They differ, however, in having a narrower 
anterior region, a larger foramen and much less convex pedicle valve. 
The development of the median septum in the interior of both the 
British and American forms is very slight. 


PSILOTHYRIS TAMARINDA (Sowerby) 
Plate 3C, figure 25 


The interior of a brachial valve is introduced for comparison with 
P. occidentalis. Note the low and reduced median septum and the 
short undivided hinge plate. 

Horizon and locality—Cretaceous (Aptian—Lower Green Sand), 
Faringdon, Berkshire, England. 


REFERENCES CONSULTED 
Bosguet, J. . 

1859. Monographie des brachiopodes fossiles du Terrain Crétacé superieur 

du Duché de Limbourg. Haarlem. 
Coppan, W. A., and Reesipe, J. B., Jr. 

1952. Correlation of the Cretaceous formations of the western interior of 
the United States. Bull. Geol. Soc. America, vol. 63, No. 10, pp. 
IOII-1044. 

Davinson, T. 

1852-1855. A monograph of the British fossil Brachiopoda. Part 2. The 
Cretaceous brachiopods. Palaeontographical Soc. 

1874. A monograph of the British fossil Brachiopoda. Vol. 4, Part 1, Sup- 
plement to the Recent, Tertiary and Cretaceous species, pp. 17-72. 

DESLONGCHAMBS, E. E. 

1884. Etudes critiques sur des brachiopodes nouveaux ou peu connus. Art. 8. 
Note sur les modifications a apporter a la classification des 
Terebratulidae. 

Extrort, G. F. 

19047. The development of a British aptian brachiopod. Proc. Geol. Assoc., 

vol. 58, pt. 2, pp. 144-159. 
Mutir-Woop, H. M. 

1934. On the internal structure of some Mesozoic Brachiopoda. Philos. 

Trans. Roy. Soc. London, ser. B, vol. 223, pp. 511-567. 





NO. 4 CRETACEOUS BRACHIOPODA, ARIZONA—COOPER 15 


RANsSOME, F. L. 
1904. Bisbee Folio, Arizona. U. S. Geol. Surv. Atlas, No. 112. 
SAHNI, M. R. 
1929. A monograph of the Terebratulidae of the British Chalk. Palaeonto- 
graphical Soc. (1927). 
Stoyanow, A. 
1949. Lower Cretaceous stratigraphy in southeastern Arizona. Geol. Soc. 
Amer. Mem. 38. 
TuHomson, J. A. 
1927. Brachiopod morphology and genera (Tertiary and Recent). New 
Zealand Board of Sci. and Art, Manual 7. 


EXPLANATION OF PLATES 


PLATE I 


A. Cyclothyris americana Cooper, new SpecieS........ccccccccsccccccecees 3 

1-5, Anterior, posterior, brachial, side, and pedicle views, respec- 
tively, X 1, of holotype U.S.N.M. No. 124193a. 6, Brachial 
view of the holotype, * 2, showing elevated rim on deltidial 
plates around the foramen. 7, Beak of young pedicle valve, 
<2, showing teeth and rim around foramen, paratype 
U.S.N.M. 124186c. 8, 9, Interior of pedicle valve x 2, and 
same tilted, showing muscle scars, thickened deltidium (del- 
tidial plates fused), and pallial marks, paratype U.S.N.M. 
No. 124186d. 10, 17, Cardinalia seen from the anterior and 
tilted, & 3, to show articulation, and anterior surface of the 
radulifer crura, paratype U.S.N.M. No. 124186a. 11, 12, Pos- 
terior of a young specimen, X 4, showing elevated rims on 
foramen before formation of the deltidium, paratype U.S.N.M. 
No. 124186e. 13, Interior of the brachial valve showing 
divided hinge plate, radulifer crura, and pallial marks, X 2, 
paratype U.S.N.M. No. 124186b. 14, Same enlarged, X 3, to 
show crura in greater detail. 15, Same tilted to the side, show- 
ing crura, X 3. 16, Same, tilted to show posterior surface of 
hinge plate, X 3. Mural limestone, from a small hill 300 yards 
east of U. S. Highway 80, NWiSW4iNE+ sec. 36, T. 23 S., 
R. 25 E., Bisbee Quadrangle, Cochise County, Arizona. 

B. Rectithyris vespertina Cooper, new SPeCicS...........ccceeecscccccccce 4 

18-22, Pedicle, posterior, brachial, side, and anterior views, re- 
spectively, of a large specimen, I, paratype U.S.N.M. No. 
124194c. 23-27, Pedicle, posterior, anterior, brachial, and side 
views, respectively, 1, of holotype U.S.N.M. No. 124194b. 
28, Brachial view, 1, of an immature specimen, paratype 
U.S.N.M. No. 124194d. 209, Brachial view of another paratype, 
< 2, U.S.N.M. No. 124187. 30, 31, Pedicle view of the pre- 
ceding specimen with window cut in pedicle valve to show 
loop, X 1 and X 2, respectively. 32, Interior of a brachial 
valve showing hinge plate, socket ridges, crural bases, and 
indistinct adductor scars, & 3, paratype U.S.N.M. No. 124188. 
33, Interior of a young brachial valve, & 3, showing a loop in 
an advanced stage, paratype U.S.N.M. No. 124196c. 34, In- 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Page 

terior view of an adult brachial valve showing fully matured 
loop, X 2, paratype U.S.N.M. No. 124195. 35, The preceding 
loop enlarged, XX 4, to show mode of attachment to crural 
bases and socket ridges, and showing the small inner hinge 
plates. 36, Interior of another specimen tilted to side with 
brachial valve broken off and showing loop with two trans- 
verse bands, X 2, paratype U.S.N.M. No. 124195a. 37, Same 
specimen tilted away from observer to show the two trans- 
verse bands, X 3. Horizon and locality same as above. 


PLATE 2 


A. Gemmarcula arizonensis Cooper, new SPeCi€S......... 2 cece eee eee cece 7 

1-5, Anterior, posterior, side, brachial, and pedicle views, re- 
spectively, of a complete specimen, 1, paratype U.S.N.M. 
No. 124197b. 6-10, Brachial, side, posterior, anterior, and pedi- 
cle views, respectively, X 2, of holotype U.S.N.M. No. 124197A. 
11, Complete specimen tilted to show interarea and deltidium 
(= united deltidial plates), 3, paratype U.S.N.M. No. 
124197d. 12,View of the interarea and disjunct deltidial plates 
of a young specimen, X 3, paratype U.S.N.M. No. 1241097¢. 
13, Posterior part of a pedicle valve tilted to show callosity 
on floor of delthyrial cavity, low median septum, and discrete 
deltidial plates, X 2, paratype U.S.N.M. No. 124197h. 14, Im- 
mature pedicle valve, & 4, showing deltidial plates just form- . 
ing and callosity on floor of delthyrial cavity, paratype 
U.S.N.M. No. 124201a. 15, Interior of an incomplete brachial 
valve, X 3, paratype U.S.N.M. No. 124197e, showing cardi- 
nalia. 16, Interior of a brachial valve younger than the preced- 
ing, X 4, showing cardinalia, paratype U.S.N.M. No. 1241097f. 
17, Interior of an old specimen showing deeply excavated 
notothyrial callosity simulating a concave hinge plate, crura 
and crural processes, X 3, paratype U.S.N.M. No. 124107. 
18, 19, Tilted and interior views of a young brachial valve 
showing hooded pre-campagiform stage of loop, x 4, para- 
type U.S.N.M. No. 124199. 20, Interior of the pedicle valve 
of the preceding specimen, with open delthyrium, X 4. 21, A 
young specimen, X 4, showing beginning of ring and attach- 
ments of descending lamellae of loop, paratype U.S.N.M. No. 
124198a. 22, A still larger specimen than the preceding show- 
ing deeper cleft in remnant of ring, X 4, paratype U.S.N.M. 
No. 124198b. 23, Young brachial valve tilted to show cleft in 
loop ring, X 4, opposite to pedicle valve shown in figure 14, 
paratype U.S.N.M. No. 124201b. 24, Specimen with pedicle 
valve partially removed to show attachment of loop to septum 
and long anterior branches of the descending lamellae, X 2, 
paratype U.S.N.M. No. 124197}. Mural limestone, from a 
small hill 300 yards east of U. S. Highway 80, NW4SW34- 
NE} sec. 36, T. 23 S., R. 25 E,, Bisbee Quadrangle, Cochise 
County, Arizona. 





NO. 4 CRETACEOUS BRACHIOPODA, ARIZONA—COOPER 17 


Page 
25-28, Interior, posterior, side tilted, and interior posteriorly 
tilted views, respectively, of an adult specimen showing 
terebrataliform loop, 2, paratype U.S.N.M. No. 124220. 
Mural limestone, Rancho Nuevo, 3 miles east of Santa Ro- 

salia, Sonora, Mexico. 

BaGemmarcuia menarat CLamarck igh {icles soicileals uo aaeiela cle Jodie dees 10 
29, 30, Brachial and side views of a complete specimen, x 1, 
for comparison with Gemmarcula arizonensis, hypotype 
U.S.N.M. No. 124223c. 31, Interarea of the pedicle valve, X 2, 
showing symphytium (or deltidium), teeth, and foramen, 
hypotype U.S.N.M. No. 124223a. 32, 33, Interior and pos- 
terior views of the brachial valve showing complicated car- 
dinal process, x 2, hypotype U.S.N.M. No. 124223b. 34, 
Brachial interior showing part of loop, & 2, for comparison 
with G. arizonensis, counterpart of pedicle valve shown by 
figure 31. Cretaceous (Green Sand), Le Mans, Sarthe, France. 


PLATE 3 


A. i\Cramscus hespersus Cooper. Mew Species. & jcc) seid. od alesieieslo wien bs a0 6s 2 

I, 2, Brachial and side views, X 1, of holotype U.S.N.M. No. 
124192. 3, Interior of the preceding, X 2, showing transverse 
muscle scars. Mural limestone, from a small hill 300 yards 
east of U. S. Highway 80, NWiSW1iNE} sec. 36, T. 23 S., 
R. 25 E., Bisbee Quadrangle, Cochise County, Arizona. 

B. Psilothyris occidentalis Cooper) new SpeCieS: ss... 3 occ Set etl ees II 

4-8, Anterior, posterior, side, brachial, and pedicle views, respec- 
tively, X 2, of holotype U.S.N.M. No. 124191. 9, 10, Brachial 
and anterior views of another individual, showing uniplicate 
anterior commissure, X 1, paratype U.S.N.M. No. 124190a. I1, 
Side view of a specimen broken to show descending branches 
of the long dalliniform loop and part of the wide ascending 
branch, 2, paratype U.S.N.M. No. 124202a. 12, Interior of 
the pedicle valve showing deltidium (united deltidial plates), 
submesothyrid foramen, and teeth, paratype U.S.N.M. No. 
124190c. 13, 14, Two small specimens in pre-campagiform 
stage, X 4, showing the median septum and pillar of an early 
juvenile stage, paratypes U.S.N.M. No. 124202e, d. 15, An- 
other juvenile specimen in the same stage as the preceding 
showing part of descending branch attached to pillar, x 4, par- 
atype U.S.N.M. No. 124202h. 16, Another juvenile specimen, 
probably in frenuliniform stage, tilted slightly to the side and 
showing descending branches attached to pillar and part of 
loop ring, X 4, paratype U.S.N.M. No. 124202g. 17, Imma- 
ture specimen in terebrataliform stage showing descending 
branches of loop attached to median septum and the deep cleft 
at the anterior of the loop, * 4, paratype U.S.N.M. No. 
1242021. 18, A specimen older than the preceding and in the 
dalliniform stage showing receded median septum but rem- 
nants of processes of attachment on the descending branches 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Page 

of the loop, X 4, paratype U.S.N.M. No. 124202f. 19, A 

slightly larger specimen than the preceding in dalliniform 

stage showing descending branches and remnantal septal at- 

tachments, 4, paratype U.S.N.M. No. 124190h. 20, Frag- 

ment of brachial valve showing hinge plate, sockets, and 

socket ridges, X 4, paratype U.S.N.M. No. 124190j. 21, In- 

terior of another brachial valve, 4, showing adductor scars 

indistinctly, paratype U.S.N.M. No. 124190e. 22, Cardinalia 

of an adult brachial valve, & 4, showing crural processes 

and hinge plate with upturned edge, paratype U.S.N.M. No. 

I24190i. 23, 24, Two views, X 4, of the interior of an adult 

brachial valve, one (23) tilted and the other not, showing 

hinge plate socket ridges, almost obsolete median septum, and 

also the dental plates of the pedicle valve, paratype U.S.N.M. 

No. 124190f. Horizon and locality same as above. | 

C. Psilothyris tamarinda. (Sowerby ) s 1.5 2010 sis 2.0) ein ds amioldee sig aiateaiete eye 14 

25, Interior of the brachial valve showing cardinalia and nearly 

obsolete median septum, X 3, figured specimen U.S.N.M. No. 

128222. Cretaceous (Aptian—Lower Green Sand), Faring- 

don, Berkshire, England. 


PLATE 4 


A. Psilothyris occidentalis Cooper, new SPeCieS... 0. 20s. .cc cere semerceses II | 
I, 2, Side and interior tilted views of a specimen in pre-campagi- 
form stage 4.5 mm. long showing the pillar before growth of 
median septum, and descending lamellae, about * 14 and & Io, 
respectively, paratype U.S.N.M. No. 124202g. Mural lime- 
stone, a small hill 300 yards east of U. S. Highway 80, NW3}- 
SW4iNE3 sec. 36, T. 23 S., R. 25 E., Bisbee Quadrangle, 
Cochise County, Arizona. 
B. Gemmarcula arigonensis Cooper, new SPeCieS............eececceececeese 7 
3, 4, Interior and side views of a brachial valve 3.6 mm. wide 
showing remnants of pre-campagiform hood, pillar, and begin- 
ning of loop ring, about X 17, paratype U.S.N.M. No. 124190. 
5, 6, Side and interior views of brachial valve showing the 
adult loop, about X 4.5, paratype U.S.N.M. No. 124220. Hori- 
zon and locality same as in text; (3) and (4) from Arizona, 
(5) and (6) from Sonora, Mexico. 


Abbreviations 


ab = ascending branch of loop g = groove of pillar 
cb = crural base p = septal pillar 
cp = cardinal process pf = remnant of pre-campagiform flange 
dl = descending branch of loop s = septum 
e = ear of loop sdl = scar of broken descending lamella 


(Drawings by Lawrence B. Isham.) 


VOL. 131, NO. 4, PL. 1 


Ze 





CYCLOTHYRIS AND RECTITHYRIS 


(SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 4, PL. 


AS 
BN 


1 


ap 





GEMMARCULA 


(SEE EXPLANATION AT END OF TEXT.) 


;THSONIAN MISCELLANEOUS COLLECTIONS 


VOL. 131, NO. 4, PL. 3 





CRANISCUS AND PSILOTHYRIS 


(SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 4, PL. 





PSILOTHYRIS AND GEMMARCULA 


(SEE EXPLANATION AT END OF TEXT.) 
. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 5 


Charles D. and Mary Waux CAalcott 
Research Fund 


Arermeeh-List OF THE FOSSIL AND 
PREHISTORIC. BIRDS. OF: NORTH 
AMERICA AND THE 
WEST INDIES 


By 
ALEXANDER WETMORE 


Research Associate, Smithsonian Institution 





(PusiicaTion 4228) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
JANUARY 25, 1956 


THE LORD BALTIMORE PRESS, INC. 
BALTIMORE, MD., U. S. A. 





Charles D. and Mary Waux Walcott Research Fund 


AYCHECK-LIST OF THE'FOSSIL; AND PRE- 
MISTORIC BIRDS" OF? NORTH AMERICA 
AND THE WEST INDIES 
By ALEXANDER WETMORE 


Research Associate, Smithsonian Institution 


The present check-list is an amplification of the one published in 
the Smithsonian Miscellaneous Collections in 1940 (vol. 99, No. 4) 
and is complete to November 1955 so far as records have come to at- 
tention. To the present time these check-lists have covered the area of 
the check-list of living birds of the American Ornithologists’ Union, 
namely North America north of México, with the addition of Baja 
California. It has seemed desirable now to include also the records, 
comparatively few in number, for México and the West Indies, since 
this information is complementary and otherwise is available only in 
widely scattered sources. Various of these latter records are of species 
of birds described from bones found during archeological excavations 
in Indian kitchen middens of pre-Columbian age or during the ex- 
ploration of caverns. The species concerned have long been extinct, so 
that the only knowledge regarding them is embodied in their skeletal 
remains. No living examples have been known. It is useful therefore 
to include them for reference with other species of fossil status, since 
they do not figure in check-lists of existing birds and since possibly 
they may be encountered at some future time in true fossil form. They 
have the same pertinence therefore as species described from Pleisto- 
cene beds whose bones have been found subsequently in Recent 
deposits. 

The considerable amount of information now available has allowed 
more detail relative to geological formations from which the various 
records have come, and these data have been brought down to date as 
far as practicable. In this I have had the advice in certain cases of 
Druid Wilson, of the U. S. Geological Survey, and also have profited 
from discussions with Dr. C. Wythe Cooke of the same service, par- 
ticularly as to formations of the southeastern United States. 

In the records from the Pleistocene there has been sufficient study 
of the deposits of this age known from the western United States to 
allow indication of position, as to whether they are considered early or 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 131, NO. 5 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


late, of most of the faunas. The situation in Florida is not so clear. 
Bone beds at Melbourne and Vero overlie the Anastasia formation, 
a marine Pleistocene deposit, and therefore are considered late Pleisto- 
cene. Apparently a newer find at Haile in Alachua County may be 
from a similar level. The Seminole Field in Pinellas County also ap- 
pears to overlie the beds of the west coast of Florida that are con- 
sidered equivalent to the Anastasia, if not exactly the same formation. 
However, Pliocene exposures are near at hand so that the sequence, 
from present knowledge, is not clear-cut as it is at Melbourne. In- 
formation relative to the localities at Bradenton, Sarasota, and on the 
Itchtucknee River is far from definite, and other deposits found in 
caverns, while evidently Pleistocene, are still more uncertain as to 
actual relationship within that period. Collecting continues actively in 
the Florida Pleistocene, and presently there should be accumulated 
sufficient data on the avifauna to permit a reasonable correlation. In 
the meantime it has seemed better to list all the Florida records as 
Pleistocene without attempt to indicate the level. To list Melbourne 
and Vero alone, for example, as late Pleistocene might be misleading. 

Recent investigations of Dr. Joseph T. Gregory (Condor, 1952, 
pp. 73-88) have changed measurably the time-honored concept in 
which the species of IJchthyornis have been associated with the 
Hesperornis group in a superorder (Odontognathae) of the Neorni- 
thes, characterized by the possession of teeth. The skull of [chthyor- 
nis always has presented an anomaly in that the teeth were in sockets 
instead of in grooves as in Hesperornis. Further, the mandible, or 
lower jaw, was unduly large in comparison with the rest of the skull 
and the body skeleton. Dr. Gregory has shown that the jaws attributed 
to Ichthyornis in reality are reptilian and are those of a small 
mosasaur. 

These conclusions destroy the main reasons for the association of 
Ichthyornis and Hesperornis in one superorder, though still leaving 
Ichthyornis apart from birds known from later periods to the present, 
in the biconcave vertebrae. In preliminary consideration it seemed 
that it might be desirable in the classification to cancel the category 
of superorders, but on further consideration it appears useful to 
emphasize the considerable and definite differences that separate 
Hesperornis, Ichthyornts, and the penguins from each other and from 
other groups of birds. This may be accomplished through a new 
superorder Ichthyornithes for the order Ichthyornithiformes, leaving 
Hesperornis and those others placed near it in the Odontognathae. 
This will serve as stated above to call attention to the existing peculi- 
arities of these groups and will give a balanced treatment. 


tl 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 3 


The family Mancallidae is added for the two species of Mancalla at 
present recognized, since resemblance between these and the great auk 
appears due to convergence. The two west-coast forms differ from 
other auks in the marked modification of the wing for use as a flipper. 
The genera Paloelodus and Megapaloelodus have been placed with the 
typical flamingos in the Phoenicopteridae, a group to which they are 
unquestionably related. Dr. Hildegarde Howard recently pointed out 
their differences in the shorter, heavier metatarsus, nonpneumatic 
femur, and different form in the tibiotarsus and has proposed the 
family Paloelodidae. To the differences outlined by Dr. Howard 
there may be added the form of the bill, which, to judge from one 
incomplete specimen of Paloelodus ambiguus Milne Edwards of the 
Oligocene of western Europe, was gooselike and not bent downward 
as in the true flamingos. It may be noted also that the toes in Paloe- 
lodus were definitely longer. 

The modern species that occur in the fossil record are distinguished 
from those not known in living form by the inclusion of a common 
name in the heading and the statement that the bird is one found in 
modern form. Most of these are listed under specific scientific names 
without regard to local race, since most subspecies may not be identi- 
fied from bones. It is extremely doubtful procedure in most instances 
to assume that Pleistocene subspecies were the same as those en- 
countered in the region today, and assumption of race is made only 
where there is reasonable certainty of the identification. The specific 
names therefore are used in an inclusive sense, though it is evident in 
wide-ranging groups that two or more subspecies may be covered in 
the fossil record, for example, in the ruffed grouse, Bonasa umbellus, 
where bones identified as this species are known from such widely 
separated localities as Maryland and California. This should be under- 
stood particularly in cases like that of the raven, Corvus corax, or 
marsh hawk, Circus cyaneus, where the range extends to other con- 
tinents. 

The present list gives the record of 189 forms still living, and of 
248 species recorded only in an extinct state, this including 11 kinds 
known only from bones in cave or midden deposits of Recent age. 
There remain the 12 additional names of uncertain status listed at the 
end under the heading INCERTAE sEpIS. The increase from the 165 
modern forms and 184 extinct species of the list of 1940 is indicative 
of the growth in knowledge in this field during the comparatively brief 
interval of 15 years but reveals only part of the increase since many 
additional records have been found for numerous living species in- 
cluded in 1940. 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Class AVES: Birps 
Subclass NEORNITHES: True Birps 
Superorder ODONTOGNATHAE: New Wortp Tooruep Birps 
Order HESPERORNITHIFORMES: HeEsperorNitTHES 
Family HESPERORNITHIDAE: HEsperorNiTHES 
Genus HESPERORNIS Marsh 


Hesperornis Marsu, Amer. Journ. Sci., ser. 3, vol. 3, 1872, p. 360. Type, by 
monotypy, Hesperornis regalis Marsh. 


Hesperornis crassipes (MarsH) 
Lestornis crassipes Marsu, Amer. Journ. Sci., ser. 3, vol. 11, 1876, p. 500. 
Upper Cretaceous (Niobrara formation) : Western Kansas. 


Hesperornis montana SHUFELDT 


Hesperornis montana SHuFe pt, Auk, vol. 32, No. 3, July 1915, p. 293, pl. 18, 
figs. 4, 6, 8, 10, 12. 


Upper Cretaceous (Claggett formation): 1 mile above mouth of 
Dog Creek, Fergus County, Montana. 
Hesperornis regalis MARSH 
Hesperornis regalis Marsu, Amer. Journ. Sci., ser. 3, vol. 3, 1872, p. 357. 
Upper Cretaceous (Niobrara formation): Smoky Hill River, 20 
miles east of Wallace (type locality), and Two Mile Creek, Smoky 
Hill River, Logan County, Kansas. 
Hesperornis gracilis MArsu 1 
Hesperorms gracilis Marsu, Amer. Journ. Sci., ser. 3, vol. 11, 1876, p. 510. 
Upper Cretaceous (Niobrara formation) : Near Smoky Hill River, 
western Kansas. 


Genus CONIORNIS Marsh 2 
Coniornis Marsu, Amer. Journ. Sci., ser. 3, vol. 45, 1893, p. 82. Type, by 


monotypy, Coniornis altus Marsh. 
Coniornis altus MArsu 


Coniornis altus Marsu, Amer. Journ. Sci., ser. 3, vol. 45, 1893, p. 82, text fig. 


Upper Cretaceous (Judith River formation) : Dog Creek, Fergus 
County, Montana. 


1 Gregory, Condor, vol. 54, No. 2, Mar. 26, 1952, p. 74, concludes that the genus 
Hargeria, erected for this species by Lucas, is not separable from Hesperornis. 

2 Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, February 1915, pp. 16, 
75, considers this a synonym of Hesperornis. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 5 


Family BAPTORNITHIDAE®: BaptornitHEs 
Genus BAPTORNIS Marsh 


Baptornis Marsu, Amer. Journ. Sci., ser. 3, vol. 14, 1877, p. 86. Type, by 
monotypy, Baptornis advenus Marsh. 


Baptornis advenus MArsH 
Baptornis advenus Marsu, Amer. Journ. Sci., ser. 3, vol. 14, 1877, p. 86. 


Upper Cretaceous (Niobrara formation): Wallace County (type 
locality), and Butte Creek, Logan County, Kansas. 


Superorder ICHTHYORNITHES: IcutTuyornis and ALLiEs 
Order ICHTHYORNITHIFORMES: Icutuyornis and ALLIEs 
Family ICHTHYORNITHIDAE: IcutTHyorNITHES 
Genus ICHTHYORNIS Marsh 

Ichthyornis Marsu, Amer. Journ. Sci., ser. 3, vol. 4, November 1872, p. 344. 

Type, by monotypy, Ichthyornis dispar Marsh. 
Ichthyornis agilis (MarsH) 
Graculavus agilis MarsH, Amer. Journ. Sci., ser. 3, vol. 5, 1873, p. 230. 


Upper Cretaceous (Niobrara formation): Butte Creek, Logan 
County, Kansas. 


Ichthyornis anceps (MarsH) 


Graculavus anceps Marsu, Amer. Journ. Sci., ser. 3, vol. 3, 1872, p. 364. 


Upper Cretaceous (Niobrara formation) : North Fork Smoky Hill 
River, Logan County, about 12 miles east of Wallace, Kansas. 


Ichthyornis dispar MArsH 


Ichthyornis dispar Marsu, Amer. Journ. Sci., ser. 3, vol. 4, 1872, p. 344. 


Upper Cretaceous (Niobrara formation): Near Solomon River, 
Kansas. 


Ichthyornis lentus (MarsH) 


Graculavus lentus Marsu, Amer. Journ. Sci., ser. 3, vol. 14, 1877, p. 253. 


Upper Cretaceous: Near McKinney, Texas. 


8 Lambrecht, Handb. Palaeorn., 1933, pp. 258-260, unites this with the family 
Enaliornithidae, on what seem insufficient grounds. As suggested by Lucas, 
Proc. U. S. Nat. Mus., vol. 26, 1903, p. 555, Baptornis probably belongs in a 
distinct order. 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Ichthyornis tener MArsH 
Ichthyornis tener Marsu, Odontornithes, 1880, pp. 151, 198, pl. 30, fig. 8. 
Upper Cretaceous (Niobrara formation) : Wallace County, Kansas. 


Ichthyornis validus MarsH 


Ichthyornis validus Marsu, Odontornithes, 1880, pp. 147, 153, 198, pl. 30, figs. 
II-14. 


Upper Cretaceous (Niobrara formation): Near Solomon River, 
Kansas. 


Ichthyornis victor Marsu 


Ichthyornis victor Marsu, Amer, Journ. Sci., ser. 3, vol. 11, 1876, p. 511. 


Upper Cretaceous (Niobrara formation): Wallace County (type 
locality), and Hackberry Creek, near Smoky Hill River, Gove 
County, Kansas. 


Family APATORNITHIDAE: ApatornitHEs 
Genus APATORNIS Marsh 


Apatornis Marsu, Amer. Journ. Sci., ser. 3, vol. 5, Jan. 21, 1873, p. 162. Type, 
by monotypy, Jchthyornis celer Marsh. 


Apatornis celer (Marsi) 


Ichthyornis celer Marsu, Amer. Journ. Sci., ser. 3, vol. 5, 1873, p. 74. 


Upper Cretaceous (Niobrara formation): Butte Creek, Logan 
County, near Smoky Hill River, Kansas. 


Superorder NEOGNATHAE: Typicat Birps 
Order CAENAGNATHIFORMES: CarenacnatHus 
Family CAENAGNATHIDAE: CarnacnatTHuus 
Genus CAENAGNATHUS Sternberg 


Caenagnathus STERNBERG, Journ. Pal., vol. 14, January 1940, p. 81. Type, by 
original designation, Caenagnathus collinsi Sternberg. 


Caenagnathus collinsi STERNBERG 4 
Caenagnathus collinsi STERNBERG, Journ. Pal., vol. 14, January 1940, p. 81, 
figs. 1-6. 


Upper Cretaceous (Pale beds, Belly River series) : Quarry No. 112, 
Steveville map area, near mouth of Sand Creek, Alberta, Canada. 


4 This interesting species, known from a nearly complete mandible, is listed in 
the above superorder tentatively. It is not absolutely certain that it is avian. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 7, 


Order GAVIIFORMES: Loons 
Family GAVIIDAE: Loons 
Subfamily GAVIINAE 
Genus GAVIA Forster 


Gavia J. R. Forster, Enchirid. Hist. Nat., 1788, p. 38. Type, by subsequent 
designation, Colymbus imber Gunnerus=C. immer Briinnich (Allen, 
1907). 


Gavia immer (BRUNNICH): Common Loon 
Colymbus Immer Brunnicu, Orn. Borealis, 1764, p. 38. 
Modern form reported from late Pleistocene (Palos Verdes sand) : 
Newport Bay, Orange County, California. 
Gavia arctica (LinNaAEuUS): Arctic Loon 
Colymbus arcticus LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 135. 
Modern form reported from late Pleistocene (Palos Verdes sand) : 
San Pedro, Los Angeles County, California. 
Gavia concinna WETMORE 
Gavia concinna WeETMoRE, Journ. Morph., vol. 66, No. 1, Jan. 2, 1940, p. 25, 


figs. I-4. 


Pliocene (Etchegoin formation): Sweetwater Canyon (type lo- 
cality), 545 miles east of King City, Monterey County, California. 
Middle Pliocene (San Diego formation): Washington Boulevard 
Freeway, San Diego, California. Pliocene (Bone Valley formation) : 
near Brewster, Polk County, Florida. 

Gavia palaeodytes WETMORE 
Gavia palaeodytes Wetmore, Proc. New England Zool. Club, vol. 22, June 23, 
1943, p. 64, figs. 1-2. 

Middle Pliocene (Bone Valley formation): Pierce (type locality) 

and Brewster, Polk County, Florida. 
Gavia howardae Bropkors 
Gavia howardae Bropxors, Condor, vol. 55, No. 4, July 20, 1953, p. 212, fig. 1B. 


Pliocene (Bone Valley formation): Pierce (type locality) and 
Brewster, Polk County, Florida. 
Subfamily GAVIELLINAE: GavieLia 
Genus GAVIELLA Wetmore 


Gaviella Wetmore, Journ. Morph. vol. 66, Jan. 2, 1940, p. 28. Type, by original 
designation, Gavia pusilla Shufeldt. 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Gaviella pusilla (SHUFELDT) 
Gavia pusilla SHuFELDT, Trans. Connecticut Acad. Arts Sci., vol. 19, Febru- 
ary IQI5, p. 70, pl. 13, fig. 106. 


Probably from Oligocene (White River formation): near Lusk, 
Wyoming.® 


Order COLYMBIFORMES: Greses 
Family COLYMBIDAE: GreseEs 
Genus COLYMBUS Linnaeus 


Colymbus LINNAEus, Syst. Nat., ed. 10, vol. 1, 1758, p. 135. Type, by subse- 
quent designation, Colymbus cristatus Linnaeus (Baird, Brewer, and 
Ridgway, 1884). 


Subgenus DYTES Kaup 


Dytes Kaur, Skizz. Entw.-Gesch. Eur. Thierw., 1829, p. 41. Type, by subse- 
quent designation, Dytes cornutus Kaup—=Colymbus auritus Linnaeus 
(Gray, 1842). 


Colymbus auritus LINNAEUS: Horned Grebe 


Colymbus auritus Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 135. 


Modern form reported from Pleistocene: Cavern deposits of Ten- 
nessee; Seminole Field, Pinellas County, and Itchtucknee River, 
Columbia County, Florida.® 


Colymbus caspicus Hapiiz_: Eared Grebe 


Colymbus caspicus Hasiiz_, Neue Nordische Beytrage, vol. 4, 1783, p. 9. 


Modern form reported from Pliocene (Ogallala formation) : Edson 
Quarry, Sherman County, Kansas. Late Pleistocene: Fossil Lake, 
Oregon; San Pedro (Palos Verdes sand, lumberyard locality), Los 
Angeles County, California; Meade County (Vanhem formation, 
Jones fauna), Kansas, 


Colymbus oligoceanus SHUFELDT 


Colymbus oligoceanus SuHuFELDT, Trans. Connecticut Acad. Arts Sci., vol. 19, 
February IQ15, p. 54. 


? Oligocene (John Day): Lower Willow Creek, Baker County, 
Oregon. 


5 See Wetmore, A., Journ. Morph., vol. 66, Jan. 2, 1940, p. 30. 

6 Specimens from Fossil Lake, Oregon, formerly included under this species 
have been found by Hildegarde Howard to represent Colymbus caspicus and 
Podilymbus podiceps. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 9 


Colymbus parvus SHUFELDT 
Colymbus parvus SHuFELDT, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 6, 
July 9, 1913, p. 136, pl. 39, fig. 477. 
Pliocene (Tulare formation): Kern County, California. Middle 


Pliocene (San Diego formation) : San Diego, California. Late Pleisto- 
cene: Fossil Lake (type locality), Oregon. 


Genus PLIODYTES Brodkorb 


Pliodytes BropKors, Ann. Mag. Nat. Hist., ser. 12, vol. 6, December 1953, 
p. 953, I fig. Type, by original designation, Pliodytes lanquisti Brodkorb. 


Pliodytes lanquisti BropKorB 


Pliodytes lanquisti BropKors, Ann. Mag. Nat. Hist., ser. 12, vol. 6, December 
1953, PD. 953, I fig. 


Pliocene (Bone Valley formation): Near Brewster, Polk County, 
Florida. 


Genus AECHMOPHORUS Coues 


ZEichmophorus Cours, Proc. Acad. Nat. Sci. Philadelphia, vol. 14, No. 5, April- 
May (Aug. 1), 1862, p. 229. Type, by original designation, Podiceps occi- 
dentalis Lawrence. 


Aechmophorus occidentalis (LAWRENCE): Western Grebe 


Podiceps occidentalis LAWRENCE, in Baird, Cassin, and Lawrence, Rep. Expl. 
and Surv. R. R. Pac., vol. 9, 1858, pp. liv, 892, 894. 


Modern form reported from late Pleistocene: Rodeo, San Francisco 
Bay region. 


Aechmophorus lucasi MILLER 


Aechmophorus lucasi L. H. Mitier, Univ. California Publ., Bull. Dept. Geol., 
vol. 6, No. 4, Feb. 4, 1911, p. 83, figs. 1-3. 


Late Pleistocene: Fossil Lake (type locality), Oregon;7 Palos 
Verdes sand, Newport Bay, Orange County, Playa del Rey, San 
Pedro, Los Angeles County, and near Manix, San Bernardino County, 
California. 


7Includes various specimens formerly listed under Colymbus grisegena and 
Aechmophorus occidentalis. Hildegarde Howard (Carnegie Inst. Washington 
Publ. 551, Jan. 25, 1946, pp. 148-151) considers lucasi the Pleistocene ancestor 
of modern A. occidentalis, listing it as Aechmophorus occidentalis lucasi, the 
relationship being expressed in the sense of distribution through time rather 
than in the geographic sense of subspecies existing simultaneously. 


Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Genus PODILYMBUS Lesson 
Podilymbus Lesson, Traité d’Orn., livr. 8, June 11, 1831, p. 505. Type, by 
monotypy, Podiceps carolinensis Latham = Colymbus podiceps Linnaeus, 
Podilymbus podiceps (LINNAEUS): Pied-billed Grebe 8 
Colymbus Podiceps LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 136. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Itchtucknee River, Columbia County, and Haile, Alachua 
County, Florida. Late Pleistocene: Fossil Lake, Oregon; Rancho 
La Brea, Los Angeles, and McKittrick, Kern County, California. 
Late Pleistocene or early Recent: Tepexpan, México. 


Order PROCELLARIIFORMES: AtsarrossEs, SHEARWATERS, 
PETRELS, and ALLIES 


Family DIOMEDEIDAE: AvpatrosseEs 


Genus DIOMEDEA Linnaeus 
Diomedia LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 132. Type, by subse- 
quent designation, Diomedea exulans Linnaeus (Gray, 1840). 
Diomedea albatrus PALLAS: Short-tailed Albatross 


Diomedea albatrus PAtvas, Spic. Zool., vol. 1, fasc. 5, 1769, p. 28. 


Modern form reported from late Pleistocene (Palos Verdes sand) : 
Newport Bay, Orange County, Playa del Rey, Los Angeles County, 
California. 


Diomedea anglica LyDEKKER 
Diomedea anglica LypEKKER, Cat. Foss. Birds Brit. Mus., 1891, p. 189, fig. 42. 
Pliocene (Bone Valley formation) : Pierce, Polk County, Florida.® 


Family PROCELLARIIDAE: SHEARWATERS and FuLMARS 
Genus PUFFINUS Brisson 1° 


Puffinus Brisson, Orn., 1760, vol. 1, p. 56; vol. 6, p. 130. Type, by tautonymy, 
Puffinus Brisson = Procellaria puffinus Briinnich. 


8 Podilymbus magnus Shufeldt, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 
6, July 9, 1913, p. 136, pl. 38, figs. 439-440, 449, has been identified as P. podiceps 
by Wetmore, California Acad. Sci., vol. 23, Dec. 30, 1937, pp. 198-199. 

9 Described by Lydekker from the Upper Pliocene at Foxhall, Suffolk, Eng- 
land. Recorded from Florida by Wetmore, Proc. New England Zodél. Club, 
vol. 22, June 23, 1943, pp. 66-67, pl. 12, figs. 10-15. 

10 Puffinus parvus Shufeldt, Ibis, October 1916, p. 632, from Recent deposits 
in the bone caves of Bermuda is considered a synonym of Puffinus lherminieri. 
Puffinus mcgalli Shufeldt, Ibis, October 1916, p. 630, from Recent deposits in the 
bone caves of Bermuda seemingly is Puffinus puffinus. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE II 


Subgenus PUFFINUS Brisson 


Puffinus griseus (GMELIN): Sooty Shearwater 
Procellaria grisea GMELIN, Syst. Nat., vol. 1, pt. 2, 1789, p. 564. 
Modern form reported from late Pleistocene (Palos Verdes sand) : 


Newport Bay, Orange County ; near San Pedro (lumberyard locality ) 
and Playa del Rey, Los Angeles County, California. 


Puffinus puffinus (BrUNNIcH): Common Shearwater 
Procellaria Puffinus BRUNNICH, Orn. Borealis, 1764, p. 29. 
Modern form reported from Pleistocene (Melbourne bone bed) : 


Melbourne, Florida. Late Pleistocene (Palos Verdes sand): San 
Pedro and Playa del Rey, Los Angeles County, California.™ 


Puffinus inceptor WETMORE 
Puffinus inceptor WETMoRE, Proc. California Acad. Sci., ser. 4, vol. 19, No. 8, 
July 15, 1930, p. 86, figs. 1-3. 


Middle Miocene (Temblor formation): Sharktooth Hill, about 7 
miles northeast of Bakersfield, California. 


Puffinus diatomicus MILLER 
Puffinus diatomicus L. H. Mitver, Carnegie Inst. Washington Publ. 349, 
August 1925, p. 111, pls. 1, 2, 7a. 


Middle Miocene (Temblor formation, Turritella ocoyana zone) : 
Lompoc (type locality). Miocene (Monterey shale): Lomita and 
San Pedro breakwater, San Pedro, California. 


Puffinus kanakoffi Howarp }2 
Puffinus kanakoffi Howarp, Carnegie Inst. Washington Publ. 584, June 22, 
1949, p. 187, pl. 2, figs. 3, 5. 


Middle Pliocene (San Diego formation) : Washington Boulevard 
Freeway, San Diego, California. 


Puffinus felthami Howarp !2 
Puffinus felthami Howarp, Carnegie Inst. Washington Publ. 584, June 22, 
1949, p. 104, pl. 2, figs. 4, 6. 
Late Lower Pliocene: 3 miles north of Corona del Mar, Orange 
County, California. 


11 The California records refer to Puffinus puffinus opisthomelas Coues, for- 
merly listed as a separate species. 
12 Subgeneric allocation provisional. 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Subgenus ARDENNA Reichenbach 


Ardenna REICHENBACH, Avium Syst. Nat., 1852 (1853), p. iv. Type, by 
monotypy, Procellaria major Faber = P. gravis O'Reilly. 


Puffinus conradi MArsu 


Puffinus conradi Marsu, Amer. Journ. Sci., ser. 2, vol. 49, 1870, p. 212. 


Middle Miocene (Calvert formation) : Maryland. 


Genus FULMARUS Stephens 


Fulmarus STEPHENS, in Shaw, Gen. Zodl., vol. 13, pt. 1, Feb. 18, 1826, p. 233. 
Type, by subsequent designation, Procellaria glacialis Linnaeus (Gray, 
1855). 


Fulmarus glacialis (LINNAEUS) : Fulmar 


Procellaria glacialis LINNAEUS, Fauna Suecica, ed. 2, 1761, p. 51. 


Modern form reported from late Pleistocene (Palos Verdes sand) : 
Newport Bay, Orange County; San Pedro, Los Angeles County, 
California. 


Family HYDROBATIDAE: Storm PETRELS 
Genus OCEANODROMA Reichenbach 


Oceanodroma REIcHENBACH, Avium Syst. Nat., 1852 (1853), p. iv. Type, by 
original designation, Procellaria furcata Gmelin. 


Oceanodroma hubbsi MILLER 


Oceanodroma hubbsi L. H. Mixer, Condor, vol. 53, No. 2, Mar. 27, 1951, p. 78, 
fig. I. 


Upper Miocene (Capistrano formation **): About 1 mile south of 
Capistrano Beach, Orange County, California. 


Order PELECANIFORMES: Tropicsirps, PELICANS, FRIGATEBIRDS, 
and ALLIES 


Suborder PELECANI: PEticans, Boostes, CorMorANTS, and DARTERS 
Superfamily PELECANOIDEA: PeticAns and ALLIES 
Family PELECANIDAE: PELicans 
Genus PELECANUS Linnaeus 


Pelecanus LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 132. Type, by subse- 
quent designation, Pelecanus onocrotalus Linnaeus (Gray, 1940). 


13 Possibly Lower Pliocene. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 13 


Subgenus CYRTOPELICANUS Reichenbach 


Cyrtopelicanus REICHENBACH, Avium Syst. Nat., 1852 (1853), p. vii. Type, 
by original designation, Pelecanus trachyrhynchus Latham = P. erythro- 
rhynchos Gmelin. 


Pelecanus erythrorhynchos GMELIN: White Pelican 


Pelecanus erythrorhynchos GMELIN, Syst. Nat., vol. 1, pt. 2, 1780, p. 571. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon ; 
Manix lake beds, near Manix, San Bernardino County, California. 
? Pleistocene: Rattlesnake Hill, Fallon, Nevada. 


Pelecanus halieus WETMORE 


Pelecanus halieus WrtMorE, Smithsonian Misc. Coll., vol. 87, No. 20, Dec. 
27, 1933, D- 3, figs. 1-2. 


Upper Pliocene (Hagerman lake beds): Near Hagerman, Idaho. 


Subgenus LEPTOPELICANUS Reichenbach 


Leptopelicanus REICHENBACH, Avium Syst. Nat., 1852 (1853), p. vii. Type, 
by original designation, Pelecanus fuscus Gmelin = P. occidentalis Lin- 
naeus. 


Pelecanus occidentalis LINNAEUS: Brown Pelican 


Pelecanus occidentalis LINNAEUS, Syst. Nat., ed. 12, vol. 1, 1766, p. 215. 


Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 


Family CYPHORNITHIDAE: CypHorniTHEs 
Genus CYPHORNIS Cope 


Cyphornis Corr, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 9, May 31, 
1894, p. 449. Type, by monotypy, Cyphornis magnus Cope. 


Cyphornis magnus Core 


Cyphornis magnus Corr, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 9, 
May 31, 1894, p. 451. 


Middle Oligocene: Carmanah Point, Vancouver Island, British 
Columbia. 
Genus PALAEOCHENGIDES Shufeldt 


Palaeochendides Suureot, Geol. Mag., n.s. 4, vol. 3, August 1916, p. 347. 
Type, by monotypy, Palaeochendides mioceanus Shufeldt. 


I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Palaeochendides mioceanus SHUFELDT 


Palaecochendides mioceanus SHUFELDT, Geol. Mag., n.s. 4, vol. 3, August 1916, 
Pp. 347, pl. 15. 


Miocene (Hawthorn formation): Near source of Stono River, 
South Carolina. 


Superfamily SULOIDEA: Boosres, Cormorants, DARTERS, and ALLIES 


Family SULIDAE: Boonies and GANNETS 
Genus SULA Brisson 


Sula Brisson, Orn., 1760, vol. 1, p. 60; vol. 6, p. 494. Type, by tautonymy, 
Sula Brisson = Pelecanus piscator Linnaeus. 


Subgenus SULA Brisson 
Sula stocktoni MILLER 
Sula stockton’ L. H. Mutter, Publ. Univ. California at Los Angeles Biol. 
Sci., vol. 1, No. 5, Mar. 12, 1935, p. 75, fig. 2. 


Middle Miocene (Monterey shale): Near Lomita, Los Angeles 
County, California. 


Sula willetti Mmm.er 
Sula willetti L. H. Miter, Carnegie Inst. Washington Publ. 349, August 1925, 
p 112 pls, 3, 8fig. 1: 


Middle Miocene (Temblor formation, Turritella ocoyana zone) : 
Lompoc, Santa Barbara County, California. 


Sula guano Bropkors 


Sula guano Bropxors, Florida Geol. Surv. Rep. Invest. No. 14, November 
1955, Pp. 9, figs. 2, 5, 8. 


Pliocene (Bone Valley formation): Near Brewster, Polk County, 
Florida. 
Sula phosphata Bropkorsp 


Sula phosphata Bropxors, Florida Geol. Surv. Rep. Invest. No. 14, November 
1955, DP. II, figs. 3, 6, 9. 


Pliocene (Bone Valley formation): Near Brewster, Polk County, 
Florida. 


Subgenus MICROSULA Wetmore 


Microsula Wetmorg, Proc. U. S. Nat. Mus., vol. 85, Jan. 14, 1938, p. 25. Type, 
by original designation, Su/a (Microsula) avita Wetmore. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 15 


Sula avita WETMORE 


Sula avita WetTMorE, Proc. U. S. Nat. Mus., vol. 85, Jan. 14, 1938, p. 22, 
figs. 2-3. 


Middle Miocene (Calvert formation) : western shore of Chesapeake 
Bay, near Plumpoint, Calvert’County, Maryland. 


Genus MIOSULA Miller 


Miosula L. H. Mitier, Carnegie Inst. Washington Publ. 349, August 1925, 
p. 114. Type, by monotypy, Miosula media Miller. 


Miosula media MILLER 
Miosula media L. H. Mruter, Carnegie Inst. Washington Publ. 349, August 
1925, p. 114, pl. 5. 


Middle Miocene (Temblor formation, Turritella ocoyana zone) : 
Lompoc, Santa Barbara County, California. 


Miosula recentior Howarp 


Miosula recentior Howarp, Carnegie Inst. Washington Publ. 584, June 22, 
1940, p. 190, pl. 2, figs. I-2a. 


Middle Pliocene (San Diego formation) : Curlew Street, opposite 
Ostego Drive, San Diego, California. 


Genus MORUS Vieillot 


Morus Viertot, Analyse, April 1816, p. 63. Type, by monotypy, Pelecanus 
bassanus Linnaeus. 


Morus loxostyla (Corr) 14 
Sula loxostyla Corr, Trans. Amer. Philos. Soc., n.s., vol. 14, December 1870, 
p. 236, fig. 53. 
Miocene: Calvert County (type locality), Maryland; New Jersey. 


Morus vagabundus WETMORE 


Moris vagabundus Wermore, Proc. California Acad. Sci., ser. 4, vol: 19, No. 8, 
July 15, 1930, p. 89, fig. 4. 


Middle Miocene (Temblor formation): Sharktooth Hill (type 


locality), about 7 miles northeast, and west branch of Granite Creek, 
Ii miles north of Bakersfield, California. 


14 Sula atlantica Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, Feb- 
ruary 1915, p. 62, pl. 15, fig. 123, from the Miocene of New Jersey, is considered 
a synonym of M. loxostyla; cf. Wetmore, Auk, 1926, p. 465. 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Morus lompocana (MILLER) 


Sula lompocana L. H. Mitter, Carnegie Inst. Washington Publ. 349, August 
1925, p. 114, pls. 4, 7b, 9. 


Middle Miocene (Temblor formation, Turritella ocoyana zone) : 
Lompoc, Santa Barbara County, California. 
Morus peninsularis Bropkors 


Morus peninsularis Brovxors, Florida Geol. Surv. Rep. Invest. No. 14, No- 
vember 1955, p. 8, figs. I, 4, 7. 


Pliocene (Bone Valley formation): Near Brewster, Polk County, 
Florida. 
Morus reyana Howarp 
Moris reyana Howarp, Condor, vol. 38, No. 5, Sept. 15, 1936, p. 213, fig. 37. 
Late Pleistocene (Palos Verdes sand): Newport Bay, Orange 


County; Playa del Rey (type locality), Los Angeles County, Cali- 
fornia. 


Family PHALACROCORACIDAE: Cormorants 
Genus GRACULAVUS Marsh 15 


Graculavus Marsu, Amer. Journ. Sci., ser. 3, vol. 3, 1872, p. 363. Type, by 
subsequent designation, Graculavus velox Marsh (Hay, 1902). 


Graculavus pumilus MarsH 
Graculavus pumilus MArsu, Amer. Journ. Sci., ser. 3, vol. 3, 1872, p. 364. 
Paleocene (Hornerstown marl): Hornerstown, New Jersey. 
Graculavus velox Marsu 
Graculavus velox Marsu, Amer. Journ. Sci., ser. 3, vol. 3, 1872, p. 363. 


Paleocene (Hornerstown marl): Hornerstown, New Jersey. 


Genus PHALACROCORAX Brisson 16 
Phalacrocorax Brisson, Orn., 1760, vol. 1, p. 60; vol. 6, p. 511. Type, by 
tautonymy, Phalacrocorax Brisson = Pelecanus carbo Linnaeus. 
Phalacrocorax wetmorei BropKors 
Phalacrocorax wetmorei BropKkors, Florida Geol. Surv. Rep. Invest. No. 14, 


November 1955, p. 12, figs. 10, IT. 


Pliocene (Bone Valley formation): Near Brewster, Polk County, 
Florida. 


15 Jimosavis Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, February 
IQI5, p. 19, proposed.as a new genus for Graculavus velox Marsh, is a synonym 
of Graculavus Marsh, as both names are based on the same species. 

16 No subgenera are recognized in recent studies of the cormorants. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 17 


Phalacrocorax auritus (Lesson): Double-crested Cormorant 

Carbo auritus Lesson, Traité d’Orn., livr. 8, June 11, 1831, p. 605. 

Modern form reported from Pliocene: Dry Creek, Malheur County, 
Oregon. Upper Pliocene (Hagerman lake beds): Near Hagerman, 
Idaho. Pleistocene: Melbourne (stratum 2), Sarasota, Bradenton, 
Seminole Field, Pinellas County, Itchtucknee River, and Vero, 
Florida. Late Pleistocene (Palos Verdes sand): Santa Monica 
and San Pedro, Los Angeles County, California. ? Pleistocene: Rattle- 
snake Hill, Fallon, Nevada. 

Phalacrocorax penicillatus (BrRANpT): Brandt’s Cormorant 
Carbo penicillatus BRANvT, Bull. Sci. Acad. Imp. Sti. St.-Pétersbourg, vol. 3, 
No. 4, Nov. 16, 1837, col. 55. 


Modern form reported from late Pleistocene (Palos Verdes sand) : 
Newport Bay, Orange County ; Santa Monica and San Pedro (lumber- 
yard locality), Los Angeles County, California. 

Phalacrocorax femoralis MILLER 
Phalacrocorax femoralis L. H. Miter, Condor, vol. 31, No. 4, July 15, 19209, 
p. 167, figs. 58-59. 

Upper Miocene (Modelo formation): Calabasas, Los Angeles 
County, California. 

Phalacrocorax idahensis (Mars) 

Graculus idahensis Marsu, Amer. Journ. Sci., ser. 2, vol. 49, 1870, p. 216. 

Pliocene: Castle Creek; Owyhee County (type locality), Idaho; 
Pliocene (Bone Valley formation): Near Brewster, Polk County, 
Florida. Upper Pliocene (Hagerman lake beds): Near Hagerman, 
Idaho. 

Phalacrocorax macropus (Core) 
Graculus macropus Corr, Bull. Geol. Geogr. Surv. Terr., vol. 4, No. 2, 1878, 
p. 386. 
Late Pleistocene: Fossil Lake, Oregon.’" 


Phalacrocorax marinavis SHUFELDT 
Phalacrocorax marinavis SHUFELDT, Trans. Connecticut Acad. Sci., vol. 109, 
February 1915, p. 56, pl. 14, figs. 114, 116-118, 122. 


? Oligocene (John Day) : Willow Creek, Oregon. 


17 Shufeldt, Auk, 1915, pp. 485-488, has identified material from the Miocene of 
Montana as this species, but examination of the specimen reveals that this is 
in error. 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Phalacrocorax mediterraneus SHUFELDT 


Phalacrocorax mediterraneus SHUFELDT, Trans. Connecticut Acad. Arts Sci., 
vol. 19, February 1915, p. 58, pl. 15, fig. 138. 


Lower Oligocene (Chadron formation): Gerry’s Ranch, Weld 
County, Colorado. 


Phalacrocorax rogersi Howarp 


Phalacrocorax rogersi Howarp, Condor, vol. 34, No. 3, May 16, 1932, p. 118, 
fig. 19. 


Early Pleistocene (Santa Barbara formation): Veronica Springs 
Stone Quarry, near Santa Barbara, California. 


Phalacrocorax kennelli Howarp 


Phalacrocorax kennelli Howarp, Carnegie Inst. Washington Publ. 584, June 
22, 1949, p. 188, pl. 3, figs. 7-8a. 


Middle Pliocene (San Diego formation) : Washington Boulevard 
Freeway, San Diego, California. 


Family ANHINGIDAE: SNAKEBIRDs 
Genus ANHINGA Brisson 


Anhinga Brisson, Orn., 1760, vol. 1, p. 60; vol. 6, p. 476. Type, by tautonymy 
and monotypy, Anhinga Brisson = Plotus anhinga Linnaeus. 


Anhinga anhinga (LinNAEUS): Anhinga 
Plotus Anhinga Linnaeus, Syst. Nat., ed. 12, vol. 1, 1766, p. 218. 


Modern form reported from Pleistocene (Melbourne bone bed) : 
Melbourne (stratum 2), Florida. 


Order CICONIIFORMES: Herons, Srorks, and ALLIES 
Suborder ARDEAE: Herons, Bitterns, and ALLIES 
Family ARDEIDAE: Herons and BitTerns 


Subfamily ARDEINAE: Herons and Ecrets 
Genus ARDEA Linnaeus 1§ 


Ardea LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 141. Type, by subsequent 
designation, Ardea cinerea Linnaeus (Gray, 1840). 


18 Ardea sellardsi Shufeldt, Journ. Geol., January-February (January) 1917, 
p. 19, described from Vero (stratum 3), Florida, proves to be based on the 
tibiotarsus of Meleagris gallopavo. See Wetmore, Smithsonian Misc. Coll., 
vol. 85, No. 2, Apr. 13, 1931, p. 32. 





NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 19 


Ardea herodias LINNAEUS: Great Blue Heron 

Ardea Herodias LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 143. 

Modern form reported from Pleistocene: Melbourne,’® Itchtucknee 
River, Bradenton and Seminole Field, Pinellas County, Florida. Late 
Pleistocene: Fossil Lake, Oregon; Rancho La Brea, Los Angeles, 
and McKittrick, Kern County, California. 

Ardea polkensis BropKorB 
Ardea polkensis Bropkors, Florida Geol. Surv. Rep. Invest. No. 14, November 
1955, p. 17, figs. 13, 14, 15. 

Pliocene (Bone Valley formation): Near Brewster, Poll County, 
Florida. 

Genus CASMERODIUS Gloger 


Casmerodius GiocEer, Hand- und Hilfsbuch Naturg., 1842 (1841), p. 412. 
Type, by subsequent designation, Ardea egretta Gmelin (Salvadori, 1882). 


Casmerodius albus (LINNAEUS): Common Egret 

Ardea alba LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 144. 

Modern form reported from Pleistocene: Melbourne, Seminole 
Field, Pinellas County, and Venice, Florida. Late Pleistocene: Rancho 


La Brea, Los Angeles, California; Bafios de Ciego Montero, Santa 
Clara Province, Cuba. 


Genus LEUCOPHOYX Sharpe 


Leucophoyx Suarre, Bull. Brit. Orn. Club, vol. 3, Apr. 30, 1894, p. xxxix. 
Type, by original designation and monotypy, Ardea candidissima Gmelin = 
Ardea thula Molina. 


Leucophoyx thula (MoLinA): Snowy Egret 
Ardea Thula Motina, Sagg. Stor. Nat. Chili, 1782, p. 235. 


Modern form reported from Pleistocene: Bradenton, Florida. 


Genus HYDRANASSA Baird 


Hydranassa Bairp, in Baird, Cassin, and Lawrence, Rep. Expl. Surv. R. R. 
Pac., vol. 9, 1858, p. 660. Type, by original designation, Ardea ludoviciana 
Wilson = Egretta ruficollis Gosse. 


Hydranassa tricolor (MULLER): Tricolored Heron 

Ardea tricolor P. L. S. Miter, Natursyst. Suppl., 1776, p. 111. 

Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 


19 The record from Vero (stratum 3) is now considered Recent. See Cooke, 
C. W., Florida Geol. Surv. Geol. Bull. 29, 1945, pp. 306-307. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Genus FLORIDA Baird 


Florida Baten, in Baird, Cassin, and Lawrence, Rep. Expl. and Surv. R. R. 
Pac., vol. 9, 1858, pp. xxi, xlv, 659, 671. Type, by monotypy, Ardea 
caerulea Linnaeus. 


Florida caerulea (LINNAEUS): Little Blue Heron 
Ardea caerulea LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 143. 
Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 
Genus BUTORIDES Blyth 


Butorides Brytu, Cat. Birds Mus. Asiatic Soc., 1849 (1852), p. 281. Type, 
by monotypy, Ardea javanica Horsfield. 


Butorides virescens (LINNAEUS): Green Heron 
Ardea virescens LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 144. 
Modern form reported from Pleistocene: Seminole Field, Pinellas 


County, Florida. Late Pleistocene: Rancho La Brea, Los Angeles, 
California. 


Genus NYCTICORAX Forster 


Nycticorax T. Forster, Syn. Cat. Brit. Birds, 1817, p. 59. Type, by tautonymy 
and monotypy, Nycticorax infaustus Forster = Ardea nycticorax Lin- 
naeus. 


Nycticorax nycticorax (LinNaEus): Black-crowned Night Heron 

Ardea Nycticorar Linnagus, Syst. Nat., ed. 10, vol. 1, 1758, p. 142. 

Modern form reported from Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leén, México; Bradenton, and Itchtucknee River, Flor- 


ida. Late Pleistocene: McKittrick, Kern County, and Rancho La 
Brea, Los Angeles, California. 


Genus NYCTANASSA Stejneger 


Nyctanassa STEJNEGER, Proc. U. S. Nat. Mus., vol. 10, Aug. 3, 1887, p. 295. 
Type, by original designation, Ardea violacea Linnaeus. 


Nyctanassa violacea (LINNAEUS): Yellow-crowned Night Heron 


Ardea violacea LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 143. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida.’ 





20 Larus vero Shufeldt, Journ. Geol., 1917, p. 18, from stratum 3 of Vero, 
Florida, is Nyctanassa violacea, according to Wetmore, Smithsonian Misc. Coll., 
vol. 85, No. 2, 1931, pp. 3, 11, and 16. Cooke, Florida Geol. Surv., Geol. Bull. 209, 
1945, pp. 306-307, considers this deposit to be of Recent age. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 21 


Genus EOCEORNIS Shufeldt 


Eoceornis SHUFELDT, Trans. Connecticut Acad. Arts Sci., vol. 19, February 
IQI5, p. 39. Type, by monotypy, Eoceornis ardetta Shufeldt. 


Eoceornis ardetta SHUFELDT 


Eoceornis ardetta SHuFELDT, Trans. Connecticut Acad. Arts Sci., vol. 10, 
February 1915, p. 39, pl. 13, fig. 102. 


Eocene (Bridger formation) : Henry’s Fork, Wyoming. 


Subfamily BOTAURINAE: Birterns 
Genus IXOBRYCHUS Billberg 


Ixobrychus BIL_BerGc, Syn. Faunae Scand., vol. 1, pt. 2, 1828, p. 166. Type, 
by subsequent designation, Ardea minuta Linnaeus (Stone, 1907). 


Ixobrychus exilis (GMELIN): Least Bittern. 
Ardea exilis GMELIN, Syst. Nat., vol. 1, pt. 2, 1780, p. 645. 


Modern form reported from late Pleistocene: Bafios de Ciego 
Montero, Santa Clara Province, Cuba. 


Genus BOTAURUS Stephens 


Botaurus STEPHENS, in Shaw, Gen. Zool., vol. 11, pt. 2, August 1819, p. 592. 
Type, by subsequent designation, Ardea stellaris Linnaeus (Gray, 1840). 


Botaurus lentiginosus (RACKETTr): American Bittern 
Ardea lentiginosa RACKETT, in Pulteney, Cat. Birds, Shells and... . Plants 


of Dorsetshire, ed. 2, May 1813, p. 14. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, and Sarasota, Florida. Late Pleistocene: Fossil Lake, Ore- 
gon; * Rancho La Brea, Los Angeles, California. 


Genus BOTAUROIDES Shufeldt 


Botauroides SHuFELDT, Trans. Connecticut Acad. Arts Sci., vol. 19, February 
IQI5, p. 33. Type, by monotypy, Botauroides parvus Shufeldt. 
Botauroides parvus SHUFELDT 
Botauroides parvus Suuretpr, Trans. Connecticut Acad. Arts Sci., vol. 10, 
February I915, p. 33. 


Eocene (? Bridger formation): “Spanish John Meadow,” Wy- 
oming. 


21 Ardea paloccidentalis Shufeldt described from Fossil Lake is based on a 
fragmentary tarsometatarsus of the American bittern. See Howard, Carnegie 
Inst. Washington Publ. 551, Jan. 25, 1946, pp. 156-157. 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Suborder CICONIAE: Srorks, Ipises, and SPooNBILLS 
Superfamily CICONIOIDEA: Srorxs and Woop IsIses 
Family CICONIIDAE: Storxs and Jasirus 
Subfamily CICONIINAE: Storxs 
Genus CICONIA Brisson 
Ciconia Brisson, Orn., 1760, vol. 1, p. 48; vol. 5, p. 361. Type, by tautonymy, 


Ciconia = Ardea ciconia Linnaeus. 
Ciconia maltha MILLER 
Ciconia maltha L. H. Mitter, Univ. California Publ., Bull. Dept. Geol., vol. 5, 
No. 30, Aug. 5, 1910, p. 440, figs. I-7. 

Upper Pliocene (Hagerman lake beds): Barbour Ranch, Snake 
River, Idaho. Pleistocene: American Falls, Idaho; Vero (stratum 2), 
Melbourne (stratum 2), Itchtucknee River, 64 miles south of Marine- 
land, Flagler County, Seminole Field, Pinellas County, and Venice, 
Florida. Late Pleistocene: Carpinteria, McKittrick, Rancho La Brea, 
Los Angeles (type locality), and near Manix, San Bernardino 
County, California; Bafios de Ciego Montero, Santa Clara Province, 
Cuba.”? 


Subfamily MYCTERIINAE: Woop IBsisEs 
Genus MYCTERIA Linnaeus 


Mycteria LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 140. Type, by monotypy, 
Mycteria americana Linnaeus, 


Mycteria wetmorei Howarp 7% 
Mycteria wetmorei Howarp, Condor, vol. 37, Sept. 15, 1935, Pp. 253, fig. 47. 


Late Pleistocene: Rancho La Brea, Los Angeles, California. 


Superfamily THRESKIORNITHOIDEA: IstseEs 
Family THRESKIORNITHIDAE: Isises and SPoonsILts 


Subfamily THRESKIORNITHINAE: Ibises 
Genus PLEGADIS Kaup 


Plegadis Kaur, Skizz. Entw.-Ges. Eur. Thierw., 1829, p. 82. Type, by mono- 
typy, Tantalus falcinellus Linnaeus. 


22 Records formerly listed as Jabiru mycteria (Lichtenstein) have all been 
assigned to the present species by Hildegarde Howard, in Carnegie Inst. Wash- 
ington Publ. 530, Jan. 19, 1042, p. 202. Jabiru weillsi Sellards, therefore, be- 
comes a synonym of Ciconia maltha. 

28 Replaces Mycteria americana as listed in Check-list of North American 
Birds, ed. 4, 1931, p. 416. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 23 


Plegadis chihi (ViemLor) : White-faced Ibis 


Numenius chihi Vie1ttor, Nouv. Dict. Hist. Nat., nouv. éd., vol. 8, March 
1817, p. 303. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 


Genus EUDOCIMUS Wagler 


Eudocimus WaGLtER, Isis von Oken, 1832, col. 1232. Type, by subsequent desig- 
nation, Scolopax rubra Linnaeus (Reichenow, 1877). 


Eudocimus albus (LINNAEUS): White Ibis 


Scolopax alba LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 145. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, and Haile, Alachua County, Florida. 


Subfamily PLATALEINAE: Spoonnitts 
Genus AJAIA Reichenbach 


Ajaia REICHENBACH, Avium Syst. Nat., 1852 (1853), p. xvi. Type, by original 
designation, Ajaia rosea Reichenbach = Platalea ajaja Linnaeus. 


Ajaia ajaja (LINNAEUS) : Roseate Spoonbill 


Platalea Ajaja LiINNAEusS, Syst. Nat., ed. 10, vol. 1, 1758, p. 140. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 


Suborder PHOENICOPTERI: FLamincos 
Family PHOENICOPTERIDAE: FLAminecos 
Genus PHOENICOPTERUS Linnaeus 


Phoenicopterus LINNAEUuS, Syst. Nat., ed. 10, vol. 1, 1758, p. 139. Type, by 
monotypy, Phoenicopterus ruber Linnaeus. 


Phoenicopterus copei SHUFELDT 


Phoenicopterus copei SHuFELDT, Amer. Nat., vol. 25, No. 297, September 1891, 
p. 820. 


Late Pleistocene: Fossil Lake, Oregon. 


Phoenicopterus minutus Howarp 


Phoenicopterus minutus Howarp, Geol. Surv. Prof. Pap. 264-J, June 1955, 
p. 202, pl. 50. 


Late Pleistocene; Manix lake beds, near Manix, San Bernardino 
County, California. 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Phoenicopterus stocki MILLER 


Phoenicopterus stocki L. H. Miter, Wilson Bull., vol. 56, No. 2, June 1944, 
Da 77 MES. sk 2: 


Pliocene (Rincon) : Chihuahua, México. 
Phoenicopterus floridanus BropkorB 


Phoenicopterus floridanus Bropkors, Chicago Acad. Sci. Nat. Hist. Misc., 
No. 124, June 9, 1953, p. I, figs. 1-2. 


Pliocene (Bone Valley formation) : Near Brewster, Polk County, 
Florida. 


Family PALOELODIDAE: PaLogeLopus and ALLIEs 
Genus MEGAPALOELODUS Miller 


Megapaloelodus A. H. Miter, Univ. California Publ., Bull. Dept. Geol. Sci., 


vol. 27, No. 4, June 22, 1944, p. 86. Type, by original designation, Megapa- 
loelodus connectens A, H. Miller. 


Megapaloelodus connectens MILLER 


Megapaloelodus connectens A. H. Mitier, Univ. California Publ., Bull. Dept. 
Geol. Sci., vol. 27, No. 4, June 22, 1944, p. 86, fig. 1. 


Lower Miocene (Rosebud formation): Flint Hill, 9 miles west- 
southwest of Martin, Bennett County, South Dakota (type locality). 
Upper Miocene (Barstow formation) : near Barstow, California. 


Order ANSERIFORMES: Screamers, Ducks, GEESE, and SWANS 
Suborder ANSERES: Ducks, GEESE, SWANs, and ALLIES 
Family PARANYROCIDAE: Paranyroca 
Genus PARANYROCA Miller and Compton 


Paranyroca A. H. Mitier and L. V. Compton, Condor, vol. 41, No. 4, July 15, 


1939, Pp. 153. Type, by original designation, Paranyroca magna Miller and 
Compton, 


Paranyroca magna MILLER and CoMPTON 


Paranyroca magna A. H. MILter and L. V. Compton, Condor, vol. 41, No. 4, 
July 15, 1939, p. 153, fig. 34 A, C, D, E. 


Lower Miocene (Rosebud formation): Flint Hill, 9 miles west- 
southwest of Martin, Bennett County, South Dakota. 


Family ANATIDAE: Ducks, GEEsE, and Swans 
Subfamily CYGNINAE: Swans 
Genus CYGNUS Bechstein 


Cygnus Becustertn, Orn. Taschenb. Deutschl., vol. 2, 1803, p. 404, footnote. 
Type, by tautonymy, Anas cygnus Linnaeus. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 25 


Subgenus STHENELIDES Stejneger 


Sthenelides STEJNEGER, Stand. Nat. Hist., vol. 4, 1885, p. 143. Type, by 
monotypy, Anas melancoripha Molina. 


Cygnus paloregonus (Cope) 74 


Cygnus paloregonus Corr, Bull. Geol. Geogr. Surv. Terr., vol. 4, No. 2, 1878, 
p. 388. 


Pleistocene: Froman’s Ferry, Idaho. Late Pleistocene: Fossil Lake, 
Oregon (type locality) .?° 


Genus OLOR Wagler 


Olor Wacter, Isis von Oken, 1832, col. 1234. Type, by subsequent designa- 
tion, Cygnus musicus Bechstein = Anas cygnus Linnaeus (Gray, 1840). 


Subgenus OLOR Wagler 
Olor columbianus (Orp): Whistling Swan 
Anas columbianus Orb, in Guthrie, Geogr., 2d Amer. ed., 1815, p. 319. 
Modern form reported from Pleistocene: Seminole Field, Pinellas 


County, Florida. Late Pleistocene: Rancho La Brea, Los Angeles, 
and McKittrick, Kern County, California. 


Subgenus CLANGOCYCNUS Oberholser 


Clangocycnus OBERHOLSER, Emu, vol. 8, pt. 1, July 1908, p. 3. Type, by mono- 
typy, Cygnus buccinator Richardson. 
Olor buccinator (RicHARDSON) : Trumpeter Swan 
Cygnus buccinator RicHARDSON, in Wilson and Bonaparte, Amer. Orn., Jame- 
son ed., vol. 4, August 1831, p. 345. 
Modern form reported from Pleistocene: Aurora, Illinois ; Itchtuck- 
nee River, Florida. Late Pleistocene: Fossil Lake, Oregon. 


Subfamily ANSERINAE: GEEsE 
Genus BRANTA Scopoli 


Branta Scoro.t, Annus I, Historico-Naturalis, 1769, p. 67. Type, by subse- 
quent designation, Anas bernicla Linnaeus (Bannister, 1870). 


24 Subgeneric allocation tentative. 

25 Specimens named Cygnus matthewi and Anser condoni by Shufeldt are now 
identified as C. paloregonus. See Howard, Carnegie Inst. Washington Publ. 551, 
Jan. 25, 1946, pp. 160, 162, 163. 


26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Branta canadensis (LINNAEUS): Canada Goose 

Anas canadensis LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 123. 

Modern form reported from Pleistocene: Santa Rosa Island, Cali- 
fornia; Seminole Field, Pinellas County, and Itchtucknee River, 
Florida. Early Pleistocene: Irvington, Alameda County, California. 
Late Pleistocene: Fossil Lake, Orgeon ; *® Potter Creek Cave, Shasta 
County; Rancho La Brea, Los Angeles, San Pedro, Los Angeles 
County, and near Manix, San Bernardino County, California. 
? Pleistocene: Rattlesnake Hill, Fallon, Nevada.*? 

Branta canadensis hutchinsii (Rrcuarpson) : Richardson’s Goose 


Anser Hutchinsii RicHarpson, in Swainson and Richardson, Fauna Bor.- 
Amer., vol. 2, 1831 (1832), p. 470. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Itchtucknee River, and Melbourne, Florida. 
Branta bernicla (LINNAEUS): Brant 


Anas bernicla LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 124. 
Modern form reported from Pleistocene: Fossil Lake, Oregon. 


Branta esmeralda Burr 


Branta esmeralda Burt, Univ. California Publ., Bull. Dept. Geol. Sci., vol. 18, 
No. 6, Mar. 19, 1920, p. 222, pl. 20. 


Upper Miocene (Esmeralda formation): Fish Lake Valley, Es- 
meralda County, Nevada. 
Branta howardae MILLER 
Branta howardae L. H. Miter, Condor, vol. 32, No. 4, July 15, 1930, p. 208, 
fig. 74. 
Lower Pliocene (Ricardo formation) : Mojave Desert area, Kern 
County, California. 


Branta dickeyi MILLER 


Branta dickeyi L. H. Mitrer, Condor, vol. 26, No. 5, Sept. 15, 1924, p. 179, 
fig. 46. 


Upper Pliocene: Dry Creek, Malheur County, Oregon. Late 
Pleistocene: McKittrick, California. 


26 Specimens from Fossil Lake range in size from modern B. c. minima to B. c. 
canadensis. 
27 Recorded as Branta canadensis canadensis. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 27 


Branta hypsibata (Cope) 78 
Anser hypsibatus Core, Bull. Geol. Geogr. Surv. Terr., vol. 4, No. 2, 1878, 
p. 387. 
Late Pleistocene: Fossil Lake, Oregon. 


Branta propinqua SHUFELDT 
Branta propinqua SHUFELDT, Journ. Acad. Nat. Sci. Philadelphia, 2d ser., 
vol. 9, sign. 53, Oct. 20, 1892, p. 407, pl. 15, fig. 17. 


Late Pleistocene: Fossil Lake, Oregon. 


Genus ANABERNICULA Ross 29 


Anabernicula Ross, Trans. San Diego Soc. Nat. Hist., vol. 8, No. 15, Aug. 24, 
1935, p. 107. Type, by monotypy, Anabernicula gracilenta Ross = Branta 
minuscula Wetmore.®° 


Anabernicula minuscula (WETMORE) 


Branta minuscula Wetmore, Proc. U. S. Nat. Mus., vol. 64, art. 5, Jan. 15, 
1924, p. 6, figs. 3-4. 

Upper Pliocene (Blancan) : Near Benson, Arizona (type locality). 
Late Pleistocene: Fossil Lake, Oregon; McKittrick, Kern County, 
and Rancho La Brea, Los Angeles, California. Quaternary: Smith 
Creek Cave, 34 miles north of Baker, White Pine County, Nevada. 


Genus PRESBYCHEN Wetmore 


Presbychen Wetmore, Proc. California Acad. Sci. ser. 4, vol. 19, No. 8, 
July 15, 1930, p. 92. Type, by original designation, Presbychen abavus 
Wetmore. 


Presbychen abavus WETMORE 
Presbychen abavus Wetmore, Proc. California Acad. Sci. ser. 4, vol. 19, 
No. 8, July 15, 1930, p. 92, figs. 5-7. 
Miocene (Temblor formation): Sharktooth Hill, Kern County, 
about 7 miles northeast of Bakersfield, California. 


Genus ANSER Brisson 


Anser Brisson, Orn., 1760, vol. 1, p. 58; vol. 6, p. 261. Type, by tautonymy, 
Anser domestica Brisson= Anas anser Linnaeus. 


28 Status doubtful. Howard, Carnegie Inst. Washington Publ. 551, Jan. 25, 
1946, pp. 167-169, indicates that this may be a synonym of Chen hyperborea. 

29 Possibly representative of a distinct subfamily. See Howard, Carnegie Inst. 
Washington Publ. 551, Jan. 25, 1946, pp. 172-173. 

80 See Howard, Condor, 1936, p. 35. 


28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Anser albifrons (ScopoL1): White-fronted Goose 
Branta albifrons Scopott, Annus I, Historico-Naturalis, 1769, p. 60. 
Modern form reported from late Pleistocene: Fossil Lake, Oregon ; 


Rancho La Brea, Los Angeles, and San Pedro ** (Palos Verdes sand), 
Los Angeles County, California. 


Genus CHEN Boie 


Chen Bote, Isis von Oken, vol. 10, Heft 5, 1822, col. 563. Type, by monotypy, 
Anser hyperboreus Pallas. 


Chen hyperborea (PALLAS): Snow Goose 
Anser hyperboreus Pauuas, Spic. Zool., vol. 1, fasc. 6, 1769, p. 25. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon ; 
Rancho La Brea, Los Angeles, and McKittrick, California.*? 


Chen rossii (CAss1n) : Ross’ Goose 
Anser Rossii “Baird,’ Cassin, Proc. Acad. Nat. Sci. Philadelphia, vol. 13, 
sign. 5-6, March-April (June 30), 1861, p. 73. 
Modern form reported from late Pleistocene: Fossil Lake, Oregon. 


Chen pressa WETMORE °% 
Chen pressa WETMORE, Smithsonian Misc. Coll., vol. 87, No. 20, Dec. 27, 1933, 
p. 9, figs. 5-8. 


Upper Pliocene (Hagerman lake beds): Near Hagerman, Idaho. 


Subfamily DENDROCYGNINAE: Treepucks 
Genus DENDROCYGNA Swainson 


Dendrocygna Swainson, Class. Birds, vol. 2, July 1, 1837, p. 365. Type, by 
subsequent designation, Anas arcuata Horsfield (Gray, 1840). 


Dendrocygna eversa WETMORE 


Dendrocygna eversa WrETMORE, Proc. U. S. Nat. Mus., vol. 64, art. 5, Jan. 
15, 1924, p. 3, figs. I-2. 


Upper Pliocene (Blancan) : Near Benson, Arizona. 


31 Specimen with size of the subspecies frontalis. 

32 Chen caerulescens recorded by Shufeldt, Bull. Amer. Mus. Nat. Hist., 
vol. 32, July 9, 1913, p. 145, on basis of scapula only, has been dropped. See 
Howard, Carnegie Inst. Washington Publ. 551, Jan. 25, 1946, p. 166? 

33 Miller, A. H., Univ. California Publ. Zool., vol. 42, No. 1, 1937, p. 41, sug- 
gests that this species may belong in the genus Nesochen. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 29 


Genus DENDROCHEN Miller 


Dendrochen A. H. Mitier, Univ. California Publ., Bull. Dept. Geol. Sci., 
vol. 27, No. 4, June 22, 1944, p. 88. Type, by original designation, 
Dendrochen robusta Miller. 


Dendrochen robusta MILLER 
Dendrochen robusta A. H. Mitier, Univ. California Publ., Bull. Dept. Geol. 
Sci., vol. 27, No. 4, June 22, 1944, p. 88, fig. 3. 


Lower Miocene (Rosebud formation): Flint Hill, 9 miles west- 
southwest of Martin, Bennett County, South Dakota. 


Subfamily ANATINAE: SurFac&-FEEDING Ducks 
Genus ANAS Linnaeus 


Anas LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 122. Type, by subsequent 
designation, Anas boschas Linnaeus = A. platyrhynchos Linnaeus (Les- 
son, 1828). 


Anas platyrhynchos LINNAEUS: Mallard 
Anas platyrhynchos LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 125. 
Modern form reported from late Pleistocene: Fossil Lake, Oregon ; 
Rancho La Brea, Los Angeles, McKittrick, and Carpinteria, Cali- 
fornia; (Palos Verdes sand): San Pedro, Los Angeles County, Cali- 
fornia ; Baiios de Ciego Montero, Santa Clara Province, Cuba. Pleis- 
tocene: Itchtucknee River, and Haile, Alachua County, Florida. 
Anas rubripes Brewster: Black Duck 
Anas obscura rubripes Brewster, Auk, vol. 19, No. 2, April 1902, p. 184. 
Modern form reported from Pleistocene: Itchtucknee River, 
Florida. 
Anas fulvigula Ringway: Mottled Duck 
Anas obscura var. fulvigula Ripaway, Amer. Nat., vol. 8, No. 2, February 


1874, p. III. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Itchtucknee River, and Bradenton, Florida. 


Anas strepera LINNAEUS: Gadwall 


Anas strepera LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 125. 


Modern form reported from late Pleistocene: McKittrick and 
Rancho La Brea, Los Angeles, California.** 


84 Listed erroneously in Check-list of North American Birds, ed. 4, 1931, p. 421, 
from Itchtucknee River, Florida. 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Anas acuta LiInNAEus: Pintail 


Anas acuta Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 126. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon. 
(Vanhem formation, Jones fauna): Meade County, Kansas. 


Anas carolinensis GMELIN; Green-winged Teal *® 


Anas carolinensis GMELIN, Syst. Nat., vol. 1, pt. 2, 1789, p. 533- 


Modern form reported from Pleistocene: Santa Rosa Island, Cali- 
fornia; Seminole Field, Pinellas County, Florida. Late Pleistocene: 
Fossil Lake, Oregon; Hawver Cave, Eldorado County, McKittrick, 
Kern County, Rancho La Brea, Los Angeles, and San Pedro, Los 
Angeles County, California; McPherson County, Kansas (Kentuck 
locality). 


Anas bunkeri (WETMORE) 


Nettion bunkeri WetTMorE, Univ. Kansas Sci. Bull., vol. 30, pt. 1, No. 9, 
May I5, 1044, p. 92, figs. 1-3. 


Upper Pliocene (Rexroad formation): Meade County, Kansas 
(type locality) ; 2 miles south of Benson, Arizona. 


Anas cyanoptera Vie1LLor: Cinnamon Teal 


Anas cyanoptera Vietot, Nouv. Dict. Hist. Nat., nouv. éd., vol. 5, December 
1816, p. 104. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon ; 
McKittrick, Kern County, California. 


Anas integra (MILLER). 


Querquedula integra A. H. Miter, Univ. California Publ., Bull. Dept. Geol. 
Sci., vol. 27, No. 4, June 22, 1944, p. 90, fig. 4. 


Lower Miocene (Rosebud formation), Flint Hill, 9 miles west- 
southwest of Martin, Bennett County, South Dakota. 


Genus MARECA Stephens 


Mareca STEPHENS, in Shaw, Gen. Zool., vol. 12, pt. 2, 1824, p. 130. Type, by 
subsequent designation, Mareca fistularis Stephens = Anas penelope Lin- 
naeus (Eyton, 1838). 


85 There are also records for the Upper Miocene or lower Pliocene of Cedar 
Mountain, Nevada, by L. H. Miller, Uniy. California Publ., Bull. Dept. Geol., 
vol. 9, Feb. 23, 1916, p. 173, and from the lower Pliocene of Hemphill County, 
Texas, by Compton, Condor, vol. 36, No. 1, January 1934, pp. 40-41, based on 
fragmentary material that is open to question as to specific identity. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 31 


Mareca americana (GMELIN): American Widgeon 
Anas americana GMELIN, Syst. Nat., vol. 1, pt. 2, 1789, p. 526. 
Modern form reported from late Pleistocene: Fossil Lake, Oregon ; 


McKittrick, Kern County; San Pedro (Palos Verdes sand, lumber- 
yard locality), Los Angeles County, California. 


Genus SPATULA Boie 


Spatula Bore, Isis von Oken, vol. 10, Heft 5, 1822, col. 564. Type, by 
monotypy, Anas clypeata Linnaeus. 


Spatula clypeata (Linnaeus) : Shoveler 

Anas clypeata LINNAEUvS, Syst. Nat., ed. 10, vol. 1, 1758, p. 124. 

Modern form reported from late Pleistocene: Fossil Lake, Ore- 
gon ; °° McKittrick, Kern County, and San Pedro (Palos Verdes sand, 


lumberyard locality), Los Angeles County, California ; Meade County 
(Vanhem formation, Jones fauna), Kansas. 


Subfamily AYTHYINAE: Divine Ducks * 
Genus AYTHYA Boie 


Aythya Bor, Tageb. Reise Norwegen, before May 1822, p. 351. Type, by 
monotypy, Anas marila Linnaeus. 


Aythya americana (Eyton): Redhead 


Fuligula americana Eyton, Mon. Anatidae, 1838, p. 155. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon; 
McKittrick, California. 


Aythya collaris (Donovan): Ring-necked Duck 
Anas collaris DonovaN, Brit. Birds, vol. 6, 1800, pl. 147. 


Modern form reported from Lower Pliocene: Cedar Mountain, 
Nevada. 


36 Shufeldt’s record of Aix sponsa from Fossil Lake is now assigned to 
Spatula clypeata. See Howard, Carnegie Inst. Washington Publ. 551, Jan. 25, 
1946, p. 176. 

87 Polysticta stelleri, Bucephala islandica, and [Histrionicus histrionicus re- 
ported from Fossil Lake by Shufeldt were wrongly identified and are eliminated 
from the list. See Howard, Carnegie Inst. Washington Publ. 551, Jan. 25, 1046, 
p. 176. 


— 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31I 


Aythya valisineria (Wirson): Canvasback 


Anas valisineria Witson, Amer. Orn., vol. 8, 1814, p. 103, pl. 70, fig. 5. 


Modern form reported from Pleistocene: Itchtucknee River, 
Florida.*® 
Aythya affinis (Eyton): Lesser Scaup 

Fuligula affinis Eyton, Mon. Anatidae, 1838, p. 157. 

Modern form reported from Pleistocene: Melbourne (stratum 2), 


Itchtucknee River, Seminole Field, Pinellas County, Venice, and cave 
deposits near Lecanto, Florida. Late Pleistocene : Fossil Lake, Oregon. 


Genus BUCEPHALA Baird 


Bucephala Baro, in Baird, Cassin, and Lawrence, Rep. Expl. Surv. R. R. Pac., 
vol. 9, 1858, pp. xx, L, 787, 788, 795. Type, by original designation, 
Anas albeola Linnaeus. 

Bucephala albeola (LINNAEUS) : Bufflehead 


Anas Albeola LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 124. 


Modern form reported from Upper Pliocene (Rexroad formation) : 
Meade County, Kansas. Pleistocene: Seminole Field, Pinellas County, 
Florida. Late Pleistocene: Fossil Lake, Oregon; McKittrick, Kern 
County, and San Pedro (Palos Verdes sand, lumberyard locality), 
Los Angeles County, California. 


Bucephala ossivallis BropKorB 


Bucephala ossivallis Bropkors, Florida Geol. Sury. Rep. Invest. No. 14 
November 1955, p. 18, figs. 16, 17. 


Pliocene (Bone Valley formation): Near Brewster, Polk County, 
Florida. 


Genus CLANGULA Leach 


Clangula Leacu, in Ross, Voy. Discovery, 1819, app., p. XLvm1. Type, by 
monotypy, Clangula glacialis Linnaeus = Anas hyemalis Linnaeus. 


Clangula hyemalis (LINNAEUS) : Oldsquaw 


Anas hyemalis LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 126. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon. 


Genus MELANITTA Boie 


Melanitta Bote, Isis von Oken, vol. 10, Heft 5, 1822, col. 564. Type, by subse- 
quent designation, Anas fusca Linnaeus (Eyton, 1838). 


88 Shufeldt’s record for Fossil Lake, Oregon, refers to Anas acuta. See 
Howard, Carnegie Inst. Washington Publ. 551, Jan. 25, 1946, p. 174. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 33 


Melanitta deglandi (BonApPARTE) : White-winged Scoter 

Oedemia deglandi Bonaparte, Rev. Crit. Orn. Europe, 1850, p. 108. 

Modern form reported from late Pleistocene (Palos Verdes sand) : 
San Pedro, Los Angeles County, California. 
Melanitta perspicillata (LINNAEUS) : Surf Scoter 

Anas perspicillata Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 125. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon ; 
San Pedro (Palos Verdes sand), Los Angeles County, California. 


Genus CHENDYTES Miller 


Chendytes L. H. Mitier, Condor, vol. 27, No. 4, July 15, 1925, p. 145. Type, 
by monotypy, Chendytes lawi Miller. 


Chendytes lawi MILLER 


Chendytes lawi L. H. Miter, Condor, vol. 27, No. 4, July 15, 1925, p. 145, 
fig. 40. 


Early Pleistocene: Sexton Canyon, near Lake Canyon, Ventura 
County. Late Pleistocene: Newport Bay, Orange County; Lomita, 
Playa del Rey, Santa Monica (type locality), San Pedro (lumberyard 
locality), Vermont and Sepulveda Boulevard, Bixby Slough near 
Hermosa Beach, and Palos Verdes, Los Angeles County, California. 
Chendytes milleri HowArp 


Chendytes milleri H. Howarn, Condor, vol. 57, No. 3, May 25, 1955, p. 137, 
fic. t a, d, e; f, 9, t, He’ 2-0,"c, tig. 3: 


Early Pleistocene: San Nicolas Island, California. 


Subfamily OXYURINAE: Ruppy and Maskep Ducxs 
Genus OXYURA Bonaparte 


Oxyura Bonaparte, Ann. Lyc. Nat. Hist. New York, vol. 2, 1828, p. 390. 
Type, by monotypy, Anas rubidus Wilson. 


Oxyura jamaicensis (GMELIN) : Ruddy Duck 
Anas jamaicensis GMELIN, Syst. Nat., vol. 1, pt. 2, 1780, p. 519. 


Modern form reported from Pleistocene: Venice, Florida. Late 
Pleistocene : Fossil Lake, Oregon ; McKittrick, Kern County, and near 
Manix, San Bernardino County, California. 


Subfamily EONESSINAE: Eonessa 
Genus EONESSA Wetmore 


Eonessa WeEtMorE, Journ. Pal., vol. 12, No. 3, May 1938, p. 280. Type, by 
original designation, Eonessa anaticula Wetmore. 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Eonessa anaticula WETMORE 
' Eonessa anaticula WetmorE, Journ. Pal., vol. 12, No. 3, May 1938, p. 280, 
figs. I-5. 


Eocene (Uinta C horizon) : Myton Pocket, Utah. 


Subfamily MERGINAE: MErGANSERS 
Genus LOPHODYTES Reichenbach 


Lophodytes REICHENBACH, Avium Syst. Nat., 1852 (1853), p. 1x. Type, by 
original designation, Mergus cucullatus Linnaeus. 


Lophodytes cucullatus (LINNAEUS): Hooded Merganser °9 
Mergus cucullatus LinNAEus, Syst. Nat., ed. 10, vol. 1, 1758, p. 120. 
Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Venice, and Itchtucknee River, Florida; Nye Sink, Beaver 
County, Oklahoma. Late Pleistocene: McPherson County (Kentuck 
locality), Kansas. 
Lophodytes floridana (SHUFELDT) 4° 
Querquedula floridana SHUFELDT, 9th Ann. Rep. Florida State Geol. Surv., 
1917, p. 36, pl. 1, fig. 4, pl. 2, fig. 25. 


Pleistocene: Vero (stratum 2, type locality), Melbourne, and 
Itchtucknee River, Florida. 


Genus MERGUS Linnaeus 


Mergus LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 129. Type, by subse- 
quent designation, Mergus castor Linnaeus = Mergus merganser Linnaeus 
(Gray, 1840). 


Mergus merganser LINNAEUS: Merganser 

Mergus Merganser LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 120. 

Modern form reported from Pleistocene: North Shore Channel, 
Chicago, Illinois.** Late Pleistocene: Fossil Lake, Oregon. 
Mergus serrator LINNAEUS: Red-breasted Merganser 


Mergus Serrator LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 120. 
Modern form reported from late Pleistocene: Fossil Lake, Oregon. 


39 Shufeldt’s record from Fossil Lake, Oregon, is based on an erroneous identi- 
fication. See Howard, Carnegie Inst. Washington Publ. 551, Jan. 25, 1946, p. 176. 

40 See Wetmore, Condor, vol. 57, No. 3, 1955, p. 180. 

41 Formerly recorded as Mergus serrator; see Wetmore, Wilson Bull., 1948, 


p. 240. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 35 


Order FALCONIFORMES: Vuttures, Hawks, and FALcons 
Suborder CATHARTAE: New Wortp VuLtures 
Superfamily NEOCATHARTOIDEA: NerocaTHARTES 
Family NEOCATHARTIDAE: NEocaTHaRTES 
Genus NEOCATHARTES Wetmore 


Neocathartes WrEtTMorrE, Auk, vol. 67, No. 2, April 1950, p. 235. Type, by 
original designation, Eocathartes grallator Wetmore. 


Neocathartes grallator (WETMORE) 


Eocathartes grallator WrETMorE, Ann. Carnegie Mus., vol. 30, May 24, 1944, 
p. 58, pls. 1-5, figs. 1-10. 


Upper Eocene (Upper Washakie beds) : Sand wash one-half mile 
north of Dobe Town Road crossing, Sweetwater County, Wyoming. 


Superfamily CATHARTOIDEA: New Wortp VuLtures 
Family CATHARTIDAE: New Wortp VuttureEs 
Genus CATHARTES Illiger 


Cathartes ILL1cER, Prodromus, 1811, p. 236. Type, by subsequent designation, 
Vultur aura Linnaeus (Vigors, 1825). 


Cathartes aura (LINNAEUS) : Turkey Vulture 42 


Vultur aura LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 86. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County,*® Melbourne, and cavern deposits near Lecanto, Florida. 
Late Pleistocene: Potter Creek and Samwel caves, Shasta County, 
Hawver Cave, Eldorado County, Carpinteria, Santa Barbara County, 
McKittrick, Kern County, Rancho La Brea, Los Angeles, and San 
Pedro (Palos Verdes sand, lumberyard locality), Los Angeles County, 
California. 


Genus CORAGYPS Geoffroy 


Coragyps GrorFrroy Ms in Le Maout, Hist. Nat. Oiseaux, 1853, p. 66. Type, 
by monotypy, Vultur urubu Vieillot = Vultur atratus Bechstein. 


42 Wetmore, Smithsonian Misc. Coll., vol. 85, No. 2, Apr. 13, 1931, pp. 4, 6, 
7, 23-24, has recorded the small Mexican turkey vulture, Cathartes aura aura, 
from Seminole Field, Pinellas County, Florida. Other reports of this species 
are mainly of the larger type, of which two races, septentrionalis and teter, are 
at present recognized in the United States. 

48 Recorded from Vero, stratum 2, erroneously by Shufeldt, 9th Ann. Rep. 
Florida State Geol. Surv., 1917, p. 36. The record from Vero (stratum 3) is of 
Recent age according to Cooke, Florida Geol. Surv. Bull. 29, 1945, pp. 306-307. 


36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Coragyps atratus (BecustTern): Black Vulture 


Vultur atratus BECHSTEIN, in John Latham’s allg. Uebers Vogel, Bd. 1, Anh., 
1793, PD. 655. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, and cavern deposits near Lecanto, Florida. Quaternary 
(probably Recent) : Rocky Arroyo, New Mexico. 


Coragyps occidentalis (MILLER) 44 


Catharista occidentalis L. H. Mitier, Univ. California Publ., Bull. Dept. Geol., 
vol. 5, No. 21, Sept. 10, 1909, p. 306. 


Pleistocene: San Josecito Cavern, Aramberri, Nuevo Leon.*® Late 
Pleistocene: Potter Creek and Samwel caves, Shasta County ; Carpin- 
teria, Santa Barbara County; McKittrick, Kern County; and Rancho 
La Brea, Los Angeles, California. 


Genus PHASMAGYPS Wetmore 


Phasmagyps WetTMorE, Proc. Colorado Mus. Nat. Hist., vol. 7, No. 2, July 
15, 1927, p. 3. Type, by monotypy, Phasmagyps patritus Wetmore. 


Phasmagyps patritus WETMORE 


Phasmagyps patritus WrtTMore, Proc. Colorado Mus. Nat. Hist., vol. 7, No. 2, 
July 15, 1927, p. 3, figs. 1-4. 


Lower Oligocene (Chadron formation): Horsetail Creek, Weld 
County, Colorado. 


Genus PALAEOGYPS Wetmore 


Palaeogyps WertMoreE, Proc. Colorado Mus. Nat. Hist., vol. 7, No. 2, July 
15, 1927, p. 5. Type, by monotypy, Palaeogyps prodromus Wetmore. 


Palaeogyps prodromus WETMORE 


Palaeogyps prodromus Wetmore, Proc. Colorado Mus. Nat. Hist., vol. 7, 
No. 2, July 15, 1927, p. 5, figs. 5-14. 


Lower Oligocene (Chadron formation): Horsetail Creek, Weld 
County, Colorado. 


44 Coragyps shastensis (Miller) is a synonym according to Miller, Condor, 
1941, pp. 140-141. 

45 Recorded also from deposits that may be late Pleistocene or early Recent 
in Pit 10 at Rancho La Brea (Howard, H., and Miller, A. H., Carnegie Inst. 
Washington Publ. 514, 1939, p. 43), Conkling Cavern, Pyramid Peak, Organ 
Mountains, Dona Ana County, New Mexico (Howard, H., and Miller, A. H., 
Condor, vol. 35, Jan. 15, 1933, pp. 15, 17), and from Smith Creek Cave, 34 miles 
north of Baker, White Pine County, Nevada (Howard, H., Condor, vol. 37, 
July 15, 1935, pp. 206-207). 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE SYA 


Genus GYMNOGYPS Lesson 


Gymnogyps Lesson, Echo du Monde Savant, ser. 2, vol. 6, Dec. 8, 1842, col. 
1037. Type, by monotypy, Vultur californianus Shaw. 


Gymnogyps amplus MIL.er 46 
Gymnogyps amplus L. H. Mrtter, Univ. California Publ., Bull. Dept. Geol., 
vol. 6, No. 16, Oct. 28, 1911, p. 390, fig. 2. 

Pleistocene: Sarasota and Seminole Field, Pinellas County, Florida ; 
San Josecito Cave, Aramberri, Nuevo Leon. Late Pleistocene: Sam- 
wel Cave (type locality) and Stone Man Cave, Shasta County; Car- 
pinteria, McKittrick, and Rancho La Brea, Los Angeles, California. 
Quaternary (probably Recent) : Rocky Arroyo, New Mexico, 


, Genus BREAGYPS Miller and Howard 


Breagyps L. H. Mitter and H. Howarp, Publ. Univ. California at Los 
Angeles, Biol. Sci., vol. 9, Feb. 18, 1938, p. 171. Type, by original designa- 
tion, Vultur clarki Miller = Sarcorhamphus clarki Miller. 

Breagyps clarki (MILLER) 

Sarcorhamphus clarki L. H. Mrtter, Univ. California Publ., Bull. Dept. 
Geol., vol. 6, No. 1, Nov. 28, 1910, p. 11, figs. 3a, 3b. 

Late Pleistocene: Rancho La Brea, Los Angeles, California. 

Quaternary (probably late Pleistocene) : Smith Creek Cave, 34 miles 
north of Baker, White Pine County, Nevada. 


Genus SARCORAMPHUS Duméril 


Sarcoramphus DuUMERIL, Zodl. Anal., 1806, p. 32. Type, by subsequent designa- 
tion, Vultur papa Linnaeus (Vigors, 1825). 
Sarcoramphus kernense (MILLER) 
Vultur kernensis L. H. MiLver, Condor, vol. 33, Mar. 18, 1931, p. 70, fig. 16, 


Pliocene: Pozo Creek, Kern River Divide, Kern County, about 9 
miles northeast of Bakersfield, California. 


Family TERATORNITHIDAE: TeErarornitHEs 
Genus TERATORNIS Miller 


Teratornis, L. H. Mitter, Univ. California Publ., Bull. Dept. Geol., vol. 5, 
No. 21, Sept. 10, 1909, p. 307. Type, by monotypy, Teratornis merriami 
Miller. 


46 Fisher, Pacific Science, vol. 1, No. 4, October 1947, p. 227, finds that all 
fossil material from western North America formerly placed under the living 
Gymnogyps californianus is properly assigned to the present bird, which is so 
slightly differentiated as to be considered the direct Pleistocene progenitor of 
the modern form. The remaining records, from Florida and Nuevo Leon, are 
placed under amplus on the basis of probability. 


38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Teratornis merriami MILLER 
Teratornis merriami L. H. Mixter, Univ. California Publ., Bull. Dept. Geol., 
vol. 5, No. 21, Sept. 10, 1909, p. 307, text figs. 1-9. 

Pleistocene: Seminole Field, Pinellas County, and Bradenton, 
Florida; San Josecito Cave, Aramberri, Nuevo Leon. Late Pleisto- 
cene: Rancho La Brea (type locality) ,*7 Los Angeles, McKittrick, 
Kern County, and Carpinteria, Santa Barbara County, California. 


Teratornis incredibilis HowArp 
Teratornis incredibilis Howarp, Bull. Southern California Acad. Sci., vol. 51, 
pt. 2, 1952, p. 51, pl. 10, figs. 1-2. 
Quaternary (probably late Pleistocene): Smith Creek Cave, 34 
miles north of Baker, White Pine County, Nevada. 


Genus CATHARTORNIS Miller +6 


Cathartornis L. H. Miter, Univ. California Publ., Bull. Dept. Geol., vol. 6, 
No. 1, Nov. 28, 1910, p. 14. Type, by monotypy, Cathartornis gracilis 
Miller. 

Cathartornis gracilis MILLER 

Cathartornis gracilis L. H. Mitier, Univ. California Publ., Bull. Dept. Geol., 

vol. 6, No. 1, Nov. 28, 1910, p. 14, figs. 4a, 4b. 


Late Pleistocene: Rancho La Brea, Los Angeles, California. 


Suborder FALCONES: Sercretary-pirps, HAwks, and FALCONS 
Superfamily FALCONOIDEA: Hawks, Fatcons, and ALLIES 


Family ACCIPITRIDAE: Hawks, Orp Worip VuLtures, and 
HARRIERS 


Subfamily AEGYPIINAE: O_p Wortp VULTURES 
Genus PALAEOBORUS Coues 


Palaeoborus Courts, Key North Amer. Birds, ed. 2, 1884, p. 822. Type, by 
original designation, Cathartes umbrosus Cope. 


Palaeoborus umbrosus (Core) *® 


Cathartes wmbrosus Corr, Proc. Acad. Nat. Sci. Philadelphia, vol. 26, Oct. 20, 
1874, p. I5I. 


Pliocene: North of Pojauque, New Mexico. 


47 Recorded also from early Recent deposits in Pit 10, at Rancho La Brea 
(Howard, H., and Miller, A. H., Carnegie Inst. Washington Publ. 514, 1939, 
p. 43). 

48 Ajlocated to Teratornithidae by Miller, L. H., and Howard, H., Publ. Univ. 
California at Los Angeles, Biol. Sci., vol. 9, Feb. 18, 1938, pp. 169-170, 173. 

49 Placed in Aegypiinae by Howard, Carnegie Inst. Washington Publ. 349, 
1932, PP. 45, 70-73, 75, 76. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 39 


Palaeoborus howardae WETMORE 
Palaeoborus howardae WeEtTMoRrE, Proc. U. S. Nat. Mus., vol. 84, No. 3, 1936, 
p. 73, fig. 13. 
Miocene: Dawes County, Nebraska. 


Palaeoborus rosatus MILLER and Compton 
Palaeoborus rosatus A. H. Miter and L. V. Compton, Condor, vol. 41, No. 4, 
July 15, 1939, p. 156, fig. 34B. 
Lower Miocene (Rosebud formation): Flint Hill, 9 miles west- 
southwest of Martin, Bennett County, South Dakota. 


Genus NEOGYPS Miller 


Neogyps L. H. Mrtter, Univ. California Publ., Bull. Dept. Geol., vol. 9, No. 9, 
Mar. I0, 1916, p. 108. Type, by monotypy, Neogyps errans Miller. 
Neogyps errans MILLER 
Neogyps errans L. H. Mitter, Univ. California Publ., Bull. Dept. Geol., vol. 90, 
No. 9, Mar. I0, 1916, p. 108, fig. 2. 

Late Pleistocene: Rancho La Brea (type locality) ,°° Los Angeles, 
Carpinteria, Santa Barbara County, and McKittrick, Kern County, 
California; San Josecito Cave, Aramberri, Nuevo Leon. Quaternary: 
Smith Creek Cave, 34 miles north of Baker, White Pine County, 
Nevada. 


Genus NEOPHRONTOPS Miller 


Neophrontops L. H. Mitier, Univ. California Publ., Bull. Dept. Geol., vol. 9, 
No. 9, Mar. 10, 1916, p. 106. Type, by monotypy, Neophrontops ameri- 
canus Miller. 

Neophrontops americanus MILLER 

Neophrontops americanus L. H. Mitver, Univ. California Publ., Bull. Dept. 
Geol., vol. 9, No. 9, Mar. 10, 1916, p. 106, fig. 1. 

Late Pleistocene: Rancho La Brea (type locality) ,° Los Angeles, 

Carpinteria, and McKittrick, California; San Josecito Cave, Aram- 
berri, Nuevo Leon. 


Neophrontops dakotensis Compton 
Neophrontops dakotensis Compton, Amer. Journ. Sci., ser. 5, vol. 30, October 
1935, PD. 344, fig. I. 
Lower Pliocene: Big Spring Canyon, 15 miles southwest of Martin, 
Bennett County, South Dakota. 


50 Recorded also from early Recent deposits in Pit 10 at this site (Howard, H., 
and Miller, A. H., Carnegie Inst. Washington Publ. 514, 1939, p. 43). 

51 Recorded also from early Recent deposits in Pit 10 at this site (Howard, H., 
and Miller, A. H., Carnegie Inst. Washington Publ. 514, 1930, p. 43). 


40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Neophrontops vetustus WETMORE 


Neophrontops vetustus WETMORE, Condor, vol. 45, No. 6, Dec. 8, 1943, p. 229, 
fig. 62. 


Middle Miocene (Sheep Creek beds) : Stonehouse Draw Quarry, 
Sioux County, Nebraska. 


Subfamily ELANINAE: Wuite-taiLep KITeEs 
Genus ELANUS Savigny 


Elanus Savicny, Descr. Egypte, vol. 1, 1809, pp. 60, 97. Type, by monotypy, 
Elanus caesius Savigny = Falco caeruleus Desfontaines. 


Elanus leucurus (VIermLLor) : White-tailed Kite 


Milvus leucurus Vie1.tot, Nouv. Dict. Hist. Nat., nouv. éd., vol. 20, May 1818, 
p. 563 [errore = 556]. 


Modern form reported from Pleistocene: San Josecito Cave, 
Aramberri, Nuevo Leon. Late Pleistocene: Rancho La Brea, Los 
Angeles, California. 


Subfamily MILVINAE: True KirTes 
Genus PROICTINIA Shufeldt 


Proictinia SHUFELDT, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 16, Aug. 4, 
1913, p. 301. Type, by monotypy, Proictinia gilmorei Shufeldt. 


Proictinia effera WETMORE 


Proictinia effera WerrTMore, Bull. Amer. Mus. Nat. Hist., vol. 48, art. 12, 
Dec. 3, 1923, p. 504, figs. 19-20. 


Lower Miocene (Lower Harrison beds): Agate Fossil Quarry, 
Sioux County, Nebraska. 


Proictinia gilmorei SHUFELDT 


Proictinia gilmorei SHUFELDT, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 16, 
Aug. 4, 1913, p. 301, pl. 55, fig. 27. 


Lower Pliocene (Ogallala formation): Long Island, Phillips 
County, Kansas. 


Subfamily ACCIPITRINAE: Birp Hawxs 
Genus ACCIPITER Brisson 


Accipiter Brisson, Orn., 1760, vol. 1, p. 28; vol. 6, p. 310. Type, by tautonymy, 
Accipiter Brisson = Falco nisus Linnaeus. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 4I 


Accipiter gentilis (LINNAEUS): Goshawk 

Falco gentilis LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 80. 

Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, and Rancho La Brea, Los Angeles, California. 
Accipiter striatus velox (Witson): Sharp-shinned Hawk 

Falco velox Witson, Amer. Orn., vol. 5, 1812, p. 116, pl. 45, fig. 1. 

Modern form reported from late Pleistocene: Samwel Cave, Shasta 
County, Carpinteria, Santa Barbara County, and Rancho La Brea, 
Los Angeles, California. 

Accipiter cooperii (BoNAPARTE): Cooper’s Hawk 
Falco Cooperii BoNaPARTE, Amer. Orn., vol. 2, 1828, p. 1, pl. 10, fig. 1. 
Modern form reported from late Pleistocene: McKittrick, Kern 


County, Carpinteria, Santa Barbara County, and Rancho La Brea, 
Los Angeles, California. 


Subfamily BUTEONINAE: Buzzarps and EacLrs 
Genus BUTEO Lacépéde 


Buteo LackéPEpE, Tabl. Ois., 1790, p. 4. Type, by tautonymy, Falco buteo Lin- 
naeus. 


Buteo jamaicensis (GMELIN): Red-tailed Hawk 

Falco jamaicensis GMELIN, Syst. Nat., vol. 1, pt. 1, 1788, p. 266. 

Modern form reported from late Pleistocene: Potter Creek Cave, 
Shasta County, McKittrick, Carpinteria, and Rancho La Brea, Los 
Angeles, California. Pleistocene: Seminole Field, Pinellas County, 
Venice, and Melbourne (stratum 2), Florida. 

Buteo lineatus (GMELIN): Red-shouldered Hawk 

Falco lineatus GMELIN, Syst. Nat., vol. 1, pt. 1, 1788, p. 268. 

Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Venice, and Melbourne, Florida. Late Pleistocene: Carpin- 
teria, Santa Barbara County, California. 

Buteo platypterus (ViemLLor): Broad-winged Hawk 
Sparvins platypterus VietLiot, Tabl. Encycl. Méth. Orn., vol. 3, 1823, p. 1273. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 


42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Buteo swainsoni BoNAPARTE: Swainson’s Hawk 

Butco swainsoni BONAPARTE, Geogr. and Comp. List, 1838, p. 3. 

Modern form reported from late Pleistocene: McKittrick, Kern 
County, and Rancho La Brea, Los Angeles, California. 
Buteo lagopus (PontoprpipANn) : Rough-legged Hawk 

Falco lagopus PoNToppiwan, Danske Atlas, 1763, p. 616. 

Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 
Buteo regalis (Gray): Ferruginous Hawk 

Archibuteo regalis G. R. Gray, Genera of Birds, vol. 1, pt. 1, May 1844, pl. 6. 

Modern form reported from late Pleistocene: Hawver Cave, EI- 
dorado County, Rancho La Brea, Los Angeles, Carpinteria, Santa 
Barbara County, and McKittrick, Kern County, California. 
Buteo fuscescens (VIEILLOT) : Buzzard Eagle 


Spizaétus fuscescens VietLtot, Nouv. Dict. Hist. Nat., nouv. éd., vol. 32, Sep- 
tember 1810, p. 55. 


52 


Modern form *? reported from late Pleistocene: Bafios de Ciego 
Montero, Santa Clara Province, Cuba. 
Buteo antecursor WETMORE 

Buteo antecursor Wetmore, Bull. Mus. Comp. Zodl., vol. 75, October 1933, 

p. 208, figs. 1-5. 

Oligocene (Brule formation): Near Torrington, Goshen County, 
Wyoming. 
Buteo grangeri WETMORE and CASE 


Buteo grangeri WrETMorE and Case, Contr. Mus. Pal. Univ. Michigan, vol. 4, 
No. 8, Jan. 15, 1934, p. 129, I pl. 


Middle Oligocene (Brule formation, Oreodon beds) : Big Badlands 
of Pass Creek, Washabaugh County, South Dakota. 


Buteo fluviaticus Mrier and Sibley 
Buteo fluviaticus A. H. MiLcer and C. G. Srstey, Condor, vol. 44, No. 1, 
Jan. 15, 1942, p. 39, fig. 12. 


Middle Oligocene (Brule formation, Oreodon beds) : Owl Creek, 
6 miles east of Carr, Weld County, Colorado. 


52 Formerly called Buteo melanoleucus (Vieillot). The modern range extends 
from the mountains ‘of Venezuela and Colombia, south through Ecuador and 
Perti to Chile, and from southeastern Brazil and Paraguay to Tierra del Fuego. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 43 


Buteo typhoius WETMORE 


Buteo typhoius WETMorE, Bull. Amer. Mus. Nat. Hist., vol. 48, art. 12, Dec. 3, 
1923, p. 489, figs. 3-5. 


Lower Miocene (Lower Harrison beds) ; Upper Miocene ** (Lower 
Snake Creek beds, type locality): south of Agate, Sioux County, 
Nebraska. 

Buteo ales (WETMORE) 


Geranoaétus ales WretTMorE, Ann. Carnegie Mus., vol. 16, No. 4, Apr. 10, 1926, 
p. 403, pl. 38, figs. 1-5. 


Lower Miocene (Lower Harrison beds): Quarry No. 2, Agate 
Springs Fossil Quarries, Sioux County, Nebraska. 
Buteo contortus (WETMORE) 


Geranoaétus contortus WrETMoRE, Bull. Amer. Mus. Nat. Hist., vol. 48, art. 
12, Dec. 3, 1923, p. 492, figs. 6-9. 


Upper Miocene ** (Lower Snake Creek beds) : Sinclair Draw (type 
locality) and Olcott Hill, Sioux County, Nebraska. 
Buteo dananus (MarsH) 
Aquila danana Marsu, Amer. Journ. Sci., ser. 3, vol. 2, August 1871, p. 125. 
Lower Pliocene (Upper Snake Creek beds): Loup Fork River, 
Nebraska. 
Buteo conterminus (WETMORE) 


Geranoaétus conterminus WrTMoRE, Bull. Amer. Mus. Nat. Hist., vol. 48, 
art. 12, Dec. 3, 1923, p. 497, figs. 11-13. 


Lower Pliocene (Upper Snake Creek beds): 20 miles south of 
Agate, Sioux County, Nebraska. 


Genus PARABUTEO Ridgway 


Parabuteo Ripcway, in Baird, Brewer, and Ridgway, Hist. North Amer. 
Birds, vol. 3, 1874, p. 250. Type, by monotypy, Buteo harrisi Audubon. 


Parabuteo unicinctus (TEMMINCK) : Harris’ Hawk 
Falco unicinctus TEMMINCK, Planch. Col. Ois., livr. 53, Dec. 25, 1824, pl. 313. 
Modern form reported from Pleistocene: San Josecito Cave, 


Aramberri, Nuevo Leon. 


53 Possibly early Pliocene; cf. Cook, H. J., and Cook, M. C., Nebraska Geol. 
Surv., Paper No. 5, 1933, p. 42. 


44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Genus CALOHIERAX Wetmore 


Calohierax WetMoreE, Bull. Mus. Comp. Zodl., vol. 80, No. 12, October 1937, 
p. 428. Type, by original designation, Calohierax quadratus Wetmore. 


Calohierax quadratus WETMORE 


Calohierax quadratus Werrmore, Bull. Mus. Comp. Zodl., vol. 80, No. 12, 
October 1937, p. 420, figs. 1-3. 


Recent (extinct) :°* Cave deposits on Great Exuma Island, Ba- 
hama Islands. 


Genus MIOHIERAX Howard 


Miohierax Howarp, Condor, vol. 46, No. 5, Sept. 27, 1944, p. 236. Type, by 
original designation, Miohierax stocki Howard. 


Miohierax stocki Howarp 
Miohierax stocki Howarp, Condor, vol. 46, No. 5, Sept. 27, 1944, p. 236, 
fig. 40. 


Late Lower Miocene (Tick Canyon formation): Near head of 
Vasquez Canyon, Los Angeles County, California. 


Genus HYPOMORPHNUS Cabanis *5 


Hypomorphnus Capants, Arch. Naturg., vol. 10, Bd. 1, 1844, p. 263. Type, by 
original designation, Falco urubitinga Linnaeus. 


Hypomorphnus enectus (WETMORE) 
Urubitinga enecta Wetmore, Bull. Amer. Mus. Nat. Hist., vol. 48, art. 12, 
Dec. 3, 1923, p. 500, figs. 14-18. 


Middle Miocene (Lower Sheep Creek beds): 20 miles south of 
Agate, Sioux County, Nebraska. 


Hypomorphnus sodalis (SHUFELDT)*@ 


Aquila sodalis SHuretpt, Amer. Nat., vol. 25, No. 297, September 18091, p. 821. 


Late Pleistocene: Fossil Lake, Oregon. 


Genus TITANOHIERAX Wetmore 


Titanohierax WetMorE, Bull. Mus. Comp. Zodl., vol. 80, No. 12, October 
1937, p. 430. Type, by original designation, Titanohierax gloveralleni 
Wetmore. 


54 Included here since it has not been found in living form, being known only 
from its bones. 

55 For the use of Hypomorphnus to replace Urubitinga see Peters, Check-list 
of the birds of the world, vol. 1, 1931, p. 244. 

56 Generic allocation questionable. See Howard, Carnegie Inst. Washington 
Publ. 551, Jan. 25, 1946, pp. 177-178. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 45 


Titanohierax gloveralleni WETMORE 
Titanohierax gloveralleni WEtMorE, Bull. Mus. Comp. Zodl., vol. 80, No. 12, 
October 1937, p. 431, figs. 4-9. 
Recent (extinct) : °7 Cave deposits on Great Exuma Island, Bahama 
Islands. 


Genus BUTEOGALLUS Lesson 
Buteogallus Lesson, Traité d’Orn., livr. 2, 1830, p. 83. Type, by monotypy, 


Buteogallus cathartoides Lesson= Falco aequinoctialis Gmelin. 
Buteogallus milleri (Howarp) 58 
Urubitinga milleri Howarp, Carnegie Inst. Washington Publ. 429, October 
1932, p. 25, pl. 2, figs. 3-3a, pl. 3, fig. 2. 
Late Pleistocene: Hawver Cave, Eldorado County, California. 


Buteogallus fragilis (MILLER) 58 
Geranoaétus fragilis L. H. Miter, Univ. California Publ., Bull. Dept. Geol., 
vol. 6, No. 12, Oct. 9, 1911, p. 315, figs. 5a, 5b. 
Late Pleistocene: McKittrick, Kern County, Rancho La Brea (type 
locality) ,°° Los Angeles, and Carpinteria, Santa Barbara County, Cali- 
fornia. 


Genus WETMOREGYPS Miller 


Wetmoregyps L. H. Mitter, Condor, vol. 30, No. 4, July 16, 1928, p. 255. 
Type, by original designation, Morphnus daggetti Miller. 


Wetmoregyps daggetti (MiLiEr) 


Morphnus daggetti L. H. Mrtrer, Condor, vol. 17, No. 5, Oct. 10, 1915, p. 179, 
fig. 63. 


Pleistocene: San Josecito Cave, Aramberri, Nuevo Leon. Late 
Pleistocene: Rancho La Brea (type locality), Los Angeles, and Car- 
pinteria, Santa Barbara County, California. 


Genus MORPHNUS Dumont 


Morphnus Dumont, Dict. Sci. Nat., vol. 1, Suppl., October 1816, p. 88. Type, 
by subsequent designation, Falco guianensis Daudin (Chubb, 1916). 


57 Included here since it has not been found in living form, being known only 
from its bones. 

58 Referred to this genus by Howard, Carnegie Inst. Washington Publ. 551, 
Jan. 25, 1946, p. 177. 

59 Recorded also from early Recent deposits in Pit 10 at Rancho La Brea 
(Howard, H., and Miller, A. H., Carnegie Inst. Washington Publ. 514, 10309, 
p. 43). And from late Pleistocene or early Recent deposits in Shelter Cave, 
Pyramid Peak, Organ Mountains, Dona Ana County, New Mexico, by How- 
ard, H., and Miller, A. H., Condor, vol. 35, 1933, pp. 16, 17. 


40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Morphnus woodwardi MILLER 


Morphnus woodwardi L. H. Miter, Univ. California Publ. Bull. Dept. Geol., 
vol. 6, No. 12, Oct. 9, 1911, p. 312, figs. 3a, 3b. 


Late Pleistocene: Rancho La Brea, Los Angeles, California.© 


Genus SPIZAETUS Vieillot 


Spizaétus Vieittor, Analyse, 1816, p. 24. Type, by subsequent designation, 
Falco ornatus Daudin (Gray, 1840). 


Spizaétus grinnelli (Mrter) ® 


Geranoaétus grinnelli L. H. Mitter, Univ. California Publ., Bull. Dept. Geol., 
vol. 6, No. 12, Oct. 9, 1911, p. 314, figs. 4a, 4b. 


Pleistocene: San Josecito Cave, Aramberri, Nuevo Leon. Late 
Pleistocene: Rancho La Brea (type locality),®* Los Angeles, Mc- 
Kittrick and Carpinteria, California. 

Spizaétus willetti Howarp 


Spizaétus willetti Howarp, Condor, vol. 37, No. 4, July 15, 1935, p. 207, fig. 40. 


Quaternary (probably late Pleistocene): Smith Creek Cave, 34 
miles north of Baker, White Pine County, Nevada. 
Spizaétus pliogryps (SHUFELDT) 


Aquila pliogryps SHureLpt, Amer. Nat., vol. 25, No. 297, September 1801, 
p. 821. 


Late Pleistocene: Fossil Lake, Oregon. 


Genus PALAEASTUR Wetmore 


Palacastur Wetmore, Condor, vol. 45, No. 6, Dec. 8, 1943, p. 230. Type, by 
original designation, Palaeastur atavus Wetmore. 


Palaeastur atavus WETMORE 
Palaeastur atavus Wetmore, Condor, vol. 45, No. 6, Dec. 8, 1943, p. 230, fig. 63. 


Lower Miocene (Lower Harrison beds) ; Stenomylus Quarry, about 
2 miles southeast of Agate Springs fossil site, near Agate, Nebraska. 


60 Recorded also from early Recent deposits in Pit 10 at this site (Howard, H., 
and Miller, A. H., Carnegie Inst. Washington Publ. 514, 1939, p. 43). 

61 Allocated in Spizaétus by Howard, Carnegie Inst. Washington Publ. 420, 
1932, PP- 33-44- 

62 Placed in Spisaétus by Howard, Carnegie Inst. Washington Publ. 551, 
Jan. 25, 1946, pp. 176-177. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 47 


Genus AQUILA Brisson ® 


Aquila Brisson, Orn., 1760, vol. 1, pp. 28, 419. Type, by tautonymy, Aquila 
Brisson = Falco chrysaétus Linnaeus. 


Aquila chrysaétos (LINNAEUS): Golden Eagle 
Falco Chrysaétos LINNAEUuS, Syst. Nat., ed. 10, vol. 1, 1758, p. 88. 


Modern form reported from Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. Late Pleistocene: Fossil Lake, Oregon; Rancho 
La Brea,** Los Angeles, Carpinteria, McKittrick, and near Manix, 
San Bernardino County, California. 


Genus HALIAEETUS Savigny 


Haliaeetus SavicNy, Descr. Egypte, Ois., vol. 1, 1809, pp. 68, 85. Type, by 
monotypy, Haliaeetus nisus Savigny = Falco albicilla Linnaeus. 


Haliaeetus leucocephalus (LinNAEuS): Bald Eagle 


Falco leucocephalus LiNNAEuS, Syst. Nat., ed. 12, vol. 1, 1766, p. 124. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Venice, Melbourne, and cavern deposits near Lecanto, Flor- 
ida; Niobrara River, near Peters, Sheridan County, Nebraska. Late 
Pleistocene: Fossil Lake, Oregon; Carpinteria, McKittrick, Rancho 
La Brea, Los Angeles, and San Pedro (Palos Verdes sand), Los 
Angeles County, California. 


Subfamily PALAEOPLANCINAE: PALaropLaNncus 
Genus PALAEOPLANCUS Wetmore 


Palacoplancus WertMorE, Smithsonian Misc. Coll., vol. 87, No. 19, Dec. 26, 
1933, p. I. Type, by original designation, Palaeoplancus sternbergi 
Wetmore. 


Palaeoplancus sternbergi WrETMORE 


Palaeoplancus sternbergi WrtMoreE, Smithsonian Misc. Coll., vol. 87, No. 10, 
Dec. 26, 1933, p- 12, figs. I-19. 


Middle Oligocene (Brule formation, Upper Oreodon beds) : East 
side of Plum Creek, Niobrara County, Wyoming. 


88 Aquila ferox Shufeldt proves to be a mammal. See Wetmore, Amer. Mus. 
Nov., No. 680, Dec. 4, 1933, pp. I-2. 

64 Howard, Auk, vol. 64, April 1947, pp. 287-201, finds that the abundant 
material from Rancho La Brea indicates a bird with longer wing, shorter leg, 
and larger skull than the living population. 


48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Subfamily CIRCINAE: Harriers 
Genus CIRCUS Lacépéde 


Circus LACEPEDE, Tabl. Ois., 17990, p. 4. Type, by subsequent designation, Falco 
aeruginosus Linnaeus (Lesson, 1828). 


Circus cyaneus (LINNAEUS): Marsh Hawk 
Falco cyaneus LINNAEUS, Syst. Nat., ed. 12, vol. 1, 1766, p. 126. 
Modern form reported from Pleistocene: San Josecito Cave, Aram- 


berri, Nuevo Leon. Late Pleistocene: Fossil Lake, Oregon; McKit- 
trick, and Rancho La Brea, Los Angeles, California. 


Family PANDIONIDAE: Ospreys 
Genus PANDION Savigny 


Pandion Savicny, Descr. Egypte, Ois., vol. 1, 1809, pp. 60, 96. Type, by 
monotypy, Pandion fluvialis Savigny = Falco haliaetus Linnaeus. 


Pandion haliaetus LINNAEUS: Osprey 
Falco Haliaetus LINNAEUws, Syst. Nat., ed. 10, vol. 1, 1758, p. OI. 


Modern form reported from Pleistocene: Melbourne (stratum 2), 
and Itchtucknee River, Florida. 


Family FALCONIDAE: Caracaras and FALcons 
Subfamily CARACARINAE: Caracaras 
Genus CARACARA Merrem 


Caracara MErreEM, in Ersch and Gruber, Allg. Encycl. Wiss. Kiinste, vol. 15, 
1826, p. 159. Type, by subsequent designation, Falco plancus Miller (Hell- 
mayr and Conover, 1949). 


Caracara prelutosus prelutosus (Howarp) 


Polyborus prelutosus Howarp, Carnegie Inst. Washington Publ. 487, July 7, 
1938, p. 226, pls. 1-3. 


Pleistocene : Seminole Field, Pinellas County, and Melbourne, Flor- 
ida. Late Pleistocene: McKittrick, Kern County; Carpinteria, Santa 
Barbara County; and Rancho La Brea (type locality), Los Angeles, 
California.® 


65 Recorded also from early Recent deposits at this site (Howard, H., and 
Miller, A. H., Carnegie Inst. Washington Publ. 514, 1939, p. 43) and from 
Quaternary deposits in Conkling Cavern, Organ Mountains, New Mexico. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 49 


Caracara prelutosus grinnelli (HowArp) 


Polyborus prelutosus grinnelli Howarp, Condor, vol. 42, No. 1, Jan. 19, 1940, 
p. 41. 


Pleistocene: San Josecito Cave, Aramberri, Nuevo Leon. 
Caracara latebrosus (WETMORE) 


Polyborus latebrosus WETMORE, Proc. Biol. Soc. Washington, vol. 33, Dec. 30, 
1920, p. 77, pl. 2, figs. 5, 6. 


Recent (extinct) : °° Cave deposits in Cueva Torajio, near Utuado, 
Puerto Rico. 


Subfamily FALCONINAE: Fatcons 
Genus FALCO Linnaeus 


Falco L1inNAEus, Syst. Nat., ed. 10, vol. 1, 1758, p. 88. Type, by subsequent 
designation, Falco subbuteo Linnaeus (A. O. U. Comm., 1886). 


Subgenus HIEROFALCO Cuvier 


Hierofalco Cuvier, Régne Animal, vol. 1, 1817 (Dec. 7, 1816), p. 312. Type, 
by monotypy, Falco candicans Gmelin. 


Falco mexicanus SCHLEGEL: Prairie Falcon 
Falco mexicanus SCHLEGEL, Abh. Geb. Zodl. Vergl. Anat., Heft 3, 1851, p. 15. 
Modern form reported from Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. Late Pleistocene: McKittrick, and Rancho La 
Brea, Los Angeles, California. 
Falco swarthi MILLER 


Falco swarthi L. H. Mitier, Condor, vol. 29, No. 3, May 15, 1927, p. 152, 
fig. 54. 


Late Pleistocene: McKittrick, California. 
Falco oregonus HowaArp 


Falco oregonus H. Howarp, Carnegie Inst. Washington Publ. 551, Jan. 25, 
1946, p. 178, pl. 1, figs. 2, 3. 


Late Pleistocene: Fossil Lake, Oregon. 


Subgenus RHYNCHODON Nitzsch 


Rhynchodon Nirzscu, Obs. Avium Art. Carot. Comm., 1829, p. 20. Type, by 
subsequent designation, Falco peregrinus Tunstall (A. O. U. Comm., 
1886). 


66 Included here since it has not been found in living form, being known only 
from bones. 


50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Falco peregrinus TUNSTALL: Peregrine Falcon 


Falco Peregrinus TuNSTALL, Orn. Brit., 1771, p. I. 


Modern form reported from Late Pleistocene: Potter Creek Cave, 
Shasta County, McKittrick, and Rancho La Brea, Los Angeles, 
California. 


Subgenus TINNUNCULUS Vieillot 


Tinnunculus Vietttor, Ois. Amér. Sept., vol. 1, 1807, p. 39. Type, by subse- 
quent designation, Falco columbarius Linnaeus (Walden, 1872). 


Falco columbarius LINNAEUS: Pigeon Hawk 


Falco columbarius LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 90. 


Modern form reported from late Pleistocene: McKittrick, and 
Rancho La Brea, Los Angeles, California. 


Falco ramenta WETMORE 


Falco ramenta WetMoreE, Proc. U. S. Nat. Mus., vol. 84, Nov. 3, 1936, p. 75, 
fig. 14. 


Miocene (Sheep Creek formation): Dawes County, Nebraska. 


Subgenus CERCHNEIS Boie 


Cerchneis Bote, Isis von Oken, vol. 19, Heft 10, October 1826, col. 970. Type, 
by monotypy, Falco rupicolus Daudin. 


Falco sparverius LINNAEUS: Sparrow Hawk 


Falco sparverius LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 90. 


Modern form reported from Pleistocene: Cavern deposits near 
Lecanto, Florida; San Josecito Cavern, Aramberri, Nuevo Leon. Late 
Pleistocene: Samwel and Potter Creek caves, Shasta County, McKit- 
trick, Carpinteria, and Rancho La Brea, Los Angeles, San Pedro 
(Palos Verdes sand), Los Angeles County, California. 


Order GALLIFORMES: Mecapopes, CurAssows, PHEASANTS, 
and Hoatzins 


Suborder GALLI: MercApropes, CurAssows, GRouUSE, and PHEASANTS 
Superfamily CRACOIDEA: Mecapopes, Curassows, and GUANS 
Family GALLINULOIDIDAE: GALLINULOIDEs 
Genus GALLINULOIDES Eastman 


Gallinuloides EAstmMANn, Geol. Mag., February 1900, p. 54. Type, by mono- 
typy, Gallinuloides wyomingensis Eastman. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 51 


Gallinuloides wyomingensis EASTMAN 


Gallinuloides wyomingensis EastMAN, Geol. Mag., n. s., vol. 7, pt. 4, No. 2, 
February 1900, p. 54, pl. 4. 


Middle Eocene (Green River formation): Fossil (type locality), 
and Henry’s Fork, Wyoming. 


Family CRACIDAE: Curassows, Guans, and CHACHALACAS 
Genus ORTALIS Merrem 


Ortalida (accusative case) —= Ortalis (nominative) Merrrem, Avium Rar. 
Icones et Descrip., vol. 2, 1786, p. 40. Type, by original designation, 
Phasianus motmot Linnaeus. 


Ortalis phengites WETMORE 


Ortalis phengites WerTMorE, Bull. Amer. Mus. Nat. Hist., vol. 48, art. 12, 
Dec. 3, 1023, p. 487, figs. 1-2. 


Lower Pliocene (Upper Snake Creek beds) : South of Agate, Sioux 
County, Nebraska. 
Ortalis tantala WETMORE 


Ortalis tantala WrtTMorrE, Condor, vol. 35, No. 2, Mar. 15, 1933, p. 64, figs. 
10-14. 


Lower Miocene (Lower Harrison beds): Carnegie Hill, Sioux 
County, Nebraska. 
Ortalis pollicaris MILLER 


Ortalis pollicaris A. H. Miter, Univ. California Publ., Bull. Dept. Geol. Sci., 
vol. 27, No. 4, June 22, 1944, p. 91, fig. 5. 


Lower Miocene (Rosebud formation): Flint Hill, 9 miles west- 
southwest of Martin, Bennett County, South Dakota. 


Genus BOREORTALIS Brodkorb 


Boreortalis BropKors, Wilson Bull., vol. 66, No. 3, September (Oct. 29), 
1954, p. 180. Type, by original designation, Boreortalis laessleit Brodkorb. 


Boreortalis laesslei BropKoRB 


Boreortalis laesslei Bropkors, Wilson Bull., vol. 66, No. 3, September (Oct. 
29), 1954, p. 182, fig. 1 (on p. 181). 


Lower Miocene (Hawthorn formation): Thomas Farm, 8 miles 
north of Bell, Gilchrist County, Florida. 


52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Superfamily PHASIANOIDEA: Grouse, Quaits, PHEASANTS, and TURKEYS 
Family TETRAONIDAE: Grouse and PTARMIGANS 


Genus DENDRAGAPUS Elliot 
Dendragapus Exttot, Proc. Acad. Nat. Sci. Philadelphia, vol. 16, No. 1, 
January-February (April 23), 1864, p. 23. Type, by subsequent designa- 
tion, Tetrao obscurus Say (Baird, Brewer, and Ridgway, 1874). 
Dendragapus obscurus (Say): Blue Grouse 
Tetrao obscurus Say, in Long, Exped. Rocky Mts., vol. 2, 1823, p. 14. 
Modern form reported from late Pleistocene: Samwel and Potter 
Creek caves, Shasta County, California. 


Dendragapus lucasi (SHUFELDT) ®7 
Pediocetes lucasi SuuFevpt, Auk, vol. 8, No. 4, October 1891, p. 367. 


Late Pleistocene: Fossil Lake, Oregon. 


Dendragapus nanus (SHUFELDT) ® 
Pediocetes nanus SHuFELptT, Amer. Nat., vol. 25, No. 297, September 1801, 
p. 821. 


Late Pleistocene: Fossil Lake, Oregon. 


Genus BONASA Stephens 


Bonasa STEPHENS, in Shaw, Gen. Zool., vol. 9, pt. 2, 1819, p. 208. Type, by 
subsequent designation, Tetrao umbellus Linnaeus (A. O. U. Committee, 


1886). 
Bonasa umbellus (LINNAEUS) : Ruffed Grouse °§ 
Tetrao umbellus LINNAEUS, Syst. Nat., ed. 12, vol. 1, 1766, p. 275. 


Modern form reported from Pleistocene: Cave near Frankstown, 
Pennsylvania ; Cumberland Cave, near Corriganville, Allegany County, 
Maryland ; caves of Tennessee. Late Pleistocene: Potter Creek Cave, 
Shasta County, California. 


Genus TYMPANUCHUS Gloger ®9 


Tympanuchus Grocer, Hand- und Hilfsbuch Naturg., 1842 (pp. 1-450, 1841), 
p. 396. Type, by monotypy, Tetrao cupido Linnaeus. 


67 Assigned to Dendragapus by Howard, Carnegie Inst. Washington Publ. 551, 
Jan. 25, 1946, p. 180. 

68 Bonasa ceres Shufeldt, Bull. Amer. Mus. Nat. Hist., vol. 32, Aug. 4, 1913, 
p. 299, pl. 55, figs. 18-20, pl. 56, figs. 45-72, from the Pleistocene of the fissure 
beds of Arkansas is possibly a synonym. On p. 300 of the reference cited the 
author alludes to it as Lagopus ceres. 

69 Records from Fossil Lake, Oregon, formerly placed under Tympanuchus 
pallidicinctus are now referred to Centrocercus urophasianus and Dendragapus 
lucasi. See Howard, Carnegie Inst. Washington Publ. 551, Jan. 25, 1946, p. 179. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 53 


Tympanuchus lulli SHUFELDT 
Tympanuchus lulli SHuretpt, Trans. Connecticut Acad. Arts Sci., vol. 10, 
February 1915, p. 69, pl. 12, fig. 90. 


? Pleistocene: 7° Hornerstown, New Jersey. 
Tympanuchus stirtoni MILLER 


Tympanuchus stirton’ A. H. Mitter, Univ. California Publ., Bull. Dept. Geol. 
Sci., vol. 27, No. 4, June 22, 1944, p. 92, fig. 6. 


Lower Miocene (Rosebud formation): Flint Hill, 9 miles west- 
southwest of Martin, Bennett County, South Dakota. 


Genus PEDIOECETES Baird 


Pedioecetes Batrp, Rep. Expl. and Surv. R. R. Pac., vol. 9, 1858, pp. xxi, xliv. 
Type, by monotypy, Tetrao phasianellus Linnaeus. 


Pedioecetes phasianellus (LINNAEUS): Sharp-tailed Grouse 


Tetrao Phasianellus Linnakus, Syst. Nat., ed. 10, vol. 1, 1758, p. 160. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon. 


Genus CENTROCERCUS Swainson 


Centrocercus SWAINSON, in Swainson and Richardson, Fauna Bor.-Amer., 
vol. 2, 1831 (1832), pp. 358, 4906. Type, by original designation, Tetrao 
urophasianus Bonaparte. 


Centrocercus urophasianus (BONAPARTE): Sage Grouse 


Tetrao urophasianus BoNAPARTE, Zool. Journ., vol. 3, No. 10, April-September, 
1827, p. 213. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon. 


Genus PALAEALECTORIS Wetmore 


Palaealectoris WETMORE, Condor, vol. 32, No. 3, May 15, 1930, p. 152. Type, 
by monotypy, Palaealectoris incertus Wetmore. 


Palaealectoris incertus WETMORE 
Palaealectoris incertus WETMoRE, Condor, vol. 32, No. 3, May 15, 1930, p. 152, 
figs. 51-53. 
Lower Miocene (Lower Harrison beds) : Agate fossil quarry, near 
Agate, Sioux County, Nebraska. 


Genus PALAEOTETRIX Shufeldt 


Palaeotetrix Suuretpt, Amer. Nat., vol. 25, No. 297, September 1891, p. 821. 
Type, by monotypy, Palaeotetrix gilli Shufeldt. 


70 Cited in the original description as “Post-Pliocene.” 


54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Palaeotetrix gilli SHUFELDT 


Palaeotetrix gilli SHuFELpT, Amer. Nat., vol. 25, No. 297, September 1801, 
p. 821. 


Late Pleistocene: Fossil Lake, Oregon. 


Genus PALAEOPHASIANUS Shufeldt 


Palaeophasianus SHUFELDT, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 16, 


Aug. 4, 1913, p. 291. Type, by monotypy, Palaeophasianus meleagroides 
Shufeldt. 


Palaeophasianus meleagroides SHUFELDT 


Palaeophasianus meleagroides SHUFELDT, Bull. Amer. Mus. Nat. Hist., vol. 
32, art. 16, Aug. 4, 1913, p. 201, pl. 58, figs. 81-84, 86-88. 


Lower Eocene (Wasatch) : Elk Creek, Big Horn Basin (type lo- 
cality). Eocene (Bridger) : Henry’s Fork, Wyoming. 


Family PHASIANIDAE: Quaits, PHEASANTS, and PEACcocKS 
Subfamily ODONTOPHORINAE: American QUAILS 


Genus COLINUS Goldfuss 


Colinus Gotpruss, Handb. Zool., vol. 2, 1820, p. 220. Type, by monotypy, 
Perdix mexicanus, Caille de la Louisiane, Pl. Enl. 149 = Tetrao virgini- 
anus Linnaeus. 


Colinus virginianus (LINNAEUS): Bobwhite 
Tetrao virginianus LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 161. 
Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Melbourne, and cavern deposits near Lecanto, Florida; caves 
of Tennessee. 
Colinus hibbardi WETMORE 


Colinus hibbardi Wetmore, Univ. Kansas Sci. Bull., vol. 30, pt. 1, No. 9, 
May 15, 1944, p. 96, figs. 4-8. 


Upper Pliocene (Rexroad fauna) : Meade County, Kansas. 


? Colinus eatoni SHUFELDT 7! 


Colinus eatoni SHUFELDT, Trans. Connecticut Acad. Arts Sci., vol. 19, Febru- 
ary 1915, p. 70, pl. 13, fig. 103. 


Geologic age uncertain: Western Kansas. 


71 Relationship uncertain. From the published figure it may possibly be an 
oscinine passeriform. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 55 


Genus LOPHORTYX Bonaparte 


Lophortyx Bonaparte, Geogr. and Comp. List, 1838, p. 42. Type, by subse- 
quent designation, Tetrao californicus Shaw (Gray, 1840). 


Lophortyx californicus (SHAW): California Quail 
Tetrao californicus SHAW, in Shaw and Nodder, Nat. Misc. vol. 9, 1798, text 
to pl. 345. 


Modern form reported from late Pleistocene: Hawver Cave, El- 
dorado County, Carpinteria, McKittrick, Rancho La Brea, Los 
Angeles and San Pedro (Palos Verdes sand), Los Angeles County, 
California. 


Genus OREORTYX Baird 


Oreortyx Batrp, Rep. Expl. and Surv. R. R. Pac., vol. 9, 1858, pp. xxi, xlv, 
638, 642. Type, by original designation, Ortyx picta Douglas. 


Oreortyx pictus (Doucras) : Mountain Quail 
Ortyx picta Douctas, Trans. Linn. Soc. London, vol. 16, pt. 1, 1829, p. 143. 
Modern form reported from late Pleistocene: Potter Creek and 
Samwel caves, Shasta County, and Hawver Cave, Eldorado County, 


California. Quaternary (probably Recent): Rocky Arroyo, New 
Mexico. 


Genus MIORTYX Miller 


Miortyx A. H. Mixer, Univ. California Publ., Bull. Dept. Geol. Sci., vol. 27, 
No. 4, June 22, 1944, p. 93. Type, by original designation, Miortyx teres 
Miller. 


Miortyx teres MILLER 


Miortyx teres A. H. Miter, Univ. California Publ., Bull. Dept. Geol. Sci., 
vol. 27, No. 4, June 22, 1944, p. 93, fig. 7. 


Lower Miocene (Rosebud formation): Flint Hill, 9 miles west- 
southwest of Martin, Bennett County, South Dakota. 


Genus CYRTONYX Gould 


Cyrtonyx GouLtp, Monogr. Odontophoridae, pt. 1, 1844, pl. and text. Type, by 
monotypy, Ortyx massena Lesson = Ortyx montezumae Vigors. 


Cyrtonyx montezumae (Vicors): Harlequin Quail 


Ortyx Montezumae Vicors, Zool. Journ., vol. 5, June 1830, p. 275. 


Modern form reported from Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. 


50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Cyrtonyx cooki WETMORE 
Cyrtonyx cooki Wetmore, Condor, vol. 36, No. 1, Jan. 15, 1934, p. 30, fig. 5. 
Upper Miocene (Upper Sheep Creek beds): 17 miles south of 
Agate, Sioux County, Nebraska. 
Cyrtonyx tedfordi MILter 72 
Cyrtonyx tedfordi L. H. Mritter, Condor, vol. 54, No. 5, Sept. 22, 1952, p. 208, 
hig. 
Upper Miocene (Barstow formation): Lake bed horizon, near 
Barstow, California. 


Subfamily PHASIANINAE: Ortp Wortp PartripcEs and 
PHEASANTS 


Genus PHASIANUS Linnaeus 
Phasianus LINNAEvS, Syst. Nat., ed. 10, vol. 1, 1758, p. 158. Type, by tau- 


tonymy, Phasianus colchicus Linnaeus. 
Phasianus alfhildae SHuFreLpt 7° 


Phasianus alfhildae SuHuFeLpt, Trans. Connecticut Acad. Arts Sci., vol. 19, 
February 1015, p. 71. 


Geologic age uncertain: 100 feet below horizon of Haystack Butte, 
Haystack Mountain, Wyoming. 


Genus ARCHAEOPHASIANUS Lambrecht 


Archaeophasianus LAMBRECHT, Handb. Palaeorn., 1933, p. 438. Type, by sub- 
sequent designation, Phasianus roberti Stone (Brodkorb, 1952). 


Archaeophasianus roberti (STONE) 


Phasianus roberti Stone, Auk, vol. 32, No. 3, July (June 29), 1915, p. 376. 


Lower Miocene (Middle John Day formation) : Paulina ™* Creek, 
6 miles from junction with Beaver Creek, Crook County, Oregon. 


? Archaeophasianus mioceanus (SHUFELDT) 75 


Phasianus mioceanus SHUFELDT, Trans. Connecticut Acad. Arts Sci., vol. 10, 
February 1915, p. 60, pl. 13, figs. 94, 96. 


Miocene; Chimney Rock and Scott’s Bluff, Nebraska. 


72 Allocation in this genus tentative. 

73 Allocation of this species to the Old World genus Phasianus follows the 
usage of the original describer, and is subject to verification. 

74 Given as “Parilina” in the original place of publication, through an error 
in reading the field label. 

75 Described from fragmentary humerus and femur from the two separate 
localities listed. Probably a composite, with neither bone coming from a bird of 
this family. Assigned to Archaeophasianus by Lambrecht. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 57 


Family MELEAGRIDIDAE: Turkeys 
Genus MELEAGRIS Linneaus 


Meleagris LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 156. Type, by tau- 
tonymy, Meleagris gallopavo Linnaeus. 
Meleagris gallopavo LINNAEUS: Turkey 76 
Meleagris Gallopavo LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 156. 


Modern form reported from Upper Pliocene (Rexroad formation) : 
Meade County Kansas. Pleistocene: Hartman’s or Crystal Hill Cave, 
near Stroudsburg, and Durham Cave, near Riegelsville, Bucks County, 
and caves near Carlisle, Pennsylvania; North Liberty, St. Joseph 
County, Indiana; Ashmore, Coles County, Illinois; caves of Tennes- 
see; fissure beds, Arkansas; Seminole Field, Pinellas County, Sara- 
sota, Bradenton, Itchtucknee River, Melbourne, and cavern deposits at 
Ocala and Lecanto, Florida; near San Antonio, Socorro County, New 
Mexico.”” 


Meleagris antiqua Marsu 
Meleagris antiquus MarsH, Amer. Journ. Sci., ser. 3, vol. 2, August 1871, 
p. 126. 
Oligocene (White River formation) : “G Ranch,” Colorado. 


Meleagris celer MArsH 


Meleagris celer Marsu, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, p. 261. 
Pleistocene: Monmouth County, New Jersey. 


Meleagris richmondi SHUFELDT 


Meleagris richmondi SuureLpt, Trans. Connecticut Acad. Arts Sci., vol. 19, 
February 1915, p. 67, pl. 2, fig. 10. 


Pleistocene: Near Mission San Jose, Alameda County, California. 


Meleagris superba Cope 


Meleagris superbus Corr, Trans. Amer. Philos. Soc., n.s., vol. 14, pt. 1, 
December 1870, p. 239. 


Pleistocene: Monmouth County (type locality), and Manalapan,”® 
New Jersey; Frankstown and Port Kennedy caves, Pennsylvania. 


76 Ardea sellardsi Shufeldt, 9th Ann. Rep. Florida State Geol. Surv., 1917, 
p. 38, pl. 2, fig. 15, from Vero (stratum 3) is a synonym of Meleagris gallopavo 
according to Wetmore, Smithsonian Misc. Coll., vol. 85, No. 2. Apr. 13, 1931, 
Ppp. 10-11, 32-33. The deposit is now considered to be of Recent age. See Cooke, 
Florida Geol. Surv. Geol. Bull. 29, 1945, pp. 306-307. 

77 Possibly Upper Pliocene. 

78 Type locality of Meleagris altus Marsh, Amer. Journ. Sci., ser. 3, vol. 4, 
1872, p. 260, which is a synonym. 


58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Meleagris tridens WETMORE 


Meleagris tridens WrTMoRE, Smithsonian Misc. Coll., vol. 85, No. 2, Apr. 
13, 1931, p. 33, fig. 13, pl. 6. 


Pleistocene: Seminole Field, Pinellas County, Florida. 
Meleagris crassipes MILLER 


Meleagris crassipes L. H. Miter, Condor, vol. 42, No. 3, May 15, 1940, p. 154, 
figs. 44-45. 


Pleistocene: San Josecito Cave, Aramberri, Nuevo Leon. 


Genus PARAPAVO Miller 


Parapavo L. H. Mitver, Univ. California Publ., Bull. Dept. Geol., vol. 9, 
No. 9, Mar. 10, 1916, p. 96. Type, by monotypy, Pavo californicus Miller. 


Parapavo californicus (MILLER) 


Pavo californicus L. H. Mitter, Univ. California Publ., Bull. Dept. Geol., 
vol. 5, No. 19, Aug. 14, 1900, p. 285, pl. 25. 


Upper Pliocene: Cita Canyon, Randall County, Texas. Pleistocene: 
York Valley site at Avenue 45 and Lincoln Avenue, Highland Park, 
Los Angeles, and southwest of La Habra near Los Angeles—Orange 
County line, California. Late Pleistocene: Carpinteria, and Rancho 
La Brea (type locality),7° Los Angeles, California. 


Order GRUIFORMES: Cranes, Rats, and ALLIES 
Suborder GRUES: Cranes, LImMpKINS, TRUMPETERS, and RAILS 


Superfamily GRUOIDEA: Cranes, Limpkins, and TRUMPETERS 


Family GERANOIDIDAE: GERANOIDES 
Genus GERANOIDES Wetmore 


Geranoides WetTMorE, Condor, vol. 35, No. 3, May 15, 1933, p. 115. Type, 
by original designation, Geranoides jepseni Wetmore. 


Geranoides jepseni WETMORE 
Geranoides jepsent WeEtTMoRE, Condor, vol. 35, No. 3, May 15, 1933, p. 115, 
fig. 22. 


Lower Eocene (Gray Bull member): South Elk Creek, Bighorn 
County, Wyoming. 


79 Recorded also from early Recent deposits in Pit 10 at this site (Howard, H., 
and Miller, A. H., Carnegie Inst. Washington Publ. 514, 1939, p. 43). Parapavo 
oklahomaensis Stovall and Sandoz, Proc. Oklahoma Acad. Sci., vol. 16, 1936, 
p. 77, is a nomen nudum. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 59 


Family GRUIDAE: Cranes 
Subfamily GRUINAE: Cranes 
Genus ALETORNIS Marsh ®° 


Aletornis Marsu, Amer. Journ. Sci., ser. 3, vol. 14, October 1872, p. 256. 
Type, by subsequent designation, Aletornis nobilis Marsh (Hay, 1902). 


Aletornis bellus Marsu 81 


Aletornis bellus Marsu, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, 
p. 258. 


Eocene (Bridger formation) : Grizzly Buttes, Wyoming. 


Aletornis gracilis MArsuH 81 


Aletornis gracilis Marsu, Amer. Journ. Sci. ser. 3, vol. 4, October 1872, 
p. 258. 


Eocene (Bridger formation) : Henry’s Fork, Wyoming. 


Aletornis nobilis Marsu 82 


Aletornis nobilis MarsH, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, p. 256. 
Eocene (Bridger formation) : Grizzly Buttes, Wyoming. 


Aletornis pernix MArsH 


Aletornis pernix Marsu, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, p. 256. 


Eocene (Bridger formation) : Henry’s Fork, Wyoming. 


Genus FULICALETORNIS Lambrecht 


Fulicaletornis LAMBRECHT, Handb. Palaeorn., 1933, p. 479. Type, by mono- 
typy, Aletornis venustus Marsh. 


80 Allocation in the subfamily Gruinae provisional. 

81 Considered by Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, Febru- 
ary 1915, pp. 32, 76, as possibly a species of Scolopacidae. 

82 Marsh in his original proposal of the genus Aletornis included in it five 
species without selecting a type. From the five in question Hay, U. S. Geol. 
Surv., Bull. 179, 1902, p. 527, designated Aletornis nobilis Marsh as genotype. 
Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, 1915, pp. 30, 31, placed 
A. nobilis in Grus, and described in the same paper (p. 77) Grus marshi. Lam- 
brecht, Handb. Palaeorn., 1933, p. 520, proposed the genus Protogrus for 
Aletornis nobilis and Grus marshi, without designating a type. Lambrecht’s 
action as regards A. nobilis obviously is erroneous as his proposed genus in- 
cludes the genotype of Aletornis. Aletornis nobilis, therefore, is to be listed 
as above, and pending study Grus marshi is included tentatively under Grus. 
Brodkorb, Condor, vol. 54, No. 3, May 21, 1952, p. 175, has designated A. nobilis, 
already the type of Aletornis through action by Hay, as the type of Protogrus. 
That generic name therefore becomes a synonym of Aletornis. 


60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Fulicaletornis venustus (MarsH) ® 


Aletornis venustus MarsH, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, 
Pp. 257. 


Eocene (Bridger formation): Henry’s Fork, Wyoming. 


Genus PARAGRUS Lambrecht 


Paragrus LAMBRECHT, Handb. Palaeorn., 1933, p. 520. Type, by monotypy, 
Gallinuloides prentici Loomis. 


Paragrus prentici (Loomis) 


Gallinuloides prentici F. B. Loomis, Amer. Journ. Sci., ser. 4, vol. 22, Decem- 
ber 1906, p. 481, figs. 1-3. 


Eocene (Wasatch): Head of Elk Creek, 10 miles west of Otto, 
Wyoming. 


Genus GRUS Pallas 


Grus Patras, Misc. Zool., 1766, p. 66. Type, by tautonymy, Ardea grus Lin- 
naeus. 


Grus americana (LINNAEUS): Whooping Crane 


Ardea americana LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 142. 


Modern form reported from late Upper Pliocene: Snake River, 
13 miles northwest of Grandview, Idaho. Pleistocene: Seminole Field, 
Pinellas County, Itchtucknee River, and Melbourne (stratum 2), Flor- 
ida. Late Pleistocene: Rancho La Brea, Los Angeles, California. 


Grus canadensis (LINNAEUS) : Sandhill Crane *4 


Ardea canadensis LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 141. 


Modern form reported from Lower Pliocene (Upper Snake Creek 
beds) : Sioux County, Nebraska. From ? Pleistocene: Niobrara River, 
Nebraska,*® and Grizzly Buttes, Wyoming. From Pleistocene: Ash- 


83 Systematic allocation provisional. Shufeldt, Trans. Connecticut Acad. Arts 
Sci., vol. 19, February 1915, pp. 31, 32, 76, placed this species in the genus 
Fulica, the principal basis for Lambrecht’s action in proposing Fulicaletornis. 

84 Grus canadensis is used as a species name to cover records of cranes of 
this type from the Pliocene and Pleistocene, including specimens that range 
in size from the modern little brown crane to the larger races of the sandhill 
crane. 

Grus minor L. H. Miller, Univ. California Publ., Bull. Dept. Geol., vol. 5, 
August 1910, p. 446, fig. 8, from the Pleistocene of Rancho La Brea, is now 
considered by the describer as a synonym of Grus canadensis. 

85 This specimen, from either Pliocene or Pleistocene deposits, is the basis 
of Grus haydeni Marsh, Amer. Journ. Sci., ser. 2, vol. 49, 1870, p. 214, con- 
sidered by Wetmore, Amer. Mus. Nov., No. 302, Feb. 29, 1928, p. 4, as a synonym 
of Grus canadensis. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 61 


more, Coles County, Illinois; Melbourne, Seminole Field, Pinellas 
County, and Bradenton, Florida. Late Pleistocene: Rancho La Brea, 
Los Angeles, and McKittrick, California. 


Grus proavus MarsH 
Grus proavus Marsu, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, p. 261. 
Pleistocene: Monmouth County, New Jersey. 


Grus nannodes WETMorRE and MARTIN 


Grus nannodes WrETMORE and Martin, Condor, vol. 32, No. 1, Jan. 20, 1930, 
p. 62, figs. 23-25. 


Middle Pliocene (Ogallala formation, Edson beds) : Sec. 25, T. 10 
S., R. 38 W., Sherman County, Kansas. 


Grus conferta MILLER and SIBLEY 


Grus conferta A. H. Miter and C. G. S1stey, Condor, vol. 44, No. 3, May 15, 
1942, p. 126, fig. 50. 


Late Lower Pliocene (Siesta formation): Black Hawk Ranch, 
southern base of Mount Diablo, Contra Costa County, California. 


Grus marshi SHUFELDT 86 


Grus marshi SHUFELDT, Trans. Connecticut Acad. Arts Sci., vol. 19, February 
1915, p. 77, pl. 15, figs. 144-147. 


Eocene (Bridger formation) : Henry’s Fork, Wyoming. 


Family ARAMIDAE: Limpxkins 


Genus ARAMUS Vieillot 


Aramus Vietttot, Analyse, 1816, p. 58. Type, by monotypy, Courliri Buf- 
fon = Ardea scolopacea Gmelin. 


Aramus guarauna LINNAEUS: Limpkin 


Scolopax Guarauna LINNAEUS, Syst. Nat., ed. 12, vol. 1, 1766, p. 242. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, and Itchtucknee River, Florida. 


Genus BADISTORNIS Wetmore 


Badistornis WrtTMorE, Journ. Morph., vol. 66, Jan. 2, 1940, p. 30. Type, by 
original designation, Badistornis aramus Wetmore. 


86 Generic allocation doubtful. See footnote under Aletornis nobilis (p. 59). 


62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Badistornis aramus WETMORE 


Badistornis aramus Wetmore, Journ. Morph., vol. 66, Jan. 2, 1940, p. 30, 
figs. 7-10. 


Oligocene (Metamynodon zone, Brule formation) : 35 miles south- 
west of Scenic, South Dakota. 


Genus ARAMORNIS Wetmore 


Aramornis WetMore, Amer. Mus. Nov., No. 211, Mar. 11, 1926, p. 1. Type, 
by original designation, Aramornis longurio Wetmore. 


Aramornis longurio WETMORE 


Aramornis longurio WretMorE, Amer. Mus. Nov., No. 211, Mar. 11, 1926, 
p. I, figs. 1-4. 


Middle Miocene (Lower Sheep Creek beds) : Snake Creek Quar- 
ries, Sioux County, Nebraska. 


Genus GNOTORNIS Wetmore 


Gnotornis WETMORE, Smithsonian Misc. Coll., vol. 101, No. 14, May 11, 1942, 
p. 1. Type, by monotypy, Gnotornis aramiellus Wetmore. 


Gnotornis aramiellus WETMORE 


Gnotornis aramicllus Wetmore, Smithsonian Misc. Coll., vol. 101, No. 14, 
May II, 1942, p. I, figs. I-4. 


Upper Oligocene (Upper Brule formation, Protoceras—Leptau- 
chenia beds) : 25 miles southeast of Scenic and 6 miles east of Rocky- 
ford, Washington County, South Dakota. 


Superfamily RALLOIDEA: Rams 

Family RALLIDAE: Ratrs, GALLINULES, and Coots 
Subfamily RALLINAE: Rats 
Genus TELMATORNIS Marsh 7 


Telmatornis MArsH, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, p. 210. 
Type, by subsequent designation, Telmatornis priscus Marsh (Hay, 1902). 


Telmatornis affinis MArsH 


Telmatornis affints Marsu, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, 
p. 211. 


Paleocene (Hornerstown marl): Hornerstown, New Jersey. 


87 Allocation in the subfamily Rallinae provisional. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 63 


Telmatornis priscus MArsH 


Telmatornis priscus MArsH, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, 
p. 210. 


Paleocene (Hornerstown marl): Hornerstown, New Jersey. 


Telmatornis rex SHUFELDT 


Telmatornis rex SHUFELDT, Trans. Connecticut Acad. Arts Sci., vol. 19, Feb- 
ruary IQI5, p. 27, pl. 13, fig. IOI. 


Paleocene (Hornerstown marl): Hornerstown, New Jersey. 


Genus PALAEORALLUS Wetmore 


Palacorallus WrtTMoreE, Condor, vol. 33, No. 3, May 15, 1931, p. 108. Type, 
by original designation, Palaeorallus troxelli Wetmore. 


Palaeorallus troxelli WETMORE 


Palaeorallus troxelli WEtTMorRE, Condor, vol. 33, No. 3, May 15, 1931, p. 108, 
figs. 26-20. 


Lower Eocene (Wasatch formation) : Northwest of Little Tatman 
Mountain, near Burlington, Wyoming. 


Genus CRECCOIDES Shufeldt 


Creccoides SuureLpt, Proc. Amer. Philos. Soc., vol. 30, Apr. 14, 1892, p. 125. 
Type, by monotypy, Creccoides osbornii Shufeldt. 


Creccoides osbornii SHUFELDT 


Creccoides osborniti SHuUFELDT, Proc. Amer. Philos. Soc., vol. 30, Apr. 14, 
1892, p. 125. 


Pliocene (Blanco fauna) : Blanco Canyon, Crosby County, Texas. 


Genus EPIRALLUS Miller 


Epirallus L. H. Mitrer, Univ. California Publ. Zool., vol. 47, Mar. 6, 1942, 
p. 43. Type, by monotypy, Epirallus natator Miller. 


Epirallus natator MILLER 


Epirallus natator L. H. Mitier, Univ. California Publ. Zool., vol. 43; Mar. 6, 
1942, p. 43, fig. Ia. 


Pleistocene: San Josecito Cave, Aramberri, Nuevo Leon. 


Genus RALLUS Linnaeus 


Rallus LinNAEus, Syst. Nat., ed. 10, vol. 1, 1758, p. 153. Type, by subsequent 
designation, Rallus aquaticus Linnaeus (Fleming, 1821). 


Rallus elegans Aupuson: King Rail 


Rallus elegans Aupuxzon, Birds Amer. (folio), vol. 3, 1834, pl. 203. 


64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, and Itchtucknee River, Florida. 
Rallus longirostris BoppAErr: Clapper Rail 

Rallus longirostris Bopparert, Table Planch. Enlum., 1783, p. 52. 

Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 
Rallus limicola ViEILLor: Virginia Rail 

Rallus limicola Viettiot, Nouv. Dict. Hist. Nat., nouv. éd., vol. 28, May 18109, 

p. 558. 


Modern form recorded from Pleistocene: Reddick, Marion County, 
Florida. Late Pleistocene: Fossil Lake, Oregon; McKittrick, Cali- 
fornia. 

Rallus prenticei WETMORE 
Rallus prenticei Wetmore, Univ. Kansas Sci. Bull., vol. 30, pt. 1, No. 9, 
May 15, 1944, p. 99, figs. 9-19. 


Upper Pliocene (Rexroad fauna): Meade County, Kansas. 


Genus PORZANA Vieillot 


Porzana Vie1tuot, Analyse, 1816, p. 61. Type, by monotypy and tautonymy, 
Marouette Buffon = Rallus porzana Linnaeus. 


Porzana carolina (LINNAEUS): Sora 
Rallus carolinus LINNAEUvS, Syst. Nat., ed. 10, vol. 1, 1758, p. 153. 
Pleistocene: Near Reddick, Marion County, Florida. 


Porzana auffenbergi BropKorB 
Porzana auffenbergi BropKors, Condor, vol. 56, No. 2, Mar. 26, 1954, p. 103, 
fig. I. 


Pleistocene (stratum 2, shell layer, Sangamon stage) : near Haile, 
Alachua County, Florida. 


Genus LATERALLUS Gray 
Laterallus G. R. Gray, Cat. Gen. Subgen. Birds, 1855, p. 120. Type, by mono- 
typy, Rallus melanophaius Vieillot. 
Laterallus guti BropkKors 
Laterallus guti BropKors, Wilson Bull., vol. 64, No. 2, June 16, 1952, p. 80, 
fig. I. 


Pleistocene: 1 mile south of Reddick, Marion County, Florida. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 65 


Genus ARAMIDES Pucheran 


Aramides PUCHERAN, Rev. Zool., vol. 8, August 1845, p. 277. Type, by origi- 
nal designation, /ulica cayennensis Gmelin. 


Aramides cajanea (MULLER): Wood Rail 
Fulica Cajanea P. L. S. MULLER, Natursyst. Suppl., 1776, p. 119. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 


Genus NESOTROCHIS Wetmore 


Nesotrochis WrtMorkE, Proc. U. S. Nat. Mus., vol. 54, Nov. 21, 1918, p. 516. 
Type, by original designation, Nesotrochis debooyi Wetmore. 


Nesotrochis debooyi WETMORE 
Nesotrochis debooyt WrEtTMorE, Proc. U. S. Nat. Mus., vol. 54, Nov. 21, 1918, 
p. 516, pl. 82. 


Recent (extinct) : 8° Archeological sites on St. Thomas ®° and St. 
Croix, Virgin Islands; and at Barrio Cafias, near Ponce; cavern 
deposits in Cueva Clara and Cueva San Miguel, near Morovis ; Cueva 
Torafio, and a cave on Hacienda Jobo, near Utuado, Puerto Rico. 


Subfamily GALLINULINAE: GALtInuLes 
Genus PORPHYRULA Blyth 


Porphyrula Brytu, Cat. Birds Mus. Asiat. Soc., 1849 (1852), p. 283. Type, 
by monotypy, P. chloronotus Blyth = Porphyrio alleni Thomson. 
Porphyrula martinica (LINNAEUS): Purple Gallinule 
Fulica martinica LINNAEUS, Syst. Nat., ed. 12, vol. 1, 1766, p. 259. 


Modern form reported from Pleistocene: Haile, Alachua County, 
Florida. 


Genus GALLINULA Brisson 
Gallinula Brisson, Orn., 1760, vol. 1, p. 50; vol. 6, p. 2. Type, by tautonymy 
Gallinula Brisson = Fulica chloropus Linnaeus. 
Gallinula chloropus (LINNAEUS) : Common Gallinule 
Fulica Chloropus LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 152. 


Modern form reported from Upper Pliocene (Hagerman lake 
beds) : Near Hagerman, Idaho. From Pleistocene: °° Seminole Field, 


88 Included here as it has not been found in living form, being known only 
from bones. Possibly the species lived until Spanish colonial times. 

89 Type locality a kitchen midden at Magen’s Bay, on the north coast of 
St. Thomas. 

80 Reported from Pleistocene at Haile, Alachua County, Florida, on basis of a 


66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


Pinellas County, and Itchtucknee River, Florida. Late Pleistocene: 
Banos de Ciego Montero, Cuba. 


Genus PALAEOCREX Wetmore 91 


Palaeocrex Wetmore, Proc. Colorado Mus. Nat. Hist., vol. 7, No. 2, July 
15, 1927, p. 9. Type, by monotypy, Palaeocrex fax Wetmore. 


Palaeocrex fax WETMORE 
Palaeocrex fax Wetmore, Proc. Colorado Mus. Nat. Hist., vol. 7, No. 2, 
July 15, 1927, p. 9, figs. 15-18. 


Lower Oligocene (Chadronian, Horsetail Creek facies) : Horsetail 
Creek, Weld County, Colorado. 


Genus EOCREX Wetmore 


Eocrex Wetmore, Condor, vol. 33, No. 3, May 15, 1931, p. 107. Type, by 
original designation, Eocrex primus Wetmore. 


Eocrex primus WETMORE 
Eocrex primus WetMore, Condor, vol. 33, No. 3, May 15, 1931, p. 107, figs. 
21-25. 
Lower Eocene (‘‘Wasatch” formation) : Near Steamboat Springs, 


Sweetwater County, Colorado (sec. 13, T. 24 N., R. 102 W., in 
Cathedral Bluffs). 


Subfamily FULICINAE: Coors 
Genus FULICA Linnaeus 


Fulica LinnNAEus, Syst. Nat., ed. 10, vol. 1, 1758, p. 152. Type, by subsequent 
designation, Fulica atra Linnaeus (Gray, 1840). 


Fulica americana GMELIN: American Coot 


Fulica americana GMELIN, Syst. Nat., vol. 1, pt. 2, 1780, p. 704. 


Modern form recorded from Upper Pliocene (Rexroad fauna) : 
Meade County, Kansas. Pleistocene: Seminole Field, Pinellas County, 
Bradenton, Itchtucknee River, and Haile, Alachua County, Florida; 
Hemphill County, Texas ; San Josecito Cave, Aramberri, Nuevo Leon. 
Late Pleistocene: Rancho La Brea, Los Angeles, and San Pedro 
(Palos Verdes formation), Los Angeles County, California. 


cervical vertebra, by Brodkorb, Wilson Bull., vol. 65, No. 1, March (Apr. 22), 


1953, P- 50. 
91 Subfamily allocation provisional. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 67 


Fulica minor SHUFELDT 22 


Fulica minor SHurFetpt, Amer. Nat., vol. 25, No. 297, September 1801, p. 820. 


Late Pleistocene: Fossil Lake, Oregon. 


Suborder CARIAMAE: CartamMas and ALLIES 
Family BATHORNITHIDAE: BatnornitHEs 
Genus BATHORNIS Wetmore 


Bathornis WeEtTMorE, Proc. Colorado Mus. Nat. Hist., vol. 7, No. 2, July 15, 
1927, p. 11. Type, by monotypy, Bathornis veredus Wetmore. 


Bathornis veredus WETMORE 
Bathornis veredus WETMORE, Proc. Colorado Mus. Nat. Hist., vol. 7, No. 2, 
July 15, 1927, p. 11, figs. 19-24. 
Lower Oligocene (Chadronian, Horsetail Creek facies) : Horsetail 
Creek, Weld County, Colorado (type locality) ; near Crawford, Ne- 
braska ; Indian Creek, Pennington County, South Dakota. 


Bathornis celeripes WETMORE 


Bathornis celeripes WWetTMorE, Bull. Mus. Comp. Zodl., vol. 75, October 1933, 
Pp. 302, figs. 6-14. 


Upper Oligocene (Brule formation): Near Torrington, Goshen 


County, Wyoming (type locality) ; 12 miles northwest of Crawford, 
Nebraska. 


Bathornis cursor WETMORE 


Bathornis cursor Wetmore, Bull. Mus. Comp. Zodl., vol. 75, October 1933, 
p. 310, figs. 15-109. 


Upper Oligocene (Brule formation): Near Torrington, Goshen 
County, Wyoming. 


Bathornis geographicus WETMORE 


Bathornis geographicus Wetmore, Smithsonian Misc. Coll., vol. 101, No. 14, 
May 11, 1942, p. 3, figs. 5-13. 


Upper Oligocene (Upper Brule formation, Protoceras—Leptau- 
chenia beds) : 25 miles southeast of Scenic and 6 miles east of Rocky- 
ford, Washington County, South Dakota. 


®2 Howard (Carnegie Inst. Washington Publ. 551, Jan. 25, 1946, pp. 182-183) 
places all Fulica records from Fossil Lake, Oregon, under this name. She con- 
siders minor the Pleistocene ancestor of modern Fulica americana, listing it as 
Fulica americana minor, the relationship indicated by the trinomial expressing 
distribution through geologic time and not the geographic range of two sub- 
species existing simultaneously. 


68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Order DIATRYMIFORMES: DtarrymMas 
Family DIATRYMIDAE: Dratrymas 
Genus BARORNIS Marsh 


Barornis Marsu, Amer. Journ. Sci., ser. 3, vol. 48, 1894, p. 344. Type, by 
monotypy, Barornis regens Marsh. 


Barornis regens MArsH 93 


Barornis regens Marsu, Amer. Journ. Sci., ser. 3, vol. 48, October 1894, p. 344, 
text fig. 


Eocene: Squankum, Monmouth County, New Jersey. 


Genus DIATRYMA Cope 


Diatryma Corr, Proc. Acad. Nat. Sci. Philadelphia, vol. 28, sign. 2, April 18, 
1876, p. 11. Type, by monotypy, Diatryma gigantea Cope. 
Diatryma ajax SHUFELDT 
Diatryma ajax SHUFELDT, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 16, 
Aug. 4, 1913, p. 287, pl. 52, figs. 4-5, pl. 53, figs. 8-10, pl. 54, figs. 13-14. 


Lower Eocene (Wasatch formation) : 3 (type locality) and 5 miles 
southeast of mouth of Pat O’Hara Creek, Clark’s Fork Basin, Wy- 
oming. 

Diatryma giganteum Cope 
Diatryma gigantea Corr, Proc. Acad. Nat. Sci. Philadelphia, vol. 28, sign. 2, 
Apr. 18, 1876, p. II. 
Lower Eocene (Wasatch formation) : New Mexico. 


Diatryma steini MATTHEW and GRANGER 


Diatryma steint MATTHEW and GRANGER, Bull. Amer. Mus. Nat. Hist., vol. 
37, art. 11, May 28, 1917, p. 322, pls. 20-33. 


Lower Eocene (Wasatch, Gray Bull member) : South Elk Creek, 
Bighorn Basin, Wyoming. 


Genus OMORHAMPHUS Sinclair 


Omorhamphus Sincratr, Proc. Amer. Philos. Soc., vol. 67, 1928, p. 51. Type, 
by monotypy, Omorhamphus storchit Sinclair. 


93 Considered a species of Diatryma by Shufeldt, Trans. Connecticut Acad. 
Arts Sci., vol. 19, February 1915, pp. 37-38. 

94 Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, February 1915, p. 34, 
refers a fragment in Peabody Museum, Yale University, from Island Point, 
North Horseshoe, Gallina, New Mexico, to this species. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 69 


Omorhamphus storchii SINCLAIR 


Omorhamphus storchit Stncuatr, Proc. Amer. Philos. Soc., vol. 67, 1928, p. 52, 
pls. 1-2, figs. 1-3. 


Lower Eocene (Lower Gray Bull horizon, Lower Wasatch): 14 
miles southeast of Dorsey Creek, about 2 miles south of Otto-Basin 
Road, Big Horn County, Wyoming. 


Order CHARADRIIFORMES: Suoresirps, Gutts, and AuKs 
Suborder CHARADRII: Sworeszirps 
Superfamily CHARADRIOIDEA: Ptovers, SANDpIPERS, and ALLIES 
Family RHEGMINORNITHIDAE: RuHEGMINOoRNIS 
Genus RHEGMINORNIS Wetmore 


Rhegminornis WETMORE, Proc. New England Zool. Club, vol. 22, June 23, 
1943, p. 61. Type, by original designation, Rhegminornis calobates Wet- 
more. 


Rhegminornis calobates Wetmore, Proc. New England Zool. Club, vol. 22, 
June 23, 1943, p. 61, pl. 11, figs. 1-5. 


Lower Miocene (Tampa limestone) : *° Thomas Farm, 8 miles north 
of Bell, Gilchrist County, Florida. 


Family HAEMATOPODIDAE: OystTERcATCHERS 
Genus PARACTIORNIS Wetmore 


Paractiornis WrEtMorE, Condor, vol. 32, No. 3, May 15, 1930, p. 133. Type, 
by monotypy, Paractiornis perpusillus Wetmore. 


Paractiornis perpusillus WETMORE 


Paractiornis perpusillus WETMorE, Condor, vol. 32, No. 3, May 15, 1930, p. 153, 
figs. 54-56. 


Lower Miocene (Harrison formation) : Carnegie Hill, Agate Fossil 
Quarry, near Agate, Sioux County, Nebraska. 


85 Cooke, Florida Geol. Surv., Geol. Bull. 29, 1945, pp. 119-120, believes that 
the specimen came from a sink in the Tampa limestone, rather than from the 
younger Hawthorn formation, to which it was ascribed by T. E. White, who 
collected it. 


7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Genus PALOSTRALEGUS Brodkorb 


Palostralegus Bropxors, Florida Geol. Surv. Rep. Invest. No. 14, November 
1955, p. 19. Type, by original designation, Palostralegus sulcatus Brod- 
korb. 


Palostralegus sulcatus BropKoRB 


Palostralegus sulcatus BropKxors, Florida Geol. Surv. Rep. Invest. No. 14, 
November 1955, p. 20, fig. 18. 


Pliocene (Bone Valley formation): Near Brewster, Polk County, 
Florida. 


Family CHARADRIIDAE: P overs, TuRNSTONEs, and SURFBIRDS 
Subfamily CHARADRIINAE: PLovers 


Genus CHARADRIUS Linnaeus 


Charadrius LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 150. Type, by tau- 
tonymy, Charadrius hiaticula Linnaeus. 


Charadrius sheppardianus Corr 


Charadrius sheppardianus Corr, Bull. Geol. Geogr. Surv. Terr., vol. 6, No. 1, 
Feb. 11, 1881, p. 83. 


Oligocene (Florissant lake beds) : Florissant, Colorado.°® 
Charadrius vociferus LInNAEUs: Killdeer 
Charadrius vociferus LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 150. 


Modern form reported from late Pleistocene: McKittrick, Kern 
County, and Rancho La Brea, Los Angeles, California. 


Genus EUPODA Brandt 


Eupoda J. F. Branpt, in Tchihatchev, Voy. Sci. Altai Orient., 1845, p. 444. 
Type, by monotypy, Charadrius asiaticus Pallas. 


Eupoda montana (TowNsenp): Mountain Plover 


Charadrius montanus J. K. TowNsENp, Journ. Acad. Nat. Sci. Philadelphia, 
vol. 7, pt. 2, Nov. 21, 1837, p. 192. 


Modern form reported from late Pleistocene: McKittrick, Kern 
County, California. 


96 Generic and subfamily allocation tentative, particularly since the Florissant 
beds now are held to be Oligocene rather than Miocene by most paleontologists. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 71 


Genus SQUATAROLA Cuvier 


Squatarola Cuvier, Régne Animal, vol. 1, 1817 (Dec. 7, 1816), p. 467. Type, 
by tautonymy, 7ringa squatarola Linnaeus. 


Squatarola squatarola (LinNAEUS): Black-bellied Plover 


Tringa Squatarola Linnagus, Syst. Nat., ed. 10, vol. 1, 1758, p. 149. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 


Genus LIMICOLAVIS Shufeldt °7 


Limicolavis SHurFELpT, Trans. Connecticut Acad. Arts Sci., vol. 19, February 
1915, p. 55. Type, by monotypy, Limicolavis pluvianella Shufeldt. 


Limicolavis pluvianella SHUFELDT 


Limicolavis pluvianella Suu¥Fetpt, Trans. Connecticut Acad. Arts Sci., vol. 
19, February 1915, p. 55, pl. 15, fig. 129. 


? Oligocene: Lower Willow Creek, Oregon. 


Family SCOLOPACIDAE: Woopcock, SNiIpEs, and SANDPIPERS 
Subfamily PALAEOTRINGINAE: PaLagorrincas 
Genus PALAEOTRINGA Marsh 


Palaeotringa Marsu, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, p. 208. 
Type, by subsequent designation, Palaeotringa littoralis Marsh (Hay, 
1902). 

Palaeotringa littoralis Marsn % 

Palaeotringa littoralis Marsu, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, 
p. 208. 

Paleocene (Hornerstown marl): Hornerstown, New Jersey. 


Palaeotringa vagans MarsH 


Palaecotringa vagans Marsu, Amer. Journ. Sci., ser. 3, vol. 3, May 1872, p. 365. 
Paleocene (Hornerstown marl): Hornerstown, New Jersey. 


Palaeotringa vetus MArsH 
Palaeotringa vetus Marsu, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, 
p. 209. 
Paleocene (Hornerstown marl): Arneytown, New Jersey. 
97 Family relationship uncertain, 


98 Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, February 1915, pp. 23, 
77, pl. 6, fig. 35, believes this to be a gull, but this is open to question. 


72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Subfamily SCOLOPACINAE: Woopcock and SNIPES 
Genus CAPELLA Frenzel 


Capella FreNzeEL, Beschr. Vogel und Eyer Wittenberg, 1801, p. 58. Type, by 
monotypy, Scolopax coelestis Frenzel = Scolopax gallinago Linnaeus. 


Capella gallinago (LINNAEUS): Common Snipe 99 
Scolopax Gallinago LinnaEvs, Syst. Nat., ed. 10, vol. 1, 1758, p. 147. 
Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 
Capella anthonyi (WeETMoRE) 
Gallinago anthonyi WEtTMoreE, Proc. Biol. Soc. Washington, vol. 33, Dec. 30, 


1920, p. 78, pl. 2, figs. 1, 2. 


Recent (extinct): Cave deposits in Cueva Catedral (type lo- 
cality) and Cueva Clara, near Morovis, Puerto Rico. 


Subfamily TRINGINAE: CurLrews, YELLOWLEGS, and ALLIES 
Genus NUMENIUS Brisson 


Numenius Brisson, Orn., 1760, vol. 1, p. 48; vol. 5, p. 311. Type, by tau- 
tonymy, Numenius Brisson = Scolopax arquata Linnaeus. 
Numenius americanus BecusteIn: Long-billed Curlew 
Nwumenius americanus BECHSTEIN, in Latham, Allgem. Uebers. Vogel, vol. 4, 
Dt 2. TO1s p;.A32: 


Modern form reported from late Pleistocene: McKittrick, Kern 
County, and Rancho La Brea, Los Angeles, California. 


Numenius borealis (Forster): Eskimo Curlew 


Scolopax borealis J. R. Forster, Philos. Trans., vol. 62, 1772, p. 431. 


Modern form reported from late Pleistocene (Kentuck locality) : 
McPherson County, Kansas. 


Numenius phaeopus (LINNAEUS): Whimbrel 2 


Scolopax Phaeopus LINNAEuS, Syst. Nat., ed. 10, vol. 1, 1758, p. 146. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 


99 Capella delicata (Ord), Wilson’s snipe, of the previous list. 

1Included here as it has not been found in living form, being known only 
from bones. 

2 Phaeopus hudsonicus (Latham), Hudsonian curlew of the previous list. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 73 


Genus PALNUMENIUS Miller 


Palnumenius L. Mixer, Univ. California Publ. Zool., vol. 43, Mar. 6, 1942, 
p. 45. Type, by monotypy, Palnumenius victima Miller. 


Palnumenius victima MILLER 


Palnumenius victima L. M1ILter, Univ. California Publ. Zoél., vol. 43, Mar. 6, 
1942, p. 45, fig. Ib. 


Pleistocene: San Josecito Cave, Aramberri, Nuevo Leon. 


Genus BARTRAMIA Lesson 


Bartramia Lesson, Traité d’Orn., livr. 7, Apr. 9, 1831, p. 553. Type, by 
monotypy, Bartramia laticauda Lesson = Tringa longicauda Bechstein. 


Bartramia longicauda (BEcHSTEIN): Upland Plover 


Tringa longicauda BECHSTEIN, in Latham, Allgem. Uebers. Vogel, vol. 4, 
pt. 2, 1812, p. 453. 


Modern form reported from late Pleistocene: Meade County (Jones 
fauna, Vanhem formation), and McPherson County (Kentuck lo- 
cality ), Kansas. 


Genus TOTANUS Bechstein 


Totanus BEecHsTEIN, Orn. Taschenb. Deutschland, vol. 2, 1803, p. 282. Type, 
by tautonymy, Totanus maculatus Bechstein = Scolopax totanus Linnaeus. 


Totanus melanoleucus (GMELIN): Greater Yellowlegs 


Scolopax melanoleuca GMELIN, Syst. Nat., vol. 1, pt. 2, 1780, p. 659. 


Modern form reported from Pleistocene: Fossil Lake, Oregon; 


Rancho La Brea, Los Angeles, and McKittrick, Kern County, Cali- 
fornia. 


Subfamily CALIDRIINAE: Sanppipers, Gopwits, and ALLIEs 
Genus CALIDRIS Merrem 


Calidris pacis Mrrrem, Lit. Zeitung, vol. 2, No. 168, June 8, 1804, col. 542. 
Type, by tautonymy, Tringa calidris Gmelin = Tringa canutus Linnaeus. 


Calidris pacis Bropkors 


Calidris pacis BropKors, Florida Geol. Surv. Rep. Invest. No. 14, November 
1955, p. 22, figs. 19, 20. 


Pliocene (Bone Valley formation): Near Brewster, Polk County, 
Florida. 


74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Genus EROLIA Vieillot 


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


Erolia penepusilla BropKorp 


Erolia penepusilla BropKors, Florida Geol. Surv. Rep. Invest. No. 14, No- 
vember 1955, p. 23, fig. 21. 


Pliocene (Bone Valley formation): Near Brewster, Polk County, 
Florida. 


Erolia alpina (LINNAEUS): Dunlin 
Tringa alpina LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 149. 


Modern form reported from late Pleistocene: McKittrick, Kern 
County, California. 


Genus LIMNODROMUS Wied 


Limnodromus Wien, Beitr. Naturg. Brasil, vol. 4, Abt. 2, 1833, p. 716. Type, 
by monotypy, Scolopax noveboracensis Gmelin = Scolopax grisea Gmelin. 


Limnodromus griseus (GMELIN): Dowitcher 


Scolopax grisea GMELIN, Syst. Nat., vol. 1, pt. 2, 1789, p. 658. 


Modern form reported late Pleistocene: McKittrick, Kern County, 
and Rancho La Brea, Los Angeles, California. 


Genus MICROPALAMA Baird 


Micropalama Barry, Rep. Expl. and Surv. R. R. Pac., vol. 9, 1858, pp. xxii, 
xlvii, 714, 726. Type, by monotypy, Tringa himantopus Bonaparte. 


Micropalama hesternus WETMORE 


Micropalama hesternus Wetmore, Proc. U. S. Nat. Mus., vol. 64, art. 5, Jan. 
15, 1924, p. 11, figs. 6-7. 


Upper Pliocene (Blancan) : 2 miles south of Benson, Arizona. 


Genus LIMOSA Brisson 


Limosa Brisson, Orn., 1760, vol. 1, p. 48; vol. 5, p. 261. Type, by tautonymy, 
Limosa Brisson = Scolopax limosa Linnaeus. 


Limosa vanrossemi MILLER 
Limosa vanrossemi L. H. Mutter, Carnegie Inst. Washington Publ. 349, 
August 1925, p. 116, pl. 6. 


Middle Miocene (Temblor, Turritella ocoyana zone): Lompoc, 
California. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 75 


Family RECURVIROSTRIDAE: Isis-BiLts, Avocets, and STILTs 
Subfamily RECURVIROSTRINAE: Avocets and Sritts 
Genus RECURVIROSTRA Linnaeus 


Recurvirostra LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 151. Type, by 
monotypy, Recurvirostra avosetta Linnaeus. 


Recurvirostra americana GMELIN: Avocet 


Recurvirostra americana GMELIN, Syst. Nat., vol. 1, pt. 2, 1789, p. 603. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon ; 
Rancho La Brea, Los Angeles, and McKittrick, Kern County, Cali- 
fornia. 


Genus HIMANTOPUS Brisson 


Himantopus Brisson, Orn., 1760, vol. 1, p. 46; vol. 5, p. 33. Type, by tau- 
tonymy, Himantopus Brisson = Charadrius himantopus Linnaeus. 


Himantopus mexicanus (MULLER): Black-necked Stilt 


Charadrius Mexicanus P. L. S. MUtier, Natursyst., Suppl., 1776, p. 117. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon. 


Family PRESBYORNITHIDAE: PresByorniTHES 
Genus PRESBYORNIS Wetmore 


Presbyornis WertTMore, Ann. Carnegie Mus., vol. 16, Apr. 10, 1926, p. 396. 
Type, by monotypy, Presbyornis pervetus Wetmore. 


Presbyornis pervetus WETMORE 


Presbyornis pervetus Wetmore, Ann. Carnegie Mus., vol. 16, Apr. 10, 1926, 
p. 396, pl. 37, figs. 10-20. 


Eocene (Lower Green River formation): White River, Utah, 2 
miles from Colorado State line. 


Family PHALAROPODIDAE: PHALAROPES 
Genus LOBIPES Cuvier 


Lobipes Cuvier, Régne Animal, vol. 1, 1817 (Dec. 7, 1816), p. 495. Type, by 
original designation, Tringa hyperborea Linnaeus = Tringa lobata 
Linnaeus. 


Lobipes lobatus (L1inNAEUS) : Northern Phalarope 


Tringa lobata LInNAEus, Syst. Nat., ed. 10, vol. 1, 1758, p. 148. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon. 


76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Suborder LARI: Sxkuas, GuLLs, TERNS, and SKIMMERS 
Family STERCORARIIDAE: Jarcers and Skuas 
Genus STERCORARIUS Brisson 


Stercorarius Brisson, Orn., 1760, vol. 1, p. 56; vol. 6, p. 149. Type, by tau- 
tonymy, Stercorarius Brisson= Larus parasiticus Linnaeus. 


Stercorarius shufeldti HowArp 


Stercorarius shufeldti H. Howarp, Carnegie Inst. Washington Publ. 551, 
Jan. 25, 1946, p. 184, pl. 2, figs. 1, 2. 


Late Pleistocene: Fossil Lake, Oregon.® 


Family LARIDAE: Gutts and TERNs 
Subfamily LARINAE: GuLts 
Genus LARUS Linnaeus 4 


Larus LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 136. Type, by subsequent 
designation, Larus marinus Linnaeus (Selby, 1840). 
Larus glaucescens NAUMANN: Glaucous-winged Gull 


Larus glaucescens NAUMANN, Naturg. Vogel Deutschl., vol. 10, 1840, p. 351. 


Modern form reported from late Pleistocene (Palos Verdes for- 
mation) : San Pedro, Los Angeles County, California. 


Larus californicus LAWRENCE: California Gull 
Larus Californicus LAwrENcE, Ann. Lyc. Nat. Hist. New York, vol. 6, 1854, 
p. 79. 
Modern form reported from late Pleistocene: Fossil Lake, Oregon. 


Larus philadelphia (Orn): Bonaparte’s Gull 
Sterna Philadelphia Orv, in Guthrie, Geogr., 2d Amer. ed., 1815, p. 310. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon.* 


Larus oregonus SHUFELDT 


Larus oregonus SuureLpt, Amer. Nat., vol. 25, No. 297, September 1891, 
p. 820. 


Late Pleistocene: Fossil Lake, Oregon. 


8 The type of Stercorarius shufeldti originally was identified by Shufeldt as 
Larus argentatus, this specimen being the basis for the record of the herring 
gull from Fossil Lake. 

4 Larus vero Shufeldt, Journ. Geol., January-February 1917, p. 18, has been 
identified by Wetmore as Nyctanassa violacca Linnaeus (Smithsonian Misc. Coll., 
vol. 85, No. 2, Apri 13, 1931, p. 16). 

5 Records of Xema sabini from Fossil Lake, so far as identified, refer to Larus 
philadelphia. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 


77 


Larus pristinus SHUFELDT ® 


Larus pristinus SHUFELDT, Trans. Connecticut Acad. Arts Sci., vol. 19, Febru- 
ary I9Q15, p. 54, pl. 14, fig. 112. 


? Oligocene (John Day) : Willow Creek, Oregon. 


Larus robustus SHUFELDT 


Larus robustus Suuretpt, Amer. Nat., vol. 25, No. 297, September 1891, 
p. 819. 


Late Pleistocene: Fossil Lake, Oregon. 
Larus elmorei BropKors 


Larus elmoret BropKors, Wilson Bull., vol. 65, No. 2, June 30, 1953, p. 94, 
fig. I. 


Pliocene (Bone Valley formation) : Near Brewster, Polk County, 
Florida. 
Genus GAVIOTA Miller and Sibley 7 


Gaviota A. H. Mitier and C. G. Sistey, Auk, vol. 58, No. 4, October 1941, 
p. 563. Type, by monotypy, Gaviota niobrara Miller and Sibley. 


Gaviota niobrara MILLER and SIBLEY 


Gaviota niobrara A. H. Mitter and C. G. Sistey, Auk, vol. 58, No. 4, Octo- 
ber 1941, p. 563, fig. 1. 


Late Upper Miocene (Barstovian, Niobrara River zone) : Niobrara 
Game Preserve, Cherry County, Nebraska. 


Subfamily STERNINAE: Terns 
Genus STERNA Linnaeus 


Sterna LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 137. Type, by tautonymy, 
Sterna hirundo Linnaeus. 


Subgenus STERNA Linnaeus 
Sterna forsteri NutTraL_: Forster’s Tern 


Sterna forsteri Nutratt, Manual Orn. U. S. and Canada, vol. 2, 1834, p. 274. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon. 


Genus CHLIDONIAS Rafinesque 


Chlidonias RAFINESQUE, Kentucky Gazette, n. s., vol. 1, No. 8, Feb. 21, 1822, 
p. 3, col. 5. Type, by monotypy, Sterna melanops Rafinesque = Sterna 
surinamensis Gmelin. 


® Generic assignation in original description tentative. 
7 Allocation to subfamily provisional. 


78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Chlidonias niger (LINNAEUS): Black Tern 


Sterna nigra LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 137. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon, 


Suborder ALCAE: Avuxs 
Family ALCIDAE: Auxs, Murres, and PUuFFINS 
Subfamily NAUTILORNITHINAE: NavtTILorniTHEs 
Genus NAUTILORNIS Wetmore 


Nautilornis Wetmore, Ann. Carnegie Mus., vol. 16, Apr. 10, 1926, p. 392. 
Type, by original designation, Nautilornis avus Wetmore. 


Nautilornis avus WETMORE 
Nautilornis avus WEtTMorE, Ann. Carnegie Mus., vol. 16, Apr. 10, 1926, p. 392, 
pl. 36, figs. 1-8. 


Eocene (Lower Green River formation): White River, Utah, 2 
miles from Colorado State line. 


Nautilornis proavitus WETMORE 


Nautilornis proavitus WrTMorE, Ann. Carnegie Mus., vol. 16, Apr. 10, 1926, 
Pp. 304, pl. 36, fig. 9. 


Eocene (Lower Green River formation): White River, Utah, 2 
miles from Colorado State line. 


Genus HYDROTHERIKORNIS Miller 


Hydrotherikornis A. H. Mitrer, Univ. California Publ., Bull. Dept. Geol. Sci., 
vol. 20, No. 3, Apr. 21, 1931, p. 24. Type, by monotypy, Hydrotherikornis 
oregonus Miller. 


Hydrotherikornis oregonus MILLER 


Hydrotherikornis oregonus A. H. Miter, Univ. California Publ., Bull. Dept. 
Geol. Sci., vol. 20, No. 3, Apr. 21, 1931, p. 24, fig. 1. 


Upper Eocene (Arago series) : Sunset Bay, near Coos Bay, Coos 
County, Oregon. 


Subfamily ALCINAE: Auxs and MurreEs 
Genus AUSTRALCA Brodkorb 


Australca Bropxors, Florida Geol. Surv. Rep. Invest. No. 14, November 1955, 
p. 25. Type, by original designation, Australca grandis Brodkorb. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 79 


Australca grandis BropkorsB 


Australca grandis Bropxors, Florida Geol. Surv. Rep. Invest. No. 14, No- 
vember 1955, p. 27, figs. 24, 29. 


Pliocene (Bone Valley formation): Near Brewster, Polk County, 
Florida. 


Genus URIA Brisson 


Uria Brisson, Orn., 1760, vol. 1, p. 52; vol. 6, p. 70. Type, by tautonymy, 
Uria Brisson = Colymbus aalge Pontoppidan. 


Uria aalge (PoNnTopPIpAN) : Common Murre 


Colymbus aalge PontTorripANn, Danske Atlas, vol. 1, 1763, p. 621, pl. 26. 


Modern form reported from late Pleistocene (Palos Verdes sand) : 
Playa del Rey, and Mussel Rock, San Mateo County, California. 


Uria affinis (MarsH) 


Catarractes affinis MarsH, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, 
p. 259. 


Pleistocene: Railroad cut on bank of Penobscot River, near Bangor, 
Maine. 


Uria antiqua (MarsH) 


Catarractes antiquus MarsH, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, 
p. 213. 


Miocene: Tarboro, Edgecombe County, North Carolina. 


Genus MIOCEPPHUS Wetmore 


Miocepphus Wetmore, Journ. Morph., vol. 66, Jan. 2, 1940, p. 35. Type, by 
monotypy, Miocepphus mcclungi Wetmore. 


Miocepphus mcclungi WETMORE 


Miocepphus mcclungi Wetmore, Journ. Morph., vol. 66, Jan. 2, 1940, p. 35, 
figs. II-I4. 


Miocene (Calvert formation, zone 12): Near the mouth of Parker 
Creek, Calvert County, Maryland.® 


Genus BRACHYRAMPHUS Brandt 


Brachyramphus M. Branot, Bull. Sci. Acad. Imp. Sci. St.-Pétersbourg, vol. 2, 
No. 22, Mar. 10, 1837, col. 346. Type, by subsequent designation, Colymbus 
marmoratus Gmelin (Gray, 1840). 


8 Two records. 


80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Brachyramphus pliocenum HowArp 


Brachyramphus pliocenus Howarp, Carnegie Inst. Washington Publ. 584, 
June 22, 1949, p. 191. 


Middle Pliocene (San Diego formation) : Washington Boulevard 
Freeway, San Diego, California. 


Genus SYNTHLIBORAMPHUS Brandt 


Synthliboramphus M. Branot, Bull. Sci. Acad. Imp. Sci. St.-Pétersbourg, 
vol. 2, No. 22, Mar. 19, 1837, col. 347. Type, by subsequent designation, 
Alca antiqua Gmelin (Gray, 1840). 


Synthliboramphus antiquum (GMELIN): Ancient Murrelet 


Alca antiqua GMELIN, Syst. Nat., vol. 1, pt. 2, 1780, p. 554. 


Modern form reported from late Pleistocene (Palos Verdes sand) : 
San Pedro, California. 


Genus PTYCHORAMPHUS Brandt 


Ptychoramphus M. Branot, Bull. Sci. Acad. Imp. Sci. St.-Pétersbourg, vol. 2, 
No. 22, Mar. 19, 1837, col. 347. Type, by monotypy, Uria aleutica Pallas. 


Ptychoramphus aleuticum (PALLAs): Cassin’s Auklet 


Uria Aleutica PALLAS, Zoogr. Rosso-Asiatica, vol. 2, 1811, p. 370. 


Modern form reported from late Pleistocene (Palos Verdes sand) : 
San Pedro, Los Angeles County, California. 


Genus CERORHINCA Bonaparte 


Cerorhinca Bonaparte, Ann. Lyc. Nat. Hist. New York, vol. 2, 1828, p. 427. 
Type, by monotypy, Cerorhinca occidentalis Bonaparte = Alca monocerata 
Pallas. 


Cerorhinca dubia MILLER 


Cerorhinca dubia L, H. Mitter, Carnegie Inst. Washington Publ. 349, August 
1925, p. 115, pl. 2. 


Middle Miocene (Temblor, Turritella ocoyana zone): Lompoc, 
California. 


Family MANCALLIDAE: Lucas Auk and ALLy 
Genus MANCALLA Lucas 


Mancalla Lucas, Science, n.s., vol. 13, Mar. 15, 1901, p. 428. Type, by original 
designation, Mancalla californiensis Lucas. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 81 


Mancalla californiensis Lucas 


Mancealla californiensis Lucas, Science, n.s., vol. 13, Mar. 15, 1901, p. 428.9 


Pliocene: Third Street Tunnel, Los Angeles (type locality), and 
Newport Bay. Middle Pliocene (San Diego formation) : San Diego, 
San Diego County, and Corona del Mar, Orange County, California. 


Mancalla diegense (MILLER) 


Pliolunda diegense L. H. Mitrer, Trans. San Diego Soc. Nat. Hist., vol. 8, 
Dec. 15, 1937, Pp. 376, 2 figs. 


Middle Pliocene (San Diego formation): Market Street, near 
Euclid Avenue (type locality), and Mission Hills district, San Diego, 
California. 


Order COLUMBIFORMES: SAnp-GRousE, PIGEONS, AND Doves 
Suborder COLUMBAE: Piceons and Doves 
Family COLUMBIDAE: Picrons and Doves 


Subfamily COLUMBINAE: Piceons and Doves 
Genus COLUMBA Linnaeus 


Columba Linnaevs, Syst. Nat., ed. 10, vol. 1, 1758, p. 162. Type, by subsequent 
designation, Columba oenas Linnaeus (Vigors, 1825). 


Columba fasciata Say: Band-tailed Pigeon 
Columba fasciata Say, in Long, Exped. Rocky Mountains, vol. 2, 1823, p. 10. 
Modern form reported from late Pleistocene: Stone Man Cave, 
Shasta County, Rancho La Brea, Los Angeles, and Carpinteria, Santa 
Barbara County, California. Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. 
Columba micula (WETMORE) 


Chlorenas micula Wetmore, Proc. U. S. Nat. Mus., vol. 64, art. 5, Jan. 15, 
1924, p. 13, figs. 8-9. 


Early Pleistocene: Curtis Ranch, 12 miles southeast of Benson, 
Arizona. 
Genus ZENAIDURA Bonaparte 


Zenaidura BONAPARTE, Compt. Rend. Acad. Sci. Paris, vol. 40, January 1855, 
p. 96. Type, by original designation, Columba carolinensis Linnaeus. 


® See also Lucas, Proc. U. S. Nat. Mus., vol. 24, Sept. 27, 1901, pp. 133-134, 
figs. I, 2. 


82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Zenaidura macroura (LINNAEUS): Mourning Dove 

Columba macroura LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 164. 

Modern form reported from Upper Pliocene (Rexroad fauna) : 
Meade County, Kansas. Pleistocene: San Josecito Cave, Aramberri, 
Nuevo Leon; Seminole Field, Pinellas County, Florida. Late Pleisto- 
cene: Carpinteria, Santa Barbara County, McKittrick, Kern County, 
and Rancho La Brea, Los Angeles, California; Meade County, Kansas 
(Vanhem formation, Jones fauna). 


Genus ECTOPISTES Swainson 


Ectopistes SwAInson, Zool. Journ., vol. 3, No. 11, September-December 1827, 
p. 362. Type, by subsequent designation, Columba migratoria Linnaeus 
(Swainson, 1837). 


Ectopistes migratorius (LINNAEUS): Passenger Pigeon 


Columba migratoria LINNAEvS, Syst. Nat., ed. 12, vol. 1, 1766, p. 285. 


Modern form reported from Pleistocene: Cave deposits of Tennes- 
see. Late Pleistocene: Rancho La Brea, Los Angeles, California. 


Genus GEOTRYGON Gosse 


Geotrygon Gossk, Birds Jamaica, 1847, p. 316. Type, by subsequent designa- 
tion, Columba cristata Latham = Geotrygon sylvatica Gosse = Columbi- 
gallina versicolor Lafresnaye (Reichenbach, 1852 = 1853). 


Geotrygon larva (WETMORE) 


Oreopeleia larva WETMORE, Proc. Biol. Soc. Washington, vol. 33, Dec. 30, 1920, | 
p. 79, pl. 3, figs. 1-2. 


Recent (extinct) :?° Cave deposits in Cueva Clara (type locality) 
and Cueva Catedral, near Morovis; Cueva Torafio, near Utuado; 
kitchen middens near Mayagiiez, and at Barrio Canfas, near Ponce, 
Puerto Rico. 


Order PSITTACIFORMES: Lortes, PArrots, PARAKEETS, and MAcAaws 
Family PSITTACIDAE: Lortrs, Parrots, and Macaws 
Subfamily PSITTACINAE: Parakeets and Macaws 


Genus ARA Lacépéde 


Ara Lactépipe, Tableaux Ois., 1799, p. 1, Type, by subsequent designation, 
Psittacus macao Linnaeus (Ridgway, 1916). 


10 Included here as it has not been found in living form, being known only 
from bones. 





NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 83 


Ara tricolor BecHSTEIN: Cuban Macaw 


Ara tricolor BECHSTEIN, in Latham, Allg. Uebers. V6g., vol. 4, Th. 1, 1811, 
p. 64, pl. 1. (Cuba.) 


Modern form recorded from late Pleistocene: Banos de Ciego 
Montero, Santa Clara Province, Cuba. 


Ara autocthones WETMORE 


Ara autocthones WetTMorE, Journ. Agr. Univ. Puerto Rico, vol. 21, No. 1, 
January 1937, p. 12, pl. 1, figs. 8, 9. 


Recent (extinct): 1 Prehistoric kitchen midden deposits at Con- 
cordia, near Southwest Cape, St. Croix, Virgin Islands. 


Genus RHYNCHOPSITTA Bonaparte 


Rhynchopsitta Bonaparte, Rev. et Mag. Zool., ser. 2, vol. 6, March 1854, 
p. 149. Type, by monotypy, Macrocercus pachyrhynchus Swainson. 


Rhynchopsitta pachyrhyncha (Swartnson): Thick-billed Parrot 


Macrocercus pachyrhynchus Swainson, Philos. Mag., n.s., vol. 1, No. 6, June 
1827, p. 439. 


Modern form reported from Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. 
Genus CONUROPSIS Salvadori 


Conuropsis SALvAporI, Cat. Birds Brit. Mus., vol. 20, 1891, pp. 146, 203. Type, 
by original designation, Psittacus carolinensis Linnaeus. 


Conuropsis fratercula WETMORE 


Conuropsis fratercula WrETMorE, Amer. Mus. Nov., No. 211, Mar. 11, 1926, 
p. 3, figs. 5-6. 


Middle Miocene (Merychippus primus zone, lower Sheep Creek 
beds) : Snake Creek Quarries, Sioux County, Nebraska. 


Order CUCULIFORMES: Prantarn-gaters and Cuckoos 
Suborder CUCULI: Cuckoos, RoApRUNNERs, and ANIS 
Family CUCULIDAE: Cucxoos, RoapRuNNERs, and ANIs 
Subfamily NEOMORPHINAE: Grounp Cuckoos 
Genus GEOCOCCYX Wagler 


Geococcyx WaGLER, Isis von Oken, vol. 24, Heft 5, May 1831, col. 524. Type, 
by monotypy, Geococcyx variegata Wagler = Saurothera californiana 
Lesson. 





11 Included here since it has not been found in living form, being known only 
from bones. 


84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Geococcyx californianus (Lesson): Roadrunner 
Saurothera californiana Lesson, Compl. Oeuvres Buffon, vol. 6, 1829, p. 420. 
Modern form reported from late Pleistocene: Rancho La Brea, 


Los Angeles, McKittrick, Kern County, and Carpinteria, Santa 
Barbara County, California. 


Geococcyx conklingi Howarp 
Geococcyx conklingi Howarp, Condor, vol. 33, No. 5, Sept. 15, 1931, p. 208, 
figs. 49-50. 


Pleistocene: Conkling Cavern (type locality), and Shelter Cave,?? 
Pyramid Peak, Organ Mountains, Dona Ana County, New Mexico; 
San Josecito Cave, Aramberri, Nuevo Leon. 


Order STRIGIFORMES: Owts 13 
Family PROTOSTRIGIDAE: Protostrrx 
Genus PROTOSTRIX Wetmore 


Protostrix WrtTmore, Amer. Mus. Nov., No. 680, Dec. 4, 1933, p. 3. Type, 
by original designation, Aquila lydekkeri Shufeldt. 


Protostrix lydekkeri (SHUFELDT) 
Aquila lydekkeri Suuretpt, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 16, 
Aug. 4, 1913, p. 208. 
Eocene (Bridger formation) : Lower Cottonwood Creek, Wyoming. 


Protostrix saurodosis (WETMORE) 
Minerva saurodosis Wetmore, Proc. Acad. Nat. Sci. Philadelphia, vol. 73, 1921 
(Apr. 6, 1922), p. 455, figs. 1-2. 


Eocene (Bridger formation) : Near Lodgepole Trail Crossing on 
Dry Creek, about 10 miles from Fort Bridger, Wyoming. 


Protostrix leptosteus (MarsH) 14 


Bubo leptosteus Marsu, Amer. Journ. Sci., ser. 3, vol. 2, August 1871, p. 126. 


Eocene (Bridger formation): Grizzly Buttes, near Fort Bridger, 
Wyoming. 


12 Possibly of Recent period. 

18 Aquila antiqua Shufeldt, type of the genus Minerva Shufeldt, formerly con- 
sidered an owl, proves to be a mammal. See Wetmore, Amer. Mus. Nov., No. 
680, Dec. 4, 1933, Pp. I, 2. 

14 See Wetmore, Condor, 1937, pp. 84-85. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 85 


Protostrix mimica WETMORE 
Protostrix mimica WeEtTMorE, Proc. U. S. Nat. Mus., vol. 85, Jan. 17, 1938, 
p. 27, figs. 4-5 


Lower Eocene (Wasatch) : South side of Ten Mile Creek, 12 miles 
northwest of Worland, Wyoming. 


Family TYTONIDAE: Barn Owts 
Subfamily TYTONINAE: Barn Owts 
Genus TYTO Billberg 


Tyto Bitiperc, Syn. Faunae Scand., vol. 1, pt. 2, 1828, tab. A. Type, by 
monotypy, Strix flammea auct. = Strix alba Scopoli. 


Tyto alba (Scopotr): Barn Owl 


Strix alba Scopott, Annus 1, Historico-Naturalis, 17609, p. 21. 


Modern form reported from Pleistocene: Cavern deposits near 
Lecanto, Florida; 1° San Josecito Cave, Aramberri, Nuevo Leon. Late 
Pleistocene: Carpinteria, Santa Barbara County and Rancho La Brea, 
Los Angeles, California. 


Tyto cavatica WETMORE 
Tyto cavatica WeETMoRE, Proc. Biol. Soc. Washington, vol. 33, Dec. 30, 1920, 


p. 80, pl. 3, figs. 3-6. 


Recent (extinct) : 1° Cave deposits in Cueva Torafio, near Utuado, 
Puerto Rico. 


Tyto ostologa WETMORE 
Tyto ostologa WETMORE, Smithsonian Misc. Coll., vol. 74, No. 4, Oct. 17, 1922, 
p. 2. 


Recent (extinct) :?° Cave deposits in Grotte San Francisco near 
St. Michel (type locality), and caves near L’Atalye, Haiti. 


Tyto pollens WETMORE 


Tyto pollens WEetTMorE, Bull. Mus. Comp. Zodl., vol. 80, No. 12, October 1937, 
p. 436, figs. 10-16. 


Recent (extinct) :1° Cave deposits on Great Exuma Island, Ba- 
hama Islands. 


1° The record from Vero (stratum 3) is now considered to be of Recent age. 
See Cooke, Florida Geol. Surv., Geol. Bull. 29, 1945, pp. 306-307. 

16 Included here as it has not been found in living form, being known only from 
bones. 


86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Family STRIGIDAE: TypicaL OwLs 
Genus OTUS Pennant 


Otus PENNANT, Indian Zool., 1760, p. 3. Type, by monotypy, Otus bakkamoena 
Pennant. 


Otus asio (LINNAEUS) : Screech Owl 

Strix asio LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 92. 

Modern form reported from Pleistocene: Cavern deposits near 
Lecanto, Florida; cave deposits of Tennessee; San Josecito Cave, 
Aramberri, Nuevo Leon. Late Pleistocene: Potter Creek Cave, Shasta 
County, Carpinteria, Santa Barbara County, and Rancho La Brea, 
Los Angeles, California. 

Otus flammeolus (KAup) : Flammulated Owl 


Scops (Megascops) flammeola Kavp, in Jardine, Contr. Orn., 1852 (1853), 
D. EIT. 


Modern form reported from Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. Late Pleistocene: Samwel Cave,’7 Shasta County, 
California. 


Otus trichopsis (WAGLER): Whiskered Owl 
Scops trichopsis WAGLER, Isis von Oken, Heft 3, March 1832, col. 276. 
Modern form reported from Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. 
Genus BUBO Duméril 


Bubo DumériL, Zool. Analytique, 1806, p. 34. Type, by tautonymy, Strix 
bubo Linnaeus. 


Bubo virginianus (GMELIN) : Horned Owl 

Strix virginiana GMELIN, Syst. Nat., vol. 1, pt. 1, 1788, p. 287. 

Modern form reported from late Pleistocene: Fossil Lake, Oregon ; 
Samwel Cave, Shasta County, Carpinteria, Santa Barbara County, 
McKittrick, Kern County, and Rancho La Brea, Los Angeles, Cali- 
fornia. Pleistocene: San Josecito Cave, Aramberri, Nuevo Leon. 
Bubo sinclairi MILLER 


Bubo sinclairi L. H. Mitter, Univ. California Publ., Bull. Dept. Geol., vol. 6, 
No. 16, Oct. 28, 1911, p. 393, figs. 4-5. 


Late Pleistocene: Samwel and Potter Creek (type locality) caves, 
Shasta County, California. 


17 Recorded originally as Micropallas whitneyi. See Miller, L. H., Trans. San 
Diego Soc. Nat. Hist., vol. 7, No. 19, Mar. 31, 1933, pp. 200-210. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 87 


Genus GLAUCIDIUM Boie 


Glaucidium Borg, Isis von Oken, Bd. 2, 1826, col. 970. Type, by subsequent 
designation, Strix passerina Linnaeus (Gray, 1840). 


Glaucidium gnoma WaAGLER: Pygmy Owl 
Glaucidium Gnoma WAGLER, Isis von Oken, vol. 25, Heft 3, March 1832, p. 275. 


Modern form reported from late Pleistocene: Samwel Cave, Shasta 
County, Carpinteria, Santa Barbara County, and Rancho La Brea, 
Los Angeles, California. 


Genus SPEOTYTO Gloger 


Speotyto Giocer, Hand- und Hilfsbuch Naturg., 1842 (1841), p. 226. Type, 
by monotypy, Strix cunicularia Molina. 


Speotyto cunicularia (Morina): Burrowing Owl 


Strix Cunicularia Mottna, Sagg. Stor. Nat. Chili, 1782, p. 263. 


Modern form reported from late Pleistocene: McKittrick, Kern 
County, and Rancho La Brea, Los Angeles, California. 


Genus CICCABA Wagler 


Ciccaba WAGLER, Isis von Oken, Heft 11, 1832, col. 1222. Type, by monotypy, 
Ciccaba huhula = Strix huhula Daudin. 


Ciccaba virgata (CaAssIN) : Mottled Owl 
Syrnium virgatum Cassin, Proc. Acad. Nat. Sci. Philadelphia, vol. 4, 1848 
(1850), p. 124. 
Modern form reported from Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. 
Genus STRIX Linnaeus 


Strix Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 92. Type, by tautonymy, 
Strix stridula Linnaeus = Strix aluco Linnaeus. 


Strix varia Barton: Barred Owl 

Strix varius Barton, Fragm. Nat. Hist. Pennsylvania, 1799, p. 11. 

Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Melbourne, and cavern deposits near Lecanto, Florida. 
Strix occidentalis (XANTUs): Spotted Owl 


Syrnium occidentale XAntus, Proc. Acad. Nat. Sci. Philadelphia, 1859 (Jan. 
10, 1860), p. 193. 


Modern form reported from Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. 


88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Strix brea Howarp 
Strix brea Howarp, Condor, vol. 35, No. 2, Mar. 15, 1933, p. 66, fig. 15. 
Late Pleistocene: Rancho La Brea, Los Angeles, California. 


Strix dakota MILLER 


Strix dakota A. H. Mitter, Univ. California Publ., Bull. Dept. Geol. Sci., 
vol. 27, No. 4, June 22, 1944, p. 95, fig. 8. 


Lower Miocene (Rosebud formation): Flint Hill, 9 miles west- 
southwest of Martin, Bennett County, South Dakota. 


Genus ASIO Brisson 


Asio Brisson, Orn., 1760, vol. 1, p. 28. Type, by tautonymy, Asio Brisson = 
Strix otus Linnaeus. 


Asio otus (LINNAEUS) : 18 Long-eared Owl 

Strix Otus LinnakEus, Syst. Nat., ed. 10, vol. 1, 1758, p. 92. 

Modern form reported from late Pleistocene: Samwel Cave, Shasta 
County, McKittrick, Kern County, and Carpinteria, Santa Barbara 
County, California.4® Pleistocene: San Josecito Cave, Aramberri, 
Nuevo Leon. 

Asio flammeus (PonTopripAN): Short-eared Owl 
Strix flammea Pontopripan, Danske Atlas, vol. 1, 1763, p. 617, pl. 25. 


Modern form reported from late Pleistocene: Rancho La Brea, 
Los Angeles, California. 


Genus AEGOLIUS Kaup 


Aegolius Kaur, Skizz. Entw.-Gesch. Eur. Thierw., 1829, p. 34. Type, by 
monotypy, Strix tengmalmi Gmelin= Strix funereus Linnaeus, 1758. 


Aegolius funereus (LINNAEUS) : Boreal Owl 


Strix funerea LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 93. 


Modern form reported from Pleistocene: *° Shelter cave, Pyramid 
Peak, Organ Mountains, Dona Ana County, New Mexico. 


18 Asio wilsonianus (Lesson) of the preceding list. 

19 According to a communication from L. H. Miller records formerly cited 
from Rancho La Brea are erroneous. 

20 Possibly of Recent age. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 89 


Aegolius acadicus (GMELIN) : Saw-whet Owl 


Strix acadica GMELIN, Syst. Nat., vol. 1, pt. 1, 1788, p. 206. 


Modern form reported from late Pleistocene: Rancho La Brea, 
Los Angeles, California. Pleistocene: San Josecito Cave, Aramberri, 
Nuevo Leon. 


Order CAPRIMULGIFORMES: Orsirps, GoatsucKeErs, and ALLIES 
Suborder CAPRIMULGI: GoatsuckeErs, Potoos, and FroGMouTHS 
Family CAPRIMULGIDAE: GoatsuckErs 
Subfamily CAPRIMULGINAE: GoatsuckeErs 

Genus PHALAENOPTILUS Ridgway 


Phalaenoptilus Ripcway, Proc. U. S. Nat. Mus., vol. 3, 1880, p. 5. Type, by 
original designation, Caprimulgus nuttallii Audubon 


Phalaenoptilus nuttallii (AupuBON) : Poor-will 


Caprimulgus Nuttalli Aupuzon, Birds Amer., octavo ed., vol. 7, 1844, p. 350, 
pl. 495. 


Modern form reported from Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. 


Order PICIFORMES: JacAmArs, BARBETS, TOUCANS, and WoopPECKERS 
Suborder PICI: Woopreckers and WrYNECKS 


Family PICIDAE: Woopprckers, WRyYNEcKS, and PICULETS 


Subfamily PICINAE: WooppecKeErs 
Genus COLAPTES Vigors 


Colaptes Vicors, Trans. Linn. Soc. London, vol. 14, pt. 3, 1826, p. 457. Type, 
by original designation, Cuculus auratus Linnaeus. 


Colaptes cafer (GMELIN): Red-shafted Flicker 
Picus cafer GMELIN, Syst. Nat., vol. 1, pt. 1, 1788, p. 431. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon ; 
Samwel and Potter Creek caves, Shasta County, Hawver Cave, EI- 
dorado County, McKittrick, Kern County, Carpinteria, Santa Barbara 
County, and Rancho La Brea, Los Angeles, California. 


go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Colaptes chrysoides (MALHERBE) : Gilded Flicker 


Geopicus (Colaptes) chrysoides MALHERBE, Rev. et Mag. Zool., ser. 2, vol. 4, 
December 1852, p. 553. 


Modern form reported from Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. 


Genus DRYOCOPUS Boie 


Dryocopus Bote, Isis von Oken, Bd. 2, 1826, col. 977. Type, by monotypy, 
Picus martius Linnaeus. 


Dryocopus pileatus (LINNAEUS): Pileated Woodpecker 


Picus pileatus LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 113. 


Modern form reported from Pleistocene: Cave deposits of Tennes- 
see. Late Pleistocene: Rancho La Brea, Los Angeles, California. 


Genus ASYNDESMUS Coues 


Asyndesmus Cours, Proc. Acad. Nat. Sci. Philadelphia, vol. 17, No. 1, Janu- 
ary-March (June 11), 1866, p. 55. Type, by original designation, Picus 
torquatus Wilson = Picus lewis Gray. 


Asyndesmus lewis (Gray) : Lewis’ Woodpecker 


Picus Lewis Gray, Gen. Birds, vol. 3, 1849, app., p. 22. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, and Carpinteria, Santa Barbara County, California. 


Order PASSERIFORMES: Percuine Birps 
Suborder PASSERES: Sone Birps 
Family ALAUDIDAE: Larxs 

Genus EREMOPHILA Brehm 


Eremophila Breum, Isis, vol. 21, pts. 3-4, 1828, p. 322. Type, by subsequent 
designation, Alauda alpestris Linnaeus (Sharpe, 1890). 


Eremophila alpestris (LinNAEUS): Horned Lark 


Alauda alpestris LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 166. 


Modern form reported from late Pleistocene: McKittrick and 
Rancho La Brea, Los Angeles, California. 


Family PALAEOSPIZIDAE: ParagospPiza 
Genus PALAEOSPIZA Allen 


Palacospiza ALLEN, Bull. Geol. Geogr. Surv. Terr., vol. 4, No. 2, May 3, 1878, 
p. 443. Type, by monotypy, Palaeospiza bella Allen. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE gI 


Palaeospiza bella ALLEN 


Palaeospiza bella ALLEN, Bull. Geol. Geogr. Surv. Terr., vol. 4, No. 2, May 3, 
1878, p. 443, pl. 1, figs. 1-2. 


Oligocene (Florissant lake beds) :** Florissant, Colorado. 


Family HIRUNDINIDAE: Swatiows 
Genus PETROCHELIDON Cabanis 


Petrochelidon Casanis, Mus. Hein., vol. 1, October (after Oct. 23), 1851, 
p. 47. Type, by subsequent designation, Hirundo melanogaster Swainson 
(Gray, 1855). 


Petrochelidon pyrrhonota (VrierLitor): Cliff Swallow 


Hirundo pyrrhonota Viexot, Nouv. Dict. Hist. Nat., nouv. éd., vol. 14, Sep- 
tember 1817, p. 519. 


Modern form reported from late Pleistocene: McKittrick, Cali- 
fornia. 


Family CORVIDAE: Jays, Macpies, and Crows 
Subfamily GARRULINAE: Jays and Macpigs 
Genus CYANOCITTA Strickland 


Cyanocitta STRICKLAND, Ann. Mag. Nat. Hist., ser. 1, vol. 15, No. 98, April 
1845, p. 261. Type, by original designation, Corvus cristatus Linnaeus. 


Cyanocitta stelleri (GMELIN): Steller’s Jay 
Corvus stelleri GMELIN, Syst. Nat., vol. 1, pt. 1, 1788, p. 370. 
Modern form reported from late Pleistocene: Samwel Cave, Shasta 


County, Hawver Cave, Eldorado County, Rancho La Brea, Los 
Angeles, and Carpinteria, Santa Barbara County, California. 


Genus APHELOCOMA Cabanis 


Aphelocoma CaBants, Mus. Hein., vol. 1, sign. 28, Oct. 15, 1851, p. 221. Type, 
by subsequent designation, Garrulus californicus Vigors (Baird, 1858). 


Subgenus APHELOCOMA Cabanis 


Aphelocoma coerulescens (Bosc): Scrub Jay 22 


Corvus coerulescens Bosc, Bull. Soc. Sci. Philom. Paris, vol. 1, pt. 1, 1795, 
p. 87. 


21 Recent studies indicate that the age may be Oligocene. 
22 Recorded as Aphelocoma californica (Vigors), California Jay, in the pre- 
ceding check-list. 


g2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Modern form reported from late Pleistocene: McKittrick, Kern 
County, Carpinteria, Santa Barbara County, and Rancho La Brea, 
Los Angeles, California. 


Genus PICA Brisson 


Pica Brisson, Orn., 1760, vol. 1, p. 30; vol. 2, p. 35. Type, by tautonymy, 
Pica Brisson= Corvus pica Linnaeus. 


Pica nuttallii (AupuBon) : Yellow-billed Magpie 
Corvus nuttallii Aupuson, Birds Amer. (folio), vol. 4, 1836, pl. 362, fig. 1. 


Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, and Rancho La Brea, Los Angeles, California. 


Subfamily CORVINAE: Crows and RAvENs 
Genus CORVUS Linnaeus 


Corvus LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 105. Type, by tautonymy, 
Corvus = Corvus corax Linnaeus. 
Corvus corax LINNAEUS: Common Raven °4 

Corvus Corax Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 105. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon ; 
Hawver Cave, Eldorado County, Carpinteria, McKittrick, Rancho 
La Brea, Los Angeles, and Playa del Rey (Palos Verdes sand), Los 
Angeles County, California. Pleistocene: San Josecito Cave, Aram- 
berri, Nuevo Leon. 


Corvus cryptoleucus Coucu: White-necked Raven 
Corvus cryptoleucus Coucu, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, No. 2, 
May 20, 1854, p. 66. 


Modern form reported from late Pleistocene: McKittrick and 
Rancho La Brea, Los Angeles, California. 


Corvus brachyrhynchos BrEHM: Crow 
Corvus brachyrhynchos C. L. BreuM, Beitr. Vogelkunde, vol. 2, 1822, p. 56. 
Modern form reported from Pleistocene: Seminole Field, Pinellas 


County, Florida. Late Pleistocene : Potter Creek Cave, Shasta County, 
and Rancho La Brea, Los Angeles, California.** 


23 Corvus shufeldti Sharpe is a synonym of C. corax. See Howard, Carnegie 
Inst. Washington Publ. 551, Jan. 25, 1946, p. 189. 
24 Record formerly given from Carpinteria refers to C. caurinus. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 93 


Corvus caurinus BAirp: Northwestern Crow 


Corvus caurinus Batrp, Rep. Expl. and Surv. R. R. Pac., vol. 9, 1858, pp. 559, 
569. 


Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, and Rancho La Brea, Los Angeles, California. 


Corvus ossifragus Witson: Fish Crow 
Corvus ossifragus Witson, Amer. Orn., vol. 5, 1812, p. 27, pl. 37, fig. 2. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 


Corvus pumilis WETMORE 


Corvus pumilis Wetmore, Proc. Biol. Soc. Washington, vol. 33, Dec. 30, 
1920, p. 81, pl. 2, figs. 3, 4. 


Recent (extinct) : *° Cave deposits in Cueva San Miguel (type lo- 
cality), near Morovis, Puerto Rico; Kitchen midden at Concordia, 
near Southwest Cape, St. Croix, Virgin Islands. 


Genus GYMNORHINUS Wied 


Gymnorhinus Wiep, Reise Nord-Amer., vol. 2, 1841, p. 21. Type, by monotypy, 
Gymnorhinus cyanocephalus Wied. 


Gymnorhinus cyanocephalus Wiep: Pifion Jay 


Gymnorhinus cyanocephalus Wien, Reise Nord-Amer., vol. 2, 1841, p. 22. 


Modern form reported from Pleistocene: Conkling Cavern, Pyra- 
mid Peak, Organ Mountains, Dona Ana County, New Mexico. 


Family SITTIDAE: NutTHatcHEs 
Subfamily SITTINAE: TypicaL NUTHATCHES 
Genus SITTA Linnaeus 


Sitta Linnagus, Syst. Nat., ed. 10, vol. 1, 1758, p. 115. Type, by monotypy, 
Sitta europaea Linnaeus. 


Sitta canadensis LINNAEUS: Red-breasted Nuthatch 


Sitta canadensis LINNAEUS, Syst. Nat., ed. 12, vol. 1, 1766, pp. 176, 177. 


Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 


25 Included here as it has not been found in living form, being known only 
from bones. Probably this small crow existed until modern times near Lares, 
Puerto Rico. 


94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Sitta pygmaea Vicors: Pygmy Nuthatch 
Sitta pygmaea Vicors, in Zool. Beechey’s Voy., 1839, p. 25, pl. 4, fig. 2. 


Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 


Family CHAMAEIDAE: WReEn-TITS 
Genus CHAMAEA Gambel 


Chamaea GAMBEL, Proc. Acad. Nat. Sci. Philadelphia, vol. 3, No. 7, January- 
February (May 7), 1847, p. 154. Type, by original designation, Parus 
fasciatus Gambel. 


Chamaea fasciata (GAMBEL): Wren-tit 


Parus fasciatus GAMBEL, Proc. Acad. Nat. Sci. Philadelphia, vol. 2, No. 10, 
July-August (Dec. 5), 1845, p. 265. 


Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 


Family MIMIDAE: TurasHers and MOocKINGBIRDS 
Genus TOXOSTOMA Wagler 


Toxostoma WaAGLER, Isis von Oken, vol. 24, Heft 5 (May) 1831, col. 528. 
Type, by monotypy, Toxostoma vetula Wagler = Orpheus curvirostris 
Swainson. 


Toxostoma bendirei (Cours): Bendire’s Thrasher 
Harporhynchus bendirei Cours, Amer. Nat., vol. 7, No. 6, June 1873, p. 330. 
Modern form reported from late Pleistocene: McKittrick, Kern 
County, California. 
Toxostoma redivivum (GAMBEL) : California Thrasher 


Harpes rediviva GAMBEL, Proc. Acad. Nat. Sci. Philadelphia, vol. 2, No. 10, 
July-August (Dec. 5), 1845, p. 264. 


Modern form reported from late Pleistocene: Rancho La Brea, 
Los Angeles, California. 


Genus OREOSCOPTES Baird 


Oreoscoptes Batrp, in Baird, Cassin, and Lawrence, Rep. Expl. Surv. R. R. 
Pac., vol. 9, 1858, pp. XIX, Xxxv. Type, by monotypy, Orpheus montanus 
Townsend. 


Oreoscoptes montanus (TOWNSEND): Sage Thrasher 


Orpheus montanus TOWNSEND, Journ. Acad. Nat. Sci. Philadelphia, vol. 7, 
pt. 2, Nov. 21, 1837, p. 192. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 95 


Modern form reported from late Pleistocene: McKittrick, Kern 
County, and Rancho La Brea, Los Angeles, California. 


Family TURDIDAE: TuHRrusHEs 
Genus TURDUS Linnaeus 


Turdus LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 168. Type, by subsequent 
designation, Turdus viscivorus Linnaeus (Gray, 1840). 


Turdus migratorius LINNAEUS: Robin 


Turdus migratorius LINNAEUS, Syst. Nat., ed. 12, vol. 1, 1766, p. 292. 


Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 


Genus SIALIA Swainson 


Sialia Swatnson, Philos. Mag., n. s., vol. 1, No. 5, May 1827, p. 360. Type, by 
monotypy, Sialia azurea Swainson = Motacilla sialis Linnaeus. 


Sialia mexicana SwWAINSON: Western Bluebird 


Sialia mexicana SwAINnson, Fauna Bor.-Amer., vol. 2, 1831 (February, 1832), 
p. 202. 


Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 


Family BOMBYCILLIDAE: Waxwincs 
Genus BOMBYCILLA Vieillot 


Bombycilla Viettuot, Hist. Nat. Ois. Amér. Sept., vol. 1, 1807 (1808), p. 88. 
Type, by monotypy, Bombycilla cedrorum Vieillot. 


Bombycilla cedrorum VierLtor: Cedar Waxwing 


Bombycilla cedrorum VietLtot, Hist. Nat. Ois. Amér. Sept., vol. 1, 1807 
(1808), p. 88, pl. 57. 


Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, and Rancho La Brea, Los Angeles, California. 


Family LANIIDAE: SurrKes 


Subfamily LANIINAE: SuHrikes 
Genus LANIUS Linnaeus 


Lanius Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 93. Type, by subsequent 
designation, Lanius excubitor Linnaeus (Swainson, 1824). 


96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Lanius ludovicianus LiInNAEuS: Loggerhead Shrike 
Lanius ludovicianus LINNAEUS, Syst. Nat., ed. 12, vol. 1, 1766, p. 134. 


Modern form reported from late Pleistocene: McKittrick, Kern 
County, and Rancho La Brea, Los Angeles, California. 


Family ICTERIDAE: Mrapow arks, BLACKBIRDsS, and TROUPIALS 
Genus STURNELLA Vieillot 


Sturnella Vierttot, Analyse, 1816, p. 34. Type, by monotypy, Stourne, ou 
Merle a fer-a-cheval Buffon = Alauda magna Linnaeus. 


Sturnella neglecta AupuBoN: Western Meadowlark 


Sturnella neglecta AupuBON, Birds Amer., octavo ed., vol. 7, 1844, p. 339, 


pl. 480. 


Modern form reported from late Pleistocene: Carpinteria, McKit- 
trick, Rancho La Brea, Los Angeles, and San Pedro (Palos Verdes 
formation), Los Angeles County, California. 


Genus AGELAIUS Vieillot 


Agelaius Vierttot, Analyse, 1816, p. 33. Type, by subsequent designation, 
Troupiale commandeur Buffon=Oriolus phoeniceus Linnaeus (Gray, 
1840). 

Agelaius phoeniceus (LINNAEUS): Red-winged Blackbird 
Oriolus phoeniceus LINNAEUS, Syst. Nat., ed. 12, vol. 1, 1766, p. 161. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 


Genus EUPHAGUS Cassin 


Euphagus Cassin, Proc. Acad. Nat. Sci. Philadelphia, vol. 18, No. 5, No- 
vember-December, 1866 (July 20, 1867), p. 413. Type, by monotypy, 
Psarocolius cyanocephalus Wagler. 


Euphagus cyanocephalus (WAGLER): Brewer's Blackbird 2° 


Psarocolius cyanocephalus Wac er, Isis von Oken, vol. 22, Heft 7 (July), 
1820, col. 758. 


Modern form reported from late Pleistocene: Fossil Lake, Oregon ; 
McPherson County, Kansas (Kentuck locality). 


26 The record by L. H. Miller from the Pleistocene of Hawver Cave, Eldorado 
County, California (Univ. California Publ. Geol., vol. 6, Oct. 28, 1911, pp. 390, 
400), was subsequently questioned by the same author (Condor, 1921, p. 130). 
In recent correspondence A. H. Miller writes that he has examined the material 
reported on from this cave and does not find this species represented. It is there- 
fore omitted from the list. Euphagus affiis Shufeldt is a synonym of E. cyano- 
cephalus. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—WETMORE 97 


Euphagus magnirostris MILLER 


Euphagus magnirostris A. H. Miter, Univ. California Publ., Bull. Dept. 
Geol. Sci., vol. 19, No. 1, Dec. 21, 1929, p. 14, pl. 1, figs. f, h. 


Late Pleistocene: Rancho La Brea, Los Angeles, California. 


Genus CASSIDIX Lesson 
Cassidix Lesson, Traité d’Orn., livr. 6, Feb. 1, 1831, p. 433. Type, by sub- 


sequent designation, Cassidix mexicanus Lesson=Corvus mesxicanus 
Gmelin (Gray, 1840). 
Cassidix mexicanus (GMELIN): Boat-tailed Grackle 
Corvus mexicanus GMELIN, Syst. Nat., vol. 1, pt. 1, 1788, p. 375. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 


Genus QUISCALUS Vieillot 


Quiscalus Vretttot, Analyse, 1816, p. 36. Type, by subsequent designation, 
Gracula quiscula Linnaeus (Gray, 1840). 


Quiscalus quiscula (LINNAEUS): Grackle 
Gracula Quiscula LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 109. 


Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 


Genus PYELORHAMPHUS Miller 


Pyelorhamphus A. H. Miter, Auk, vol. 49, No. 1, January 1932, p. 39. Type, 
by original designation, Pyelorhamphus molothroides Miller. 
Pyelorhamphus molothroides MILter 
Pyelorhamphus molothroides A. H. Miter, Auk, vol. 49, No. 1, January 
1932, Pp. 39, pl. 4. 


Quaternary (? Pleistocene) :?7 Shelter Cave, Pyramid Peak, Organ 
Mountains, Dona Ana County, New Mexico. 


Genus PANDANARIS Miller 


Pandanaris A. H. Miter, Condor, vol. 49, No. 1, Feb. 6, 1947, p. 22. Type, 
by original designation, Pandanaris convera A. H. Miller. 
Pandanaris convexa MILLER 
Pandanaris convera A. H. Miter, Condor, vol. 49, No. 1, Feb. 6, 1947, p. 22, 
fig. 4 a-d. 


Late Pleistocene: Pit “A,” Rancho La Brea, Los Angeles, Cali- 
fornia. 


*7 The deposits in which this extinct species was found are possibly of Recent 
age. 


98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Family FRINGILLIDAE: Grospeaks, FINCHES, SPARROWS, and 
BuUNTINGS 
Subfamily RICHMONDENINAE: Carpinats and ALLIES 
Genus PHEUCTICUS Reichenbach 


Pheucticus REICHENBACH, Av. Syst. Nat., June 1, 1850, pl. 78. Type, by sub- 
sequent designation, Pitylus aureoventris Lafresnaye and d’Orbigny (Gray, 
1855). 


Pheucticus melanocephalus (Swainson): Black-headed Grosbeak 


Guiraca melanocephala Swatnson, Philos. Mag., n. s., vol. 1, No. 6, June 
1827, p. 438. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 


Subfamily CARDUELINAE: Purp_e FINCHES, GOLDFINCHES, and 
ALLIES 


Genus HESPERIPHONA Bonaparte 


Hesperiphona Bonaparte, Consp. Gen. Avium, vol. 1, sign. 64, 1850 (Feb. 3, 
1851), p. 505. Type, by original designation, Fringilla vespertina W. 
Cooper. 


Hesperiphona vespertina (Cooper): Evening Grosbeak 


Fringilla vespertina W. Cooper, Ann. Lyc. Nat. Hist. New York, vol. 1, pt. 2, 
1825, p. 220. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 


Genus CARPODACUS Kaup 


Carpodacus Kaur, Skizz. Entw.-Gesch. Eur. Thierw., 1829, p. 161. Type, by 
subsequent designation, Loxia rosea Pallas (Gray, 1842). 


Subgenus BURRICA Ridgway 


Burrica Rweway, Man. North Amer. Birds, 1887, p. 390. Type, by original 
designation, Fringilla mexicana Miller. 


Carpodacus mexicanus (MULLER): House Finch 


Fringilla mexicana P. L. S. MU ter, Natursyst., Suppl., 1776, p. 165. 


Modern form reported from late Pleistocene: McKittrick, Kern 
County, California. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 99 


Genus SPINUS Koch 


Spinus Kocu, Syst. Baier. Zool., vol. 1, 1816, p. 233. Type, by tautonymy, 
Fringilla spinus Linnaeus. 


Spinus pinus (Wirson): Pine Siskin 
Fringilla pinus Witson, Amer. Orn., vol. 2, 1810, p. 133, pl. 17, fig. I. 


Modern form reported from Pleistocene: Carpinteria and Rancho 
La Brea, Los Angeles, California. 


Spinus tristis (LinNAEuS): American Goldfinch 


Fringilla tristis LiInNaEus, Syst. Nat., ed. 10, vol. 1, 1758, p. 181. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 


Genus LOXIA Linnaeus 


Loxia LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 171. Type, by subsequent 
designation, Loxia curvirostra Linnaeus (Gray, 1840). 


Loxia curvirostra LINNAEUS: Red Crossbill 


Loxia Curvirostra LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. I7I. 


Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 


Subfamily EMBERIZINAE: Sparrows and BunrtTINGs 
Genus PALAEOSTRUTHUS Wetmore 


Palaeostruthus WETMorE, Bull. Mus. Comp. Zodl., vol. 67, May 1925, p. 192. 
Type, by original designation, Palaeospiza hatcheri Shufeldt. 


Palaeostruthus hatcheri (SHUFELDT) 


Palaeospiza hatcheri SuHurevpt, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 16, 
Aug. 4, 1913, p. 301, pl. 55, fig. 28. 


Middle Pliocene: Near Long Island, Kansas. 


Genus PIPILO Vieillot 


Pipilo Viewxot, Analyse, 1816, p. 32. Type, by monotypy, Pinson aux yeux 
rouges Buffon = Fringilla erythrophthalma Linnaeus. 


Pipilo maculatus Swainson: Spotted Towhee 


Pipilo maculata Swainson, Philos. Mag., n. s., vol. 1, 1827, p. 434. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, and Carpinteria, California. 


I0O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Pipilo fuscus Swainson: Brown Towhee 


Pipilo fusca SWAINSON, Philos. Mag., n. s., vol. 1, 1827, p. 434. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, and Carpinteria, California. 


Pipilo angelensis DAwson 


Pipilo angelensis DAwson, Condor, vol. 50, No. 2, Mar. 16, 1948, p. 39, fig. 16. 


Late Pleistocene: Rancho La Brea, Los Angeles, California. 


Genus CALAMOSPIZA Bonaparte 


Calamospiza Bonaparte, Geogr. and Comp. List, 1838, p. 30. Type, by mono- 
typy, Fringilla bicolor J. K. Townsend=Calamospiza melanocorys 
Stejneger. 


Calamospiza melanocorys STEJNEGER: Lark Bunting 


Calamospiza melanocorys STEJNEGER, Auk, vol. 2, No. 1, January 1885, p. 49. 


Modern form reported from late Pleistocene: Meade County, 
Kansas (Jones fauna, Vanhem formation). 


Genus AMMODRAMUS Swainson 


Ammodramus SwAInson, Philos. Mag., n. s., vol. 1, No. 6, June 1827, p. 435. 
Type, by monotypy, Ammodramus bimaculatus Swainson. 


Ammodramus savannarum (GMELIN): Grasshopper Sparrow 


Fringilla savannarum GMELIN, Syst. Nat., vol. 1, pt. 2, 1789, p. 921. (Jamaica). 


Modern form reported from Pleistocene: Near Haile, 4 miles north- 
east of Newberry, Alachua County, Florida. 


Genus POOECETES Baird 


Pooecetes Batrp, in Baird, Cassin, and Lawrence, Rep. Expl. Surv. R. R. Pac., 
vol. 9, 1858, pp. xx, xxxIx. Type, by monotypy, Fringilla graminea 
Gmelin. 


Pooecetes gramineus (GMELIN): Vesper Sparrow 


Fringilla graminea GMELIN, Syst. Nat., vol. 1, pt. 2, 1789, p. 922. 
Modern form reported from Pleistocene: Rancho La Brea, Los 
Angeles, California. 
Genus CHONDESTES Swainson 


Chondestes Swatnson, Philos. Mag., n. s., vol. 1, No. 6, June 1827, p. 435. 
Type, by monotypy, Chondestes strigatus Swainson. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE IOI 


Chondestes grammacus (Say): Lark Sparrow 
Fringilla grammaca Say, in Long, Exped. Rocky Mts., vol. 1, 1823, p. 139. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 


Genus AMPHISPIZA Coues 


Amphispiza Cougs, Birds Northwest, 1874, p. 234. Type, by original designa- 
tion, Emberiza bilineata Cassin. 


Amphispiza bilineata (Cassin): Black-throated Sparrow 
Emberiza bilineata Casstn, Proc. Acad. Nat. Sci. Philadelphia, vol. 5, No. 5, 
September-October (Dec. 7), 1850, p. 104, pl. 3. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 
Amphispiza belli (Casstn): Bell’s Sparrow 
Emberiza Belli Cassin, Proc. Acad. Nat. Sci. Philadelphia, vol. 5, No. 5, 
September-October (Dec. 7), 1850, p. 104, pl. 4. 


Modern form reported from late Pleistocene: McKittrick, Kern 
County, and Rancho La Brea, Los Angeles, California. 


Genus SPIZELLA Bonaparte 


Spizella Bonaparte, Giornale Arcadico, vol. 52, October-December 1831 
(1832), p. 205. Type, by monotypy, Fringilla pusilla Wilson. 


Spizella passerina (BECHSTEIN) : Chipping Sparrow 
Fringilla passerina BECHSTEIN, in Latham, Allgem. Uebers. Vogel, vol. 3, 
pt. 2, 1798, p. 544, pl. 120, fig. 1. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 


Genus ZONOTRICHIA Swainson 


Zonotrichia SwAINSON, in Swainson and Richardson, Fauna Bor.-Amer., 
vol. 2, 1831 (February 1832), p. 493. Type, by subsequent designation, 
Fringilla pensylvanica Latham = Fringilla albicollis Gmelin (Bonaparte, 
1831). 


Zonotrichia leucophrys (Forster): White-crowned Sparrow 
Emberiza leucophrys J. R. Forster, Philos. Trans., vol. 62, art. 29, 1772, 
p. 426. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 


102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


Genus PASSERELLA Swainson 


Passerella Swainson, Nat. Hist. and Class. Birds, vol. 2, July 1, 1837, p. 288. 
Type, by monotypy, Fringilla iliaca Merrem. 


Passerella iliaca (MERREM): Fox Sparrow 


Fringilla iliaca MerrEM, Avium Rar. Icones et Descrip., vol. 2, 1786, p. 37, 
pl. ro. 


Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, and Carpinteria, California. 


Genus MELOSPIZA Baird 
Melospiza Batrp, in Baird, Cassin, and Lawrence, Rep. Expl. Surv. R. R. 
Pac., vol. 9, 1858, pp. XX, XL, 440, 476. Type, by original designation, 
Fringilla melodia Wilson. 
Subgenus MELOSPIZA Baird 
Melospiza melodia (Witson): Song Sparrow 
Fringilla melodia Witson, Amer. Orn., vol. 2, 1810, p. 125, pl. 16, fig. 4. 
Modern form reported from late Pleistocene: Rancho La Brea, Los 
Angeles, California. 
INCERTAE SEDIS 
Genus CIMOLOPTERYX Marsh 28 


Cimolopteryx Marsu, Amer. Journ. Sci. ser. 3, vol. 38, 1880, p. 83, foot- 
note. Type, by monotypy, Cimolopteryx rarus Marsh. 


Cimolopteryx rarus MarsH 


Cimolopteryx rarus MarsH, Amer. Journ. Sci., ser. 3, vol. 38, July 1880, 
p. 83, footnote. 


Upper Cretaceous (Lance formation) : Niobrara County, Wyoming. 
Cimolopteryx retusus MarsH 


Cimolopteryx retusus Marsu, Amer. Journ. Sci., ser. 3, vol. 44, August 1892, 
p. 175. 


Upper Cretaceous (Lance formation) : Niobrara County, Wyoming. 


28 Lambrecht, Handb. Palaeorn., 1933, pp. 586-587, lists this genus at the end 
of the Ichthyornithiformes. He suggests that the two species belong in separate 
genera, possibly in different families. See also Shufeldt, Trans. Connecticut 
Acad. Arts Sci., vol. 19, February 1915, pp. 11, 12, and 76. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 103 


Genus EOPTERYX Meyer 
Eopteryx Meyer, Ber. Senckenberg. Nat. Ges. Frankfurt am Main, 1887, 
p. 14. Type, by monotypy, Eopteryx mississippiensis Meyer. 
Eopteryx mississippiensis MEYER 29 


Eopteryx mississippiensis Meyer, Ber. Senckenberg. Nat. Ges. Frankfurt am 
Main, 1887, p. 14, pl. 2, figs. 22a-22c. 


Eocene: Jackson, Mississippi. 


(Genus uncertain) 


Falco falconellus SHUFELDT °° 
Falco falconella SHUFELDT, Trans. Connecticut Acad. Arts Sci., vol. 19, Feb- 
ruary 1915, p. 40, pl. 15, figs. 139-143. 


Eocene (Bridger formation) : Dry Creek ?, Wyoming. 


Genus FONTINALIS Lesquereux 


Fontinalis LESQUEREUX, Rep. U. S. Geol. Surv. Terr., vol. 8, 1883, p. 135. 
Type, by monotypy, Fontinalis pristin Lesquereux. 


Fontinalis pristina LESQUEREUX °1 
Fontinalis pristina LESQUEREUX, Rep. U. S. Geol. Surv. Terr., vol. 8, 1883, 
p. 135, pl. 21, fig. 9. 
Oligocene (Florissant lake beds): Florissant, Colorado. 


Genus HEBE Shufeldt 


Hebe SHUuFELDT, Journ. Geol., vol. 21, October-November (Nov. 1), 1913, 
p. 644. Type, by monotypy, Hebe schucherti Shufeldt. 


Hebe schucherti SHUFELDT ®2 


Hebe schucherti SHUFELDT, Journ. Geol., vol. 21, October-November (Nov. 1), 
1913, p. 644, fig. 10, a, b. 


Eocene: ** 5 miles west of Green River, Wyoming. 


29 Described from a fragmentary vertebra. 

80 Not a falcon; relationships doubtful. See Wetmore, A., Proc. U. S. Nat. 
Mus., vol. 84, Nov. 3, 1936, pp. 77-78. 

81 Type a fragment of a fossil feather, described originally as a species of 
moss. See Knowlton, Proc. U. S. Nat. Mus., vol. 51, Nov. 24, 1916, p. 245, and 
Wetmore, Bull. Mus. Comp. Zo6l., vol. 67, May 1925, p. 184. Possibly of Oligo- 
cene age. 

82 Said to be a passeriform bird with four notches in the posterior border of 
the sternum; of uncertain affinity. Hebe Shufeldt, 1913, is preoccupied by Hebe 
Risso, 1826 (applied to a genus of crustaceans), so that should the form here 
under consideration be definitely identified it may require a new generic appella- 
tion. There is no necessity for action at this time in view of its uncertain rela- 
tionships. 

88 From data furnished by Dr. M. R. Thorpe, of the Peabody Museum, Yale 
University. 


104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Genus IGNOTORNIS Mehl 


Ignotornis MEHL, Amer. Journ. Sci., ser. 5, vol. 21, May 1931, p. 443. Type, 
by monotypy, Ignotornis mcconnelli Mehl. 


Ignotornis mcconnelli MEHL 34 


Ignotornis mcconnelli Ment, Amer. Journ. Sci., ser. 5, vol. 21, May 1931, 
Pp. 444, fig. 1. 


Cretaceous (Dakota sandstone): About 14 miles northwest of 
Golden, Colorado. 


Genus LAOPTERYX Marsh 


Laopteryx Marsu, Amer. Journ. Sci., ser. 3, vol. 21, April 1881, p. 341. Type, 
by monotypy, Laopteryx priscus Marsh. 


Laopteryx priscus Marsu 35 
Laopteryx priscus Marsu, Amer. Journ. Sci., ser. 3, vol. 21, April 1881, p. 341. 


Upper Jurassic (Morrison formation): Quarry 9, Como Bluff, 
southern Wyoming. 


Genus LAORNIS Marsh 


Laornis MarsH, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, p. 206. Type, 
by monotypy, Laornis edvardsianus Marsh. 


Laornis edvardsianus Marsu °6 


Laornis edvardsianus Marsu, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, 
p. 206. 


Paleocene (Hornerstown marl): Near Birmingham, New Jersey. 


Genus PALAEONORNIS Emmons 


Palaeonornis Emmons, Amer. Geol., pt. 6, 1857, p. 148. Type, by monotypy, 
Palaeonornis struthionoides Emmons. 


Palaeonornis struthionoides Emmons 37 


Palaconornis Struthionoides Emmons, Amer. Geol., pt. 6, 1857, p. 148, fig. 114. 


? Triassic: Anson County, North Carolina. 


84 Described from fossil impressions of 4-toed footprints, apparently with webs 
connecting the three anterior toes. 

85 J. D. Dana, Amer. Journ. Sci., ser. 5, vol. 12, July 1926, pp. 3, 4, considered 
the avian affinity of this supposed species as not definitely certain. 

36 Doubtfully related to Anseriformes. Lambrecht, Handb. Palaeorn., 1933, 
pp. 526-527, has placed it uncertainly after the Aramidae. 

87 Affinity doubtful: possibly not avian. 


NO. 5 CHECK-LIST OF FOSSIL BIRDS—-WETMORE 105 


Genus UINTORNIS Marsh 


Uintornis MarsH, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, p. 250. 
Type, by monotypy, Uintornis lucaris Marsh. 


Uintornis lucaris MArsH 38 


Uintornis lucaris MarsH, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, p. 259. 


Eocene (Bridger formation): Near Henry’s Fork, Wyoming. 


Genus YALAVIS Shufeldt 


Yalavis SHUFELDT, Journ. Geol., vol. 21, October-November (Nov. 1), 1913, 
p. 649. Type, by monotypy, VYalavis tenuipes Shufeldt. 


Yalavis tenuipes SHUFELDT *? 


Yalavis tenuipes SHUFELDT, Journ. Geol., vol. 21, October-November (Nov. 1), 
1913, p. 649, figs. 11c and 12c. 


Geologic age and locality of occurrence not known. 


88 According to Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, February 
IQI5, pp. 50-52, 77, pl. 6, fig. 42, this species is of uncertain affinity, and is not a 
woodpecker as suggested by Marsh. 

39 Said in the original description to be a passeriform bird of uncertain affinity. 


- 


elacuiarl 





SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 6 


Charles D. and Mary Waux Talcott 
Research Fund 


PALEOCENE MAMMALIAN FAUNAS OF THE 
BISON BASIN IN SOUTH-CENTRAL 
WYOMING 


(WirH 16 PuLatEs) 


By 
C. LEWIS GAZIN 


Curator, Division of Vertebrate Paleontology 
United States National Museum 
Smithsonian Institution 











° AME-INCRS 
S DIEEWO Ase 
Ps See AYO, £ 








(PuBLICATION 4229) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
FEBRUARY 28, 1956 


/ 


THE LORD BALTIMORE PRESS, INC. 
BALTIMORE, MD., U. S. A. 


CONTENTS 


; Page 

en EL OCUIGEIO My care crci crete) avec) ore. arelevale oeicalfade Vase tavestatane, ofeteveliclsi evel olejsicra%sfalal ela: e “arelaraiciers I 
PBMC ITIENIES Noe 5 os a alain ets te scot oaishorasercue aise srciatereveie eielsiaie wi4)s'skarelstels 2 
Riastory. Of Myesh@ation. « sass easyer aa qeesisbls Fosse sc eweeweoescas ees 2 
Occurrence and preservation, of material... oc .cccccciccceesenncneeeeds 3 
PIR ea ESISOM ASIN LalIMAS «soya ore c.ctesore la OMNI T Nleva a araia wi'eistevoitoralsseneyaays!oye'cie/e cloves) « 4 
Environment and relationships between the Bison basin faunas.......... Fi 
Mime mand correlation Of the tauNase.. sive no s.ccices 6.0.0 4 85 0b 4 Sie Seine so wae 10 
Systematic description of vertebrate remains. ..........cccccsccsccccees 12 
PRG IDEL Teds Pavers cvcrecs, ores. 2) arate Cron Syenaote rayelayoucole)cieks svorelersl esi ale! ale! sye/oyeve ¢,6icin'si 12 

SS ULE Lede Seat g steep HA Toa ote naueTeY POR Na/ eis ONG SieVelw a aysl evolfer srelel crocs. 1stenotejotse 12 

NSTI SUG AG:, tr cs'os aks a yavel suapeley he letot Vast oVous (ove ole: ein shoves: e; enn cle) 8) or'sh 87s exarele)@ 12 

Micon ended caigra fate: cance ots ‘sis vovereleso7s fav sheyeyev sn ey fis Siek she el exejo leavers jouelas opehal eve ocacsyoie 12 
Mal tittberctilatas reversreteroe ecciya ove teres stole sor sher enters sje lslovereie iciersvera yi 12 
PEMOCONHGAG g2 eine. surerciesie ws win cunts a sels eraoie everett Symeiarecnls Ome 12 

Marstptalia: << 5 rs,c SARC TS VRAVSTO GCE SECS ohSLSIGTELO' 6 SiGl Seo BLO are eo MoLSgT OS 14 
idelphtdae’” Sere oe syeiciers ooops «, goters ss tersceueinl sae ers cont oiduaies efaleveretes 14 

TTSCCEV OTA eres atte cach ts tate cia Se aOR or sia Gs fate cleisions a bia leysrere eis ore 15 

ANGE NTE RCAC circle Se hata Bionie ey aera sruwere oa tis pa clglere clap nites 15 
Pantolestidae: teracjen & cat a tehetarolote, Naveiolat sie send Slaves diaue-s stslety ete 17 

IP Attia te Sipecrer rakes cove eere tas tecr eve esrnste ter sehen eioy cictchs cater aeeeio ta fuse) evshexarete enstele 19 

PESTA AIAG selene a oicieng e8i vars e(dip/e! vies eibsis se #5 farelare sania’ ¢ sclars 19 

Garmivana wee atti e ee ere cichatere ciate ere hci e reas crsteneke Ain ere hina were are area 25 
Arctocyonidae ........ Stealers SP accreted avatar eitia eas ale Saha avs vena’ s 25 
IVES oRr ye IGae sy aie cats taretaicht, tiene eat Oaveseie se tia, he lasthaya rel eiohste aloes 35 

INA Teh 11 Ba cvss cs Svetea cottnsy tee ate kakok crave vetovstonela Nanere: dralteha tararvieverstes ottarare 35 
REGIA AME cisions c'd/5,az01a' slotetaceiacatere isa « laieuival oie miei Siem Wave, cieaLS 36 

Ely OpsGdOntid aes vert axctire aveteirsiatere tt evele aioteh o4eie\ ay alts aheretonsters! ators ouers 36 
Phenacodonttdae wey. syacvertscte oh chee athe sieciatevessreld otele weiere sexeiae 42 

Panta Ontaaiei ac cre cis vate etalon tatelete eters ere a tae neato cstele bisa tl aare watts 47 

Cory PHOGONGae i teresa pare wtidee oe, cdiaslet aotenades Lsatade 47 

PRELERONICES meter tai ecoichel ste os oy ous elalecers Spa desearet aha der eyeteis Gisreahlah Slavs Gis sales Mare selon ates 5I 
MPT MAN AEN Tas GIL | COEALCS 605.0 o o,0tn/ee' 4, ece¥Ain cays ovale aot Wid Wale a1 sieiB laiaioya ep aleinin. 4/eve/3 54 


ili 


ON AN HRW DN HH 


ILLUSTRATIONS 
PLATES 


(All plates following page 58.) 


. Multituberculates and insectivores from the Bison basin Paleocene. 
. Primates and marsupials from the Bison basin Paleocene. 

. Pronothodectes from the Bison basin Paleocene. 

. Plesiadapis from the Bison basin Paleocene. 

. Tricentes and Chriacus from the Bison basin Paleocene. 

. Thryptacodon from the Bison basin Paleocene. 

. Claenodon from the Bison basin Paleocene. 

. Litomylus and Protoselene? from the Bison basin Paleocene. 
. Haplaletes and Gidleyina from the Bison basin Paleocene. 

. Phenacodus? from the Bison basin Paleocene. 

. Condylarths and Titanoides from the Bison basin Paleocene. 
. Caenolambda from the Bison basin Paleocene. 

. Caenolambda from the Bison basin Paleocene. 

. Caenolambda from the Bison basin Paleocene. 

. South rim of Bison basin showing fossil localities. 

. Two fossil localities in the Bison basin. 


FIGURES 
Page 
. Map of western Wyoming and portions of adjacent States............ 5 
. Histogram: of length of Ms of: Claenodon 25. Occ ec esn en ces anew ak 33 


iv 


Charles D. and Mlary Waux Walcott Research Fund 


PALEOCENE MAMMALIAN FAUNAS OF THE 
BISON BASIN IN SOUTH-CENTRAL 
WYOMING 


By C. LEWIS GAZIN 


Curator, Division of Vertebrate Paleontology 
United States National Museum 
Smithsonian Institution 


(Wirx 16 PLatEs) 


INTRODUCTION 


One of the more interesting developments relative to the investi- 
gation of early Tertiary mammals in the Rocky Mountain region 
during the past several years was the discovery in 1952 by Dr. R. W. 
Brown, Harold Masursky, and H. R. Christner of the U. S. Geologi- 
cal Survey of the occurrence of Paleocene mammal remains in the 
Bison basin of Wyoming. The Bison basin is in the Sweetwater 
drainage to the north of the Red Desert, and its south rim forms part 
of the Continental Divide, separating inland drainage of the Continen- 
tal Divide basin from that of the Missouri River system. The gray 
and buff to reddish silty clays and sandstones of the Paleocene are 
here exposed at intervals along the escarpment bounding the basin. 
There are four principal fossil localities—two in the exposures below 
the south rim of the basin, one in the southwestern part, and one at 
the western extremity. These have been determined as lying within 
sections 28 and 29 of T. 27 N., R. 95 W., in Fremont County, but 
very near the southern boundary. 

Slight differences in age would appear to be indicated by the faunas 
represented at the different localities, but most if not all may be in- 
cluded within the early or lower part of Tiffanian upper Paleocene. 
Similarities to the Torrejonian fauna of the Montana Fort Union are 
noted, but these are in part attributed to a possible similarity in rather 
general environmental conditions. A resemblance is evident in the 
variety of carnivores and condylarths, modified by certain genera 
which are regarded as indicative of Tiffanian time. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 131, NO. 6 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


ACKNOWLEDGMENTS 


In addition to the Geological Survey personnel above mentioned as 
having discovered the occurrence of Paleocene mammals in the Bison 
basin, acknowledgment is due George N. Pipiringos, James Mac- 
Lachlan, Dr. J. R. Hough, and Robert DeMar for having given field 
aid in 1953. Dr. Paul O. McGrew aided in turning over, for the pur- 
poses of this study, collections obtained for the University of Wyo- 
ming in 1953. Particular mention may be made of interest shown in 
this work by Dr. Roland W. Brown in having first called my atten- 
tion to the occurrence, and in his marked contribution to the field col- 
lecting in both 1952 and 1953, as well as being an original discoverer. 

Acknowledgment is made of aid no less important from Drs. George 
G. Simpson, Edwin H. Colbert, Bobb Schaeffer, and Mrs. Rachel H. 
Nichols in permitting me to examine and make comparisons with vari- 
ous Paleocene collections in the American Museum and from Dr. 
Glenn L. Jepsen in making available type and other materials in the 
extensive Polecat Bench Paleocene collections at Princeton University. 

The drawings depicting the specimens shown in plates I to II were 
made by Lawrence B. Isham; those of Caenolambda pattersoni in 
plates 12 to 14 by William D. Crockett. 


HISTORY OF INVESTIGATION 


Following discovery of the fossil materials by the U. S. Geological 
Survey party in July 19527 and their reference to me for study and 
report, the results of a preliminary examination were presented before 
the Cambridge meeting of the Society of Vertebrate Paleontology in 
November. During the later part of 1952 and early in 1953, Wallace 
G. Bell, a graduate student at the University of Wyoming, engaged 
in a thesis study of the geology of a rather general area including the 
3ison basin, and Paul McGrew made collections at certain of the fos- 
siliferous sites. Agreement was early reached whereby the University 
of Wyoming party would join with the Smithsonian Institution—U. S. 
Geological Survey expedition in the search of these exposures in the 
summer of 1953, and that I would be permitted to study and describe 
the collections as a whole. 

In June 1953 a party from Washington consisting of Dr. Roland 
W. Brown, Franklin Pearce, and myself was accompanied in the field 
by Messrs. Pipiringos and MacLachlan. We were joined at the fossil 


1A very cursory examination of these beds was made by the writer in 1951, 
accompanied by C. Li Jenks, Jr., of the Shell Oil Co., but on the north side of 
the basin where exposures are evidently quite barren of fossils. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN——-GAZIN 3 


locality by Dr. Hough and Mr. DeMar of the U. S. Geological Survey 
and by Dr. McGrew and his party of students from the University. 
The combined efforts of the group were largely concentrated in a 
search of the vicinity of the small saddle or discovery locality below 
the south rim of the basin, and a site found by Mr. Bell in the south- 
western part of the basin. Later, a third fossiliferous site was en- 
countered by Dr. Brown, Mr. Pearce, and myself at the western ex- 
tremity of the basin, and Pearce located a fourth along a ledge on 
the south escarpment between the saddle and Bell’s Titanoides locality. 
All sites were revisited by Pearce and me with continued success 
in 1954 but with diminishing returns, as it appears that the original 
richness of the sites was due largely to a residual concentration of 
materials, and the interval between successive field seasons is evidently 
too short for erosion to afford profitable collecting. Moreover, there 
seemed to be no one place where the concentration of bone might be 
regarded as great enough to warrant quarrying operations. 


OCCURRENCE AND PRESERVATION OF MATERIAL 


The four principal fossil occurrences (see pls. 15 and 16) are re- 
ferred to in the following discussion as the saddle or discovery lo- 
cality, the ledge locality, Bell’s or the Titanoides locality, and the 
west-end locality. The small badland saddle where Brown and others 
of the Geological Survey first encountered bone is located about mid- 
way north and south in the eastern half of sec. 28, T. 27 N., R. 95 W., 
according to information from Bell’s mapping furnished me by Mc- 
Grew. The richest concentration of the smaller forms was in the 
saddle, determined by means of a hand level to be about 58 feet below 
the rim or top of the escarpment immediately to the south. The beds 
here have a dip of approximately 9° southward. Fossils were found 
scattered for about a couple of hundred feet in either direction from the 
saddle and stratigraphically near the same level, although a couple of 
specimens of Plesiadapis jepseni in the University of Wyoming col- 
lections came from possibly 50 feet higher. The material rather gen- 
erally consists of incomplete jaws and maxillae and a good number of 
isolated teeth. A single skull, that of the new pantodont Caenolambda 
pattersont, was encountered in a nodule a little distance away but at 
about the stratigraphic level of the saddle. 

Approximately a quarter of a mile or more to the west, apparently 
in the western half of section 28, a northwest-facing exposure ex- 
hibits a prominent ledge about 25 or 30 feet below the rim. A fair 
concentration of jaw and maxillary portions and isolated teeth was 
found for a hundred feet or less along the ledge and in the soft clay 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


for a few feet immediately above. The ledge locality would seem 
almost certainly to be stratigraphically much higher than the saddle. 
The locality is nearer the rim, hence topographically higher, and the 
dip of the beds may be a little more southwesterly than at the saddle 
so that the ledge would appear to be stratigraphically higher than any 
portion of the escarpment above the saddle. A considerable extent, 
however, of the exposures between the two localities is obscured by 
talus and vegetation, and so the relationship could not be determined 
precisely. 

The locality that I am informed Mr. Bell discovered, and from 
which the University of Wyoming secured upper teeth of Titanoides 
primaevus, is still farther west, about midway north and south in the 
eastern half of adjacent section 29. The general locality is in the 
southwestern part of the basin and just west of the most westerly of 
the wagon trails crossing the south rim of the basin. Fossils, though 
comparatively few, were found ranging from very near the top of the 
escarpment to 30 or 40 feet stratigraphically lower. The beds here 
appear to have a greater southerly dip than at the saddle or ledge lo- 
calities, and the stratigraphic position relative to the more easterly 
localities is not readily evident from field relations, as much of the 
escarpment between this locality and the ledge is obscured. The 
faunas discussed in the following part of this paper would suggest 
that it is still higher. 

The most westerly locality is an east-facing gravel-capped exposure 
at the west end of the basin, evidently about midway east and west 
across section 29, close to the northern line. Fossils were discovered 
here in a very restricted zone around a low hill set out from near the 
base of the exposure and at about its most southeasterly extent. Jaws 
and isolated teeth were encountered over an area of only a few hun- 
dred square feet on the slopes extending out from the base of this hill. 
The west-end exposure is well isolated by grass and sagebrush slopes 
from the Titanoides locality to the south, but there seems no doubt 
from the dip at the latter locality that the west-end site must be con- 
siderably lower stratigraphically, unless the intervening rock is com- 
plicated by changes in dip or faulting. The relative position of the 
horizon represented with respect to that at the ledge or saddle is un- 
certain, but there is some evidence from the fossils that it may not be 
far removed in time from that represented at the ledge. 


THE BISON BASIN FAUNAS 


The following tabulation pertains to the four principal localities in 
the Bison basin from which collections were obtained. The figures 


S 
e 
5 





Fig. 1—Map of western Wyoming and portions of adjacent States showing Eocene 
sedimentary basins, with nearby Paleocene fossil localities numbered as follows : 
1, Bison basin Tiffanian; 2, Polecat Bench Puercan to Clarkforkian; 3, Buckman 
Hollow Clarkforkian (Almy) ; 4, Wagonroad ridge and Dragon Canyon, Puercan and 
Dragonian (North Horn). Map reproduced from Osborn, U. S. Geol. Surv. Monogr. 


55, figs. 9, 49, 1920. 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


used refer to the number of specimens encountered, giving an indica- 
tion of the extent of the material upon which identifications are based, 
as well as some, though generally meager, information on the rela- 
tive abundance of the various forms within and between the faunas. 
In certain instances species names are repeated but on a less certain 
or comparative basis where differences of a minor sort, possibly 
variant or subspecific in value, are noted between related materials 
from two different localities. 

From the tabulated data it is seen that the known collections total 
about 236 generally determinable specimens. Of these 53 are in the 
collections of the University of Wyoming, with occurrence divided 
between the vicinity of the saddle and the Titanoides locality. The 
larger collections are in the U. S. National Museum and represent 
all four sites. 


g g 
3 — = g 
ies aan rs 
oF Opes 8 
REPTILIA : Sens 8 
Sauria: Grae = & 
PA SANIG UTIGEE 3 wean x vininie.g 3:5 sce ateb evista sees + getters 6 I 
MAMMALIA: 
Multituberculata: 
Cf. Ptilodus montanus Douglass...........0..002% I 
Cf. Ectypodus musculus Matthew and Granger.... I .. 
Ci. Ectypodus hazeni Jepsen......ccssccecscssses I 
Cf. Anconodon russelli (Simpson)............... Ti oacaeate 
Marsupialia : 
Peradectes elegans Matthew and Granger......... I 
Peradectes pault, new SPeCi€S........cccecsscaves 2 
Insectivora : 
Diacodon pearcet, new SPECieS.........cccseccenes I 
Bisonalveus browni, new genus and species........ 4 
Primates: 
Pronothodectes, cf. matthewi Gidley............6 4 
Pronothodectes simpsoni, new species.........+44- ie 9 
Pronothodectes, cf. simpsoni, new species......... SF baa: ae 
Plesiadapis, cf. fodinatus Jepsen........seseceeves es I 2 3 
Plestadapis jepseni, new SPECIES........e0ceceeves tig 2 
Plesiadapis, cf. jepsent, new specieS.........+.05 os coin Os 2 
Carnivora: 
Tricentes fremontensis, new SPECi€S.........eeeee II 20. 
Chriacus, near C. pelvidens (Cope)......seeeeeee Bincace I 
CHAISE FEB SSSI Yi os 50 os cto keene ee ene I iS ee 
Thryptacodon, cf. australis Simpson...........008 «+ I I 
Thryptacodon demari, new SpecieS...........00006 ie 6 I 
Thryptacodon, cf. demari, new species........+++. 2 I I 
Thryptacodon belli, new specieS.........+..eeeee- II geen 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN Zi 
Stine 
ay ie ate ails 
ana 8 
t as to $ s 
MAMMALIA—continued Chee ae 2 
Carnivora—continued a A 5 
Claenodon, cf. procyonoides (Matthew).......... 3 
Claenodon, cf. montanensis (Gidley)...........+. 5 : 
Clacnadow, ck ferox (COPE) idss'scsctacsscccese Rens 3 I 
Claenodon acrogenius, new species............06- 6 I 
TDESSGEUSY SP Us eta sie a ioe a re elte Eshsfele oreielaal che resele a.6.1bs 2 
Didymictis, near D. tenuis Simpson............++ I 
Condylarthra: 
Promioclaenus pipiringost, new species..........-. I 
Cf. Promioclaenus pipiringosi, new species........ i 2 
Promtoclaenus? Sp. 0.) cise sedis ole 0 Rh aides erautacna a I 
Litomylus scaphicus, new SPecieS.........eeeeeeee 2 I 
Litomylus scaphiscus, new SPeCi€S........eeee ees a gk 2 
Haplaletes pelicatus, new SpecieS........0.--.006. 4 
Hablaletes sertor, new SPCCIES. ..65..ccsccsccecsss oo eee eye I 
Protoselene? novissimus, new SpeCieS..........46- 2 
Estolestes lacunatus, New''SPeCieS if ...0. ccc cece s 08 Ai I 3 
Cf. Litolestes lacunatus, new species............6. EV ithe 
Gidleyina wyomvingensis, NEW SPECIES..........008 0s 10 9 
Gidleyina, cf. wyomingensis, new species.......... Be es alee 3 
Phenacodus? bisonensis, new specieS...........00. 28 2 } ; 
PACHECEOMS . SPs CIALRE) osc gcisae <5 ebice sare. ces 58.4 ae I I 2 
Pantodonta : 
Titanoides primaevus Gidley.............0see0ees eOe tion, FEMS I 


Caenolambda pattersoni, new genus and species.... I. 
Cf. Caenolambda pattersoni, new genus and species. I LR Ug ees 
Pantodont. undetsa(tooth frags.) ssccec see's oe cee 2 5 3 I 


4 ae of these are from the vicinity of the saddle locality but approximately 50 feet 
igher. 


ENVIRONMENT AND RELATIONSHIPS BETWEEN THE 
BISON BASIN FAUNAS 


Although differences are noted between the faunas represented at 
the four principal localities, all four faunas appear to be Tiffanian, 
and for the most part, if not all, a comparatively early part of this 
time interval. Whether these differences are essentially a matter of 
chance collecting, of facies or environmental differences with time as 
a minor factor, or of change resulting in part from evolution of cer- 
tain persisting kinds, but complicated by migration involving the in- 
troduction and disappearance or local extinction of others, is not en- 
tirely clear. Each, though, is likely to have contributed to the picture. 

Chance collecting is unquestionably an important factor where the 
number of specimens of each form is small, but it cannot be predicted 
with any assurance that further collecting would increase the faunal 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


resemblance between sites or further emphasize dissimilarities. Un- 
doubtedly certain missing forms would appear, increasing the num- 
ber of genera and species common to two or more localities, but better 
representation of populations of each would likely point out per- 
sistent differences. 

A stratigraphic difference seems evident between at least three of 
the four sites so that differences due to environment or facies would 
not be unexpected, whereas this would be unlikely were the same 
horizon represented at each in so restricted a geographic area. Chance 
collecting may be appealed to as distorting the picture with respect to 
environmental differences, particularly where the numbers of speci- 
mens are small; nevertheless, with the same collecting personnel in- 
volved at each of the sites, attention may be directed to certain con- 
trasting features observed. The saddle locality for example shows 
evidence of a fauna containing a wealth of smaller mammalian forms. 
Scant numbers of specimens show a variety of multituberculates, 
marsupials, and insectivores not represented at the other localities. 
Still better materials include representation of primates, creodonts, 
and condylarths, with equivalent or closely related forms known at 
the other levels. Almost all the genera here peculiar to the saddle 
locality, with the exception of Bisonalveus and Protoselene ?, are else- 
where known in later faunas, so that one may postulate in addition 
to time an environmental difference possibly only of local significance 
which, were it not for the persistence of the primates, would suggest 
a more open or less sylvan environment for the later levels. The 
saddle genera missing from the higher levels I suspect are forest- 
dwelling types. On the other hand, all but two, Dissacus and Tita- 
noides, of the genera known in the four faunas as a whole are rep- 
resented in the saddle collections, indicative of a cosmopolitan 
assemblage of a type perhaps better known only from the Crazy Moun- 
tain Fort Union and evidently also from the Polecat Bench. In all 
probability the Torrejon fauna is of a more open terrain, although 
points of resemblance are seen in the Carnivora. Nevertheless, the 
condylarths in particular and many of the other forms from the Bison 
basin seem more closely allied to faunas of the Crazy Mountain Fort 
Union as well as to those of the Polecat Bench series. No doubt much 
of this resemblance is regional in significance and perhaps generally 
characteristic of the extensive Fort Union, which the Wyoming as 
well as the Montana sites have been regarded as representing. These 
in turn are geographically remote with a rather distinctive differ- 
ence in latitude from the Nacimiento deposits in the San Juan basin 
of New Mexico. 


NO. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 9 


There would seem to be little doubt that some differences observed 
between the Bison basin faunas may be attributed to time. That there 
is a time difference seems evident from the stratigraphic relations, and 
the localities in the foregoing tabulation are arranged from left to 
right in ascending order as far as I can determine, except that the 
relative position of the west end with respect to the ledge locality is 
entirely uncertain. The more marked differences between the various 
localities, such as in the genera represented, would, between what ap- 
pear to be closely related faunas, have less significance as a time fac- 
tor than the differences observed between the most closely related 
types. Change resulting from the evolution of certain forms, or the 
superseding of primitive by more advanced though related types, may 
be noted in at least three of the orders. 

Among the primates Pronothodectes is represented in both the sad- 
dle and ledge localities, but evidently not higher, and the more primi- 
tive Pronothodectes matthewi has been found only at the saddle level. 
Plesiadapis is recognized at all localities, and the smaller of these, 
P. jepseni, is best or more typically represented at the saddle and 
ledge, whereas large P. cf. fodinatus has not been found in the saddle 
and most of the material is from the presumably highest or Titanoides 
locality. 

Among the Carnivora, Tricentes was encountered frequently at the 
saddle, scantly at the ledge and not higher. Particularly striking with 
regard to the change in Carnivora is the development of Thryptaco- 
don. The small T. belli found in the saddle is replaced at the ledge 
by distinctly larger T. demari, almost certainly through development 
in situ. Material of Thryptacodon, which appears structurally a 
little different than the foregoing and resembling more closely 7. aus- 
tralis, is found only at the west-end and Titanoides localities, although 
associated there with scant materials that appear to indicate one or 
both of the other species. These latter, however, may possibly repre- 
sent variation within a population of a single species in which the 
mean is rapidly changing. On the other hand, 7. australis may have 
appeared from elsewhere during the later time here represented. The 
impressive display in range of size for Claenodon is seen only at the 
saddle, and only the typically large form compared to Claenodon 
ferox and possibly a single specimen of Claenodon acrogenius occur 
at higher levels. It is possible that Tricentes and the smaller forms 
of Claenodon may have become extinct during the interval included 
by the Bison basin faunas. 

Most of the small condylarths are found only in the saddle or ledge 
faunas, and there appears in these no comment-worthy differences be- 


10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


tween the two faunas; however, a single specimen of Haplaletes en- 
countered at the Titanoides locality is of a surprisingly larger form 
than the Haplaletes represented at the saddle. On the other hand, a 
single specimen from the saddle referred tentatively to Litolestes 
lacunatus is scarcely different than the typical material from the Tita- 
noides locality. Among the larger condylarths the form described as 
Gidleyina wyomingensis from the ledge and west-end localities may 
be a little more progressive than indicated by material referred to it 
from the saddle. Abundant Phenacodus? bisonensis is almost re- 
stricted to the saddle level although two specimens came from the 
ledge. This seems replaced by a considerably larger, at the same time 
much more rare, species from the ledge and higher. The span of time 
represented by the Bison basin faunas may have witnessed the extinc- 
tion of such forms as Promioclaenus (which includes much that had 
been grouped before under Ellipsodon), Protoselene, and Litomylus. 

It is not known to what extent the times of Caenolambda and Tita- 
noides may have overlapped. Typical materials of each were found 
as single specimens at the saddle and Titanoides localities respec- 
tively. Undetermined pantodont tooth fragments were collected at all 
localities. 


AGE AND CORRELATION OF THE FAUNAS 


In consideration of the age or ages represented by the Bison basin 
faunas we may deal first with that represented at the saddle locality, 
rather clearly the oldest of the four levels. In regarding this as Tif- 
fanian somewhat greater emphasis is given to the appearance of forms 
known to characterize later horizons than to the presence or survival 
of older genera. For example, Plesiadapis, Thryptacodon, Litolestes, 
Gidleyina, and Phenacodus may be regarded as Tiffanian in first ap- 
pearance, whereas Pronothodectes, Tricentes, Claenodon, Promioclae- 
nus, Litomylus, Haplaletes, and Protoselene have been rather generally 
thought to be Torrejonian. Species represented of certain of the lat- 
ter genera are not clearly separable from those of the Torrejonian 
levels in the Nacimiento and Fort Union and might be regarded as 
long lived, but others in this group such as Pronothodectes simpsoni, 
Claenodon acrogenius, Litomylus scaphicus, and Haplaletes pelicatus 
are distinctly advanced. 

The interpretation that there is an admixture of materials of rather 
different horizons at the saddle locality, though not impossible, may 
be discarded as a serious possibility inasmuch as nearly all the genera 
of both aspects are found together at the ledge locality where collect- 
ing was limited to a narrow zone above a ledge and very near the top 
of the escarpment. Moreover, among the Tiffanian genera repre- 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN II 


sented at the saddle an early stage of development seems clearly indi- 
cated in Plesiadapis jepseni, Thryptacodon belli, and the material re- 
ferred to Gidleyina wyomingensis, as well as in the close approach 
that Phenacodus? bisonensis makes to Tetraclaenodon. 

With due regard to the presence of forms of older aspect in the 
fauna an age assignment of early or lower Tiffanian rather than later 
Torrejonian seems indicated. This is further supported by the re- 
semblance or similarity that persists between the Bison basin faunas, 
although above the saddle and particularly above the ledge locality 
the Torrejonian aspects appear to be lost. Direct comparison of the 
saddle fauna with that of the Melville is difficult because of the spar- 
sity of Carnivora and condylarths so well represented in the Bison 
basin; nevertheless the saddle level may not be much older than the 
Melville, generally regarded as lower Tiffanian. 

The fauna from the Titanoides locality is rather limited, but evi- 
dence is seen for a somewhat more typical Tiffanian stage. The pres- 
ence of Titanoides primaevus suggests a definite relationship only with 
a horizon in the type Fort Union in western North Dakota, although 
somewhat smaller forms from the Melville and Silver Coulee have 
been referred to this genus, as well as “Sparactolambda”’ looki from 
the DeBeque beds. A large species of Plesiadapis regarded as closest 
to P. fodinatus from the Silver Coulee level of the Polecat Bench se- 
ries suggests an equivalent horizon in Tiffanian time, but the Bison 
basin form in at least one individual retains a second lower premolar 
not seen in any of the typical P. fodinatus material. Material close to 
Plesiadapis jepseni also occurs at the Titanoides locality, suggesting 
a closer tie with the older levels in the Bison basin. The Thryptacodon 
here is evidently to be compared closely with that from the Tiffany 
beds in Colorado. The presence of Claenodon cf. ferox is not signi- 
ficant in view of the still later occurrence of this genus in beds of 
Clarkforkian age elsewhere. Haplaletes serior would possibly suggest 
a comparatively early horizon, but this species is so much larger than 
that from the Lebo, or even than the form described from the saddle 
locality in the Bison basin, that its significance seems lost. Litolestes 
is a Tiffanian genus here represented by a species which appears to 
differ only in being of larger size than that from the Melville and 
much larger than the Silver Coulee genotype. Gidleyina and the mate- 
rial of the comparatively large form of Phenacodus? at the Titanoides 
locality suggest little other than Tiffanian. 

From the foregoing it would seem that the Titanoides locality fauna, 
in the absence of forms of Torrejonian aspect, might be regarded as 
somewhat higher in the Tiffanian than Melville but, from considera- 
tion of the primates only, possibly not so late as Silver Coulee. 


12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Except for their intermediate positions, no particular additional 
evidence is forthcoming from the ledge and west-end locality faunas, 
other than that the ledge would seem almost certainly closer to the 
saddle than to the Titanoides locality in age. 


SYSTEMATIC DESCRIPTION OF VERTEBRATE REMAINS 
REPTILIA 
SAURIA 
ANGUIDAE 


The only nonmammalian specimens encountered during the collect- 
ing were four fragmentary dentaries, two portions of maxillae, and a 
premaxilla of a lizard. These were examined by Dr. David H. Dun- 
kle and recognized as belonging to a small anguid type. The genus 
represented could not be determined from the material at hand, but 
Peltosaurus has been recognized in horizons as early as Lance and 
Fort Union. Nevertheless Glyptosaurus and Xestops also include 
diminutive species and these genera are known in the Eocene. All 
but one of the specimens came from the saddle locality. A single 
fragmentary dentary was found at the Titanoides locality. 


MAMMALIA 
MULTITUBERCULATA 


PTILODONTIDAE 


Although the multituberculates appear to be comparatively rare, to 
judge by the frequency with which their remains are encountered, 
nevertheless they must have been highly diversified, because each of 
the four fragmentary specimens known evidently represents a different 
form. The materials in each case are too incomplete to indicate with 
certainty the genus represented but, of the forms tentatively identified, 
two suggest Torrejonian and two Tiffanian, although three of the spec- 
imens came from the small saddle locality discovered by Dr. R. W. 
Brown. The fourth specimen, that compared with the Tiffanian 
Ectypodus musculus, came from a short distance away but regarded 
as the same stratigraphic horizon as the saddle. 


Cf. PTILODUS MONTANUS Douglass, 1908 
Plate 1, figure 1 


A relatively large ptilodont multituberculate is represented by a 
single incomplete left Py, U.S.N.M. No. 20877. There is no certainty 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 13 


that the form represented is Ptilodus, as no other portions of the 
dentition were found and the anterior margin of the tooth was broken 
away. In size and outline, as well as in the spacing of the serrations, 
of which there were at least 13, the tooth rather closely resembles P, 
in Ptilodus montanus. The preserved portion measures 8.1 mm., but 
estimated from complete specimens of P. montanus this tooth in its 
entirety would have been about 8.8 or 8.9 mm. long and within the 
upper limit of measurements for P. montanus. 


Cf. ECTYPODUS MUSCULUS Matthew and Granger, 1921 
Plate 1, figure 3 


A small jaw fragment with M, and the alveoli for M., U. of Wyo. 
No. 1105, would appear to represent Ectypodus musculus. The size 
of the included molar, 2.5 by 1.25 mm., is near that given by Granger 
and Simpson (1929, p. 655) for E. musculus, although the Wyoming 
specimen would appear to be about a quarter of a millimeter broader. 
The cusp formula, 9: 5 or 6, is comparable to the 8: 6 cited by Jepsen 
(1940, p. 307) as well as by Matthew and Granger, particularly as 
one of the outer cusps in No. 1105 is scarcely distinct. This tooth 
may well belong to the form represented by the Py, U.S.N.M. 
No. 20878, compared below with Ectypodus hazeni but is appreciably 
shorter than the 3.2 mm. cited by Jepsen for the length of M, in 
E. hazeni, although the cusp formula for this tooth is the same as that 
for E. musculus. 


Cf. ECTYPODUS HAZENI Jepsen, 1940 


Plate 1, figure 2 


An isolated though complete P,, U.S.N.M. No. 20878, compares 
very closely to this tooth in the Silver Coulee Ectypodus hazeni. It 
resembles this form in the size (5 mm. long) and outline of the tooth 
but has only 11, or possibly 12, serrations rather than the 13 listed 
by Jepsen (1940, p. 307). No. 20878 also resembles P, in Mimetodon 
churchilli, which is indicated as having 12 serrations, but the tooth 
has perhaps a somewhat more convex profile, with the straight pos- 
terior section a relatively shorter part of the entire profile. The an- 
terior margin of the tooth is deeply notched and pocketed, suggesting 
the presence of P;, but the absence of other associated material pre- 
cludes certain generic identification. 


I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Cf. ANCONODON RUSSELLI (Simpson), 1935 
Plate 1, figure 4 


Anconodon russelli may be represented by a fragment of the right 
mandibular ramus showing the root portion of the incisor and P,, 
U. of Wyo. No. 1065. The preserved premolar is close in size to that 
referred above to Ectypodus hazeni, but its profile is slightly more 
convex dorsally and has a long straight front edge more as in An- 
conodon. FP, is 5.2 mm. in length, which is about midway in the 
range of 4.9 to 5.4 mm. given by Jepsen (1940, p. 291) for Anconodon 
russelli, The number of serrations is not certainly determined but 
would appear to be about 13 or 14. Fourteen serrations are noted for 
several of the Gidley Quarry specimens from the Crazy Mountain 
Fort Union, but 15 or 16 prevail in the Rock Bench material accord- 
ing to Jepsen. 


MARSUPIALIA 


DIDELPHIDAE 
PERADECTES ELEGANS Matthew and Granger, 1921 
Plate 2, figure 6 


A rather well preserved right mandibular ramus with the posterior 
three molars, U. of Wyo. No. 1104, corresponds so closely in direct 
comparison with the type of Peradectes elegans from the Tiffany of 
Colorado that there seems no doubt that the species are the same. 
Lower molars are, of course, amazingly conservative in didelphids, 
but the near identity in various measurements of the teeth in the Bison 
basin jaw and the type leave no alternative but recognition of this 
species in the upper Paleocene of Wyoming. Measurements of the 
teeth have been incorporated below with those of the following species 
believed to be distinct. 


PERADECTES PAULI,? new species 
Plate 2, figures 4, 5 


Type.—Portion of left mandibular ramus with last two molars, 
U.S.N.M. No. 20879. 

Horizon and locality—Bison basin Tiffanian, saddle locality, below 
south rim of Bison basin, sec. 28, T. 27 N., R. 95 W., Fremont 
County, Wyo. 

Specific characters—Appreciably smaller teeth than Peradectes 
elegans and lower molars with outer cusps slightly less elevated and 
talonids relatively more abbreviated. 


2 Named for Paul O. McGrew. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 15 


Discussion—In addition to being of smaller size than Peradectes 
elegans, it was noted that lower molars of P. pauli show a protoconid 
which, though higher than the paraconid and metaconid, is not so 
much elevated with respect to these cusps. Also, the hypoconid is a 
comparatively lower cusp. Moreover, the talonid basin is relatively 
both a little narrower and shorter than in P. elegans and the entoconid 
and hypoconulid a little less widely separated. 

With the distinctions between species of didelphid marsupials gen- 
erally including little more than size so far as characters of lower 
molars are concerned and with very limited information on the varia- 
bility of observed structural differences in these earlier forms, con- 
clusions as to generic identity based on lower molars are not entirely 
satisfactory. As to whether the cited differences preclude reference 
of P. pauli to Peradectes there is no certainty. Although the some- 
what lower protoconid might suggest Peratherium, this is not sup- 
ported by the more abbreviated talonids, and the entoconid is not 
nearly so prominent. Furthermore, Peratherium has not been cer- 
tainly recognized in pre-Eocene deposits. In the absence of any repre- 
sentation of the upper dentition, about which on a generic level evi- 
dence of a somewhat more satisfactory nature has been developed 
(Simpson, 1935a), the species is assigned to Peradectes. 


MEASUREMENTS IN MILLIMETERS OF LOWER TEETH IN SPECIES OF Peradectes 
Peradectes elegans Peradectes pauli, n. sp. 
A.M.N.H. 


U.S.N.M. 
No. 17376 U.of Wyo. No. 20879 U.S.N.M. 
(type) No. 1104 (type) No. 20880 


M,, anteroposterior diameter ............ 1.75 gah chase 1.6 
transverse diameter of trigonid...... 0.9 mise dare 0.7 
transverse diameter of talonid....... 0.95 nes nee 0.75 

M2, anteroposterior diameter ............ 1.75 1.7 aa 
transverse diameter of trigonid...... 1.0 1.0 Shs 0.8 
transverse diameter of talonid....... 1.0 1.0 Neve 

M:, anteroposterior diameter ............ 1.75 1.7 1.55 
transverse diameter of trigonid...... 1.05 1.05 0.95 
transverse diameter of talonid....... 1.05 1.0 0.8 

Ma, anteroposterior diameter ............ 075 1.65 1.6 
transverse diameter of trigonid...... 0.95 0.95 0.85 
transverse diameter of talonid....... 0.8 0.7 0.6 

INSECTIVORA 
LEPTICTIDAE 


Simpson (1937b) presented a logical arrangement of the earlier 
leptictids which went far toward clarifying the complexity and di- 
versity of these forms. Nevertheless, a review of the various mate- 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


rials in connection with the study of the leptictid form represented in 
the Bison basin Paleocene has indicated the need for certain further 
modification. 

Attention (Gazin, 1952) was called to the rather distinctive char- 
acters observed in the type of Diacodon alticuspis, and I am now con- 
vinced that Cope’s Ictops bicuspis should not have been referred to 
Diacodon and that Matthew’s earlier disposition of this species under 
the name Palaeictops should be revived. As well as Palaeictops bicus- 
pis (Cope), this genus apparently should include Palaeictops tauri- 
cineret (Jepsen) and Palaeictops pineyensis (Gazin) from among 
the lower Eocene forms, and possibly also Palaeictops minutus (Jep- 
sen) from the Silver Coulee (Tiffanian) Paleocene. 

The genus Prodiacodon was named by Matthew as a subgenus re- 
placing Palaeolestes (preoccupied) for the species P. puercensis of 
the Torrejon horizon. This form, though generically distinct, is, I be- 
lieve, more closely allied to Palaeictops bicuspis than to typical Dia- 
codon or D. alticuspis. In 1935 Simpson (see 1937b) described Pro- 
diacodon concordiarcensis from the upper Lebo (Torrejonian) and 
expressed some doubt as to the correctness of referring it to that genus. 
In view of the somewhat later but closely related form encountered in 
the Bison basin fauna, and of the particular characteristics, rather 
generally overlooked, of Diacodon alticuspis, I am placing both the 
Lebo and Bison basin forms in Diacodon. These then become Dia- 
codon concordiarcensis (Simpson) and Diacodon pearcei, new species. 


DIACODON PEARCEI,? new species 
Plate 1, figure 6 


Type.—Left ramus of mandible with P;-M,, U.S.N.M. No. 20970. 

Horizon and locality—Bison basin Tiffanian, small saddle below 
south rim of Bison basin, sec. 28, T. 27 N., R. 95 W., Fremont 
County, Wyo. 

Specific characters.—Diacodon pearcei closely resembles Diacodon 
concordiarcensis (Simpson) from the Crazy Mountain Fort Union 
(upper Lebo) in the structure of the teeth, but is distinctly larger, 
about intermediate between D. concordiarcensis and D. alticuspis. 
P, is seen to be about 14 percent longer and 27 percent wider than in 
D. concordiarcensis and about 20 percent shorter and 22 percent nar- 
rower than D. alticuspis. The paraconid of this tooth is higher than 
in the Lebo species. 


8 Named for Franklin L. Pearce. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN vy 


Discussion—The closeness of the resemblance between D. con- 
cordiarcensts and D. pearcei convinces me that whatever disposition 
is made of one, so far as the genus is concerned, the other must like- 
wise be assigned. These two are characterized by comparatively high 
trigonids and abbreviated molar talonids, associated with a progres- 
sive Py, with a likewise abbreviated though basined talonid. The com- 
bination of characters seems most closely approximated in the San 
Jose and Knight material of Diacodon alticuspis. In Palaeictops Ps 
is comparable in progressiveness, but the talonid in this tooth and in 
the lower molars is broad and long, comprising a greater proportion 
of the tooth crown. The molar trigonids, though also high in certain 
species, are relatively shorter anteroposteriorly. Prodiacodon is like- 
wise characterized by comparatively large talonids. A small cuspule 
about halfway down the posterior slope of P, and P; in Prodiacodon 
puercensis and much better developed in species of Palaeictops is 
not seen on the P, of D. concordiarcensis or the P; of D. pearcei. On 
the other hand, an anterior cuspule, well developed and high on these 
teeth in the latter two species, is absent or weak and low in Prodtaco- 
don and Palaeictops. 

Among the other early leptictids, Leptacodon has a less progressive 
P, than Diacodon, with a strong but low paraconid and in particular 
a comparatively weak and more posterior metaconid. Moreover, the 
molar trigonids appear lower and the talonids comparatively larger 
than in Diacodon. Myrmecoboides has a large P, with paraconid for- 
ward as in Diacodon and Palaeictops, though lower; however, the 
greatly elongated talonids exhibited in the molars and P, immediately 
distinguish this genus. The abbreviation of the talonid on P, and the 
lower molars of Xenacodon is suggestive of Diacodon, but P, would 
appear to be distinctly less progressive in that the paraconid is small 
and the talonid not basined. 


MEASUREMENTS IN MILLIMETERS OF LOWER TEETH IN TYPE SPECIMEN 
or Diacodon pearcet, U.S.N.M. NO. 20970 


P:, anteroposterior diameter : greatest transverse diameter .......... 1.8: 1.0 

P,, anteroposterior diameter : transverse diameter of trigonid........ 2.4: 1.4 

M:, anteroposterior diameter : transverse diameter of trigonid........ 2.2: 1.6 
PANTOLESTIDAE 


BISONALVEUS,‘ new genus 


Type.—Bisonalveus browni, new species. 
Generic characters.—Resembling Aphronorus, but P, much smaller 


4 Bison + alveus, basin—for Bison basin. 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


and exhibiting a slightly more noticeable paraconid and a better de- 
veloped and more posterolingual entoconid. Paraconids of lower 
molars higher and more lingual, but hypoconulid of M, and M, in- 
distinct and less prominently projecting on M;. Entoconid on M, and 
M, more forward in position. Small cuspule on crest anterolingual 
to hypoconid on Ms. 

Discussion.—The structure of P, and the general form of the molars 
suggest an alliance of this form with Aphronorus and hence with the 
pantolestids, although the weakness of the hypoconulid would seem 
to negate such a relationship. The molars, though exhibiting com- 
paratively acute cusps, might by themselves have been regarded as 
condylarth. 

P, of Bisonalveus has a slightly better developed paraconid at the 
anterolingual margin of tooth, and a more pronounced and postero- 
lingually located entoconid so that this portion of the talonid crest is 
not so depressed or so nearly oblique. The metaconid is only a little 
lower than the protoconid and slightly posterior to it. The shape of 
these two cusps is rather like that in Aphronorus. 

The lower molars show elevated trigonids, somewhat less so than 
in Aphronorus, but the paraconid is almost as high as the metaconid. 
The paraconid, moreover, is more lingual in position than it is in 
Aphronorus. The talonids of the molars are basined much as in 
Aphronorus, but the arcuate posterior crest of the first two shows 
little or no evidence of a hypoconulid. It may be noted that the 
hypoconulid on molars of Aphronorus is relatively weak in compari- 
son with middle Eocene Pantolestes but is nevertheless clearly de- 
fined. In Bisonalveus, furthermore, the entoconid has a more forward 
position on the crest of the talonid in M, and M, than in Aphronorus. 
In M, the entoconid and hypoconulid are closer together. An- 
terior to the hypoconid on Mz; (only) there is a distinct cuspule, much 
as seen in some leptictid molars. 

Bisonalveus lower molars differ from those of Bessoecetor in much 
the same way as they do from those of Aphronorus. The fourth lower 
premolar, however, is entirely different from that of Bessoecetor and 
has more nearly the form of that in Aphronorus. 


BISONALVEUS BROWNI,° new species 
Plate 1, figure 5 
Type.—Left ramus of mandible with Py-Ms, U.S.N.M. No. 20928. 
Horizon and locality—Bison basin Tiffanian, saddle locality, south 


5 Named for Dr. Roland W. Brown. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN Ig 


rim of Bison basin, sec. 28, T. 27 N., R. 95 W., Fremont County, 
Wyo. 

Specific characters——In size of teeth Bisonalveus browni is dis- 
tinctly smaller than Aphronorus fraudator Simpson. The length of 
the lower molar series is about 13 percent shorter, whereas P, is about 
45 percent less in length, and 36 percent narrower. Specific characters 
are not otherwise distinguished from those of the genus. 

In addition to the type there are in the collections of National Mu- 
seum a fragmentary left mandibular ramus with M; (No. 20929) and 
an isolated P, (No. 20930). A jaw fragment with a much worn P, 
and M, in the University of Wyoming collections (No. 1067) may 
also represent this species. 


MEASUREMENTS IN MILLIMETERS OF LOWER TEETH IN TYPE SPECIMEN 
oF Bisonalveus browni, U.S.N.M. NO. 20928 


Length of lower cheek tooth series, Ps-Ma, incl..............ceeeee0ee 9.5 
Iwength ot lower molar series, Ms-DMe Incl... 0. cece sccetoceceescuc 7.5 
P,, anteroposterior diameter : greatest transverse diameter........... 2 2ehies 
M, anteroposterior diameter : transverse diameter of trigonid......... IS he, 
M; anteroposterior diameter : transverse diameter of trigonid......... 26°20 
Ms anteroposterior diameter : transverse diameter of trigonid......... 2601.5 
PRIMATES 
PLESIADAPIDAE 


In numbers of jaws and, to a lesser extent, maxillae, the plesia- 
dapids are surprisingly well represented. Not less than four species, 
presumably divided between two genera, are recognized. Not all, 
however, are found associated at any one locality. The most primi- 
tive, Pronothodectes cf. matthewi, and presumably the most progres- 
sive, Plesiadapis cf. fodinatus, are not found together, but both are 
associated with the forms or variants of the forms that might be re- 
ferred to as intermediate in development, Pronothodectes simpsoni 
and Plesiadapis jepseni. Pronothodectes is generally regarded as a 
forerunner of Plesiadapis, which it undoubtedly is, but their occur- 
rence together here is unquestionable in two restricted localities. 
Pronothodectes cf. matthewi and a small variant of Pronothodectes 
simpsoni are found associated with Plesiadapis jepseni at the saddle 
locality and typical Pronothodectes simpsoni is associated with Plesia- 
dapis jepsent and Plesiadapis cf. fodinatus (a single specimen of a 
small individual) at the stratigraphically much restricted ledge locality. 
At the more westerly and probably higher localities Plesiadapis cf. 
fodinatus is found with scant material referred, perhaps questionably, 
to Plesiadapis jepseni. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


PRONOTHODECTES, cf. MATTHEWI Gidley, 1923 
Plate 2, figures 1, 2 


A decidedly small plesiadapid is represented in the collections by 
four lower jaw portions and a maxillary fragment. One of the jaws, 
U.S.N.M. No. 20758, includes P,-M, and part of M3, and a second 
specimen, U. of Wyo. No. 1062, exhibits P;-M,. The maxillary por- 
tion, U. of Wyo. No. 1099, with two molars, is very tentatively re- 
ferred but would seem to belong with this material rather than the 
larger Pronothodectes simpson. 

There is little doubt that the genus represented is Pronothodectes 
rather than Plesiadapis, as the dental formula includes all the lower 
premolars. The species represented is very close to the small Pro- 
nothodectes matthewi, which Gidley (1923) described from the Mon- 
tana Fort Union. The lower premolars and molar trigonid cusps, 
though sloping, are less so than in Plesiadapis material of the P. anceps 
type, and the trigonids are moderately compressed anteroposteriorly 
as in Pronothodectes matthewi. Only a slightly greater transverse 
width to the teeth was noted in the Bison basin material. 

All the specimens of this form were obtained from the saddle lo- 
cality at the south rim of the Bison basin, associated with Pronotho- 
dectes cf. simpsoni and Plesiadapis jepseni, but not the advanced 
Plesiadapis cf. fodinatus. 


MEASUREMENTS IN MILLIMETERS OF TEETH REFERRED TO 
Pronothodectes matthewi GIDLEY 


U. of Wyo. 
No. 1099 
Me SANterOpOSLEMIOr GiamMeleN & f.ciaarcs ass ais-aicarteiaraisiainjels’, Peie wien 2.5 
Mes aAtLErOGOSLCHION |. GIAITICEEN, «5 «.u, c1njsten ui slsie a'nin's ostticle on tarseieais 2.5 
U.S.N.M. U. of Wyo. 
No. 20758 No. 1062 
P;, anteroposterior diameter : transverse diameter .... ...... TGS Ley 
P,, anteroposterior diameter : transverse diameter .... 2.1: 1.7 2.2: 1.8 
M,, anteroposterior diameter : transverse diameter 
Gate OLTClere rc ayctoe te Rene Mer hehe Hotete Sit ote ete als cxtere hele 2.4: 2.0 25) 201 
M:, anteroposterior diameter : transverse diameter 
CHER Ca OMNIA pte tania etacare octets chen erarhcteer aie ierociceromtoe aera 255,223 0 nes cite 
M,,. anteroposterio® Giamieter 60/05, <ic,3, «00's os -aiew awikwlom nie Beg | fase dele 


@ Approximate. 
PRONOTHODECTES SIMPSONI,® new species 
Plate 3 


Type.—Right ramus of mandible with P,-M;, U.S.N.M. No. 20754. 
Horizon and locality.—Bison basin Tiffanian, ledge locality at south 


® Named for Dr. G. G. Simpson in appreciation of his work on the early 
primates. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 21 


rim of Bison basin, W4 sec. 28, T. 27 N., R. 95 W., Fremont County, 
Wyo. 

Specific characters ——Size nearly intermediate between Pronotho- 
dectes matthewit and Plesiadapis gidleyi, closer to the latter. Py, 
pressed close to incisor, posterior lower premolars and molar trigonids 
of more inflated appearance than in P. matthew. 

Discussion—The more typical materials of this species are from 
the ledge locality about a quarter of a mile west of the saddle locality 
and include about nine jaws besides the type in the collections of the 
U. S. National Museum. About four specimens in each of the Na- 
tional Museum and University of Wyoming collections from the 
saddle locality would seem to represent a variant somewhat smaller 
in size, although evidently closer to this species than to Pronotho- 
dectes matthewi. None of the material of this species was found at 
the more westerly and possibly higher horizons in the basin. 

Like the material referred to Pronothodectes matthewi, that of this 
species would appear by definition to be Pronothodectes rather than 
Plesiadapis, because in all specimens where the dental formula can 
be determined all the lower premolars were represented, P,; and P; 
being, of course, single-rooted teeth as in P. matthewi. P. simpsoni, 
as noted in the diagnosis, is characterized by much larger teeth, about 
18 to 31 percent larger in length of lower molar series in materials 
from the ledge locality, and possibly as low as 12 to about 24 percent 
larger than P. matthewi in the materials referred to P. simpsoni from 
the saddle locality. 

Pronothodectes simpsoni differs from P. matthewi, in addition to 
its greater size, by exhibiting teeth of a more Plesiadapis-like appear- 
ance. This is noticeable in the more typical materials from the ledge 
locality in the comparatively inflated appearance of the cusps. Per- 
haps it is more noticeable in the trigonid, which is distinctly less an- 
teroposteriorly compressed in M, and M;. The variant from the sad- 
dle locality overlaps in size range that represented at the ledge locality 
and is less obviously different from P. matthewi in size and appear- 
ance of the cusps, but would seem to be closer to P. simpsoni. A 
lower jaw (U. of Wyo. No. 1057, pl. 3, fig. 2) of P. cf. simpsoni with 
P, to M, from the saddle locality exhibits a disproportionately long 
M, (buccally incomplete), but other specimens with M, from the 
saddle show this tooth to be normally proportioned. M, and Mz, in 
No. 1057 are scarcely distinguished from these two teeth in the smaller 
specimens of P. simpsoni from the ledge (i.e., U.S.N.M. No. 20770, 


pl. 3, fig. 3). 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


MEASUREMENTS IN MILLIMETERS OF LOWER TEETH IN SPECIMENS OF 
Pronothodectes simpsom 


U.S.N.M. 
No. 20754 U.S.N.M. U. of Wyo. 
(type) No. 20770 No. 1057 * 
Length of lower molar series............. 9.8 8.9 9.4 
Ps, anteroposterior diameter : transverse 
Giameter Worcinescameiee tise cule eons Wie BS seiQuire Matec 22°20 
M,, anteroposterior diameter : greatest 
transverse diameter .............+20+ 2.8: 2.4 2.6: 2.3 2.5925 
Ma, anteroposterior diameter : greatest 
transverse diameter .........sssee00: 3.13,2.6 20-25 B77 OG 
Ms, anteroposterior diameter : greatest 
transverse diameter <.c(. acti. cca ses 4.3:2.5 ZO 22 BAA. 


* Small variant but with large Ms 


PLESIADAPIS, cf. FODINATUS Jepsen, 1930 
Plate 2, figure 3 


Three specimens in the collections of the National Museum and 
three in those of the University of Wyoming are believed to repre- 
sent the comparatively large plesiadapid that Jepsen (1940) described 
from the Silver Coulee horizon in the Polecat Bench series of north- 
western Wyoming. Represented among these are P, to Mg, and only 
one (U. of Wyo. No. 1085) of the specimens is a maxilla, exhibiting 
M, and M;. The specimens, with one exception, are from the more 
westerly localities and probably higher stratigraphically than the sad- 
dle locality and possibly higher than the ledge. One jaw with teeth 
a trifle smaller than in the others came from the ledge but is believed 
to be closer to Plesiadapis fodinatus than it is to the new form, 
P. jepseni. 

The teeth in the Bison basin materials referred to P. fodinatus cor- 
respond so closely to those of the Silver Coulee form that there would 
seem to be no serious doubt as to the correctness of the assignment. 
The resemblance is very close in all proportions of the molar teeth, 
and like P. fodinatus the teeth do not show such markedly sloping 
labial walls as in correspondingly large Plesiadapis rex and related 
P. anceps. It was noted, however, that in one of the jaws, which has 
preserved the alveoli of the anterior cheek teeth, a small P, had been 
present, although there was no evidence of a P;. Pz is not present in 
the type or other observed materials of P. fodinatus from the Prince- 
ton Quarry but is present in P. gidleyi and almost always present, 
though not invariably so, in the material described below as the new 
species, Plesiadapis jepseni. Its presence in the Bison basin jaw com- 
pared with P. fodinatus may not be significant. There is, moreover, a 
suggestion in this particular jaw of somewhat smaller premolars and a 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 23 


shorter diastema behind the incisor than in typical P. fodinatus. The 
length of the diastema, though, is uncertain as the bone is incomplete. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMENS OF Plesiadapis, CF. 
fodinatus JEPSEN 


U. of Wyo. 
No. 1085 
M®*, anteroposterior diameter : greatest transverse diameter.... 3.8: 5.9 
Ma Jeredtest trausverserdiameter ssc lsisildeedild sesinsioonigeetss Js 5.5 
U. of Wyo. U.S.N.M. U.S.N.M. 
No. 1082 No. 20783 No. 20784 
Ps, anteroposterior diameter : greatest 
transverse diameter 20) i jcebet ede ss magmas ir ity Ceti! SAAS ee 
M;, anteroposterior diameter : greatest 
tLals verse .CiamMeters-.7..<.sjcicie ds.0 seiemis's 3.6: 2.9 B2IZG *\ dow hiesus 
Mz, anteroposterior diameter : greatest 
THALISVERSE \CIAMELET © o/s nis cineis oa ties ales wise: 3.6: 3.4 3.90: 3.6 
Ms, anteroposterior diameter : greatest 
transverse atameter 2 7S ce ceene Siocon lt ode 5.5: 3-5 


PLESIADAPIS JEPSENI,’ new species 
Plate 4 


Type.—Left ramus of mandible with P,-M;, U.S.N.M. No. 20760. 

Horizon and locality.—Bison basin Tiffanian, ledge locality at south 
rim of Bison basin, W3 sec. 28, T. 27 N., R. 95 W., Fremont County, 
Wyo. 

Specific characters —Close in size to Plesiadapis gidleyi and Plesi- 
adapis anceps. P, almost always present and hypoconulid portion of 
talonid of M; broad as in P. gidleyi. Lower teeth relatively broad 
with outer walls decidedly sloping as in P. anceps and P. rex. Conule 
weak to scarcely discernible on lingual slope of primary cusp of P* 
but prominent on P*. Mesostyle absent or very weak on upper molars, 
M® slightly more expanded posterointernally than in P. anceps. 

Discussion.—Plesiadapis jepsent is one of the better represented 
forms in the Bison basin collection, exceeded in number of specimens 
only by Phenacodus? bisonensis and Gidleyina wyomingensis. About 
23 specimens, mostly lower jaws, are for the most part divided be- 
tween the saddle and ledge localities below the south rim of the basin. 
Two, however, came from the locality at the west end of the basin and 
two from near the Titanoides primaevus locality in the southwestern 
part of the basin. Three mandibular portions and one maxilla are 
in the collections of the University of Wyoming. 


7Named for Dr. Glenn L. Jepsen in appreciation of his work on the 
Plesiadapidae. 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


A small part of this collection, that secured by Dr. Roland Brown 
and others, was originally cataloged and described by me in a prelimi- 
nary unpublished manuscript, as well as in a report to the U. S. Geo- 
logical Survey, as representing Plesiadapis anceps, which it most 
nearly resembles in the general structure of the lower molars. The 
resemblance is noticeable in the relative breadth of the teeth and 
strongly sloping outer walls, a feature also noted in the larger Plesia- 
dapis rex. The protoconid in particular has a long posterolateral slope 
quite unlike P. gidleyi and P. fodinatus or the later P. dubius and 
P. rubeyt. This slope is characteristic of the posterolateral wall of 
the primary cusp in premolars as well as in the molars of P. jepseni 
and P. anceps. Plesiadapis jepseni is unlike P. anceps and more nearly 
resembles P. gidleyi in the expansion of the posterior portions of the 
third upper and lower molars. The posterolingual portion of M°, 
though somewhat more expanded than in P. anceps, is possibly not 
so distinctive in this respect as P. gidleyi; however, the third lobe or 
hypoconulid portion of M; is generally as much expanded as in P. gid- 
leyi. Moreover, in about eight of the lower jaws in which the presence 
or absence of P, can be determined, it is certainly missing in only 
one. This tooth is apparently not present in P. anceps, typical 
P. fodinatus, and later plesiadapids. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMENS OF 
Plesiadapis jepseni 


U.S.N.M. U.S.N.M. 
No. 20781 No. 20780 
Length of cheek tooth series, P*-M*...............c00. 12.00 VP ae 
Length of upper molar series, M*-M®............00-00- S.gavi 1 I 
P*, anteroposterior diameter : greatest transverse 
SRIAITIGH OES dustin shay el ieiin(- ays oaioas sheso ate ova fare eaimeys cat oka areal 2.0.2.3 eT 2E2G 
P*, anteroposterior diameter : greatest transverse 
CIATELET ater ces torehetels op ascPete bicvel oravcievatahs elcterey reroute rast 22 Stk 22353 
M’, anteroposterior diameter : greatest transverse 
IA TEL ete ts mereters Cit itt otc a chk eis, oe a clare Reha eres 2.9: 4.1 2.8: 4.0 
M’, anteroposterior diameter : greatest transverse 
FVEN a2 2) SUR a TERR AE er AERP pice eB fee 3.0: 4.7 3.0: 4.6 
M®*, anteroposterior diameter : greatest transverse 
CRAIC ERNE sretarg et erieln 1b. ote Wrhle ale a0 ays oiee «ppare Sie aieesha lens PS SG Ts Vries 
U.S.N.M. 
No. 20760 U.S.N.M. 
(type) No. 20586 
Length of cheek tooth series, Ps-Ms............000200 T2ZOet Dakar 
Length of lower molar series, Mi-Ms.............00005 LOM WH? ok ave 
Ps, anteroposterior diameter : greatest transverse 
CHATTIGEER) ts cM Stole, scat eevee 00: wine. vi ale ciave eis Srahe Soe Cie ea ee ener 2.4: 1.9 


Ps, anteroposterior diameter : greatest transverse 
IAMeLER certs tie tee aie cletean be sree SR ene TOR 2.4:2.8 25124 


NO. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 25 


M,, anteroposterior diameter : greatest transverse 


CSAUNVISE Cert oppor ey ov che oants for eke ededaler sbaleeetwloyas |e ish steve el 'e.0 3.0: 3.0 Z'2213:0 
M2, anteroposterior diameter : greatest transverse 
RUTATNCLEE, watctavctotsersysteteorsicieh dois Ore ate Gishsiee/eiapeie! Sores B39 2 a sis2 
M;, anteroposterior diameter : greatest transverse 
CHIAITICLEI BO IEEE acs rater ante Beeld tree Metlre das itie aretents AAP Bol WATE ES te 
CARNIVORA 
ARCTOCYONIDAE 


Creodonts are well represented in the collections, comprising a 
diversity of forms not hitherto recorded in the Tiffanian. Most of 
the forms are of arctocyonid types, and among these are species of 
Tricentes and Claenodon, suggesting affinities with the earlier Tor- 
rejonian faunas, together with Thryptacodon better known in later 
horizons. Chriacus, having a comparatively great range in geologic 
time, is represented by a species rather similar but possibly a little 
more progressive than that of the Torrejon. So far as Claenodon is 
concerned, although the species are difficult to distinguish from those 
of the Torrejon, undescribed material of the genus has been obtained 
from Paleocene deposits as late as Clarkforkian, and the presumably 
descendant Anacodon is, of course, found in the lower Eocene. 


OxyYCLAENINAE 
TRICENTES FREMONTENSIS,® new species 
Plate 5, figure 4 


Type.—Left ramus of mandible with M,-M3, U.S.N.M. No. 20582. 

Horizon and locality—Bison basin Tiffanian, saddle locality at 
south rim of Bison basin, sec. 28, T. 27 N., R. 95 W., Fremont 
County, Wyo. 

Specific characters——Teeth close in size to those of Tricentes sub- 
trigonus but anterior portion of lower dentition reduced, with trigonid 
of M, narrow and premolars smaller. 

Discussion—The above new specific name is proposed with some 
hesitancy, as the Torrejon species, Tricentes subtrigonus, shows an 
astonishing amount of variation in characters of the lower teeth. Vari- 
ation in size, relative proportion of teeth, and cusp development makes 
any attempt at detailed comparison nearly futile. Marked variability 
was also noted in the material of Tricentes fremontensis. Neverthe- 
less, Ps and P,, as observed in U.S.N.M. No. 20584, are smaller than 
in any of the specimens of T. subtrigonus I have examined, and in 


8 Named for Fremont County, Wyo. 


26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


the dozen or more other specimens of T. fremontensis at hand the 
relative narrowness of the anterior molars, particularly the trigonid 
of M,, may be distinctive. The type, U.S.N.M. No. 20582, is a com- 
paratively large individual with teeth relatively wider than nearly all 
others from this locality. Their width, however, in proportion to their 
length (more evident in M,), though matched in certain individuals of 
T. subtrigonus, is rather less than the average in the Torrejon ma- 
_ terials observed. Other specimens of T. fremontensis appear to be 
outside the range of T. subtrigonus in this respect. Moreover, the 
paraconid on M, and Ms is placed low on the trigonid of lower molars 
in T. fremontensis and is weaker than generally seen in T. subtri- 
gonus. In none of the posterior lower molars of the Bison basin form 
is this cusp so conspicuously developed as it is in so much of the 
Torrejon material. It should be noted, however, that the difference 
is one in average for the material at hand, as teeth of T. subtrigonus 
can be found in which there is scarcely a trace of the paraconid on Ms. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMENS OF 
Tricentes fremontensis 


U.S.N.M. 
No. 20584 
P;, anteroposterior diameter : transverse diameter ..........ee.00. 4.0: 2.4 
P,, anteroposterior diameter : transverse diameter ..............0- 4.5: 2.9 
U.S.N.M. 
No. 20582 
(type) 
M,, anteroposterior diameter : transverse diameter of trigonid...... 6.2:3.8 
IM; transverse. diameter, of ttalomidstee wieictitehers aves c o1epeisheie.e <fereioisieiavete 4.5 
M;, anteroposterior diameter : greatest transverse diameter......... 6.4: 4.9 
M;, anteroposterior diameter : greatest transverse diameter......... 6.6: 4.3 


CHRIACUS, near C. PELVIDENS (Cope), 1881 
Plate 5, figures I, 2 


About four fragmentary jaws of a species of Chriacus are included 
in the collections of the National Museum. Unfortunately, only one 
of these (U.S.N.M. No. 20983) has as many as two complete teeth. 
The form represented is undoubtedly close to Chriacus pelvidens of 
the Torrejon, with the anteroposterior diameter of the lower teeth 
about the same as in that species. Their width, however, in two of 
these is a little greater than in any of the C. pelvidens material at 
hand. Moreover, the metaconid on P,, in one of the two specimens 
that retains this tooth, is distinctly better developed, as it is in small 
Spanoxyodon latrunculus. Although a distinct species of Chriacus, 
or even possibly Spanoxyodon, may well be represented here, the evi- 
dence is not conclusive and no satisfactory diagnosis can be made. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 27 


P, in U.S.N.M. No. 20983 measures 6.1 mm. long by 3.7 mm. wide. 
M, in this specimen is 7.3 by 5.4 mm. 

An isolated upper molar, probably M? (U.S.N.M. No. 21003), 
would also appear to represent a species of Chriacus about the size of 
C. pelvidens. The outer styles of this tooth are not noticeably de- 
veloped, but lingually the cingulum carries a prominent hypocone and 
a likewise prominent though less developed protostyle at the antero- 
lingual margin of the tooth. This tooth measures 6.3 mm. long by 
8.8. mm. wide transversely. 


CHRIACUS, sp. 
Plate 5, figure 3 


A single upper molar from the saddle locality, U.S.N.M. No. 210109, 
presumably M?, is much smaller, approximately 25 percent less in 
general proportions than the M® discussed in the foregoing section. 
In size it would appear to be more nearly comparable to Chriacus 
truncatus, approximately that of Thryptacodon belli. The rectangu- 
lar appearance of this tooth and the prominence of the anterointernal 
cusp or protostyle would seem to remove it from consideration as a 
form of Thryptacodon. The tooth measures 5.2 mm. long by 6.3 mm. 
wide transversely. 


THRYPTACODON, cf. AUSTRALIS Simpson, 1935 
Plate 6, figure 5 


A fragmentary left mandibular ramus, including P,, M,, and Ms, 
in the collection obtained by the University of Wyoming (No. 1076) 
from the Titanoides locality, in details of the teeth closely approxi- 
mates that of the species of Thryptacodon named by Simpson (1935c) 
from the Tiffany beds of Colorado. The teeth are a trifle larger than 
in the type as may be seen from the dimension Simpson has given, but 
the rudimentary condition of the metaconid on P, and the prominently 
isolated hypoconulid on Ms; suggest possibly a closer relationship to 
Thryptacodon australis than to T. demari described as new in the 
following section. 

The trigonid of M; in U. Wyo. No. 1076 seems rather broadly 
basined and the paraconid distinctly weak. Moreover, the hypo- 
conulid, in addition to being prominent, is rather distinctly set off 
from the entoconid, and there is a low crest between the entoconid and 
hypoconid. The hypoconulid portion of Msg is believed to be highly 
variable in Thryptacodon, but T. australis material observed shows 
this cusp rather better defined than in much of the Wasatchian 


28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


T. antiquus material exhibiting M;. A slightly more posteriorward 
position for the hypoconulid is indicated also for M,, somewhat remi- 
niscent of Chriacus, though not nearly so distinctive in this respect 
and, of course, the teeth in general are not nearly so high cusped. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMEN REFERRED TO 
Thryptacodon australis SIMPSON, U. OF WYO. NO. 1076 


Ps, anteroposterior diameter : greatest transverse diameter........... 5.06:2:0 
M,, anteroposterior diameter : greatest transverse diameter........... 6.7: 4.6 
Ms, anteroposterior diameter : greatest transverse diameter........... PETE SSS 


THRYPTACODON DEMARI,? new species 
Plate 6, figures 2, 3 


Type.—Right ramus of mandible with P,-M;, U.S.N.M. No. 200985. 

Horizon and locality.—Bison basin Tiffanian, ledge locality at south 
rim of Bison basin, W4 sec. 28, T. 27 N., R. 95 W., Fremont County, 
Wyo. 

Specific characters.—Size a little smaller than Thryptacodon aus- 
tralis. P, isolated by longer diastemata. Metaconid of P, much bet- 
ter developed. Proportions of M, about as in T. australis but pos- 
terior molars comparatively short and broad. Hypoconulid of M; 
variable but may be much reduced. 

Discussion.—About 11 specimens in the National Museum col- 
lections and one or two in the collection made by the University of 
Wyoming are recognized as pertaining to the new species Thryptaco- 
don demari. The lower molar series of this form is only a little shorter 
than in Thryptacodon australis Simpson from the Tiffany beds of 
Colorado, and on the basis of size alone would probably not be dis- 
tinct from that species. In many respects T. demari shows points of 
resemblance to the distinctly more robust appearing T. antiquus. 
The lower premolars, though slender, are well spaced anteriorly, more 
as in T. antiquus. P4, however, has a better developed metaconid than 
in either of these. M, is about the same size as in more nearly con- 
temporary 7. australis, but M, and Ms; are shorter and relatively 
broader. In M,; the shortness may be effected largely by the more 
reduced hypoconulid in some specimens which, somewhat as illus- 
trated by Matthew for the type of 7. antiquus, may be more closely 
joined to the entoconid, and the talonid basin opened posteriorly be- 
tween the hypoconid and entoconid. In the Bison basin specimen 
thought to represent T. australis, and as evident in Simpson’s illustra- 


® Named for Robert DeMar, who aided materially in the collecting of 1953. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 29 


tion of the type, the talonid basin of M; is confined posteriorly by a 
low crest between the hypoconid and entoconid and the hypoconulid 
is sharply separated from the entoconid. It should be noted, however, 
that an approach to this condition is made in certain specimens of T. 
demari, hence possibly not of diagnostic significance. 

Differences from T. antiquus, in addition to the development of the 
metaconid on Py, include a little less difference in width between M, 
and the trigonid of M;, with M; relatively much shorter. Moreover, 
teeth of Thryptacodon demari show a cingulum, usually discontinuous, 
external to the hypoconid, but it appears not to be developed external 
to the protocone to the extent seen in T. antiquus. Also, it is not nearly 
so expanded posterior to the hypoconid on M3. 

A fragmentary maxilla, U.S.N.M. No. 20984, referred to this 
species, has M‘! and M? preserved, and a second maxilla, U.S.N.M. 
No. 20992, has only M?. Also, there is an isolated M?* in the Uni- 
versity of Wyoming collection which may represent this species. 
M? exhibits an anteriorly projecting and weakly cusped parastyle. 
The cingulum is evenly continuous around the anterior and lingual 
portions of the tooth and the hypocone is a simple conical cusp rising 
from the cingulum posterointernal to the the protocone. There is no 
protostyle and there are no particularly distinct accessory cuspules 
adjacent to the hypocone as observed in the Eocene materials. M? 
lacks the distinctive parastyle, and in No. 20984 (but not in No. 
20992) there is a very rudimentary protostyle anterolingual to the 
protocone where the cingulum is somewhat more sharply deflected 
around the margin of the tooth than in M*. As in M?, however, there 
are no clearly distinguishable accessory cuspules adjacent to the 
hypocone. 

The right M? described by Simpson (1928) as Thryptacodon 
pseudarctos in the Bear Creek Paleocene fauna of southern Montana 
is larger and apparently has a more robust protocone than in No. 20984 
considered to be Thryptacodon demari. The type, T. pseudarctos, 
measures 6.9 by 8.6 mm. Measurements of teeth in T. demari are in- 
cluded with those of T. belli. 


THRYPTACODON BELLI,)° new species 
Plate 6, figures I, 4 
Type.—Right ramus of mandible including M,-M;, U. of Wyo. 
No. 1045. 


10 Named for Wallace G. Bell, engaged in the geologic mapping of the Bison 
basin area. 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Horizon and locality——Bison basin Tiffanian, saddle locality at 
south rim of Bison basin, sec. 28, T. 27 N., R.95 W., Fremont County, 
Wyo. 

Specific characters——Length of lower molar series is about 13 per- 
cent less than in the type of Thryptacodon demari and lower jaw 
much shallower. Upper and lower molars similar to those in T. de- 
mart, except that M; is relatively much narrower. 

Discussion.—In addition to the type there are two other specimens 
representing this species in the University of Wyoming collection and 
about nine in the collections of the National Museum. No difficulty 
was encountered in distinguishing these materials from that repre- 
senting Thryptacodon demari and upon completing the segregation of 
the species it was found that all the material except one specimen 
of T. belli was derived from the saddle locality at the south rim, 
whereas all but two of the specimens of T. demari were from the 
localities farther to the west, and apparently a little higher strati- 
graphically. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN TWO NEW SPECIES OF Thryptacodon 


T. demari T. belli 
U.S.N.M. U.S.N.M. 
i . No. 20984 No. 20986 
M’, anteroposterior diameter buccally : transverse 
CiaMetCl sweet eros oe re hes oe Cee eT ea ease e ee 7 Oa: (at ARPES 
M*, anteroposterior diameter buccally : transverse 
diameters: tae kre wid sl erhewers olga erenleneelgetts 6.5°: 8.0 52Gek 
U.S.N.M. U. of Wyo. 
No. 20985 No. 1045 
(type) (type) 
P,, anteroposterior diameter : transverse diameter ... 5.2:2.7. = aaeaee 
M,, anteroposterior diameter : transverse diameter ... 6.7: 4.3 5.6: 3.8 
M:, anteroposterior diameter : transverse diameter ... 6.2: 5.3 5.4:4.7 
Ms, anteroposterior diameter : transverse diameter ... 6.2:4.8 5-3: 4.0 


@ Approximate. 


ARCTOCYONINAE 


The approximately 30 specimens of claenodonts in the Bison basin 
collections nearly run the gamut in size of teeth from a little smaller 
than in Claenodon procyonoides to possibly a little larger than in the 
largest Claenodon ferox, as represented in the Torrejon collections. 
No one, I believe, would seriously contend that a single species is rep- 
resented, nor does it seem possible to arrange them logically, with the 
material at hand, into less than about four species. Possibly a larger 
collection would show a different distribution as to species and would 
probably represent not more than three. Simpson (1936) has shown, 
for example, that the amount of material now known from the Tor- 
rejon in New Mexico has resolved the complex there into only two 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 31 


determinable species, the larger of which exhibits a surprising range 
in size. A complex similar to that in the Bison basin picture is seen 
in the fewer though distinctly better materials encountered in the Fort 
Union of the Crazy Mountain field in Montana. Simpson was there 
faced with the necessity of recognizing five species, but undoubtedly 
this arrangement would also be somewhat simplified if an adequate 
sample could be obtained. 

Except for the largest form in the Bison basin fauna, there seem 
to be no characters but size by which the various species may be 
recognized. Using Simpson’s histogram (1937b, fig. 35) for the 
length of M, in the Torrejon materials in the American Museum, I 
have, in figure 2, added to the number individuals in each size group 
according to information derived from Torrejon collections in the 
National Museum, and included a similar histogram for the Bison 
basin teeth. In the latter, columns are extended by dashed lines in 
instances where in the absence of M, the size of an adjacent molar is 
indicative of one group or another. Specific assignments made, mostly 
tentative, are also shown. I rather supect that with further material 
a single intermediate group will eventually be indicated where com- 
parison is now made with C. montanensis and C. ferox, although the 
pattern shown in the Torrejon materials would suggest great varia- 
tion in a large species. Nevertheless, the differences between the new 
species, C. acrogenius, and that referred to C. ferox are rather marked 
and would appear to include more than size of teeth alone. 


CLAENODON, cf. PROCYONOIDES (Matthew), 1937 
Plate 7, figure 5 


A decidedly small species of Claenodon is represented at the saddle 
locality by a lower jaw, U.S.N.M. No. 20630, including the molars 
M,-Ms, a jaw portion retaining only P, and P;, U.S.N.M. No. 21007, 
and an isolated M;. The proportions of the teeth in No. 20630 are 
very close to those in the type of Claenodon procyonoides from the 
New Mexico Torrejon. The isolated molar represents an individual 
slightly smaller. I was unable to find any characters of significance in 
these specimens by which the Bison basin form could be determined 
as distinct from the earlier C. procyonoides. 


MEASUREMENTS IN MILLIMETERS OF LOWER TEETH IN SPECIMENS OF 
Claenodon, cr. procyonoides (MATTHEW), U.S.N.M. NO. 20630 


Renotn Of lower aiotat, Series, Mi- Mois tig «ain ts. acne sia o00c0igons.s oink 24.0 
M,, anteroposterior diameter : greatest transverse diameter........... Oa 
M2, anteroposterior diameter : greatest transverse diameter........... 8.0: 6.7 


M:, anteroposterior diameter : greatest transverse diameter..........- 8.7: 5.6 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


CLAENODON, cf. MONTANENSIS (Gidley), 1919 
Plate 7, figure 4 


A range in size of teeth indicated by about five claenodont speci- 
mens in the Bison basin suite includes the proportions of the type of 
Claenodon montanensis from the Torrejon stage of the Montana Fort 
Union. All these tentatively referred materials, as well as those of 
the smaller form discussed above, were derived from the vicinity of 
the saddle locality below the south rim of the Bison basin. There 
would appear to be no characters of significance in the rather frag- 
mentary materials of this intermediate species which would serve to 
distinguish it from that of the earlier C. montanensis. Moreover, I 
suspect that additional material may render difficult its clear separa- 
tion from that represented by the materials referred to C. procyo- 
noides. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMEN OF 
Claenodon, cr. montanensis (GIDLEY), U.S.N.M. NO. 20574 


M2, anteroposterior diameter : greatest transverse diameter.......... 9.57275 
M:, anteroposterior diameter : greatest transverse diameter.......... 9.8: 6.3 
* Approximate. 


CLAENODON, cf. FEROX (Cope), 1883 
Plate 7, figures 2, 3 


A somewhat larger series, including at least nine specimens, is most 
nearly comparable to the materials of Claenodon ferox that in the New 
Mexico collections were earlier distinguished as Claenodon corruga- 
tus, or the lower portion of the size range for C. ferox. As well as 
jaw portions, there are in this group three fragmentary maxillae, each 
with two molars, and an isolated P*. In the portion of the histogram 
representing this material the three individuals indicated by dashed 
lines have M; preserved rather than M,, and proportions of the latter 
are estimated by comparison with teeth in a better preserved indi- 
vidual of this group, U.S.N.M. No. 20633. 

Whether or not the form represented by this group of specimens is 
the same as that tentatively referred to C. montanensis, there seems 
no certain evidence ; nevertheless, the extremes in size when combined 
are strikingly far apart, and any attempt to group them together with- 
out rather conclusive evidence would seem an incompatible arrange- 
ment. Moreover, it should be particularly noted that although the 
actual size range of the individuals in such a lumped arrangement 
might be no more than in the C. ferox material of the Torrejon, the 
percentage of difference in the series is very much greater. For this 


INDIVIDUALS 


OF 


NUMBER 


No. 6 PALEOCENE FAUNAS OF BISON BASIN-—GAZIN 33 






C. ocrogenius n.sp. 


C.cf.montanensis C.ct.ferox 













2 C.cf. 
procyonoides 
Bbrigccks 
75 80 85 9.0 95 100105 110 115 12.0 12.5 13.0 13.5 140 14.5 15.0 
C. ferox 
+ 
6 
5 
4 
: C.procyonoides 
2 


73 78 83 88 93 98103 108 13 118 123 128 13.3 13.8 143 148 153 


Fig. 2—Histogram of length of Mz of Claenodon from Bison basin Paleocene, 
above, and Torrejon (modified from Simpson, 1937) of New Mexico, below. 


reason a histogram using a linearly arranged grouping of sizes is 
misleading unless this characteristic is understood. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMENS OF 
Claenodon, cF. ferox (COPE) 
U.S.N.M. 


No. 20797 
M?, anteroposterior diameter : transverse diameter* ...........-- 10.7: 15.9 
M®, anteroposterior diameter : transverse diameter .............. $2): EL.T 
U.S.N.M. 
No. 20633 
M2, anteroposterior diameter : transverse diameter ...........+.- 11.4: 10.4 
Ms, anteroposterior diameter : transverse diameter ...........+-- 12:75 Q.1 


* Transverse diameter taken lingually to base of enamel. 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


CLAENODON ACROGENIUS,1! new species 
Plate 7, figures 1, 6 


Type.—Right ramus of mandible with P,, M,, and Ms, U.S.N.M. 
No. 20634. 

Horizon and locality—Bison basin Tiffanian, saddle locality at 
south rim of Bison basin, sec. 28, T. 27 N., R. 95 W., Fremont 
County, Wyo. 

Specific characters——Size comparable to the very largest indi- 
viduals of Claenodon ferox. Jaw very deep, particularly beneath posi- 
tion of anterior premolars. Canine large and anterior premolars sepa- 
rated by a marked diastema. Ps. possibly absent or reduced to single 
rooted tooth. 

Discussion.—Claenodon acrogenius is represented by approximately 
seven specimens in the collections of the National Museum and pos- 
sibly by a molar talonid in the University of Wyoming collection. 
Among the referred specimens are the posterior portions of three 
other lower jaws with one molar each, two maxillary fragments ex- 
hibiting M+, one of which also includes most of P*, and an isolated P,. 

Although the size of the teeth in this material is within, or nearly 
within, the upper limits of the size range for Claenodon ferox as 
recognized in the Torrejon materials, the depth of the jaw, as shown 
in the type specimen, appears to be exceedingly great, particularly to- 
ward the forward extremity. Moreover, the type specimen has a large 
saberlike canine, as indicated by the root portion, and a diastema of 
very considerable length anterior to P3. P, is close to the canine and 
P, is missing. A short distance posterior to P, there is a depression 
that may have been an alveolus for a root of P,. It is uncertain, of 
course, but this tooth may have been lost during the life of the animal 
and the alveoli nearly obliterated. In any case, the length of the 
diastema between P, and Py, is very much greater than would be 
required for a Ps. 

The two molars preserved in this jaw are much worn and do not 
include M.. These teeth are a little smaller than those in the other 
more fragmentary deep-jawed specimens and are interpreted as repre- 
senting the smaller size group of C. acrogenius as shown in the 
histogram. 


MEASUREMENTS IN MILLIMETERS OF TYPE OF Claenodon acrogenius, 
U.S.N.M. NO. 20634 


Length of cheek tooth series, posterior margin of alveolus of canine to 
Posterior Maree OF Meas saiscsst «adesan sheen eae ake ea aeeee 87." 
Diastemat between: Ps, and (Pavstacticiis. 68 ce nuid.  eeeiek cae eee 15.0 


11 From Greek, akrogeneios = with prominent chin. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 35 


Depth of jaw beneath diastema between P: and Ps................. 31.0 
P;, anteroposterior diameter : transverse diameter ................ Bn 122 
Ps, anteroposterior diameter atialveoli if... cet cece cnc ccee 11.0 
Pe anteroposterior, diameter, atpal veoltcs, cyais ye, ssyeioys « cysre areree ayetoinss oie 13.5 
Mi, anteroposterior diameter : transverse diameter ................ 12:2) 12.0. @.7; 
Ms, anteroposterior diameter : transverse diameter ................ 15.0°: 9.9 

@ Approximate. 

e Estimated. 

MESONYCHIDAE 


DISSACUS, sp. 


Mesonychid creodont material is exceedingly scarce in the Bison 
basin collections, as only two incomplete teeth have been encountered. 
One of these is the outer portion of an upper cheek tooth, not identi- 
fied as to position, but showing the high conical paracone, somewhat 
lower metacone, and a prominent parastyle characteristic principally 
of Ps-Mz in Dissacus. In size it is a trifle larger than M, in D. nava- 
jovius as illustrated by Matthew (1937, figs. 16,17). The other speci- 
men consists of about the posterior two-thirds of a lower cheek tooth. 
The anterior and medial portions of the protoconid, including the posi- 
tion of a possible metaconid, are missing. The shearing talonid is 
slightly longer than in the Torrejon D. navajovius tooth material in 
the National Museum collections. The material, however, is not ade- 
quate for specific diagnosis, although there would seem to be no doubt 
as to the genus represented. 


MIACIDAE 
DIDYMICTIS, near D. TENUIS Simpson, 1935 


A lower jaw fragment with M, and the talonid of M, in the Uni- 
versity of Wyoming collection (No. 1063) would appear to be the 
only determinable miacid material so far encountered in the Bison 
basin collecting. The specimen is from the vicinity of the saddle lo- 
cality below the south rim of the basin. The species is clearly a minute 
form of Didymictis and the talonid of M, has proportions almost the 
same as in the type of Didymictis tenuis from the Gidley quarry in 
the Crazy Mountain Fort Union. M, is not preserved in the D. tenuis 
material, but the root portions shown in the type indicate a tooth 
slightly longer than that of the Bison basin specimen. However, this 
difference alone would not warrant recognition of a separate species. 
Nevertheless, it is probable, in view of the difference in age of the 
horizons represented, that the species are not the same. Mz, in No. 
1063 measures 2.6 mm. long by 1.4 mm. wide. The talonid of M, is 
about 1.5 mm. wide. 


36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


CONDYLARTHRA 


HYOPSODONTIDAE 
PROMIOCLAENUS PIPIRINGOSI,!2 new species 
Plate 11, figures 1, 2 


Type.—Right ramus of mandible with M, and M,, U.S.N.M. 
No. 20571. 

Horizon and localtty—Bison basin Tiffanian, saddle locality at 
south rim of Bison basin, sec. 28, T. 27 N., R. 95 W., Fremont County, 
Wyo. 

Specific characters ——Close in size of molar teeth to Promuoclaenus 
lemuroides (Matthew), but lower premolars noticeably smaller. Pre- 
molars simple and but slightly inflated. P. and P, without parastylid 
and without talonid cusps or crest. P, with only a vestige of a para- 
stylid, no metaconid, but exhibiting two small cusps at posterior mar- 
gin of a very short talonid. Molars relatively narrow transversely 
without entoconid, and hypoconulid on talonid rim of M, and M, well 
defined. 

Discussion.—In addition to the type, a fragmentary jaw with P.-P, 
(U.S.N.M. No. 21021) and a maxilla with M?-M* (U.S.N.M. 
No. 21022) are believed to represent Promioclaenus pipiringosi. The 
lower molars exhibited in the type are only slightly shorter anteropos- 
teriorly than in the Torrejon Promuoclaenus lemuroides material at 
hand, but distinctly narrower relatively. Though worn, the cusps on 
the marginal crest of the talonid appear comparatively well defined, 
rather more as in teeth of the distinctly smaller Promioclaenus aqui- 
lonius of the Montana Fort Union. The lower premolars, if No. 21021 
is correctly referred, are small and not so inflated as in P. lemuroides, 
though less slender than in P. aqguilonius. Moreover, the anterior pre- 
molars are without parastylid or any talonid cusps. P,, however, 
shows a slight parastylid and a pair of cusps on the talonid ; neverthe- 
less, there is no evidence of a metaconid so generally observed on this 
tooth in P. aquilonius. The two upper molars in the tentatively re- 
ferred maxillary fragment are much worn and exhibit few characters 
of significance. The individual represented is a little smaller than the 
type. The external cingulum is prominent between the paracone and 
metacone and divided about midway. 

Use of the generic designation Promioclaenus Trouessart, rather 
than Ellipsodon, for these forms, is in conformity with Dr. R. W. 
Wilson’s findings (1952) with respect to the genotype Ellipsodon 
inaequidens. Trouessart proposed Promioclaenus for the two species 


12 Named for George N. Pipiringos, of the U. S. Geological Survey. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN a7. 


P. acolytus and P. lemurotdes. Presumably P. acolytus (Cope), the 
first of the two listed by Trouessart, is to be regarded as the genotype. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMENS OF 
Promioclaenus pipiringosi 


U.S.N.M. 
No. 21021 
P:, anteroposterior diameter : greatest transverse diameter.......... 2.716 
Ps, anteroposterior diameter : greatest transverse diameter.......... S320 
Ps, anteroposterior diameter : greatest transverse diameter.......... 3.8: 2.8 
U.S.N.M. 
No. 20571 
(type) 
M,, anteroposterior diameter : greatest transverse diameter.......... AL 34 
M2, anteroposterior diameter : greatest transverse diameter.......... AA237 


PROMIOCLAENUS? sp. 


A very large species of Promioclaenus may be represented by a 
fragmentary right mandibular ramus (U.S.N.M. No. 21020), having 
preserved only the posterior portion of M, and part of the trigonid as 
well as the roots of Ms. The form is close in size to Litaletes disjunc- 
tus Simpson of the Montana Fort Union. M3; would appear to be 
fully as large as in L. disjunctus, a relative size quite unlike typical 
Ellipsodon. There is a distinct possibility that this is Litaletes rather 
than Promioclaenus; however, the cusps included in the preserved 
portion of M,, though somewhat worn, suggest a lower crowned tooth 
as in Promtoclaenus, distinctly less inflated than in Mioclaenus. 


LITOMYLUS SCAPHICUS,!* new species 
Plate 8, figures 2, 4 


Type.—Right ramus of mandible with M, and M3, U.S.N.M. 
No. 21014. 

Horizon and locality—Bison basin Tiffanian, saddle locality at 
south rim of Bison basin, sec. 28, T. 27 N., R. 95 W., Fremont 
County, Wyo. 

Specific characters—Lower molars larger and relatively wider than 
in Litomylus dissentaneus. Cusps relatively lower and more inflated, 
with paraconid more reduced. 

Discussion —There would seem no doubt but that the genus Litomy- 
lus, originally described by Simpson on the basis of the species L. dis- 
sentaneus in the Torrejonian of the Montana Fort Union, is repre- 
sented by two distinct species in the Bison basin Tiffanian. Litomylus 


18 Skaphikos, from Greek skaphos = anything hollowed out as a basin, in allu- 
sion to the Bison basin. 


38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


scaphicus, the larger, is represented at the saddle locality by the type 
specimen, and at the ledge locality, about one-quarter mile to the west, 
by a second jaw portion, U.S.N.M. No. 21015, almost identical to it. 

An upper molar, U.S.N.M. No. 21013, possibly M?, from the saddle 
locality is structurally very much like the first or second upper molars 
in L. dissentaneus, except that the protoconule and metaconule are 
less well defined. 

Measurements of the teeth of this form are included with those of 
the following species. 


LITOMYLUS SCAPHISCUS,!* new species 
Plate 8, figure 5 


Type.—Right ramus of mandible with P;, M, and M,, U.S.N.M. 
No. 21010. 

Horizon and locality.—Bison basin Tiffanian, ledge locality at south 
rim of Bison basin, W4 sec. 28, T. 27 N., R. 95 W., Fremont County, 
Wyo. 

Specific characters——Size of teeth close to those in Litomylus dis- 
sentaneus, distinctly less than in L. scaphicus, Ps with parastylid and 
posterior cusps much better developed than in L. dissentaneous. Para- 
conid on molars more reduced. Talonid basin a little less deeply 
pocketed. 

Discussion—A second jaw fragment with M, and part of Mb, 
U.S.N.M. No. 21011, is also from the ledge locality and a single 
lower molar of this small species was encountered at the saddle but 
no material was obtained from the west end of the basin. 

Litomylus scaphiscus resembles the larger L. scaphicus in the more 
reduced paraconids of the lower molars but is strikingly close in size 
of both premolars and molars to L. dissentaneus. It differs from 
L. dissentaneus essentially in the better development of the cusps of 
Ps, the weaker paraconids of the molars (particularly M,), and the 
less deeply basined molar talonids. Moreover, the depth of the pocket 
is greatest nearer the lingual margin than in L. dissentaneus. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN TYPE SPECIMENS OF 
TWO NEW SPECIES OF Litomylus 
L. scaphicus L. scaphiscus 
U.S.N.M. U.S.N.M. 


No. 21014 No. 21010 
: ; (type) (type) 
P;, anteroposterior diameter : greatest transverse 
iatIeter if. 2b vans Uh a apmsna bile anki cs (aemaneieniett haul 3.7: 1:65 
P,, anteroposterior diameter : greatest transverse 
GIAITICTEE goeiae et WR earch fers wie, sis ci e’eic'v'aislepare ceeunis rere tress CRT e pie tTetece 3.8°:— 


14 Skaphiskos, from Greek skaphos (diminutive form) = hollowed out as a 
basin, in allusion to the Bison basin. 


NO. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 39 


Mu, anteroposterior diameter : greatest transverse 


TATE LE Tiamat ases is ol Sette a) Tate ade eceitetouetaaueke isl slis wrap laye Ayety 0161, 0) Stolsiehe 328212 
M2, anteroposterior diameter : greatest transverse 

PLEAS ER OTN eis cia-s situaicutis sain ie Siorsioa aie sie aarsiahis’ ays, © 6.076 BOeasr i, He A 
Ms, anteroposterior diameter : greatest transverse 

GIAINELEE | Mate mretde s wiclee ide tids Shae caekactaiete otis lye « Z8926L8 vires 


¢ Estimated. 


HAPLALETES PELICATUS,15 new species 
Plate o, figure 1 

Type.—Left ramus of mandible with P;-M;, U.S.N.M. No. 21008. 

Horizon and locality—Bison basin Tiffanian, saddle locality at 
south rim of Bison basin, sec. 28, T. 27 N., R., 95 W., Fremont 
County, Wyo. 

Specific characters—Length of lower molar series about one-fifth 
greater than that of Haplaletes disceptatrix. Premolars more inflated. 
Metaconid of P, weaker. Paraconid weak on M,, vestigial or absent 
on M, and M3. External cingulum weak and discontinuous or absent 
on lower molars. 

Discussion.—The type of Haplaletes pelicatus is an excellent lower 
jaw with the cheek-tooth series in a nearly unworn state. Among 
the referred materials are three lower-jaw fragments each with one 
molar, all from the same locality as the type. Two of the latter are in 
the collections of the University of Wyoming. 

The teeth in U.S.N.M. No. 21008 bear a striking resemblance to 
those in the type of H. disceptatrix from the earlier or Torrejonian 
equivalent of the Fort Union, particularly in the form and slope of the 
molar cusps. The premolars, however, are a little more inflated and 
as a consequence the metaconid of P, is not so prominent. Haplaletes 
is rather distinctive among condylarths, and the posterior molars, 
particularly M;, look surprisingly like these teeth in Thryptacodon. 
The first molars, however, bear little resemblance. 

Comparison of Haplaletes pelicatus with Haplaletes diminutivus 
Dorr (1952) is not feasible inasmuch as the latter is represented by 
a partial upper dentition ; nevertheless the very small size of the Dell 
Creek form leaves no doubt as to their distinctness, 

Measurements of teeth in H. pelicatus are included with those of 
the following species. 


HAPLALETES SERIOR,!* new species 
Plate 9, figure 2 
Type——Left ramus of mandible with Mz, and M;, U. of Wyo. 
No. 1078. 


15 From Greek, pelike = basin, in allusion to the Bison basin. 
16 Serior = later, with reference to its stratigraphic position. 


40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


Horizon and locality.—Bison basin Tiffanian, Titanoides locality, 
southwestern portion of basin, sec. 29, T. 27 N., R. 95 W., Fremont 
County, Wyo. 

Specific characters—Lower molar teeth about 15 percent longer 
and nearly 22 percent wider transversely than in Haplaletes pelicatus. 
Paraconid on M, and Ms; vestigial. Talonid basin shallow. Hypo- 
conulid of M,; broader and less protruding posteriorly. No external 
cingulum. 

Discussion.—The type of Haplaletes serior is the only known speci- 
men and comes from near where the Titanoides upper teeth were dis- 
covered, about a mile west of the saddle locality, and apparently a 
little higher stratigraphically. 

Haplaletes serior is a larger form than H. pelicatus with relatively 
wider molars. The form appears, from the limited material, to repre- 
sent the same genus as H. pelicatus but would seem more distinct 
from H. disceptatrix. However, the difference from the latter is for 
the most part a rather marked disparity in size. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN TYPE SPECIMENS OF 
TWO NEW SPECIES OF Haplaletes 


H., pelicatus H. serior 


U.S.N.M. U. of Wyo. 
No. 21008 No. 1078 
(type) (type) 
P;, anteroposterior diameter : greatest transverse 
MiAMEtEN OS irals ascyaherele's tos ose Se ieareeta ok 26° i.6F Bh Wine 
Ps, anteroposterior diameter : greatest transverse 
CATTICLER gees oie i6 cyst fayeralon ete Ste sya ebucin scl s ele iersuspnioke iene B23 20 ae erecta 
M,, anteroposterior diameter : greatest transverse 
Giamleter Greene riaie acter nme ae aie son etre moe 2102s eee 
M2, anteroposterior diameter : greatest transverse 
(idimeterahe hs 050. ces coo eens 3.2: 2.8 Bg 
M;, anteroposterior diameter : greatest transverse 
CIATMELE Ii sie oh =. shave scolvgcteversuelehers seus enereibapsharel cre eaters 3.2.2.6 BUF aur 


« Approximate. 


PROTOSELENE? NOVISSIMUS,!7 new species 
Plate 8, figures I, 3 


Type—Left ramus of mandible with M, and M3, U.S.N.M. 
No. 20572. 

Horizon and locality—Bison basin Tiffanian, saddle locality at 
south rim of Bison basin, sec. 28, T. 27 N., R. 95 W., Fremont 
County, Wyo. 


17 Novissimus = youngest or latest, with reference to the stratigraphic horizons 
for Protoselene. 


NO. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 4I 


Specific characters—Lower molars a little smaller and relatively 
more slender than in Protoselene opisthacus. Paraconid on M, and M; 
slightly lingual to midposition and distinctly isolated from both proto- 
conid and metaconid. Talonid crest, particularly the crista obliqua, 
lower, and basin a little shallower. 

Discussion.—A single M, (U.S.N.M. No. 21023) is known in ad- 
dition to the type and was found at the same locality. 

Protoselene? novissimus may not represent this genus but is much 
closer to it than to any other known condylarth. It has elongate 
molars approximating the selenodonty exhibited in P. opisthacus, but 
the crests are lower and consequently the basins a little more shallow 
appearing. The paraconid is located in about the same position, but 
on the posterior molars is more definitely isolated from the adjacent 
cusps. The extent to which this cusp is joined by a crest to the pro- 
toconid, however, is variable in the Torrejon form. M, in the type 
specimen measures 5.4 mm. long by 3.8 mm. wide. M3; is 5.5 by 
3.2 mm. 

The isolated M, can be nearly matched in material of P. opisthacus 
but in each case the trigonid and talonid basins are a little shallower 
and the crista obliqua between the hypoconid and trigonid is a little 
more depressed. M, measures 5.6 mm. long by 3.6 mm. wide. 


LITOLESTES LACUNATUS,!§ new species 
Plate 11, figures 3, 4 


Type—Left ramus of mandible with P, and M,, U.S.N.M. 
No. 21016. 

Horizon and locality—Bison basin Tiffanian, Titanoides locality, 
southwestern portion of Bison basin, sec. 29, T. 27 N., R. 95 W., 
Fremont County, Wyo. 

Specific characters—Approximately a third larger than Litoléstes 
notissimus in size of lower molars and well outside the range given by 
Simpson (1937a). Lower premolars relatively larger. P, with small 
anterolingual parastylid, metaconid weak and close to primary cusp, 
and two small talonid cuspules very close together. Paraconid weak 
or absent on M, and M;. Metaconid and protoconid about equal on 
M, and Mz, and metaconid higher on M3. Entoconid comparatively 
high on all three molars. 

Discussion—The form herein described as Litolestes lacunatus is 
the smallest of the condylarths recognized in the Bison basin collec- 
tions, yet it is distinctly larger than either of the previously described 


18 Lacunatus = hollowed out, with reference to the Bison basin. 


42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


species of this Tiffanian genus. The genotype Litolestes ignotus Jep- 
sen is from the Silver Coulee horizon in the Polecat Bench series and 
L. notissimus from the Melville portion of the Crazy Mountain Fort 
Union. Litolestes lacunatus is represented in the Bison basin collec- 
tions by two additional specimens from the same locality as the type: 
a lower jaw fragment with M, and M, (U. of Wyo. No. 1083) 
and a jaw fragment with M; and part of Mz (U. of Wyo. No. 1079). 
A lower jaw portion with P;, P,, and a much worn M, (U. of Wyo. 
No. 1059) from the saddle locality may represent this species, but 
P, is lower and wider and lacks any evidence of a metaconid; more- 
over, the details of M, are rather obscured by wear. A jaw fragment 
with only P, (U.S.N.M. No. 21017) corresponds very closely to the 
type but came from the locality at the extreme west end of the basin. 

A maxilla (U.S.N.M. No. 20931) from the west-end locality ex- 
hibiting P*-M? has rather well worn molars; nevertheless there seems 
no doubt that it represents Litolestes lacunatus. P* and P* are nearly 
similar to those teeth in L. notissimus, but the parastyle, a very small 
cusp on P* in L. notissimus, is absent on this tooth in L. lacunatus, 
however, that on P, is more robust in L. lacunatus. The parastyle 
and perhaps the metastyle on the molars are not so outstanding buc- 
cally as in L. notissimus. Moreover, the talon portions of the molars 
appear to be a little broader anteroposteriorly in the Bison basin 
specimen. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMENS OF 
Litolestes lacunatus 


U.S.N.M. 
No. 21016 
(type) 
P,, anteroposterior diameter : greatest transverse diameter......... 3.2: 1.9 
M,, anteroposterior diameter : greatest transverse diameter......... 2.7: 2.0 
U. of Wyo. 
No. 1079 
Wy, (Erarisverse Clameter: Of. CALONIG «wis «djs svn sie alee wie wa Mean ae mately 2.2 
M;, anteroposterior diameter : greatest transverse diameter......... ZAS 1 


PHENACODONTIDAE 
GIDLEYINA WYOMINGENSIS, new species 
Plate 0, figures 3, 4 
Type.—Right ramus of mandible with P,;-M,, U.S.N.M. No. 20790. 
Horizon and locality—Bison basin Tiffanian, locality at west end 


of Bison basin, N4 sec. 29, T. 27 N., R. 95 W., Fremont County, 
Wyo. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 43 


Specific characters.—Size of P, and molars close to that in Gid- 
leyina silberlingi. P; much smaller and with simple posterior median 
crest and single talonid cusp. Paraconid of lower molars variable but 
generally low and weak. Trigonid not so prominently basined as in 
Gidleyina superior. Upper molars with prominent styles, and crests 
of protocone distinctive. 

Discussion.—Gidleyina wyomingensis is better represented in the 
more westerly and stratigraphically somewhat higher levels than at 
the saddle locality. The type is from the locality at the west end of 
the basin, as are about eight other specimens, although most of these 
are isolated upper and lower teeth. The material from the ledge lo- 
cality seems entirely similar to that from the west-end locality, and 
among the 10 specimens from the ledge is the upper dentition 
(U.S.N.M. No. 20795) figured in plate 9 and about three jaws with 
two or more molars. About five specimens from the saddle locality, 
including portions of upper and lower dentitions, might well represent 
a slightly smaller variant, though probably not specifically distinct 
from that represented in the material from the more westerly collec- 
ing sites. 

Doubt may be logically entertained as to the advisability of recog- 
nizing Gidleyina as distinct from Ectocion. Comparison of the Tif- 
fanian materials with the genotype of Ectocion, E. osbornianum, from 
the lower Eocene would seem to justify separate recognition but, as 
may be expected, the Clarkforkian materials, particularly those from 
the Almy, in no way simplify this arrangement. As noted by Simp- 
son, the upper premolars in Gidleyina are less progressive and the 
upper molars show better development of crests from the protocone 
to the protoconule and metaconule. No upper premolars appear to be 
included in the Bison basin collections but the molars exhibit the 
protocone crests as mentioned above, and in comparison with G. mon- 
tanensis have perhaps somewhat better developed external styles. 
P, in both the Montana and Wyoming Gidleyina material would ap- 
pear distinctive when compared with Eocene material of Ectocion, 
principally in that the trigonid is elongate in comparison: with the 
talonid length, whereas in EF. osbornianum this relationship is rather 
generally reversed with the trigonid often short and broad and the 
talonid usually, though not invariably, better developed. Moreover, 
in some individuals of E. osbornianum the talonid of P, looks quite 
molariform, with a surprisingly well developed entoconid. 

With regard to the forms of Gidleyina known from the Melville unit 
of the Montana Fort Union, I strongly suspect that Gidleyina sil- 
berlingi is a synonym of G. montanensis. Simpson (1937b) called 


44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


attention to this possibility at the time he published Gidley’s descrip- 
tions. There is some difference in the stratigraphic levels attributed 
to the two, but not as much difference as between either of them and 
Gidleyina superior. Nevertheless, to judge by the variation in molar 
structure noted for both Gidleyina and Ectocion, G. superior may be 
no more than a variant of G. montanensis. The possibility also re- 
mains that G. wyomingensis is likewise not distinct, but possible 
synonomy here awaits demonstration that the lower premolars ex- 
hibited in the type of G. silberlingi are atypical. 

A form which Simpson (1935c) described as Phenacodus gidleyt in 
the Tiffany fauna has teeth only a trifle larger than in G. wyomingen- 
sis, but the trigonid portions of the lower teeth in the type of P. gidley1 
represent a little greater proportion of the tooth length than in 
G. wyomingensis. I am unable to determine whether P. gidleyi repre- 
sents Phenacodus or Gidleyina. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMENS OF 
Gidleyina wyomingensis 


U.S.N.M. 
No. 20795 
Leneth ot wppet.molar, series, M2—-MF sis escoceicteiics ot aesstamste dale wivinrget 20.1 
M’, anteroposterior diameter : greatest transverse diameter........ TAT AQ 
M?, anteroposterior diameter : greatest transverse diameter........ 2 TOA 
M®, anteroposterior diameter : greatest transverse diameter........ he gti a | 
U.S.N.M. 
No. 20790 U.S.N.M. 
- : (type) No. 20793 
P;, anteroposterior diameter : greatest transverse 
AiamIGter ee Seer es Boas Sota se ee iene ate a cine ak Res Slt Gite 
Ps, anteroposterior diameter : greatest transverse 
CUNAITICEEY “he 8 chs ctraisias vin i3 sos oie, 674 O90 & 5 bytiaisalcine asia ate G8 AF yn eee 
M,, anteroposterior diameter : greatest transverse 
GiamMetererenn sce onsale ss chin sees ne See eee O:555.2 6.6: 5.2 
M2, anteroposterior diameter : greatest transverse 
iatHeeE EG os HE IL 8, LOC Othe ATEN SICES . CM ee ee 6.8: 5.6 
M,,' transverse: diameter® of ‘trigonid: syoijei. os. Snide So ieleleltnele «08 4.8 


@ Approximate. 
PHENACODUS? BISONENSIS,!9 new species 
Plate 10, figures 1-3 
Type.—Right maxilla with P*-M?, U.S.N.M. No. 20564, and prob- 


ably a left maxilla with M* and M? believed to be from the same 
individual. 

Horizon and locality—Bison basin Tiffanian, vicinity of saddle lo- 
cality at south rim of Bison basin, sec. 28, T. 27 N., R. 95 W., Fre- 
mont County, Wyo. 


19 Named for the Bison basin. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 45 


Specific characters.—Size very close to that of Phenacodus almien- 
sis, about intermediate between that of Phenacodus? grangeri and 
Phenacodus? matthewi of the Tiffany beds. P* and P* with tritocone 
distinct, but much less progressive than in P. almiensis. Mesostyle 
prominent on M? but variable on M?. Lower premolars comparatively 
simple and unprogressive. P* trigonid with paraconid low and for- 
ward, and talonid weakly basined, with entoconid generally distinct 
though small. 

Discussion——Approximately 30 specimens of this form are at 
hand and nearly all are from the vicinity of the saddle locality. Two 
specimens, however, were secured from the ledge locality statigraphi- 
cally a little higher. 

Uncertainty exists as to whether the species represented by this 
material should be referred to Phenacodus or to Tetraclaenodon. Its 
allocation to Phenacodus is entirely arbitrary and scarcely more than 
an impression. As noted by Granger (1915), there are actually no 
clear-cut characters by which the genera may be separated. Although 
there are differences between them in degree of development for a 
number of characters, they are in the nature of average differences, 
lacking in the consistency generally expected at the generic level. 
Granger attempted a definition based on the development of the 
mesostyle, but certain upper molars of Tetraclaenodon puercensis 
show a rather surprising prominence in this style. The shift of the 
metaconule posteriorward would seem evident for Phenacodus pri- 
maevus but not diagnostic for such Paleocene forms as Phenacodus 
almiensis or Phenacodus? grangeri. I note a decreasing prominence 
of the protoconule and metaconule with respect to the primary cusps 
in rising above the Torrejon level, to the extent that in some 
Wasatchian material of Phenacodus the metaconule is entirely missing 
on M? and M®. Nevertheless, this is variable in populations of the 
better known species of Phenacodus as well as in Tetraclaenodon 
puercensis, and, like the increasing significance of the tritocone of the 
upper premolars, is a difference in degree not readily defined. 

The lower teeth do not appear to present characters of significance 
on a generic level. Certainly the development or reduction of the 
paraconid is too highly variable. The talonid of P, would seem to 
become more molariform in time and the entoconid better developed 
but this cusp is occasionally prominent in material of Tetraclaenodon 
puercensis, and a decidedly primitive appearing P, structure has been 
observed in material representing certain of the smaller species of 
Phenacodus in the Eocene. 

Phenacodus? bisonensis would appear to be Tetraclaenodon in the 
subordinate appearance of the tritocone on P* and P* and its proximity 


46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


to the primary cusp, and in this respect is certainly distinct on a specific 
level from the Clarkforkian materials, such as Phenacodus almiensis. 
Nevertheless, these teeth show deuterocone portions more suggestive 
of Phenacodus, and P* has a rather conspicuously developed postero- 
internal talon basin not observed in U. S. National Museum Torrejon 
materials. P.? bisonensis, moreover, resembles P. almiensis in the 
slightly more crested appearance of the upper molar cusps and in the 
lesser significance of the conules in comparison with Tetraclaenodon 
puercensis. The mesostyle of the upper molars is distinct and moder- 
ately prominent in all referred materials ; however, in the type speci- 
men, although prominent on M}, it is very weak on M?. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMENS OF 
Phenacodus? bisonensis 


U.S.N.M. 
U.S.N.M. No. 20564 
No. 20566 (type) 
P*, anteroposterior diameter : transverse diameter ... 7.8:6.5 ....+.. 
P*, anteroposterior diameter : transverse diameter ... 7.2: 8.0 8.3: 9.2 
M’, anteroposterior diameter : greatest transverse 
GiamMetes sores. te. aqshits. Bech dee Ste. ve 2th .eseter : 9.4: 11.0 
M’, anteroposterior diameter : greatest transverse 
(cl ci cao 12) SEE Py See RE IE IER SE ee eee Ler 9.4: 12.8 
U.S.N.M. U.S.N.M. 
No. 20567 No. 20569 
Length of lower molar series.........cccccccccees Mite. 28.7 
P;, anteroposterior diameter : greatest transverse 
PAS EER i she’ oid ne toy 2 win patintc ia Ye capoav ope iege Eviowels jetenneig via AcObe Reta eaeteiee 
P,, anteroposterior diameter : greatest transverse 
MATTEL Me Wirig si otetsianeo1d 5 0s ais w aides lei ene artes iss SOc Se Neceaeee ‘ 
M,, anteroposterior diameter : greatest transverse 
Giameterilit.. ced. ee EE ahd tree teeta eke 8.9: 7.6 8:2:'7.3 
M2, anteroposterior diameter : greatest transverse 
CiaMetert bakrscresece si Mbl lesiunatize etc aette ext eaten foe niOeare 2 rae 9.6: 8.2 
M;, anteroposterior diameter : greatest transverse 
PAL ANIIOU RL table oicseiu ics /a, bind ea ee oe pie a hee evavage | Syaererene 10.5: 7.2 


@ Approximate. 


PHENACODUS? sp. (large) 
Plate 10, figures 4, 5 


A fragmentary right mandibular ramus with Ms;, U.S.N.M. 
No. 21025, an isolated P*, M°, and an incomplete lower molar (the 
latter two in the collections of the University of Wyoming) are of a 
species much larger than Phenacodus? bisonensis. The range of size 
represented in materials of P.? bisonensis is surprisingly limited, cer- 
tainly in comparison with such forms as T. puercensis and P. primae- 
vus, so that the teeth here indicated as of a distinct species stand out 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 47 


conspicuously in the collections. The form represented may be 
Phenacodus? grangeri which Simpson (1935c) described from the 
Colorado Tiffany, but the measurements where equivalent materials 
are present noticeably exceed those of the more southern animal. 
M; which measures 13.6 by 9.2 mm., for example, is 19 percent longer 
and 12 percent wider. This difference in a form such as Phenacodus 
might not be important. The significance would, of course, depend 
on the position of these two examples with regard to their respec- 
tive but unknown means. 

The third lower molar apparently reveals no information as to 
whether references should be made to Phenacodus or Tetraclaenodon. 
P’, however, as in P.? bisonensis has a well-developed posterointernal 
basin and the tritocone is apparently better defined than usual in 
Tetraclaenodon puercensis. The isolated M® has a distinct but small 
mesostyle and the hypocone is particularly small. 


PANTODONTA 


CORY PHODONTIDAE 
TITANOIDES PRIMAEVUS Gidley, 1917 
Plate 11, figure 5 


The finding in North Dakota by a party under the direction of 
Dr. Glenn L. Jepsen of portions of the skull including the upper denti- 
tion belonging beyond doubt to the type of Titanoides primaevus was 
unusually good fortune, so that no uncertainty now exists as to the 
characteristics of the superior dentition of this upper Paleocene panto- 
dont. Patterson early (1933) described new materials from the Pla- 
teau Valley beds of Colorado as representing Titanoides, but upon later 
‘discovery of at least three pantodont forms from these beds, with 
distinguishing features in the upper dentition, was forced to regard 
all as distinct from Titanoides and the species at first referred to Tita- 
noides was given the new generic name Barylambda. It now develops, 
with the finding of Titanoides upper teeth, that while Barylambda and 
Haplolambda are clearly distinct, Sparactolambda *° is the form which 
I believe must now be regarded as the synonym of Titanoides. 

In the University of Wyoming material from the Bison basin there 


20 Jepsen’s discovery has likewise permitted us to determine correctly the 
identity of an excellent pantodont skull collected by Dr. T. E. White in 
McKenzie County, N. Dak., and, like the type of T. primaevus, from the general 
area of the type Fort Union. This skull, originally and with apparent correct- 
ness, determined by White as representing Sparactolambda, is now seen to belong 
beyond doubt to Titanoides primaevus. 


48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


is a right maxilla (U. of Wyo. No. 1093) with three little-worn upper 
molars. The resemblance of these teeth to those in the type of 
Titanoides primaevus is striking so that no doubt exists as to the 
identity. The teeth in No. 1093 are slightly smaller in all dimensions, 
except M’, and are nearly similar in form. Observable differences 
include a better developed metacone on the molars, and M® shows, 
in addition to a slightly larger size, the parastyle directed somewhat 
more laterally in No. 1093, not so forward as in the type. Very slight 
differences noted are a less developed protoconule on M? and meta- 
conule on M®. 


MEASUREMENTS IN MILLIMETERS OF UPPER MOLARS IN SPECIMEN OF 
Titanoides primaevus GIDLEY, U. OF WYO. NO. 1093 


Length ot aipper,molar series, .Mi=Me ss sd:scicpnmivetse nemidscuaceceines 66.8 
M,, anteroposterior diameter : transverse diameter * .............-. 20.8 : 23.3 
M2, anteroposterior diameter : transverse diameter * ............... 25.3: 27.9 
M;, anteroposterior diameter : greatest transverse diameter......... 20.4: 34.0 


_* Anteroposterior diameters of M, and Mg taken across outer styles and transverse 
diameters perpendicular to line between outer styles. Anteroposterior diameter of Mg taken 
perpendicular to anterior face. 


CAENOLAMBDA,?! new genus 


Type.—Caenolambda pattersoni, new species. 

Generic characters—Skull with elongate cranium, strong, arched 
sagittal crest, broad frontals, narrow nasals and heavy canines re- 
sembling the Titanoides group. Upper cheek teeth, though compara- 
tively small, are anteroposteriorly shortened and transversely broad 
as in the Barylambda-Haplolambda group, but with molars M' to M® 
about equaling one another in size. 

Discussion.—Caenolambda presents a rather unusual combination 
of characters and does not closely resemble any of the previously de- 
scribed genera. Nevertheless, in a general way, the skull is apparently 
more like Titanoides than Barylambda or Haplolambda. This is 
noticed in the relatively elongate cranium and strong, arched sagittal 
crest. It resembles the cast of the paratype of “Sparactolambda” looki 
in narrowness of the nasals, although the nasal cavity is apparently 
not so large. As in the latter and the type of Titanoides primaevus 
the canine is very well developed. The upper cheek teeth, however, 
are decidedly different. The teeth are distinctly shortened anteropos- 
teriorly, particularly the lingual portions, and very broad transversely. 
The external styles at the anterior and posterior angles of the teeth 
do not project laterally to such an extent and the primary external 
cusps are somewhat closer to the labial margin of the tooth, so that 


21 From Greek kainos, recent or new, + /ambda, the Greek letter—named in 
analogy with Pantolambda, Barylambda, Archaeolambda, and others. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 49 


the characteristic “V” and “W” shapes to the outer cusps are not so 
transversely extended as in Titanoides, or the more extreme condi- 
tion seen in Archaeolambda. The relatively great width is composed 
largely of the talon, and in the molars, as clearly shown in M?’, there 
is a strong ledgelike cingulum about the lingual and posterior margin. 
On Titanoides molars the lingual cingulum is weak, although better 
developed along the posterior margin of the premolars than in Caeno- 
lambda. The molars of Caenolambda would appear to be about equal 
to one another in size as in Pantolambda, not showing the marked 
increase from M?! to M® seen in Titanoides, or the reduction of M* 
noted in Barylambda and Haplolambda. 

The skull of Barylambda is large and relatively broader than that 
of Caenolambda. The nasal cavity is larger and the nasal bones much 
wider. The frontals are broad in both forms, but Barylambda does 
not exhibit a sagittal crest so heavy and prominently arched as in 
Caenolambda. The teeth of Barylambda are transversely broad in 
comparison to their anteroposterior dimension, as in Caenolambda, but 
the talon of molars, particularly M1? and M?, is not nearly so slender ; 
moreover, the cingulum is weak or absent lingually rather than shelf- 
like. M*, as noted above, is much reduced in Barylambda. 

The comparisons between Haplolambda and Caenolambda are 
rather similar to those between Barylambda and Caenolambda, 
although the species Haplolambda quinni and Caenolambda pattersonm 
are more nearly comparable in size. The cranial portion of Haplo- 
lambda is shorter and the sagittal crest not so arched as in Caeno- 
lambda, but the nasals are wider and the nasal cavity larger although 
the frontals are not so broad. The cheek teeth resemble those of 
Caenolambda in their relative width and the distinctly labial position 
of the primary external cusps, but again as in Barylambda the molar 
talons are not so slender, the cingulum is lingually weak, and M® is 
reduced. In Haplolambda, moreover, M* would appear to be larger 
than M?, quite the reverse of Titanoides. The comparatively small 
canine in Haplolambda would appear to be a striking difference from 
both Caenolambda and Titanoides, but the size of this tooth is so often 
a matter of dimorphism that one hesitates to stress the character. 
Nevertheless, if there is any dimorphism in this respect within species 
of Coryphodon, it is certainly much less evident. 


CAENOLAMBDA PATTERSONI,?? new species 
Plates 12-14 


Type—Skull, lacking zygomatic arches and mandible, U.S.N.M. 
21036. 


22 Named for Bryan Patterson in appreciation of his work on the pantodonts. 


50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Horizon and locality.—Bison basin Tiffanian, vicinity of saddle lo- 
cality at south rim of Bison basin, sec. 28, T. 27 N., R. 95 W., Fre- 
mont, Wyo. 

Specific characters —Length of skull greater than that of Haplo- 
lambda quinni but less than Titanoides primaevus. Much smaller than 
Barylambda faberi. Cheek teeth comparatively small. Other charac- 
ters not distinguished from those discussed above as characterizing 
the genus. 

Discussion.—Except for a few isolated teeth or tooth fragments 
which may represent this species, there are no determinable materials 
other than the type. Moreover, the more fragmentary specimens can- 
not be allocated as between this form and Titanoides primaevus, the 
latter having been certainly encountered only at the Titanoides lo- 
cality, at a level believed to be higher stratigraphically than the saddle. 

The skull designated the type of Caenolambda pattersoni is ad- 
vanced in maturity so that the teeth are rather well worn, with the 
characters of M* almost obliterated. Moreover, the sutures are nearly 
all obscured so that little is revealed of the surface extent of the sepa- 
rate elements of the skull. This situation was further complicated 
by the fact that the skull was discovered in a dense limestone nodule 
and during its preparation much difficulty was experienced determin- 
ing the actual boundary between bone and matrix. As a result much 
in the way of important detail cannot be discerned. 


MEASUREMENTS* IN MILLIMETERS OF SKULL, U.S.N.M. NO. 21036, 
TYPE SPECIMEN OF Caenolambda pattersoni 


Length of skull from the anterior margin of premaxillae to posterior 


marpinior occipital condyles sjc.5 .u'dieas av coe Ses. BEMLS Web db oat 320. 
Length from anterior margin of canine alveolus to posterior margin 

GEAOCCII tal ;CONGYIES of ote) ste.c sie pla piaccusuce mebisicsatasie oe bicvelate eisseieetaie 300. 
Distance from posterior margin of palate at posterior narial aperture 

to posterior margin of occipital condyles. .........2c2csecceeesns 170. 
Wadth across ‘postorbital ‘processes... oe ee tes ot te neale emt TI0. 
Width across nasals about midway of length..........-..eceesceeeee 24. 
Length of upper dentition, C (at alveolus) to M§, incl.............. . 135. 
Length of upper cheek tooth series, P? to M%, incl..........c002ceee: 92. 
Length .ot. upper molar series; Mo to MM... ..ns..+axeesmtnnaemeedeenas 52. 
C, anteroposterior diameter (at alveolus) : greatest transverse diameter 26.0: 16.0 
P’, anteroposterior diameter : transverse diameter ............+-+- 13.0: 19.0 
P*, anteroposterior diameter : transverse diameter ..........+-.++- 13.0: 23.0 
P*‘, anteroposterior diameter : transverse diameter ..............+- 23.53 235 
M’, anteroposterior diameter : transverse diameter ...........e-+e- 18.5 : 27.0 
M°, anteroposterior diameter : transverse diameter ...........+.2++ 19.0 : 30.0 
M®*, anteroposterior diameter : transverse diameter .........-.++-.- 16.0: — 


* Measurements are nearly all ae owing to fracturing and distortion of skull, 
and teeth are much worn and nearly all slightly damaged at styles. Tooth measurements 
include styles and are taken parallel to and at right angles to direction of tooth row. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 5!i 


REFERENCES 
Core, Epwarp D. 

1881. Notes on Creodonta. Amer. Nat., vol. 15, pp. 1018-1020. 

1883. First addition to the fauna of the Puerco Eocene. Proc. Amer. Philos. 
Soc., vol. 20, pp. 545-563. 

Dorr, JouHN A., JR. 

1952. Early Cenozoic stratigraphy and vertebrate paleontology of the 
Hoback basin, Wyoming. Bull. Geol. Soc. Amer., vol. 63, pp. 59-94, 
figs. 1-7, pls. 1-7. 

Douctass, Ear. 

1908. Vertebrate fossils from the Fort Union beds. Ann. Carnegie Mus., 

vol. 5, pp. 11-26, pls. 1, 2. 
Freroy, K. K. 

1952. Pantodonty (Pantodonta), sobrannye mongol’skoi paleontologicheskoi 
ekspeditsiel Akademii Nauk SSSR. Paleontologicheskii Institut. 
Trudy, vol. 41, pp. 43-50, 1 pl. 

Gazin, C. Lewis. 

1942. Fossil Mammalia from the Almy formation in western Wyoming. 
Journ. Washington Acad. Sci., vol. 32, No. 7, pp. 217-220, figs. 1, 2. 

1952. The lower Eocene Knight formation of western Wyoming and its 
mammalian faunas. Smithsonian Misc, Coll., vol. 117, No. 18, 
pp. 1-82, figs. 1-6, pls. 1-11. 

GIpLEy, JAMEs W. 

1909. Notes on the fossil mammalian genus Ptilodus. Proc. U. S. Nat. Mus., 
vol. 36, pp. 611-626, figs. 1-9, pl. 70. 

1915. An extinct marsupial from the Fort Union with notes on the 
Myrmecobidae and other families of this group. Proc. U. S. Nat. 
Mus., vol. 48, No. 2077, pp. 395-402, pl. 23. 

1917. Notice of a new Paleocene mammal, a possible relative of the titano- 
theres. Proc. U. S. Nat. Mus., vol. 52, pp. 431-435, fig. 1, pl. 36. 

1919. New species of claenodonts from the Fort Union (Basal Eocene), of 
Montana. Bull. Amer. Mus. Nat. Hist., vol. 41, art. 14, pp. 541-555, 
figs. I-10, pl. 28. 

1923. Paleocene primates of the Fort Union, with discussion of relation- 
ships of Eocene primates. Proc. U. S. Nat. Mus., vol. 63, art. 1, 
No. 2460, pp. 1-38, figs. 1-4, pls. 1-5. 

GRANGER, WALTER. 

1915. A revision of the lower Eocene Wasatch and Wind River faunas. 
Part 3.—Order Condylarthra. Families Phenacodontidae and 
Meniscotheriidae. Bull. Amer. Mus. Nat. Hist., vol. 34, art. 10, 
pp. 320-361, figs. 1-18. 

GRANGER, WALTER, and Simpson, GEORGE G. 

1929. A revision of the Tertiary Multituberculata, Bull, Amer. Mus. Nat. 

Hist., vol. 56, art. 9, pp. 601-676, figs. 1-43. 
JEPSEN, GLENN L. 

1930a. New vertebrate fossils from the lower Eocene of the Bighorn basin, 
Wyoming. Proc. Amer. Philos. Soc., vol. 69, pp. 117-131, No. 4, 
pls. 1-4. 

1930b. Stratigraphy and paleontology of northeastern Park County, Wyo- 
ming. Proc. Amer. Philos. Soc., vol. 69, No. 7, pp. 463-528, figs. 1-4, 
pls. I-10. 


52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


1934. A revision of the American Apatemyidae and the description of a 
new genus, Sinclairella, from the White River Oligocene of South 
Dakota. Proc. Amer. Philos. Soc., vol. 74, No. 4, pp. 287-305, 
figs. 1-4, pls. 1-3. 
1940. Paleocene faunas of the Polecat Bench formation, Park County, 
Wyoming. Proc. Amer. Philos. Soc., vol. 83, No. 2, pp. 217-340, 
figs. 1-22, pls. I-5. 
MATTHEW, WILLIAM D. 
1g1sa. A revision of the lower Eocene Wasatch and Wind River faunas. 
Part 1—Order Ferae (Carnivora). Suborder Creodonta. Bull. 
Amer. Mus. Nat. Hist., vol. 34, art. I, pp. 1-103, figs. 1-87. 
191sb. A revision of the lower Eocene Wasatch and Wind River faunas. 
Part 4.—Entelonychia, Primates, Insectivora (part). Bull. Amer. 
Mus. Nat. Hist., vol. 34, art. 14, pp. 429-483, figs. 1-52, pl. 15. 
1917. The dentition of Nothodectes. Bull. Amer. Mus. Nat. Hist., vol. 37, 
art. 33, pp. 831-839, pls. 99-102. 
1918. A revision of the lower Eocene Wasatch and Wind River faunas. 
Part 5.—Insectivora (continued), Glires, Edentata. Bull. Amer. 
Mus. Nat. Hist., vol. 38, art. 16, pp. 565-657, figs. 1-68. 
1937. Paleocene faunas of the San Juan basin, New Mexico. Trans. Amer. 
Philos. Soc., n.s., vol. 30, pp. i-viii, 1-510, figs. 1-85, pls. 1-65. 
MatrHew, WILLIAM D., and GRANGER, WALTER. 
1921. New genera of Paleocene mammals. Amer. Mus. Novitates, No. 13, 
pp. I-7. 
PATTERSON, BRYAN. 
1933. A new species of the amblypod Titanoides from western Colorado. 
Amer. Journ. Sci., vol. 25, pp. 415-425, figs. 1-4. 
1934. A contribution to the osteology of Titanoides and the relationships 
of the Amblypoda. Proc. Amer. Philos. Soc., vol. 73, No. 2, 
pp. 71-101, figs. I-13, 2 pls. 
1935. Second contribution to the osteology and affinities of the Paleocene 
amblypod Titanoides. Proc. Amer. Philos. Soc., vol. 75, No. 2, 
Pp. 143-162, figs. 1-6. 
1937. A new genus, Barylambda, for Titanoides faberi, Paleocene amblypod. 
Geol. Ser. Field Mus. Nat. Hist., vol. 6, No. 16, pp. 229-231. 
1939. New Pantodonta and Dinocerata from the upper Paleocene of west- 
ern Colorado. Geol. Ser. Field Mus. Nat. Hist., vol. 6, No. 24, 
Pp. 351-384, figs. 100-111. 
RussEL1, Loris S. 
1929. Paleocene vertebrates from Alberta. Amer. Journ. Sci., vol. 17, pp. 
162-178, figs. 1-5. 
1932. New data on the Paleocene mammals of Alberta, Canada. Journ. 
Mammalogy, vol. 13, No. 1, pp. 48-54, figs. I-12. 
Scott, WILLIAM B. 
1892. A revision of the North American Creodonta with notes on some 
genera which have been referred to that group. Proc. Acad. Nat. 
Sci. Philadelphia, vol. 44, pp. 291-323. 
Simpson, GeorceE G. 
1927. Mammalian fauna and correlation of the Paskapoo formation of 
Alberta. Amer. Mus. Novitates, No. 268, pp. 1-10, figs. 1-7. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 53 


1928. A new mammalian fauna from the Fort Union of southern Montana. 
Amer. Mus. Novitates, No. 297, pp. 1-15, figs. I-14. 

1929a. Third contribution to the Fort Union fauna at Bear Creek, Mon- 
tana. Amer. Mus. Novitates, No. 345, pp. I-12, figs. 1-5, 1 chart. 

1929b. A collection of Paleocene mammals from Bear Creek, Montana. 
Ann. Carnegie Mus., vol. 19, No. 2, pp. 115-122, figs. 1-4. 

1935a. The Tiffany fauna, upper Paleocene. 1.—Multituberculata, Marsupi- 
alia, Insectivora, and ? Chiroptera. Amer. Mus. Novitates, No. 795, 
pp. I-19, figs. 1-6. 

1935b. The Tiffany fauna, upper Paleocene. 2.—Structure and relationships 
of Plesiadapis. Amer. Mus. Novitates, No. 816, pp. 1-30, figs. I-II. 

1935¢c. The Tiffany fauna, upper Paleocene. 3.—Primates, Carnivora, Con- 
dylarthra, and Amblypoda. Amer. Mus. Novitates, No. 817, pp. 
1-28, figs. I-14. 

1935d. New Paleocene mammals from the Fort Union of Montana. Proc. 
U. S. Nat. Mus., vol. 83, pp. 221-244. 

1936. A new fauna from the Fort Union of Montana. Amer. Mus. Novi- 
tates, No. 873, pp. 1-27, figs. 1-16. 

1937a. Additions to the upper Paleocene fauna of the Crazy Mountain field. 
Amer. Mus. Novitates, No. 940, pp. I-15, figs. I-4. 

1937b. The Fort Union of the Crazy Mountain field, Montana, and its mam- 
malian faunas. U. S. Nat. Mus. Bull. 169, pp. 1-287, figs. 1-80, 
pls. I-10. 

1937c. Notes on the Clark Fork, upper Paleocene, fauna. Amer. Mus. Novi- 
tates, No. 954, pp. 1-24, figs. 1-6. 

1940. Studies on earliest Primates. Bull. Amer. Mus. Nat. Hist., vol. 77, 
art. 4, pp. 185-212, figs. 1-8. 

Witson, Rospert W. 

1952. Paleocene mammalian genus Ellipsodon. Bull. Geol. Soc. Amer., 

vol. 63, No. 12, pt. 2, pp. 1315-1316 (abstract). 


EXPLANATION OF PLATES 
PLATE I 
MULTITUBERCULATES AND INSECTIVORES FROM THE BISON BASIN PALEOCENE 


Fig. 1. Cf. Ptilodus montanus Douglass: Left Ps (U.S.N.M. No. 20877), lateral 
view. Four times natural size. 

Fig. 2. Cf. Ectypodus hazeni Jepsen: Left Ps (U.S.N.M. No. 20878), lateral 
view. Four times natural size. 

Fig. 3. Cf. Ectypodus musculus Matthew and Granger: Left ramus of mandible 
with M, (U. of Wyo. No. 1105), occlusal view. Four times natural size. 
Fig. 4. Cf. Anconodon russelli (Simpson): Right ramus of mandible with Ps 

(U. of Wyo. No. 1065), lateral view. Four times natural size. 

Fig. 5. Bisonalveus browni, new genus and species: Left ramus of mandible 
(U.S.N.M. No. 20928), type specimen, lingual and occlusal views. Four 
times natural size. 

Fig. 6. Diacondon pearcei, new species: Left ramus of mandible (U.S.N.M. 
No. 20970), type specimen, lingual and occlusal views. Four times natural 
size. 


PLATE 2 
PRIMATES AND MARSUPIALS FROM THE BISON BASIN PALEOCENE 


Figs. 1, 2. Pronothodectes, cf. matthewi Gidley: 1, Left ramus of mandible 
(U.S.N.M. No. 20758), lateral and occlusal views; 2, right ramus of man- 
dible (U. of Wyo. No. 1062), lateral and occlusal views. Three times 
natural size. 

Fig. 3. Plesiadapis, cf. fodinatus Jepsen: Left ramus of mandible (U.S.N.M. 
No. 20784), lateral and occlusal views. Three times natural size. 

Figs. 4, 5. Peradectes pauli, new species: 4, Left ramus of mandible (U.S.N.M. 
No. 20879), type specimen, lingual and occlusal views; 5, left ramus of 
mandible (U.S.N.M. No. 20880), lingual and occlusal views. Four times 
natural size. 

Fig. 6. Peradectes elegans Matthew and Granger: Right ramus of mandible 
(U. of Wyo. No. 1104), lingual and occlusal views. Four times natural size. 


PLATE 3 
PRONOTHODECTES FROM THE BISON BASIN PALEOCENE 


Figs. 1, 3. Pronothodectes simpsoni, new species: 1, Right ramus of mandible 
(U.S.N.M. No. 20754), type specimen, lateral and occlusal views; 3, right 
ramus of mandible (U.S.N.M. No. 20770), lateral and occlusal views. Three 
times natural size. 

Fig. 2. Pronothodectes, cf. simpsoni, new species: Left ramus of mandible 
(U. of Wyo. No. 1057), lateral and occlusal views. Three times natural 
size, 


54 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 55 


PLATE 4 
PLESIADAPIS FROM THE BISON BASIN PALEOCENE 


Figs. 1-3. Plestadapis jepseni, new species: 1, Left ramus of mandible 
(U.S.N.M. No. 20586), lateral and occlusal views ; 2, left maxilla (U.S.N.M. 
No. 20781), occlusal view; 3, left ramus of mandible (U.S.N.M. No. 20760), 
type specimen, lateral and occlusal views. Three times natural size. 


PLATE 5 
TRICENTES AND CHRIACUS FROM THE BISON BASIN PALEOCENE 


Figs. 1, 2. Chriacus, near C. pelvidens (Cope): 1, Right ramus of mandible 
(U.S.N.M. No. 20983), lateral and occlusal views; 2, left M*? (U.S.N.M. 
No. 21003), occlusal view. Two and one-half times natural size. 

Fig. 3. Chriacus, sp: Left M®? (U.S.N.M. No. 21019), occlusal view. Two and 
one-half times natural size. 

Fig. 4. Tricentes fremontensis, new species: Left ramus of mandible (U.S.N.M. 
No. 20582), type specimen, lateral and occlusal views. Two and one-half 
times natural size. 


PLATE 6 
THRYPTACODON FROM THE BISON BASIN PALEOCENE 


Fig. 1. Thryptacodon belli, new species: Right ramus of mandible (U. of Wyo. 
No. 1045), type specimen, lateral and occlusal views. Twice natural size. 
Fic. 2. Thryptacodon, cf. demari, new species: Left maxilla (U.S.N.M. 
No. 20984), occlusal view. Twice natural size. 

Fig. 3. Thryptacodon demari, new species: Right ramus of mandible (U.S.N.M. 
No. 20985), type specimen, lateral and occlusal views. Twice natural size. 

Fig. 4. Thryptacodon, cf. belli, new species: Left maxilla (U.S.N.M. No. 20986), 
occlusal view. Twice natural size. 

Fig. 5. Thryptacodon, cf. australis Simpson: Left ramus of mandible (U. of 
Wyo. No. 1076), occlusal and lateral views. Twice natural size. 


PLATE 7 
CLAENODON FROM THE BISON BASIN PALEOCENE 


Figs. 1, 6. Claenodon acrogenius, new species: 1, Left ramus of mandible 
(U.S.N.M. No. 20575), occlusal view, natural size; 6, right ramus of man- 
dible (U.S.N.M. No. 20634), type specimen, lateral view, one-half natural 
size. 

Figs. 2, 3. Claenodon, cf. ferox (Cope): 2, Left ramus of mandible (U.S.N.M. 
No. 20633), occlusal view; 3, right maxilla (U.S.N.M. No. 20797), occlusal 
view. Natural size. 

Fig. 4. Claenodon, cf. montanensis (Gidley) : Left ramus of mandible (U.S.N.M. 
No. 20574), occlusal view. Natural size. 

Fig. 5. Claenodon, cf. procyonoides (Matthew): Right ramus of mandible 
(U.S.N.M. No. 20630), lateral and occlusal views. Twice natural size. 


56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


PLATE 8 
LITOMYLUS AND PROTOSELENE? FROM THE BISON BASIN PALEOCENE 


Figs. 1, 3. Protoselene? novissimus, new species: 1, Left ramus of mandible 
(U.S.N.M. No. 20572), type specimen, lateral and occlusal views; 3, right 
ramus of mandible (U.S.N.M. No. 21023), lateral and occlusal views. Four 
times natural size. 

Fig. 2. Litomylus, cf. scaphicus, new species: Right M* or M?® (U.S.N.M. 
No. 21013), occlusal view. Four times natural size. 

Fig. 4. Litomylus scaphicus, new species: Right ramus of mandible (U.S.N.M. 
No. 21014), type specimen, lateral and occlusal views. Four times natural 
size. 

Fig. 5. Litomylus scaphiscus, new species: Right ramus of mandible (U.S.N.M. 
No. 21010), type specimen, lateral and occlusal views. Four times natural 
size. 


PLATE 9 “ 
HAPLALETES AND GIDLEYINA FROM THE BISON BASIN PALEOCENE 


Fig. 1. Haplaletes pelicatus, new species: Left ramus of mandible (U.S.N.M. 
No. 21008), type specimen, lateral and occlusal views. Four times natural 
size, 

Fig. 2. Haplaletes serior, new species: Left ramus of mandible (U. of Wyo. 
No. 1078), type specimen, lateral and occlusal views. Four times natural 
size. 

Figs. 3, 4. Gidleyina wyomingensis, new species: 3, Right ramus of mandible 
(U.S.N.M. No. 20790), type specimen, lateral and occlusal views; 4, right 
maxilla (U.S.N.M. No. 20795), occlusal view. Twice natural size. 


PLATE 10 
PHENACODUS? FROM THE BISON BASIN PALEOCENE 


Figs. 1-3. Phenacodus? bisonensis, new species: 1, Right ramus of mandible 
(U.S.N.M. No. 20567), lateral and occlusal views, 14 times natural size; 
2, right maxilla (U.S.N.M. No. 20564), type specimen, occlusal view, 14 
times natural size; 3, right maxilla (U.S.N.M. No. 20566), occlusal view, 
twice natural size. 

Figs. 4, 5. Phenacodus? sp. (large): 4, Right ramus of mandible (U.S.N.M. 
No. 21025), occlusal view, 14 times natural size; 5, right P* (U.S.N.M. No. 
21038), occlusal view, twice natural size. 


PLATE II 
CONDYLARTHS AND TITANOIDES FROM THE BISON BASIN PALEOCENE 


Figs. 1, 2. Promioclaenus pipiringosi, new species: 1, Right ramus of mandible 
(U.S.N.M. No. 20571), type specimen, lateral and occlusal views; 2, right 
ramus of mandible (U.S.N.M. No. 21021), lateral and occlusal views. Four 
times natural size. 

Figs. 3, 4. Litolestes lacunatus, new species: 3, Left ramus of mandible 
(U.S.N.M. No. 21016), type specimen, lateral and occlusal views; 4, left 
ramus of mandible (U. of Wyo. No. 1079), lateral and occlusal views. Four 
times natural size. 


No. 6 PALEOCENE FAUNAS OF BISON BASIN—GAZIN 57 


Fig. 5. Titanoides primaevus Gidley: Right upper molars (U. of Wyo. No. 
1093), occlusal view. Natural size. 


PLATE 12 
CAENOLAMBDA FROM THE BISON BASIN PALEOCENE 


Caenolambda pattersoni, new genus and species: Skull (U.S.N.M. No. 21036), 
type specimen, dorsal view. One-half natural size. 


PLATE 13 
CAENOLAMBDA FROM THE BISON BASIN PALEOCENE 


Caenolambda pattersoni, new genus and species: Skull (U.S.N.M. No. 21036), 
type specimen, lateral view. One-half natural size. 


PLATE 14 
CAENOLAMBDA FROM THE BISON BASIN PALEOCENE 


Caenolambda pattersoni, new genus and species: Skull (U.S.N.M. No. 21036), 
type specimen, ventral view. One-half natural size. 


PLATE I5 
SOUTH RIM OF BISON BASIN SHOWING FOSSIL LOCALITIES 


View westward along escarpment forming south rim of Bison basin. Fossil 
localities indicated are as follows: a, saddle locality; b, ledge locality; 
c, Titanoides locality ; and d, west-end locality. 


PLATE 16 
TWO FOSSIL LOCALITIES IN THE BISON BASIN 


Above, view eastward of ledge locality (b). Below, view southwestward of west- 
end locality (d). 


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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 6, PL. 1 





MULTITUBERCULATES AND INSECTIVORES FROM THE BISON BASIN PALEOCENE 


(SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 6, PL. 2 








PRIMATES AND MARSUPIALS FROM THE BISON BASIN PALEOCENE 


SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 6, PL.3 





PRONOTHODECTES FROM THE BISON BASIN PALEOCENE 


(SEE EXPLANATION_AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 132, NO. 6, PL. 4 





PLESIADAPIS FROM THE BISON BASIN PALEOCENE 


(SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131. NO:.'6,.PL. 5 





TRICENTES AND CHRIACUS FROM THE BISON BASIN PALEOCENE 


(SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 131, NO. 6, PL. 6 





\ at) p= 


THRYPTACODON FROM THE BISON BASIN PALEOCENE 


(SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 6, PL. 7 





a 


Ber \- 
ee 


iw 
/ 








CLAENODON FROM THE BISON BASIN PALEOCENE 


(SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 6, PL. 8 


= 


(eR La nals usiade ac HSM 





LITOMYLUS AND PROTOSELENE? FROM THE BISON BASIN PALEOCENE 
(SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 6, PL. 9 





HAPLALETES AND GIDLEYINA FROM THE BISON BASIN PALEOCENE 


(SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 6, PL. 10 








PHENACODUS? FROM THE BISON BASIN PALEOCENE 


SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 6, PL. 11 





CONDYLARTHS AND TITANOIDES FROM THE BISON BASIN PALEOCENE 


(SEE EXPLANATION AT END OF TEXT.) 


MITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 6, PL. 12 





CAENOLAMBDA FROM THE BISON BASIN PALEOCENE 


SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 6 





CAENOLAMBDA FROM THE BISON BASIN PALEOCENE 


(SEE EXPLANATION AT END OF TEXT.) 


PE 


1 


3 


VOL, 131, NO, 6, PL. 14 


2 


MISCELLANEOUS COLLECTIONS 


ONIAN 


SMITHS 





CAENOLAMBDA FROM THE BISON BASIN PALEOCENE 


NATION AT END OF TEXT.) 


SEE EXPLA 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


esate 


(‘LXAL AO GNA Lv NOILVNW1dX4 33S) 


S3ILITVWDO7 TISSO4 DNIMOHS NISvVg NOs!ig AO WIM HLNOS 





SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 6, PL. 16 


ee 





Two FOSSIL LOCALITIES IN THE BISON BASIN 


SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 7 


Charles BD. and Mary Waux Talcott 
Research Fund 


THE UPPER PALEOQCENE MAMMALIA 
FROM THE ALMY FORMATION IN 
WESTERN WYOMING 


(Wirn 2 Prates) 


By 
C. LEWIS GAZIN 


Curator, Division of Vertebrate Paleontology 
United States National Museum 
Smithsonian Institution 





(Pusnication 4252) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
JULY 31, 1956 





THE LORD BALTIMORE PRESS, INC. 


BALTIMORE, MD., U.S.A, 
aight? 





CONTENTS 


Pntraduction and history Of. isivestiSations we cel oiisicals erases 6 csccveecwe 
Geologic relations and occurrence of remains..............0ccccceeeeees 
BINED eee AN ara Vputich LITT oA reravs ecetovefee av eresevele aie eraterceres (ofeach ore te ocioie: ott cia oavekn, ices 
Eee vang Correlation Of the fase so. cane s.0oc eons ce aieses cin esis Koss ws sie 
Systematic description of the mammalian remains...................04- 
WATARIARES) Ponca ee 6 cree eis wesldas Te ea ee alo ore Baia inlois ba nuh bac ae 

Be eeaeAes TRE er ie Rrart ei kai ek shaken aia ola wake ee wid olaa.d Bee etete nahin sé 
Gre ae ae atts hte ut, seh ae pula cat als 8 acide Aware ws ale arte 

BEA EMA WO eiatcre om Nard eins ettoins RNS RIEU eed IOI owed plaiseieeo Siew oe 
PPE LORY ORIG A, Sat are re Bale ate cw ale stern oie ever sialaahd ole ete Sintec Mes 
esompcniciae yt Cheers «eisai cle es cis vik tha bed wis hep we Seeds areas 
MSA A AES ag arcs ch a io tos aie Apts ats 1e:ar aarctniere Vn aueiec ovale oo whe ator elavasajarsinie aca 

MRAM EANNREN Vide fog ashes veretes costal coeses ant asaies at elav ovale nis al SR ee Daw 8 ces 
APPEAL O AAG © oie tae adscns tie. aaa ool ass sie ieraiden ol oelo tie. eiova Wlaieea ethic e 

PRA ACA aie ore einen. ce sie cites ns asa RI ee MeCN tee ao ee ee relte eee 


ILLUSTRATIONS 
PLATES 
(Following p. 18) 


1. Primates, Anacodon? and Probathyopsis? from the Almy Paleocene. 
2. Condylarths from the Almy Paleocene. 


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Charles BD. and Mary Waux Walcott Research Fund 


THE UPPER PALEOCENE MAMMALTA FROM 
THE ALMY FORMATION IN 
WESTERN WYOMING 


By C. LEWIS GAZIN 


Curator, Division of Vertebrate Paleontology 
United States National Museum 
Smithsonian Institution 


(WirtH 2 PLATES) 


INTRODUCTION AND HISTORY OF INVESTIGATION 


Repeated, intensive search of a comparatively small exposure area 
of the Almy formation in western Wyoming has, over the past 15 
years, resulted in a faunal representation of about a dozen mammalian 
species. This is, no doubt, a rather meager sample of the probable 
fauna although based on a little over 70 determinable specimens. It is, 
nevertheless, an interesting increase, from the original five forms 
recognized (Gazin, 1942) on but nine specimens. The Clarkforkian 
upper Paleocene age interpreted for the scant, earlier materials now 
seems clearly indicated by the collections as a whole. 

The locality consists of a small cluster of closely adjacent exposures 
on the north side of La Barge Creek about 7 miles due west of the 
town of La Barge, formerly Tulsa P. O., in Lincoln County, Wyo. 
The most productive of these has been one in the vicinity of a topo- 
graphic saddle, bare of vegetation, at the head of a ridge along the 
southeast side of Buckman Hollow (see advance sheet, U.S.G.S. 
La Barge quadrangle) in NW4NE} sec. 12, T.26 N., R.114 W. 
Other localities worthy of mention are on the southeast side of the 
above ridge, nearer the highway, and on the ridges to the northwest 
of Buckman Hollow in the vicinity of Spring Creek. 

Discovery of these localities, as has been previously noted, was 
made by J. B. Reeside, Jr., B. N. Moore, and W. W. Rubey of the 
U. S. Geological Survey in 1936. Discovery by Rubey and John 
Rodgers in 1939 of Plesiadapis material at one of the sites provoked 
our interest, and in 1941 an additional small collection was made by 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 131, NO. 7 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


G. F. Sternberg, Franklin Pearce, and myself. The result of these 
early searches was the nine specimens described in the preliminary 
note of 1942. Smithsonian Institution parties revisited the localities 
in 1948, 1949, 1951, 1953, and 1954. On most of these expeditions 
I was assisted by the chief of our laboratory of vertebrate paleon- 
tology, Franklin L. Pearce. In 1948, I was aided by my wife, Elisa- 
beth, and son, Chester. Chester Gazin also assisted Pearce and me 
in 1949. 

The excellent pencil drawings of specimens shown in the two plates 
accompanying this report were made by Lawrence B. Isham, scien- 
tific illustrator for the Department of Geology in the National 
Museum. 


GEOLOGIC RELATIONS AND OCCURRENCE OF REMAINS 


Although the fossil-bearing beds on La Barge Creek have been 
mapped by A. R. Schultz (1914) as Almy, and are so regarded by 
Rubey + in his recent investigations of the region, it should be noted 
that the type section for the Almy formation, in the vicinity of Evans- 
ton, Wyo., is in a separate, although adjacent, basin of Tertiary dep- 
osition and there may have been no actual continuity between the 
two lithologically somewhat similar deposits. 

The Almy formation is mapped in the Upper Green River Basin 
as a nearly continuous band along the east flank of the Wyoming 
Range from the vicinity of La Barge Creek to Fall River in the 
Hoback Basin. At La Barge Creek it appears and is shown by 
Schultz to be in depositional contact with lower Paleozoic rocks to 
the east and with upper Paleozoic and Triassic rocks forming the 
front of the range to the west. The Almy area immediately to the 
north of La Barge Creek is separated from the Eocene of the Green 
River Basin on the east by faulting, shown as a thrust by Schultz in 
which the lower Paleozoic beds underlying the Almy have ridden out 
over the younger rocks to the east. To the south of La Barge Creek 
the Paleozoic rocks which make up La Barge or Hogsback Ridge, 
together with the trace of the thrust fault, disappear beneath the 
Eocene of the Green River Basin, with the Knight formation extend- 
ing westward to contact with the Almy, as it does again some distance 
to the north. 

In the vicinity of the fossil occurrences the Almy beds are a red- 
dish, pebbly clay, partly conglomeratic, dipping steeply to the south- 
west toward La Barge Creek. They appear to be nearly conformable 


1 Oral communication. 


NO. 7 MAMMALIA FROM THE ALMY FORMATION—GAZIN 3 


with the underlying Paleozoic limestones in Buckman Hollow, a rela- 
tionship, of course, of a strictly local character. The various fossil 
sites are nearly all very low in the section and at the topographic 
saddle formed at the head of the ridge bounding Buckman Hollow on 
the southeast scattered remains were found to within only a few feet 
of the underlying limestone. A locality on the southeast side of this 
ridge and nearer the road, which produced the type of Phenacodus 
almiensis, would appear to be a little higher in the section. P. al- 
miensis, however, is well represented by materials from the lowest 
levels so that the stratigraphic difference in this instance would not 
appear to have faunal significance. 


THE ALMY FAUNA 


There follows a listing of the forms encountered in the Almy col- 
lections and an indication of the number of specimens recognized as 
representing each: 


PRIMATES : 
Plesiadapidae: 
Fhe HGHOESS  FMUEYS NAZI He 2)F soto Sale 8 LIPO Le S's ARRAS Se AE SG 3's ceeds az 
LEStOdG Pes SCOOK EY I CDSEI < sicForcts oioterale ciate hole sieiateswit o'er dich ye ayeyeuerersiore 3 
Plesiadapis? ,pearcet, MEW SPCCIES....« <is'evssiceincde nas pesiceescds aes 2 
Carpolestidae : 
ae HOIESIPS, (Ch. AUIS NEDSEM « ocicion'e-<iaarwe.s inlaws o.8 veiss vas deste ties I 
CARNIVORA: 
Arctocyonidae: 
ANGLODONE REGUS MEW: SPECIES ).a)d « a.s}ere,cicistetoyaserreieyoe stele ainrs oveiey else I 
Mesonychidae: 
NOUSICSES: SINE Cia A tatard aii ile ats N50 Sts Site 5 Up ain nibs athe a9, «6 4, ai'aiel @ 3/8. 2 
Miacidae: 
WHE PINICHLST ¢ SDS Saeed tee cee cere eats CE Ses te tate gai ota I 
CoNDYLARTHRA: 
Phenacodontidae : 
EGVOCALTR. TOUS ON ENStSe GLAMIS Clie. etavare ehciehounsarenes0 ei oxcyaitiniotens eisie cfeceragahets 13 
Eglorian, Ct. OSPOTMUIEN CCODE) siny os ss piccg cata seine ans Anigaies 4 
PEMACOMUS.  CLINGENSUS GAZA s fe erat cis ce dics ielsidinie vie ciavevae 94 ese eis 32 
EP ieeaCOIUs Preetg CONG io kaa e s:sie win j0<:6;d) 5 aiaidio)s Bios a: alld dela, wl.ace 10 * 
DINOCERATA : 
Uintatheriidae : 
EPODELAY OP SIS ca SD cre tent nici lated 4c pretateietas iaiolat eiestelersreitelsisie'aattyatele's 2 


* Eight of these are of a smaller form tentatively regarded as P. p., cf. intermedius. 


AGE AND CORRELATION OF THE FAUNA 


The fauna above listed is beyond doubt a Clarkforkian assemblage. 
It is interesting to note, moreover, that all the genera, except Car- 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


polestes, and certain of the species are also lower Eocene. Neverthe- 
less, this fact, together with the relative abundance of condylarths and 
Plesiadapis, and absence of the more common Eocene forms such as 
Hyracotherium, Homogalax, Hyopsodus, Diacodexis, Pelycodus, etc., 
is regarded as characterizing Clark Fork time. 

It is of further interest that all the genera here recognized are also 
either Tiffanian or, as Ectocion and Anacodon, arbitrarily distin- 
guished from Tiffanian ancestral forms. This is regarded as further 
characteristic of Clark Fork time, i.e., the bulk of the Clark Fork 
fauna is a survival of a certain selection of known Tiffanian lines with 
the appearance of very few new stems from elsewhere. It would 
appear then that recognition of Clarkforkian time as distinct from 
Tiffanian on a generic level is, as in comparison with the Eocene, 
somewhat negative in character, partly depending upon the absence 
of a number of forms comparatively common in the earlier beds but 
presumed to have become extinct. Nevertheless, the species for the 
most part are advanced over those of Tiffanian time and we have in 
the Almy, for example, such forms as Plesiadapis cooket, Anacodon? 
nexus, Ectocion ralstonensis, E., cf. osbornianum, Phenacodus almien- 
sis and P. primaevus. From the Clark Fork beds in the Big Horn 
Basin there may be added to this list of progressive species such forms 
as Thryptacodon antiquus, Didymictis protenus, Haplomylus spetrianus, 
and a species of Coryphodon. 

The further evidence given the distinctiveness of Clark Fork time 
by the first known appearance of tillodonts and palaeanodonts calls 
attention to the comparatively few new lines that appear, evidently 
introduced from some other area, in contrast with the strikingly large 
and important part of the Eocene fauna that appeared at the end of 
Clark Fork time. This emphasizes the appropriateness and, undoubt- 
edly, formed part of the reasoning followed in regarding Clarkforkian 
as upper Paleocene rather than Eocene, thereby permitting factors of 
a regional or perhaps greater importance that must have affected the 
faunal distribution to be correlated with an important time boundary. 


SYSTEMATIC DESCRIPTION OF THE MAMMALIAN REMAINS 
PRIMATES 
PLESIADAPIDAE 
PLESIADAPIS RUBEYI Gazin, 1942 
Plate 1, figure 10 


No additional material of this species has been found since descrip- 
tion of the original Geological Survey collection. Included is the type 


NO. 7 MAMMALIA FROM THE ALMY FORMATION—GAZIN 5 


(U.S.N.M. No. 16696), a right mandibular ramus with P;-Mz, but 
lacking the trigonid of M,, and a left M® tentatively referred to 
Plesiadapis rubeyt. 

P. rubeyi clearly belongs to the group of species that includes 
P. gidleyi, P. fodinatus, P. dubius, and probably P. cooket. It is 
remote from the distinctive P. jepseni—P. anceps—P. rex group or 
subgenus. It is, moreover, rather close to P. fodinatus which Jepsen 
(1930) described from the Silver Coulee horizon of the Polecat Bench 
sequence. There is a possibility that P. rubeyi is not specifically dis- 
tinct from P. fodinatus; however, in view of the distinctly small size 
of M,, the anteroposteriorly shorter appearing summit of the trigonid 
of M., and the comparatively slender premolars showing an incipient 
metaconid on P, (as in P. dubius rather than P. fodinatus), the 
species P. rubeyi would seem to be valid. Moreover, P. fodinatus is 
typically Tiffanian in age, regarded as represented in the Bison Basin 
deposits and Fossil Basin Evanston(?) as well as in the Polecat 
Bench, and survival of this species into Clarkforkian time, though 
likely, awaits demonstration. 

The tentatively referred last upper molar (U.S.N.M. No. 16697) 
is distinctly large for the type lower jaw of P. rubeyi and is postero- 
lingually expanded somewhat as in P. fodinatus. It is possible that 
this tooth represents P. fodinatus, but the evidence is rather meager 
and would not seem to justify separate listing. The tooth measures 
4.3 mm. anteroposteriorly by 6.0 for the greatest transverse diameter. 


MEASUREMENTS IN MILLIMETERS OF LOWER TEETH IN TYPE SPECIMEN OF 
Plesiadapis rubeyi, U.S.N.M. No. 166096 


Ps, anteroposterior diameter: transverse diameter..............eeeeees 2.8: 1.9 
P,, anteroposterior diameter: transverse diameter............+--eeeeee- 2G a2 
ie itausverse diameter OF LAIOWIGs pave. ccd eauis jcctoeccieg ea verse sencis 27 
M;, anteroposterior diameter : transverse diameter of talonid........... B72 


PLESIADAPIS COOKEI Jepsen, 1930 
Plate 1, figures 5-8 


In addition to the lower jaw of Plesiadapis cookei (U.S.N.M. 
No. 16698) found in 1941, a second lower jaw with all three molars 
and an isolated M® were found by Franklin Pearce while in the field 
with me in 1954. Plesiadapis cookei is truly gigantic in comparison 
with other Paleocene primates and is nearly as large as the upper 
Bridgerian Notharctus robustior. Direct comparison of these jaw 
materials with the type specimen in the collections of Princeton Uni- 
versity shows near identity in size and character of the teeth for the 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


I94I specimen, verifying the tentative assignment of the Buckman 
Hollow Almy form made in 1942. U.S.N.M. No. 20785 has lower 
molars a little broader than in the type, perhaps more noticeable in 
M;, but no doubt this is within the range of individual variation. 

The last upper molar (U.S.N.M. No. 21281) is considerably larger 
than that (No. 16697) tentatively assigned to P. rubeyi but relatively 
does not show so marked a posterior extension of the posterolingual 
portion. It is also less expanded in this respect than in the type ma- 
terial of P. cookei. Its measurements are 6.8 mm. anteroposteriorly 
by 9.8 for the greatest diameter. 


MEASUREMENTS IN MILLIMETERS OF LOWER TEETH IN SPECIMENS OF 
Plasiadapis cookei 


U.S.N.M. U.S.N.M. 
No. 16698 No. 20785 
Ps, anteroposterior diameter: transverse diameter... 5.2: 3.7 
Py canteropostenion) GiamMeter’ «.o:..0:5.¢.cs0is6,areiapoicts sole epsieie 53 
M,, anteroposterior diameter: transverse diameter of 
PAIOBIG! <a ai crojorn.s a:0)6)0's.as's's nase Gs new oe ee case eetmae ©, wane siete 6.1: 5.6 
Ms, anteroposterior diameter: transverse diameter of 
Halonidl ails act ietat tes de coeie tre ates Oats ee ineieioes 6.4: 5.5 6.4: 6.3 
Ms, anteroposterior diameter: transverse diameter of 
ETA ORIG risa 5 bo le o.0:s sites erioin vis. ale ole eve ave Giaretelamte TOSS siete 10.0% : 6.2 


® Approximate. 


PLESIADAPIS? PEARCEI,? new species 
Plate 1, figure 9 


Type.—Right ramus of mandible (U.S.N.M. No. 20787), with Mi 
and M,. 

Horizon and locality—Buckman Hollow Clarkforkian Paleocene 
on La Barge Creek, NW4NE} sec. 12, T.26 N., R.114 W., Lincoln 
County, Wyo. 

Specific characters.—Size close to that of Plesiadapis dubius, but 
trigonids of lower molars narrower and talonids wider than in that 
species. Apices of cusps on trigonid more widely spaced transversely 
and entoconid of M, and M, distinctly more posterolingual in position. 

Discussion —The two lower jaw portions considered to represent 
this peculiarly distinctive form were at first allocated to P. rubeyt 
but their smaller size coupled with the lingually and backward-jutting 
entoconid position apparently precludes this possibility. The position 
of the entoconid gives the talonid of the first two lower molars a 
relatively marked width in contrast with the narrow trigonid, com- 


2 Named for Franklin L. Pearce who found the type specimen. 


INO: 57, MAMMALIA FROM THE ALMY FORMATION—GAZIN 7. 


pared, for example, with P. dubius, which this form approaches in 
length of lower molars. Moreover, although the trigonid is narrower 
at its base than in P. dubius, the apices of the cusps are more widely 
spaced transversely, and in M, the paraconid is farther forward. In 
Mz, however, the paraconid is not farther forward with respect to 
the metaconid than in P. dubius. 

The peculiarities outlined above tempt speculation on the possibility 
that an undescribed genus is represented. I believe, however, that the 
differences here noted are probably of no greater significance than 
(and quite opposite in general tendency to) the markedly sloping 
outer walls of lower cheek teeth seen in the P. jepseni—P. anceps— 
P. rex group, presumably no more than subgeneric in importance. 


MEASUREMENTS IN MILLIMETERS OF LOWER TEETH IN SPECIMENS OF 
Plesiadapis? pearcei 
U.S.N.M. 


No. 20787 U.S.N.M. 
Type No. 20786 
IMaanteroposterion diameter .iics\. ccrcs cicvereielo odierewaie vieisiaieis sce ar 3:2 
My transverse diameter of trigonididiis.3.bes icc casesseccs ar 2.0 
M;, transverse diameter of talonid 0 oi... c dois lect cece 2.5 2.5 
Rs StECrOpOsteniOr CIAMIGLET saci co tietisice ceed se cbse canna 3.4 
M,, transverse diameter of trigonid...06.i.c5ccsccccedaceas 2.4 
M,. transverse’ diameter of ‘talonid.. . é «ss: edi dares dslnemwae 2.8 


CARPOLESTIDAE 
CARPOLESTES, cf. DUBIUS Jepsen, 1930 
Plate 1, figure 4 


A carpolestid P,; (U.S.N.M. No. 21280) in the collection can be 
closely matched in size by specimens of Carpolestes dubius. The tooth 
shows a high, uniformly convex crest in lateral view with scarcely 
discernible vertical ridges. There would appear to be about eight 
feeble serrations in advance of the position of the heel which is broken 
away. In lingual view the vertical ridges are a little more visible and 
the height of the crown is less, but with possibly less difference in 
height between the two sides than in the Polecat Bench material. The 
posterior portion of the lingual surface is gently concave, whereas the 
labial wall is slightly convex in vertical profile. In a dorsal view the 
crown appears slightly bilobed with the greatest width across the 
posterior portion. There is no distinct cingulum labially, and lingually 
a cingulum is perhaps feebly defined posteriorly. 

The Almy tooth is distinctly larger and higher crowned than the 
corresponding tooth in Carpodaptes hazelae. It also has a greater 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


number of serrations of smaller size and the associated ridges are less 
clearly defined. U.S.N.M. No. 21280 is certainly much closer to 
Carpolestes dubius than to any of the other known carpolestids, and 
it seems, moreover, that C. dubius is somewhat more removed from 
Carpodaptes in the form of P, than is the genotype Carpolestes 
nigridens. 

Carpolestes dubius is recorded by Jepsen (1930) from the Clark 
Fork beds as well as the Tiffanian portion of the Polecat Bench 
sequence. 

The Almy P, measures 2.9 mm. from its anterior margin to the 
posterior root at the alveolus. The width is approximately 1.8 mm. 
across the posterior portion of the base of the tooth. 


CARNIVORA 
ARCTOCYONIDAE 
ANACODON? NEXUS,? new species 
Plate 1, figure I 


Type.—Left ramus of mandible (U.S.N.M. No. 21282) with M, 
and Ms. 

Horizon and locality—Buckman Hollow Clarkforkian Paleocene 
on La Barge Creek, NW4NE3% sec. 12, T.26 N., R.114 W., Lincoln 
County, Wyo. 

Specific characters.—Size considerably smaller than Anacodon ursi- 
dens, but teeth only slightly larger than in type of Claenodon mon- 
tanensis. Primary cusp pattern of lower molars better defined than 
in Anacodon ursidens or A. cultridens, but trigonid less elevated above 
talonid than in Claenodon montanensis. Also, anterior crest from 
hypoconid low but joins protoconid at a completely lingual position 
so that inner wall of first two lower molars shows little flexure mid- 
way of its length. 

Discussion—Anacodon? nexus is considered as possibly repre- 
senting that genus rather than Claenodon as earlier (Gazin, 1956 and 
in press) supposed, because of the lowness of the trigonid on M, as 
well as Mz. Also the crista obliqua has entirely lost its oblique char- 
acter or has been overshadowed by the development of a distinctly 
lateral spur or crest extending forward from the hypoconid to the 
posterolateral surface of the protoconid, one of the several possibili- 
ties afforded by the crenulated character of the principal cusps in the 


8nexus (L.)=tie, bind, with reference to its intermediate position between 
Claenodon and Anacodon. 


NO. 7 MAMMALIA FROM THE ALMY FORMATION—GAZIN 9 


Claenodon line. The result of this is a broadening of the basin of the 
talonid giving it a rather different appearance than, for example, in 
Claenodon montanensis. Nevertheless, the lower molars have retained 
clear definition of the principal cusps, showing the Claenodon pattern, 
which is nearly lost in the crenulate character of the more flattened 
tooth crowns of Eocene Anacodon. 

The character of the anterior portion of the jaw in Anacodon? 
nexus cannot be determined, nevertheless the reduction of the anterior 
premolars, the development of a diastema behind the canine, and a 
flange on the lower jaw below the symphysis characteristic of Ana- 
codon has already been anticipated in Claenodon acrogenius of the 
lower Tiffanian in the Bison Basin. However, in C. acrogenius the 
flange is comparatively incipient and the lower canine is enlarged 
rather than reduced. Moreover, the lower molars of C. acrogenius, 
except for size, would appear to be indistinguishable from those in 
other species of Claenodon. I suspect that the anterior portion of the 
lower jaw of Anacodon? nexus was deepened and exhibited a di- 
astema behind the canine, although this is not certain, and there 
remains the possibility that A.? nexus is a survival of more typical 
Claenodon with shallow symphysis and unreduced premolars, but 
with the tooth pattern advancing parallel to that leading to Anacodon. 

M, in No. 21282 of Anacodon? nexus measures approximately 
9.5 mm. in length by 7.8 mm. across the talonid. Mz is about 10.7 mm. 
long and 8.5 mm. across the trigonid. 


MESONYCHIDAE 
DISSACUS, sp. 


The upper tooth portion (U.S.N.M. No. 16699) including the pro- 
tocone and metacone, previously (Gazin, 1942) listed as a “creodont, 
gen. and sp. undet.,” may well be an anterior molar of Dissacus. A 
second tooth fragment, the posterior portion of a lower premolar, 
possibly P, or P;, also suggests Dissacus. These are evidently of a 
form not greatly different in size from the Torrejonian Dissacus 
navajovius, clearly smaller than Dissacus praenuntius Matthew of 
the Clark Fork beds. 


MIACIDAE 
DIDYMICTIS?, sp. 


A left M, may well belong to a species of Didymictis, but is very 
much smaller than contemporary Didymictis protenus proteus from 


IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


the Clarkforkian of the Big Horn Basin. It is only a little smaller 
than Torrejonian Didymictis haydenianus, but not to be compared 
with D. microlestes or D. tenuis. Possibly the Almy form is a fore- 
runner of one of the other miacid genera of the Eocene, but the tooth 
in question is not too well preserved so that detailed comparison is 
unwarranted. 


CONDYLARTHRA 
PHENACODONTIDAE 
ECTOCION RALSTONENSIS Granger, 1915 
Plate 2, figures 1 and 2 


Ectocion, though by no means as abundantly represented in number 
of specimens as Phenacodus, has in the material comprising it cer- 
tainly the best specimen in the Almy collection. The skull and jaws 
(U.S.N.M. No. 20736) referred to Ectocion ralstonensis in general 
lack only the right and posterior elements of the cranium and the 
posterior portion of the right ramus of the mandible. The rostrum 
and left side of the cranium and mandible are comparatively well 
preserved, though fractured and slightly distorted. The teeth are for 
the most part in excellent condition and only the upper incisors and 
P! and P? of the right side are missing from the skull. The lower 
jaws have P;-M; preserved in both rami. An incomplete humerus and 
ulna were found associated with the skull. There are, in addition 
to this specimen, about 12 others consisting of jaw and maxillary 
portions and isolated teeth referred to or tentatively identified as 
E. ralstonensis in the Almy collection. 

In a lateral aspect the Ectocion ralstonensis skull reveals certain 
details of interest regarding the foramina, so often not ascertainable 
in Paleocene materials. The anterior opening of the infraorbital 
foramen is directly above the anterior root of P* and well forward 
of the anterior margin of the orbit. Posteriorly this foramen opens 
in the orbital cavity at the anterior apex of the large triangular-shaped 
orbital plate of the maxilla. Superior and somewhat medial to the 
posterior opening of the infraorbital foramen and separated from it 
by a backward and medially extending ridge, which may coincide with 
the sutural ridge of the maxilla, is an aperture believed to be a spheno- 
palatine foramen. Above this and somewhat lateral to it is the lachry- 
mal foramen, concealed in lateral view by the margin of the orbit. 

In the posterior portion of the orbital cavity, the optic foramen is 
well forward—a little less than a centimeter—of the sphenoidal fis- 


NO. 7 MAMMALIA FROM THE ALMY FORMATION—GAZIN re 


sure. About a half centimeter posterolateral and somewhat ventral 
to the sphenoidal fissure is an aperture which is surely the anterior 
opening of an alisphenoid canal. The posterior opening is clearly de- 
fined well forward and ventromedial to the foramen ovale. I am 
unable to determine the presence or absence of a foramen rotundum, 
possibly opening into the alisphenoid canal. According to W. K. 
Gregory (Orders of Mammals, p. 354), a foramen rotundum opened 
into the alisphenoid canal in Phenacodus; however, Simpson (1933), 
in describing an endocranial cast of Phenacodus, shows both first and 
second branches of the trigeminal nerve as having passed through the 
sphenoidal fissure. This would seem to preclude the possibility of a 
distinctly separate foramen rotundum in Phenacodus, which is re- 
garded as closely related to Ectocion. 

Ventrally, the posterior palatine foramen is about opposite the 
posterolingual portion of M,. There is a small, blunt pterygoid proc- 
ess of the maxilla, and opposing it medially is a somewhat everted 
lateral portion of the anterior margin of the posterior narial aperture. 
The nasal cavity is closed below posteriorly to a position about even 
with the posterior margin of the last molar. The previously mentioned 
posterior opening of the alisphenoid canal faces more ventrally and 
well ahead of the foramen ovale, a relative distance nearly as great as 
in Meniscotherium. The postglenoid foramen is large and placed 
posteromedial to the postglenoid process, and the space for the audital 
tube behind the postglenoid process is shallow and broadly open. 

The teeth in U.S.N.M. No. 20736 show the anterior premolars, 
above and below, to be separated from each other and from the canine 
by diastemata, the greatest separation being between the first and 
second premolars, about 4 mm. above and 3 mm. below. The anterior 
premolars above are simple and 2-rooted, whereas P, has but one root. 

The essential difference between Ectocion ralstonensis, as exempli- 
fied by No. 20736, and the Ectocion osbornianum material in the U. S. 
National Museum from the Gray Bull is to be found, in addition to a 
slightly smaller size of the teeth, in the less progressive character of 
the posterior premolars of E. ralstonensis. The tritocone in both P* 
and P* is distinctly less developed and less well separated from the 
primary cusp. This is particularly noticeable in P*. Moreover, the 
anterointernal cusp or protoconule is less developed. It is not present 
on P* and comparatively weak on P* of E. ralstonensis. In E. os- 
bornianum material at hand, the protoconule is generally prominent 
and may be thrust to a decidedly anterolingual position in both P* and 
P*, There is no tetartocone on the posterior upper premolars of 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


No. 20736, but it may be moderately developed on the cingulum of 
P* and sometimes P* in E. osbornianum. 

The lower premolars appear relatively more slender in Ectocion 
ralstonensis than in FE. osbornianum, and P* has a less molariform 
appearing talonid with, as noted by Granger (1915, p. 353), a much 
weaker entoconid than usually seen in E. osbornianum; also E. ral- 
stonensis exhibits a shallower mandible. 

Upper and lower molars of Ectocion ralstonensis are apparently not 
distinctive in comparison with EF. osbornianum, except for the greater 
average size in the latter. However, comparison with Gidleyina wy- 
omingensis (Gazin, 1956) shows that, as earlier stated, the crests 
from the protocone to the protoconule and metaconule of the upper 
molars are better defined in the latter. Moreover, in some specimens 
of Gidleyina wyomingensis and in the types of G. silberlingi and G. 
superior the parastylid crest of the lower molars tends to join the 
metaconid, suggestive of Phenacodus. As noted by Granger, the 
parastylid crest of Ectocion, as far as observed, is separate from the 
metaconid in the lower molars. 


MEASUREMENTS IN MILLIMETERS OF DENTITION IN SPECIMEN OF 
Ectocion ralstonensis, U.S.N.M. NO. 20736 


Length of upper dentition from anterior margin of canine (at alveolus) 


to. posterior miargig Of Ms 2s Pee tice ces heen eee ct ss taken 49.7 
Length of upper cheek tooth series, P*-M*, inclusive.................. 41.8 
Length of upper premolar series, P*-P*, inclusive..............eeeee0: 24.0 
Length of upper molar series, M*-M®, inclusive.............sseeeecees 18.0 
C, anteroposterior diameter at alveolus: transverse diameter at alveolus. 4.1: 28 
P*, anteroposterior diameter: transverse diameter.............eeeeeeee 3.0: 1.5 
P*, anteroposterior diameter: transverse diameter.............eeeceees 4.0: 2.2 
P*, anteroposterior diameter: transverse diameter.............eeeeee0> 5715.5 
P*, anteroposterior diameter: transverse diameter.............20e-ees 6,057.3 
M’, anteroposterior diameter: transverse diameter across anterior por- 

CIE ccs ooctoee tae oo sera «getts vie cia.sias ook eiteielnie Rant ae aerate 6.2: 8.5 
M?, anteroposterior diameter: transverse diameter across anterior por- 

tonnes Pr rere hed rue” ay ee cbr at Be rene Ree 6.2: 9.1 
M®, anteroposterior diameter: greatest transverse diameter............. 4.9: 7.5 
Length of lower cheek tooth series P: (at alveolus)-Ms, inclusive...... 428 
Length of lower premolar series, P: (at alveolus)-Ps, inclusive......... 23.3 
Length of lower molar series, M:i-Ms, inclusive............eeeseeeeees 20.0 
Ps, anteroposterior diameter: transverse diameter.............-e+esee-- 5-5: 3.2 
P,, anteroposterior diameter: transverse diameter..................4-- 6.7: 4.1 
M,, anteroposterior diameter: transverse diameter of talonid........... 6.5 sigan 
M;, anteroposterior diameter: transverse diameter of trigonid.......... 6.5: 5.4 


Ms, anteroposterior diameter: transverse diameter of trigonid.......... 6.7: 4.5 


NO. 7 MAMMALIA FROM THE ALMY FORMATION—GAZIN 13 


ECTOCION, cf. OSBORNIANUM (Cope), 1882 


A right lower jaw fragment with P,-M, (U.S.N.M. No. 20645) 
has teeth more robust than in the jaw belonging to the E. ralstonensis 
skull, and in addition Ps is more progressive with a better developed 
talonid basin and a large entoconid. P,, moreover, has a rather dis- 
tinctly developed paraconid or parastylid. This specimen is tentatively 
regarded as representing the Gray Bull species E. osbornianum. There 
are in addition three other jaw fragments, each with a comparatively 
large molar which may likewise be referred. 

Although Granger (1915) recognized three species of Ectocion in 
the Clark Fork beds, including both E. ralstonensis and E. osborn- 
tanum, Simpson (1937b), in his treatment of the material, believed 
(except for rare E. parvus) that a single species was represented in 
which there was a shift in the mean size, the length of M, for ex- 
ample, between successive horizons from Clark Fork to Lost Cabin 
time. While this seems evident in the demonstration given, I am, 
nevertheless, concerned about the more progressive P, in the larger 
Almy specimen. The character of P, might likewise show marked 
variation within a species, but there are three small-toothed or 
E. ralstonensis specimens which have P, preserved, and in each of these 
this tooth is distinctly less progressive. The correlation may be a 
coincidence, but if not, I am inclined to believe that in this instance 
a distinct species is actually represented. 


MEASUREMENTS IN MILLIMETERS OF TEETH IN THE SPECIMEN OF 
Ectocion, cf. osbornianum, U.S.N.M. NO. 20645 


P,, anteroposterior diameter: transverse diameter of talonid........... PASis Cay 
M,, anteroposterior diameter : transverse diameter of trigonid.......... 6.8: 5.9 
M2, anteroposterior diameter: transverse diameter of trigonid.......... mT O.5 


PHENACODUS ALMIENSIS Gazin, 1942 
Plate 2, figures 3 and 4 


A relatively small species of Phenacodus, P. almiensis, is clearly 
the most abundantly represented form in the fauna. The 32 speci- 
mens in the collection referred to it comprise about 44 percent of the 
total. The type specimen, U.S.N.M. No. 16691, consists of maxillae 
with the canines and P* to M* in a scarcely worn state, together with 
certain limb and vertebral portions. Although collected in 1941, it 
remains after six subsequent collecting trips the best specimen of this 
species extant. 


I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


P. almiensis is much smaller than Phenacodus p. intermedius but 
appreciably larger than P. copei. It differs essentially from the Phena- 
codus primaevus group, other than in size, in better developed ex- 
ternal styles, particularly the parastyle, and in exhibiting slightly more 
crescentic cusps. The protocone in the upper molars, for example, is 
united by better defined crests to the protoconule and metaconule and 
generally with the hypocone as well. The metaconule is about on a 
line between the metacone and hypocone, not posterior to this, as 
frequently observed in P. primaevus, nor so forward as in Ectocion. 

P. almiensis is significantly larger than any of the P. copei material 
observed, and although the latter exhibits fairly prominent external 
styles on the upper molars, the cusps, particularly the protocone, have 
less developed crests than in P. almiensis. Granger (1915) noted that 
the metaconules were weak or absent in P. copei. These are appar- 
ently not reduced in P. almiensis. Moreover, the upper premolars, 
strangely enough, appear more advanced than in P. copet. P® has a 
well-defined and separate tritocone, described as weak in P. copet, and 
this tooth in P. almiensis also has incipient to clearly defined conules 
and tetartocone. P* is distinctly molariform in appearance, and is 
recognized among isolated teeth by the absence of a mesostyle and 
by the somewhat less developed, though by no means weak, hypocone 
(or tetartocone, in upper premolar nomenclature). Both conules are 
present and well defined. 

Compared to earlier species, P. almiensis is distinctly larger than 
P. matthewi, as well as P. gidleyi, and not nearly so robust as 
P. grangeri among the species known from the Colorado Tiffany. 
Moreover, the teeth are relatively not so broad transversely as in 
P. grangeri. The premolars are decidedly more advanced than in 
Phenacodus bisonensts. 

As noted earlier (Gazin, 1942), the teeth of P. almiensis show some 
resemblance to Ectocion in the development of the external styles and 
somewhat crescentic appearance of the cusps; however, I do not 
believe that Ectocion is represented because of the markedly elongate 
(anteroposteriorly) and relatively narrow upper molars, the position 
of the metaconule, and the comparatively unreduced condition of the 
hypocone of M%. Also, in the lower molars the anterior crest joins 
both the protoconid and metaconid, and the hypoconulid is not so 
close to the entoconid as it usually is in Ectocion. 


NO. 7 MAMMALIA FROM THE ALMY FORMATION—GAZIN 15 


MEASUREMENTS OF UPPER TEETH IN SPECIMENS OF 
Phenacodus almiensis 


U.S.N.M. 
No. 16691 U.S.N.M. 
Type No. 21286 

Length of cheek tooth series, P*-M*, inclusive.......... 42.7% 41.0% 
Length of molar series, M*-M®, inclusive............... 26.1% 26.7 
P*, anteroposterior diameter: transverse diameter....... 8.2: 7.4 77"'!S.6 
P*, anteroposterior diameter: transverse diameter....... 8.5: 8.5 7.9: 9.2 
M’, anteroposterior diameter: transverse diameter *..... 9.0: 10.0 9.0: II.0°% 
M*, anteroposterior diameter : transverse diameter *..... 922113 8.8 12'5* 
M®, anteroposterior diameter: transverse diameter *..... 7.7: 10.8 8.93 1035 


@ Approximate. 
* Anteroposterior diameter of upper molars taken perpendicular to anterior margin and 
transverse diameter across anterior portion. 


PHENACODUS PRIMAEVUS Cope, 1873 
Plate 2, figure 5 


Two specimens in the collection may well represent typical Phena- 
codus primaevus. One of these, U.S.N.M. No. 21287, is a lower jaw 
with P; to Mag, inclusive, and the other an incomplete lower molar. 
The teeth in No. 21287 are comparable in size to those in the Clark 
Fork material referred to P. primaevus. The length of the lower 
molars is near the lower limit of the range given for each (Simpson, 
1937b, p. 18) and the widths are nearer the upper limit, suggesting 
relatively broad teeth, not otherwise distinguished from P. primaevus. 

About eight specimens of smaller size, though not comparable to 
P. almiensis, correspond in general proportions to Gray Bull materials 
earlier regarded as Phenacodus intermedius. The dimensions of teeth 
in one of these (U.S.N.M. No. 20644), evidently the largest of the 
group, are given in the accompanying table. In this and others having 
comparable lower molars the teeth are observed to be relatively slen- 
der, particularly in comparison with the larger, broad-toothed form 
discussed above. A single specimen encountered by Simpson (1937b, 
p. 19) in the Clark Fork collections, representing a smaller group 
which approximates the intermediate-sized form in the Almy fauna, 
was regarded by him as Phenacodus primaevus, small var., cf. inter- 
medius. The Almy materials may be treated in a similar manner, for 
taxonomic convenience, because, although the limited Almy materials 
might appear to be clearly defined, I find it difficult to distinguish 
P. intermedius from P. primaevus in the Gray Bull collections. Never- 
theless, I feel rather strongly opposed to a concept which recognizes 
more than one subspecies of the same form coexisting in time and at 
the same geographic locality. 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


MEASUREMENTS IN MILLIMETERS OF CERTAIN LOWER TEETH IN JAWS OF 


Phenacodus 

Pp Ck. Px D3 

P. almiensis intermedius primaevus 

U.S.N.M. U.S.N.M. 

No. 20643 No. 20644 No. 21287 
P,, anteroposterior diameter ......... 9.6 TTS 12.7 
Ps, transverse diameter of talonid..... 6.2 7.8 10.2 
Mz, anteroposterior diameter......... Q.1 11.8 Eur 
M,, transverse diameter of talonid.... 7.4 9.8 in? 
M2, anteroposterior diameter ......... seers 3 theia 12:55 
M2, transverse diameter of trigonid.... ... ana 12.6 

@ Approximate. 
DINOCERATA 
UINTATHERIIDAE 


PROBATHYOPSIS?, sp. 
Plate 1, figures 2 and 3 


Two upper premolars, possibly both P*, or P* and P*, but of differ- 
ent individuals as indicated by wear, are evidently of Probathyopsis. 
They are, however, significantly larger than Probathyopsis praecursor 
Simpson (1929) of the Clark Fork beds. They correspond closely in 
size to a P® belonging with a partial skeleton of cf. Bathyopsis fissi- 
dens Cope from the New Fork upper Wasatchian (Gazin, 1952, 
p. 64), but are slightly more brachydont. The unworn Almy premolar 
(U.S.N.M. No. 21283) measures 14.6 mm. long perpendicular to 
anterior margin by 16.5 mm. wide perpendicular to outer wall. P* in 
the type of P. praecursor measures 11.5 by 13.3 mm. in the same 
directions. 

The proportions of the upper premolars are comparable to those of 
the earlier Bathyopsoides harrisorum Patterson (1939) from the 
Plateau Valley beds, although the unworn Almy premolar is a little 
shorter anteroposteriorly and broader transversely than the B. harris- 
orum P*. Nevertheless, the transverse lophs have about the same 
proportions. Although Patterson has indicated certain differences in 
cusp pattern of M2, it would seem from the evidence presented by 
Dorr (1952, p. 89) that Bathyopsoides is possibly a male Probath- 
yopsis. Better evidence with regard to this situation should be forth- 
coming in the more detailed study of the Hoback Basin material 
contemplated by Dorr. 


NO. 7 MAMMALIA FROM THE ALMY FORMATION—GAZIN 17 


REFERENCES 
Core, Epwarp D. 
1873. Fourth notice of extinct Vertebrata from the Bridger and Green 
River Tertiaries. Palaeont. Bull. No. 17, pp. 1-4. 
1882. Contribution to the history of the Vertebrata of the lower Eocene of 
Wyoming and New Mexico, made during 1881. 1. The fauna of the 
Wasatch beds of the basin of the Big Horn River. Proc. Amer. 
Philos. Soc., vol. 20, pp. 139-191, I fig. 
Dorr, JoHN A. 
1952. Early Cenozoic stratigraphy and vertebrate paleontology of the 
Hoback Basin, Wyoming. Bull. Geol. Soc. Amer., vol. 63, pp. 59-94, 
figs. 1-6, pls. I-7. 
Gazin, C. LEwIs. 
1942. Fossil Mammalia from the Almy formation in western Wyoming. 
Journ. Washington Acad. Sci., vol. 32, No. 7, pp. 217-220. 
1952. The lower Eocene Knight formation of western Wyoming and its 
mammalian faunas. Smithsonian Misc. Coll., vol. 117, No. 18, pp. 
1-82, figs. 1-6, pls. I-II. 
1956. Paleocene mammalian faunas of the Bison Basin in south-central 
Wyoming. Smithsonian Misc. Coll., vol. 131, No. 6, pp. 1-57, 
figs. 1-2, pls. 1-16. 
The occurrence of Paleocene mammalian remains in the Fossil Basin 
of southwestern Wyoming. Journ. Paleont. (in press). 
GRANGER, WALTER. 
1915. A revision of the lower Eocene Wasatch and Wind River faunas. 
Part 3—Order Condylarthra. Families Phenacodontidae and Menis- 
cotheriidae. Bull. Amer. Mus. Nat. Hist., vol. 34, art. 10, pp. 320- 
361, figs. 1-18. 
JePsEN, GLENN L. 
1930. Stratigraphy and paleontology of the Paleocene of northeastern Park 
County, Wyoming. Proc. Amer. Philos. Soc., vol. 69, pp. 463-528, 
figs. 1-4, pls. I-10. 
MattTHeEw, WILLIAM D. 
1915a. A revision of the lower Eocene Wasatch and Wind River faunas. 
Part 1—Order Ferae (Carnivora), Suborder Creodonta. Bull. 
Amer. Mus. Nat. Hist., vol. 34, art. 1, pp. 4-103, figs. 1-87. 
1915b. A revision of the lower Eocene Wasatch and Wind River faunas. 
Part 4.—Entelonychia, Primates, Insectivora (part). Bull. Amer. 
Mus. Nat. Hist., vol. 34, art. 14, pp. 429-483, figs. 1-52, pl. 15. 
PATTERSON, BRYAN. 
1939. New Pantodonta and Dinocerata from the upper Paleocene of west- 
ern Colorado. Geol. Ser. Field Mus. Nat. Hist., vol. 6, No. 24, 
pp. 351-384, figs. 100-111. 
ScHuLtz, ALFRED R. 
1914. Geology and geography of a portion of Lincoln County, Wyoming. 
U. S. Geol. Surv., Bull. 543, pp. 1-141, figs. 1-8, pls. 1-11. 
Srmpson, Georce G. 
1928. A new mammalian fauna from the Fort Union of southern Montana. 
Amer. Mus. Nov., No. 297, pp. 1-15, figs. 1-14. 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


1929. A new Paleocene uintathere and molar evolution in the Amblypoda. 
Amer. Mus. Nov., No. 387, pp. 1-9, figs. 1-9. 

1933. Braincasts of Phenacodus, Notostylops, and Rhyphodon. Amer. Mus. 
Nov., No. 622, pp. 1-19, figs. I-3. 

1935. The Tiffany fauna, upper Paleocene. 3.—Primates, Carnivora, Con- 
dylarthra, and Amblypoda. Amer. Mus. Nov., No. 817, pp. 1-28, 
figs. I-14. 

1936. A new fauna from the Fort Union of Montana. Amer. Mus. Nov., 
No. 873, pp. 1-27, figs. 1-16. 

1937a. The Fort Union of the Crazy Mountain field, Montana, and its 
mammalian faunas. U. S. Nat. Mus. Bull. 169, pp. 1-287, figs. 1-80, 
pls. I-10. 

1937b. Notes on the Clark Fork, upper Paleocene fauna. Amer. Mus. Nov., 
No. 954, pp. 1-24, figs. 1-6. 


EXPLANATION OF PLATES 
PLATE I 


PRIMATES, ANACODON ps AND PROBATHYOPSIS? FROM THE ALMY PALEOCENE 


Fig. 1. Anacodon? nexus, new species: M: and M2 (U.S.N.M. No. 21282), 
type specimen, occlusal view. Natural size. 

Figs. 2 and 3. Probathyopsis?, sp.: 2, Upper premolar (U.S.N.M. No. 21283), 
occlusal view; 3, upper premolar (U.S.N.M. No 21284), occlusal view. 
Natural size. 

Fig. 4. Carpolestes, cf. dubius Jepsen: Ps (U.S.N.M. No. 21280), labial (left) 
and lingual views. Six times natural size. 

Figs. 5-8. Plesiadapis cookei Jepsen: 5 and 7, Left ramus of mandible 
(U.S.N.M. No. 20785), (5) occlusal view, twice natural size, and (7) lat- 
eral view, natural size; 6 and 8, left ramus of mandible (U.S.N.M. No. 
16698), (6) occlusal view, twice natural size, and (8) lateral view, natural 
size. 

Fig. 9. Plesiadapis? pearcei, new species: Right ramus of mandible (U.S.N.M. 
No. 20787), type specimen, occlusal and lingual views. Three times natural 
size. 

Fig. 10. Plesiadapis rubeyi Gazin: Right ramus of mandible (U.S.N.M. No. 
16696), type specimen, occlusal and lingual views. Three times natural 
size. 


PLATE 2 
CONDYLARTHS FROM THE ALMY PALEOCENE 


Figs. 1 and 2. Ectocion ralstonensis Granger: 1, Skull (U.S.N.M., No. 20736), 
lateral and ventral views. Natural size; 2, left ramus of mandible 
(U.S.N.M. No. 20736), lateral and occlusal views (M: restored from right 
side). Natural size. 

Figs. 3 and 4. Phenacodus almiensis Gazin: 3, Right upper cheek tooth series 
(U.S.N.M. No. 16691), type specimens, occlusal view (M”* restored from 
left side) ; 4, left ramus of mandible (U.S.N.M. No. 20643), occlusal view. 
Natural size. 

Fig. 5. Phenacodus primaevus Cope: Left ramus of mandible (U.S.N.M. 
No. 21287), occlusal view. Natural size. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131,"NO. 7, PL. 1 





PRIMATES, ANACODON?, AND PROBATHYOPSIS? FROM THE 
ALMY PALEOCENE 


(SEE EXPLANATION AT END OF TEXT.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.131,0NO:.7,, PES.2 





CONDYLARTHS FROM THE ALMY PALEOCENE 
(SEE EXPLANATION AT END OF TEXT.) 
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SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 8 


Charles D. and Mary Waux Galcott 
Research Fund 


THE GEOLOGY AND VERTEBRATE 
PLE ONTOLOGY OF WPPER "EOCENE 
STRATA IN THE NORTHEASTERN 
PART OF THE. WIND RIVER 
BASIN, WYOMING 


PART 2. THE MAMMALIAN FAUNA OF THE 
BADWATER AREA 
(WitH 3 PLaTEs) 


By 
C. LEWIS GAZIN 


Curator, Division of Vertebrate Paleontology 
United States National Museum 
Smithsonian Institution 





(PusiicaTIon 4257) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
OCTOBER 30, 1956 


THE LORD BALTIMORE PRESS, INC. 
BALTIMORE, MD., U. S. A. 





CONTENTS 


Page 
MEIER MIE CAOIEE Vc xsl aha is sc 2 vor awie'e esis) sacs Om aeaitayalsiee steiaisie ale aitiers ¥ieie'¥ bles als el ai I 
NGI OW IEC SITIEMESA i cjeyatey st sjsheucts efe\ciovels oyoteraeroriavsteleinis eve ioisierslellciaverets) > s/arsle: syelsve 2 
ELISEOGY OL MAMVESES ALI OM stovoa cts Sraleio sists le eis tavalelaalcfalcneisiclovele oie cieisievese' elevate clsiste 2 
Occurrence and: preservation Of material . os cjccecescecne csc sicsocs vice 3 
se) Frac Water Palsiai- tic.c.ctac ae.cts hod Sui aiealbi ale ah’e gist eisisie @ie's'e a's Siu/ae/e.0isieisais 4 
Worselation and age Of the: fauna (5.502516 sos tebe sing. dees ceils nee oeisieaee 5 
Systematic description of the Mammalia occ 6isc 60 anes s,ciecdiecsviewsessse 0 
BLS AP MeANA A 8 dans crssnysy Spoke de tal eters opevetatmtarebe ofa ors! ale, <16, 0, diana sscim cieieie.e.daia 9 
PVIGCIPINGAEL a5) soi, oiesisersae so aisle as,0 a)e sieie- 9.2.6 ¥-¥'6ia. 6 66ers sareeiae 9 
APOE | osc aissc Bale weir ee OLR ewe we alelec der wiatkl ew ese.ave eo sisidieversierety 9 
PROROTIAE.. (3.2 ctisscsrsiaim esc ae ersieeraais ous cbieisia's estein eye casa eens 9 
CMSINE TAs Sc 03 ateua s & aj tia cehec tea ici tater here eie eieiaiait Oe whale ara whee ies ona ais weenie 10 
ROPERNEV ON Pyaar ato ta' cts reha)watcasin at ue Syst faa, Dies valnl a stalere St elarasa a Goa abs) RO eoharsaxora'e 10 
PAMINOCVOMMGAC a. ,sa ee ec cee ae Cots renin eae ey SONa Oa chap wers 10 
MACTDAC  siacs cikgtate ee a eiaine Macatee tle Oe eb WS ois Oks Rae STAG ales 10 
MG Gotatiar cat ana 5. Siac < oie ao erevane atencreh rete Meare ee wee bial Sie'e od eahain ged bNlars II 
EA VOMSOUMGHUIGAC sic je-chavei ad Sra iencioha erence evailoreiaw ciate ola os e-cinnies aie acs II 
Pea ESSOUACES La nee) craiah seca ta bracaiil alavers re oie e eiata) Ba latadioiaiere gral antrene s-@:ersinl¥ javarovace 12 
EET ae yet tela reel atetolatelord che chee toucloue elas areiene esa eveuerevelwiatwie mis Sisiae eke 12 
EPEOMILOLACTIMLAG’, a.6:3)orstcin vie raters. oa 'ale eis vere a\e.a-b/ela piaibie(aldle wieleie/stereieiae 13 
RTC EMP ENGARM, lrg i.' Sera.) tts Sevduare, dbrelerd ea ejhiale, daoeioain'e.o-ofsloune’s 13 
VACUA Gee oto hus ceNaiere aaah e aisraleiwial o sina ais,auesateuayalesstPiasa a, Sie ersislale tains 16 
EISWACOMONITICIAE vole tities wine tecveie tare vere ale aye GING, waracarOle Sade ees SSO 23 
RE MMAAL EM s eteeheac wet ae static a Zea cis nik Ce Casale slone aaa ow aN Gia veces 23 
DT CHODUMIG ACN iia Ne cree her iste SIS HE a ee ane erate fed 23 
PN UC IOCRACLAL OF sts aieisto ie ashen teterals Sait oeiciaratoiola Soave wid oe e Miers aia aio 26 
ORCOMEENIAAE WE. 12) «ors oroalassarapalataters sree chelate wu ernie ala Ricfaltrave ate ales atte 30 
BENE ORIE TY CIM AC!seskue poeoetele AoyevasSecie ah oti er Ro nto ees phecoee ale eiesios Chloe 30 
aR RIES POR Mars icici se oon, aspirant data, i srelie 6 Mydaneo gid'ns wv ayaudin ie ack meomaceeha nl eehzie 31 
RMU POLAEMEMGTA LAE) SPARES 6 oyu chee cle ato nl eik sig crater Seore nie a sccit die oheiy a wi aitmid wradin's 34 

ILLUSTRATIONS 

PLATES 


(All plates follow page 35.) 


~ 


. Lagomorph, carnivore, condylarth, and perissodactyls from the Badwater 
upper Eocene. 

2. Perissodactyls from the Dry Creek and Badwater upper Eocene. 

3. Artiodactyls from the Badwater upper Eocene. 


CHART 


_ 


. Suggested phylogenetic arrangement of North American tapiroids..... 16 


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slats 


- | : 
rT ; 


Charles D. and Mary Waux Walcott Research Fund 


THE «GEOLOGY ANDm«VERTEBRATE,: PA- 
PEONTOEOGY’T OR UEPER (LOCENE 
STRATA IN THE NORTHEASTERN 
PART OF THE WIND RIVER 
BASIN, WYOMING 


PART 2.1 THE MAMMALIAN FAUNA OF 
THE BADWATER AREA 


By C. LEWIS GAZIN 


Curator, Division of Vertebrate Paleontology 
United States National Museum 
Smithsonian Institution 


(WITH 3 PLATEs) 


INTRODUCTION 


The significance of the Wind River Basin in contributing informa- 
tion on mammalian faunas of upper Eocene time has been appreciated 
only during comparatively recent years. Although a rather meager 
fauna had been known from beds of Uintan equivalence below the 
Beaver Divide along the south side of the basin for many years, it is 
rather surprising that the occurrences on the north side were not 
earlier discovered, particularly in view of the long history of collecting 
associated with the adjacent lower Eocene Wind River formation. 
Discovery of the occurrence of upper Eocene mammalian remains 
along Badwater Creek near the site of the old Badwater Post Office 
by Wood, Seton, and Hares in 1936 was followed by investigations of 
others, notably those of Harry A. Tourtelot for the U. S. Geological 
Survey and parties for the Smithsonian Institution. 

The present study stems largely from an interest in Eocene tapi- 
roids, the upper Eocene representatives of which are so well repre- 
sented here, and is in part a sequel to an earlier review of artiodactyls 


1 Part 1 of this paper is a study of the geologic relations, in preparation by 
Harry A. Tourtelot. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 131, NO. 8 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


of this age. Misunderstandings in the past as to the characteristics and, 
in some instances, the age of related types, largely from inadequate 
original descriptions of forms in these two ungulate groups, eariy 
confused the picture and resulted in incorrect identifications and evi- 
dently misleading conclusions as to the horizon represented by the 
Badwater assemblage. It is hoped that the present review and revision 
will clarify the record and render more useful the information to be 
derived from this occurrence. 


ACKNOWLEDGMENTS 


I am particularly indebted to Dr. G. Edward Lewis of the U. S. 
Geological Survey for relinquishing to me for restudy this very inter- 
esting assemblage, and assuring me that no embarrassment would 
ensue. I wish also to acknowledge the helpful information furnished 
me by Harry A. Tourtelot both in the office and in the field. Tourtelot, 
obligingly and with contagious enthusiasm, showed me the more sig- 
nificant and likely collecting sites during our 1946 field exploration. 

Investigation was immeasurably aided by my being permitted to 
examine and study related materials in universities and other museums, 
and by the loan of pertinent specimens in these collections for direct 
comparison. Acknowledgment is particularly due Dr. J. LeRoy Kay at 
the Carnegie Museum, Dr. Glenn L. Jepsen at Princeton University, 
Dr. George G. Simpson at the American Museum, and Dr. Joseph T. 
Gregory at Yale Peabody Museum. Dr. H. E. Wood, II, aided in 
furnishing information on materials in the original Badwater col- 
lection. 

The exquisite pencil drawings depicting selected materials shown in 
plates I-3 were prepared by Lawrence B. Isham, staff artist for the 
Department of Geology in the U. S. National Museum. Mr. Isham 
also prepared the chart showing the tapiroid sequence. 


HISTORY OF INVESTIGATION 


Discovery of upper Eocene vertebrate materials in the Badwater 
area was made by Wood, Seton, and Hares, as reported by them in 
1936. Moreover, this would appear to be the first record for the oc- 
currence of upper Eocene on the north side of the Wind River Basin. 
Recognized by Wood, Seton, and Hares were Amynodon advenus, 
Telmatherium, cf. cultridens, and a crocodile. Collections later (1942) 
secured by J. D. Love and G. E. Lewis from Lysite Mountain to the 
north of Badwater Creek, for Yale University, include remains iden- 
tified by Lewis as Telmatherium, cf. cultridens, and an indeterminate 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 3 


helaletid (this is Dilophodon). Nevertheless, significant collections, 
more representative of the fauna, were not obtained from these beds 
until Harry A. Tourtelot and his assistants secured for the U. S. 
Geological Survey in 1944 and 1945 the materials discussed in his 
maps and reports of 1946, 1948, and 1953. Identification of the 
Geological Survey material was made by G. E. Lewis and reported by 
him in 1947. Collections for the Smithsonian Institution were made 
by F. L. Pearce, Chester Gazin, and myself in 1946, and Pearce and 
I revisited the localities with good results in 1953. Other known col- 
lections include that made by A. E. Wood in 1948 for Amherst Col- 
lege, the small mammals represented having been described by him in 
1949. Further collecting was done by Tourtelot and the unusual 
Malaquiferus tourteloti skull was found by him near Dry Creek in 
1948. Materials secured by the U. S. Geological Survey also included 
a collection made by J. R. Hough in 1950, and in her 1955 report on 
the Sage Creek occurrence comparisons are made with portions of the 
Badwater fauna. 


OCCURRENCE AND PRESERVATION OF MATERIAL 


The principal occurrences for materials of the Badwater fauna are 
the low gray-green exposures along the south side of Badwater Creek 
between 24 and 34 miles almost due northwest of the site of the now 
abandoned Badwater Post Office. These are immediately to the south 
and to the southeast of the mouth of Clear Creek in the southeast part 
of section 14, the southwest part of section 13, and the northwest 
part of section 24, T. 39 N., R. 89 W. The above, together with 
other scattered localities, are shown on both the 1946 and 1953 maps 
of Tourtelot, as well as his map accompanying part 1 of this study. 

The discovery of Badwater vertebrate remains by Wood, Seton, and 
Hares was made at a locality south of Badwater P.O. about 3 miles 
to the southeast of the above exposures and in section 32, T. 39 N., 
R. 88 W. Determinable remains have likewise been encountered on 
Lysite Mountain to the north of the Badwater area by Lewis and 
Love, probably in section 25, T. 42 N., R. 90 W. The Dry Creek 
exposures, almost certainly the same age as those on Badwater Creek, 
are about 20 miles due west and include the sites for the Malaquiferus 
and Eomoropus skulls, in the NW4 sec. 14 and the SE sec. 9, re- 
spectively, T. 39 N., R. 92 W. 

Much of the fossil material encountered has been rather fragmen- 
tary although there are five comparatively good skulls in the National 
Museum-Geological Survey collections from there. Two of these are 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I3I 


of Diplobunops and the others are of Protoreodon, Malaquiferus, and 
Eomoropus. It is particularly noteworthy that although often much 
fractured, the remains show almost no distortion, a condition unusual 
in collections of Eocene age, particularly those from the Uinta basin. 
The bone for the most part is a light buff color and the teeth amber to 
dark brown. The beds themselves do not resemble exposures of the 
Uinta formation in Utah but much more closely resemble the light 
gray-green middle Eocene reworked volcanic ash of the Bridger for- 
mation in southwest Wyoming. The upper Eocene fossil-bearing beds 
in the Badwater area have been named the Hendry Ranch member 
by Mr. Tourtelot and regarded as a part of the Tepee Trail forma- 
tion. Definition and description of these beds are included in part 1 
of this paper. 


THE BADWATER FAUNA 


MARSUPIALIA?: 
Didelphidae ? : 
Peratherium?, sp. 
LAGOMORPHA : 
Leporidae : 
Mytonolagus wyomingensis A. E, Wood 
RODENTIA * : 
Paramyidae : 
Rapamys?, sp. 
Sciuravus dubius A. E. Wood 
Paramyid indet. (large) 
Paramyid indet. (small) 
Eomyidae: 
Protadjidaumo?, sp. 
Cricetidae : 
Cricetid indet. 
CARNIVORA : 
Limnocyonidae : 
Limnocyon?, sp. 
Miacidae: 
Miacis, cf. robustus (Peterson) 
CONDYLARTHA: 
Hyopsodontidae : 
Hyopsodus, cf. uintensis Osborn 
PERISSODACTYLA : 
Equidae: 
Epihippus, cf. gracilis (Marsh) 
Epthippus, cf. parvus Granger 
Brontotheriidae : 
Brontotheriid indet. 
Chalicotheriidae : 
Eomoropus anarsius, new species 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 5 


Helaletidae: 
Desmatotheriwm woodi, new species 
Dilophodon, cf. leotanus (Peterson) 
Hyracodontidae : 
Epitriplopus?, sp. 
Amynodontidae* : 
Amynodon advenus (Marsh) 
ARTIODACTYLA : 
Dichobunidae : 
Pentacemylus?, sp. 
Apriculus praeteritus, new genus and species 
Agriochoeridae: 
Protoreodon, cf. petersoni Gazin 
Protoreodon, near P. pumilus (Marsh) 
Protoreodon pearcei, new species 
Diplobunops, cf. matthewi Peterson 
Oromerycidae: 
Malaquiferus tourteloti Gazin 
Leptomerycidae: 
Leptotragulus, cf. medius Peterson 
Leptoreodon?, sp. 


* Rodents are as described by A. E. Wood from material in the collections at Amherst 
College and not represented in collections at the U. S. National Museum. The Amynodon 
material is that identified by H. E. Wood, II, in the collection of Wood, Seton and Hares. 


CORRELATION AND AGE OF THE FAUNA 


The fauna listed above is, of course, by present standards upper 
Eocene in age. Moreover, there would seem to be no doubt but that 
it is Uintan. Apparently not any of the forms here recognized are 
characteristically or exclusively Duchesnean. The general association 
of forms seen in the assemblage and the development reached in 
certain groups such as the agriochoerids rather strongly suggest an 
upper Uintan stage close to that at Myton pocket. 

Considering first the reasons for not regarding the fauna as 
Duchesnean, only the rodent which A. E. Wood (1949) cited as 
questionably Protadjidaumo might be interpreted as this age. Never- 
theless, this form is also older than Lapoint in age, as Kay (1953, 
p. 24) cites it as occurring in the Randlett fauna, and the latter I re- 
gard as but scarcely distinct from that of Myton, including it ? (Gazin, 
1955, chart 1) in the Uintan. The remainder of the fauna is composed 
of genera that so far as Duchesnean is concerned are characteristically 
earlier or common to both Uintan and Duchesnean. Thus, besides 


2 As currently being proposed by the Committee on Nomenclature and Cor- 
relation of North American Continental Tertiary of the Society of Vertebrate 
Paleontology. 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Protadjidaumo, only Epihippus, Amynodon, and Protoreodon are 
common to the two, and the genera Mytonolagus, Rapamys, Eomoro- 
pus, Epitriplopus, Pentacemylus, Diplobunops, Leptotragulus, Lep- 
toreodon, and possibly Desmatotheriwm and Dilophodon are charac- 
teristic of the Uintan. Some of these almost certainly gave rise to later 
types in the Oligocene but the genera in the latter group are not ac- 
tually known in Duchesnean time. The genera Sciuravus, Limnocyon, 
and Hyopsodus are survivals from Bridgerian time and Miacis ranges 
through most of the Eocene. 

The evidence for regarding the fauna as upper rather than lower 
Uintan pertains to the presence of Mytonolagus, the possible Pro- 
tadjidaumo, and particularly to the stage of development shown in the 
Protoreodon and Diplobunops material. On the other hand, the pres- 
ence of Sciuravus (doubtfully this genus according to Wood), Lim- 
nocyon, and Eomoropus might suggest an earlier horizon, but these 
are comparatively rare forms in Uintan deposits and their absence 
heretofore in beds as, late as Uinta C is not nearly so significant as 
the fact that the agriochoerids are distinctly advanced over those of 
Uinta B time. 

Question as to whether the Badwater fauna should be correlated 
with that from Myton pocket or with that known from the Randlett 
member may well have little significance. Protadjidaumo is not known 
from Myton pocket but the Badwater specimens are stated by A. E. 
Wood to consist of incisors only, so can scarcely merit serious debate. 
Mytonolagus is known from both levels but the Badwater form is a 
different species. Dilophodon (“Heteraletes”’) might suggest a rela- 
tionship to the Randlett, but Uinta collections in the U. S. National 
Museum show that this form is present also in the Myton fauna. 
A slight evidence favoring the Myton fauna is seen in the artiodactyl 
species represented. Of the Badwater forms, Protoreodon pumilus 
is evidently present in all three occurrences, but P. petersoni and 
probably P. pearcei are known only in the Myton fauna. Also, the 
Diplobunops from Badwater resembles the Uinta form D. matthewi 
more closely than it does the broad-skulled D. crassus. It is entirely 
possible that, although a difference in stratigraphic level has been 
described for the Myton pocket and Randlett occurrences, the differ- 
ences that may be pointed out are of ecologic significance, as sug- 
gested by the rather different nature of the deposits. The beds at 
the Myton pocket and Randlett occurrences received sedimentary 
materials from quite different rock sources. I have been unable to 
detect any change which can be regarded as evolutionary between 
forms common to the two levels. 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN a 


Lack of uniformity of opinion regarding the source of Douglass’s 
Eocene materials from the Sage Creek areas makes comparison with 
the fauna or faunas represented there decidedly unsatisfactory. I have 
not had the opportunity of studying the field occurrence firsthand so 
am unable to contribute any information to the stratigraphic picture. 
Nevertheless, from the materials that I have examined in the collec- 
tions of Kay and of Hough, understood to be from a single horizon 
in the Eocene sequence, I find a comparatively close relationship be- 
tween their fauna and the Badwater assemblage. While I do not con- 
cur in several of the identifications cited in Hough’s (1955) paper, 
nor do I agree with the Duchesnean age assignment, there would ap- 
pear to be a near equivalence in time, possibly also in environment, 
considering the similarity in faunal representation. With regard to 
the Douglass collection, I have seen only the helaletid and am reason- 
ably convinced that it represents an advanced dilophodont distinct 
from the Dilophodon in Kay’s collection. If, as Horace E. Wood 
(1934, p. 255) postulates, Douglass’s amynodont might have weath- 
ered from the overlying Cook Ranch Oligocene, it is not impossible 
that the dilophodont did likewise and is a distinctly small and perhaps 
unprogressive species of Protapirus. In any case, its stage of develop- 
ment in the line of true tapirs postulated elsewhere in this paper would 
appear to be later than Uinta B. Recognition of the amynodont re- 
mains as Amynodon advenus by Wood in both the Douglass and Bad- 
water collections would suggest a near equivalence in time. As to 
Hyrachyus douglassi, it would not appear to be as late as upper Uintan. 
H. douglassi and Chasmotheroides, cf. intermedius may well be Uinta 
B, or even earlier. 

There remains consideration of the faunas from the Swift Current 
Creek beds of Saskatchewan and the Tapo Ranch horizon of the Sespe 
in California. Although the collections known from the Swift Current 
Creek beds consist of decidedly fragmentary materials there is sug- 
gestion of an age which might not be far removed from that at Bad- 
water. Contributing to this is the association of lagomorph and 
Hyopsodus seen in both assemblages. 

Of the horizons represented in the Sespe sequence, the Badwater 
would appear to be nearest to that represented at Tapo Ranch or 
C.I.T. locality 180. Although the species and most of the genera are 
not the same, the ages are probably not too different. The distinctive 
nature of the Tapo Ranch fauna may be largely due to its geographic 
remoteness, 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 132 


SYSTEMATIC DESCRIPTION OF THE MAMMALIA 


MARSUPIALIA? 
DIDELPHIDAE? 
PERATHERIUM?, sp. 


The isolated lower molar, Amherst No. 10019, which A. E. Wood 
(1949) regarded as questionably representing Metacodon does not 
seem to me to be insectivore. His careful drawing of this tooth sug- 
gests possibly a closer relationship to the marsupials. I am particularly 
impressed by the lingual position of the hypoconulid, and the posterior 
deflection of the crest on which this cusp is located, away from the 
entoconid. The talonid appears to be quite different from the structure 
seen in Jctops and is unlike that, for example, in M, of Metacodon 
mellingeri where the hypoconulid is closely connected to the entoconid. 
For these reasons I have tentatively assigned this specimen to Pera- 
therium?, sp. 


LAGOMORPHA 
LEPORIDAE 
MYTONOLAGUS WYOMINGENSIS Wood, 1949 
Plate 1, figure 1 


A. E. Wood (1949) has described several isolated teeth of 
Mytonolagus from the Badwater area and a comparatively unworn 
P® was designated the type of Mytonolagus wyomingensis. A right 
maxilla (U.S.N.M. No. 21090) with P*-M? collected by F. L. Pearce 
undoubtedly represents the same species. P* in this specimen, however, 
is more worn than in the type, although the teeth in general appear to 
be less worn than in the type of Mytonolagus petersoni which Burke 
(1934) described from Uinta C at Myton pocket. The teeth are strik- 
ingly like those in the type of M. petersoni, but it is noted that the 
hypostriae on M, and particularly M, are more persistent, extending 
nearly to the upper limit of the enamel lingually. The comparative 
weakness of the lingual fold toward the root of M* in M. peterson 
was further noted in upper-tooth material of the Myton form in the 
collections of the National Museum. At least the hypostria extends 
nowhere near the upper limit of the enamel. Wood has regarded M. 
qwyomingensis as perhaps more primitive than M. petersont. 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 9 


RODENTIA 


Description of the known Rodentia in the Badwater fauna has been 
covered by A. E. Wood (1949). The collection described by him is 
at Amherst College. 


CARNIVORA 
LIMNOCYONIDAE 
LIMNOCYON?, sp. 


A maxillary fragment (U.S.N.M. No. 21088) with only P* may 
represent Limnocyon, but this is not certain. The specimen shows 
the infraorbital foramen immediately above and anterointernal to the 
anteroexternal root of P*, much as observed in Limnocyon. The tooth 
would appear to be a trifle smaller than in Limnocyon douglassi to 
judge by Peterson’s (1919) illustration of this form. The Badwater 
tooth measures 9.6 mm. long by 9.4 transversely to base of enamel 
on the deuterocone. 


MIACIDAE 
MIACIS, cf. ROBUSTUS (Peterson), 1919 
Plate 1, figure 2 


A comparatively large miacid is represented by a lower jaw exhibit- 
ing the teeth P, to Mz inclusive. Miacis would appear to be indicated 
by the distinctly basined form of the relatively small talonid of M.. 
The talonid of M, may likewise have been basined, although most 
of the superior surface of this portion of the carnasial is damaged 
so that its precise form is uncertain. It is, nevertheless, as in M., 
short and decidedly narrower than the trigonid. Mz; is missing, repre- 
sented by a single alveolus. 

From measurements given by Peterson (1919), the type of Miacis 
robustus from the Uinta at Myton pocket is a little larger than the 
Badwater specimen. Peterson regarded P, and M, as subequal in size 
so that P, is evidently both relatively and actually larger in the type. 
This tooth, however, in U.S.N.M. No. 21087 closely resembles that of 
the type in the presence of a prominent anterior cusp and a strong 
talonid cusp, followed posteriorly by a well-developed cingulum. The 
abbreviation of the talonid in both M, and M, likewise suggests M. 
robustus. 

The type of Miacis uintensis Osborn (1895) from Uinta B would 
appear from the scale of Osborn’s illustration to be a little shorter 


IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


in length of cheek teeth and distinctly shallower jawed than the Bad- 
water form. Moreover, P, in the type of Osborn’s species apparently 
lacks the anterior cusp but has one more cusp on the posterior crest, 
to judge by the illustration (fig. 2), resembling more closely the re- 
ferred tooth, A.M. No. 1895. M; is relatively smaller, and M, in 
Osborn’s type has a higher trigonid and a less distinctly basined 
talonid. 

Miacis gracilis Clark (1939) is, of course, a decidedly smaller 
species and P, is evidently characterized by two posterior cusps in ad- 
dition to the cingulum. 


MEASUREMENTS IN MILLIMETERS OF LOWER TEETH 
IN Miacis, cr. robustus, U.S.N.M. NO. 21087 


P,, anteroposterior diameter: transverse diameter...............0000. 10.5: 5.4 

Mu,, anteroposterior diameter: transverse diameter of trigonid......... T3.05°7:5 

M2, anteroposterior diameter: transverse diameter of trigonid......... 5.6: 4.3 
CONDYLARTHRA 
HYOPSODONTIDAE 


HYOPSODUS, cf. UINTENSIS Osborn, 1902 
Plate 1, figure 3 


A single Hyopsodus upper molar, U.S.N.M. No. 21089, may well 
represent H. uintensis, although some doubt may be entertained as 
comparisons involving such limited material cannot be entirely satis- 
factory. The tooth is about intermediate in size between M* and M? 
in the type, A.M. No. 2079, but resembles M? more closely than M?. 
The Badwater molar measures 4.3 mm. long by 5.6 transversely. This 
is too small to occlude properly with the type lower molar of Hyopso- 
dus fastigatus Russell and Wickenden (1933) from the Canadian 
Swift Current Creek beds. 

It is interesting to note that although the type, and presumably the 
two referred lower jaws mentioned by Osborn, are from the Uinta 
C of Utah, there is in the collections of the U. S. National Museum 
a lower jaw from Uinta B at White River pocket. 

Mention may also be made of an upper Eocene occurrence of 
Hyopsodus at the Beaver Divide. The specimen, an upper molar, 
comparable in size to the Badwater tooth, was collected by Van Houten 
in beds he early regarded as representing the Beaver Divide conglome- 
rate. The locality in question is some distance away from the critical 
Wagonbed Springs section and Van Houten has since doubted * the 


8 Personal communication. 


No. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN Tr 


correlation so that the tooth may well have originated in the Uinta 
equivalent present in the sequence. 


PERISSODACTYLA 
EQUIDAE 
EPIHIPPUS, cf. GRACILIS (Marsh), 1871 
Plate 1, figure 5 


The rather scant material representing Epihippus was first en- 
countered in the Badwater localities in 1953. A maxillary fragment, 
U.S.N.M. No. 21092, including P?, P*, and part of P* and a single 
lower molariform tooth, U.S.N.M. No. 21094, possibly Mz, represent 
an equid approximately the size of Epihippus gracilis. 

P? in No. 21092 is advanced over Orohippus in the development of 
the lingual portion, but not nearly so molariform as in Mesohippus. 
The anterointernal cusp in this tooth appears weaker than in the type 
of Epihippus parvus as figured by Granger (1908), being scarcely 
more than a low crest, extending lingually from the lingual surface 
of the paracone rather than from a position anterior to the paracone. 
There is no evidence of a mesostyle on P?. P* would appear to be en- 
tirely molariform. The second premolar measures 6.7 mm. long by 
5.8 transversely. 

The lower molar, in comparison with Uintan horses, shows little 
of diagnostic importance other than size which is close to that of the 
preserved molar (M,) in the type of Epihippus uintensis (Marsh), 
placed by both Marsh and Granger in synonymy with E. gracilis. The 
tooth is a little smaller, though scarcely if any more brachydont than 
Epihippus (Duchesnehippus) intermedius. The V-shaped crests of 
the lower molar, however, are a little less acute than in the Duchesnean 
horse. The metaconid and metastylid are separate at the apex but this 
has been noted in molars as well as premolars of both the Uintan and 
Duchesnean Epihippus. The tooth measures 9.0 mm. long by 6.3 wide. 


EPIHIPPUS, cf. PARVUS Granger, 1908 
Plate 1, figure 6 


The material of a smaller horse in the Badwater fauna likewise in- 
cludes a maxillary portion with P? and P*, U.S.N.M. No. 21091, and 
an isolated lower molariform tooth, U.S.N.M. No. 21093. There is, 
in addition, the greater part of an isolated molariform upper cheek 
tooth. 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


P? in the specimen compared with £. parvus is distinctly more pro- 
gressive looking than in No. 21092 helieved close to E. gracilis. The 
anterointernal cusp is clearly defined on a crest extending postero- 
lingually from a position anterior to the lingual surface of the para- 
cone, somewhat as it appears in the type of E. parvus, but with the 
long diameter of the cusps directed a little more transversely than in 
the latter, so that the anterior portion of the tooth seems broader. 
Moreover, the outer wall shows evidence of an incipient mesostyle, 
better developed in the type, but no trace of which was found in P? 
of No. 21092. P? in No. 21091 measures 6.1 mm. long by 5.1 wide 
transversely ; P* is 6.7 by 6.9. 

The lower molariform tooth is quite like that (No. 21094) com- 
pared to E. gracilis but distinctly smaller. It measures 7.8 mm. long 
by 6.3 wide, comparing favorably in length, but a little broader than 
molariform lower teeth in A.M. No. 2066 referred to E. parvus. 


BRONTOTHERIIDAE 


A fragmentary maxillary portion without teeth but showing root 
portions of the canine and first two premolars would appear to be of a 
titanothere. Speculation as to the genus represented would be un- 
warranted. Enamel fragments of large teeth in the collection may also 
be titanothere, but this is uncertain as they might equally well repre- 
sent an amynodont rhinoceros. 

The Badwater specimen cited by Wood, Seton, and Hares (1936) 
as Telmatherium, cf. cultridens is half of a lower molar which W. K. 
Gregory, in a note to Wood, observed, “Granger and I found this tooth 
to be close to M, of referred specimens of Telmatherium cultridens.” 
There is, of course, a close resemblance; nevertheless, from the very 
fragmentary nature of the specimens it is extremely doubtful if among 
the several genera of Uintan titanotheres all can be excluded from 
consideration. The specimen from Lysite Mountain identified by 
G. E. Lewis (in Tourtelot, 1948) as Telmatherium, cf. cultridens I 
have not seen. It could not be located in the collections at Yale Pea- 
body Museum. 


CHALICOTHERIIDAE 
EOMOROPUS ANARSIUS,* new species 
‘ Plate 2, figures 1-3 


Type.—Greater part of left side of skull and left ramus of mandible, 
U.S.N.M. No. 25097. 


* Anarsios (Gr.), incongruous, strange—in allusion to the large and unex- 
pected canine. 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 3 

Horizon and locality—Hendiy Ranch member of Tepee Trail for- 
mation on Dry Creek, SE4 sec. 9, near line between secs. g and 16, 
T. 39 N., R. 92 W., Wind River Basin, Wyo. 

Specific characters —Teeth only slightly larger than in Eomoropus 
amarorum, but skull proportions and depth of lower jaw much greater. 
Parastyle of upper molars increasingly prominent from M!? to M%, 
considerably more extended anteroexternally than in Eomoropus an- 
nectens, and evidently more so than in E. amarorum. 

Discussion.—One of the more important discoveries in the upper 
Eocene of the Wind River Basin is the skull and jaw material of the 
chalicothere, Eomoropus. The specimen (U.S.N.M. No. 21097) con- 
sists of the left half of the skull and left ramus of mandible, and was 
found by F. L. Pearce in exposures on an eastern tributary of the 
east fork of Dry Creek about 20 miles west of the Badwater Creek 
localities. The deposits here were mapped by Tourtelot ° as the same 
formation as that exposed along the south side of Badwater Creek 
and are believed to be the same age. 

The species represented was earlier (Gazin, 1955, p. 77) thought 
to be Eomoropus amarorum, but subsequent direct comparison with 
the type, A.M. No. 5096, would seem to preclude this possibility. 
E. amarorum was described by Cope (1881) from a specimen consist- 
ing of the posterior portion of a skull, a lower jaw, and certain other 
portions of the skeleton illustrated by Osborn (1913), and derived 
from the Washakie Basin. According to Osborn, Cope’s specimen 
probably came from near the base of Washakie B, or the upper 
Washakie. I suspect that the horizon represented is from higher in 
the Washakie than suggested, inasmuch as E. amarorum would appear 
to be more progressive than Uinta B Eomoropus annectens. 

Comparison of Eomoropus anarsius with the type of E. amarorum 
shows similarities in the orbital region but the depth of the face below 
the lower margin of the orbit is conspicuously greater, also the post- 
orbital process of the frontal appears less prominent and overhanging. 
The lateral view of the squamosal is similar in the two with the rela- 
tive position of the external auditory meatus with respect to the 
glenoid surface much the same. However, the distance between the 
last molar and the glenoid surface is about 30 percent greater in 
E. anarsius. Moreover, the depth of the lower jaw is also nearly 30 
percent greater and the masseteric fossa n. e deeply impressed and 
better defined. The two animals would appear to be at about the same 
stage of maturity, with E. amarorum possibly a little older, to judge 


°U.S.G.S. Oil and Gas Investigations Map OM 124, sheet 1. 


14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


by wear of M, (compare height of cusps shown in lateral view, pl. 2, 
fig. I, with Osborn’s fig. 3A on p. 267, 1913). 

Unlike the type of E. amarorum, the cranial portion of the £. 
anarsius skull is poorly preserved, but fortunately the side of the 
rostrum, missing in the former, is present in the Dry Creek specimen. 
Perhaps the most striking feature to be revealed by the E. anarsius 
rostrum is the enlarged canine, evidently not included in the dentition 
of Moropus. Absence of this tooth has been generally regarded as 
characterizing the family although in certain forms the complete 
formula is not known. The upper premolars were not preserved in 


MEASUREMENTS IN MILLIMETERS OF DENTITION IN TYPE OF 
Eomoropus anarsius, U.S.N.M, NO. 21097 


Length of upper molar series, M*-M* inclusive, parallel to tooth row...... 54.4 
M,, anteroposterior diameter perpendicular to anterior margin........... 14.9 
M,, greatest transverse diameter across parastyle and protocone.......... 10.7 
M2, anteroposterior diameter perpendicular to anterior margin............ 19.0 
M2, greatest transverse diameter across pafastyle and protocone.......... 23'5" 
Ms, anteroposterior diameter perpendicular to anterior margin........... 19.6 
Mz, greatest transverse diameter across parastyle and protocone......... 25.0 
Length of preserved lower cheek tooth series, P:-Ms, inclusive.......... 78.0 
Bencth, Of fOwer PECMIOIALS bac bre. sia mie 3 o:k’eimipisinrnjas A einbs einai otwin thin are inion 24.0 
Reneti-or lower molars: Ma-Ms i. c.cicet ss tame cea cena ne vows tauren ney ee 54.0 
P;, anteroposterior diameter : transverse diameter of talonid.......... 12.8:7.8 
P,, anteroposterior diameter : transverse diameter of talonid.......... 11.2: 8.1 
M,, anteroposterior diameter: transverse diameter of talonid......... 13.9: 9.0 
M2, anteroposterior diameter: transverse diameter of trigonid........ 16.9: 10.5 
Ms, anteroposterior diameter : transverse diameter of talonid......... 24.0: 10.5 


@ Approximate. 


No. 21097, but the three molars are complete. Eomoropus upper 
molars are characterized by a lophoid protocone and hypocone, more 
elongate than in Moropus, and with a distinct protoconule. The ex- 
ternal wall exhibits a strikingly developed parastyle and a prominent, 
anteroposteriorly compressed mesostyle and paracone rib. The rib 
on the metacone is weak or wanting. M* exhibits a spurlike metacone 
directed nearly at right angles to the similarly developed mesostyle. 
Comparison with upper teeth in Cope’s type is limited to the lingual 
portion of M® and no significant differences are observed. Neverthe- 
less, the anteroexternal root for M? and M® can be observed in Cope’s 
specimen, and its position is not nearly so forward and outward as in 
E. anarsius, suggesting rather less extension of the parastyle in the 
molars. Comparison with the excellent upper cheek tooth series pre- 
served in the type of Eomoropus annectens, which Peterson (1919) 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 15 


described from the Uinta B horizon, shows that the Dry Creek speci- 
men has molars rather similar, except that the parastyles are strikingly 
more outstanding and the teeth are about 20 percent larger. Moreover, 
the rib on the paracone, in keeping with the parastyle, is better de- 
veloped. 

The lower teeth of FE. anarsius are quite similar to those in F. 
amarorum and about the same size, although the anterior premolars 
may be relatively a little larger. E. amarorwm includes all the lower 
cheek teeth from P, to Ms, inclusive. In No. 21097 only Pz. of this 
series is missing, although the inner walls of M. and M; are not com- 
plete. As Osborn has shown, these teeth are much like those in 
Moropus; however, in the earlier form Pz» is a relatively larger tooth 
and M; retains a prominent hypoconulid. Moreover, as observed in 
occlusal view of the Dry Creek specimen, the crista obliqua in all the 
cheek teeth following P, joins the posterior wall of the trigonid some- 
what more buccally and lower than in Moropus, so that the W-pattern 
is not so well developed. 


HEELALPETIDAE 


Simpson (1945) included the Helaletidae in the Tapiroidea, an ar- 
rangement which is distinctly preferable to including it, as Scott 
(1941) has, in the Rhinocerotoidea. Scott, moreover, included the 
hyrachyids in the Helaletidae, and although there is much to be said 
for such a grouping, as the hyrachyids are in many ways intermediate 
between the tapiroids and rhinocerotids and show certain marked 
resemblances to Helaletes, nevertheless the family distinction as the 
Hyrachyidae within the Rhinocerotoidea, as advocated by H. E. Wood, 
II (1934) and retained by Simpson (1945), may well be more de- 
sirable. 

The Eocene tapiroids are structurally a comparatively conservative 
group, distinguished from one another by relatively small and seem- 
ingly unimportant differences. Nevertheless, at least two families, the 
Helaletidae and Isectolophidae, should probably be recognized for 
North American forms. 


DESMATOTHERIUM Scott, 1883 


Originally described by Scott as coming from the Bridger Eocene, 
it is now understood that the type of Desmatotherium guyoti came 
from the Washakie beds (see Granger, 1909, p. 22) and is in all likeli- 
hood upper Eocene rather than Bridgerian in age. Peterson (1919, 
p. 127) was evidently in error in citing the locality for this specimen 
as ‘‘Henry’s Fork, Wyoming.” 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


TAPIRIDAE 
OLIGOCENE Protapirus Colodon 


oe 
ah 
Randlett 


DUCHESNEAN 













| \oonaten pe 
’ il 4 
A Helaletes se af Parisectolophus 
ane 
Sty 
Bridger C Heloletes © !  Parisectolophus 
1 i 
fu 
pel Kd ( Parisectolophus 
Soo eee 
\ 
Lysitean H 


crati Homogalax 


EO GE NE 


BRIDGERIAN 








Tel 


%Exoct horizon uncertain 


Fic. 1.—Suggested phylogenetic arrangement of North American Eocene 
tapiroids. 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 17 


Scott’s generic comparisons were made largely with Hyrachyus, 
from which it clearly differs. Iam convinced, however, that a distinctly 
closer relationship to Helaletes is indicated. The Badwater, Sage 
Creek, and Washakie materials together provide the evidence showing 
the sequence from Bridgerian Helaletes through Uintan Desmato- 
therium to Oligocene Colodon. As a consequence, Desmatotherium 
should be included in the Helaletidae, as Simpson (1945) has in- 
dicated, not with Hyrachyus as Peterson (1919) placed it. On the 
other hand, the subfamily separation of the Colodontinae from the 
Helaletinae made by Wortman and Earle (1893), which Simpson has 
preserved, cannot now be reconciled with the sequential arrangement 
indicated above. 


DESMATOTHERIUM WOODI,*® new species 
Plate 2, figure 4 


Type.—Right maxilla with P?-M* (P* incomplete), U.S.N.M. No. 
20200. 

Horizon and locality —Hendry Ranch member of Tepee Trail for- 
mation on south side of Badwater Creek, SW cor., SE} sec. 14, 
T. 39 N., R. 89 W., Wind River Basin, Wyo. 

Specific characters—Upper molar teeth approximately 20 percent 
smaller than in Desmatotherium guyotu Scott or Desmatotherium kayi 
Hough. Upper premolars smaller than in these species but relative 
size intermediate between them and closer to D. guyotit. 

Discussion.—F our incomplete upper dentitions and a number of iso- 
lated teeth, including some from the lower series, all from the Bad- 
water Creek localities, represent the species D. woodi. Two of these, 
part of the type and two upper premolars (part of U.S.N.M. No. 
20202) were figured by Hough (1955, pl. 8, figs. 6 and 9) as material 
referred to the Sage Creek species D. kayi. I have examined all the 
Sage Creek specimens together with the Badwater material and find 
there is no overlap in observed size range for each. The type of the 
Sage Creek species is nearly 20 percent larger than that of the Bad- 
water form. D. kayi was described as close in size to D. “guyotit” 
but with smaller premolars. The premolars of D. woodi are a little 
smaller than in D. kayi, but the ratio of their size to that of the molars 
is more nearly as in D. guyotit. 

The principal feature of the upper dentition of Desmatotherium, 
distinguishing it from Hyrachyus, is the more progressive condition 
of the premolars. The divided lingual portion of P* and P* gives these 


6 Named for H. E. Wood, II, in appreciation of his work on the Hyrachyidae. 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


teeth a distinctly more molariform appearance. Resemblance is im- 
mediately seen to the earlier Helaletes nanus (genotype) which in a 
similar way is distinguished from Helaletes bodps. Washakie D. 
guyotu is a much larger form than Helaletes nanus, but D. woodi is 
nearly intermediate. Certain of the larger Bridger individuals with 
progressive premolars, referred to H. nanus, make a close approach to 
D. woodi in size but the separation of the lingual cusps of P* and P* 
in any case is clearly not so well effected. 

Resemblance of Desmatotherium upper teeth to those of Colodon 
is perhaps even more striking; nevertheless, Colodon can with little 
doubt be defended as distinct from Desmatotherium. The premolars 
of Colodon, particularly P*, would appear to be more progressive and, 
as shown in illustrations given by Scott (1941, pl. 81) of Colodon occi- 
dentalis, the posterior upper premolars, noticeably P*, would appear 
to have better defined, more clearly separated, transverse lophs. More- 
over, a comparison of measurements shows that although D. guyotiu 
is comparable to Colodon occidentalis in size, the latter has distinctly 
wider teeth both in the premolar and molar series. This is perhaps 
most noticeable in the appearance of the posterior loph of the anterior 
molars which is decidedly longer in the illustration of Colodon. 

Lower teeth of Desmatotherium are rather poorly represented, 
except in the Sage Creek collections. They are not known for D. 
guyotu and only certain isolated teeth and tooth fragments are included 
in the materials of D. woodi. Characters of the lower teeth of D. kayi 
were briefly discussed by Hough but somewhat further description, 
particularly a comparison with the earlier Helaletes, seems indicated. 

Lower premolars of Desmatotherium in comparison with those of 
Helaletes are noticeably shortened anteriorly and relatively broad. 
Particularly significant is the distinctly larger entoconid which in 
Colodon is quite as large as the hypoconid. There is little evidence for 
an entoconid in lower premolars of Hyrachyus. The progressive de- 
velopment of the entoconid toward Colodon, and shortening of the 
trigonid, give the premolars a more molariform appearance, but the 
entoconid remains distinct from the hypoconid so that a completely 
lophoid posterior crest as in the molars is never reached. 

The lower molars of D. kayi, as in Helaletes and unlike Hyrachyus, 
show clear-cut transverse lophs with only a very subdued crista be- 
tween them, a tooth form already realized in Heptodon. The para- 
stylid, particularly on M3, seems more reduced in Desmatotherium 
than in Helaletes and much more reduced than in Hyrachyus. The 
hypoconulid on Ms; may be slightly more reduced than in either 
Helaletes or Colodon. The lower molars of Colodon, in addition to 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 19 


their relatively greater width, are more nearly symmetrical bilaterally, 
exhibiting a slight crest forward from both the entoconid and meta- 
conid, quite matching those of the labial side. 

As previously noted, there seems little doubt but that a phyletic 
sequence is indicated from Helaletes through Desmatotherium to 


MEASUREMENTS IN MILLIMETERS OF UPPER DENTITIONS 
oF Desmatotherium woodi 


U.S.N.M. 
No. U.S.N.M. U.S.N.M. 
20200 No. No. 
Type 20201 20202 
Length of upper premolar series, P’-P*, in- 
LEAS TV Cmevetsieee eros syera. si eee cektreferexa ee orat yetaberen Masa ararbce ahah doa tera hehe 33.3 
P’, anteroposterior diameter : greatest trans- 
VERSE GIAITICLSE. fsck c Nehe a eteha Na aceite bie cha PITTS Shak hed 7.5:6.0 
P*, anteroposterior diameter.............0. eee CAA Wiest, aoe 8.3:— 
P*, anteroposterior diameter: greatest trans- 
MECH COIAMIGLER Oice sche. <'p ljaisiatyaleta stoi elie oes 9.0: 11.7 8.9: 11.8 
P*, anteroposterior diameter: greatest trans- 
MCCS MIAINELER se Sccstaes sence te 8.8: 12.4 0.2::12.7 9.1: 12.6 
U.S.N.M 
No. 
20204 
Length of upper molar series, parallel to 
EQOEHE TOW, slonie feiiciawran sateine one seers aaa (WAN cheese Malo achene tate 
M,, anteroposterior diameter perpendicular 
ei EAE TAS IO. 6.) ce <r n,e 2 alone she 350" 11.0 TNO OP eneivenicse 
M,, transverse diameter across parastyle and 
PEOLORO ME raters siche lois ote GANS eho ios Soueie 14.0 TOG eae 
M2, anteroposterior diameter perpendicular 
COU ATRCEAOE THAT RID 0.5 oc ics we wis sere 6 esl TQ WON eaeicte oes 12.8 
Mz, transverse diameter across parastyle and 
PIDLOCOME? dans Sins eee nese e cee LGN ts sdeacee 15.0 
Ms, anteroposterior diameter perpendicular 
FOP ANLEL IONS MIALS 1il15)4.4 siaiete slave eiersiei eres ove TO eK per Bt 13.0 
M;, transverse diameter across parastyle and 
DLOLORON Ee clack e chalet rere Sie cisie aes merreiers Se ein ears 15.1 


Colodon, and this may logically include Heptodon in the lower Eocene 
which, in addition to a much reduced P,, has only slightly less pro- 
gressive premolars than Helaletes. The four genera are not readily 
separated on the basis of molar teeth but a progressive change in the 
premolars is noted, more precocious in tapiroid character than in con- 
-temporary isectolophodont and dilophodont forms (as well as hy- 
rachyid). Nevertheless, this line evidently did not give rise to true 
tapirs. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


DILOPHODON Scott, 1883 


Dilophodon was described by Scott (1883) in the same publication 
as Desmatotherium and, as in the case of D. guyotii, the type of 
Dilophodon minusculus was attributed to the Bridger Eocene. It is 
clear that Scott regarded the Washakie beds as Bridger and it is from 
the Washakie Basin rather than the Bridger Basin, as indicated by 
Granger (1909, p. 22), that the D. minusculus type originated. Granger 
has the species listed as representing Washakie A, but I suspect that 
the horizon for this, as well as D. guyotii, is B, particularly since both 
are known from the upper Eocene elsewhere and neither has turned 
up in the rather extensive collections known from the Bridger proper. 

Dilophodon is clearly related to Helaletes but represents a line 
separate from that of Desmatotherium, possibly derived from the 
species Helaletes bodps, having the less progressive premolars or, as 
seems more than likely, from a somewhat earlier stage. It is not cer- 
tainly demonstrated that Dilophodon gave rise to Protapirus but, as 
far as can be determined, this upper Eocene form possesses all, or 
nearly all, the requirements in the structure of the teeth that might be 
sought for in the Eocene ancestor of the true tapirs. 


DILOPHODON, cf. LEOTANUS (Peterson), 1931 
Plate 1, figures 7, 8 


The type of Peterson’s Heteraletes leotanus from the Randlett lo- 
cality exhibits beyond doubt an immature dentition so that the char- 
acters attributed to the premolar series, particularly the “molariform 
P,,” apply to the deciduous series, and hence do not serve to dis- 
tinguish Heteraletes from Dilophodon. 

In the Badwater collection there is a right mandibular ramus 
(U.S.N.M. No. 20207, figured by Hough, 1955) including all the 
lower cheek teeth, and both maxillae of a skull (U.S.N.M. No. 21098) 
with P*-M® represented, although P*, M*, and M? are not complete. 
There are in addition almost a dozen isolated teeth or portions of teeth. 
Comparison between the Badwater and Randlett materials is limited 
to M, and M,. These teeth in No. 20207 are close in size to those in 
the type, although possibly a trifle wider, and have similar completely 
lophoid transverse crests with practically no development of a crista 
obliqua. 

The Badwater form clearly represents a species with smaller teeth 
than the Washakie Dilophodon minusculus, but the lower jaw is deeper 
and a little more robust. Moreover, the symphysis is broader and ex- 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 21 


tends posteriorly to a position much farther back than in D. minuscu- 
lus. The width of the lower teeth is not significantly different but 
those in the Badwater form are a little shorter, particularly in the pre- 
molar region. A peculiar parallel with the Desmatotherium line is 
noted in the anteroposterior shortening of the anterior or trigonid por- 
tion of the premolars, suggesting that the Badwater form is a little 
more advanced than D. minusculus. This is not an unreasonable sug- 
gestion since the Washakie horizon represented by the latter, though 
possibly earlier than Uinta B, is certainly not later. D. minusculus 
lower teeth, compared in turn with those in Helaletes, are seen to re- 
semble them very closely. I note only the somewhat more progressive 
premolars with distinctly more basined talonids, and the presence of a 
hypoconulid on M3. 

Dilophodon leotanus, though having lower premolar trigonids short- 
ened from the Helaletes stage, has these portions developed for the 
most part about as in Protapirus, not so abbreviated as in Colodon. 
However, P. in the D. leotanus specimen at hand is relatively un- 
developed. Although this tooth shows characters which are probably 
variable, the paraconid and metaconid are scarcely more than crests, 
somewhat as in Colodon. Nevertheless, the talonid is more nearly 
similar to that in Protapirus in that the crest of the hypoconid appears 
to be more median in position as it approaches the trigonid, produc- 
ing a rather distinctive labial fold or depression. 

In the lower molars the parastyle development is rather similar to 
that of Protapirus, although the crosslophs seem more clean-cut. 

Significant features are seen in the upper teeth of the Badwater 
species, and except for M°, these teeth were hitherto not known for 
Dilophodon. It may be noted in particular that P* and P* (P? and P? 
are not known) have a single, undivided lingual cusp or deuterocone 
as in Helaletes bodps, not divided as in the Desmatotherium—Colodon 
line, and that in M* and M? the metacone, though exhibiting a heavy 
cingulum externally, is not concave but distinctly convex labially, so 
that the metacone has a little more conical appearance. Its form, how- 
ever, is not quite comparable to that in Homagalax or in the middle 
and upper Eocene isectolophids which, as Hatcher (1896) pointed 
out, are not entirely suited in this detail as potential ancestors of 
Protapirus. The form of the metacone is unlike Desmatotherium or 
Colodon and different than in most of the Helaletes material examined, 
although in some specimens of the latter the concavity is not empha- 
sized and much of the Heptodon material would not be excluded as 
potentially ancestral. 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I3I1 


The combination of characters seen in the upper dentition is highly 
suggestive of Protapirus and the possibility of an ancestral relation- 
ship is not precluded by the characters of the lower dentition, as it so 


MEASUREMENTS IN MILLIMETERS OF DENTITIONS IN SPECIES OF 


Dilophodon 
Ds ck. 
leotanus 
We . 
21098 
Length of upper dentition, P?-M* inclusive... 02.20. cckecle-saecnse 39.0° 
Length of upper premolars, P?-P* inclusive, at alveoli.............. 17.0° 
Length of upper molar series, M’* (at alveolus)-M® inclusive........ age 
P*, anteroposterior diameter: transverse diameter...............0.. 5.7: 8.2 
M’, anteroposterior diameter perpendicular to anterior margin...... 8.2° 
M*, transverse diameter across parastyle and protocone............. 9.8° 
M®*, anteroposterior diameter perpendicular to anterior margin...... 8.7 
M*, transverse diameter across parastyle and protocone............. 10.4 
D. minus- D,, ct. 
culus leotanus 
P.U. No. U.S.N.M. 
10019 No. 
Type 20207 
Length of lower cheek tooth series, P2-M: inclusive...... 46.6 41.1 
Length of lower premolar series, P2-Ps inclusive........ 18.7 16.2 
Length of lower molar series, M:-Ms inclusive........... 28.3 25.4 
P:, anteroposterior diameter: greatest transverse diam- 
EOL sia ivstarete se Sievoil Wal sieis isin vis ices retelea ai tee vousTepelay ciara lence eke 5.6: 3.4 4.8 : 3.5 
P;, anteroposterior diameter: greatest transverse diam- 
eteric. TOU SEE ae SCARS eo atitoainc.s beets ete aee 6.5: 4.6 Baas 
Ps, anterioposterior diameter: greatest transverse diam- 
GEOR CEA Paras ears eects tere tik ctotne Rare Gan Nah Sees weber eee 6.8: 5.5 6.1: 5.2 
My, anteroposterior diameter: transverse diameter of tri- 
OTM ce stare wavstete eichoereiereie iets sic cretarthans stake plete mien steratces 8.1: 5.2 7.2: 5.2 
M2, anteroposterior diameter: transverse diameter of tri- 
SOW: eh Recta: Uareeaola tis ty fest tetw SRLS haw os She hem aette SS 9.4: 5.9 8.8: 6.1 
Ms, anteroposterior diameter: transverse diameter of tri- 
GORI” 2a. 5r6. he je Be Se OG a ies 6 bible Be eras eee 10.6: 6.3 10.1: 6.5 


¢ Estimated. 


clearly is in the isectolophid line. The phyletic position of Dilophodon 
with respect to Protapirus had been suspected by Peterson (1919, p. 
113) on the basis of the lower dentition, and the likelihood of such 
a relationship seems greatly strengthened by information furnished 
by the upper cheek teeth of Dilophodon, cf. leotanus. 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 23 


HYRACODONTIDAE 
EPITRIPLOPUS?, sp. 
Plate 1, figure 4 


A lower jaw fragment with a well-worn cheek tooth, U.S.N.M. 
No. 21099, evidently a molar, and fragments of two lower molars 
belonging to another specimen are surely rhinocerotid and would ap- 
pear to be hyracodont rather than hyrachyid. I am, nevertheless, un- 
able to determine whether the form represented is Prothyracodon, 
Triplopus, or Epitriplopus. The Badwater teeth are closer in size to 
those in Epitriplopus uintense than they are to those in Prothyracodon 
obliquidens. The teeth also strongly resemble, but are much smaller 
in size than in the Lapoint hyracodont which Peterson unfortunately 
named Mesamynodon medius. The lower tooth in No. 21099 measures 
16.2 mm. long by 11.1 mm. wide. 

Dr. H. E. Wood concurs with me that, of the various possible alloca- 
tions which may be made of this material, Epitriplopus is the most 
probable. 


ARTIODACTYLA 
DICHOBUNIDAE 
PANTACEMYLUS?, sp. 


A homacodont that may well be Pentacemylus is represented by the 
posterior two-thirds of M;. There is no certainty, however, that the 
form is not Mytonomeryx. The tooth conforms closely in form of 
cusps and is only very slightly smaller than in Pentacemylus pro- 
gressus. It does not appear to represent the smaller Bunomery-. 


APRICULUS,’ new genus 


Type.—Apriculus praeteritus, new species. 

Generic characters—Simple conical cusps on upper molars as in 
Helohyus, but these teeth more nearly quadrate with large lingually 
placed metaconule on all three. Cingulum continuous around molars 
without external styles and without evidence of a hypocone. Proto- 
conule slightly better defined than in Helohyus. P* with single primary 
cusp and deuterocone. 

Discussion.—A priculus is almost certainly a descendant of Bridg- 
erian Helohyus and belongs in the Helohyinae, but its trend has been 


7 Apriculus, diminutive of Aper (L.), wild boar. 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


more conservative and along a line independent of that for Achaeno- 
don. Change from Helohyus has been the increase in size of the meta- 
conule, and the shift to a more lingual position. This is particularly 
noticeable for M* which, rather than having a triangular outline, nas 
achieved the quadrate form of the anterior molars with the metaconule 
equally well developed. There is, however, no evidence of the devel- 
opment posteriorward of M® as in Perchoerus; nor do any of the 
molars show development of a “pseudometaconule” anterolateral to 
the metaconule. 

Distinction from Achaenodon is seen not only in the very much 
smaller size of Apriculus so far as known, but in the retention and 
marked development of the protoconule. 


APRICULUS PRAETERITUS,® new species 
Plate 3, figure 1 


Type.—Right maxilla with P*-M* (teeth incomplete), U.S.N.M. 
No. 21100. 

Horizon and locality—Hendry Ranch member of Tepee Trail for- 
mation on south side of Badwater Creek, SW cor., SE} sec. 14, 
T. 39 N., R. 89 W., Wind River Basin, Wyo. 

Specific characters.—Size of upper molars very close to that for 
Helohyus plicodon. Other differences included in description of genus. 

Discussion—The type maxilla with somewhat damaged upper teeth 
is the only known specimen of this comparatively late survival of a 
Helohyus-like dichobunid. The upper molars, though comparable in 
size to those of Helohyus plicodon, are perhaps a little narrower trans- 
versely, as the basal slope of the protocone and cingulum median to it 
do not extend so far lingually. The anterointernal and posterointernal 
portions of the molars are more nearly equal in lingual extent. 

Precise measurements of the individual teeth in the type of Apric- 
ulus praeteritus cannot be obtained because of breakage; nevertheless 
over-all dimensions can be determined and the length of the pre- 
served portion of the upper cheek tooth series, P*-M* inclusive, is 
found to be 30 mm. The molar series is about 24.5 mm. long. M* 
is estimated to be about 8.7 (anteroposteriorly) by 10.6 mm. 

In size, upper molars of Apriculus praeteritus are only a little larger 
than the dichobunid tooth figured by Russell and Wickenden (1933). 


8 Praeteritus, overlooked or passed over—overlooked in recent review of col- 
lections for upper Eocene artiodactyl study. 


~ 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 25 


The Swift Current Creek ®° specimen, however, is distinctly less quad- 
rate, and the metaconule, rather than occupying the posterolingual 
angle of the tooth, is between the protocone and metacone. More- 
over, there appears to be a hypocone, or at least a well-developed 
crest, posterior to the protocone. The structural resemblance of this 
tooth to both the leptochoerids and diacodexids was noted by Russell 
and Wickenden. Possibly further evidence bearing on the relation- 
ship suggested (Gazin, 1955) for these two groups is to be found in 
the Swift Current Creek beds when the fauna from there is better 
known. 

Apriculus praeteritus will not be confused with the Helohyus?, sp. 
described by Peterson (1934) from the Lapoint Duchesnean. Although 
direct comparison in details of teeth is precluded by the different 
nature of the known material, the disparity in size is alone conclusive, 
at least as far as species are concerned. The Lapoint specimen, as 
indicated by Peterson, is rather close in size to Helohyus lentus, and 
although the premolar, disregarding the small parastylid, is rather like 
that in Bridger H. lentus, the molar is not particularly close. The 
paraconid in M,, as shown in Peterson’s illustration, is much too far 
forward. Helohyus in general does not show the crest extending 
posteriorly from the protoconid or the triangular-shaped basin formed 
between this crest and crista obliqua observed in the Lapoint M,. In 
Helohyus the low crista obliqua extends forward to a much more 
buccal position on the trigonid, with a well-formed basin posterolingual 
to this crest. 

I strongly suspect that the Lapoint specimen is a very small en- 
telodont. The crest pattern of the molar which Peterson so clearly 
described and as outlined above, while unlike that of Helohyus, can 
be clearly, though weakly, discerned in unworn first and second lower 
molars of Archaeotherium. In a footnote Peterson called attention to 
the resemblance of P, to that in Archaeotherium, but discredited such 
a relationship by the presence of a paraconid on M,. This reasoning 
I cannot understand, as the lingual portion of the trigonid of not too 
well worn lower molars in the Oligocene form usually shows two 


9 Omitting consideration of the Saskatchewan Swift Current Creek beds and 
fauna in my review of upper Eocene artiodactyls was a regrettable oversight and 
should in no way be regarded as implying a lack of significance. The materials, 
though fragmentary, give us the only glimpse so far obtained of the nature of 
the upper Eocene fauna at a latitude so far north. It is only from such Canadian 
discoveries that speculation by various paleontologists on the possibility of more 
northern origins of early Tertiary groups with obscure ancestry may be 
evaluated. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


clearly defined and well-separated cusps. Whether the anterior of 
these originated by twinning or from the cingulum, or whether the 
two cusps are actually the metaconid and metastylid, would not seem 
to alter the picture. If subsequent material should demonstrate that 
the Lapoint specimen is indeed of a small primitive entelodont the 
possibility of the entelodonts having originated early in the Helohyinae 
would not seem to be precluded. Although Peterson’s specimen is 
unlike Helohyus in details seemingly on a generic level, the bunodont 
form of the Lapoint molar could possibly be regarded as a modifica- 
tion from that of Helohyus or Lophiohyus. The interval from Bridger 
to Lapoint time would seem entirely adequate. 

I agree with Peterson that the Lapoint specimen is probably inade- 
quate as a type, nevertheless its possible new-born significance may 
warrant a name, if for no other purpose than as a handle for discus- 
sion purposes. I propose the new name Dyscritochoerus *° lapointen- 
sis.11 The type is the lower jaw portion described by Peterson, C.M. 
No. 11912.'? It might tentatively be aligned with the entelodonts. If 
Dyscritochoerus is in truth a link between the entelodonts and the 
helohyids its position in the uppermost Eocene is reasonable and would 
not necessarily imply an Oligocene age for the Lapoint horizon. 


AGRIOCHOERIDAE 
PROTOREODON, cf. PETERSONI Gazin, 1955 
Plate 3, figure 4 


Not more than four specimens represent a distinctly small species 
of Protoreodon in the Badwater fauna. Three of these are isolated 
teeth, but one, U.S.N.M. No. 21101, is a right maxilla including M*- 
M®*. The teeth correspond closely in size to those in Uinta C Pro- 
toreodon petersoni, and, like that species, the protoconule is rather 
weak, not so emphasized as in earlier Uinta C Protoreodon minor. 
The molar series in No. 21101 is 20.0 mm. long. 


PROTOREODON, near P. PUMILUS (Marsh), 1875 


There are about 18 specimens of a comparatively large form of 
Protoreodon. Most of these are isolated teeth, although a few are 
jaw or maxillary fragments with two or three teeth, generally broken. 


10 Dyskritos (Gr.), hard to determine or doubtful, and choiros (Gr.), pig. 

11 Japointensis, from the town and horizon Lapoint. 

12 Dr. Kay informs me that he has been unable to locate this specimen in the 
collections at the Carnegie Museum. 


No. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 27 


These teeth are all of about the size of those in Uintan P. pumilus. 
There is, moreover, no certain evidence that another species such as 
Protoreodon primus is not represented. 


PROTOREODON PEARCEI,!® new species 
Plate 3, figures 7, 8 


Type.—Skull, jaws, and other portions of skeleton, U.S.N.M. No. 
20305. 

Horizon and locality—Hendry Ranch member of Tepee Trail for- 
mation on south side of Badwater Creek, SW4 sec. 13 near line be- 
tween sections 13 and 24, T. 39 N., R. 89 W., Wind River Basin, 
Wyo. 

Specific characters—Larger and more robust than Protoreodon 
pumilus, very close in size to Diplobunops matthewi. Marked diastema 
between canine and P?. 

Discussion.—Although this specimen had been early regarded as 
Protoreodon primus (see Hough, 1955) it is readily distinguished 
from this species and the advanced Protoreodon pumilus annectens by 
its distinctive size. It is much more easily confused with the equally 
large Diplobunops matthewi. There are several isolated teeth and in- 
complete dentitions that appear to represent this very large protoreo- 
dont, but a number, more fragmentary or too well worn, cannot be al- 
located as between this form and Diplobunops, cf. matthew. 

The skull of Protoreodon pearcei differs rather noticeably from that 
of Diplobunops, cf. matthewi in the Badwater collection (see Gazin, 
1955, pls. 10-12) in the anterior extremity of the rostrum. Although 
the canines are actually larger and as far apart in P. pearcet, the snout 
extremity does not appear so bluntly expanded, evidently because the 
palate is not so noticeably constricted behind P+. There is a diastema 
between the canine and P? about the same length as in the Diplobunops 
specimen but there is no diastema behind P?, and the premolars are 
distinctly crowded. P* shows a slight basin posterointernally but 
no deuterocone. P* has a smaller deuterocone but a distinctly better 
defined basin posterointernally than in the Diplobunops material, and 
in P* there is clearer evidence of a tritocone. The upper molars show 
more lingually sweeping outer crescents and the protoconule may be a 
little weaker. This is particularly true of M°. 

The lower jaw is not so constricted through the symphysial portion 
as it is in the Diplobunops material figured by Scott (1945, pl. 5, fig. 


13 Named for Franklin L. Pearce, who found the type specimen. 


28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


2). P, is a very large caniniform tooth, closely followed by the suc- 
ceeding, noticeably overlapping premolars. I have not observed signifi- 
cant differences in the lower premolars and molars although from the 
material at hand the lower cheek teeth of P. pearcei are a little nar- 
rower than in D., cf. matthew. 

Protoreodon pearcei makes a close approach to Agriochoerus 
antiquus in size and in the presence of a short though distinct 
diastema between the upper canine and P*. There is, however, no 
diastema between P, and P,. Moreover, the posterior premolars above 
and below are not nearly so progressive. Also, the protoconule, though 
very weak on M®, is clearly defined on M? and M?. 

I am not certain that this species is represented in Uinta collections, 
but a robust jaw from Leland Bench Draw with closely crowded pre- 
molars and no diastema between P; and P2 may represent P. pearcei 
rather than Diplobunops matthewt. Measurements of this jaw were 
given (Gazin, 1955, p. 64) in comparison with the type materials, 
showing the shortness of the space occupied by the premolar sequence. 

Protoreodon pearcei is apparently represented in the Sage Creek 
area collections by a skull which has the Carnegie Museum number 
8927. It was collected by J. L. Kay in 1940 and the catalog card 
carries the information “Spring Gulch, Sage Creek.” The information 
“Oligocene (Cook Ranch)” also appears on the label, but this informa- 
tion is surely a misinterpretation of the horizon represented. The 
skull was figured by Hough (1955, pl. 8, fig. 8) as “Mesagriochoerus, 
cf. primus” and the catalog number is incorrectly cited as “9827.” 

Measurements for the teeth in the type of P. pearcei are given with 
those for Diplobunops, cf. matthew in the following section. 


DIPLOBUNOPS, cf. MATTHEWI Peterson, 1919 
Plate 3, figure 6 


In contrast to the rather small ratio of Diplobunops to Protoreodon 
specimens encountered in the Uinta basin, Diplobunops is almost as 
abundant in the Badwater collection as Protoreodon. Remains of this 
comparatively large agriochoerid include some of the better preserved 
materials representative of the fauna and the least distorted known 
for the genus. Two excellent skulls were collected by Harry A. 
Tourtelot, one of these, U.S.N.M. No. 20303, has been previously 
figured (Gazin, 1955, pls. 10-12). 

The distinction between Diplobunops and Protoreodon on the basis 
of isolated teeth is difficult to make, particularly in the Badwater 
materials, because with the recognition of the equally large Pro- 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 29 


MEASUREMENTS IN MILLIMETERS OF DENTITIONS IN SPECIMENS OF 
Protoreodon pearcet AND Diplobunops, cr. matthewi 


D., cf. matthewi P. pearcei 
—Oe U.S.N.M. 
U.S.N.M. U.S.N.M. No. 
No. No. 20305 
20304 20303 Type 
Length of upper dentition, C (at alveolus) - 

Mes HACHISEV ERS ets dc Aces Kom ecaaiean! Gee vaiere ose 06.5 90.5 
Length of upper dentition, P*-M*, inclusive. ........ 83.0 74.0 
Upper premolar series, P*-P*, inclusive.... ........ 44.0 36.5 
Upper molar series, M*-M®, inclusive...... 30.5 40.0° 30.6 
C, anteroposterior diameter (at alveolus) : 

greatest transverse diameter.......... Nevercrarerers TG Gel 11.0: 8.7 
P’, anteroposterior diameter : greatest trans- 

DErse -GIAMIELEL. wives tec ae eos chee os 9.0: 3.8 Wie Aig 
P?, anteroposterior diameter : greatest trans- 

yerse Giatieter! Voc... s ses Ree 9.6: 5.9 10.3 : 6.4 17123365" 
P*, anteroposterior diameter : transverse di- 

AIMELOE Fi os iccinccais ese ca geee Leper 10.2:10.2 10.3:9.8 II.1: 9.5 
P*, anteroposterior diameter : transverse di- 

BEIGSTCL sin c/aton& Sats Sa a acchaca travers Pl or 10.7 $}F2)3 \ | 50.013 "12.9 10.2: 12.5 
M’, anteroposterior diameter : transverse di- 

USICLEL, Gel hate hiv sfexehe Scr sihcis adereintatensenciontoe 12.8:15.1 12.07: — 12.5) 13:5 
M’, anteroposterior diameter : transverse di- 

AMELE EA e or tec ae corns echoes chat facle owas sete 1383160, 613:8:2.17:5 13.72>16:6 
M*, anteroposterior diameter : transverse di- 

PEPER stele wicen, elena isis cists wore lera éjea lm esate 13-72 19.0 14.6°2 19.5", 15.07 18:3 
Length of lower cheek tooth series, P: (at 

BIVEOIUS )— Dis MICINSIVE. Sos cbs fee fale cite d's MEL fees. eats ec 82.5 
Length of lower premolar series, P: (at 

AIVERNUS = Eay  THCMUSIVE ia ciciarcitjarsitjainiaeoieiele ees 43.2 38.3 
Length of lower molar series, M:-Ms, in- 

MIS VEL Meade eta cbse cbere (ein $e» cherace fe ahs dpsed siete ahr pea abee'’s hint over 44.0 
Px, anteroposterior diameter (at alveolus) : 

PLCALESE LLATISVERSE Ciametety se cee aie ocrlts cakeicletah ec elaisietelererdls 10.0: 8.3 
P,, anteroposterior diameter : greatest trans- 

VEUSE WOIAIICLER, jsaraa ti ethe etan tiers crecosteys cifarersi els avb ave 10.0: 5.3 90.5: 4.6 
P;, anteroposterior diameter : greatest trans- 

MEESEMOIAIMELEEN, Ware eaataue Peccrere bis os Gisieiavuelenierere is ET0;:'7.0 10.8 : 6.2 
P,, anteroposterior diameter : greatest trans- 

WEMGE TGATNCLEE? Fisica cence ciate ceeds Vsetaae ss 12.2: 8.0 1153 7.2 
M,, anteroposterior diameter : transverse di- 

AINETEDHOL \talONIG'. 2:22 sc/ers seve oes viaetena betaine  Lesieshtanctae 10.9: 8.1 
M;, anteroposterior diameter : transverse di- 

AMeLereOe talOnid’ s,.../aed adale sjagidcee aps da welace TAA 1-3" “1F0Fs0.3 
Ms,, anteroposterior diameter : transverse di- 

AMMELEL TOL THISOMIM kaki siclerdewhactiede! ete sane we —:10.7 20.3: 10.4 


2 Approximate. 

* Measurements of posterior upper premolars are taken anteroposteriorly across outer 
portion and transversely perpendicular to outer margin. Those of upper molars are taken 
anteroposteriorly perpendicular to anterior margin and transversely across anterior portion 
of tooth, lingually to base of enamel or cingulum. 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


toreodon pearcet size is no longer an aid. Teeth of Diplobunops are 
slightly less selenodont. The inner and outer crests of the upper molars 
are seemingly farther apart with the outer cusps a trifle more erect and 
their apices more buccal or not so lingually directed. The cusps or 
crests of the lower molars have a slightly more inflated or obtuse look 
and may be relatively broader. There is, of course, variation in both 
forms ; moreover, these characters of the teeth are scarcely of generic 
significance and seem somewhat distinctive only between contempo- 
raries in these groups in the higher horizons of Uintan time. 
Diplobunops, so far as known, did not progress much beyond the 
Protoreodon tooth structure characterizing Uinta B time. The genera 
are, as I have attempted to show earlier (1955), very closely related 
and their skulls may be distinguished essentially on the different char- 
acter of the anterior part of the snout. 


OROMERYCIDAE 
MALAQUIFERUS TOURTELOTI Gazin, 1955 
Plate 3, figure 2 


The type of this form is from the Dry Creek occurrence some 20 
miles to the west of the Badwater Creek localities. Its description and 
illustration were included in an earlier paper (1955, pl. 16), and need 
not be repeated here. An isolated oromerycid upper molar (U.S.N.M. 
No. 21102) in the Badwater collection exhibits the same rectangular 
form, rugosity, posteriorly bifurcate protocone, and outstanding ribs 
on the outer cusps as in Malaquiferous tourteloti. It corresponds 
closely in form and size, and in the possession of a slightly outward- 
deflected metastyle to M*. It differs in having a heavier, antero- 
posteriorly developed mesostyle. The shape of this tooth is entirely 
different, although structurally related to that in Oromeryx plicatus. 
Likewise, it would not be confused with the comparatively large 
Protylopus? annectens. 

This tooth lends support to the belief that the Dry Creek occurrence 
is probably equivalent in time to that at Badwater, a conclusion earlier 
reached on the basis of lithology, and the general upper Eocene indica- 
tion by the presence of Eomoropus. 


LEPTOMERYCIDAE 
LEPTOTRAGULUS, cf. MEDIUS Peterson, 1919 


Plate 3, figure 5 


Two lower jaw portions, and probably several of the isolated molars, 
represent a form close or identical to the upper Uintan Leptotragulus 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 31 


medius. One of the jaw portions (U.S.N.M. No. 21104) with P, and 
M, falls within the size range of the Myton material, but the other 
(U.S.N.M. No. 21103), which has P;-M,, is a little larger than any 
in the above series. Nevertheless the latter is closer in size of teeth 
to L. medius than it is to L. proavus. The structure of the lower pre- 
molars in these jaws corresponds very closely to that regarded as 
characterizing Leptotragulus. 

The teeth in both Nos. 21103 and 21104 are significantly smaller 
than in the type of Leptotragulus? significans Russell from the Kish- 
enehn beds in British Columbia. Although I have not examined 
the Kishenehn specimen, from Russell’s clear description and stereo- 
scopic illustrations I would favor referring his form to Leptomeryx 
rather than Leptotragulus. As a consequence, there would seem to be 
somewhat better evidence for the Oligocene age postulated by Russell 
as an alternate possibility for the Canadian occurrence. 


LEPTOREODON?, sp. 
Plate 3, figure 3 


A couple of isolated premolars in the collection, a little smaller than 
in Leptoreodon marshi but not greatly different in size from Lepto- 
tragulus medius, exhibit a well-defined metaconid. In one of these 
the metaconid is opposite the protoconid and shows a well-defined 
groove between them anteriorly, much as in characteristic material 
of Leptoreodon. The parastylid, however, is a well-developed column 
distinct from the anterior crest of the protoconid, suggestive of 
Leptomeryx. Nevertheless, as in Leptoreodon and unlike Leptomery., 
the talonid basin is formed by the posteroexternal crest swinging 
lingually near its posterior extremity, joined only by a weak spur 
from the metaconid. In Leptomeryx, the hypoconid and entoconid 
in P, are in most cases sharply separated and join forward with the 
protoconid and metaconid respectively. The second isolated P, ex- 
hibits a parastylid much as in Leptotragulus, but the metaconid, 
though weak, is distinct and unlike Leptotragulus. This tooth has a 
very primitive look and may not represent any known leptotragulids. 


REFERENCES 


Burke, JoHN J. 
1934. Mytonolagus, a new leporine genus from the Uinta Eocene series in 
Utah. Ann. Carnegie Mus., vol. 23, pp. 390-420, pl. 50. 
CLarK, JOHN. 
1939. Miacis gracilis, a new carnivore from the Uinta Eocene. Ann. Car- 
negie Mus., vol. 27, pp. 349-370, figs. 1-2, pls. 34-37. 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Corr, Epwarp D. 
1881. The systematic arrangement of the order Perissodactyla. Proc. Amer. 
Philos. Soc., vol. 19, pp..377-401, figs. I, 2. 
Gazin, C. Lewts. 
1955. A review of the upper Eocene Artiodactyla of North America. Smith- 
sonian Misc. Coll., vol. 128, No. 8, pp. 1-96, charts 1-2, pls. 1-18. 
GRANGER, WALTER. 
1908. A revision of American Eocene horses. Bull. Amer. Mus. Nat. Hist., 
vol. 24, pp. 221-264, figs. 1-5, pls. 15-18. 
1909. Faunal horizons of the Washakie formation of southern Wyoming. 
Bull. Amer. Mus. Nat. Hist., vol. 26, art. 3, pp. 13-23. 
HatTcHER, JOHN B. 
1896. Recent and fossil tapirs. Amer. Journ. Sci., ser. 4, vol. 1, pp. 161-180, 
figs. I-2, pls. 2-5. 
HovucH, JEAN R. 
1955. An upper Eocene fauna from the Sage Creek area, Beaverhead 
County, Montana. Journ. Paleont., vol. 29, No. 1, pp. 22-36, figs. 
1-3, pls. 7-8. 
Kay, J. LeRoy. 
1953. Faunal lists of vertebrates from Uinta Basin, Utah. Guide Book, 5th 
Ann. Field Conf., Soc. Vert. Paleont. in N.E. Utah, pp. 20-24. 
Lewis, G. Epwarp. 
1947. Tertiary geology of northwestern Wind River Basin, Wyoming (ab- 
stract). Journ. Washington Acad. Sci., vol. 37, p. 371. 
Love, Joun D. 
1939. Geology along the southern margin of the Absaroka Range, Wy- 
oming. Geol. Soc. Amer. Spec. Pap. No. 20, pp. 1-134, figs. 1-3, 
pls. I-17. 
MarsH, OTHNIEL C. 
1871. Notice of some new fossil mammals from the Tertiary formation. 
Amer. Journ. Sci. and Arts, vol. 2, pp. 35-44. 
1875. Notice of new Tertiary mammals—IV. Amer. Journ. Sci. and Arts, 
vol. 9, pp. 239-250. 
MatrHew, WILLIAM D. 
1909. Faunal lists of the Tertiary Mammalia of the West. U. S. Geol. Surv. 
Bull. 361, pp. 91-138. 
Osszorn, Henry F. 
1895. Fossil mammals of the Uinta Basin. Expedition of 1894. Bull. Amer. 
Mus. Nat. Hist., vol. 7, pp. 71-105, figs. 1-17. 
1902. American Eocene Primates, and the supposed rodent family, Mixo- 
dectidae. Bull. Amer. Mus. Nat. Hist., vol. 16, art. 17, pp. 169-214, 
figs. I-40. 
1913. Eomoropus, an American Eocene chalicothere. Bull. Amer. Mus. Nat. 
Hist., vol. 32, pp. 261-274, figs. 1-9. 
PETERSON, Oar A. 
1919. Report upon the material discovered in the upper Eocene of the 
Uinta basin by Earl Douglas in the years 1908-1900, and by O. A. 
Peterson in 1912. Ann. Carnegie Mus., vol. 12, pp. 40-168, figs. 
I-19, pls. 34-47. 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 33 


1931. New species from the Oligocene of the Uinta. Ann. Carnegie Mus., 
vol. 21, pp. 61-78, figs. 1-12, pl. 1. 
1934. List of species and description of new material from the Duchesne 
River Oligocene, Uinta Basin, Utah. Ann. Carnegie Mus., vol. 23, 
PP. 373-380, figs. 1-8. 
RusseELL, Loris S. 
1954a. The Eocene-Oligocene transition as a time of major orogeny in west- 
ern North America. Trans. Roy. Soc. Canada, ser. 3, vol. 48, sec. 
4, pp. 65-69. 
1954b. The mammalian fauna of the Kishenehn formation, southeastern 
British Columbia. Ann. Rep. Nat. Mus. Canada (1952-1953), Bull. 
132, pp. 92-111, figs. 1-8, pls. 1-3. 
Russet, Loris S., and WICKENDEN, R. T. D. 
1933. An upper Eocene vertebrate fauna from Saskatchewan. Trans. Roy. 
Soc. Canada, ser. 3, vol. 27, sec. 4, pp. 53-65, fig. 1, pl. 1. 
Scott, WILLIAM B. 
1883. On Desmatotherium and Dilophodon, two new Eocene lophiodonts. 
Contr. E. M. Mus. Geol. and Archaeol. Princeton College, Bull. 
No. 3, pt. 4, pp. 46-53, pl. 8. 
1941. The mammalian fauna of the White River Oligocene. Trans. Amer. 
Philos. Soc., vol. 28, pt. 5. Perissodactyla, pp. 747-980, figs. 137-157, 
pls. 79-100. 
1945. The Mammalia of the Duchesne River Oligocene. Trans. Amer. 
Philos. Soc., vol. 34, pt. 3, pp. 209-253, pls. 1-8. 
Simpson, Georce G. 
1945. The principles of classification and a classification of mammals. 
Bull. Amer. Mus. Nat. Hist., vol. 85, pp. i-xvi, 1-350. 
TourTELOT, Harry A. 
1946. Tertiary stratigraphy in the northeastern part of the Wind River 
Basin, Wyoming. U. S. Geol. Surv. Oil and Gas Invest. Prelim. 
Chart 22. 
1948. Tertiary rocks in the northeastern part of the Wind River Basin, 
Wyoming. Guide Book, 3rd Ann. Field Conf., Soc. Vert. Paleont. 
in S.E. Wyoming, pp. 53-67. 
1953. Geology of the Badwater area. U. S. Geol. Surv. Oil and Gas. Invest. 
Map OM124 (2 sheets). 
Woop, Apert FE. 
1949. Small mammals from the uppermost Eocene (Duchesnean) near 
Badwater, Wyoming. Journ. Paleont., vol. 23, No. 5, pp. 556-565, 
figs. 1-24. 
Woop, Horace E., II. 
1934. Revision of the Hyrachyidae. Bull. Amer. Mus. Nat. Hist., vol. 67, 
pp. 181-295, figs. 1-51, pls. 20-24. 
Woon, Horace E., II; Seton, Henry; and Hares, CHArLes J. 
1936. New data on the Eocene of the Wind River Basin, Wyoming (ab- 
stract). Proc. Geol. Soc. Amer. for 1935, pp. 394-395. 
WortMAn, Jacos L., and EarLe, CHARLES. 
1893. Ancestors of the tapir from the lower Miocene of Dakota. Bull. 
Amer. Mus. Nat. Hist., vol. 5, pp. 159-180, figs. 1-7. 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


EXPLANATION OF PLATES 
PLATE I 


LAGOMORPH, CARNIVORE, CONDYLARTH, AND PERISSODACTYLS FROM 
THE BADWATER UPPER EOCENE 


Fic. 1. Mytonolagus wyomingensis A. E. Wood: Right maxilla (U.S.N.M. 
No. 21090), occlusal view of teeth. 4Xnatural size. Badwater upper 
Eocene, Wind River Basin, Wyo. 

Fic. 2. Miacis, cf. robustus (Peterson): Left ramus of mandible (U.S.N.M. 
No. 21087), occlusal and lateral views. Natural size. Badwater upper Eo- 
cene, Wind River Basin, Wyo. 

Fic. 3. Hyopsodus, cf. uintensis Osborn: Right upper molar (U.S.N.M. No. 
21089), occlusal view. 4 Xnatural size. Badwater upper Eocene, Wind 
River Basin, Wyo. 

Fic. 4. Epitriplopus?, sp.: Left lower molar (U.S.N.M. No. 21099), occlusal 
view. Natural size. Badwater upper Eocene, Wind River Basin, Wyo. 
Fic. 5. Epihippus, cf. gracilis (Marsh): Right maxilla (U.S.N.M. No. 21092), 
occlusal view. Twice natural size. Badwater upper Eocene, Wind River 

Basin, Wyo. 

Fic. 6. Epihippus, cf. parvus Granger: Right maxilla (U.S.N.M. No. 21091), 
occlusal view. Twice natural size. Badwater upper Eocene, Wind River 
Basin, Wyo. 

Fics. 7, 8. Dilophodon, cf. leotanus (Peterson): 6, Right upper cheek teeth 
(U.S.N.M. No. 21098), occlusal view (incomplete P* reversed from left 
side). 7, Right lower cheek teeth (U.S.N.M. No. 20207), occlusal view. 
One and one-half times natural size. Badwater upper Eocene, Wind River 
Basin, Wyo. 


PLATE 2 
PERISSODACTYLS FROM THE DRY CREEK AND BADWATER UPPER EOCENE 


Fics. 1-3. Eomoropus anarsius, new species: 1, Skull and left ramus of mandi- 
ble (U.S.N.M. No. 21097), type specimen, lateral view. Two-fifths natural 
size. 2, Left upper molars (U.S.N.M. No. 21097), type specimen, occlusal 
view. Natural size. 3, Left lower cheek teeth (U.S.N.M. No. 21097), type 
specimen, occlusal view. Natural size. Dry Creek upper Eocene, Wind 
River Basin, Wyo. 

Fic. 4. Desmatotherium woodi, new species: Composite right upper cheek 
tooth series; premolars (U.S.N.M. No. 20202), occlusal view; molars 
(U.S.N.M. No. 20200), type specimen, occlusal view. Natural size. Bad- 
water upper Eocene, Wind River Basin, Wyo. 


PLATE 3 


ARTIODACTYLS FROM THE BADWATER UPPER EOCENE 


Fic. 1. Apriculus praeteritus, new genus and species: Right maxilla (U.S.N.M. 
No. 21100), type specimen, occlusal view. Twice natural size. Badwater 
upper Eocene, Wind River Basin, Wyo. 


no. 8 MAMMALIAN FAUNA, BADWATER AREA—GAZIN 35 


Fic. 2. Malaquiferus tourteloti Gazin: Right upper molar (U.S.N.M. No. 
21102), occlusal view. Twice natural size. Badwater upper Eocene, Wind 
River Basin, Wyo. 

Fic. 3. Leptoreodon?, sp.: Left Ps (U.S.N.M. No. 21105), occlusal view. 
Twice natural size. Badwater upper Eocene, Wind River Basin, Wyo. 

Fic. 4. Protoreodon, cf. petersoni Gazin: Right maxilla (U.S.N.M. No. 211or), 
occlusal view. Twice natural size. Badwater upper Eocene, Wind River 
Basin, Wyo. 

Fic. 5. Leptotragulus, cf. medius Peterson: Right ramus of mandible 
(U.S.N.M. No. 21103), occlusal and lateral views. Twice natural size and 
natural size, respectively. Badwater upper Eocene, Wind River Basin, Wyo. 

Fic. 6. Diplobunops, cf. matthewi Peterson: Right upper cheek tooth series 
(U.S.N.M. No. 20304), occlusal view (P? and P* restored from left side). 
Natural size. Badwater upper Eocene, Wind River Basin, Wyo. 

Fics. 7, 8. Protoreodon pearcei, new species: 7, Right upper cheek teeth 
(U.S.N.M. No. 20305), type specimen, occlusal view. Natural size. 
8, Right lower cheek teeth (U.S.N.M. No. 20305), type specimen, occlusal 
view (P; reversed from left side). Natural size, Badwater upper Eocene, 
Wind River Basin, Wyo. 





SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 8, PL. 1 





LAGOMORPH, CARNIVORE, CONDYLARTH, AND PERISSODACTYLS FROM 
THE BADWATER UPPER EOCENE 


(See explanation of plates at end of text.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOEScISi5 INO. 8) Pln2 





PERISSODACTYLS FROM THE DRY CREEK AND BADWATER UPPER EOCENE 


(See explanation of plates at end of text.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 8, PL. 3 





ARTIODACTYLS FROM THE BADWATER UPPER EOCENE 


(See explanation of plates at end of text.) 


a 


~ 





SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 9 


3REEDING AND OTHER HABITS OF CASQUED 
JORNBILLS (BYCANISTES SUBCYLINDRICUS) 


(Wir 6 PLaTEs) 


By 
LAWRENCE KILHAM 
Bethesda, Md. 











Steed, 

*% |}. A te Pi . é 

REF HSON OF" ° 
See HINGTOW Se 


C2000 00?* 


(PusticaTion 4259) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
NOVEMBER 8, 1956 


THE LORD BALTIMORE PRESS, INC. 
BALTIMORE, MD., U. S. A. 


PREFACE 


I went to Uganda at the invitation of the East African High Com- 
mission to carry on virus research as a visiting scientist at the Virus 
Research Institute, Entebbe, where I worked from August 1954 until 
mid-May 1955. My ornithological observations were made as an ama- 
teur in the early mornings and evenings, and on weekends. It had been 
my hope to study some particular field problem in addition to making 
a general acquaintance with African bird life. The nature of the prob- 
lem was determined soon after my arrival. In my bird notes, black- 
and-white casqued hornbills [Bycanistes subcylindricus (Sclater) | 
soon took up more pages than any other species. They came to our 
garden frequently. In addition, a pair of them roosted and carried 
on courtship activities in a tree above our house. When I discovered 
a concentration of hornbill nests in the Mpanga Research Forest, it 
was apparent that I had an unusual opportunity to study the natural 
history of casqued hornbills. Present studies did not begin until many 
females were already walled in. A few pairs of late-nesting hornbills, 
however, enabled me to witness the beginning stages of nesting ac- 
tivity. Observations on 16 nesting pairs gave, in the aggregate, a 
rounded picture of breeding and other habits of these birds. As far 
as I am aware, this is the first detailed description published on the 
natural history of Bycanistes subcylindricus. Moreau (1936), how- 
ever, has written of a related species, Bycanistes brevis. His account 
is based on the histories of two nests that he observed in Usambara, 
Tanganyika. 

Acknowledgments.—The writer is grateful to the following indi- 
viduals for help contributed in various ways: Dr. A. J. Haddow, 
director, and Dr. W. H. R. Lumsden, assistant director, of the East 
African Virus Research Institute ; H. C. Dawkins, ecologist, Uganda 
Forest Department; Charles Sandison, curator, Botanical Gardens, 
Entebbe, for identification of fruits and seeds; Dr. V. G. L. Van 
Someren, Ngong, Kenya, for identifying insect remains; and Dr. 
Herbert Friedmann, curator, division of birds, U. S. National Mu- 
seum, for aid and encouragement in preparation of the manuscript. 
Two sketches of hornbills by their nests were contributed by my wife, 
Jane Kilham. The avian scientific nomenclature used is that given by 
Mackworth-Praed and Grant, 1952. 

LK. 


iii 





CONTENTS 


Page 
eT Ce eee epee ea oy tres Sahat RG eee eve aT aT aT et bnb celal rac ohies Shovel a eseelie ayocabe\wis a's iil 
MEIER OG UCTIONS «art ccyrersis eis er atsk isso sister ctoveserehete rer ave) eVoid susie ayers areatels © sieraverets)e.sleve ie I 
IRS E EE OS Beate ciara zicte lw ceeiahere are STe Levee he Ste Eo sla eue) shaves aie) Sielaharetwrel'ara vole o'2 o.bleia) 2 2 
Generalvhabitstorehornbills areas oe rareeietoie wateke aici al clare eiaimiaie le ered slevaleleisiore 4 
ESE CONSEDUCE Of i cpcinis clare araie erat Sroferera her siets avatars isiforeiatel scaimlarel adit die\ejessisieve ina erat 8 
PNGHIVITIES s Ole TCSEITIS PAILS cca clofnh cxe/etstelsveraravetste. ciel ctelers Mines isles! <jales\ syaieseps"s eters 17 
BS OSHVOUN et and NESE OPENING Sine ais steven eferedevaialetslévewiareveisis cuseisiereisisaitiere ayes 20 
Territory, and relations of hornbills with one another.................. 24 
eT ALINAS AUTEM ORIEL: DINGS a's erase oases cteia eis as winaiaeloms eaten e Misia pals wdboreled 29 
WES vO Rupes arera ta ee tayo: teh ciety otencvc) sy ahene arate tos aie red ceteite soe nfanst overalay shore are obalade « dcece craven 29 
Some anatomical features in relation to function.................e0000: 32 
Discussion of factors controlling hornbill populations.................... 34 
Engiparative studies. of other Hornbill. <6 s/5asce ecco ccc. cists cesac sees 35 
DHSCHESIOM OL HOLIDIL DIOLORY.. ae tiv icicles oaleeictelsiere «os viele di eieis vin niereviote wa at 40 
SSPE 17 os ols, ol giarare's) 6 weiss Siaiaiha@vlaeooe wei: divas Poldateiets saeiatecla ac dass 44 
MSPECRGEGY rote ciied btu o tre sip ais! Casd’ecats/ areialale eilshald sie:alesd Sehela Si ejas Aisa wie oak 45 





BREEDING, AND) OTHER. HABITS, OF ;CASOUED 
HORNBILLS (BYCANISTES SUBCYLINDRICUS) 


By LAWRENCE KILHAM 
Bethesda, Md. 
(WitTH 6 PLaAtTEs) 


INTRODUCTION 


Description of area.—Casqued hornbills were studied in an area 
extending from Entebbe, Uganda, to the Mpanga Research Forest 
located 13 miles to the west. This area is situated on the north flank 
of Lake Victoria. It is a few miles north of the Equator, at an alti- 
tude of approximately 3,750 feet. The rainfall is about 50 inches a 
year, with a peak in November and a peak of heavier rains in April. 
Temperatures vary little from a range of 60° to 85° F. Entebbe is 
the administrative center of British Government and is beautifully 
situated on a peninsula in Lake Victoria. Its extensive gardens and 
natural features make it attractive to an incredibly rich bird fauna. 
There are few trees, however, large enough for nesting hornbills. The 
Botanical Gardens have a small patch of forest where a single pair 
nested. 

Eastern Uganda consists principally of rolling hills covered with 
small native farms or shambas. Excellent climate and abundant rain- 
fall enable natives to raise crops continuously and in wide variety, 
including bananas, coffee, sugarcane, cotton, and cassava. Small 
fingers of forest persist along bays of the lake and swampy valleys. 
Zika Forest, 7 miles from Entebbe, is somewhat more extensive and 
consists of medium-sized trees. Most of these forest patches are 
under attacks from natives. Africans are continually collecting fire- 
wood or trying to enlarge their shambas. The Mpanga Research 
Forest was the only place I visited with any concentration of large 
trees furnishing suitable nesting sites. In a sense it is an island, pre- 
served from encroachment of the ever increasing native shambas. 
The forest is largely second growth. Some of its trees, however, are 
150 feet in height. A network of well-kept trails enabled me to move 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 131, NO. 9 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


rapidly and quietly from one nest tree to another. Most of the nest 
trees I discovered were within a quarter mile of the headquarters 
clearing. There were undoubtedly more at a greater distance. The 
forest covers 1.75 square miles and is 24 miles long. An African 
ranger and his assistants who lived at the headquarters clearing (pl. 1, 
fig. 1) were most helpful in putting up ladders and erecting scaffolds. 

Life in Mpanga Forest—A remarkable feature of the tropical 
forest was that during many hours I saw and heard few living things, 
whether I was sitting or moving quietly about. Throughout the day, 
especially in the first part of the nesting season, casqued hornbills were 
noisy and conspicuous. Other wildlife activity reached a crescendo 
early in the morning and again late in the afternoon, set off by the 
screaming of gray parrots (Psittacus erithacus). Great blue turacos 
(Corythaeola cristata) often came to feed on fruit of the same trees 
as the casqued hornbills. Their rolling calls were tremendous in vol- 
ume. On the other hand black-billed turacos (Tauraco schiittw), run- 
ning squirrel-like along high branches, were quiet and difficult to find. 
Two other, smaller hornbills (Tockus alboterminatus and Tockus 
fasciatus) occasionally came through the forest in small groups. I 
never saw or heard an owl at Mpanga. Hawks and eagles were not 
frequent, but they raised a commotion among hornbills whenever they 
appeared. Most magnificent was the crowned hawk eagle (Stepha- 
noaétus coronatus). The harrier hawk (Polyboroides typus), some- 
what vulturine in appearance, would search crevices and holes of dead 
trees for birds’ nests and other prey. Lastly, I encountered the great 
sparrow hawk (Accipiter melanoleucus) for some weeks in the horn- 
bill area. It made a continual shrill call, “ker-kee-kee.” I had a strong 
suspicion, but could not prove, that this powerful bird sometimes 
preyed on casqued hornbills. Smaller birds were rather retiring. It 
usually took some searching to see such birds as the West African 
nicator (Nicator chloris), the yellowbill (Ceuthmochares aereus), 
and Narina’s trogon (Apaloderma narina). Among mammals, troops 
of redtail monkeys (Cercopithecus ascanius schmidti) were much in 
evidence at the extremes of the day. 


METHODS 


Finding nests —Knowledge of hornbill habits facilitated the finding 
of nests. The various ways in which 16 nests were located, with the 
number of nests discovered by each method, may be summarized as 
follows: Search for the largest tree in an area where hornbills were 
suspected of nesting (5 nests) ; chance observation of a male at the 


NO. 9 CASQUED HORNBILLS—KILHAM 3 


nest hole (3 nests) ; hearing the feeding chuckle of the male and fol- 
lowing it through the forest (2 nests) ; rattle of female bill in nest 
opening (1 nest); shrill screaming of female hornbill from behind 
wall when her nest was approached by foreign hornbills (1 nest) ; 
commotion of a group of hornbills surrounding an eagle (1 nest). 
Two other methods involved observations of a male bird: When on 
a direct, purposeful flight into the forest (1 nest) ; and following a 
bird after it picked up dirt from the ground (1 nest). Finally, one 
nest located in the Botanical Gardens was first noted by other ob- 
servers. A helpful clue in finding an actual nest tree, once the general 
territory had been localized, was the presence of the elliptical stones 
of Canarium schweinfurthu on the ground. This fruit is a main item 
of hornbill diet. Feces were of little help as signs. They are mostly 
brown in color, disintegrate rapidly on vegetation, and are expelled 
away from the nest. 

Identification of individuals—Adult hornbills, as well as young 
emerging from the nest, have the same pattern of black and white 
plumage. Males, however, are readily distinguishable from females. 
As adults they are a third larger and have the huge, forward-projecting 
casque on the upper bill. Young males, even at time of nest leaving 
(pl. 1, fig. 2), have a larger bill than adult females. There is an ivory- 
white patch at the base of the upper mandible. As illustrated by my 
young hornbill (pl. 2, fig. 1), this patch is very large and is well 
supplied with blood vessels. It is probably an area of growth. In adult 
males it is smaller, but can be seen at a distance and, owing to varia- 
tions in size and configuration, it proved to be of considerable value 
in recognition of individual birds. It is unknown at what age a young 
male develops a forward projection of his casque. 

Watching hornbills—Observations on nesting hornbills were made 
from the ground close to nest trees, using 8 x 50 Zeiss binoculars. 
A few males were shy and rarely seen at the nest. Ordinarily, how- 
ever, males came to feed their mates if I sat still and waited. A blind 
was not essential. I usually watched from the spot which gave the 
best possible view. Along some trails in Mpanga Forest and in the 
Botanical Gardens the hornbills were accustomed to seeing people 
passing below. Hornbills were less shy when constructing nests, 
possibly because of preoccupation with the work. Many nests were 
inconveniently located. I therefore concentrated my watching on nests 
most favorably situated. Only one nest was located low enough for 
construction of a scaffold, reached by ladders tied in place. One 
could look inside by pointing a flashlight through the aperture. Ob- 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


servations were all made by myself except in two instances, both of 
which I was able to check to some extent on a subsequent occasion. 

Captive hornbills—Field studies have been supplemented with ob- 
servations on four young captive hornbills, three of which were re- 
moved from nests when approximately 6 to 7 weeks of age. A male 
(pl. 2, fig. 1) and a female (pl. 2, fig. 2), named “Mpanga” and 
“Zika” respectively, were both taken from nests and have lived in 
my house for a year. This paper does not present full observations 
on these captive birds. It is hoped to make a more complete study 
over a number of years. 


GENERAL HABITS OF HORNBILLS 


Flight —Casqued hornbills, with large bills and black and white 
plumage, were conspicuous birds in the vicinity of Entebbe, especially 
when they flew over open spaces. They were usually in pairs, the male 
flying about 20 feet in front of the female. Occasionally she took the 
lead. Their flight was remarkable. There would be a series of wing- 
beats, then a glide with head and bill held well up. These glides could 
be without apparent loss of altitude. If a bird was going downhill, 
as from our hilltop garden toward the lake, a glide might extend 200 
or 300 yards. Either phase of flight was noisy. The wingbeats made 
a “wush, wush” noise and the glide a prolonged “woo-oosh.”’ These 
noises were helpful, especially in Mpanga Forest, as they enabled me 
to know, even at some distance, when a male hornbill was returning to 
his nest. 

Roosting.—There were several opportunities within the Institute 
Compound for observing roosting habits. From August, when we 
arrived, until October, a pair of hornbills spent every night in a tree 
in our garden. They would come in with fair regularity at about 
6:50 p.m. and sit together for Io or 15 minutes in the dusk. Then 
they would separate to roost on individual perches. These perches 
were at the periphery of the tree where branches were about one 
inch in diameter. They were about 20 feet apart and 25 feet above 
the ground. The male always used his own perch and the female 
hers. In the period of perching together, either one of the two perches 
might be used. During my first nights at Entebbe, I was mystified by 
strange noises coming from the tree, not knowing that they came 
from hornbills. There would be an occasional “woof” or a whacking 
of bills on bark. By dawn at 6 a.m. noises increased, especially the 
bill whackings. The hornbills again perched side by side but were 
in no hurry to leave. They would finally move to other branches of 


NO. 9 CASQUED HORNBILLS—-KILHAM 5 


the tree, then fly off about 6:45 a.m. They thus spent nearly 12 hours 
roosting. When I returned from a short safari on October 19, I 
found that the pair had left. I presumed they had started to nest. 
Unfortunately, I had not discovered at this time that male hornbills 
can be identified by the white patch on the bill. I had spent many 
evenings watching the pair and wondered if they would return. Later 
I had some evidence that the male continued to roost alone in the 
Institute Compound. This evidence was most suggestive. On the 
evening of January 19 I heard a familiar bill whacking outside of 
my window. A male hornbill was roosting on the same male perch 
observed early in October. He spent only one night. I now recog- 
nized, by the white patch on his bill, that he was the same bird I had 
observed some weeks before roosting in an unusual place just beyond 
our garden. At night he was perched on a bare limb 20 feet over a 
driveway. On the nights of January 28 and 29, he was on the same 
perch, silhouetted against the sky. On the evening of January 31, 
this hornbill again came to our big tree, alighting first on the female 
perch, then settling on the male perch. Apparently the lone male 
alternated roosting places. Although territory among casqued horn- 
bills was not obvious much beyond the vicinity of the nest tree, it 
would appear possible that the area used for roosting might be more 
permanent. This situation would be worth further investigation. My 
captive hornbills, Mpanga and Zika, became extremely nervous at the 
approach of the evening. This was true even when in a room with 
artificial illumination. They appeared to have a strong desire to be 
on a roost they were used to and where they felt secure. Each bird 
perched on exactly the same place on the roosting pole night after 
night. 

Courtship behavior and the pair bond.—Casqued hornbills remain 
closely paired the year around, as far as I could determine. Some of 
their activities prior to nesting appeared to be courtship behavior. 
The pair of hornbills that roosted in our garden from August to Oc- 
tober could be readily observed. They would come flying in about 
dusk. Soon after they were perched side by side, the male would 
jerk his head and pop a cherry-sized fruit to his bill tip. Then he 
would bend over and try to feed the “cherry” to his mate. Usually 
she would not accept. This never discouraged his efforts. For ex- 
ample, on September 18 he hung his head and with a few slight heaves 
ejected a “cherry” from his gullet. He held it delicately in the tip 
of his huge bill as he offered it to his mate. She refused. The male 
then opened his bill with an upward toss which sent the “cherry” fly- 
ing back into his throat. In a few minutes he again produced the 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


fruit and again she refused to take it. She finally touched the “cherry” 
with her bill. The male then swallowed the fruit for the fourth and 
last time. Although this “cherry” presentation was observed on six 
evenings between September 25 and October 5, she accepted only 
once. On other evenings the ceremony climaxed in a touching of 
bills. There was never more than a single fruit involved. 

Presentation of a piece of bark, a stick, or a leaf was commonly 
observed during the nesting period. It was almost always the male 
who did the offering. On one day, November 20, I observed the re- 
verse process. A male perched in Mpanga Forest was joined by his 
mate who held a large winged insect in her bill. She gave it to him. 
He then gave it back and she swallowed it. Possibly she simply liked 
to have things handed over, even if she had to provide the objects 
herself. 

Mutual preening was another late evening activity of the two horn- 
bills in our garden. On September 15, the female sidled up to the male 
on his perch. Preening now went on for 20 minutes. The male nibbled 
at the feathers of his mate’s head and neck. She appeared to enjoy 
this attention. Her head moved slowly backward until the occiput 
rested on her back and her bill pointed upward. The male meanwhile 
kept nibbling at her exposed throat. Later it was his turn and she 
went over the feathers of his head and neck. He did not put his head 
way back as she had. I seldom saw males do this, once being on 
March 7 in the Botanical Gardens. A pair of hornbills were together 
for the first time in 4 months. The recently emerged female, in soiled 
plumage, was perched close to her mate as she nibbled at the feathers 
of his exposed throat. Mpanga and Zika, my captive hornbills, were 
preening each other at 3 months of age. Zika has always enjoyed hav- 
ing her throat tickled gently. Even when sitting in my lap, her head 
has gone way back in the manner of the wild bird which roosted in 
our garden. Mpanga has often invited preening. He does this by 
turning the back of his head to Zika, then ruffling up the feathers. 
In going over each other’s feathers, hornbills take special delight in 
finding small bits of horny material. They may stop to toss these 
about, small as they are. 

Playing with sticks and bill whacking.—Bill whacking was pre- 
dominantly a male activity. It was usually done after a male had fed 
his mate on the nest, but might take place on waking up at dawn or 
most any time of day. The huge bill would resound like a castanet 
as it was whacked from side to side on a limb. Females scrape their 
bills on a perch after feeding. 

A favorite occupation of resting male hornbills was to toss a stick 


NO. 9 CASQUED HORNBILLS—KILHAM 7 


in their bills, continually clamping on it to get a fresh grip. On No- 
vember 20 I saw one knock off a piece of bark and juggle it about 
until it dropped. He then fell straight from his perch toward the 
ground and retrieved the bark with surprising agility. On November 
29 a male, after much knocking on dead wood, finally broke off a 
piece 10 inches long. This fell toward the ground and the bird 
swooped down 50 feet but failed to catch it. My captive hornbills, 
male and female, would seize a stick or piece of crumpled paper with 
great gusto. Their clamping and tossing, however, would soon cause 
them to loose it. They were amazingly quick at catching any piece of 
food thrown at them, and when placed by a sunny window, they would 
try to seize bits of floating dust. 

Notes and calls.—In addition to noises made by wings in flight and 
whacking of bills, casqued hornbills made a din with their notes and 
calls. At times they sounded somewhat like domestic hens. A pair, 
perched in separate trees, would keep in touch with a series of hoarse 
“cuk, cuk’s.” At times they made single notes such as “ugh” or 
“woof.” Most lugubrious noises might come from a male in search 
of or temporarily separated from a mate. Thus on September 24 I 
saw a lone male and a nearby pair of hornbills. The single bird made 
“ka-ka-ka” and “ka-wack, ka-wack, ka-wack” noises of considerable 
volume. He broke off a small stick, and when he bounced along a 
limb with it in his bill, the pair flew away at his approach. The male 
of nest 1 made Mpanga Forest resound with his cries when his mate 
lost interest in nest building after weeks of futile effort. He was ap- 
parently trying to entice her back to the nest hole. Calls associated 
with nesting were of help in studying hornbills, for I could hear much 
farther than I could see in the forest. A feeding visit might be 
announced by a croak when a male came to perch near the nest. Then, 
when clinging to the entrance and transferring food, males usually 
made a low, rapid feeding chuckle as bills touched. Females sometimes 
responded with low guttural notes. These birds, walled within the 
nests, had a repertoire of their own. Two different noises were made 
with the bill. The first was a rapid, woodpecker-like tapping made 
with the tip of the bill, used broadside. Females tapped not only in 
constructing the original nest wall, but also when walled inside the 
tree. Sometimes they were repairing the nest wall, but at other times 
I conjectured that they were tapping on the sides of the hollow tree 
by way of idle amusement, for the persistence of tapping in some 
nests was greater than could be expected from repair work alone. My 
captive hornbills tapped in similar fashion on the walls of their cage 
when 7 and 8 months old. Two further noises of nesting females 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


expressed alarm. One was a rattling of the slightly opened bill in 
the aperture of the mud wall. The bill was visible from the ground. 
A second alarm noise was a loud, wailing screech. I never heard this 
call except from a nesting female in distress. Nest 15 was first lo- 
cated by following such screaming through 700 yards of forest. A 
foreign pair of hornbills was by the nest when I arrived. Such in- 
trusions were a frequent cause of screaming. Similar screams were 
heard from another female whose mate, early in the nesting season, 
fed her comparatively infrequently, and from another female whose 
mate had been recently killed. I seldom heard chirping of the young 
inside a nest. A newly hatched chick observed in nest 10 made a 
“chirpee, chirpee” note. Older young ones, as I learned from my 
captives, can make an assortment of chittery, whistling, and screaming 
noises. The whistling somewhat resembled that of a smaller species 
of hornbill, Tockus fasciatus. 


NEST CONSTRUCTION 


Hole hunting.—I first noted nesting activities of hornbills late on 
the afternoon of October 13. A male flew to the top of a high tree by 
the lake shore and peered into a hole. He was soon joined by his mate, 
who took her turn, looking into the hole for 10 minutes. Then she 
went inside and excavated pieces of rotten wood up to 8 inches long 
which she tossed out. The male hung his head down to watch what 
was going on. When his mate finally came out, he started to inspect 
a second hole nearby. He changed his mind, however, and flew away 
with a dismal wailing to alight by a hole in another tree. Here he 
called to his mate with a succession of “caks” and “ughs.” He put 
his head into the hole and pecked at the sides. Within a few minutes 
his mate joined him. It was apparent from this episode that the male 
pioneered exploration of possible nesting holes and enticed the female 
to follow. This observation was borne out by subsequent experience. 
On November 11, again late in the afternoon, I noticed a male hornbill 
perched next to a likely looking hole. He flew away and shortly re- 
turned with his mate. For the next 5 minutes she kept hanging her 
head down into the hole and pecking at the entrance. Then she lit on 
the lower rim, putting head and body inside. When she flew up by the 
male, he hopped down for another inspection. In Mpanga Forest, I 
had other examples of the lead taken by the male. Nest 1 had too 
large an opening for successful nesting. A pair of hornbills spent 
weeks trying to wall up the aperture, the male’s interest persisting 
longer than that of his mate. On November 7 J found the pair inspect- 


NO. 9 CASQUED HORNBILLS—KILHAM 9 


ing a hole in a stump 25 feet above ground. I suspected that they were 
trying to find an alternative to their other nest cavity. They both 
lit on the rim together, but he kept bending in to remove bits of rotten 
wood up to 2 inches long. She took a few of these from his bill, but 
let him continue the excavating. In January I observed further pio- 
neering by a male under unusual circumstances. The male of nest 5 
had been killed and his mate had broken out by January 2. I arrived 
shortly after 9 a.m. To my surprise, I saw a male picking up dirt in 
the forest clearing. I had seen no signs of nest construction for many 
weeks. The male’s flight led me to the abandoned nest. Here he 
perched by his mate, then clung to the lower rim of the hole and spent 
some minutes poking his bill about inside. His mate scrutinized the 
hole carefully before flying to it. She clung to the rim momentarily, 
but dropped away as if frightened. This desultory type of inspection 
went on for 3 weeks. After losing interest in the hole, presumably 
due to lateness of the season, the pair continued to use the tree as a 
perch. 

Location of nests ——Locations of nests are summarized in table 1. 
It was apparent that casqued hornbills preferred the largest trees and 
a hole as far from the ground as possible. Very large trees were 
scarce, even in Mpanga Forest. Nest 3 was in one of the finest trees 
(Antiaris toxicaria) in the forest, a huge specimen 6 feet in diameter 
at breast height and possibly 150 feet tall. The large branches sup- 
ported a growth of epiphytic plants and were draped with lianas. Text 
figure I is a sketch of the nest opening in a limb 85 feet above the 
ground and shows the male as he always perched preparatory to bend- 
ing over to feed his mate. Such large trees, free of limbs for 60 or 
70 feet, were impossible to climb. Nests were often located where a 
large limb had broken off, exposing an area of decay. Such a site is 
illustrated by text figure 2. Plate 3, figure 1, is a photograph of a large 
tree in the open, showing location of a nest entrance in the stub of a 
broken branch. Nest 5 was located in a huge arching limb of a 
Piptadenia, as shown in plate 3, figure 2. Some few nests were in less 
favorable sites. Nest 16, for example, was only 30 feet from the 
ground in a comparatively small tree. It was relatively easy for an 
African to climb up and open it. Three nest trees were isolated and 
in the open and the remainder were in forests. Nest 2 was in a unique 
position. The cavity was located in a crotch in the top of an exceed- 
ingly tall, dead stump. Since the entrance faced vertically upward, 
one wondered what might happen in a heavy rain storm. 

Gathering dirt for building at nest 1—I learned the most about wall 
building from a pair of hornbills in Mpanga Forest. Their hole was 


VOL. 131 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


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2. Young male hornbill 2 days after emerging from nest 5. 





1. “Mpanga,” hand-reared male hornbill 10 months of age. 





“Zika.’ hand-reared female hornbill 10 months of age. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131, NO. 9, PL 





1. Location (shown by arrow) of nest 13 in stub of broken limb. 





>, Nest 5 (location shown by arrows) in Piptadenia tree in 


Botanical Gardens, lentebbe. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE.°237, INO 9) Pia 





1. Termite mound at headquarters clearing, Mpanga Research Forest, 


where female hornbill collected earth. 





2. Entrance of nest 10, showing cement wall 


NO. 9 CASQUED HORNBILLS—-KILHAM il 


ideally located. It was 70 feet up in the trunk of a huge tree (Celtis 
saoyouxit), as illustrated by a sketch (text figure 2). Unfortunately 
the hole was about a foot in diameter and apparently too large. The 
sketch shows the small wall built across the lower portion. Its rate of 
construction was exceedingly slow, even though the female worked for 





N 
Fic. 1.—Nest 3, Mpanga Forest. 


many hours, usually in the morning. On October 24 the pair flew to 
the hole at 8:45 a.m. The female went inside and the male perched 
close by. He offered her a mud pellet 5 or 6 times, but she was occu- 
pied and paid no attention. At 9 a.m. the male flew to the forest 
clearing, then down to a path among the shambas. Here he picked 
two gobs of damp earth, swallowing the first and holding the second, 
an inch in diameter, in his bill. Then he flew back in stages to the 
nest tree. I watched him cough up and pass three small mud pellets 


12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


to his mate. She took them at her bill tip in rapid succession. The 
male then remained quietly by for some time while she continued 
working inside. Twenty minutes later the male again flew to the for- 
est clearing, alighting in a patch of maize. I crept up to within 30 feet 
and found him perched on a stump several feet above the ground. 





Fic. 2.—Nest 1, Mpanga Forest, never completed. 


He was bending over repeatedly, and I could see that he was picking 
up lumps of earth and swallowing them. To my surprise the female 
flew over. Both birds now perched together at the forest edge while 
he coughed up and transferred 5 or 6 pellets to the bill of his mate. 
Then he hopped away a few feet, broke off a 2-inch piece of bark, and 
bounced back to offer it to her. She refused it. When the pair had 
flown into the forest with loud squawking, I examined the stump. 
A termite nest clung to the side of it. Freshly opened tunnels, now 


NO. 9 CASQUED HORNBILLS—KILHAM 13 


lined by soldiers, showed where the hornbill had collected earth. On 
October 27 the male again visited the stump. At 12:30 noon he flew 
from his nest tree to the maize patch where I was able to observe him 
from a distance of 25 feet by using a screen of corn stalks. This time 
he picked up considerable earth from the stump and a little from the 
ground. Then he flew back to his nest hole, where he spent some time 
perched on the rim, moving his head about inside where his mate was 
working. Both birds were silent. So far it appeared that his job was 
to gather dirt and hers to build with it. However, at 8:20 a.m. on 
October 31 both birds flew to the forest clearing. She flew to the 
ground and hopped behind a mud-wattle hut. I moved around to see 
her bounce up against the wall and knock off a piece of dry mud the 
size of a plum, which she swallowed. She next hopped around a 
corner and knocked off another piece. From here she flew up to rejoin 
her mate. Shortly afterward she followed him to a distant pawpaw 
tree, where he fed her some fruit. Within 15 minutes they were back 
at the nest. By 9:10 a.m. the female was working on the nest wall, 
making a rapid tapping noise like a woodpecker. The male flew away 
for a short time, apparently to collect dirt. On his return, he perched 
beside his mate outside the hole, then gave her 11 pellets which she 
accepted at the tip of her bill and swallowed. She did not use them 
immediately, for the pair flew away from the nest hole and were gone 
for a half hour or more. 

These observations showed that female hornbills as well as males 
collect dirt, although they probably do so less often. The female from 
nest 1 did not always use pellets immediately after swallowing them. 
Dirt might be retained in her gullet for half an hour or more. The 
same was true of the male. Since earth was generally hard and dry 
when collected, the period of retention would give it a chance to be- 
come moistened by glandular secretions and by juices from such fruit 
aS paw-paws retained in the gullet at the same time. A larger part of 
the moistening may be done by the male. I noticed with my captive 
hornbills that the male could swallow more than twice as much as the 
female. He is, of course, a third larger in size. It was of interest that 
hornbills came to termite nests to gather building material, as these 
nests are exceedingly durable; each grain of earth is selected and 
coated with a cement substance by the termites. I saw the birds from 
nest I visit a termite nest on another occasion. The pair flew to the 
edge of the forest clearing at 9:08 a.m. on November 7. The female 
flew to a low tree, then over the shambas to the tall termite mound 
shown on plate 4, figure 1. Clinging to its steep sides, she took up 15 
chunks of dirt and swallowed them. Finally her mate flew over to 


14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


join her. Developments which took place in the next few minutes 
were unexpected. 

Coition——The pair had been nest building for at least 2 weeks 
when they perched together on the termite mound. There was a 
touching of bills, and the male acted as though he were trying to give 
her a few extra pellets, but he apparently had none to give. At 
9:15 a.m. the female flew from the termite nest, closely followed by 
the male who made a loud, wailing noise. The two headed toward 
the nest tree, but lighted on a branch at the forest edge. I had to run 
across the clearing for a better view. Within this short time the male 
had mounted the female. He came off a few moments later. Then he 
mounted on her back again, without hurry, and got securely placed. 
There was no noise. She had her tail held way over to one side. When 
he mounted he pressed his tail downward and somewhat under her 
body. His wings were kept closed. After some moments he got off 
and flew alone to the rim of the nest. She continued to hold her tail 
to one side. I could see against a background of white feathers that 
her vent was extroverted but was retracted shortly afterward. From 
at least 9:45 a.m. until 12:20 p.m. the female apparently worked from 
inside the nest on the wall, as she had on previous mornings. 

Gathering of dirt at other nests—At 5 p.m. on November 7 I was 
watching nest construction by a pair of hornbills in the Botanical 
Gardens. The male flew toward me and lighted on the ground 35 feet 
away. Here he picked up 15 or more chunks of earth. After he had 
flown away, I examined the spot which proved to be a low termite 
mound. Freshly opened tunnels showed where the bird had been 
working. Although hornbills appeared to be especially attracted to 
termite nests they are not invariably so. Sometimes they pick up 
ordinary dirt. On October 31 the male from nest 4 lighted on a stick 
close to the ground of a native shamba. He repeatedly bent over, took 
up chunks of black cultivated soil, and swallowed them. His next 
move was to an adjacent banana tree. Here he tore off pieces of leaf 
and bark, 1 to 2 inches long, three of which he swallowed. His subse- 
quent flight to the forest led to the discovery of his nest hole. The 
wall of this nest was unusually black. 

Construction of the nest wall—At various times I watched four 
pairs of hornbills constructing their nest walls. There was variation 
in the time of day when work took place. Three pairs worked in the 
morning and one pair, from the Botanical Gardens, late in the after- 
noon. Experience at nest 2 was typical. On October 24, the male 
returned to his tall, isolated nest stump at noon. He leaned over the 
nest opening as he heaved up pea-sized pellets of dirt. These he 


NO. 9 CASQUED HORNBILLS—KILHAM 15 


passed with his bill tip to that of his mate directly below. At least 
10 pellets were passed in rapid succession. The male then bounced 
along to another part of the stump and tried to whack off bits of dead 
wood. In a few moments he bounced back to the hole. He shook his 
head from side to side, with bill half open, as though a pellet had 
gotten stuck. Then for a while he held his head low over the hole to 
watch what was going on. A week later I found him carrying on 
much the same. He spent considerable time looking down and even 
putting his head and neck through the hole. Sometimes his mate 
accepted two pellets and refused a third. At such periods of active 
construction, the only sound was the rapid tapping of the female’s 
bill, which could be heard some distance away. 

Male hornbills did no work on the walls, although they might peck 
and explore about a nest entrance. A male usually sat by like a brick- 
layer’s helper. He would fetch building material and supply it to his 
mate as needed. The pair at nest 1 would spend the larger part of a 
morning in this fashion, settling down to work at about 8:30 a.m. 
Work at the Botanical Garden nest was done from midafternoon on. 
At this time the sun shone directly into the nest entrance. Possibly 
this pair was taking advantage of the illumination. I could see the 
female’s bill tapping inside. Details of how the tapping was done 
could not be perceived readily in high nests. I had a better opportunity 
at nest 7 which was comparatively low. The female was already 
walled in but on January 29 she was repairing the entrance with 
material from the nest floor. She applied this material with rapid 
tapping of the side of her bill tip. Tapping was again seen to good 
advantage in my captive pair of hornbills. In September 1955, when 
9 months of age, both birds would get in a corner of their cage and 
plaster the wall with any litter, feces, or food matter which might 
serve as mud. Both sexes did the tapping. This activity was carried 
on daily, as judged by the appearance of the wall and the amount of 
tapping we heard. 

Abortive nesting—On November 7, following coition, the pair of 
hornbills from nest 1 had worked all morning on the nest wall. By 
the following weekend their interest had obviously declined. The hole 
was too large to wall in. On the afternoon of November 13 the male 
kept returning to the nest rim. Then he would fly back to the forest 
to join his mate. Sometimes the pair would fly together to the nest 
tree, making a tremendous noise. It was a dismal wailing. The male, 
however, would look into the nest alone. On the following day I 
observed the same behavior from 7:15 a.m. until 12:30 p.m. These 
two were the noisiest hornbills in the forest. It appeared that he was 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


anxious to continue with the nest, but that she had lost interest, as 
during the whole morning she never came near it. He would put half 
of his body inside and make low grunts as he poked about with his 
bill. The same performance continued on a following weekend. She 
entered the nest several times, started tapping, then flew out again 
after a short time. The male persisted for two more weeks in his 
efforts to have her return. December 4 was the last time I ever saw 
her in the nest. This was 6 weeks after I had first found her at work. 
My final view of the pair was on December 12. They were perching 
near the nest cavity but showed no interest in it. 

Structure of wall and the inside of the nest.—Nest entrances were 
usually elongate slits, 14 to 2 inches wide and 4 to Io inches high, 
depending on the size of the natural cavity. These measurements 
are approximate. Plate 4, figure 2, a photograph of nest 10, shows a 
representative nest entrance. Walls were remarkably strong. Neither 
Africans nor myself, by putting a hand into the slit and pulling hard, 
could budge or loosen them in three nests opened for removal of 
young. Much hacking and prying with a curved bush knife were 
needed to effect an entrance. Another indication of the stoutness of 
walls was provided at the time of natural nest openings. One-half of 
the entrance cement of 2 nests (3 and 12) was knocked out entire, 
apparently by the emerging female. These pieces fell 60 to 85 feet 
to the ground, where I picked them up unbroken. They were roughly 
4 inches long, 3 inches in width and thickness, and were built in con- 
centric layers. Possibly each layer represented a day’s work. Odd 
bits of insects, bark, and plant material were incorporated in the 
cement in haphazard fashion. In two of the three pieces, one side 
was dark brown. It did not run with the concentric layers and it had 
faced the inside of the nest. Females had probably made this addition 
after being walled in. For lack of earth they had used fruit stones, 
seeds, and what appeared to be darker fecal material. It did not 
appear that feces was an integral part of the main cement structure 
in any of the five nest walls I examined closely. 

There was nothing organized about the inside of a nest. Nests 10 
and 14, examined at time of opening for removal of young, had frag- 
ments of rotten wood at the bottom. Nest 10 also contained many 
large contour feathers. When I pulled away the cement from this 
nest, hundreds of small ants swarmed out from behind the edges. 
My hand and arm were covered with them as I explored the cavity 
which was roughly a foot in diameter. An African who broke open 
nest 14 for me had a similar experience. These ants are presumably 
attracted by feces and other debris, for an amazing amount of fruit 


NO. 9 CASQUED HORNBILLS—KILHAM 17 


appears to go right through a hornbill’s intestinal tract undigested. 
This is especially true of young birds, as I discovered with my captive 
specimens. Feces from younger individuals does not always clear the 
entrance. This was observed from the scaffold by nest 10. There was, 
therefore, ample organic matter to attract ants, and their presence in 
great numbers may explain why I failed to find insects, in the nature 
of parasites, scavengers, or others, in examinations of nest debris. 


ACTIVITIES OF NESTING PAIRS 


Walling in of female.—As far as I can determine, I was watching 
nest 5 on the evening when the female began her 4 months of volun- 
tary confinement. On November 7, the male was active late in the 
afternoon bringing termite earth to his mate, and giving her pellets 
as she worked. On the following day I arrived at 5:15 p.m. to find 
him bending over the entrance. The sun shone into the hole. I could 
see that the nest wall was complete and that no more building was in 
progress. For the next hour the male stayed close by. At 6:30 p.m. 
he flew to a large tree (Canarium schweinfurthi) and picked some 
fruit, then returned to his nest with loud wailing. He gave his mate 
five or six fruits and she made low guttural notes. Dusk was now 
coming on. The male flew to a limb 15 feet away, then took a long 
flight which carried him out of sight over the brow of the hill. His 
mate remained alone. It was not until March 7 that I was to see her 
again. 

Feeding and other attentions of male to mate in nest—Once walled 
in the nest, the female was entirely dependent on the male for food 
until she emerged with her young one some months later. Nest 5 was 
well situated for observations. I would station myself below the huge 
Piptadenia tree in the Botanical Gardens (pl. 3, fig. 2) every few days 
during the week, before breakfast and again late in the afternoon. 
The male was accustomed to people passing beneath the arching limb 
containing the nest hole. Some days I would wait nearby an hour 
without seeing him approach the nest. I discovered, however, that 
there was some regularity to his visits, one usually taking place close 
to 7:30 am. Forty feeding visits were observed in the course of 
4 months. A visit on November 13 was typical. At 5:40 p.m. the 
male came flying over the open lawns, head held high, and wings 
making a “woo-ooshing” noise. He held a chip of wood 5 inches long 
in his beak. It looked as though he were flying with a cigar. He 
landed on the big limb, then bounced sideways until he was on the 
bole above the nest entrance. Then he leaned over and pushed the 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


stick of wood through the entrance. When his mate had taken it, he 
coughed up eight blackish fruits in succession, swinging his head down 
each time to place them in the tip of her bill. By the time the feeding 
was done, the stick had come out of the hole. He picked it up and 
pushed it in again. Then he flew to a perch 15 feet away. At such 
times he would usually whack his bill loudly back and forth on a limb 
as one would whet a knife. This male, like most of the others I 
observed, did not linger in the nest tree after a feeding visit. 

Stick or bark presentation was a common prelude to feeding. It 
occurred in 13 of the 40 visits observed. Objects presented ranged 
from curled pieces of bark, 1 by 6 inches, to smaller bits an inch in 
diameter. Discarded and dropped pieces accumulated on the ground 
below the nest. The male was persistent about these offerings. On 
December 8 he lighted above the nest and swung his bill down into 
the entrance 23 times to offer a piece of bark. His mate gave no re- 
sponse. When she accepted on the twenty-fourth try, he fed her four 
fruits. 

The male usually made a feeding chuckle when a fruit was trans- 
ferred. Numbers of fruits offered varied from 2 to 17 per visit, but 
counting was often difficult. Thus on February 7 the male fed his 
mate 17 small fruits. Some of these were offered 4 or 5 times before 
she accepted. On February 28 he fed her 2 “cherries.” A third one, 
however, had to be held down 11 times before she took it. Possibly the 
female is at times occupied with the young chick, so that she is not in 
a position to accept. Fruits brought to the nest ranged from the size 
of a pea to that of a small plum. 

In addition to bringing sticks and fruit, the male of nest 5 cleared 
away accumulations from the entrance. The approach to the nest, 
formed by a broken-off limb, sloped slightly upward from the wall. 
It thus collected fecal and other matter expelled from the nest. The 
male lingered to clear away this debris after 14 of 40 feeding visits. 
Sometimes he swallowed a few items. More often he would pick up 
small bits with his bill and toss them outward in a rapid and system- 
atic fashion. On February g he did this 30 times after one visit and 
on February 28, 25 times. Usually he made only a few tosses before 
flying away. Nest 7 was the only other nest where I observed a male 
clearing the entrance. 

Observations at other nests, while generally similar to those made 
in the Botanical Gardens, differed with the individual character of 
hornbill pairs. Sometimes physical peculiarities of the nest cavity led 
to differences of behavior. Nest 6 was in the straight trunk of a huge 
tree and the female often rested with her tail protruding from the 


130) 


VOL. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 





Female hornbill at time she was removed from nest 10, approximately 


I. 


period. 


oO 
d 


two-thirds of the way through nestin 





months of age. 


> 


when approximately 


“Zika”’ 


and 


a 


“Mpaneg 


> 





NO. 9 CASQUED HORNBILLS—KILHAM 19 


entrance. When the male bent over to offer food, she would not always 
bother to turn around. One day he gave her a fresh green leaf before 
coughing up four yellow fruits. At nest 12, also at Mpanga, the 
female would put her whole bill out of the entrance to take food. 
Possibly females in these last two nests were crowded for space. 
Some of the holes, such as those of nests 4 and 7, were on straight 
trunks without boles. Consequently, the visiting males had to cling 
to the lower rim of the nest with tail fanned out against the trunk. 
An occasional visiting male would bring bark to his mate but no food. 

Expulsion of feces, and other activities of female in nest—Watch- 
ing and listening from the ground gave some insight into activities 
of nesting females. At infrequent intervals one might see a stream of 
fecal matter shoot 2 to 3 feet out of an entrance hole, glisten in the 
sun, then land with a splash on the leaves below. The white feathers 
of the female’s rear end were, in some nests, clearly visible as she 
maneuvered her vent to the opening. Hornbill vents are protrusible 
and mobile. This could be well seen in our young birds 6 to 8 weeks 
of age when, standing way up on their legs, they would back up over 
the edge of the box they were in and expel feces on the floor. Expul- 
sion was not as forceful as seen in some hawks. Tree trunks and 
foliage below nests were not appreciably stained by expelled feces. 
This was partly due to the dark color of the droppings resulting from 
a fruit diet. Some streaking of white appeared in feces with develop- 
ment of the young. On January 8, as I was sitting on the scaffold be- 
side nest Io in Mpanga Forest, the 4-weeks-old chick backed to the en- 
trance and deposited a cylinder of feces 4 inches long on the lower 
cement. This feces had a white film over one end. The ladder leading 
to nest 10 became increasingly spattered with feces as weeks went on. 
Observations on my captive birds indicate that the white substance 
in the feces increased with ingestion of animal protein. I saw one 
nesting female toss debris out of the entrance with her bill. Doubtless 
this method also contributes to nest sanitation. 

Females within the nest did not lose their constructive instincts. 
Bill tapping continued, but was carried on far more by some females 
than others. I frequently heard tapping from nests 7 and 10, which 
were only 120 feet apart, at the same time. Much of the time I could 
not see a bill in the aperture. It is conceivable that these birds plaster 
debris against their nest cavities either because habit is strong or just 
by way of idle amusement. My young captive hornbills plastered the 
walls of their cage, possibly for the same reasons. Entrance walls, 
however, sometimes needed repair. On November 28, the female of 
nest 6 was repairing her nest entrance at noontime. I could see her 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


bill tapping rapidly on either side of the lower aperture. The shape 
of this aperture changed somewhat from one week to another and 
the repaired areas were darker in color. Presumably feces and other 
debris present by the opening were used, for I later obtained half of 
the cement from this nest and found that the dark areas had seeds and 
fruit stones embedded in it. 

Nesting females may enlarge their nest cavities by pecking at rotten 
wood surrounding them. The female of nest Io had an escape attic 
above her nest. I could hear her scuttling into it when I climbed up 
the ladder, and on looking through the opening all I could see was the 
tip of her tail. When nest 14 was opened on January 21, there was no 
female in sight. The African who had removed the chick swept the 
whole length of his arm inside without encountering the mother bird. 
His position was too precarious for him to look inside. It seemed 
probable that the mother had crawled into some remote recess. 


EGGS, YOUNG, AND NEST OPENINGS 


Eggs.—Nest 10 was in a dead tree 30 feet above the ground in 
Mpanga Forest. On December 4 I climbed the scaffold to this nest 
for the first time and peered through the aperture, using a flashlight. 
The mother bird was almost out of sight in her escape attic. There 
were two white eggs, similar to those of a domestic fowl in size and 
shape. My next visit was on December 11. The forest ranger said 
that he had climbed to the nest at 6 a.m. and had seen two eggs. I 
approached the nest tree at 1:30 p.m. and saw two-thirds of an egg- 
shell on the ground directly below the nest hole. The shell was so 
fresh that ants were still swarming over its moist inner surface. I 
climbed the scaffold to find the mother hornbill facing me at the 
entrance. This was the only time she ever did so. As far as I could 
determine before she climbed to her escape attic, she had a complete 
plumage. When she left I saw one egg and one blind, completely 
naked, rather blue young one. This was a first view of my subsequent 
pet, Mpanga. When I looked in on the following morning, I could 
not see him, but he soon emerged from under some debris, giving a 
feeble ““chirpee, chirpee.” His lower bill was larger than the upper one. 
Early in the morning of December 14 the ranger found the second 
egg chipped, and by afternoon he saw the shell on the ground and a 
second chick in the nest. I was able to visit the nest two days later 
and see the two chicks together. The larger one was chirping lustily. 
He had brown mash over his bill and throat, and there was more mash 
in the nest. I wondered whether the mother hornbill had regurgitated 
food onto the nest floor and then fed it to her offspring. 


NO. 9 CASQUED HORNBILLS—KILHAM 2I 


It was January 1, 1955, before I was able to visit the nest again. 
There was now a single young one, the size of a plucked pigeon, 
which begged and peeped a few times when I looked in. The forest 
ranger reported that the second chick had disappeared a week after 
hatching. The remaining chick had its eyes open. They were dull but 
mobile. Pinfeathers were just beginning to emerge on its head and 
wings. The entrance hole was becoming stained with feces, whereas 
it had been clean previously. On January 8, the young bird had black 
pinfeathers one-fourth of an inch long on head, neck, and wings. 
There were smaller, colorless pinfeathers on back, tail, and in two 
tracks bordering the breast bone. Feet and an inch-long fleshy tail ap- 
peared large for the size of the bird. The chick seized my finger when 
I pushed it in. He also chewed pieces of wood. This bird was removed 
from its nest when 6 weeks old and has lived well in captivity for 
over a year. 

Periodic inspections of the ground below nest trees gave clues as 
to the number of eggs and approximate time of hatching in four other 
nests. For example, on November 27 I found pieces amounting to 
two-thirds of an eggshell, with its membranes, below nest 4. I care- 
fully removed all pieces, and on November 28 there were no further 
eggshells. Six days later, however, I found a second eggshell, three- 
fifths intact, with an additional one-fifth in pieces. It appeared that 
two eggs had been laid and that they had probably hatched on different 
days. This had happened at nest 10. There, it may be recalled, the 
eggshell was tossed from the nest soon after the hatching of the chick. 
On the ground below nest 9, I removed most of an eggshell on Novem- 
ber 28, and three-fourths of a shell on December 4. These and other 
data are tabulated in table 1. Data from two other nests were less 
complete. I found over half of an eggshell below nest 3 on November 
28, but it was not until January 1 that a second shell turned up. Pos- 
sibly it had been covered with debris, either in the nest or on the forest 
floor. On December 9 there were pieces of one shell below nest 12. 
I did not, as in the other cases, know how long they had been there. 

Breaking open nests to obtain young. Molting of female.—Inacces- 
sibility made it difficult to study the molt in nesting females, but I was 
able to gather some information. Nest 10 could be reached by ladders. 
The female was usually in her escape attic and I did not want to 
interfere with her by making an opening. The nest was well lined by 
remiges when I looked in on December 4. In retrospect I should have 
removed them with a pair of long forceps for arrangement and count- 
ing. All I could see of the mother bird was her tail. The tail feathers 
remained soiled with no evidence of renewal. On December 11 I had 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


my only full view of the mother when she briefly defended her chick. 
Her plumage appeared complete, but I did not see her outstretched 
wings. My next view of her was when I broke open the nest on 
January 22. After putting the 6-weeks-old chick in a bag, I reached 
into the hollow trunk and pulled the mother bird down. She was kept 
in captivity for a few days of observation. Plate 5, figure 1, shows 
that her plumage was complete. The only sign of molting was one tail 
feather, a few inches long, which was still enclosed in a sheath. She 
was not shy in captivity, but she remained motionless, as if stunned, 
and refused to eat. I liberated her on January 24. In spite of a long 
period of confinement in the nest and having had no food for 2 days, 
she flew to a tree, squawked a few times, then took a flight of 300 to 
4oo yards. She was headed back in the direction of Mpanga Forest, 
13 miles away. 

It was apparent that this female, viewed when roughly one-third, 
and again at two-thirds through the nesting period, had not experi- 
enced any sudden or complete molt. On January 30, we opened nest 16 
to remove a chick 6 or 7 weeks of age. The mother bird struggled 
vigorously, striking the African who held her a sharp blow on the 
chin, so that he fell over backward. When I took hold of her it was 
obvious that she was in no weakened condition. Her plumage ap- 
peared to be complete except that her tail feathers, although well 
grown, had sheaths at the base. She flew readily to a tree when 
liberated. 

Premature escape of female due to loss of mate-——When I entered 
Mpanga Forest on the afternoon of January 1, I heard the wailing 
screech of a female hornbill in distress. The calls were given twice a 
minute. I followed them to nest 4 where I found a pair of foreign 
hornbills. These flew away at my approach. The female in the nest 
kept screeching for the next 2 hours in a most pitiful manner, but 
her mate failed to return. I examined the ground below the nest tree 
and found that he had been killed. There were two large patches of 
feathers directly below his usual perch. These patches were 2} to 
4 feet across. One consisted principally of small body feathers and 
the other of large feathers from wings and tail. I suspected that the 
hornbill had been struck from his perch by some bird of prey, and, 
after falling directly to the ground, had been plucked on the spot. The 
female was still screeching when I left the forest late in the afternoon. 
On the following morning I reached the nest shortly after 9 a.m. The 
mud wall was partially broken out. A new pair was inspecting the 
nest and it was evident that the original female was no longer there. 

Natural nest openings.—I observed how nests were opened naturally 


NO. 9 CASQUED HORNBILLS—KILHAM 23 


in five nests with entrances visible from the ground. In each the 
cement had been knocked away from one side of the aperture. This 
left ample room for the mother and young to emerge. I was interested 
to find that the missing cement was lying in an intact piece on the 
ground below three of the five nests. These five nests (Nos. 3, 6, II, 
12, and 13, table 1) all opened between January 1 and February 5. 
Some of these may have been open for a week before I noticed them. 
Nest 6, however, was closed on the afternoon of January 22 but open 
by 9:15 a.m. on the following day. There were no hornbills in the 
vicinity. 

Emergence of mother and young.—On February 5 I noticed that 
the female of nest 7, which I had had under observation for 84 days, 
was still walled in. By the following morning she had left. I began 
to search the adjacent forest and was able to locate the pair 100 
or more feet from the nest tree. The female was recognizable 
by her soiled plumage, the white patches of which were muddy. The 
male, recognizable by his bill markings, sat close by her. For the next 
2 hours I hunted back and forth through uncut jungle. It was raining 
hard and I thought that if I could find the young, I could probably 
catch it if its plumage was water soaked. The parents expressed great 
alarm, coming down within 20 feet of my head. Unfortunately I could 
not find the young. I wondered if it had crawled into some hollow 
limb. 

I had better success in the Botanical Gardens. The female was 
walled in on November 8, 1954, and had emerged with her young one 
on March 7, 1955. She was confined for 119 days, with a possible 
error of 2 days. There were no signs of activity by the nest on the 
morning of March 7. Late in the afternoon, however, I found the 
male perched by his mate 50 yards from the nest tree. He made 
continuous noises. Several times he hung his head way back, allowing 
her to nibble the feathers of his throat. Her plumage was in poor con- 
dition. The white parts of her feathers were soiled, her tail rumpled, 
and the small feathers on the back of her head and neck were worn. 
There was no sign of the young one. At 7:15 a.m. on March 8 I 
located the pair by cries coming from a patch of forest. They were 
together in a tall tree, and a young hornbill was close by. His plumage 
was in fine condition, pure black and white, his tail nearly full length, 
and his upper bill had the large, light-colored growing patch of a male 
(pl. 1, fig. 2). He made squawks similar to those of my captive birds. 
Everything appeared well when I left. 

The tragedy that overtook the family during the morning may be 
reconstructed from the chance observations of another bird watcher, 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Mrs, Iris Darnton. At 10:30 a.m. Mrs. Darnton was attempting to 
photograph the parent hornbills where I had seen them earlier. The 
young one was perching on a low branch by a roadway. He flew with 
some difficulty to a higher perch. At this moment an intruding female 
hornbill attacked the young one and the two fell grappling to the 
ground. The parents made a great commotion. Their young one lay 
flat on the road, but soon flew onto the lawn, then into a low tree. 
After 5 p.m. I came to the gardens and found the family where last 
seen by Mrs. Darnton. The young bird was perched precariously near 
the top of a spindly tree and one foot hung limp and useless. He was 
using half-spread wings to maintain his perch. The male parent made 
a great noise when he saw me, but soon quieted down, hopped closer, 
and fed the young one four fruits. Ten minutes later he tried to feed 
him again, but without success. The mother bird did not attempt to 
feed the young one. She remained inactive. A foreign female hornbill 
stayed about 50 feet away. On the following morning I found the 
parents in the same area, but the young bird was not in the trees, so I 
searched the underbrush and found him on the ground. When placed 
on the lawn, he was unable to fly. The male parent swooped repeatedly 
at my head. I was reluctant to take away the young bird, but it was 
obvious that any passing dog or individual could kill it. I therefore 
took it home. Plate 1, figure 2, shows his appearance 2 days after 
leaving the nest. I estimated that he was Io to 11 weeks of age, using 
as a guide my captive male of known age. One of his feet was broken. 
When placed in a splint, it healed completely in 3 weeks. This bird 
was the only one of my four captives that did not become tame. 
Parental devotion.—No large hornbills had come to our garden 
regularly since departure of the roosting pair in October. However, 
from April 1 until May 15, when we left, a pair of hornbills came 
every day, often remaining for some time. I soon recognized the 
male. It was the one I had watched for 4 months in the Botanical 
Gardens, which were 2 miles away. The parent hornbills had located 
their young in his outdoor cage, and our garden resounded off and on 
all day with their wailing and commotion. They perched on adjacent 
trees, and frequently swooped down low over the wire. “Mutesa,” 
as we called the young one, never responded in any perceptible way. 


TERRITORY, AND RELATIONS OF HORNBILLS WITH 
ONE ANOTHER 


Specific interference-—Nesting hornbills were interfered with by 
members of their own species to a surprising degree. Experiences at 
nest 5 illustrate the persistence of such interference. On November 6 


NO. 9 CASQUED HORNBILLS—KILHAM 25 


the pair of hornbills were working on their nest late in the afternoon. 
An adult male kept coming into the tree and the male in possession 
repeatedly drove him away. By November 8 the female was walled 
in, and a more serious attempt at interference was now made by a 
foreign female. I first noticed her on November 19. She was follow- 
ing the male and lighted in the nest tree when he lighted above his 
nest hole. On November 23 the same course of events took place, 
except that the male was less tolerant. He fed his own mate, then 
drove the intruder away. A week later I again saw her fly in close 
behind the male and light 25 feet from the nest hole. The male gave 
his mate a piece of bark followed by some fruit, and then bounced 
from one branch to another toward the foreign female. The intruder 
called and the female within the nest screamed a number of times. 
I wondered whether the interloper could seduce the male, but from 
subsequent observations it seemed unlikely that she would. The male 
returned again to the nest hole, and a few minutes later was in the 
upper part of the tree knocking about on dead branches until he dis- 
lodged a piece of bark. He clamped his bill on the bark until it was 
largely fragmented. Then he moved toward the foreign female. If he 
presented the bark, one would suppose that she had some attraction 
for him. After a moment, however, he changed his direction, flew 
down to the big limb below, bent over the nest hole, and gave the 
token to his mate, accompanied by a feeding chuckle. Subsequently 
he returned to perch quietly within 8 feet of the intruding female. 
At 7:30 a.m. the two of them flew away together. As the nesting 
season progressed, he became less tolerant of her intrusions. On my 
next visit, a week later, he made several swoops in an effort to drive 
her away, but she was not discouraged. On February 3 I again 
watched her fly in behind the male and alight in the nest tree, making 
considerable noise. The male stopped feeding his mate, swooped at 
the interloper and drove her down toward the ground. However, 
when he flew away, she followed a short distance behind. It generally 
appeared that her interest was in him rather than in the nest. On 
March 2 I observed a more serious situation. Late in the afternoon I 
found a foreign female clinging to the nest entrance. This time she 
was alone. She worked industriously, removing debris from the en- 
trance and knocking from the cement wall chunks which she broke 
in her bill before dropping them. There was no noise. After 5 minutes 
the male arrived and she flew a short distance away. He tossed some 
debris from the entrance, then drove the foreign female to another 
tree, flying at her so hard that he knocked leaves from intervening 
branches. He returned to his nest with a small stick held like a cigar. 


26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


His mate, who had remained silent, now began her wailing screeches. 
I also heard her bill tapping. The intruding female, persistent as 
usual, had followed the male back to the nest tree. In a few minutes 
he flew at her again, flying faster than hornbills usually do as he 
chased her from one tree to another. Five days later, mother and 
young emerged from the nest. As already related, a foreign female 
attacked the young bird and apparently broke its foot. After I had 
picked up the helpless young one on March 9g, I returned to the 
Botanical Gardens late in the afternoon. The pair of hornbills were 
perched side by side in their nest tree. Not long after I heard a great 
flutter of wings. I looked back to see both members of the pair pur- 
suing a foreign female. This was the last I saw of her. When the 
parents later came to our garden, she did not follow. I have presumed 
that the same foreign female was involved in all these incidents 
relating to nest 5. This presumption was based on her consistent 
behavior, general appearance, and bill shape. I never saw another 
female with which to compare her near the tree. 

At 5 p.m. on January 26 I witnessed an intrusion by a pair of 
hornbills. A foreign female was on the lower rim of the nest entrance, 
poking her bill about the aperture. She made no noise. After some 
minutes a foreign male lighted on a limb above. He had a fruit in his 
bill tip. The female moved toward him, took the fruit, and kept 
offering it down inside the hole. It was not accepted. The foreign 
female would toss the fruit about in her bill, then try again. Finally 
the rightful male returned, drove the intruding pair away, and fed 
his mate a number of fruits. The whole incident appeared odd. I 
wondered whether the foreign pair were unsuccessful nesters, who, 
having a strong, though thwarted instinct to feed something, dropped 
in on the female in nest 5. 

Interference by foreign hornbills was not limited to the nest in the 
Botanical Gardens. It happened not infrequently at other nests. A 
pair of hornbills were involved in each of the following intrusions. 
On November 19 a foreign pair were perched by nest 4 in Mpanga 
Forest. The female flew to the entrance, clinging to the lower rim 
with tail outspread for support. She then gave some hard pecks 
against the mud wall and grappled at bill point with the female inside 
the nest. Neither bird made any noise. However, when the intruder 
withdrew, the nest owner rattled her bill in the entrance. The foreign 
male sat quietly by without participating. In a period of Io minutes 
the intruding female attacked the nest entrance 12 times, but did no 
significant damage. In the next 5 minutes she attacked only twice. 
Then the rightful male returned and drove the trespassers away. 


NO. 9 CASQUED HORNBILLS—KILHAM 27 


Since this episode took place early in the nesting period, I conjectured 
that the foreign pair had, perhaps, not found a suitable nesting site 
and the female was trying to take possession of one already occupied. 
A second episode was difficult to interpret. It took place late in the 
nesting period, on January 23. I saw a foreign pair fly into the tree 
containing nest 15. The male repeatedly bent over the nest rim and 
there was a rattling of bills. He produced a “cherry” at his bill tip. 
Then he either gave it to the nesting female or dropped it into her 
nest. The female rattled her bill at the strange male. Ten minutes 
later the foreign female swung dramatically on a long tangle of 
epiphytic roots, then landed on the nest rim. This was the only time 
I ever saw a pair of foreign hornbills perched together on a nest. 
The intruding female waggled her bill vigorously in the opening. 
A few minutes later the owning male swooped in and drove the in- 
truders away. He fed his mate some yellow fruit. She now screamed 
repeatedly. 

On one occasion I saw a lone male attacking a nest. This was on 
November 21 at nest 6 in Mpanga Forest. The foreign male came 
quietly to a limb above the nest, then dropped to the nest rim. He 
appeared wary, bending his head to one side, then to the other, as he 
hung his head down to look through the entrance. The female had 
her bill ready but made no noise. He finally struck at the cement, then 
sparred with the female within the nest through the opening. I could 
hear their bills clashing. After 12 minutes the returning male owner 
drove the trespasser away. He had a leaf in his bill tip which he gave 
to his mate along with some fruit. It should be mentioned that a male 
may attack his own nest. At midmorning on November 28 the male 
from nest 7 flew down and rattled his bill in the opening. He was 
apparently in a bellicose mood for he next flew to a limb directly 
over my head, which was unusual. I could see his bill markings 
clearly. Meanwhile, his mate rattled her bill in the entrance. 

Lone females were the most frequent intruders at hornbill nests. 
I often saw one at nest 11. She had a favorite perch 10 feet from 
the entrance where she would sit for some periods. The nesting 
female would rattle her bill and scream repeatedly, but her mate, on 
feeding visits, paid little attention to the intruder. On December 12 
a foreign female perched calmly on the bole above nest 6. She repeat- 
edly leaned down into the opening, giving the feeding chuckle eight 
times as she did so. The female in the nest rattled her bill. After 
I5 minutes the male returned and drove the foreign female away. A 
final and most unusual case of interference occurred at nest 16, which 
was 30 feet above the ground. On January 30 we had placed some 


28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


ladders and an African was preparing to climb up and open the nest. 
As we stood below, a foreign female flew in and lighted on the edge 
of the nest. I thought that the mother had already emerged, but it 
was later discovered that the mother and young were inside. 

Territory —Neighboring pairs of hornbills tolerated each other 
well. For example, nests 7 and 10 in Mpanga Forest were within 
120 feet of each other and a third pair made persistent attempts to 
nest within an equal distance. Within this triangle I could watch 
activities of all three pairs at one time. They paid no attention to each 
other, their sense of territory apparently being limited to the nest tree 
and its immediate vicinity. Some trespassing was accidental and with- 
out interest in the nest itself. Thus nest 4 was centrally located in 
Mpanga Forest so hornbills engaged in their various activities fre- 
quently came near it. The male from this nest did more chasing than 
any other I had under observation. He had a peculiar habit of perch- 
ing during the day within 10 feet of his nest hole. No other male 
perched close to its nest. Some might occasionally perch within 100 
feet, but usually I saw males near their nest trees only on feeding 
visits. Females were frequently alone for 45 minutes to an hour at 
a time, and during these periods the nest tree was open to intruders. 
Hornbills may have a sense of territory in relation to roosting areas ; 
I did not make sufficient observations to determine whether this 
was so. 

Aggregations of hornbills —Sometimes a number of hornbills would 
come together, usually owing to a common attraction such as a flight 
of insects, a fruiting tree, or a passing hawk. This did not appear to 
be true flocking. Nine was the largest number of hornbills I ever saw 
together. They were in our garden on August 26. Other observers 
told me that they had seen larger gatherings. On November. 28 I 
watched two males and four females closely besetting a harrier hawk. 
A week later I came across what appeared to be the same group in 
the same area of Mpanga Forest. This time a band of redtail monkeys 
were working along the forest edge. Six female and two male horn- 
bills followed along with them sitting in the same trees but making 
no noise. I believe that the association was an idle one, for the horn- 
bills were playing with sticks and showed no alarm. The excess of 
females was of interest. Pairs of hornbills were also frequent during 
the nesting season. I wondered if there had not been enough nesting 
sites to go around. On February 15 I saw a curious association of 
two adult male hornbills. They came to our garden and hunted to- 
gether closely, going over cracks and crevices in a big tree. Two 


NO. 9 CASQUED HORNBILLS—KILHAM 29 


weeks later I found the same pair a mile away and still closely 
associated. 


RELATIONS WITH OTHER BIRDS 


Hornbills became much disturbed when a hawk or eagle appeared 
in Mpanga Forest. A crowned hawk eagle perched on a tall tree at 
the edge of the headquarters clearing. Then he sailed into the forest 
and was lost from view. An hour later I heard a great noise and 
found the hawk eagle surrounded by casqued and the smaller pied 
hornbills (Tockus fasciatus). None came closer than 20 feet. On 
November 28 I was watching nest 8 when I heard a number of horn- 
bills making short flights from one perch to another. This drew my 
attention to a harrier hawk in a dead tree. Two female hornbills were 
perched within a few feet of him, one on either side. Two males were 
in the same tree. When the hawk flew, all four hornbills followed 
him closely but made no noise. The bird that upset hornbills the most 
was a great sparrow hawk. On December 11 he flew up close to me 
in Mpanga Forest, calling “ker, kee, kee” in plaintive fashion. Three 
male hornbills accompanied him. None of them made any noise. 
Whenever the hawk circled and returned, the hornbills pursued closely 
and even swooped at him. On January 2 I again heard the cry of the 
great sparrow hawk. When he lighted above me, a male hornbill 
lighted within 6 feet of him, and when he flew, two hornbills followed 
within 20 feet. Hornbills are occasional predators themselves. Their 
presence, however, seldom caused any disturbance among smaller 
birds. I saw one hornbill momentarily beset by sunbirds and colies 
when he was robbing a nest of the latter. Broad-billed rollers 
(Eurystomus afer) would pursue hornbills passing by the lake shore. 
These aggressive birds attack everything from anhingas to starlings. 


FOOD 


Fruit—Food brought by male hornbills to their nests consisted 
largely of fruits, ranging in size from a pea to an olive. Some fruits, 
such as figs and pawpaw, were brought in as amorphous pieces. The 
elliptical fruits of Canariwm schweinfurthii were conspicuous and 
prevalence of their stones on the ground were a helpful clue to the 
location of nest trees. I was able to collect various seeds and fruit 
stones by cleaning the ground below nests. Following is a list of all 
fruits identified. Such indigestible matter passes through the digestive 
tract of the hornbills and is expelled with the feces. This was observed 
in both wild and captive birds. I have never seen hornbills go near 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


water and my captives do not seem to know what it is. Apparently 
they get enough water from fruit. 


FRUITS IDENTIFIED FROM SEEDS, STONES, AND PIPS RECOVERED FROM 
FECES BELOW HORNBILL NESTS 


Canarium schweinfurthii Engl. Antiaris toxicaria (Pers.) Lesch. 
Pycnanthus angolensis (Welw.) Exell. Chlorophora excelsa (Welw.) Benth. 
Pseudospondias microcarpa (A. Rich.) | Eugenia jambolana Lam. 

Engl. Dracaena steudneri Engl. 
Maesopsis eminii Engl. Ficus natalensis Hochst. 


Animal food.—Bannerman (1953), writing of Bycanistes subcylin- 
dricus, states that “this bird lives entirely upon fruit, as indeed do 
most of the large hornbills.” During initial observations I had no 
reason to doubt this statement. By closer watching, however, I found 
that hornbills take a wide variety of animal prey. On December 6 
a male went from one low perch to another among our garden trees, 
sometimes only 7 feet from the ground. Five minutes later I saw him 
fly up from the foot of a jacaranda with a 5-inch lizard squirming in 
his bill. He flew to a perch over a native shamba. Here he tossed 
the lizard about in his bill for some time, holding it by the tip of the 
tail, then chewed along until he reached the head. Finally the hornbill 
lost hold and the lizard fell. In a feat of acrobatics, the hornbill fell 
down after his prey, disappearing from sight in the vegetation. Fif- 
teen minutes later the bird was back in our garden. A completely 
limp lizard hung from his bill as he flew over the hill in what I 
suspected was the direction of his nest. On January 31 another male 
hornbill was hunting in our garden. He hopped onto a perch, looked 
around slowly in all directions, then hopped to another perch and did 
the same. After some minutes he flew to a thick bushy tree, where 
he scrutinized the foliage carefully, then hopped directly to the end 
of a branch where a mouse bird had its nest. The hornbill picked up 
a small egg with his bill tip, sent it flying back into his gullet with a 
toss of his head, then did the same with a second egg. To finish off, 
he seized some nest material and dropped it. What followed was an 
example of the delicate control casqued hornbills have with their bills. 
The male coughed up one egg from his gullet and held it again in his 
bill tip. By this time his mate had arrived in a tree across the lawn. 
He flew over to her, still holding the egg, and settling beside her, 
presented her with the egg; then heaved and presented her with the 
other, both intact. She swallowed both. On February 15 I watched 
two male hornbills hunting in our garden. A double-toothed barbet 
(Lybius bidentatus) was excavating in a tree when the hornbills flew 


NO. 9 CASQUED HORNBILLS—KILHAM 31 


directly to the hole. One male repeatedly drove his bill into the exca- 
vation. I was not sure whether the barbet was inside or not. For the 
next 15 minutes the hornbills quietly examined the hole, knocked off 
dead bark, and searched leaves and seed pods. Their hunting was not 
successful while in the garden. However, hornbills probably catch 
other birds on occasion. On January 29 a male in Mpanga Forest 
glided to his nest with a sparrow-sized bird, chewed beyond recogni- 
tion, hanging from his bill. He saw me and quickly flew away. My 
captive hornbills have been maintained to a large extent on left-over 
laboratory mice which they swallow whole. They appear to thrive on 
animal protein. 

Hornbills catch insects both large and small. I found the remains 
of some insects which they had fed upon by examining fecal matter 
below nest 5 in the Botanical Gardens. Dr. V. G. L. van Someren 
was able to identify the following: 


REMAINS OF INSECTS RECOVERED FROM FECES BELOW HORNBILL NEST 5 


Dicranorrhina micans (Drury) 

Longicornis beetles of cerambycid group 
Rhyparobia grandis (Sauss.)—large cockroach 
Long-horn cricket 

Tenebrionid beetle, Metallonotus 
Slender-winged mantis 


Some were not adequate for species identification. Large bark beetles 
were a frequent finding. Activities of a male hornbill observed on 
February 1 indicated how these might be captured. For a half hour 
in midmorning he worked on the dead portion of a large tree. He 
would strike slanting blows to loosen a piece of bark, then pry under 
it and appear to pick out something from underneath. The next step 
was to knock the loose bark to the ground. This hornbill showed 
acrobatic skill, often leaning way over on its long legs, with head and 
neck outstretched, in an effort to reach more bark. He removed at 
least 3 square feet before flying away. Praying-mantis nests were 
not infrequent in accumulations below nest 5. They had come through 
the digestive tract more or less intact, as indicated by adherent feces. 

I would not have supposed that casqued hornbills could catch small 
insects on the wing. They are, however, quite resourceful in this 
respect. On April 18 three hornbills were perched on a casuarina tree 
late in the afternoon. They were gulping at a close swarm of insects 
that were whirling about in a small cloud near the treetop. These 
insects were black-winged termites. Some were still moving upward 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


from the ground. The hornbills made a clapping noise as they snapped 
at the passing termites and were at the game for over 10 minutes, 
On May 1 I witnessed a similar spectacle, again late in the afternoon. 
Compact swarms of small insects (not lake flies) were hovering at 
the tops of tall trees adjacent to the Institute Compound. Smaller 
birds were catching them, mostly by perching on a topmost spray. 
These birds included splendid starlings (Lamprocolius splendidus), 
bulbuls (Pycnonotus tricolor), Abyssinian gonoleks (Lanarius eryth- 
rogaster), and didric cuckoo (Chrysococcyx caprius). Two casqued 
hornbills were catching insects along with the smaller birds. For over 
20 minutes they kept turning their heads to snap at the swarm around 
them. 

Dr. W. H. R. Lumsden has contributed an observation which fur- 
ther indicates the agility of these large hornbills. On September 6, 
1953, he was in the woods of the Botanical Gardens. Three or four 
hornbills were perched about 60 feet from the ground. They would 
swoop down across an open space, pick up something in midair, then 
rise to a perch in an opposite tree. They were after winged ants which 
were swarming on ground and vegetation below the clearing. 


SOME ANATOMICAL FEATURES IN RELATION TO FUNCTION 


Some peculiarities of hornbill anatomy came to have more signifi- 
cance with continued watching. The large eyes are unusually mobile 
for a bird. Casqued hornbills can look up and down to a moderate 
extent without cocking their heads as many birds do. This gives them, 
by human interpretation, a more intelligent expression. The upper 
and lower eyelids are continuous and in sleeping this fused eyelid is 
pulled over the eye from back to front. The combined eyelids are 
white in adult females. Considering that the eye is dark and sur- 
rounded by blackish feathers, I have wondered whether these white 
eyelids enable the male to see his mate better when looking into a 
dark nest cavity. The head is covered by fluffy feathers, 14 to 2 inches 
long. These are used in emotional expressions and when fully erect 
the head is like a small, round feather duster. From front view the 
topmost feathers, which may be the only ones erected, may resemble 
two horns. My young captive hornbills demonstrate how these 
feathers may be used. If I toss grapes to them, Mpanga may grab 
them all. Zika, the female, then feels left out. This is obvious by her 
expression. Her head feathers stand straight out in all directions as 
though to say “Where do I come in?” When alarmed or excited, her 
head feathers lie tightly back. If she next investigates some object, 


NO. 9 CASQUED HORNBILLS—KILHAM 33 


like a crumpled piece of paper, they stand out partially again. In 
young birds the feathers just above the eyes and forming the horns 
are brown. By the age of 10 months these are almost entirely replaced 
by black feathers. 

The bill tip can be used as delicately as a pair of forceps. In fe- 
males, only the tips may be in apposition, leaving a slightly open space 
for several inches behind. This space is more exaggerated in older 
females and may suggest, to a mild degree, the bill of an open-bill 
stork (Anastomus lamelligerus). At first I thought the space was due 
to wear. However, my captive Zika had this space at an early age 
when wear was not apparent. The bill tip itself is very sensitive. It 
is, for example, continually used to investigate strange objects. If I 
give my captive hornbills a fruit they have not seen before, they will 
toss and squeeze it in their bill tips for some time before swallowing. 
The bill of the male is huge, and that of a young one is larger than a 
female’s before he leaves the nest. This is shown in the photograph 
(pl. 5, fig. 2) of two young hornbills, male and female, at 7 to 8 
weeks of age. Plate 6, figure 1, shows Mpanga’s bill at 6 months of 
age and plate 2, figure 1, at 10 months. The white patch is apparently 
an area of growth. With some transillumination one can see that it 
is full of blood vessels. The patch becomes smaller in older males. 
No one knows, as far as I am aware, how long it takes the bill of the 
male to reach full development, with a forward projecting knob. At 
present I can only speculate on the function of this huge structure. 
It would appear to have no strictly practical use, for the smaller bill 
of the female serves more immediate functions. Her bill not only 
is used to plaster the wall of the nest, but is also a formidable weapon 
for defending the nest opening. Its inner capacity is no less than that 
of the male’s. In addition, males and females are equally adept at 
catching small objects with their bills. I wonder, therefore, if the 
casqued bill is not chiefly sexual in function. Possibly it is com- 
parable to the mustache stripe of the male flicker or the red breast 
of the cock robin. In this sense it would serve to release behavior 
patterns in the female which promote successful breeding and pair 
formation. | 

The feet of hornbills, with three front toes somewhat webbed, do 
not grasp tightly. I have never felt any real grip from the birds 
perching on my arm. Hornbills can, however, hang down from a 
perch, almost parrotlike, without losing their hold. The long tail is 
remarkable in that it can be neatly folded over the back. This adapta- 
tion is convenient for females walled within nest cavities, as is the 
fact that they are a third smaller than males. 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


DISCUSSION OF FACTORS CONTROLLING HORNBILL 
POPULATIONS 


Birds such as hornbills which rear a single young one are, one may 
suppose, relatively safe from enemies. This was probably true before 
the original rain forests had been cut. A female walled in a nest 70 
feet above the forest floor, in the trunk of a huge tree without lower 
branches, is in an almost impregnable position. Such trees, however, 
are now entirely absent over large areas. Natives’ shambas, elephant 
grass, and patches of second growth cover the countryside, leaving 
only thin fringes of large trees along lake shore and swamp. Eastern 
Uganda is good agricutural country and the native population is 
rapidly expanding. Interference by man’s activities is, I believe, the 
greatest factor limiting hornbill populations. Mpanga Research Forest 
remains as a needed refuge. Even here, observations suggested that 
suitable nesting sites were way below the demand. Some pairs of 
hornbills were nesting in unfavorable situations. For example, nests 
to and 16 (table 1) were only about 30 feet from the ground and 
were easy to reach. Also, I continually saw pairs of hornbills that were 
not nesting during the nesting period. Two pairs tried without suc- 
cess to build nests in unsuitable locations. When nest 4 suddenly 
became vacant owing to the death of the male, another pair of horn- 
bills took it over immediately. Some of the incidents of specific inter- 
ference already narrated indicate the degree of competition. The 
disastrous effects of forest destruction on casqued hornbills is well 
described by Capt. C. R. S. Pitman (1955, personal communication). 
He writes that “ever since I first went to Entebbe in 1925 forest 
destruction in the vicinity of Entebbe and Kisubi, and in fact all along 
the NTB-Kampala Road, has been on such an appalling scale, that 
annually large numbers of trees, with the best nesting sites, are being 
destroyed. Bycanistes therefore is constantly having to move farther 
and farther afield to find suitable nesting sites. When I first went to 
Entebbe there must have been at least two dozen Bycanistes nests 
within a 2-mile radius . . . but now good nest sites are fewer and far 
between and Bycanistes resorts to hollows, some readily accessible, 
which it would have ignored in the past.” 

Fortunately Africans in eastern Uganda do not molest birds to any 
extent. Ease of growing food and comparative prosperity probably 
puts less pressure on them to do so. But in Bwamba, where hornbills 
were considered fair prey, I continually came across Pygmies and 
other natives wandering about with slingshots and small bows and 
arrows. Under these conditions I found the birds more wary and 
difficult to observe than near Entebbe. 


NO. 9 CASQUED HORNBILLS—KILHAM 35 


COMPARATIVE STUDIES OF OTHER HORNBILLS 


Genus Tockus.—There were two other species of hornbills in the 
vicinity of Entebbe, the crowned hornbill (Tockus alboterminatus) 
and the pied hornbill (Tockus fasciatus). These two smaller horn- 
bills are somewhat similar in size and appearance. I could never dis- 
cover any basic difference in their habits. Their high piping cries, 
erratic type of flight with many rises and dips, and greater concentra- 
tion on insect food readily distinguished them from casqued hornbills. 
All three species occurred in the same stretches of open country and 
forest. 

On March 20, 1955, I noticed a crowned hornbill flying through 
Zika Forest. He lighted on a treetop, then suddenly dropped down- 
ward. Searching the area, I found a leaning tree with a bole, 4o feet 
above the ground, with a 2-inch hole in the center. There was almost 
no suggestion of a mud wall. I watched for 20 minutes. At one time 
white feathers closed the entrance as the female pushed her vent to 
the opening, and a stream of excreta shot out. The maneuver was 
the same as I had witnessed with Bycanistes. Later the male returned 
to perch on the bole and feed his mate a large insect (mantis?). He 
did not linger, the briefness of his visits apparently being due to the 
fact that he carried only one item in his bill tip ; there was no heaving 
up of fruits from the gullet such as characterized visits of male 
casqued hornbills to their nests. On March 25 an African, well 
trained at the Institute, climbed up and inspected the nest for me. 
There were three white eggs. The mother bird, when poked, backed to 
the rear of the cavity. Unfortunately, preparations for leaving Africa 
prevented an adequate study of these birds. I am indebted to Dr. 
Friedmann (1925) for the following account, hitherto unpublished, 
of the opening of a nest of this species in Kenya Colony. 


On April 7, at Taveta, some natives cut down a large tree in which there was 
a nest of this hornbill containing the adult female and two young birds. The nest 
was about 50 feet up in the tree and was in a large hole, the entrance of which 
measured roughly 10 inches in long diameter and 3 inches wide. This entrance 
was plastered up with dry mud, bird feces, and bits of bark all mixed together, 
leaving an opening about 2 inches long and 1 inch wide. As I picked away at 
the mud the adult female pecked at me with its bill, about an inch and a half of 
which could protrude through the opening. When finally I opened the nest and 
took out the birds I found that the two young birds were of different ages, the 
older of the two [pl. 6, fig. 2] being feathered on the back, wings, sides, legs, 
and crown, while the tail feathers were free of their sheaths for their distal thirds 
and the sheaths of the neck and breast feathers were beginning to burst. The other 
bird was less well feathered, the wings and flanks being the only parts really well 
covered. The tail feathers were about the same as in the older bird and the under 


36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


tail coverts in both were well developed. In both nestlings the bill showed no trace 
of a casque and was a yellowish-horn color; the feet were dark lead color; the 
skin light pinkish; the iris bluish gray. The tail folded up against the back so 
well as to look like a definite adaptation to living in crowded quarters. In fact 
it seemed to be muscularly easier for the birds to hold their tails up than to 
straighten them out [pl. 6, fig. 2]. One of the nestlings when put on the ground 
fell over forward on its head and breast and the tail remained sticking straight 
up in the air as though the bird were unable to drop it into what would be con- 
sidered the normal position. The adult female when about to enter a nest before 
egg-laying usually begins to molt and is for some time thereafter in quite a help- 
less condition. New feathers grow in while the eggs are incubating and the 
young growing to the flying stage. The female taken from the nest had all the 
new tail feathers well developed but all of them were still basally enclosed in 
their sheaths. The bird was still missing the outermost secondaries and innermost 
primaries but the other remiges were there, most of them more or less still in 
their sheaths basally. The bird could fly only very poorly and seemed dazzled by 
the light. Several times I let it go and each time it flew or rather half fluttered, 
half flopped through the air very laboriously for a short distance and stopped by 
smashing against a tree or the side of my tent. 


I had some evidence that pied hornbills also breed in March. From 
November on I had been observing a pair of casqued hornbills in the 
Botanical Gardens and had kept watch on a squirrel hole 50 feet up 
in a nearby tree. It was not until March 1 that I noticed a pair of 
pied hornbills showing any interest. At 7:30 a.m. a pair were preen- 
ing nearby. Between them they made Io visits to the hole, poking 
their bills inside. When a crowned hornbill appeared, they chased it 
away. The following day the pair were at the hole morning and eve- 
ning. On March 8 I saw them putting their bills into the hole and 
tossing out debris. I had no subsequent evidence that the pair nested. 
The hole may have been occupied by a squirrel which I had seen using 
it previously. Apparently smaller hornbills may compete with hole- 
nesting mammals. On February 18 I was driving near Kaboona, in the 
arid country of Karamoja, when I noticed a pair of Jackson’s horn- 
bills (Tockus jacksoni) catching insects and flying to a 2-inch hole 
in a dead tree. When I returned 4 days later the pair were still in- 
specting the nest hole. Thinking young hornbills might be in the tree, 
I cut it down. To my surprise, the cavity contained a mother bush 
baby (Galago senegalensis) with a mouse-sized young one. These 
limited observations may have interest because I could find no breeding 
dates for these three species of Tockus in eastern Uganda. 

The investigations of Gordon Ranger (1949-52) offer an opportunity 
to compare the habits of Bycanistes with those of Tockus in some 
detail. These investigations on African hornbills are the most com- 
plete known to me. They have extended over many years and concern 


NO. 9 CASQUED HORNBILLS—KILHAM 37 


another crowned hornbill (Tockus alboterminatus) which occurs in 
South Africa. Comparisons will be made first in regard to differences 
of behavior, then to points of similarity with Bycanistes. All observa- 
tions and quotations on the crowned hornbill are from Ranger’s 
publications. 

Differences in behavior between Bycanistes and Tockus.—(a) 
Crowned hornbills have a definite territory which is fairly extensive, 
is defended against trespassing hornbills, and is maintained year after 
year by the same pair which temporarily share it with the offspring 
of each season. I found little evidence that Bycanistes subcylindricus 
maintains a definite territory other than the immediate vicinity of the 
nest tree. 

(b) Crowned hornbills live more extensively on insects. This 
greater consumption of animal protein is reflected in their white 
excreta. In feeding his mate at the nest, the male carries the food, 
usually a single insect, at his bill tip. He does not load his gullet, then 
cough up fruits one at a time as do male casqued hornbills. Further- 
more, crowned hornbills make casts of indigestible seeds, pips, and 
hard parts of insects. Casqued hornbills, on the other hand, pass every- 
thing out in the feces—even large fruit stones, whole baby mice, and 
mantis nests. 

(c) There are a number of differences in the manner of plastering 
nest walls. Crowned hornbills make plaster of feces, finer soil from 
the floor of the nest, and insect remains. According to Ranger “the 
female does not swallow anything for the purpose of disgorging it 
in the form of plaster,’ and “the male plays no part whatever in 
plastering the nest hole.” Bycanistes collect soil and lumps of earth 
from the ground. Both sexes do this, but the male brings the most 
and is a kind of “bricklayer’s helper,” supplying his mate who does 
the actual plastering. 

(d) A distinction between Bycanistes and Tockus lies in the time 
of emergence of the female from the nest. Ranger (1955, personal 
communication) has extensive data on this subject. He has found that 
the female may emerge 62 to 74 days after being walled in. At this 
time the precocious young reseal the entrance. Both parents then 
feed the young which emerge 19 to 34 days later. Moreau (1936) 
has collected similar information in regard to Tockus deckeni and 
T. alboterminatus. 

Similarities in behavior between Bycanistes and Tockus.—A close 
relationship between the two genera of hornbills is indicated by simi- 
larities in their behavior patterns. Many of Ranger’s descriptions 
(1949-52) of the habits of Tockus alboterminatus australis apply 


38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL: I3t 


equally well to Bycanistes subcylindricus. I have quoted a number of 
these verbatim. 

(a) Food. The crowned hornbill’s handling of larger prey is the 
same as for Bycanistes. Speaking of a grasshopper, Ranger writes 
“the hornbill . . . subjects the creature to prolonged chewing and 
pulping between the mandibles before swallowing, turning it about 
and tossing it to secure new holds. . . .”” Chameleons and nestling 
birds are treated in the same manner. Among insects taken by 
crowned hornbills, those as diverse as winged termites, long-horned 
beetles (cerambycids) and, curiously enough, mantid egg cases were 
all fed on by Bycanistes. After feeding, cleaning “is performed by 
scraping and wiping the outside of the bill against a branch.” 

(b) Roosts. The crowned hornbill has roosting sites which are 
used in rotation. Each member of the family, however, has its own 
private perch. The two casqued hornbills in our garden always used 
the same individual perch each night. Like Bycanistes, the crowned 
hornbill is not an early riser. The birds stretch and make gruff utter- 
ances to each other for some time in the morning before leaving their 
roosts. 

(c) Play and agility. The following odd traits are also true of 
casqued hornbills. Ranger wrote that the crowned hornbill reveals 
“its dexterity when by diving it recovers an object that falls from its 
bill before it reaches the ground.” Speaking of a captive bird he 
writes that ““Conkie was adept at catching objects cast at her over 
intervals of many yards.” The bill whacking of male casqued horn- 
bills was a characteristic sound in the forest. Ranger wrote as follows 
of the crowned hornbill: “The meaning of the exaggerated scraping 
of the bill against a branch, indulged in more particularly by the male, 
has not become apparent.” 

(d) Basking. “The foliage bath is followed by basking, advantage 
being taken of the sun’s appearance in a clouded sky, but basking is 
independent in purpose. The body with wings extended is relaxed and 
spread limply upon branch or foliage, the head and neck upturned. 
Conkie assumed the most limp, lifeless, unbirdlike attitudes, neck 
curled with throat uppermost, eyes obscured by the relaxed third 
eyelid.” Such postures are the ones assumed by my pet hornbill, Zika. 
It is not a matter of drying her plumage but love of sunshine for its 
own sake. As soon as the sun comes from behind a cloud, whether 
she is indoors by a window or outside, Zika assumes the grotesque 
attitudes so well described by Ranger. 

(e) Courtship. In describing breeding habits of Bycanistes, I have 
included various activities under a heading of courtship and main- 


NO. 9 CASQUED HORNBILLS—KILHAM 39 


tenance of the pair bond. This is a matter of interpretation. Ranger 
uses other phraseology, but the activities he describes are similar. For 
example, he wrote of the following behavior as having taken place 
26 days before final entry. “Investigation of a knocking and rat- 
tling near Site I disclosed the hornbill pair, one striking its bill 
with vibratory rapidity against a branch. The side of the terminal 
part of the bill was used, and the point, vibrating, traveled around the 
surface of the branch till in turn the opposite side came into play 
. . . then the other bird .. . became enlivened and extending its 
bill performed the same rattling action.” Ranger believed this rattling 
was the same motion employed in plastering and made special note 
that both sexes were involved. I am not sure whether this perform- 
ance is entirely related to the onset of nesting in Bycanistes. My 
captives, Mpanga and Zika, do a good deal of tapping. They began 
when 9 to 10 months of age and sometimes do it against my clothes. 
Ranger has stated further that his crowned hornbills made increasing 
visits to the nest tree as the season progressed. Such flights were 
initiated by the male. A new behavior was noted 19 days before final 
entry into the nest when the male began to present food to the female. 
This was done anywhere, not necessarily near the nest tree. Finally, 
bark presentation was frequent among crowned hornbills. Ranger 
found that the female would take bark with ready interest from her 
mate, then bite it to pieces. 

(f{) Intruders. I have described intrusions on nesting casqued 
hornbills by members of their own species. Apparently a similar 
phenomenon takes place among crowned hornbills. Speaking of a 
feeding visit Ranger wrote “the male and a young intruder arrived, 
and this drew a single sharp cry from the female. . . . The male then 
delivered an item and resumed his chasing of the intruder.” This male 
subsequently delivered “13 items of food and bark, but all the time 
was worried by the young trespasser who followed him again and 
again to the nest. . . .”’ I was unable to tell whether the female in- 
truders I saw by Bycanistes nests were young birds or not. The male 
intruders were all adults. Ranger has also written of the nesting 
female rattling her bill in the entrance. He describes this “habit 
rattling” as useless activity. This was not true of casqued hornbills. 
Every time I saw a female rattling her bill there was some cause, 
such as presence of intruders, to evoke this alarm signal. 

(g) Plastering. Photographs of nest entrances presented by 
Ranger show that the cement walls look much the same as those con- 
structed by Bycanistes. The female crowned hornbill has the same 
technique of plastering. ‘Always the bill works rapidly in vibratory 


40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


fashion, the side of the end portion . . . beating against the surface, 
to which the moist dung is applied.” Ranger has also noted that the 
cement walls are built up in layers. 

Ranger has been fortunate in having many years in which to study 
crowned hornbills. If I had had at least a second year to study 
Bycanistes I should have been specially interested in finding out (1) 
whether these hornbills remain paired from one season to another ; 
and (2) whether the same pair returns to the same nest tree in suc- 
ceeding years. Both of these situations, true for the Tockus albotermi- 
natus, presumably hold for casqued hornbills. 

Ground hornbills—tI had only casual views of the huge ground 
hornbills in Karamoja and in Murchison Falls National Park. These 
form the third main group of hornbills that occur in British East 
Africa. The following unpublished account of Bucorvus cafer 
(Schlegel) is contributed by Dr. Friedmann. It is of interest from the 
point of view of comparative biology. 


This giant hornbill was seen in rather small numbers in the open bushveldt at 
Taveta, Kenya Colony, during March and April. The birds were usually seen 
walking around on the ground in loose groups of three to six individuals. They 
really walk, not hop. In East Africa they are protected as scavengers and 
are not molested by big-game hunters and settlers. Although they feed on the 
ground they sleep high up in tall trees and can fly remarkably well for their bulk. 
The original “take-off” seems to give them some little difficulty, but when once 
under way they fly more directly than do most hornbills, their heavy wings 
causing a very audible woof woof with every stroke. The call note is a deep 
boom boom, a rather hollow, and reverberating note. During the mating season 
the birds become more vociferous and call to each other with great frequency. 

The natives in Kenya Colony have a story to the effect that the female ground 
hornbill says, “boom boom, I’m going home; boom, I’m going home” and the 
male counters with, “you always say that; boom, you always say that; I’m tired 
of hearing it; go on home; boom boom.” It was, therefore, with considerable 
interest that I learned from Mr. Rudyerd Boulton that the natives in Angola have 
another interpretation of the calling of these birds. They say that the female 
says, “boom boom, I’m going home, I’m going home,’ while the male replies 
with, “you must not do that, you must hold up the corn.” 

Like all hornbills these birds feed by picking up bits of food with the bill, then 
tossing it in the air and catching it far down in the bill or even in the open 
mouth as it descends. 


DISCUSSION OF HORNBILL BIOLOGY 


An early impression at Entebbe was that many of the nonmigratory 
tropical birds, from hadadas (Hagedashia hagedash) to red-bellied 
shrikes (Laniarius erythrogaster), remained paired throughout the 
year. Casqued hornbills were usually encountered in pairs. They 
are presumably mated for life and one would like to know when pair 


NO. 9 CASQUED HORNBILLS—KILHAM 4I 


formation takes place. It may have no immediate relation to the breed- 
ing season. I had three young captives, hand-reared and approxi- 
mately of the same age, in a cage at Entebbe. Mpanga and Zika were 
definitely paired before they were 3 months of age. Zika would work 
through Mpanga’s throat feathers as he let his head fall back, then 
he would do the same for her. The other hornbill, and later a fourth, 
both males, led independent lives except for roosting. Unnatural 
conditions undoubtedly favored this early pairing. Young birds, 
however, do not necessarily pair up when confined together. This was 
shown by three magpies (Pica pica hudsonia), taken before they left 
the nest, which I kept in a large cage in the same manner as the horn- 
bills. They showed no inclination to pair. 

Maintenance of close pair formation demands mutual attentions. 
When casqued hornbills are perching in different trees, the members 
of a pair are almost always in communication with each other, some- 
times only with single notes such as “‘cak” or “ugh.” When together, 
mutual preening, in which the female may take the lead, is a common 
activity. This preening about the head and nibbling of feathers under 
the throat went on regardless of the time of year. I saw it going on 
at dusk in the pair which roosted in our garden in October and again 
with the pair in the Botanical Gardens, on the day the female emerged 
with her young one in March. It took place early in the life of 
Mpanga and Zika. 

I have interpreted as courtship, activities which bring a pair of 
hornbills into the rhythm needed for the close cooperation involved 
in nesting. The lead is taken by the male. He feeds his mate and 
presents her with sticks and pieces of bark. In addition he becomes 
noisier in his calls and wailing. Similar activities are not uncommon 
to the courtship of many groups of birds. The male hornbill also takes 
the lead in exploring possible nest holes. By his cries and wailings, 
and his flights back and forth, he tries to induce his mate to look at 
them. 

Stonor (1937) has given an interesting account of a pair of 
trumpeter hornbills (Bycanistes buccinator) which attempted to breed 
in the London Zoological Gardens. Courtship consisted principally 
of the male feeding the female. She would fly down to the feeding 
dish, then wait expectantly for him to feed her. Sometimes he would 
do so. At other times he would swallow the food himself. Then, as 
if stricken with remorse, when she would fly to a higher perch, he 
would at once follow to feed her a morsel. Stonor wrote of a “curious 
ceremony, wherein the female flew up from the ground with food in 
her beak which she passed to the male, who then returned it to her, 


42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


when she swallowed it.” I have previously described an almost identi- 
cal situation which took place in Mpanga Forest. 

Important differences in appearance of hornbills are located about 
the head, the region which can be seen best through a nest opening. 
Head feathers in birds of all ages express emotions. In young birds 
the feathers at the base of the upper mandible are brown instead of 
black. The huge, forward-projecting casque of the male is his chief 
sexual characteristic and white skin around the eye is a peculiarity of 
the female. Many African hornbills have brightly colored patches 
of skin and wattles about the head and neck. These, however, are en- 
tirely lacking in Bycanistes subcylindricus. One would like to know 
what part these bright colors may play in courtship performances. 

Coition in one pair of casqued hornbills took place without any 
special courtship other than some touching of bills. The pair were 
returning to their nest, after gathering termite earth. Moreau (1936) 
found that copulation took place in Bycanistes brevis just after the 
female had emerged from her morning’s work and about to days 
before the nest wall was complete. 

Good nesting sites are probably used annually. Pitman (personal 
communication, 1955) believed that the nest hole that I watched in 
the Botanical Gardens had been used in 1947 and in 1949. At nest 1 
the pair tried for weeks to close the opening. Interest, however, began 
to fall off a week after coition, a situation that paralleled one de- 
scribed by Moreau (1936) in Usambara. Pairs of Bycanistes brevis 
tried for 2 years to nest at one site without success. “In both years,” 
Moreau wrote, “building continued after copulation had taken place, 
and when work had ceased, both birds still showed a keen interest in 
the nest hole.” Failure at the nest in Mpanga Forest may have been 
due to the large size of the opening. Other factors could have been 
operative also. The pair, or perhaps only the female, for example, 
may have been young and inexperienced. It is difficult to follow 
Moreau’s hypothesis that in Usambara, failure to complete nests was 
due to the male’s running out of saliva. 

Casqued hornbills probably lay two eggs to insure that a single 
healthy chick is produced. The young bird becomes so large that the 
nest might be overcrowded if two chicks survived. Crowned hornbills 
have two to three young. The mother, however, leaves the nest some 
weeks ahead of time. This not only makes more room for the young 
but enables her to help in the feeding. 

The length of time a female is walled in a nest (119 days for nest 
5) does not appear unusual for a bird of hornbill size to lay eggs, 
incubate, and rear a young one. One can use Wahlberg’s eagle 


NO. 9 CASQUED HORNBILLS—KILHAM 43 


(Aquila wahlbergi) for comparison. It is approximately the size of a 
casqued hornbill and has been studied by Leslie H. Brown (1952) in 
Kenya Colony. He observed an incubation period of 46 days and a 
fledgling period of 72 days at a nest where a single eaglet was raised. 
This gave a total of 118 days. The total time is about the same as for 
the casqued hornbill, which, I have estimated, leaves the nest when 
10 to 11 weeks of age. I would have supposed that young hornbills 
would grow more slowly on a fruit diet—they had animal food only 
occasionally. However, as indicated by white matter in the feces, 
they may have had more animal protein, particularly in the form of 
smaller insects, than I realized. It was almost impossible to feed my 
young captive hornbills on fruit alone. The volume required was 
exhausting. We reduced the number of feedings, first by coating 
pieces of pawpaw with powdered milk, then by giving each bird six to 
eight half-grown mice a day. 

Intrusions of foreign hornbills on nesting pairs of their own species 
presented an interesting study. In a few instances the intruders came 
in pairs. There were many free pairs of hornbills throughout the 
nesting season and I wondered if these were not an index of an in- 
creasing shortage of suitable nest trees. Intrusions by single female 
hornbills were more difficult to explain. At one nest the same female 
apparently stayed around for months, and possibly attacked and 
crippled the young one soon after it left the nest. Several explana- 
tions suggest themselves. First, intruding females may have been 
offspring of the season before, unwilling to leave their parents or, 
second, they may have been unmated adults attracted by a seemingly 
lone male; possibly they fell into both categories. Some of them 
seemed to be more attracted to the nest itself and others to the male, 
coming and going with him as he made his feeding visits. I had an 
impression that there was an excess of females in the hornbill 
population. 

In conclusion, the pleasure of watching hornbills comes from their 
love of play, unexpected agility, clownishness, and seeming intelli- 
gence. This last quality is difficult to assess. The intelligence of the 
crow family is well recognized. I have kept tame blue jays, crows, 
and magpies and rate my captive hornbills on the same level. Both 
groups are playful and curious, examining new objects with interest. 
They have a wide range of vocal expression. The large eyes of horn- 
bills, together with expressive movements of head feathers, give an 
impression of intelligence which is hard to disregard. Whatever their 
mental capacity, however, it is difficult to see how these birds can 
adapt themselves to civilization, as they are destined to inhabit large 


44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


forest trees. My unusual opportunity to study their breeding and 
other habits in the vicinity of Entebbe has been most fortunate. 


SUMMARY 


Sixteen nesting pairs of casqued hornbills (Bycanistes subcylin- 
dricus) have been studied in the vicinity of Entebbe, Uganda. An 
unusual concentration of nests was found in the Mpanga Research 
Forest. 

In courtship the male presented his mate with food and bits of 
bark or sticks. Mutual preening and calls back and forth were im- 
portant in maintenance of the pair bond. 

The male did the pioneering in exploration of possible nest sites 
and tried to entice the female to them with wailing cries. Plastering 
was done by the female from the inside of the nest cavity, using a 
rapid tapping with the side of her bill tip. Both sexes flew to the 
ground to collect dirt for building. Most of this, however, was done 
by the male. He attended the female, furnishing her with pellets for 
construction of the nest wall. Termite earth was preferred for 
building. 

Two eggs are laid. Although both may hatch, only one chick is 
raised. 

Male hornbills feed their mates about every 30 to 60 minutes, 
heaving up fruits held in their gullets. Often a stick or piece of bark 
is presented first. 

Nest sanitation is accomplished by female and chick expelling feces 
through the entrance. The female may also toss debris out with her 
bill. If such debris collects outside of the entrance, it is regularly 
cleared away by the male. 

Ants which swarm in some hornbill nests may act as scavengers 
and keep down insect fauna. 

Females walled within nests can be extremely noisy if alarmed. 

Observations made at one-third and two-thirds through the nesting 
period indicate that females have a gradual molt. Two females re- 
moved when two-thirds through their nesting could fly readily. 

The total period a female was walled in one nest was I119+2 days. 
A young hornbill, captured two days after emerging, was in full 
plumage. 

The majority of casqued hornbills in eastern Uganda probably begin 
nesting in September and break out in January. 

A hornbill territory is confined largely to the vicinity of the nest 
tree. 


NO. 9 CASQUED HORNBILLS—KILHAM 45 


Occasionally a foreign pair of Bycanistes would visit a nesting 
female and attempt to feed her. 

Lone female hornbills interfered persistently with a number of 
nesting pairs. At one nest this interference had serious consequences. 

Male hornbills fed their mates largely on fruit but also caught in- 
sect and vertebrate prey. Lists are given of such fruits and insect 
remains as could be identified. 

Roosting habits are described for one pair before and for a lone 
male during the nesting season. 

Casqued hornbills were much alarmed by hawks and eagles. 

They prefer the largest of forest trees to nest in. Rapid destruction 
of forests in Uganda threatens the future of these birds. 

Three hornbills, removed from nests when 6 to 7 weeks of age, 
have been reared in captivity, largely on a diet of animal protein. The 
exact age of one captive was known. Two of them have remained 
closely paired from the age of 3 months. Bill tapping and plastering 
against the walls of their cage were done by the male and the female, 
beginning at 9 to 10 months of age. 

A comparison has been made of Bycanistes and Tockus. There are 
many points of similarity in the habits of the two genera. 


REFERENCES 


BANNERMAN, D. A. 

1953. The birds of West and Equatorial Africa. 2 vols., 1,526 pp., 144 figs., 

54 pls. 
Brown, Les.ie H. 

1952. On the biology of the large birds of prey of the Embu district, Kenya 

Colony. Ibis, vol. 94, pp. 577-620. 
MackwortH-Praep, Cyrit W., and Grant, C. H. B. 

1952. Birds of eastern and northeastern Africa. xxv + 836 pp., 53 col. pls., 

6 black-and-white pls., hundreds of text figs. and maps. 
Moreau, R. E. 

1936. The breeding biology of certain East African hornbills (Bucerotidae). 
Journ. East African and Uganda Nat. Hist. Soc., vol. 13, Nos. 1 
and 2, pp. 1-28. 

RANGER, G. 

1949a. Life of the crowned hornbill, Lophoceros suahelicus australis. Terri- 
torialism, family life, and breeding. Ostrich, vol. 10, pp. 54-65. 

1949b. Life of the crowned hornbill (Pt. II). Ostrich, vol. 20, pp. 152-167. 

1950. Life of the crowned hornbill (Pt. III). Ostrich, vol. 21, pp. 2-14. 

1951. Life of the crowned hornbill (Pt. IV). Ostrich, vol. 22, pp. 77-93. 

1952. Life of the crowned hornbill (Pt. V). Ostrich, vol. 23, pp. 26-36. 

Stonor, C. R. 

1937. On the attempted breeding of a pair of trumpeter hornbills (By- 
canistes buccinator) in the Gardens in 1936, together with some 
remarks on the physiology of the moult in the female. Proc. Zool. 
Soc. London, vol. 107A, pp. 89-95. 


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SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 10 


CRUSTACEAN METAMORPHOSES 


By 
R. E. SNODGRASS 


Collaborator of the Smithsonian Institution and of the 
U.S. Department of Agriculture 





(PusiicaTion 4260) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
OCTOBER 17, 1956 


THE LORD BALTIMORE PRESS, INC. 
BALTIMORE, MD., U. S. A. 


CONTENTS 


MER OCUCEION: o's .c\a/s aye, sieve store ssa icicicl alerts iebe ltrs ars sues 

ie Evolution o1 the*arthropodS2:/s.e. tees. ciere'e cies 
II. The nauplius and the metanauplius............. 
III. Examples of crustacean metamorphoses......... 
Bravichtopodal << jac,cceyesosia sehecise/etere estes eae 
Ostracoda. swe sslowis she ccusa leis aoe ise se Se 

CODED OA ichaevesteciss dt aeiorcioste amines oi aw oe 
CirKipediay Se Ps we 5 homer nears seraiorearetes 

PSOPOGAs Micra seteleiciarsiotererseieto seajetel a sie sia thale. ster 

IF tgp paAiSta CCD dive laraaroneyalereresval orate tel avovcuele- serie 
Wecanoda: yee.s ais teins wie oie Sera cisidislow os. ttoretelorers 
StOMALOPOGA: | sie cievsist cverusnasis dele iacrwievs aula! sii 

IV. Structure and evolution of arthropod appendages 
RELEKENCES). te cistarccoets cist oahetereis nial cletole oh werk stores 


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CRUSTACEAN METAMORPHOSES 
By R. E, SNODGRASS 


Collaborator of the Smithsonian Institution and of the 
U.S. Department of Agriculture 


INTRODUCTION 


The review of crustacean metamorphoses given in this paper con- 
tains little that will be new to carcinologists, except perhaps a few 
accompanying unorthodox ideas. The paper is written for students 
in general zoology and is recommended reading for entomologists, who 
commonly think of metamorphosis as a phenomenon pertaining par- 
ticularly to insects. It is true that the metamorphoses of insects and 
of crustaceans have no relation to each other, and have little in com- 
mon, but a preliminary discussion of both will help in arriving at a 
general understanding of the nature of metamorphosis as it occurs in 
the arthropods. 

The first treatise on metamorphosis was written by Ovid in about 
the year A.D. 7, but the metamorphoses that Ovid described were 
mostly the transformations of members of the human species into 
animals, trees, or rocks, willed by the ancient gods or goddesses in 
revenge against some mortal that had offended them. The meta- 
morphoses imposed on animals by nature are not punishments, ex- 
treme as they may be in some cases, but are beneficent changes of form 
to better accommodate the individuals of a species temporarily to a 
more advantageous way of living. The young butterfly, for example, 
transformed in the egg into a wormlike caterpillar, is not an elegant 
creature as are its parents, but from a practical standpoint the cater- 
pillar is perfectly adapted to its chief function, which is that of feeding. 

The metamorphoses of Crustacea differ essentially from those of 
insects in that they pertain to a much earlier stage of development. 
The young insect hatches from the egg usually with the definitive 
number of body segments. The insects are thus epimorphic; but if 
the young insect has taken on a metamorphosed form in its embryonic 
development, it appears on hatching as a creature quite different from 
its parents. Yet a caterpillar, for example, is actually a winged juvenile 
stage of the butterfly corresponding with the so-called nymphal stage 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 131, NO. 10 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


of a grasshopper. The principal difference between the two is that the 
wings of the young grasshopper develop externally, and those of the 
caterpillar grow internally within pockets of the skin beneath the 
cuticle. Likewise, a “legless” fly maggot has legs developing in 
pouches of the skin covered by the cuticle. A young grasshopper goes 
over directly into a mature grasshopper; the caterpillar, the maggot, 
and others of their kind, when full grown with plenty of food stored 
in their bodies, must undergo a second transformation in a pupal stage 
to be restored to the parental form. This is the usual course of 
metamorphosis among the insects. 

Most of the Crustacea, on the other hand, hatch at an early stage of 
embryonic development, though at varying periods of immaturity, 
when they have only a few body segments and corresponding ap- 
pendages. During their development after hatching they successively 
add new segments and appendages until the definitive number is 
attained. The majority of crustaceans are thus anamorphic in their 
manner of postembryonic growth, though a few are epimorphic. 

Anamorphosis involves a change of form during development, but 
it is merely a way of growing, common to crustaceans, diplopods, and 
some chilopods. It should not be confused with changes of form that 
have nothing to do with progressive development toward the adult; 
such changes constitute a true metamorphosis. The metamorphoses 
of Crustacea are changes of form that the growing animal may take 
on at successive stages of its anamorphic growth, including the sex- 
ually mature stage of many parasitic species. In such cases, meta- 
morphosis has been superposed on anamorphosis. As Gurney (1942) 
has said, “it may be assumed that development in the Crustacea was 
primitively a continuous process of growth and addition of somites 
and limbs, as we find it to be in some branchiopods, and that abrupt 
changes between successive moults leading to the origin of definable 
phases are secondary responses to changes in the habit of life of the 
larva and adult.” Gurney notes an apparent exception to this rule in 
the Euphausiacea and some Penaeidae, in which the larva and the 
adult lead much the same kind of life. The successive phases of de- 
velopment in these two groups, however, are mainly stages of ana- 
morphic growth; their only metamorphosis is the adaptation of the 
larval appendages for swimming. 

Insect larvae may undergo metamorphic changes of form during 
their growth, but with the insects this larval heteromorphosis, com- 
monly called “hypermetamorphosis,” affects the fully segmented young 
insect, and is therefore not comparable to the heteromorphic larval 
growth of most Crustacea. Some metamorphosed young insects trans- 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 3 


form directly into the adult, but most of them first undergo a recon- 
struction in a special, proimaginal pupal stage. Among the Crustacea 
there is no transformation stage strictly comparable to the insect pupa. 

True metamorphic forms are not recapitulations of phylogenetic 
stages in the evolution of a species. An insect larva, though often . 
wormlike in appearance, does not represent a worm stage in the an- 
cestry of insects. A caterpillar has a modern insect head and mouth 
parts, a well-developed tracheal system, and wings growing beneath 
its cuticle. No worm, ancestral or otherwise, was ever thus equipped. 
Among the Crustacea also most juvenile forms assumed during the 
larval growth are temporary adaptations to a changed mode of life 
and are not phylogenetic recapitulations. Yet, it is true that former 
ancestral characters discarded somewhere along the line of evolution 
may appear in the ontogeny of the individual, and it is often difficult 
to determine what phases of development are recapitulatory and what 
are metamorphic aberrations. The following hypothetical example 
will make clear the distinction between the two, and will lead to a 
practical definition of metamorphosis. 

If the eggs of birds regularly hatched into reptilelike creatures, 
which later transformed into feathered fowls, the change of form 
would literally be a metamorphosis ; but, since birds have been derived 
from reptilian ancestors, it might be specifically a case of phylogenetic 
recapitulation. On the other hand, if there issued from the bird’s egg 
a creature having no relation to anything in the avian line of adult 
evolution, but which still finally transformed into a bird, the change 
of form would be one of quite a different nature, and it is this kind 
of change that will be regarded as metamorphic in the following dis- 
cussions. As here defined, therefore, metamorphosis is a structural 
change at any time in the life history of an animal that may be re- 
garded as an aberration from the ancestral direct line of adult develop- 
ment which followed approximately the phylogenetic course of evolu- 
tion of the species. In this case metamorphosis may affect the embryo, 
the larva, or the adult. Simple development without metamorphic 
interpolations might then be termed orthomorphosts. 

In the higher Crustacea there is a tendency for hatching to take 
place at later and later stages of ontogeny, leaving a correspondingly 
lesser amount of development to be accomplished after the larva leaves 
the egg. Finally a condition is reached when body segmentation and ap- 
pendage formation are complete or almost so at hatching ; the animal 
then becomes epimorphic in its development. In an epimorphic arthro- 
pod, the embryonic development may proceed by the method of ana- 
morphosis, or the entire body may be first laid down as a germ band. 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


In the second case segmentation appears later, usually progressing 
from before backward, suggesting that it represents a former ana- 
morphic mode of segment formation in which the anterior segments 
are the oldest. Since anamorphic growth, either in the larva or the 
embryo, is characteristic of the annelid worms and recurs in so many 
of the arthropods, it was probably the primitive method of growth in 
the annulate animals. 

The most immature larval form among the arthropods is the crus- 
tacean nauplius. For practicable purposes early hatching must be 
given up by terrestrial animals, unless they go back to the water to 
lay their eggs, as do the land crabs, frogs, and toads. The anamorphic 
myriapods do not quit the egg until they have acquired the adult type 
of structure and are equipped with a sufficient number of legs for 
terrestrial locomotion. The completely epimorphic spiders and insects 
are best fitted to cope at once on hatching with the conditions of their 
environment, and they have become the most successful of the land 
arthropods. Though some insects lay their eggs in the water and the 
young are aquatic, they are simply terrestrial forms that have become 
secondarily adapted in the larval stage for life in the water; they 
hatch at the same stage as their relatives on land. 

The Crustacea are primarily aquatic animals; only a few have be- 
come adapted to a permanent life on land. The eggs of most species 
are laid in the water, and the newly hatched young must be capable 
of swimming; the adults can later adopt a bottom habitat if they ac- 
quire ambulatory legs. Considering the uniformity of the water 
environment of a swimming larva, there is relatively little inducement 
for a young aquatic animal to undergo adaptive metamorphoses. The 
metamorphoses of most crustacean larvae, therefore, are relatively 
simple as compared with those of insect larvae, which have a great 
diversity of habitats open to them. Parasitic crustaceans, however, 
are a conspicuous exception to this generalization. 

As a rule small animals in the water are eaten by larger animals, 
but the small creatures have one recourse against their possible preda- 
tors and that is to become parasitic on them. Parasites, however, 
have to be structurally adapted to a parasitic life, and consequently 
most parasites undergo metamorphic changes. Many of the smaller 
crustaceans have adopted parasitism, and the most extreme degrees 
of crustacean metamorphosis are found among such species, especially 
if the adults themselves remain parasites. Such adults in some cases 
have lost all resemblance to the ancestral forms of their race, even 
every mark of their crustacean ancestry. Moralists may cite the 
“degeneration” of such parasites as a warning of what parasitism may 


NOs /LO CRUSTACEAN METAMORPHOSES—SNODGRASS 5 


lead to, but actually parasites are highly specialized for the life they 
lead by a simplification of structure and the elimination of all un- 
necessary organs, which were indispensable to their free-living an- 
cestors. In fact, no sympathy need be wasted on “degenerate’’ para- 
sites; give them credit for having found a simple and easy way of 
living, though at the expense of another creature. They have discarded 
all useless equipment, and some of them have devised most ingenious 
ways of attacking the host. 

The control of metamorphosis by hormones has been extensively 
studied in insects, but apparently no comparable studies have been 
made on the role of hormones in the metamorphosis of crustaceans. 
It is well known that hormones are produced in the eyestalks of 
decapods, and the source of the eyestalk hormones has usually been 
referred to two organs known as the sinus gland and the X organ. 
However, from recent investigations (see Bliss and Welsh, 1952; 
Passano, 1953) it is now known that the so-called sinus “gland” is 
not a gland but a complex of the enlarged ends of nerve fibers pro- 
ceeding from the X organ and from numerous neurosecretory cells 
in the brain, in the ganglia of the optic lobe, and in the thoracic 
ganglia. The sinus “gland” is therefore a receiving and distributing 
center for various hormones. Functions that have been attributed to 
these hormones include the movement of pigment in the compound 
eye, regulation of chromatophore activity in the integument, control 
of moulting, and the rate of development of the ovaries. Knowles 
(1953) gives evidence that the chromatophores are activated also by 
neurosecretory cells in the region of the tritocerebral commissure and 
the postcommissural nerves. The control of moulting by lengthening 
the period between moults was attributed by Passano to the X organ, 
which is itself a neurosecretory tissue in the proximal ventral part of 
the medulla terminalis of the optic lobe. Removal of both sinus 
“glands” has no effect on moulting since the hormone can escape from 
the cut ends of the nerves. Panouse (1946) also, in a study of 
Leander, had claimed that the “sinus gland” produces a hormone that 
normally blocks the growth of tissues and thus causes a lengthening 
of the intermoult period and retards the maturing of the ovaries. 

From later work by Gabe (1953) and Echalier (1954), however, 
it now appears that moulting, at least in the Malacostraca, is controlled 
by a pair of ductless glands in the antenno-maxillary region. These 
glands, discovered by Gabe, are named by him the Y organs, and were 
demonstrated to be present in 58 malacostracan species, ranging from 
Nebalia to the decapods and stomatopods. In species in which the 
excretory gland is maxillary, the Y organs are in the antennal seg- 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


ment ; in those having antennal glands they lie in the second maxillary 
segment. Each gland is implanted on the epidermis by an enlarged 
base and is innervated from the suboesophageal ganglion ; in form it is 
conical, lenticular, or foliaceous. From their histological structure and 
changes during the intermoult period, Gabe shows that the Y organs 
are comparable to the thoracic endocrine glands of holometabolous 
insects, and he suggests that they have something to do with moulting. 
Following this suggestion, Echalier (1954) made experimental tests 
by removing the organs. He found that bilateral ablation of the 
glands, when not made too late after they had already discharged their 
secretion, resulted in a very great lengthening of the intermoult period, 
far in excess of the usual time between moults. Echalier, therefore, 
contends that the Y organs are crustacean endocrine glands for the 
control of moulting. That they do not disappear in the adult as do 
the thoracic glands of insects, Gabe points out, follows from the fact 
that the crustaceans continue to moult in the adult stage. 


I. EVOLUTION OF THE ARTHROPODS 


In any discussion of arthropod metamorphosis the question of re- 
capitulation always comes up in relation to the larval forms. If there 
is any ancestral recapitulation in ontogeny, it then becomes necessary 
to have at least a theoretical concept of the evolution of the arthropods 
and some idea of what ancestral forms they had that might be re- 
capitulated in the development of the individual. 

The evolutionary origin of the arthropods is hidden in remote Pre- 
Cambrian times, so probably we shall never know the facts from visual 
evidence. There is, however, ample evidence from a study of modern 
forms to indicate that the early progenitors of the arthropods were 
closely related to the progenitors of the annelid worms, and that these 
two groups of annulate animals had a common ancestor. The funda- 
mental characters preserved in the annelid—arthropod organization 
are: an elongate segmented body, an alimentary canal extending 
through the length of the body, a paired ventral nerve cord with seg- 
mental ganglia, a somatic musculature, and mesodermal coelomic sacs. 
We may therefore visualize the primitive annulate as a very simple, 
wormlike creature having these features. The mode of development 
was anamorphic, new segments being formed in a subterminal zone 
of growth. From this primitive segmented worm the annelids have 
been directly evolved with little addition other than the development 
of segmental groups of lateral bristles, which in the polychaetes have 
been carried outon movable lateral lobes of the segments, the so-called 
parapodia, that serve for swimming and burrowing. 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 7. 


By a different type of specialization for locomotion, members of 
another branch from the ancestral stock developed ventrolateral, 
lobelike outgrowths of the body segments, and thus became walking 
animals. These primitive legs eventually evolved into the jointed 
appendages of modern arthropods, the lobelike origin of which is still 
recapitulated in the embryo. At the lobopod stage of evolution 
(fig.1 A) the animals resembled a modern onychophoran, and are 





Fic. 1.—Theoretical evolutionary stages of the arthropods. 


A, a primitive lobopod, common ancestral form of the Onychophora and 
Arthropoda. B, a derived form with longer and slenderer legs. C, a primitive 
arthropod with sclerotized integument, jointed legs, and gill lobes on the coxae. 
D, a fairly generalized modern crustacean, Anaspides tasmaniae. 

rAnt, first antenna; 2A4nt, second antenna; Mxpd, maxilliped; Plpds, pleo- 
pods; Prpds, pereiopods; Tel, telson; Urpd, uropod; IJ-XV III, body segments. 


perhaps represented by such fossils as the Pre-Cambrian Xenusion 
and the Cambrian Aysheaia. The modern Onychophora are probably 
direct descendants from these early lobopods, and have structurally 
not progressed much beyond them. Others, however, acquired a 
sclerotization of the integument, which allowed the legs to become 
longer and slenderer (B), and finally jointed (C) for more efficient 
action in locomotion. These jointed-legged forms were the first true 
arthropods. The segmentation of the legs early took on a definite 
pattern, which has been preserved in both fossil and living arthropods, 
most of which retained the walking mode of locomotion, though 
some may also swim or fly. 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


From these early Pre-Cambrian arthropods (fig. 1 C) in which all 
the appendages were fully segmented ambulatory legs, the trilobites 
branched off by specialization of the body structure, but with no es- 
sential differentiation of the appendages. In the other derivative 
groups, however, the appendages took on different forms adapting 
them to various uses, but the number retained for walking is charac- 
teristic of the several modern arthropod groups. The myriapods use 
most of their postoral appendages for progression ; the Malacostraca 
(D) use five or more pairs for walking, except where some of these 
have been modified for grasping ; Limulus and the arachnids use four 
pairs, the insects three. That the ambulatory limbs, when limited in 
number, should in all cases be those of the middle part of the body, 
though not necessarily the same appendages, ‘follows from the me- 
chanical necessity of balance. The anterior appendages become sen- 
sory and gnathal in function; those of the abdomen have been modi- 
fied for various purposes, such as respiration, silk spinning, copulation, 
egg laying, or swimming. 

The modern arthropods comprise two distinct groups, the Chelicer- 
ata and the Mandibulata. In the chelicerates the first postoral ap- 
pendages are a pair of pincerlike chelicerae that serve for feeding, 
and the ancestors of this group were probably closely related to the 
ancestors of the trilobites. The principal feeding organs of the 
mandibulates are a pair of jaws, the mandibles, formed of the second 
postoral appendages. The Mandibulata, including the crustaceans, 
the myriapods, and the insects, are certainly a monophyletic group, 
but their origin and their interrelationships are obscure. 

Among the Crustacea the malacostracan type of organization (fig. 
1 D), in which the thoracic appendages are typically ambulatory and 
the abdominal appendages natatory, would appear to be more primitive 
than the entomostracan types because it more closely conforms with 
the structure of other arthropods, and could be more directly derived 
from that of a primitive walking arthropod (C). The entomostracan 
forms, therefore, have been secondarily reconstructed for a purely 
pelagic life by a readaptation of the thoracic appendages for swimming. 

If we accept the premise that the original arthropod (fig. 1 C) was 
a simple animal with jointed legs along the entire length of a uniformly 
segmented body, the crustaceans were derived from this common 
arthropod ancestor by specializations that established the generalized 
crustacean structure (D). Developmental recapitulation of adult 
crustacean structures, therefore, can go back only to the beginning 
of adult crustacean evolution. The embryo, however, starts its de- 
velopment from a single cell and the free larva completes development 


NO. I0 CRUSTACEAN METAMORPHOSES—SNODGRASS 9 


up to the adult. The embryo and the early larva, therefore, represent 
pre-crustacean stages of arthropod evolution. The embryo, however, 
must reproduce its parental form. Hence the crustacean characters 
appear at a very early stage of ontogeny, but the resulting embryonic 
or larval stages are not recapitulations of adult crustacean evolution. 
The crustacean characters are simply precociously imposed on the 
anamorphic stages of ontogeny. Finally, if the embryo is set free as a 
larva at an early stage of development, it must be structurally adapted 
to a free life, and in its subsequent growth other adaptations may 
be necessary. Thus it comes about that metamorphosis still further 
complicates the course of ontogeny. The life histories of parasitic 
larvae best demonstrate that larval forms are metamorphic adaptations 
to a way of living, since the nonparasitic adult ancestors of such spe- 
cies can hardly be supposed to have had the larval form. Where a 
specialized adult structure has arisen since the crustaceans became 
crustaceans, there may be a true recapitulation of an earlier adult 
form, as in the megalops of the crabs. A further discussion of the 
nature of larval forms will be given in connection with the life history 
of a penaeid (p. 54). 


II. THE NAUPLIUS AND THE METANAUPLIUS 


Since among the crustaceans the young hatch at different periods of 
development, the youngest larvae may have very diverse forms in the 
various orders, representing different ontogenetic stages according to 
the degree of development they undergo within the egg. The earliest 
hatched larval form is the nauplius, which is particularly characteristic 
of the Entomostraca, but occurs also in the Euphausiacea and Pe- 
naeidea among the Malacostraca. The nauplius is usually followed by 
a metanauplius, which is the first stage of postembryonic growth. 
From the metanauplius on, development may be merely a matter of 
regular anamorphic growth by the successive addition of new segments 
and appendages, but in many species the larva takes on different forms 
as it develops. These ontogenetic changes differ so much in the various 
orders that no general description can be given, hence a discussion of 
them will be left to the next section of this paper. Special attention, 
however, must be given to the nauplius and the metanauplius. 

The nauplius—The nauplius is a minute creature, highly variable 
in form in different species, but typically ovoid or pyriform in shape 
with the larger end anterior (fig. 2 A). It has a pair of uniramous an- 
tennules, or first antenna (1Ant), typically biramous second antennae 
(2Ant) and mandibles (/d), and a median eye of two or more parts. 


Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31T 


The antennae serve for locomotion. The internal organization includes 
usually an alimentary canal, a muscular system, a nervous system, and 
a pair of antennal excretory glands. The alimentary canal when fully 
developed consists of an endodermal mesenteron and an ectodermal 
stomodaeum and proctodaeum. The mouth is concealed above a large 
labrum; the anus is usually formed at a later stage. The nervous 
system includes three pairs of ganglia corresponding to the append- 
ages. Though there is no visible segmentation in the ectoderm, the 
presence of appendages and ganglia shows that the nauplius is at 





Fic. 2—Nauplius and metanauplius of Apus cancriformis Bosc (from Claus, 
1873). 

A, nauplius, with first antennae (7Ant), second antennae (2Ant), and man- 
dibles (Md), rudiments of teloblastic appendages seen through cuticle. B, 
metanauplius, appendages of teloblastic segments (thSegs) exposed after first 
moult. 


least a partly segmented stage of development. The region of the body 
behind the mandibles is that in which later the other segments will be 
formed, and their rudiments may be seen beneath the naupliar cuticle. 
When these segments are formed, however, they are generated by a 
different method from that which formed the anterior segments. 

The nauplius is derived from a very early stage of embryonic de- 
velopment, represented in species that hatch at a later period by a 
simple embryo with rudiments of three pairs of appendages. The 
embryo still in the egg at this stage is clearly more simple in its 
structure than is the nauplius. The nauplius, therefore, is not merely 
an early hatched embryo—it has undergone a metamorphosis before 
hatching to adapt it to a free life in the water. 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS Il 


Our chief interest in the nauplius is the question of its theoretical 
value in phylogeny. The nauplius has been likened to the polychaete 
trochophore, and has been regarded as representing a primitive an- 
cestral form of the Crustacea. The trochophore, however, is entirely 
unsegmented and does not have the internal organization of the 
nauplius. Later it becomes segmented by a direct division of the 
posterior part of its body into a few primary somites. Likewise the 
very young trilobite, known as a protaspis, at first shows no sign 
of segmentation, but it soon becomes marked by transverse grooves 
that divide it into a few primary segments corresponding with the 
segments in the prosoma of the adult. A similar early direct segmenta- 
tion occurs also in the ontogeny of the Xiphosurida. The nauplius, 
therefore, would appear to represent the same stage of primary seg- 
mentation in crustacean ontogeny, though metamerism has not yet 
affected the ectoderm. It is reasonable then to infer, as contended by 
Iwanoff (1928), that the first somites in both the annelids and the 
arthropods were formed directly in the previously unsegmented body 
of the animal. The later extension of the body took place by the 
teloblastic generation of secondary somites from a subterminal zone 
of growth. The annelid and arthropod ancestors did not diverge until 
this method of anamorphic growth was fully established. 

While the three larval forms discussed above do have a basic simi- 
larity of structure, which is primitive, it is evident that distinctive 
characters of more recent phylogenetic evolution have been impressed 
separately on each. The protaspis shows distinctly the definitive 
trilobite type of structure, the nauplius is clearly a crustacean, the 
trochophore is a young worm. The trochophore and the nauplius, 
moreover, are adapted in quite different ways for swimming at an 
early ontogenetic stage. The trochophore is not an adult ancestral 
form of the annelids, nor is the nauplius an ancestral form of the 
Crustacea. 

The metanauplius—The nauplius is the direct product of em- 
bryonic development. The further growth of the larva, or of the 
embryo if hatching occurs at a later stage, proceeds from a subterminal 
zone of growth, which becomes active before the naupliar cuticle is 
shed, so that rudiments of the new segments may be seen in the 
posterior part of the body of the nauplius (fig. 2A). In the meta- 
nauplius (B), which appears after the last ecdysis of the nauplius, 
the posterior part of the body is much lengthened; it is now distinctly 
segmented and bears the rudiments of several pairs of new append- 
ages. The postmandibular somites are the teloblastic segments 
(tbhSegs). 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Most studies that have been made on the larval development of 
Crustacea describe and picture the newly forming postnaupliar somites 
and appendages as they appear externally, without giving any in- 
formation as to how they are formed. A paper by Fransemeier 
(1939), however, describes the segment formation in the free-swim- 
ming metanauplius of the branchiopod Artemia salina, and papers 
by Sollaud (1923) and by Manton (1928, 1934) give details of the 
corresponding segmentation in the embryos of Leander, Hemimysis, 
and Nebalia. 

In the anterior part of the nauplius the embryonic ectoderm is 
already differentiated into the tissues derived from it, and the meso- 
derm has been formed from the embryonic mesoblasts. The ectoderm 
of the body region behind the mandibles, however, is still undiffer- 
entiated and there is here no mesoderm distinguishable at this stage. 
At the posterior end of the body of Artemia the ectoderm forms a 
circumanal fold, the cells of which are the ectodermal teloblasts that 
will form the ectoderm of the new segments. From the ectodermal 
teloblasts, according to Fransemeier, cells are given off into the in- 
terior of the body that become the mesodermal teloblasts, which will 
generate the secondary mesoderm. The naupliar mesoderm and the 
postnaupliar mesoderm of Artemia are thus distinct in their origin, 
though the formation of the second takes place 10 to 15 hours before 
the hatching of the nauplius. The teloblasts constitute the zone of 
growth, from which the new segments will be generated forward. 
The first segments formed from the teloblasts are said by Fransemeier 
to be those of the first and second maxillae. As other segments are 
generated the anus-bearing region is carried posteriorly as a permanent 
telson. The proliferation zone remains active until the last segment is 
formed, when it is fully exhausted. The alimentary canal apparently 
simply lengthens posteriorly, the proctodaeum having been formed 
in the nauplius. 

In the young naupliar embryo of the palemonine Leander, as de- 
scribed by Sollaud (1923), the postmandibular part of the body is 
a small anus-bearing lobe, or caudal papilla, which subsequently 
lengthens and projects free from the body in front of it and bends 
forward. A transverse row of large cells becomes differentiated in 
the ectoderm of the lobe before the anus, and later encircles the lobe. 
These cells are the ectodermal teloblasts. Below and a little before 
them is formed a corresponding ring of mesodermal teloblasts, which, 
according to Sollaud, are derived from the blastopore. The teloblasts 
generate the secondary segments in the usual manner, but in Leander, 
Sollaud says, the two maxillary segments are formed directly in the 


NO. I0 CRUSTACEAN METAMORPHOSES—SNODGRASS 13 


larval body and not from the teloblasts, the first teloblastic segment 
being that of the first maxillipeds. In the Pericarida, however, he 
says the boundary between the primary tissue and the secondary 
tissue is between the segment of the mandibles and that of the first 
maxillae. 

The accounts given by Manton (1928, 1934) of the embryonic 
process of secondary segmentation in Hemimysis and Nebalia are 
essentially the same as those of Sollaud for the embryo of Leander 
and of Fransemeier for the larva of Artemia, Manton agrees with 
Fransemeier that the teloblastic segments include both maxillary seg- 
ments. In Hemimysis, she says, the naupliar and postnaupliar meso- 
derms are at first some distance apart, but later the teloblastic ecto- 
derm and mesoderm extend forward as far as the first maxillary 
segment inclusive. The teloblasts of Nebalia are differentiated at the 
sides of the posterior blastoporic area, and the ectodermal teloblasts 
eventually form a complete circle around it. The mesodermal telo- 
blasts, according to Manton, in agreement with Sollaud, are formed 
from the mesendodermal mass at the blastopore; Fransemeier says 
they are proliferated from the ectodermal teloblasts. The ectodermal 
teloblasts, according to Manton, join the naupliar ectoderm between 
the mandibular and first maxillary segments, so that “all segments 
between the mandibular segment and the telson are formed by the 
teloblasts.” The rudiment of each segment arises from one transverse 
row of descendants from the original ectodermal and mesodermal 
teloblasts. When the last abdominal segment is completed the telo- 
blasts disappear in both Hemimysis and Nebalia. 

Since the teloblastic generation of secondary somites added to the 
primary segmented body of the young larva or embryo is characteris- 
tic of the annelid worms and recurs in many of the arthropods, it must 
have been a way of lengthening the body developed in the very primi- 
tive wormlike ancestors of the two groups. The annelids and the 
arthropods, therefore, did not diverge until this method of growth 
was well established. Elsewhere the writer (1938) has suggested that 
telogenesis may have originated as a means of increasing the repro- 
ductive function by distributing the germ cells from the zone of 
growth through a larger number of segments. 


III. EXAMPLES OF CRUSTACEAN METAMORPHOSES 


The metamorphoses of Crustacea are so diverse that in a brief re- 
view of the subject we can include only a few examples representative 
of some of the principal orders. Since crustaceans that hatch at an 
early stage of ontogeny go through anamorphic phases of development 


I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


by the successive addition of segments and appendages, many of their 
changes are merely those resulting from the anamorphic manner of 
growth. In nearly all cases, however, there is some degree of meta- 
morphosis superposed on the anamorphic stages, varying from a 
mere adaptation of the appendages for swimming to a total recon- 
struction of the animal for a parasitic way of life. The most striking 
examples of crustacean metamorphosis, therefore, occur in parasitic 
species. Among the Crustacea metamorphosis evidently has been de- 
veloped separately in each order, and often independently in different 
members of the same order. There is no type of metamorphosis 
characteristic of large groups of orders, as in the holometabolous 
orders of insects. Moreover, since crustacean metamorphosis affects 
the juvenile anamorphic stages, except where it is carried over into 
the adult, the metamorphosis of Crustacea has no relation to that of 
the epimorphic insects. A brief but interesting account of the life- 
history problems of crustacean larvae is given by Gurney (1926). 
For much assistance in preparing this section of the paper the writer 
is indebted to Dr. Fenner A. Chace, Jr., and his associates in the di- 
vision of marine invertebrates of the U. S. National Museum. 


BRANCHIOPODA 


The branchiopods undergo few changes during their larval de- 
velopment that are not the result of simple anamorphic growth by 
which the body and the appendages are completed and brought to the 
adult condition through successive instars. The branchiopods are 
thus of interest in showing a simple progressive development from 
nauplius to adult, which is accompanied, however, by a specialization 
of the postgnathal appendages for swimming. As an example we may 
take the life history of Branchinecta occidentalis Dodds as described 
by Heath (1924). 

The newly hatched larva of Branchinecta is a typical nauplius 
(fig. 3 A) with three pairs of appendages, a median simple eye, and 
a large labrum, but the oval, unsegmented posterior part of the body 
is more than usually constricted from the forepart. The large second 
antennae are the principal swimming organs. Between the nauplius 
and the second instar, or metanauplius (B), a very considerable 
change takes place. Lateral compound eyes are now conspicuous by 
their pigmentation. The posterior part of the body has greatly length- 
ened, and bears rudiments of maxillulae, maxillae, and six or seven 
following pairs of appendages. In the third instar (C) the post- 
maxillary appendages have lengthened and the more anterior pairs 


NO. 10 CRUSTACEAN METAMORPHOSES—SNODGRASS 15 


have taken on a leglike form; their mesal margins are indented, and 
each limb bears a conspicuous lobe, or flabellum, just proximal to an 
apical point. The body is more lengthened behind the appendages and 





Fic. 3.—Branchiopoda. Branchinecta occidentalis Dodds, developmental stages 
(from Heath, 1924) and a thoracic limb of the adult. 


A, newly hatched nauplius, length 0.4 mm. B, second instar. C, third instar. 
D, fifth instar. E, eighth instar, 29 mm. F, sixth left thoracic limb of adult 
male, with six endites (1-6) and a movable terminal lobe (Dactpd). G, head of 
adult male, anterior, with large second antennae (2Ant). 


shows lines of further segmentation. After two more moults, the 
larva in the fifth instar (D) reaches an average length of 1.6 mm. 
The second antennae are relatively much shortened, but the legs have 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


increased in length and are more distinctly indented on their mesal 
margins. The slender posterior part of the body bears rudiments of 
four new appendages, and its apex is split into a pair of small caudal 
lobes. The postmaxillary appendages continue to develop through 
the sixth and seventh instars until 11 pairs are present. In the eighth 
instar (E) they have lost their leglike form and have become broad 
flat phyllopodia with large flabella and slender apical lobes. At this 
stage, as the thoracic appendages take over the swimming function, 
the second antennae are much reduced in size and are directed for- 
ward. Heath enumerates 17 instars in the larval life of Branchinecta, 
but development beyond the eighth instar merely brings about refine- 
ments toward the adult structure. 

The larval stages of Artemia described by Heath (1924) are very 
similar to those of Branchinecta, as are those of Branchipus described 
by Oehmichen (1921). In the Concostraca and Cladocera the larval 
development is complicated by the formation of a bivalved shell. 

The development of the branchiopod appendages is of interest be- 
cause it suggests that the natatory phyllopodium has been evolved 
from a segmented ambulatory leg. The mature appendage of Branchi- 
necta (fig. 3 F) is cut on its mesal margin into a number of lobes, of 
which five (7-5) are commonly described as endites, while the large, 
so-called flabellum (6) is interpreted as the endopodite, and the mov- 
able apical lobe as the exopodite. The same structure is seen in the 
limbs of Branchipus (fig. 27 A,B) and other anostracans. Since 
endites in general are lobes of the limb segments, the six mesal lobes 
of the phyllopodium suggest that they represent six leg segments, 
coxopodite to propodite. The movable, independently musculated 
apical lobe (Dactpd), therefore, should be the dactylopodite. There 
is thus in the phyllopodium evidence of the presence of the seven 
segments characteristic of the crustacean walking legs. In the second 
maxilliped of Apus (fig. 27 C) seven segments, including a terminal 
dactylopodite, are plainly evident, and each of the first six segments 
except the ischiopodite bears an endite. We can hardly escape the 
conclusion, therefore, that the phyllopodial limbs of the branchiopods 
have been evolved from 7-segmented walking legs. The metamorpho- 
sis of the appendages, therefore, has taken place since the crustaceans 
became crustaceans, and is recapitulated in the larval ontogeny. A 
more extensive discussion of the nature of the primitive arthropod 
limbs is given in section IV of this paper. 

About the only metamorphosis in the life history of Branchinecta 
is the temporary adaptation of the antennae for swimming. It is 
hardly to be supposed that the primitive crustaceans swam with their 


NO. I0 CRUSTACEAN METAMORPHOSES—SNODGRASS 17 


antennae. The nauplius has only three pairs of limbs, and, since it 
must swim, it has no choice but to use what appendages it has. As 
the body lengthens and the postmandibular appendages become broad 
and flat, these appendages assume the function for which they were 
modified in the branchiopod ancestors. The antennae then revert to 
a more simple form (fig. 3 E), and in the adult they are again modi- 
fied, in the male (G) for grasping. 


OSTRACODA 


The ostracods, being enclosed in a bivalve shell from the time they 
leave the egg, go through no body changes of form that might be 
termed a metamorphosis ; their appendages, however, give an impres- 
sive example of the extreme degree of structural modification that an 
ordinary segmented leg may take on. 

The newly hatched ostracod larva is in the nauplius stage of de- 
velopment (fig. 4 A), since it has only the three usual pairs of naupliar 
appendages. It is not a typical nauplius, however ; the antennae and 
mandibles are uniramous, and the body is already enclosed in a shell 
formed in the egg. Here is a good demonstration, then, that the 
crustacean nauplius, in addition to its primitive features, can take on 
a specialized structure characteristic of the order to which it belongs. 
During the postnaupliar stages, as shown in the series of drawings 
(fig. 4) here copied from Schreiber (1922) on the development of 
Cyprinotus incongruens, the postmandibular appendages are succes- 
sively added until the definitive number of seven in all is present in 
the eighth instar (F), in which the larva has attained essentially the 
adult structure. 

There is no question that the naupliar appendages are the anten- 
nules, the antennae, and the mandibles, but there has been some 
difference of opinion as to the identity of the postnaupliar appendages. 
In the Cypridae the first appendage after the mandible (fig. 5 B, 4) 
bears a large, flat, fringed lobe projecting upward in the shell cavity, 
and this appendage is commonly regarded as the maxilla. The next 
appendage (5) Schreiber termed the maxilliped. These two append- 
ages on each side in Cypris arise side by side on the arm of the hypo- 
stome (D, 4, 5), and Cannon (1926) regarded them as the maxillula 
and the maxilla, respectively. In Limnocythere inopinata (A), how- 
ever, as in other Cytheridae and in Nesideidae, appendage 5 is a 
typical leg well separated from 4. If, therefore, appendage 7 is 
interpreted as the maxilla, appendages 5, 6, and 7 are thoracic legs, 
and Kesling (1951) says this is now the accepted interpretation of 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


the ostracod limbs. Since we are here not particularly concerned with 
the identification of the appendages, they have simply been numbered 
on the drawings for purposes of comparison. 





Fic. 4.—Ostracoda. Larval stages of Cyprinotus incongruens Ramdohr (from 
Schreiber, 1922). 

A, nauplius, with three pairs of appendages. B, second instar, with one pair 
of added appendages (4) and caudal furca (f). C, fourth instar, with fifth ap- 
pendages (5). D, fifth instar, with sixth appendages (6). E, sixth instar, with 
seventh appendages (7). F, eighth instar, essentially adult structure. 


Inasmuch as in such forms as Limnocythere (fig. 5 A) the antennae 
and the last three pairs of appendages have the form of segmented 
legs, and in Cypris (B) the sixth and seventh appendages are typical 
legs (E), it may be inferred that the primitive ostracod appendages 


NOF LO CRUSTACEAN METAMORPHOSES—SNODGRASS 19 





Fic. 5.—Ostracoda. 


A, Limnocythere inopinata (Baird), eighth instar (from Scheerer-Ostermyer, 
1940). B, Cypris testudinaria Sharpe, adult, left shell removed. C, Philomedes 
globosa (Lilljeborg), adult, left shell removed. D, Cypris testudinaria, fourth 
and fifth appendages of left side and hypostome (Hst), posterior. E, same, sixth 
appendage. F, Philomedes globosa, fourth appendage. G, same, fifth appendage. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


were all ambulatory limbs. Moreover, since the legs of Limnocythere 
can be correlated with appendages of a very different structure in 
other genera, the ostracods give an actual demonstration of the 
changes that legs can undergo in adaptation to new uses. The very 
unleglike fifth appendage of Cypris (D, 5), for example, must have 
been evolved from a typical leg such as its representative in Limnocy- 
there (A, 5). This same appendage in Philomedes (C, 5) is again 
quite different from the corresponding appendage in Cypris. The 
maxilla of Cypris (D, 4) is represented in Philomedes (C, F, 4) by 
an appendage still suggestive of its leg origin. The fifth appendage 
of Philomedes (C, G, 5), however, has no resemblance to its counter- 
part in Limnocythere (A, 5), though the presence of three small 
marginal lobes and an apical tooth might be taken as evidence of a 
former segmentation. The sixth appendage (C, 6) has likewise three 
small marginal lobes and a broad, fringed apical lobe, but otherwise 
it has departed far from the structure of a leg (A,6). The seventh 
appendage of Philomedes (C, 7) has lost all semblance of a leg; it 
has become a long, flexible, vermiform cleaning organ armed with 
an apical brush of recurved bristles. The corresponding appendage 
in Cypris (B, 7)is likewise used for cleaning the shell chamber, but 
the only concession it has made to its function is an inversion of 
position. The sixth appendage of Cypris testudinaria (fig. 5 E) looks 
like a typical 7-segmented crustacean limb, counting the long terminal 
claw as the dactylopodite. The fourth and fifth podomeres of this 
appendage, however, are perhaps not true segments, since in Cypridop- 
sis vidua Kesling (1951, fig. 20) shows that the muscles from the sixth 
podomere have their origins in the base of the fourth podomere. 

The ostracods give no support to the theoretical phyllopod origin 
of crustacean limbs, and show clearly how simple segmented legs 
can be modified into very unleglike structures. 


COPEPODA 


The copepods include marine and fresh-water free-swimming spe- 
cies and a large number of parasitic species. They are nearly all very 
small crustaceans, mostly from 0.50 mm. to Io mm. in length in the 
adult stage. The simpler free-swimming copepods seem to approach 
more closely the typical shrimplike form of the higher crustaceans 
than do any of the other entomostracans. The body of a generalized 
form such as the marine Calanus (fig. 6) is divided into a cephalo- 
thoracic region bearing the appendages, and a slender limbless ab- 
domen. The cephalothorax includes an anterior unsegmented part 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 21 


(7H) known as the head, or cephalosome, and a posterior thoracic 
region of five segments. The head carries the two pairs of antennae, 
the mandibles, two pairs of maxillae, and the first pair of legs, or 
maxillipeds. The five segments of the thoracic region bear each a pair 
of legs, but the legs of the last pair may be much reduced. The 
genital ducts open on the basal segment of the abdomen. 

The free-swimming copepods occur in such vast numbers in the 
ocean and in some inland lakes that they constitute a most important 
food source for many other aquatic animals from arrowworms to 
whales, but particularly for fishes. Being minute creatures themselves, 
the free copepods feed on the microscopic plant life of the water, 





2Mx Mxpd' 


Fic. 6.—Copepoda. Calanus cristatus Kroyer, adult. 


TAnt, first antenna; 2Ant, second antenna; H, “head’; Md, mandible; 1M x, 
first maxilla; 2M-s, second maxilla; Mapd, maxilliped; VJ, VII, XI, body 
segments. 


which, elaborated in their own bodies, is thus passed on as food for 
the larger animals. It would seem, however, that the copepods have 
retaliated on the animals that eat them, since many species have be- 
come parasites of their potential enemies. Though fish are their favor- 
ite hosts, the parasitic copepods are not discriminative and attack 
almost every kind of creature that lives in the ocean. On the other 
hand, the copepods themselves are infested by numerous parasites, 
even by some of their own kind. Evidently life in the ocean is not 
a happy existence for either the predators or their victims. 

The nonparasitic copepods go through no changes of form in their 
life histories that can truly be called a metamorphosis. Their environ- 
ment is practically the same at all periods of their lives, and there 
is no call for adaptive modifications in either the larval or the adult 
stage. The successive developmental stages are merely steps in growth 
from youth to maturity. As an example, we may take the fresh-water 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Cyclops and follow its life history as described by Dietrich (1915) 
and by Ziegelmayer (1925). 

The Cyclops larva hatches from the egg as a typical nauplius (figs. 
7 A, 8A), which is followed by a second nauplius instar and four 
metanaupliar instars (fig. 8 B, C, D, E) in which the body lengthens, 
and finally (E) five pairs of appendages are added beyond the mandi- 
bles, including the maxillipeds (Mpd) and two pairs of legs (1L, 





Fic. 7—Copepoda. Nauplius and two metanaupliar instars of Cyclops (outlines 
from Ziegelmayer, 1925). 
A, nauplius, ventral, showing three pairs of appendages and labrum (Lm). 
B, first metanaupliar instar. C, fourth metanaupliar instar. 


2L). At the next moult the larva (F) begins to take on the form and 
structure of the adult (G) and is now termed a copepodid, the ending 
id signifying that at this stage the larva has become copepodlike. The 
first copepodid acquires a third pair of legs; with further growth it 
passes through six copepodid instars until at last it becomes a sexually 
mature adult (G). Among Cyclops species there is thus no abrupt 
change between the various stages of growth, but new segments are 
added and the appendages develop from simple rudiments to their 
definitive forms. According to Ziegelmayer the segments formed 
after the nauplius stage are generated in a subterminal zone of growth. 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 23 


The genus Calanus, a typical free-swimming marine copepod of 
the suborder Gymnoplea, likewise develops from nauplius to adult 
by ordinary anamorphic growth without any metamorphic changes 





Fic. 8.—Copepoda: Podoplea. Developmental stages and adult of Cyclops. 
(A-F from Dietrich, 1915; G from Claus, 1863). 


A, Cyclops strenuus Fischer, nauplius, 0.119 mm. B, same, first metanauplius. 
C, same, second metanauplius. D, same, third metanauplius. E, same, fourth 
metanauplius. F, same, first copepodid, 0.303 mm. G, Cyclops coronatus Claus, 
adult female with eggs, 3.50 mm. 

1Ant, first antenna; 2Ant, second antenna; 1L, 2L, 3L, legs; Md, mandible; 
IMz, 2Mx, first and second maxillae; Mxrpd, maxilliped. 


adaptive to different ways of living at different stages. The life history 
of the common Calanus finmarchicus has been described by Lebour 
(1916). The first six instars the author calls nauplii, but some of them 
would ordinarily be regarded as metanauplii, since two posterior 
segments and indications of a third segment are said to appear in the 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


fifth stage, with traces of the fifth and sixth pairs of appendages. In 
the sixth instar, judging from related species, there are present second 
maxillae, maxillipeds, and two pairs of swimming legs. The next 
instar is that of the first copepodid, which has three pairs of legs; the 
fourth legs appear in the second copepodid instar, and the definitive 
number of five is present in the third copepodid. The fifth and last 
copepodid is essentially like the adult. The free-swimming copepods, 
therefore, have a typical anamorphic development. Being crustaceans, 
they are primarily constructed for life in the water, and so long as 
they maintain a free existence there is no need of metamorphic 
adaptations to any other way of living. 

When now we turn to the parasitic copepods, the story is very 
different. An aquatic animal that hatches as a freely swimming larva 
and then becomes sedentary on another animal from which it extracts 
its food changes its environment and its mode of living in a very 
radical way. In some manner difficult to understand metamorphic 
changes of structure have been evolved that adapt the parasitic animal 
to its life of parasitism, and in many cases the transformation has 
been carried so far that the adult parasite could not be identified, or 
even recognized as a crustacean, if its early stages were not known. 

A few copepods appear to be transitional in their habits between a 
free life and one of parasitism. Such species are termed semiparasitic 
by Wilson (1921b), who says they are found on worms, mollusks, 
echinoderms, and in the gill chambers of crabs. These species are 
capable of swimming freely in the water, and their residence on any 
one host may be temporary. Their mouth parts, according to Wilson, 
are not suitable for either chewing or sucking and appear to be adapted 
for licking nourishment from the animals to which they attach them- 
selves. A species with biting mouth parts, however, could hardly 
resist sampling the blood of its host and then becoming an habitual 
parasite. 

The truly parasitic copepods include a large number of species, all 
of which undergo striking metamorphic adaptations to the nature 
of the host or the part of the host attacked, and some of them lead 
a double life on two different species of hosts. Some parasitic copepods 
undergo their metamorphoses during the larval development and 
become again free living in the adult stage; others remain on the host 
and attain their highest degree of metamorphosis as adults. Most of 
them, however, hatch from the eggs as typical nauplii, and in this 
stage or the following copepodid stage they must find their proper 
hosts. 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 25 


As an example of the life history of a parasitic copepod that returns 
to a free life in the adult stage, we may take the monstrillid Cymba- 
soma rigidum Thompson, described by Malaquin (1901) as Haemo- 
cera danae (Claparéde), which in its larval stages lives in the blood 
vessel of the serpulid worm Salmacina dysteri Huxley. The nauplius 
(fig. 9 A) has the usual three pairs of naupliar appendages, but the 
mandibles are recurved hooks, and the young larva has no mouth or 
alimentary canal. It is poorly fitted for swimming, and Malaquin 
suggests that the females probably sow their eggs over a colony of 
the serpulids. When in contact with a worm the nauplius attaches 
itself by its mandibular hooks to the worm’s integument, but it has 
no special organs for penetration. The skin of the worm, however, 
is delicate, and, a puncture once effected, the nauplius does a most 
surprising thing; it casts off its own cuticle and its appendages and 
forces its soft nude body into the host. Within the latter it becomes 
a shrunken, oval mass of undifferentiated cells (B), as if it had re- 
turned to an early embryonic condition to begin development all over 
again. In this form the parasite traverses the coelom of the host and 
makes its way into the ventral blood vessel. Here it secretes a new 
cuticle and then from its ventral side anteriorly there grow out two 
tapering, armlike processes (C) that extend posteriorly in the blood 
vessel of the worm and will serve the parasite as food-absorbing 
organs. Here, therefore, we see a metamorphic development adapting 
the parasite to its life in the host that certainly had no counterpart in 
the presumed free-living ancestors of its species. It is hard enough 
to believe the facts themselves, and we can speculate in vain as to 
how they all came about in evolution. The nauplius is prepared in 
advance for the life it is to lead by being provided with hooked 
mandibles, but what induces it to shed its cuticle and appendages and 
to squeeze itself into the worm? 

With the growth of the young larva in the worm (fig. 9H) the 
nutritive arms increase in length (D, E), the new cuticle is drawn 
out into a rostrum in front (E, Rk), and on the enlarged conical 
posterior part of the body it becomes armed with circles of spines 
directed forward. The organs of the future adult now gradually de- 
velop within the cuticle of the larva (F), and the abdomen forms as 
a ventral flexure (Ab) of the posterior part of the body. At an early 
stage the first antennae are regenerated (F, G, 1Ant) and eventually 
penetrate into the rostrum (1) when the head tissue has receded from 
the latter. From this point on the parasite develops normally into the 
adult form within the cuticular sheath of the larva. Finally, when its 
development is almost completed (1), the parasite becomes strongly 


fh + 


LU 


DWN bs 
LTE 





Fic. 9—Copepoda: Monstrillidae. Larval stages and adult of Haemocera 
danae (Claparéde) parasitic in the polychaete Salmacina dysteri Huxley (from 
Malaquin, 1901). 

A, free nauplius. B, after penetration into host. C, same, with nutritive ten- 
tacles (t). D, same, later stage. E, later stage, with cuticular envelope, rostrum 
(R), and spines. F, beginning transformation to adult inside cuticular sheath, 
rudiment of abdomen (Ab) bent forward. G, later stage of male, showing testis 
(Tes). H, specimen of Salmacina with two parasites in ventral blood vessel. 
I, male parasite almost adult. J, adult female, free after shedding the sheath and 
leaving the host; rAnt, first antenna, ef, egg filaments, gSeg, genital segment, 
Ov, ovary. 


26 


NO. ILO CRUSTACEAN METAMORPHOSES—SNODGRASS 27 


active, doubling and straightening upon itself with the result that it 
ruptures both its enclosing sheath and the integument of the host. 
Then it escapes, leaving behind in its late host its spiny cuticle and 
its nutritive arms, which will no longer be needed. The monstrillid 
thus, according to Malaquin, makes during its life only two moults, 
one on entering the host, the other on leaving it. With its liberation 
the adult becomes at once an active free-swimming copepod (J). It 
now has only one pair of antennae and four pairs of swimming legs, 
and it lacks a complete alimentary canal. The body of the female, 
however, is mostly filled with a great mass of eggs (J, Ow) ; the busi- 
ness of the adult is the procreation of more parasites. 

Members of the family Caligidae, mostly parasitic on fish, are also 
free in the adult stage, but, though the adults are at liberty to leave 
the host and are equipped with swimming legs, they still depend for 
their food on the host that nourished them as larvae or on some other 
fish of the same kind. They, therefore, live largely as free external 
parasites. The structure and habits of many species of Caligidae have 
been described by Wilson (192Ia), and a detailed account of the 
larval stages of Caligus curtus (O. F. Miller) is given by Heegaard 
(1947). 

In Caligus curtus, according to Heegaard, there are two naupliar 
instars, the second of which goes over directly into a first copepodid 
without an intervening metanaupliar stage. The first copepodid is 
followed by a second copepodid, and then come five larval stages in 
a form known as a chalimus before the individual becomes adult. The 
actively swimming first copepodid has the responsibility of finding 
a host, which will be a codfish. It grasps a scale or a fin ray of the 
fish by means of its clawed second antennae, and holds on with the 
maxillipeds. After attachment the copepodid moults into the second 
copepodid (fig. 10 A). In this stage a gland in the head produces a 
secretion which will be discharged from the frontal region as a fila- 
ment (B), which becomes firmly fixed to a scale or a fin ray of the 
host. The parasite now becomes quiescent and takes no food as it 
hangs motionless on its attachment line, while within its cuticle a 
development takes place that will transform the copepodid into the 
first chalimus. This quiescent period of the copepod (B) is termed 
by Heegaard and some other writers a “pupa,” but, though motionless 
and nonfeeding, it is not comparable to the pupa of an insect. The 
insect pupa is a stage in itself during which the metamorphosed larva 
reverts to the parental form. Each larval instar of any arthropod 
begins its development within the loosened cuticle of the preceding 
instar. The copepod “pupa,” therefore, is merely the second copepodid 


28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


after completion of its own growth when the first chalimus is de- 
veloping beneath its cuticle. An insect larva in a corresponding stage 
becomes quiescent and ceases to feed, but it is not a pupa. This con- 
cealed period in which any instar begins its development within the 





Fic. 10.—Copepoda. Larval and adult stages of fish parasites. (A, B from 
Heegaard, 1947; C, D from Wilson, 1905; E, F from Wilson, 1921a; G from 
Wilson, 1917.) 

A, Caligus curtus (O. F. Miiller), second copepodid. B, same, transforming 
stage of second copepodid. C, same, mature male chalimus. D, same, adult male. 
E, Trebius latifurcatus Wilson, adult female. F, Blakeanus corniger Wilson, 
adult female. G, Haemobaphes cyclopterina (Fab.), adult female. 


cuticle of the preceding instar has been termed by Hinton (1946) 
the pharate, or cloaked, phase of development. 

The young chalimus that emerges from the copepodid cuticle is not 
particularly different from the copepodid, though it is somewhat more 
advanced in development. Its first concern is to reattach itself to the 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 29 


host, since the old filament remained with the discarded copepodid 
skin. Heegaard (1947) gives an interesting account of how the 
young chalimus with the pointed frontal lobe of its head bores a hole 
in a fin ray of the host. Into the wound thus formed is injected the 
secretion from the head gland, which hardens and holds fast, while 
the chalimus backs away and draws it out into a filament that secures 
the parasite to the host, but still allows it to move about on its tether. 
According to Heegaard each of the four succeeding chalimus stages 
reattaches itself in the same manner. The chalimus (C) was given its 
name because when first discovered it was thought to be the adult of 
an unknown species. Since the chalimus stages progressively de- 
velop from the second copepodid to the adult (D), they evidently 
represent the later copepodid stages of free-living copepods. 

The adults of Caligus curtus (fig. 10D) have pretty much the 
form and structure of an ordinary copepod, but, having no attachment 
to the host, both the males and the females are free to swim away. 
The egg-carrying female of another species with similar habits is 
shown at E of the figure. Since these copepods are dependent on a 
host for food in the adult stage, they retain their parasitic habits and 
are generally found crawling and feeding on the host, though they 
have not become specially modified in structure for a life of parasitism. 
This condition of dependence on a host, however, Wilson (1915) 
points out, constitutes the first step toward adult degeneration. If 
the adult parasite finds it advantageous to remain on the host, organs 
of locomotion become unnecessary, and in the end all that is needed 
are organs of nutrition and reproduction. The species shown at F, 
parasitic in an ascidian, still retains its appendages and a segmented 
abdomen, but the thorax has taken on a strange shape. The female 
at G, however, a permanent parasite on the gills of a fish, has de- 
generated from the copepod structure almost to the limit of simpli- 
fication. Yet, as already noted, “degeneration” is merely adaptation 
by the elimination of unnecessary organs. 

An example of an intermediate degree of degenerative simplification 
is seen in the lernaeopodid fish parasite Achtheres ambloplitis (fig. 
11) described by Wilson (1911). In this copepod, Wilson says, the 
naupliar and metanaupliar stages are completed in the egg, and the 
larva hatches as a copepodid (A). During the egg stage the head 
gland produces a filament, which is still coiled in the head of the 
emerging copepodid (A, f). The young larva has two pairs of feath- 
ery swimming legs, and its maxillipeds (M-apd) are armed with strong 
hooks. It swims actively in search of a host, which must be a fish 
of the surface-swimming Centrarchidae. That the young copepod 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


deliberately presents itself to the head end of a fish has perhaps not 
been observed, but the fish unwittingly engulfs the copepod as food, 
which is of course just what the prospective parasite wanted it to do. 
To save itself from being swallowed the copepod grasps a gill arch of 
the fish with the hooks of its maxillipeds. Then it pushes its head into 





Fic. 11.—Copepoda: Lernaeopodidae. Developmental stages of fish parasites 
(from Wilson, 1910, 1915). 

A, Achtheres ambloplitis Kellicott, first copepodid, with filament (f) in head. 
B, same, second copepodid. C, same, with filament extruded and attached. D, 
same, adult male. E, same, adult female. F, Salminicola siscowat (Kellicott), 
egg-carrying female. 


the soft skin of a gill, which act breaks the cuticle over its head and 
releases the filament. The filament protrudes into the wound of the 
gill and the end spreads out into a disc that anchors the parasite inside 
the gill chamber of the fish. 

The first copepodid larva of Achtheres (fig. 11 A) undergoes a 
moult and enters a second copepodid instar (B), which is decreased 
in size and has taken on a different shape. The swimming legs, being 
now useless organs, are greatly reduced and later disappear (C). 
The mandibles have become toothed piercing organs for feeding. The 


NO. 10 CRUSTACEAN METAMORPHOSES—SNODGRASS 31 


large second maxillae (C, 2M) are much thickened and securely 
grasp the base of the attachment filament (f) by means of hooks 
imbedded in apical depressions. Then the larva backs away and draws 
the filament out to its full length, and thus maintains its hold on the 
gill with sufficient freedom of movement for feeding. At the next 
moult the sexes are mature. The female grows to a length of 4 or 
5 mm., but the male remains a pygmy not over I mm. long. 

In the adult female (fig. 11 E) the maxillae are greatly lengthened, 
but the filament (f) is contracted so that only a short stalk projects 
beyond the maxillae. The maxillae of the male (D) are relatively 
not so long as those of the female, but the filament is unshortened. 
The filament, being a product of an internal head gland, is not shed 
and renewed at the moults; it retains its attachment and thus allows 
the parasite to complete its life in security within the gill chamber 
of the fish. The long filament of the small adult male permits the 
male to swing around on his tether until he comes in contact with a 
female, whom he grasps with his maxilliped claws and then lets go his 
hold on the filament, which remains attached to the gill. The female 
of another similar species of the genus Salminicola (F) is depicted 
by Wilson (1915) carrying her extruded eggs (es) in two long 
cylindrical sacs projecting from the gonopores while still attached to 
the gill of the fish. The newly hatched young presumably are carried 
out of the gill chamber in the expiratory currents of water. 

A good example of a parasitic copepod that inhabits two hosts dur- 
ing its life is the well-known fish parasite Lernaeocera branchialis 
(L.), a member of the Lernaeopodidae. This species during its larval 
life is an attached parasite on the gills of a flounder, but when adult 
both the male and the female become free and leave the flounder. The 
male undergoes no further transformation, and, after mating with a 
female still on the flounder, his purpose is accomplished. The female, 
on the other hand, is not yet sexually mature, and some instinct now 
urges her to leave the flounder and to seek a cod on which to com- 
plete the development of her ovaries. Once attached in the gill 
chamber of a cod she goes through an adult metamorphosis by which 
she is functionally reduced to the bare essentials necessary for feeding 
and egg production. For an account of the life history of Lernaeocera 
branchialis we may draw on the work of Pedaschenko (1898), Scott 
(1g0o1), Wilson (1917), Schuurmans-Stekhoven (1936), Sproston 
(1942), and Capart (1948). 

There is some difference of opinion concerning the nature of the 
early forms of this species. Pedaschenko says the first larva is a 
metanauplius (fig. 12 B) ; Scott and Sproston observed only one early 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


UAE GCPM ECU URAC AAC a AC RC CCC 





Fic. 12—Copepoda. Developmental stages of the fish parasite Lernaeocera 
branchialis (L.). (A, C, D, E, G from Sproston, 1942; B from Pedaschenko, 
1898; F, I from Capart, 1948; H, J from Scott, 1901.) 

A, nauplius, 0.37 mm. B, metanauplius. C, free-swimming copepodid. D, 
third instar of chalimus stage on flounder. E, free-swimming adult male, 1.55 
mm. F, young inseminated female on flounder. G, female on cod, beginning 
metamorphosis to penella stage. H, female in fully developed penella stage. 
11.4 mm, I, adult egg-carrying female on cod, 40 mm. J, section of mature 
female. 

An, anus; AlCnl, alimentary canal; 1Ant, first antenna; 2Ant, second an- 
tenna; CmGld, cement gland; es, egg string; Mth, mouth; 2M, second maxilla; 
M-pd, maxilliped; Odct, oviduct; Ov, ovary; Sphr, spermatophore. 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 33 


stage, and called this stage a nauplius (A) ; while Wilson and Capart 
record both a nauplius and a metanauplius. The matter is of no 
particular importance for us in a study of the metamorphosis of the 
species. Whatever the larva that hatches from the egg may be, it 
moults into a free-swimming copepodid (C). Though the copepodid 
is only about half a millimeter in length, it has the responsibility of 
finding a flounder and of fixing itself to the gills of the fish, for which 
latter purpose it is provided with strongly chelate second antennae. 
Its hold on the gill, Sproston says, is never relinquished, and becomes 
the anchorage of the parasite until the free-swimming adult stage is 
reached. The gill filaments, however, are grasped also by the second 
maxillae in order to bring the mouth parts into close contact with the 
tissues on which the parasite feeds. 

When the copepodid moults the larva becomes a chalimus (fig. 
12 D), but there is little change in form or structure. The chalimus, 
however, in its first instar acquires an additional attachment on the 
host in the form of a filament secreted by a gland in the head, which 
is anchored in the gill by two diverging branches that penetrate into 
punctures in the gill tissue. The rest of the secretion from the gland, 
Sproston says, falls back on the head of the larva where it hardens 
into a conical hood. The chalimus goes through four instars, and 
with each moult but the last a new hood is formed while the old ones 
remain, so that there are thus formed a set of overlapping caps corre- 
sponding in number with the moults. The third instar of the chalimus, 
to be identified as such by its three hoods, is illustrated at D of figure 
12, redrawn from Sproston. The copepodid and the chalimus are 
metamorphic larval forms adapted to their respective functions of 
swimming and parasitic feeding. During its four instars the chalimus 
gradually approaches the adult structure, which is attained at the 
fourth moult after the copepodid stage. 

The adult male of Lernaeocera (fig. 12 E) leaves the old attachment 
filament with the castoff chalimus cuticle hanging on the gill of the 
flounder, and goes off in search of a female. The female (F), how- 
ever, awaits the coming of a male before she relinquishes her hold on 
the flounder. When the male finds a female still attached, mating 
takes place; two large spermatophores are inserted into the genital 
ducts of the female and are eventually lodged in her lengthened genital 
segment (F, Sphr). The female, still not sexually mature, then frees 
herself from the flounder and swims away to look for her second 
host, which should be a cod. On attaining a prospective victim, the 
female fixes herself to the bases of the gills by her second antennae, 
and now begins her metamorphosis into the final egg-producing stage. 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


After attachment on the cod the head of the female undergoes a 
curious transformation. Large branching, hornlike processes grow 
out from it and sink into the host tissue as anchoring devices (fig. 
121). The proboscislike mouth region penetrates deeply in the flesh 
at the base of the gill until a large blood vessel is reached, from which 
the female will draw a rich nourishment for the maturing of her eggs. 
The first change of the body is a lengthening of the abdomen, princi- 
pally the genital segment, which grows out in a twisted wormlike 
form (G), and finally (H) becomes a long, straight, slender ap- 
pendage hanging from the thorax. At this stage the female is known 
as a penella from her resemblance to another adult copepod of that 
name. In the figure the penella stage shown at H is, of course, drawn 
on a much smaller scale than is the female at F or G. Next, the abdo- 
men swells into a great, elongate, twisted bag (1). The female in 
her final stage is said by Wilson (1917) to attain a length of 4o 
millimeters when fully extended. From now on she is merely an egg- 
producing organism. Her internal organs (J) consist principally of 
the enlarged alimentary canal (AlCnl), the ovaries (Ov) and oviducts 
(Odct), and a pair of cement glands (CmGld) that form the casings 
for the eggs. The eggs are discharged in two long coiled strings (es), 
which, Wilson says, reach a lenth of 150 to 200 millimeters. Consider- 
ing the number of eggs that the species produces, any flounder or cod 
may consider itself lucky if it escapes infestation. According to 
Schuurmans-Stekhoven there is only one generation of the parasite 
each year. 

The metamorphosis of Lernaeocera branchialis affects principally 
the female on the secondary host. The less modified chalimus instars 
carry on the developmental processes while attached on the flounder 
as do the copepodid stages of free-living species. The adult male and 
the adult female on leaving the flounder are normal, swimming cope- 
pods. The transformation of the female on the cod involves, on the 
one hand, a simplification of the thorax until it becomes indistinguish- 
able from the abdomen, except for the retention of the appendages ; 
but, on the other hand, there is a new development of anchoring 
process on the head, and a great overgrowth of the reproductive part 
of the body. The metamorphosis of the female, therefore, is both 
recessive and progressive in an anatomcial sense. A study of the de- 
velopment and metamorphosis should take into consideration not only 
the anatomical changes that the individual goes through, but also the 
changes in its instincts. The copepodid of Lernaeocera, for example, 
must have an instinctive urge to attach itself to a flounder; the adult 
female instinctively leaves the flounder and looks for a cod. 


NO. 10 CRUSTACEAN METAMORPHOSES—SNODGRASS~ . 35 


Copepod fish parasites are not all content with attacking the scales, 
fins, or gills of the host. Some make their abode in the nostrils of the 
fish; others penetrate through the skin into the body cavity where 
they attack the vital inner organs. The worst of them are members 
of the genus Phrixocephalus, several species of which are described 
by Wilson (1917). These parasites bore into the eyes of their victims 
in order to feed from blood vessels at the back of the organs. Para- 
sites seem to have been endowed by nature with great versatility, but 
the life of a fish is nothing to be envied. 


CIRRIPEDIA 


The cirripeds include the familiar barnacles and several groups of 
parasitic species. The first-stage larvae in most cases are nauplii 
usually characterized by the presence of a pair of lateral frontal horns 
on the anterior part of the body. In some species the horns are merely 
short spines (figs. 14 B, 16 A, fh), in others they are long and either 
straight or curved, but when present the horns identify the nauplius as 
a young cirriped. The nauplius becomes a metanauplius; the meta- 
nauplius transforms into a free-swimming larval stage known as a 
cypris because its body is enclosed in a bivalve shell with a closing 
muscle, and thus resembles the ostracod of the same name. The 
cirriped cypris (fig. 14 C) has six pairs of swimming legs, a simple 
median eye, compound lateral eyes, and a pair of antennules project- 
ing from the anterior end of the shell. After swimming freely for 
some time the cypris of most species attaches itself by the antennules 
to some solid object on which it remains permanently fixed and here 
develops into the adult form. 

The barnacles in the adult stage (fig. 14 F, H) are sedentary on 
rocks, clam shells, wooden piles, ship bottoms, whales, or almost any- 
thing else in the ocean, and they get their food from the water. The 
parasitic cirripeds attach themselves to other animals and derive their 
sustenance from the host. The adult barnacles retain enough of their 
ancestral structure to be recognized as crustaceans ; some of the para- 
sitic cirripeds, on the other hand, undergo such extreme degrees of 
adult metamorphosis that their crustacean derivation is known only 
from their early larval stages. 

The Ascothoracica.—The members of this suborder are of particu- 
lar interest because as adults they appear to be equivalent to the cypris 
stage of other cirripeds. If they truly are cirripeds, therefore, they 
evidently are a primitive group of the order, and suggest that the 
cirripeds have been derived from cyprislike ancestors, perhaps re- 


36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


lated to the bivalved Ostracoda. From the standpoint of metamorpho- 
sis the Ascothoracica are of small interest, since whatever modifica- 
tions some of them do undergo effect principally a simplification of 
the cypris structure. They are all minute creatures parasitic on 
Actinozoa and Echinodermata. 

The least modified member of the Ascothoracica is Synagoga mira 
Norman (fig. 13 A), which lives externally on the black corals 
Antipathes, clinging to the host by the large antennules. Since Syna- 
goga has well-developed setigerous legs, however, it appears probable 
that it can relax its hold and swim from one host to another. The 
species is known only from a few specimens described by Norman 
(1913). The head and thorax are enclosed in a large, oval bivalve 
shell, 4 millimeters in length, provided with strong adductor muscles, 
but the slender, five-segmented abdomen projects freely from the 
shell and bears a pair of long uropods. The large antennules (7Ant) 
are armed with apical hooks; the six pairs of thoracic legs bear long 
setae and are evidently adapted for swimming. The mouth parts as 
described by Norman are slender piercing organs enclosed in a large 
conical proboscis (Prb). Of all the Ascothoracica, Synagoga mira 
alone appears to have no metamorphosis and to have retained the 
ability to swim; no other species, therefore, has so good a claim to 
being a primitive cirriped. 

A related member of the Ascothoracica is described by Okada 
(1938) as Synagoga metacrinicola (fig. 13 B). This species has the 
entire body enclosed in the shell, the abdomen being relatively short, 
but otherwise it is similar to S. mira. Okada finds well-differentiated 
males and females in S. metacrinicola, the sexes being separate in most 
of the Ascothoracica, in which the males are much smaller than the 
females. He reports that Norman’s specimens, supposed to be females, 
are found on reexamination by sections to be males with mature 
spermatozoa. Okada thus demonstrates that the known examples of 
Synagoga are adult forms and not larvae of an otherwise unknown 
species, as some writers had suggested they might be. 

The other Ascothoracica that are parasitic on horny corals appear 
as small budlike bodies on the coral stems. The shells are of various 
shapes and in some species are enclosed in a tunic derived from the 
host. In most of these forms the legs are more or less reduced and 
lack swimming setae. An ascothoracid described by Heegaard (1951) 
as Ascothorax bulbosa, found in specimens of an ophiuroid, or brittle 
starfish, has an oval shell (fig. 13 C), the small males being attached 
dorsally on the females beneath the cuticle of the latter. The body 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 37 


of the animal (D, E) is somewhat deformed and the thoracic legs are 
reduced. 
The greatest modification among the Ascothoracica occurs in the 





Fic. 13.—Cirripedia: Ascothoracica. (A from Norman, 1913; B from Okada, 
1938; C, D, E from Heegaard, 1951; F, G from Knipowitsch, 1890.) 

A, Synagoga mira Norman, adult. B, Synagoga metacrinicola Okada. C, 
Ascothorax bulbosa Heegaard, shell of female with small male on top, internal 
parasite of ophiuroid. D, same, female. E, same, male. F, Dendrogaster asteri- 
cola Knipowitsch, cypris larva. G, same, adult enclosed in branched mantle, 
internal parasite of starfish. 


Dendrogasteridae, which are internal parasites of echinoderms. Den- 
drogaster astericola, described by Knipowitsch (1890), is enclosed in 
a voluminous mantle (fig. 13 G) with large lateral lobes, which are 
penetrated by diverticula of the stomach. The cypris larva (F), 


38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


however, is a typical cirriped cypris, much resembling the adult of 
Synagoga (A). A species figured by Fisher (1911) as Dendrogaster 
arbusculus, found in a Californian starfish, has an elaborately branched 
structure. 

The known nauplii of the Ascothoracica, according to Okada, differ 
from the nauplii of other cirripeds in the absence of the usual frontal 
horns, another feature that sets the ascothoracicans off as a primitive 
branch of the cirripeds. Some species hatch as nauplii, others as meta- 
nauplii, and still others in the cypris stage. 

The Thoracica.—To this suborder belong the barnacles, which in 
the adult stage are enclosed in calcareous shells. Some are conical and 
sit flat on the substrate (fig. 14 F), others are flattened and sup- 
ported on stalks (H). When either kind is broken open, however, 
there is exposed within the shell a shrimplike creature (G) lying on 
its back or standing on its head with its cirruslike feet, when active, 
sticking out of the top or side (H) with a waving movement. 

The nauplius of a common barnacle such as Balanus, described by 
Runnstrém (1924-1925), has the typical naupliar structure (fig. 14 A) 
except for the pair of small horns (B, fh) on the anterior part of 
its body. Runnstrom describes two naupliar stages, but since the sec- 
ond becomes elongate and acquires rudiments of three postmandibular 
appendages it would ordinarily be called a metanauplius. After a few 
hours of swimming, the metanauplius abruptly transforms into a 
cypris (C) with a bivalve shell and long seta-bearing legs, wherewith 
it is better equipped for a pelagic life. Eventually the cypris fixes 
itself to a support by its first antennae (1Ant), each of which (E) 
is provided with an adhesive cup on the third segment. A cementing 
substance discharged through the antennae from glands in the head 
gives the cypris a permanent attachment. Then the cypris withdraws 
the hind part of its body and its legs into the shell, and now begins 
the formation of the plates of the adult barnacle. According to 
Runnstrom, the plates are first formed as chitinizations of the mantle 
and only later become calcified. When the plates have the essential 
adult pattern (D) the cypris shell is cast off, and with the moult the 
legs of the cypris are replaced by the cirri of the barnacle. 

The metamorphosis of the cypris into the barnacle is not excessive. 
It is a structural adaptation to the permanently sessile condition within 
the shell, and the eyes are absorbed as now useless organs. The 
changes that take place in the body have been described by Doochin 
(1951). The shell-closing muscle of the cypris is retained (fig. 14 G, 
mcl), and the mantle supporting the plates of the shell is attached to 


NO. 10 CRUSTACEAN METAMORPHOSES—SNODGRASS 39 


the body only around the ends of the muscle. The peduncle of the 
stalked barnacles is a product of the head and becomes occupied by 
connective tissue and muscles. The barnacles are hermaphroditic, but 
they generally live in crowded colonies and cross fertilization is made 
possible by a long, tubular penis arising at the base of the vestigial 
abdomen. 

The Rhizocephala.—In this suborder of parasitic cirripeds we en- 
counter the strangest metamorphic phenomena known in the whole 





Fic. 14.—Cirripedia: Thoracica. (A-E from Runnstrom, 1924-1925.) 

A, Balanus balanoides (L.), nauplius. B, same, anterior end of body with 
median eye and frontal horns (fh). C, same, cypris larva. D, same, later stage, 
barnacle plates formed inside cypris shell. E, same, first antenna of cypris with 
attachment cup on third segment. F, Balanus eburneus Gould, group of adults. 
G, Lepas anserifera L., adult animal in natural position removed from shell. 
H, same, stalked shell. 


animal kingdom. The rhizocephalans include a number of genera, of 
which the best known are crab parasites of the genus Sacculina. The 
visible external evidence that a crab is parasitized by a sacculinid is 
the presence of a large saclike body attached ventrally on the crab at 
the base of the abdomen (fig. 15 A). This external sac is the re- 
productive part of the parasite containing the ovaries and the testes, 
but from it long, rootlike processes extend into the body of the crab 
and serve for the nutrition of the parasite. The eggs are fertilized 
and hatch within the external sac, giving rise to nauplii, which trans- 
form into typical cirriped cypris larvae. The free-swimming cypris 


40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


larvae escape through a hole in the sac, find another crab, and enter 
the latter after undergoing extraordinary transformation processes. 
The life history of Sacculina carcini Thompson was fully described 
and illustrated by Delage in 1884, and Delage’s account has been veri- 
fied, at least in part, by G. Smith (1906) and Veillet (1945). It still 
remains as the authentic history of a Sacculina, and the following 
story of the life and metamorphosis of this parasite is based on the 
papers by Delage and Smith, with illustrations taken from both. 

The nauplius of Sacculina (fig. 16 A) has the characteristic frontal 
horns (fh) of cirriped nauplii, but it lacks an alimentary canal and 
has neither a mouth nor an anus. After several moults the nauplius 
becomes a cypris larva (B) with a length of 0.20 mm. On leaving 





Fic. 15.—Cirripedia: Rhizocephala. External parasitic stages on crabs. 


A, Loxothylacus texanus Boschma, a sacculinid on Callinectes sapidus Rath- 
bun. B, Thompsonia on Thalamita prymna (Herbst) (from Potts, 1915). 


the brood sac on the crab, the cypris leads a free life in the ocean for 
several days. Finally, on finding a young crab that has just moulted, 
it attaches itself to the latter by one of its antennules (C, rAnt), which 
are provided with small suction cups. The point of attachment is 
usually in the membrane at the base of a hair (Hr). When firmly 
secured the cypris begins violent swinging movements of the body, 
which detach the thorax (7h) along with the legs and the abdomen 
and throw the whole rear part of the body out of the shell (Sh). 
From the large hole thus left in the head end of the cypris are now 
expelled most of the internal tissues, leaving only a mass of cells con- 
taining the reproductive elements. Later the hole closes. 

While this process of elimination has been going on, other changes 
take place. The body of the larva separates from the shell (D) and 
contracts to a sac walled by the ectoderm, which is much smaller than 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 4I 


the original cypris (B). Soon a new cuticle is secreted on the surface 
of the sac (D, rCt) in continuity with the cuticle of the attached 
antenna, and the larva becomes a compact oval body still within the 
shell but now entirely free from it. Again, as if preparing for a moult, 
a second cuticle (2Ct) is formed beneath the outer one, and a small 
point (d) grows out from its anterior end into the hollow of the 
antenna. The body of the larva then retracts within the outer cuticle 
(E), and as it does so the cuticular point elongates into a long, hollow 
dartlike tube (d) with the narrow end cut off obliquely like the point 
of a hypodermic needle, and its widened base embedded in the body of 
the retracted tissue of the larva. This newly formed organ Delage 
called the dart, and the larva armed with the dart he termed a kentro- 
gon (from Greek kentron, a dart, and gonos, a larva). The shell 
together with its loose inclusions is now thrown off, leaving the kentro- 
gon, still enclosed in the outer cuticle, attached to the erab by the 
antenna (F). 

The body of the larva again expands and pushes the dart into the 
antenna (fig. 16 F) until its tip comes into contact with the integument 
of the crab. Since the parasite is held fast by the antenna, the dart 
pierces the integument instead, pushing the larva away from it, and 
finally (G) projects into the body of the crab. Now the soft tissues of 
the larva contract away from the cuticle but remain still connected 
with the base of the dart. The remains of the larva thus have a free 
passageway into the body of the crab through the narrow channel of 
the dart, the orifice of which is said by Delage to be 3 to 6 microns 
in diameter. Though Delage says he did not observe the actual passage 
of the larval substance through the dart, globules are seen inside the 
dart and the parasite is next found inside the crab. By the method of 
the Sacculina a mouse might get into the pantry through the keyhole 
of the door, but once inside it would have to devise a new way of 
eating. This problem the Sacculina solves very easily—it simply 
adopts the feeding method of a plant by sending out absorbent roots 
among the organs of the crab. 

Inside the crab the parasite becomes a small oval body consisting 
of a mass of cells enclosed in an ectodermal epithelium. It finds its 
way to the ventral side of the crab’s intestine and here becomes at- 
tached. Now the principal concern of the parasite is to obtain nourish- 
ment from the host for maturing the germ cells which it has brought 
with it from the cypris stage. Incidentally, this will be the first food 
from an external source that the larva itself has had, since it was 
hatched without an alimentary canal. The larval body expands 
against the intestine of the crab (fig. 17 A) and sends out branching 


42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 





Fic. 16.—Cirripedia: Rhizocephala. Metamorphosis of Sacculina carcint Thomp- 
son (from Delage, 1884). 


A, nauplius. B, free-swimming cypris stage ready to moult. C, cypris fixed 
by antenna at base of hair of crab, shell separated, thorax detached and thrown 
off with internal tissues. D, larva still in shell has formed a new cuticle (1Ct). 
E, shell being shed, larval body retracted within cuticle, with long, hollow “dart” 
(d) extended toward base of antenna. F, larva with a second inner cuticle 
(2Ct), the dart extended into antenna. G, larval body expanded, the dart has 
pierced the hair membrane of the crab. 

Ab, abdomen; rAnt, first antenna; rCt, outer cuticle; 2Ct, inner cuticle; d, 
dart; fh, frontal horn; Hr, hair of crab; Sh, cypris shell; Th, thorax. 


NO; LO CRUSTACEAN METAMORPHOSES—SNODGRASS 43 


rootlike processes (rhizai), which continue to grow, branch, and unite 
until a network surrounds the intestine (B), from which branches 
penetrate between the other organs and extend out into the append- 
ages. The roots do not enter the tissue of the crab, but Delage says 
only the heart and the gills are not attacked. These are the organs 
necessary for maintaining the life of the host and therefore that of 
the parasite, but how did the parasite ever learn to discriminate? The 
Sacculina at this stage has been aptly likened to a fungus. That a 
crustacean can be so transformed shows the unlimited potentialities 
of metamorphosis. 

When nutrition has been fully provided for, attention must be given 
to the reproductive function. If the eggs were allowed to hatch inside 
the crab, the young larvae would find themselves in a prison from 
which there would be no escape. The body of the parasite, therefore, 
emerges through the ventral integument of the crab and becomes a 
brood chamber in which the eggs mature and from which the larvae 
are liberated into the ocean. The pressure of the parasite’s body 
causes a dissolution of the crab’s epidermis beneath it, and prevents 
the formation of cuticle at this point. Consequently at the next moult 
of the crab the Sacculina body containing the reproductive cells 
emerges and becomes external, but is still connected with the crab 
by a short peduncle giving passage to the feeding roots. The place of 
emergence is at the middle of an abdominal segment ; if it were inter- 
segmental, movements of the abdomen might constrict the peduncle 
and shut off the food supply of the parasite. It seems that the simpler 
a creature may be in its organization, the more does nature guard 
it against emergencies. It is interesting to note that the species shown 
at A of figure 15 is exactly modeled to fit into the pocket between the 
under surface of the thorax of the crab and the reflexed abdomen 
beneath it. 

The external parasite, as seen in section (fig. 17 C, D) consists of 
a central mass of cells contained in a tunic suspended from the 
peduncle, and of an outer mantle (mn) that encloses a peripheral 
brood chamber (bc). The figures at C and D, taken from G. Smith 
(1906), depict a species of Peltogaster, but the structure is essentially 
the same in Sacculina. The cells of the central mass are the eggs in 
the ovary (Ov) ; above them is a pair of tubular testes (Tes) anda 
single nerve ganglion (Gung). The ripe eggs are discharged into the 
mantle cavity and here fertilized by spermatozoa from the testes, the 
parasites being necessarily hermaphroditic. The larvae escape in the 
cypris stage from an opening (D, op) in one end of the brood 
chamber. Successive lots of eggs are discharged and fertilized, and 
after each brood of larvae the cuticular lining of the brood chamber 


44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


is shed. The maturation of the sperm and the eggs and the fertiliza- 
tion of the latter are fully described by Smith, but here ends our 
discussion of the metamorphosis of Sacculina. 





Fic. 17.—Cirripedia; Rhizocephala. Internal and external parasitic stages. (A-D 
from G. Smith, 1906; E, F from Potts, rors.) 


A, Sacculina neglecta attached on intestine of crab Inachus scorpio. B, same, 
later stage with root system developed. C, Peltogaster sp., diagrammatic cross 
sction of parasite after emergence on ventral side of crab. D, same, longitudinal 
section. E, Thompsonia sp., part of root system in tail fan of crab Synalpheus 
brucei, with external brood sacs. F, same, external sacs on chela of Thalamita 
prymna. 

a, internal reproductive buds; b, external brood sac containing cypris larvae; 
be, brood cavity; c, external sac with all but a few larvae escaped through 
terminal aperture (op) ; Gng, ganglion; mn, mantle; Od, oviduct; of, external 
opening of brood cavity; Ov, ovary; r, nutritive roots; Jes, testis. 


The parasitization of the crab by Sacculina adversely affects the 
gonads and results in structural changes of the host called parasitic 
castration. At the moult accompanying the emergence of the parasite, 
the male crab takes on certain female characters and the female suffers 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 45 


a change from normal. Inasmuch as Sacculina produces only one re- 
productive body, the parasite has no concern with what happens to the 
host. 

Peltogaster socialis, another rhizocephalan, differs from species of 
Sacculina in that a number of parasites, 2 to 30 of them, all in about 
the same stage of development, are found on the outside of one host. 
In his investigation of this species, G. Smith (1906) reported that 
each external parasite appeared to have its individual root system in 
the crab. Potts (1915') questioned the accuracy of Smith’s observa- 
tion, and suggested that more probably the several external parasites 
arise from a common root system, pointing out that Peltogaster 
socialis is a comparatively rare species and that it would seem unlikely 
that so many cypris larvae should attack the same crab at the same 
time. 

That many external reproductive sacs may arise from one internal 
system of roots has been amply demonstrated by Potts (1915) in 
his study of the genus Thompsonia. Species of this genus, parasitic 
on various crabs, reach the ultimate in the conversion of an adult 
crustacean to the status of a fungus. The parasite within the host has 
the form of an extensive and intricate network of fine branching and 
anastomosing threads distributed principally on the ventral wall of the 
abdomen at both sides of the nerve cord, but also entering the thorax 
where the branches may extend up on the lateral and dorsal walls. 
The root threads, according to Potts, are from 10 to 20 microns in 
thickness. From the central network branches penetrate into the 
thoracic and abdominal appendages and into the lobes of the tail fan. 

On the branches in the appendages are developed small budlike 
processes (fig. 17 E,a) that project outward against the integument. 
These buds contain the germ cells that will become ova. At the next 
moult of the crab they break through the soft new cuticle and become 
small external sacs (E, b, and F) standing on the surface. The sacs 
may be so numerous that the appendages, especially the legs, are 
loaded with them (fig. 15 B). These external sacs are the reproductive 
organs of the parasite, and might be likened to the spore-bearing 
bodies of a fungus nycellium. Since Thompsonia produces no male 
elements, the eggs are apparently parthenogenetic. They hatch di- 
rectly into young cypris larvae (fig. 17 E), which, before the next 
moult of the crab, escape from the sac through an apical perforation 
(op). The empty sacs are carried off on the exuviae at the following 
moult of the crab. The development of the eggs, therefore, is so regu- 
lated that the larvae reach maturity during the time between moults 
of the host. At each moult a new crop of egg sacs breaks out on the 


46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


surface. Thompsonia, unlike Sacculina, appears to do no specific 
damage to its host, so that it can continue its parasitic life and indefi- 
nitely repeat its reproductive processes. The inoculation of the host 
by the free-swimming cypris has not been observed. 

The Thompsonia-infested crab presents one of the most curious 
anomalies in the whole realm of nature. Here are two crustaceans, 
one inside the other, the crab a highly developed arthropod, the para- 
site, a crustacean relative of the crab, spread out inside the latter in 
the form of a network of filaments. Both host and parasite are adult 
animals, each being the reproductive stage of its species. Progressive 
and regressive evolution could hardly reach a greater degree of 
divergence. 

Thompsonia is known to be a crustacean because it produces free- 
swimming cypris larvae, it is known to be a rhizocephalan because of 
its likeness to Sacculina, and Sacculina is known to be a cirriped 
because of the character of its nauplius. The barnacles and the 
rhizocephalans have in common the habit of attaching themselves to 
a support by the antennules in the cypris stage. From this point on 
they widely diverge. It would be highly interesting to know how the 
Sacculina larva learned to attach itself at the base of a hair on a crab, 
how it acquired the urge to get into the crab, and how it ever de- 
veloped a self-reducing method for doing it. Halfway measures 
would be useless. Clearly there are problems in evolution for which 
natural selection does not offer a ready solution. 


ISOPODA 


Most of the Malacostraca are too large to be parasites. The ma- 
jority are predatory, and few of them exhibit any considerable degree 
of metamorphosis. Most of them, moreover, hatch at a later period 
of development than do the Entomostraca, and some of them are al- 
most completely epimorphic. A prominent exception to the general 
free mode of life, however, occurs among the isopods, a few species 
of which have adopted parasitism, and have become structurally 
adapted to a parasitic life in a degree equal to that of some of the 
entomostracans. This fact shows how readily metamorphosis can 
crop out independently in species that have adopted a new way of 
living. 

The isopods in general are a conservative group in which the young 
hatch at a late stage of development with complete body segmentation 
and most of the appendages present. Among those that have become 
parasitic, however, varying degrees of adaptive metamorphosis occur 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 47 


in the life history. Species that feed temporarily on the host only 
during the larval stages may undergo but little structural adaptation. 
On the other hand, species that are permanently parasitic are likely 
to go through a high degree of metamorphosis both in the larval and 
the adult stages. The two species described in the following pages, 
one belonging to the Gnathiidea, the other to the Epicaridea, may be 
taken to illustrate the two extremes of parasitic metamorphosis found 
among the isopods. 

Paragnathia formica (Hesse).—This isopod, parasitic in its larval 
stage on fishes, gives us a good example of a parasite that undergoes 
but a minimum of metamorphic adaptation to life on its host. The 
developmental life history of Paragnathia formica has been amply 
described by Monod (1926) and the following account with accom- 
panying illustrations (fig. 18) is taken from Monod’s work. 

The adult males and females live together in small burrows exca- 
vated in semihard mud banks of stillwater estuaries below the mean 
level of the ocean. Here the pregnant females in late summer or early 
fall give birth to active larvae. The newborn larvae leave the burrow, 
swimming with great speed by movements of the abdomen. Once in 
the open water they lose no time in attaching themselves to a fish; 
most any fish will do. The time between birth and attachment is a 
period of dispersal, during which the larva takes no food, subsisting 
on the remains of yolk in its alimentary canal. The larva attaches itself 
on the fish with its second maxillipeds, and the attack is made at any 
place that will readily yield blood, such as the membrane between the 
rays of a fin, the gills, or the mouth. 

The swimming larva (fig. 18 A) is a fully segmented young isopod 
with large compound eyes and a complete equipment of appendages. 
In its embryonic development it has been provided in advance with 
efficient piercing mouth parts and a sucking apparatus. The mouth 
parts (G) are enclosed in a large conical proboscis projecting forward 
from the head, formed of the epistome (Epst) above and the first 
maxillipeds (rMxpd) below. The long, strongly toothed mandibles 
(Md) are but little movable ; they serve as harpoons to hold the para- 
site close to the fish while the sharp-pointed, freely movable first 
maxillae (1M), supported by the paragnaths (Pgn) beneath them, 
puncture the integument. The much reduced second maxillae (2/7) 
have no recognized function in feeding. 

When the young larva (fig. 18 A) has once established itself on a 
fish and has begun to feed on the blood of the host, its form changes ; 
the change is said by Monod to be effected without the intervention 
of a moult. The thorax lengthens, accompanied by a swelling of the 


48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 






x 


‘ 
‘ 


Pgn. 1Mxpd 1Mx SMx ( 


Fic. 18.—Isopoda: Gnathiidea. Paragnatha formica (Hesse) (from Monod, 
1926). 

A, first free-swimming larva. B, second form of larva parasitic on a fish. 
C, attitude of feeding larva. D, adult male. E, adult female. F, head of adult 
male, dorsal. G, section of larval head showing piercing mouth parts. 

Epst, epistome; Md, mandible; 1M+x, first maxilla; 2Mx, second maxilla; 
M-xpd, maxilliped; Pgn, paragnath. 





NO. 10 CRUSTACEAN METAMORPHOSES—SNODGRASS 49 


last three segments (B) ; the segmental limits disappear owing to the 
unfolding of the previously deeply infolded intersegmental mem- 
branes. This is the feeding stage of the parasite (B, C), called the 
pranize by Monod (L., prandium, lunch). Its meal lasts about six 
months. 

At the end of winter or the beginning of spring the fully fed para- 
sites leave the host and return to the bank of the estuary. The males 
individually dig burrows or take possession of empty ones in advance 
of the coming of the females. The completed burrows are 1.5 to 
2.5 cm. in depth, sloping downward from the mouth to an inner 
chamber 4 or 5 mm. in diameter. When the females arrive they enter 
burrows already inhabited by a male; as many as IO or more may 
consort with a single male. Within the burrows both the male and 
the females undergo their first and only moult, accompanied by a 
small degree of metamorphosis. The cuticle splits crosswise over the 
thorax, and the two ends are cast off separately. The sexes are now 
differentiated and the isopods enter their third functional stage, which 
is that of reproduction. The male (fig. 18D) retains a relatively 
slender figure, but the female (E) becomes greatly distended with 
the development of the ovaries. The mouth parts of both sexes are 
reduced, except the mandibles of the male (F), which are long prongs 
perhaps used for digging or for holding the female in mating. Sub- 
sistence is now at the expense of the food consumed during the para- 
sitic stage. 

The eggs develop into mature larvae within the ovaries of the fe- 
male, which become distended into a pair of large, saclike uteri, com- 
pressing the empty alimentary canal between them. On the ventral 
surface of the female’s thorax are several pairs of small overlapping 
oostegite plates, and above them is a large atrial cavity, into which the 
oviducts open, but this cavity does not serve as a brood chamber. 
When the young issue from the uteri through the oviducts into the 
atrium, the oostegites open and the larvae precipitate themselves head 
first through the aperture directly into the water, where they at once 
begin active swimming. After giving birth to the young, the females 
quickly die, but the males are longer lived and their metamorphosis 
is not so closely correlated with the season. 

There is clearly in the life history of Paragnathia formica little that 
can be called a true metamorphosis. The change of form between the 
two larval phases is merely a distention and elongation of the thorax 
resulting from the unfolding of the intersegmental membranes. The 
metamorphosis at the moult to the adult stage involves principally a 
reduction of the mouth parts which are no longer used for feeding. 


50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


Since most isopods have biting and chewing mouth parts, the con- 
version of the mouth parts in the embryo of Paragnathia into piercing 
organs may be regarded as an embryonic metamorphosis preparing the 
future larva for its prospective life as a parasite. 

Danalia curvata Fraisse-——This isopod belongs to the suborder 
Epicaridea, the members of which are parasitic on other crustaceans. 
It gives us an example of the sex versatility of some of the epicaride- 
ans in which the animal is first a functional male and then a functional 
female. In its female role Danalia curvata attaches itself to a crab 
infested with a rhizocephalan and feeds either on this parasite in its 
external state or on its roots in the host. Here the female is in- 
seminated by a young free-swimming male, after which the male 
attaches to the crab and becomes a female. In this manner, though 
the species is hermaphroditic, it avoids self-fertilization. The follow- 
ing outline of the life history of Danalia curvata is taken from the 
work of G. Smith (1906) and of Caullery (1908). 

The mature female (fig. 19 H) has no likeness whatever to a 
crustacean; she is little more than a sac full of eggs attached to the 
crab by a narrow stalk inserted into the crab’s body. The young on 
hatching leave the brood pouch of the mother and become free-swim- 
ming larvae. At this stage the larva (A) is recognizable as an im- 
mature isopod, and is called a microniscus. The larva is distinctly 
segmented, has two pairs of antennae, five pairs of thoracic append- 
ages, and five pairs of pleopods, but eyes are absent and the mouth 
parts are reduced to a pair of styliform mandibles enclosed in a small 
buccal cone. The microniscus larva may adopt a copepod as a tempo- 
rary host, as do most of its relatives. After several moults it takes 
on a different form (B, C) and is now termed a cryptoniscus, pre- 
sumably because its isopod characters are less evident. The body is 
more elongate and eyes have been developed, the appendages are 
retained ; the cryptoniscus is a free-swimming stage. Within its body 
is a pair of large hermaphroditic sex organs (B), each of which con- 
tains in its anterior end a small ovary (Ov) and in its posterior part 
a large testis (Tes). The testes rapidly develop and become filled with 
an abundance of spermatozoa. The larva is now a functional male. 

The male cryptoniscus seeks out a crab parasitized by a sexually 
mature female of his own species (G). After accomplishing the 
insemination of the female the larval male attaches himself to the 
crab or to the Sacculina on the crab by the first two pairs of his 
chelate pereiopods. Then a moult takes place, the cuticle being shed 
in two pieces from the opposite ends of the body, and it is then seen 
that the larva has undergone a radical change of structure within the 


NO. I0 CRUSTACEAN METAMORPHOSES—SNODGRASS 51 


cryptoniscus cuticle. The body has become a small cylindrical sac 
(D) about one and a quarter millimeters in length in which all trace 
of segmentation has disappeared. The eyes are gone, and all the 





Fic. 19.—Isopoda: Epicaridea. Life history of Danalia curvata Fraisse. (A, C-H 
from Caullery, 1908; B from G. Smith, 1906.) 

A, first instar larva. B, second free larval stage, with hermaphroditic sex 
organs containing small ovaries and large testes. C, larva with testes fully 
developed. D, parasitic larva on crab. E, same, with proboscis elongated. F, 
functional female stage, with testes degenerated, ovaries fully developed. G, 
female containing brood sac. H, female in final stage. 

Mth, mouth; Ov, ovary; Prb, proboscis; 2Prpd, second pereiopod; Tes, testis. 


appendages have been cast off with the exuviae except a pair of small 
hooklike second pereiopods (2Prpd) with which the parasite main- 
tains its hold on the host. A small conical proboscis (Prb) bears the 
mouth on its end. The testes, now that they have performed their 


52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


function, degenerate and the ovaries begin to develop, so that the 
former male larva thus changes functionally to a female. 

As a female, the parasite begins to increase in size. First the 
proboscis elongates (fig. 19 E) and, though it is armed with only a 
pair of minute teeth, it penetrates the integument of the crab. Inside 
the host the proboscis stretches out to a long neck (F, Prb) until the 
mouth (Mth) at the end comes in contact with the roots of the 
Sacculina, and four diverging processes grow out around the mouth 
to anchor the proboscis in the tissues of the crab. The body of the 
newly feminized individual then takes on a saclike form (G). The 
ovaries (Ov) are now fully developed; the oviducts open on two 
pairs of ventral papillae. At this stage the female is inseminated by 
a cryptoniscus larva still in the male phase of development (C). The 
fertilized eggs are discharged into a large incubation chamber be- 
neath the cuticle of the female. The process of forming the chamber 
is somewhat complex as described by Caullery, but essentially it ap- 
pears that two lateral ingrowths of the ventral ectoderm extend 
inward around the sides of the body, and eventually close over the 
orifices of the oviducts. When the eggs are discharged into the incu- 
bation chamber, the female ceases to feed, doubles on herself in the 
form of a U (H) and becomes a mere inert sac in which the eggs 
complete their development. 


EUPHAUSIACEA 


The Euphausiacea and some of the Penaeidae are exceptional 
among the Malacostraca in that they are hatched as nauplii. They are 
both marine and entirely pelagic. The euphausiids go through many 
moults before reaching the adult stage. Students of the group com- 
monly distinguish five immature stages in the life history of an indi- 
vidual. The first two are the nauplius and the metanauplius, the fol- 
lowing three stages are termed the calyptopis, the furcillia, and the 
cyrtopia. These forms, however, are merely stages of growth charac- 
terized by different degrees of differentiation toward the adult struc- 
ture (fig. 20 A-G). Except for the successive specialization of differ- 
ent groups of appendages for swimming there are few metamorphic 
changes involved in the development. The following condensed ac- 
count of the typical life history of a euphausiid species is based on 
the papers by Heegaard (1948) and Lebour (1925), with illustrations 
taken from both. The order includes only a single family, the 
Euphausiidae. 

The newly hatched euphausiid larva (fig. 20 A) is a typical nauplius 
of simple form with the usual three pairs of appendages, a simple 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 53 


median eye, and a large labrum. The metanauplius (B) acquires three 
additional pairs of appendages, which are the first and second maxillae 
and the first pair of legs (rL), or maxillipeds. The mandibles (Md) 


we ite 





Fic. 20.—Euphausiacea. Life-history stages. (A-E from Heegaard, 1948; F, G 
from Lebour, 1925.) 


A, Meganyctiphanes norvegica Sars, nauplius. B, same, metanauplius. C, 
same, first calyptopis instar. D, same, third calyptopis instar. E, same, first 
furcillia instar. F, Nyctiphanes couchii Bell, last (12th) furcillia instar. G, 
Meganyctiphanes norvegica, first cyrtopia instar. 


have become jawlike. The metanauplius is followed by the calyptopis 
stage, which at an early instar (C) is characterized by the distinct 
development of the carapace and the elongation of the abdomen. The 
median eye is replaced by sessile rudiments of compound eyes con- 
cealed beneath the carapace. The appendages are those of the meta- 
nauplius. At a later calyptopis instar (D) the abdomen has become 


( », 


54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


segmented and the uropods are developed. In the furcillia stage 
(E, F) the larva begins to resemble the adult. The eyes are now 
stalked and project from beneath the carapace. The first furcillia 
instar (E) has still only the appendages of the metanauplius, but after 
the first moult the pereiopods appear as simple papillae, which later 
enlarge (F, G) and finally become biramous appendages. At the same 
time the pleopods are formed. According to Lebour (1925) in 
Nyctiphanes and Meganyctiphanes there are 12 furcillia instars sepa- 
rated by moults. In the final cyrtopia stage (G), after 8 to 13 moults 
according to the species, the young euphausiid acquires the adult 
structure with a complete set of appendages and luminescent organs. 


DECAPODA 


The decapod crustaceans include the shrimps, lobsters, crayfishes, 
and crabs. None of them exhibits any pronounced metamorphic 
changes during development or in the adult stage, but most of them 
go through stages of growth characterized chiefly by the successive 
development of sets of appendages. Only in the Penaeidae is there 
a free nauplius and a metanauplius. Most species hatch in a form 


.,called a zoea, in which the appendages following the second maxilli- 
~~. peds are as yet undeveloped or are present as rudiments. With the 


functional completion of the pereiopods the larva is known as a mysis 
from its fancied resemblance to a member of the Mysidacea. Some 
species, however, go through the zoea stage in the egg and hatch as 
a mysis, and a few are almost completely developed in the adult form 
on leaving the egg. 

The decapod larvae are free swimming, and in general are fairly 
uniform in structure with a fully developed carapace and a long seg- 
mented abdomen. A few, however, take on unusual forms. Among 
the Sergestidae many of the larvae are characterized by a great de- 
velopment of long, often profusely branched spines on the thorax and 
abdomen. The rounded carapace of the palinuran Polycheles larva 
looks like a spiny burr, and others of the same group, known as 
phyllosome larvae, are broad, flat, and leaflike in shape. Presumably 
such forms are adaptations to buoyancy or floating. 

The Penaeidea.—In this order the family Penaeidae is of particular 
interest because it includes the only decapods that begin life as free- 
swimming nauplii. The fact that among the Malacostraca both the 
penaeids and the euphausiids hatch from the egg as nauplii may be 
taken as evidence that primarily all the crustaceans hatched at this 
early stage of development, and that later hatching among the higher 
Malacostraca is secondary, resulting from the earlier stages being 


NO. 10 CRUSTACEAN METAMORPHOSES—SNODGRASS 55 


completed for better security in the egg. The life history of Penaeus 
setiferus (L.) is now well known from the studies of Pearson (1939) 
and Heegaard (1953), and will here be briefly reviewed from the 
papers by these two authors. The penaeid life history, moreover, will 
serve also as a good subject for a discussion of the significance of 
crustacean larval forms. 

Both the nauplius and the metanauplius of Penaeus (fig. 21 A, B) 
have long swimming appendages, but the alimentary canal is not yet 
developed and the larvae in these stages subsist on the yolk derived 
from the egg. In the metanauplius (B), however, the mandibles have 
acquired gnathal lobes on their bases, and rudiments of four pairs of 
postmandibular appendages are present, the last being those of the 
second maxillipeds (2M«pd). The metanauplius goes over into the 
zoeal stage, in which there are three instars. In the first zoea (C) 
the carapace has developed, the mandibles have become functional 
jaws, and the larva now takes its first external food. The following 
appendages have developed into biramous limbs, and the abdomen 
shows a faint trace of segmentation, but the larva apparently still 
swims by means of the antennae. In the third zoeal instar (D) the 
larva takes on something of the adult form (G). The carapace 
covers the thorax, and rudiments of the pereiopods (D, Prpds) are 
present, the abdomen is fully segmented but pleopods have not yet 
appeared, and the antennae are still the chief organs of propulsion. 
The third zoea is followed by the so-called mysis stage, which goes 
through two instars. In the first mysis (E) the pereiopods are all 
present and have long seta-bearing exopodites, which now assume 
the locomotor function, and the antennae are reduced. The abdomen 
has well-developed uropods, but pleopods are as yet absent. 

The next stage (fig. 21 F), known as the postmysis, or postlarva, 
more nearly resembles the adult. The pereiopods have lost their 
exopodites, and those of the first three pairs are chelate. Slender 
uniramous pleopods are present on the abdomen and are now the 
swimming organs as they are in the adult. In the adult (G) the 
pleopods have acquired the typical biramous structure, and a long 
filamentous flagellum arises from each second antenna. 

The life-history stages of Lucifer, as described by Brooks (1882), 
are similar to those of Penaeus, except that the larva hatches as a 
metanauplius and the animal takes on a different form in its preadult 
and mature stages. Numerous examples of the bizarre larvae of 
Sergestidae, characterized by long, branched spines on the thorax and 
abdomen, are illustrated by Gurney (1924). 

There has been much discussion among carcinologists as to whether 
or not the forms of decapod larvae have a phylogenetic significance. 


56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


Gurney (1942) has pointed out that “the larval stages of today pro- 
vide evidence for phylogeny, but indirectly,” since the ontogeny of 
an animal recapitulates the ontogeny of its ancestors. 





Fic. 21.—Decapoda: Penaeidea. Developmental stages of Penaeus setiferus (L.). 
(A-F from Pearson, 1939.) 
A, first nauplius. B, metanauplius. C, first protozoea. D, third protozoea. E, 
first mysis. F, first postmysis. G, adult. 
2M xpd, second maxillipeds ; Prpds, rudiments of pereiopods. 


It is true that ontogenetic stages of a species may represent in a 
modified way adult ancestral stages of phylogenetic evolution. The 
adult ancestry of a crustacean, however, can go back only as far as 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 57 


the primitive adult arthropod from which the Crustacea were evolved. 
Life-history stages representing adult crustacean ancestors, therefore, 
can be recapitulations only of forms that intervened in evolution 
between the primitive arthropod and the modern crustacean. 

On the assumption adopted in the early part of this paper as a basic 
concept, the primitive arthropod is presumed to have been an elongate, 
segmented animal with a pair of similar jointed appendages on each 
body segment (fig. 1C). From such a progenitor all the modern 
arthropods were evolved by special modifications, particularly of the 
appendages, according to the adopted way of living. Anaspides (D) 
may be taken as an example of a fairly generalized modern crustacean, 
but other crustaceans go through no developmental stage resembling 
Anaspides or any other form that might be intermediate between 
their adult structure and that of a primitive arthropod. The megalops 
of a crab undoubtedly represents an early crab form, but there is 
little evidence that the Crustacea in general recapitulate adult stages 
of crustacean ancestry or the adults of other species of lower rank 
in taxonomy. There is no reason to believe that the likeness of the 
“mysis” stage of the penaeid (fig. 21 E, F) to an adult Mysis is any- 
thing more than a superficial resemblance. Foxon (1936) has shown 
that the decapod larvae do not go through a typical euphausiid or 
mysid stage, and that neither the structure nor the function of the 
mysid appendages is recapitulated in other groups. The precocious 
development of the uropods before the pleopods are formed is ex- 
plained by Foxon (1934) as an adaptation to reverse movement. 

Most crustaceans develop by anamorphosis, but the anamorphic 
method of growth was established in the remote progenitors of the 
arthropods before the arthropods became arthropods. The embryo 
in the egg goes through the preanamorphic stages of its ancestors, 
and if it is hatched as a nauplius, the following ontogenetic stages 
recapitulate the anamorphic steps of precrustacean evolution. The 
larva, however, is destined to be a crustacean, it carries the genes of 
its species, and its crustacean destiny is thus stamped on it before it 
leaves the egg. Hence, from the beginning of its development the 
larva takes on crustacean characters, but the forms it assumes are 
ontogenetic and not recapitulations of adult crustacean evolution. 
When the larva is set free at a very immature stage it must be struc- 
turally adapted to the exigencies of an independent life, and it may be 
modified for a way of living that was not at all that of its ancestors. 
Thus the normal ontogenetic stages may take on metamorphic aberra- 
tions having no relation to anything in the past history of the animal 


58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


or to its own future adult stage. Such nonancestral forms are par- 
ticularly evident in parasitic species. 

Gurney (1942), referring to the progressive shift of the swimming 
function in the larva from the antennae to the pereiopods and finally 
to the pleopods, has expressed the idea that “the fundamental fact 
which determines the organization of the larva is the mode of locomo- 
tion.” However, it is to be presumed that the use of the pleopods for 
swimming was first established in the adult ancestors of such species. 
The nauplius naturally cannot swim in this ancestral manner, and 
must use what appendages it has. As the larva grows by the addition 
of segments and appendages it can more efficiently swim by making 
use of the pereiopods, and finally when the pleopods are developed it 
can swim in the adult manner. It is the progressive organization of 
the larva, therefore, that determines the mode of locomotion. 

The Macrura.—The macruran decapods are the lobsters and the 
crayfishes. The lobster, Homarus, according to S. I. Smith (1871- 
1873) undergoes its early development in the egg and hatches at a 
stage when all the pereiopods are present and are equipped with 
feathery exopodites. This first free stage of the lobster (fig. 22 A), 
therefore, corresponds with the mysis stage of Penaeus (fig. 21 E). 
In the next instar the larva increases somewhat in size, and rudiments 
of pleopods appear on the abdomen. In the third instar (fig. 22 B) 
the young lobster attains a length of 12 to 13 mm. and much re- 
sembles the adult; the chelae are well developed, the pleopods are 
biramous, but the exopodites are still present on the pereiopods. 
Smith suggests that there is probably another instar intervening be- 
tween the third and the adult when the exopodites are lost, as in the 
postmysis of Penaeus (fig. 21 F). 

The fresh-water crayfishes, Astacus and Cambarus, hatch at a later 
stage of development than Homarus, when they have practically the 
adult structure except for the lack of the first and sixth pleopods. 

The Brachyura.—The brachyurans, or “short-tailed” decapods, are 
the ordinary crabs, so named because of the small size of the abdomen, 
which in the adult is carried bent forward beneath the thorax. The 
zoeal larvae are characterized in most species (fig. 23) by the presence 
of a long dorsal spine on the thorax and by the spinelike form of the 
rostrum, the two often projecting in a straight median line from 
opposite ends of the back. Some have also lateral spines. The larva 
swims with the large first and second maxillipeds, and the spines are 
supposed to assist in directing the course of the larva in the water 
or to help keep it afloat. The spines are absent in only a few species, 
as in the genus Ebalia and in members of the Pinnotheridae. The last 


NO. I0 CRUSTACEAN METAMORPHOSES 





SNODGRASS 59 


zoea transforms into a preliminary crablike stage known as a megalops. 

The life history of the blue crab of the Chesapeake Bay, Callinectes 
sapidus Rathbun, has been studied by Churchill (1942), Hopkins 
(1944), and Sandoz and Hopkins (1944), and is typical of the de- 
velopment of most of the Brachyura. The young crab is sometimes 
hatched in a final embryonic stage called by Churchill a prezoea (fig. 
23 A). It is still enclosed in a thin, transparent, closely fitting cuticle 





Fic. 22.—Decapoda: Macrura and Brachyura. Young stages. (A, B from S. I. 
Smith, 1871-1873; C from Cano, 1891.) 
A, Homarus americanus H. Milne Edw., first larval instar, zoea. B, same, 
third instar. C, Pilwmnus, a brachyuran crab, metazoea with partly developed 
chelipeds and pereiopods. 


that covers the spines, which will be exposed at the first moult. Sandoz 
and Hopkins say that emergence in the prezoeal stage results from 
unfavorable conditions at the time of hatching. The first free larva 
is a typical crab zoea (B) about 0.85 mm. in length. It has a short, 
rounded carapace and a long, slender, segmented abdomen. The last 
appendages are the large first and second maxillipeds, the exopodites 
of which are equipped with terminal fans of long featherlike bristles. 
The sixth segment of the abdomen is still united with the telson. In 
the second zoea (C) there is no essential change of structure, but the 


60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


body and appendages have increased in size. Churchill describes five 
zoeal instars in Callinectes sapidus, but his figures of the third, fourth, 
and probably the fifth instar are said by Hopkins (1944) to be larvae 
of some other crab. The differences, however, are slight and pertain 
mostly to the number of setae on the appendages and spines on the 
abdomen. In the fifth instar (D) the last abdominal segment is sepa- 
rated from the telson and pleopods are present. About the only 





Fic. 23.—Decapoda: Brachyura. Larval stages of Callinectes sapidus Rathbun 
(from Churchill, 1942). 
A, prezoea. B, first zoea. C, second zoea. D, fifth zoea. E, megalops. 


metamorphic features of the crab zoea are the development of the 
dorsal and rostral spines and the adaptation of the maxillipeds for 
swimming. 

During the zoeal stage buds of the third maxillipeds and of the 
pereiopods appear on the thorax beneath the carapace and increase in 
length in successive instars, but they are not seen in Churchill’s 
figures (fig. 23). It seems hardly likely that the zoea shown at D of 
the figure could go over directly into the megalops (E). In the final 
zoea of other crabs, sometimes called a metazoea, the appendages be- 
hind the second maxillipeds are well developed, as shown by Cano 


NO. 10 CRUSTACEAN METAMORPHOSES—-SNODGRASS 61 


(1891) in the metazoea of Pilumnus (fig. 22C). The first and sec- 
ond maxillipeds still have the zoeal structure, but they are followed 
by the third maxillipeds and five pairs of pereiopods, of which the 
first are strongly chelate. Moreover, all these newly developed ap- 
pendages except those of the last two pairs support branchial lobes 
on their bases. Similar advanced larval instars are shown for several 
other species of Brachyura by Lebour (1928). Hence, we should 
assume that there must be in Callinectes a metazoeal instar in which 
the thoracic appendages are in a state of development that could go 
over at one moult into the appendages of the megalops. In the life 
history of the crab there is no form corresponding to the mysis stage 
of Penaeus (fig. 21 E) or that of Homarus (fig. 22 A), but the meta- 
zoea might be likened to the postmysis of Penaeus. 

The megalops (fig. 23 E) is clearly a young crab, though it is only 
a few millimeters in length. The dorsal spine of the zoea has been 
shed with the larval cuticle (fig. 22 C) and the rostrum is shortened 
to the ordinary length. The swimming maxillipeds are transformed 
into feeding organs, and the other appendages are those of the adult. 
The prominent stalked eyes give the megalops its name (“bigeye”). 
An important feature of the megalops, however, is the extension of 
the abdomen from the thorax, which suggests that the megalops 
represents an adult ancestral form of the crab before the latter 
permanently flexed its abdomen forward beneath the thorax. 

The adult crab on issuing from the cuticle of the megalops is still 
a minute creature and goes through a large number of instars before 
becoming sexually mature, after which it may continue to moult at 
intervals. The habits of adult crabs are more various than those of 
the larvae. While most adult crabs live in the ocean and crawl on the 
bottom, some of them live in the shells of mollusks, in echinoderms, 
in cavities of corals, and in tubes of worms. Others have left the 
water for the land, where they dig deep burrows in the sand above high 
water, and still others go freely inland, even invading human habita- 
tions. The famous anomuran robber crab of the South Sea Islands 
is said to climb cocoanut trees for their nuts. Regardless of their 
habits or the nature of their dwellings, however, the brachyuran crabs 
have undergone little structural adaptation. They vary in size and 
shape, in the relative size of the chelae, and in the length of their 
legs, but in general they retain the typical crab structure. Among the 
Anomura, however, a pronounced adaptive modification of the body 
occurs in the hermit crabs that live in snail shells. The carapace of 
these crabs is weak and flexible. The abdomen is a long, soft sac that 


62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


fills the cavity of the snail shell ; pleopods are present generally on the 
left side only, but the uropods are strong, recurved appendages evi- 
dently serving to secure the crab in its house. 


STOMATOPODA 


The stomatopods are an individualistic group of malacostracans 
having some relatively primitive features in combination with so many 
structural specializations that it is difficult to give them a definite 
place in taxonomy. The head of the adult animal (fig. 24 G), pro- 
jecting from beneath a small rostral lobe of the carapace, has a com- 
plex structure not found in any other crustacean. The short, narrow 
carapace covers only the gnathal region and the first four thoracic 
segments. The other four free segments of the thorax are symmetri- 
cal with the large abdomen, and appear to be a part of it except for 
the leglike appendages borne on the last three. The limbs of the first, 
third, fourth, and fifth thoracic segments are turned forward and each 
bears a small apical chela; but those of the second segment (2L) are 
huge raptorial organs in which the terminal segments are long, strongly 
toothed claws, each closing tightly against the penultimate segment, 
giving the stomatopod its likeness to the insect praying mantis (which 
is not responsible for its name). The large abdomen has five pairs of 
pleopods, and the stomatopod gills are borne on the pleopods. The 
uropods are large, biramous appendages; the telson is a broad spiny 
plate. 

The adult stomatopods are mostly littoral in their habits. Though 
they swim freely, they live principally in burrows in the sand or mud 
of the bottom. The females lay their eggs in a mass beneath the fore 
part of the body, where they are held between the raptorial legs by 
the four small chelate legs of the thorax. The eggs are carried in this 
manner until the young larvae emerge, a period said by Giesbrecht 
(1910) to last for Io or 11 weeks. 

The young stomatopods are hatched in two different larval forms, 
which seem to have no developmental relation to each other. Our 
best source of information on the larval stages will be Giesbrecht’s 
(1910) elaborate monograph on Mediterranean species. Gurney 
(1946) gives descriptions and good illustrations of various stomato- 
pod larvae, but no full account of the life history of any one species. 
Alikunhi (1952) describes and figures particularly the last-stage 
larvae of Indian species. 

The simpler first-stage larval form pertains to species of Lysio- 
squilla and Coromida, and is termed by Giesbrecht an antezoea, This 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 63 


larva (fig. 24 A) is from 2 to 2.25 mm. in length. The thorax is fully 
segmented and is entirely covered by the carapace. The abdomen 
(AbD) is either unsegmented and entirely united with the telson (Tel) 





aig 
TSN 
2Ant-f LEGA y / ff 
a {i 
s TS, pe 


2L - 





Fic. 24.—Stomatopoda. Larval stages and an adult. (A-E from Giesbrecht, 
1910; F from Alikunhi, 1952.) 

A, an antezoea larva. B, Squilla mantis Latr., first propelagic stage. C, same, 
second propelagic stage. D, same, first pelagic stage, dorsal. E, same, first 
pelagic stage, lateral. F, Squilla latreillei, last pelagic larval stage. G. Squilla 
mantis, adult male. 

Ab, abdomen; 1Ant, first antenna; 2Ant, second antenna; E, eye; 1L,2L,5L, 
8L, first, second, fifth, and eighth thoracic appendages; Tel, telson. 


in a wide, fan-shaped plate, or one or two anterior segments may be 
free. The eyes (£) are large but sessile. The first five thoracic seg- 
ments bear each a pair of small, biramous appendages (7L, 5L) used 
for swimming. The antezoeal larva is pelagic. During subsequent 


64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


stages of its growth, the abdominal segments are successively sepa- 
rated from the telson and acquire pleopods. The five thoracic ap- 
pendages lose their exopodites and take on the adult form, those of 
the second segment becoming typical raptorial fangs in the fourth 
instar. Later the appendages appear on the last three segments of 
the thorax. In the second instar the eyes are stalked. 

The other first-larval form, termed a pseudozoea (fig. 24 B), occurs 
in species of Squilla, Gonodactylus, and probably of other genera. 
The eyes in this form are stalked at hatching. The thorax is fully 
segmented, but only the first two segments bear appendages, and those 
of the second segment are raptorial fangs. The short carapace has 
small spines on its anterior and posterior angles, and leaves four 
posterior thoracic segments uncovered. The abdomen has five free 
segments, of which the first four bear pleopods, but the sixth is still 
united with the telson. 

Squilla mantis, according to Giesbrecht, goes through 10 larval 
instars. The first two live on the bottom, but after the second instar 
the larva becomes pelagic, swimming with the pleopods. In the sec- 
ond propelagic instar (fig. 24 C) there is little change from the first 
(B) except for an increase in size and a lengthening of the posterior 
carapace spines, which in the first pelagic instar (D, E) become much 
longer and widely divergent. In the third pelagic instar rudiments of 
the third, fourth, and fifth thoracic appendages appear, and become 
longer in the next stage, when also the appendages of segments six, 
seven, and eight are developed. The sixth segment of the abdomen 
becomes free from the telson in the seventh instar. The last pelagic 
larva (F) has essentially the structure of the adult (G) except for the 
large carapace, which now covers all but one of the thoracic segments. 
After about the fifth instar, Giesbrecht says, the two larval forms, 
originating with the antezoea and the pseudozoea, become struc- 
turally alike. 

The principal structural changes during the life of the stomatopod 
take place at the transformation of the larva (fig. 24 F) into the adult 
(G). Even here, however, the only essential change affects the 
carapace, which is much shortened and narrowed and loses its pos- 
terior spines. Instead of covering most of the thorax as in the larva 
(D, F) the carapace of the adult leaves the last four thoracic seg- 
ments exposed. In this respect the carapace reverts to its relative 
length in the first propelagic larva (B). It is evident, therefore, that 
the larval development of the back shield is a metamorphic adaptation 
to the pelagic life of the larva, probably to assist in keeping the larva 
afloat. The relative length of the larval carapace varies in different 


NO. I0 CRUSTACEAN METAMORPHOSES—SNODGRASS 65 


species. In some forms the last four thoracic segments are not cov- 
ered, as in the adult of Squilla (G), in others such as Squilla latreillet 
(F) only the eighth segment is exposed in the larva, while in species 
of Lysiosquilla the carapace of the last larva may cover the entire 
thorax and the first two abdominal segments. Probably these varia- 
tions in the length of the larval carapace are only differences in the 
extent to which the free posterior margin is produced beyond the 
attachment of the plate on the third or fourth segment of the adult 
thorax. Otherwise the changes during the growth of the larva are 
‘merely developmental stages of growth and have no metamorphic 
value. It is difficult even to see any functional reason for the differ- 
ences between the two larval forms on hatching. 


IV. STRUCTURE AND EVOLUTION OF 
ARTHROPOD APPENDAGES 


Inasmuch as changes in the form and function of the appendages 
are important features in the metamorphoses of Crustacea, and various 
conflicting views have been held concerning the primitive nature and 
the evolution of arthropod limbs, we must give some attention to this 
controversial subject. 

Most studies on the comparative structure of the arthropod ap- 
pendages, and deductions as to the origin and primitive form of the 
limbs give the impression that conclusions have resulted too much 
from an attempt to fit the facts into a preconceived theory. Widely 
accepted has been the idea that the primitive appendage was a 
biramous limb; and many carcinologists would derive all kinds of 
arthropod appendages from an original phyllopodial type of limb, 
such as that of the branchiopod crustaceans. 

The trilobites are among the oldest known arthropods, and, with 
respect to their appendages, they are the most generalized, since all the 
postoral limbs are fully segmented legs. The base of each leg bears a 
branched lateral process (fig. 25 A, Eppd), which, arising on the coxa, 
is clearly an epipodite and hence cannot be an equivalent of the 
crustacean exopodite, which by definition is an exite of the basipodite. 
The trilobite limb, therefore, is not “biramous” in the manner of a 
crustacean limb, and hence does not relate the trilobites to the 
Crustacea. Raymond (1920), however, explicitly states the opposite 
view. “The trilobites,” he says, “are themselves crustaceans, as is 
amply proven by their biramous appendages.” More recently, Hee- 
gaard (1947) has argued that the trilobite limb is truly biramous, in 
spite of the evident coxal position of the “exopodite,” and he further 
attempts to show that remnants of a primitive biramous structure are 


66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


to be found in various modern arthropods other than the Crustacea. 
His discussion, however, takes too many liberties with simple visible 
facts in an endeavor to fit them into a consistent scheme of structure. 
The studies of trilobite appendages by Stormer (1939) leave little 
doubt that the trilobite leg (fig. 25 A) is simply a uniramous, seg- 
mented limb with a coxal epipodite that was perhaps a gill. Stormer’s 
contention, however, that a narrow ring at the base of the coxa is a 





Fic. 25.—Examples of segmentation of arthropod legs. 


A, leg of a trilobite (from Stérmer, 1939). B, leg of Marella (adapted from 
Walcott, 1931). C, leg of Burgessia (from Walcott, 1931). D, leg of solpugid 
arachnid. E, leg of a chilopod, Lithobius. F, leg of a decapod, Cambarus. 

Crppd, carpopodite (tibia) ; Capd, coxopodite (coxa) ; Dactpd, dactylopodite 
(pretarsus) ; Eppd, epipodite; Mrpd, meropodite (femur) ; Pat, patella; Propd, 
propodite (tarsus). 


precoxal segment is questionable. The coxa of other arthropods is 
often marked by a circular groove near the base that forms an internal 
strengthening ridge giving attachment to the body muscles of the 
limb. In the trilobite leg the large coxopodite should be the movable 
basal segment of the limb and not the narrow “precoxa.” 

The idea that the primitive arthropod limb was a flat, lobulated 
appendage of the phyllopodial type has been accepted by some carcin- 
ologists regardless of the fact that the limbs of the trilobites (fig. 
25 A) and of associated fossil forms such as Marella (B) and 
Burgessia (C) are slender jointed legs, as are those of nearly all 
modern arthropods (D, E, F), including the Malacostraca (F). 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 67 


Walcott (1931), for example, in discussing his Burgess Shale fossils 
seems to accept this theory without question when he says: “The 
biramous limb of Marella, like that of the trilobite, undoubtedly 
passed through the foliaceous or multiramous limb stage in its evolu- 
tion, probably in pre-Cambrian time.” There is no disproving this 
idea, which should apply to the other arthropods as well, but such 
implicit faith in a theory is hard to understand. 

On the other hand, Raymond (1920) says the theory of the phyllo- 
pod origin of the arthropod limb “has been completely upset” by the 
finding of such “undoubted branchiopods” as Burgessia in the Middle 
Cambrian with trilobitelike legs. The same idea has been expressed 
by Heegaard (1947) in his statement that the “undoubted branchio- 
pods” found by Walcott in the Middle Cambrian having trilobite legs 
show that “it can no longer be held that the phyllopodial limbs are 
primitive.” The writer fully agrees with this conclusion, but for 
different reasons than those given by Raymond and Heegaard. Such 
fossils as Burgessia and Marella are certainly not “undoubted” 
branchiopods. Walcott (1931) says of Marella that it is a less primi- 
tive form than the Apodidae and more primitive than the trilobites, 
but is nearer to the latter than to the former. Among the Middle 
Cambrian fossils, however, is a form, Opabina regalis Walcott, par- 
ticularly studied by Hutchinson (1930), which evidently 7s an anostra- 
can branchiopod with foliaceous appendages. 

Another popular belief concerning the derivation of the arthropod 
limb, taken to support the theory of its biramous phyllopodial origin, 
is that the limb was evolved from the polychaete parapodium. Reasons 
have already been given in section I of this paper for believing that 
the annelids have only a remote connection with the arthropod pro- 
genitors. Certainly the arthropods can have no relation to modern 
polychaetes, which are highly specialized annelids and could give rise 
only to more polychaetes. The appendages of the worm, though they 
are bilobed flaps, have a lateral position on the body (fig. 26 A), and 
there is nothing in their structure having any likeness to an arthropod 
limb at any stage of its development. The parapodium bears two 
bundles of bristles supported on a pair of long internal rods giving 
attachment to muscles. Its only common feature with an arthropod 
limb is that, being a locomotor organ, it is movable forward and back- 
ward by body muscles. In short, the idea that.the arthropod append- 
ages were derived from annelid parapodia appears to be just another 
case of excessive zeal for generalization. 

Among modern wormlike animals those closest to the arthropods are 
the Onychophora; some zoologists have even included the ony- 


68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


chophorans in the Arthropoda. Though a modern onychophoran 
shows no external segmentation of the body in the adult stage, the 
segmented repetition of internal organs and the complete body seg- 
mentation of the embryo leave no doubt that the Onychophora are 


AlCnl DV 
\ / 






B 


Fic. 26.—Diagrammatic transverse sections of Nereus (A) and Peripatus 
(B), showing comparative structure of the appendages of a polychaete annelid 
and an onychophoran. 

AlCnl, alimentary canal; Com, nerve commissure; DDph, dorsal diaphragm ; 
dm, dorsal muscles; DS, dorsal sinus; Dsp, dissepiment; DV, dorsal blood 
vessel; dum, dorsoventral muscles; NC, lateral nerve cord; Nph, nephridium ; 
Npr, nephropore; Papd, parapodium; S/mGld, slime gland; vm, ventral muscles ; 
VNC, ventral nerve cord. 


fundamentally metameric animals. The body cavity is undivided by 
dessepiments, the primitive coelom is represented only by the lumina 
of the nephridia and the gonads, and the embryogeny of the Onycho- 
phora gives the key to the early embryonic development of the arthro- 
pods. The onychophoran legs have a lateroventral position on the 
body (fig. 26B) as in the arthropods, in contrast to the lateral posi- 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 69 


tion of the polychaete parapodia (A). The nephridia (B, Nph) and 
the primitive genital ducts open mesad of the leg bases suggestive of 
their openings on the coxae in many of the arthropods. Though 
modern Onychophora are terrestrial animals, there can be little doubt 
that they had aquatic ancestral relatives represented by the Cambrian 
Aysheaia of the Burgess Shale, and perhaps by the Pre-Cambrian 
Xenusion described by Heymons (1928). 

The arthropod limbs are developed on the embryo from latero- 
ventral budlike rudiments that lengthen and become segmented. We 
may therefore suppose that from the ancestral onychophorans (fig. 
1 A) a form was evolved with longer legs (B), which later, with 
sclerotization of the integument, became the jointed appendages of the 
ancestral arthropods (C). It then required a long period of Pre- 
Cambrian evolution to produce a trilobite on the one hand, and some 
ancestral form of crustacean on the other. The differentiation between 
the two groups, however, was first in the form of the body, not in 
that of the appendages, as seen in the legs of a trilobite (fig. 25 A) 
and those of Marella and Burgessia (B, C). Though there is no valid 
reason for regarding the primitive arthropod appendage as being a 
biramous limb, the crustacean appendages later acquired their charac- 
teristic biramous structure, which is usually lost in the ambulatory 
limbs (F). 

Many carcinologists hold the view that the phyllopodial type of 
limb is primitive, at least for the Crustacea, and this concept has been 
well elucidated by Borradaile (1926a, 1926b). It is supposed that the 
primitive crustacean appendage was a flat, unsclerotized lobe with a 
fringe of hairs on the mesal border. Then the inner margin was 
broken up by the development of a series of endites. Next, the limb 
became more rigid by a sclerotization of the integument, but this 
necessitated lines of flexibility that led to a system of jointing, and 
naturally the joints were formed between the endites. Thus the 
endites are explained as the precursors of the later developed limb 
segments. Finally, with the departure from the phyllopodial form and 
the suppression of the endites, some of the limbs became slender, seg- 
mented, leglike appendages. In favor of this theory it may be noted 
that in many of the branchiopod appendages there are six endites on 
the mesal margin and a free lobe at the apex (fig. 27 A,B). If all 
the parts of such a limb became segments there would be seven seg- 
ments in all, the terminal lobe becoming the dactylopodite, which gives 
the usual number of limb segments in the Crustacea generally, though 
Borradaile holds that the maximum number is nine, which would 
include the doubtful “precoxa” of the trilobite. 


7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


eZ 


| 
| 


— 


OS 





Fic. 27,—Examples of branchiopod appendages. 

A, Branchipus stagnalis, Anostraca, thoracic limb (from Claus, 1873). B, 
Branchipus serratus, male, eighth thoracic limb. C, Apus longicaudata, Noto- 
straca, second maxilliped. D, same, first maxilliped. E, same, thoracic limb 
from middle of body. F, Daphnia magna, Cladocera, third thoracic appendage 
(from Hansen, 1925). G, Estheria clarkii, Conchostraca, second thoracic limb. 
H, same, terminal segment. I, same, left limb from middle of body. 

Bspd, basipodite; Crppd, carpopodite; Capd, coxopodite; Dactpd, dactylopo- 
dite; Iscpd, ischiopodite; Mrpd, meropodite; Propd, propodite; 1-6, endites. 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS 7a 


There are two chief objections to this phyllopod theory of the origin 
of jointed crustacean limbs. First, it gives no explanation of the origin 
of the similarly jointed legs of other arthropods, except by the wholly 
unsupported assumption that they likewise were developed from 
phyllopodial limbs. Second, the ontogenetic development of the crusta- 
cean appendages themselves gives no evidence of a phyllopodial origin, 
and suggests, on the contrary, that the phyllopodium has been evolved 
from an ambulatory leg. 

The study by Heath (1924) of the postembryonic development of 
the branchiopod Branchinecta occidentalis shows very clearly that the 
limbs arise as simple, lateroventral lobes of the body segments (fig. 
3B). Instead of taking on a phyllopodial shape, the rudiments grow 
out first in a slender leglike form (C, D). On the inner margins of 
the appendages at this stage there are indentations suggestive of an 
incipient segmentation, and at the apex is a terminal lobe. Only at a 
later stage (E) do the appendages become broad overlapping flaps. 
Finally in the adult (F) the appendages have taken on the form of 
typical unsegmented phyllopodia with three large flat exites, six 
endites, and a free, independently musculated terminal lobe. Clearly, 
these appendages in their ontogenetic development undergo a meta- 
morphosis from an ambulatory leg into a phyllopodium. Though 
Heath himself did not have this phase of the subject in mind, his 
pictures speak for themselves. 

Conversely, as seen in Heegaard’s (1953) account of the post- 
embryonic stages of the decapod crustacean Penaeus setiferus, the 
rudiments of the pereiopods develop directly into legs without under- 
going any stage suggestive of a phyllopodial origin. The pereiopods 
appear during the second protozoeal stage as simple lobes on their re- 
spective body segments (fig. 28 A). In the third protozoea they take 
on a biramous structure (B), in which the protopodite, at first un- 
divided, bears a short unsegmented endopodite and a longer exopodite. 
In the second mysis stage (C) the limbs attain a fully segmented struc- 
ture by the division of the protopodite into two segments and the 
endopodite into five, with a terminal chela on each of the first three. 
The exopodites are now large seta-bearing branches of the basipodites 
used for swimming. In the postmysis (D) the pereiopods have become 
essentially uniramous by the reduction of the exopodites to small lobes, 
and the swimming function has been taken over by the pleopods. This 
condition is retained in the adult. If the pereiopods of Penaeus had a 
phyllopodial origin in their phylogeny, there is nothing to suggest it in 
their ontogeny. The mouth-part appendages proceed along their own 
lines of development to serve the special functions they have assumed 


72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


as organs of feeding. The pleopods (E) and the uropods (I), on the 
other hand, appear to remain in an early stage of development repre- 
sented by the simple, unsegmented biramous stage of the pereio- 
pods (B). 

The swimming appendages of the anostracan branchiopods so regu- 
larly have six mesal lobes (figs. 3 F, 27 A, B) and a movable terminal 





Fic. 28.—Development of the pereiopods and pleopods of Penaeus setiferus (L.) 
(from Heegaard, 1953). 

A, rudiments of pereiopods on thoracic segments of second protozoea. B, 
pereiopod of third protozoea. C, pereiopod of second mysis instar. D, third 
pereiopod of postmysis. E, second pleopod of young adult. F, uropod of post- 
mysis. 

Bspd, basipodite; Cxpd, coxopodite; Endpd, endopodite; Expd, exopodite; 
Prpds, pereiopod rudiments ; Prtpd, protopodite. 


lobe as to suggest that the six endites represent the first six segments 
of a leg (coxopodite to propodite) and the independently musculated 
apical lobe the dactylopodite. Yet, the sixth endite is commonly inter- 
preted as the endopodite and the apical lobe as the exopodite. In the 
notostracan Apus, however, the second maxilliped (fig. 27 C) is a 
seven-segmented leg ending with a clawlike dactylopodite (Dactpd) 
and having an endite on each of the other segments except the 
ischiopodite. The first maxilliped of Apus (D) is somewhat simpli- 


NO. I0 CRUSTACEAN METAMORPHOSES—SNODGRASS 73 


fied, but the swimming appendages (E) clearly retain the structure 
of the second maxilliped. In other branchiopods the appendages may 
be variously reduced (F, G, H, 1) obscuring the basic leg structure. 

The segmentation of the arthropod legs is surprisingly constant ; 
variations result from the elimination of segments, seldom from addi- 
tion, though the propodite (tarsus) is generally rendered flexible by 
subdivision. If all the podomeres in the legs of the trilobite (fig. 
25A) and Marella (B) are true musculated segments, the ancient 
arthropods had eight limb segments, including a small apical dactylopo- 
dite, or pretarsus, and thus possessed all the segments that are present 
in any of the legs of modern arthropods. Among the latter, eight 
segments are present in the Pycnogonida and in some of the legs of 
the arachnid Solpugidae (D), but in most of the arachnids the leg 
has only seven segments by the elimination of the third segment from 
the base. The segment beyond the knee bend (D, Pat), which is the 
fifth segment in the trilobite leg (A), is called the patella, though it 
might appear to correspond with the carpopodite (tibia) in the leg 
of a centipede (E) or a decapod (F). Yet there are three segments 
beyond it in the spider leg, and only two in the other arthropods. In 
the latter, therefore, either two original segments in the distal part 
of the leg are united, or one has been eliminated. The legs of the 
chilopods and the decapods (E, F) have seven segments; the insect 
leg has only six segments because of the apparent union of the 
ischiopodite (second trochanter, or prefemur) with the meropodite 
(femur). 

Though the primitive arthropods (fig. 1C) undoubtedly were 
aquatic, they were walking animals provided with jointed limbs, and 
probably lived on plants in shallow water near the shore. Their habits 
may have been similar to those of the modern Anaspides (D). The 
typical jointed ambulatory leg has been retained in all modern arthro- 
pods, except in those crustaceans in which it has been modified for 
swimming, but even the phyllopodium preserves evidence of the seven- 
segmented structure of a walking leg. It would appear that the primi- 
tive arthropods had more legs than they needed for walking, and 
because of this fact their descendants have been able to reconstruct 
many of them into the great variety of appendicular organs possessed 
by modern forms. The arthropods owe what they are, as well as their 
name, to their jointed appendages. 


REFERENCES 
ALIKUNHI, K. H. 
1952. An account of the stomatopod larvae of the Madras plankton. Rec. 
Indian Mus., vol. 49, pp. 239-310, 25 figs. 


74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Buiss, Dororuy E., and WEtsH, J. H. 

1952. The neurosecretory system of brachyuran Crustacea. Biol. Bull. 

vol. 103, pp. 157-169, 9 figs. 
BorrRADAILE, L. A. 

1926a. Notes upon crustacean limbs. Ann. Mag. Nat. Hist., ser. 9, vol. 17, 
Pp. 193-213, 5 pls. 

1926b. On the primitive phyllopodium. Ann. Mag. Nat. Hist., ser. 9, vol. 18, 
pp. 16-18. 

Brooks, W. K. 

1882. Lucifer: A study in morphology. Philos. Trans. Roy. Soc. London, 

vol. 173, pp. 57-137, I1 pls. 
Cannon, H. G. 

1926. On the feeding mechanism of a freshwater ostracod, Pionocypris vidua 

(O. F. Miller). Journ. Linn. Soc. London, Zool., vol. 36, pp. 325- 
335, 5 figs., 1 pl. 

CANo, G. 

1891. Sviluppo postembrionale dei Cancridae. Bull. Soc. Ent. Italiana, 

vol. 23, pp. 146-158, 2 pls. 
Capart, A. 

1948. Le Lernaeocera branchialis. La Cellule, vol. 52, pp. 159-212, 22 text 

figs., 7 pls. 
CAULLERY, M. 

1908. Recherches sur les Liriopsidae, Epicarides cryptonisciens parasites des 
Rhizocéphales. Mitt. Zool. Stat. Neapel, vol. 18, pp. 583-643, 8 figs., 
pl. 26. 

CHURCHILL, E. P. 

1942. The zoeal stages of the blue crab, Callinectes sapidus Rathbun, Chesa- 

peake Biol. Lab., Publ. No. 49, 25 pp., 4 pls. 
CLaus, C. 

1863. Die freilebenden Copepoden. 230 pp., 37 pls. Leipzig. 

1873. Kenntnis des Baues und der Entwicklung von Branchipus stagnalis 
und Apus cancriformis. Abh. Konig. Ges. Wiss. Gottingen, vol. 18, 
pp. 1-48, 8 pls. 

DELAGE, Y. 

1884. Evolution de la Sacculine (Sacculina carcini Thomps.). Arch. Zool. 

Exp. Gén., sér. 2, vol. 2, pp. 417-736, pls. 22-30. 
DretricH, W. 

1915. Die Metamorphose der freilebenden Siisswasserkopepoden. I. Zeit- 

schr. wiss. Zool., vol. 113, pp. 252-324, 19 figs. 
DoocuHin, H. D. 

1951. The morphology of Balanus improvisus Darwin and Balanus amphi- 
trite niveus Darwin during initial attachment and metamorphosis. 
Bull. Marine Sci. Gulf and Caribbean, vol. 1, pp. 15-30, 7 figs. 

EcHALIER, G. 

1954. Recherches expérimentales sur le rdle de “l’organe Y” dans la mue de 
Carcinus moenas (L.), Crustacé Décapode. Compt. Rend. Acad. 
Sci. Paris, vol. 238, pp. 523-525. 

FisHeEr, W. K. 

to11. Asteroidea of the North Pacific and adjacent waters. U.S. Nat. Mus. 

Bull. 76, Pt. 1, 419 pp., 122 pls. 


NO. IO CRUSTACEAN METAMORPHOSES—SNODGRASS | 75 


Foxon, G. E. H. 

1934. Notes on the swimming methods and habits of certain crustacean 
larvae. Journ. Marine Biol. Assoc. United Kingdom, vol. 19, 
pp. 829-850, 7 figs. 

1936. A note on recapitulation in the larvae of the decapod Crustacea. Ann. 
Mag. Nat. Hist., ser. 10, vol. 18, pp. 117-123, 4 figs. 

FRANSEMEIER, LUISE. 

1939. Zur Frage der Herkunft des metanauplialen Mesoderms und die Seg- 
mentbildung bei Artemia salina Leach. Zeitschr. wiss. Zool., 
vol. 152, pp. 439-472, 17 figs. 

Gabe, M. 

1953. Sur l’existence, chez quelques Crustacés Malacostracés, d’un organe 
comparable a la glande de la mue des Insectes. Compt. Rend. Acad. 
Sci. Paris, vol. 237, No. 18, pp. 1111-1113. 

GIESBRECHT, W. 

1910. Stomatopoden, Ist part. Fauna Flora Golfes Neapel, Monogr. 33, 

239 pp., 12 figs., 11 pls. 
Gurney, R. 

1924. Decapod larvae. British Antarctic (Terra Nova) Expedition, 1910. 
Zool., vol. 8, No. 2, pp. 37-202, 78 figs. 

1926. Life-history problems of crustacean larvae. Discovery Reports, vol. 7, 
PP. 350-353, 7 figs. 

1942. Larvae of decapod Crustacea. 306 pp., 122 figs. Ray Society, London. 

1946. Notes on stomatopod larvae. Proc. Zool. Soc. London, vol. 116, pp. 
133-175, 14 figs. 

Hansen, J. H. 
1925. Studies on Arthropoda. II. Crustacea. 176 pp., 8 pls. Copenhagen. 
Heatu, H. 

1924. The external development of certain phyllopods. Journ. Morph., 

vol. 38, pp. 453-483, 5 pls. 
HEEGAARD, P. 

1947. Contribution to the phylogeny of the arthropods. Copepoda. Skrif. 
Univ. Zool. Mus., Kgbenhavn, vol. 8, 236 pp., 27 pls. 

1948. Larval stages of Meganyctiphanes (Euphausiacea) and some general 
phylogenetic remarks. Medd. Komm. Danmarks Fiskeri og Havun- 
ders6gelser, ser. Plankton, vol. 5, No. 1, 27 pp., 4 pls. 

1951. Antarctic parasitic copepods and an ascothoracid cirriped from brittle- 
stars. Vid. Medd. Dansk Naturh. Foren., vol. 113, pp. 171-190, 
7 figs., 2 pls. 

1953. Observations on spawning and larval history of the shrimp, Penaeus 
setiferus (L.). Publ. Inst. Marine Sci., vol. 3, pp. 74-105, 133 figs. 

Heymons, R. 

1928. Uber Morphologie und verwandtschaftliche Beziehungen des Xenusion 
auerswalde Pomp. aus dem Algonkium, Zeitschr. Morph. und Okol. 
Tiere, vol. 10, pp. 307-329, 7 figs. 

Hinton, H. E. 

1946. Concealed phases in the metamorphosis of insects. Nature, vol. 157, 

PP. 552-553, I fig. 


76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


Hopkins, S. H. 
1944. The external morphology of the third and fourth zoeal stages of the 
blue crab, Callinectes sapidus Rathbun. Biol. Bull., vol. 87, pp. 145- 
152, 2 pls. 
Hutcuinson, G. E. 
1930. Restudy of some Burgess Shale fossils. Proc. U.S. Nat. Mus., vol. 78, 
art. II, pp. I-24, 5 figs., I pl. 
Iwanorr, P. P. 
1928. Die Entwicklung der Larvalsegmente bei den Anneliden. Zeitschr. 
Morph. und Okol. Tiere, vol. 10, pp. 62-161, 40 figs., 7 pls. 
KEsLING, R. V. 
1951. The morphology of ostracod molt stages. Illinois Biol. Monogr., 
vol. 21, 324 pp., 34 figs., 96 pls. 
KwnipowItTscu, N. 
1890. Dendrogaster astericola nov. g. et sp., eine neue Form aus der Gruppe 
Ascothoracica. Biol. Centralb., vol. 10, pp. 707-711, 3 figs. 
KNow Les, F. G. W. 
1953. Endocrine activity in the crustacean nervous system. Proc. Roy. Soc. 
London, ser. B, vol. 141, pp. 248-267, 8 figs., 3 pls. 
Lespour, Marie V. 
1916. Stages in the life history of Calanus finmarchicus (Gunnerus), experi- 
mentally reared by Mr. L. R. Crawshay in the Plymouth laboratory. 
Journ. Marine Biol. Assoc. United Kingdom, vol. 11, pp. 1-17, 5 pls. 
1925. The Euphausiidae in the neighborhood of Plymouth. II. Nyctiphanes 
couchi and Meganyctiphanes norvegica. Journ. Marine Biol. Assoc. 
United Kingdom, vol. 8, pp. 810-829, 9 pls. 
1928. The larval stages of the Plymouth Brachyura. Proc. Zool. Soc. 
London, 1928, pp. 473-560, 5 figs., 16 pls. 
Mataguin, A. 
1901. Le parasitisme €évolutif des Monstrillides (Crustacés Copépodes). 
Arch. Zool. Exp. Gén., sér. 3, vol. 9, pp. 81-232, 8 figs., 7 pls. 
Manton, S. M. 
1928. On the embryology of a mysid crustacean, Hemimysis lamornae. 
Philos. Trans. Roy. Soc. London, ser. B, vol. 216, pp. 363-463, 
32 figs., 5 pls. 
1934. On the embryology of the crustacean Nebalia bipes. Philos. Trans. 
Roy. Soc. London, ser. B, vol. 223, pp. 163-238, 17 figs., 7 pls. 
Monon, T. 
1926. Les Gnathiidae. Mém. Soc. Sci. Nat. Maroc, vol. 13, 668 pp., 275 figs., 
I pl. 
Norman, A. M. 
1913. Synagoga mira, a crustacean of the order Ascothoracica. Trans. 
Linn. Soc. London, Zool., vol. 11, pp. 161-166, 3 pls. 
OEHMICHEN, A. 
1921. Die Entwicklung der dusseren Form des Branchipus grubei Dyb. 
Zool. Anz., vol. 53, pp. 241-253, 18 figs. 
Oxana, Y. K. 
1938. Les cirripédes ascothoraciques. Trav. Stat. Zool. Wimereux, vol. 13, 
pp. 489-512, 18 figs., 1 pl. 


NO. I0 CRUSTACEAN METAMORPHOSES—SNODGRASS 77 


Ovip (Pustius Oviprus Naso). 

A.D. 7. Metamorphoses. 

PANOUSE, J. B. 

1946. Recherches sur les phénoménes humoraux chez les Crustacés. Ann. 

Inst. Océanographique, vol. 23, pp. 65-147, 49 figs. 
Fassano, L. M. 

1953. Neurosecretory control of molting in crabs by the X-organ sinus 

gland complex. Physiol. Comp. et Oecol., vol. 3, pp. 155-189, 6 figs. 
PEARSON, J. C. 

1939. The early life histories of some American Penaeidae, chiefly the com- 
mercial shrimp Penaeus setiferus (Linn.). U.S. Bur. Fisheries 
Bull. 30, 73 pp., 67 figs. 

PEDASCHENKO, D. 

1898. Embryonalentwicklung und Metamorphose von Lernaea branchialis L. 
Trav. Soc. Imp. Nat. St. Pétersbourg (Trudy), vol. 26, livr. 4, 
No. 7, 307 pp., 6 pls., German text, pp. 247-307. 

Ports, F. A. 

1915. On the rhizocephalan genus Thompsonia and its relation to the evolu- 
tion of the group. Pap. Dept. Marine Biol. Carnegie Inst. Wash- 
ington, vol. 8, pp. 1-32, 12 figs., 2 pls. 

Raymonp, P. E. 

1920. Phylogeny of the Arthropoda with especial references to the trilobites. 

Amer. Nat., vol. 54, pp. 398-413. 
RUNNSTROM, S. 

1924-1925. Zur Biologie und Entwicklung von Balanus blanoides (Linné). 

Bergens Mus. Aarb., 1924-25, No. 5, 46 pp., 22 figs. 
Sanpoz, Mitprep, and Hopkins, S. H. 

1944. Zoeal larvae of the blue crab Callinectes sapidus Rathbun. Journ. 

Washington Acad. Sci., vol. 34, pp. 132-133. 
SCHEERER-OSTERMEYER, ELSE. 

1940. Beitrag zur Entwicklungsgeschichte der Siisswasserostrakoden. Zool. 

Jahrb., Anat., vol. 66, pp. 349-370, 5 figs. 
SCHREIBER, ERNA. 

1922. Beitrage zur Kenntnis der Morphologie, Entwicklung und Lebens- 
weise der Siisswasser-Ostracoden. Zool. Jahrb., Anat., vol. 43, 
pp. 485-538, 24 figs. 

SCHUURMANS-STEKHOVEN, J. H. 

1930. Beobachtungen zur Morphologie und Physiologie der Lernaeocera 
branchialis L., und der Lernaeocera lusci Bassett-Smith (Crustacea- 
Parasitica). Zeitschr. Parasitenkunde, vol. 8, pp. 659-6096, 28 figs. 

Scott, A. 
1901. On the fish parasites, Lepeophtheirus and Lernaea. Trans. Liverpool 
Biol. Soc., vol. 15, pp. 188-241, 5 pls. 
SmiTH, G. 
1906. Rhizocephala. Fauna Flora Golfes Neapel, Monogr. 29, 123 pp., 
24 figs., 8 pls. 
SmiTH, S. I. 

1871-1873. The early stages of the American lobster (Homarus americanus 
Edwards). Trans. Connecticut Acad. Arts and Sci., vol. 2, pp. 351- 
381, 5 pls. 


78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 131 


Snoperass, R. E. 

1938. Evolution of the Annelida, Onychophora, and Arthropoda. Smith- 

sonian Misc. Coll., vol. 97, No. 6, 159 pp., 54 figs. 
SoLLaup, E. 

1923. Recherches sur l’embryogénie des Crustacés décapodes de la sous- 
famille des ‘““Palemoninae.” Bull. Biol. France et Belgique, Suppl. 5, 
234 pp. 17 figs., 5 pls. 

Sproston, N. 

1942. The developmental stages of Lernaeocera branchialis (Linn.). Journ. 

Marine Biol. Assoc. United Kingdom, vol. 25, pp. 441-466, 5 figs. 
STORMER, L. 

1939. Studies on trilobite morphology. I. The thoracic appendages and their 
phylogenetic significance. Norsk Geol. Tidsskr., vol. 19, pp. 143-273, 
35 figs., 12 pls. 

VEILLET, A. 

1945. Recherches sur le parasitisme des crabes et des galathées par les 
Rhizocéphales et les Epicarides. Ann. Inst. Océanographique, 
vol. 22, pp. 193-341, 31 figs. 

Watcott, C. D. 

1931. Addenda to description of Burgess Shale fossils. (With notes by C. E. 
Resser.) Smithsonian Misc. Coll., vol. 85, No. 3, 46 pp., 11 figs., 
23 pls. 

WIitson, C. B. 

1905-1907. North American parasitic copepods belonging to the family 
Caligidae. Pt. 1, Proc. U.S. Nat. Mus., vol. 28, pp. 479-672, 51 figs., 
25 pls.; pt. 2, vol. 31, pp. 669-720, 19 figs., 6 pls.; pts. 3 and 4, 
vol. 33, pp. 223-490, 18 figs., 27 pls. 

1911. North American parasitic copepods. Pt. 9. The Lernaeopodidae. 
Proc. U.S. Nat. Mus., vol. 39, pp. 189-226, 18 pls. 

1915. North American parasitic copepods belonging to the Lernaeopodidae, 
with a revision of the entire family. Proc. U.S. Nat. Mus., vol. 47, 
pp. 565-720, 15 figs., 32 pls. 

1917. North American parasitic copepods belonging to the Lernaeidae, with 
a revision of the entire family. Proc. U.S. Nat. Mus., vol. 53, 
pp. 1-150, 4 figs., 21 pls. 

192I1a. New species and a new genus of parasitic copepods. Proc. U.S. Nat. 
Mus., vol. 50, pp. I-17, 7 pls. 

1921b. The North American semiparasitic copepods of the genus Clausidium. 
Proc. U.S. Nat. Mus., vol. 59, pp. 425-431, 2 pls. 

ZIEGELMAYER, W. 

1925. Metamorphose und Wachstum der Cyclopiden. Zeitschr. wiss. Zool., 

vol. 126, pp. 493-570, 20 figs., 2 pls. 





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SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 11 


(Enp oF VOLUME) 


THE VENTRAL INTERSEGMENTAL 
THORACIC: MUSCLES: OF 
COCKROACHES 


By 
L. E. CHADWICK 


Department of Entomology 
University of Illinois 











7 wre 
W; THSO oO po” F 
ST] OnN 

was HINO ee 


(PusiicaTIon 4261) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
JANUARY 15, 1957 


THE LORD BALTIMORE PRESS, INC. 
BALTIMORE, MD., U.S. A. 


THE VENTRAL INTERSEGMENTAL THORACIC 
MUSCLES OF COCKROACHES 


By L. E. CHADWICK 1 


Department of Entomology 
University of Illinois 


The cockroaches, which have persisted in much their present out- 
ward form since the Carboniferous, are admittedly rather primitive 
in many structural respects. Although one dares not take for granted 
that their musculature also retains a primitive configuration, there 
is a good a priori chance that this is the case; and comparisons with 
other primitive types, such as Grylloblatta (Walker, 1938, 1943), 
the larvae of Dytiscus (Speyer, 1922) or Cybister, and larval or 
adult Megaloptera, to my mind leave no doubt that the muscular 
pattern of blattids displays many primitive characteristics. 

In this paper attention will be called to certain of these features, 
as they are seen in the ventral intersegmental musculature of the 
thorax. This system of muscles, though far from homogeneous 
morphologically, provides a convenient segment of the total thoracic 
musculature for analysis. It may also be regarded as itself a relatively 
primitive component of pterygote anatomy, for study of the more 
recently evolved, highly specialized flying insects shows the ventral 
intersegmental muscles of the thorax gripped in an evolutionary trend 
that has already led to the drastic reduction of these muscles and that 
may ultimately result in their total disappearance. In contrast, the 
cockroaches and other less advanced forms still exhibit a wealth of 
muscles in this category, and thus afford some conception of this 
portion of the ancestral basis from which the more adept flying insects 
of today have developed. 

Descriptions of the thoracic musculature have already been pub- 
lished for three blattid species: Blatta orientalis L., by Miall and 
Denny (1886) ; Periplaneta australasiae (L.), by Maki (1938) ; and 
P. americana (L.), by Carbonell (1947). Miall and Denny purposely 
gave only a general account ; and comparisons of the reports by Maki 
and Carbonell discloses more numerous and in some instances more 


1 Formerly Chief, Entomology Branch, Chemical Warfare Laboratories, Army 
Chemical Center, Md. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 131, NO. 11 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


striking differences among the ventral intersegmental muscles than 
one would ordinarily expect from members of the same order, not to 
say genus. Since a precise knowledge of the distribution of the ventral 
muscles is essential if one is to draw from them conclusions concern- 
ing thoracic evolution, a reinvestigation of these species was made, 
in the course of which it became apparent that familiarity with a 
wider variety of blattids would be helpful. Eventually 19 species 
were examined. The discussion below attempts to extract from this 
material information that illuminates certain important facets of the 
evolution of the thorax in winged insects. 


METHOD AND MATERIAL 


Specimens from culture were etherized, pinned venter-down in a 
wax dish, and covered with water. Under binoculars, the dorsal 
integument was carefully cut away from thorax and abdomen, after 
which the ventral system of muscles was gradually exposed by re- 
moving structures that interfered with the view. A few details were 
checked from other angles. Liberal staining with 1 percent methylene 
blue from time to time in the course of dissection proved helpful, and 
brief hardening in 70 percent alcohol was occasionally resorted to. 
The magnification used was 12.5 to 50 times. At various stages of 
dissection drawings were made to scale on squared paper with the 
aid of a micrometer eyepiece. 

P. americana (L.), Blaberus craniifer (Burm.), Blatta orientalis 
L., Blattella germanica (L.), and Supella supellectilium (Serv.) 
were available in our laboratory. Cultures of the following species 
were supplied through the generosity of F. H. Babers, J. H. Fales, 
W. L. Nutting, L. M. Roth, P. R. Ruck, C. N. Smith, and E. R. 
Willis: P. australasiae (L.), P. brunnea Burm., P. fuliginosa 
(Serv.), Blaberus giganteus (L.), Blattella vaga (Heb.), Cryptocer- 
cus punctulatus Scud., Diploptera dytiscoides (Serv.), Eurycotis 
floridana (Walk.), Leucophaea maderae (Fabr.), Nauphoeta cinerea 
(Oliv.), Neostylopyga rhombifolia (Stoll), Parcoblatta pennsylvanica 
(DeGeer), and Pycnoscelus surinamensis (L.). A single preserved 
specimen of Macropanesthia rhinocerus Sauss. also was dissected. 
The 19 species investigated were chosen mainly on the basis of avail- 
ability and are but a small fraction of the more than 3,500 species of 
cockroaches that (fide Rehn, 1951) have been described. 

Nymphs and adults of both sexes were examined for most species, 
although few differences attributable to stage or sex were found 
among the msucles to which the present investigation was confined. 
For comparison, data were obtained from the literature or from the 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK 3 


writer’s dissections for representatives of a number of other orders. 
All observations cited in this paper without a statement as to source 
are from my own work. 

Each morphologically distinct muscle was given a designation 
formed by hyphenating the accepted abbreviations for the skeletal 
parts between which the muscle is stretched. Under this system, if 
an attachment is segmental, the segment is identified by an arabic 
or roman postsubscript for the thorax or abdomen, respectively, while 
the designations of intersegmental structures are preceded by the 
appropriate arabic numeral, beginning with o for the cervical inter- 
segment. Exception: the customary abbreviations rcv, 2cv . . . for 
the cervical sclerites, and 1ax, 2ax . . . for the axillary sclerites of 
the wing, the latter with segmental subscripts, are retained. Cruciate 
muscles, with origin and insertion on opposite sides of the longitudinal 
body axis, are distinguished by adding X to the usual designation. The 
skeletal abbreviations used are for the most part those given currency 
by Snodgrass (1929, and numerous other publications). 

Examples: 2sps-epss, a muscle stretched between the second (post- 
mesothoracic) spina(sternite) and the metepisternum; fus-sya, the 
longitudinal ventral muscle from the metafurca to the second abdomi- 
nal sternum ; epS-cx,X, a cruciate muscle of the procoxa, with origin 
on the contralateral mesepisternum. 

A glossary of abbreviations is given at the end of the text. 


OBSERVATIONS AND INTERPRETATION 


The ventral intersegmental muscular system of the blattid thorax 
includes elements with primary attachments on the spinae (sps) or 
on the intersegmental laterosternites (ils), as well as furcal (fw) 
muscles that run between successive segments. This report is divided 
accordingly into three main sections. 


I. THE SPINAE 


Cockroaches have two authentic spinae (isps, 2sps), and in addi- 
tion possess in the third thoracic intersegment a common junction of 
serial homologs of the more anterior spinasternal muscles that lacks 
the median connection with the integument but obviously represents 
a postmetathoracic spina. This junction (“3sps”) is attached by 
fibrous ligaments (figs. 4, 6, 7, 9, 10, 18:27) to the bases of the 
metathoracic furcal arms (fus), between which it floats above the 
nerve cord. Comparative evidence leaves little doubt that these non- 
contractile ligaments, which now usually appear as fus-fus, have been 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


derived from former muscles, 3sps-fug. A true third spina, which oc- 
curs in some Apterygota (Maki, 1938), is known among pterygote 
insects only in Grylloblatta (Walker, 1938, 1943), but an arrange- 
ment much like that of blattids has been reported for larval Dytiscus 
(Speyer, 1922) and has also been seen in larval Cybister, in Zoo- 
termopsis (fig. 6), and in larval and adult Corydaius. Other vestiges 
of the third spina and its musculature have been found but not recog- 
nized as such by several students in a number of other insects. Alto- 
gether, the facts constitute strong evidence that a third spina was 
present in the ancestral Pterygota, and probably in early hexapods 
generally. 

More thoroughly documented, since much of the testimony is still 
available in a variety of living forms, is a general tendency toward 
loss of the remaining spinae and their associated musculature during 
the post-Carboniferous evolution of the pterygote thorax. Certain 
cockroaches, however, have gone contrary to this trend, and have 
experienced a prodigious extension of the first spina and, to a lesser 
extent, the second, in the direction of the body axis. Compare, for 
example, figure 1 with figure 2, or 17 with 18. This spinal elongation 
is related, in part, to hypertrophy of the transverse spinal musculature 
(Isps-epSe, 2sps-eps3) ; and it is probably no coincidence that sev- 
eral species that manifest this development have also exceptionally 
large transverse muscles of the first abdominal intersegment, syz4- 
sna (figs. 1,8, 11, 12: 34). My judgment that these characteristics are 
secondary rests partly upon the fact that they are peculiar to some 
blattids, being unparalleled in others and absent, so far as I know, in 
any other group of insects; and partly upon the presence in the mus- 
culature of these same cockroaches of other trends away from a primi- 
tive condition, such as a tendency toward loss of certain spinacoxal 
muscles. (See c, this section, below.) 

Table 1 provides a composite list of the spinasternal muscles of 
blattids, as these are now known, and is so arranged as to indicate 
the probable serial homologies. Most species possess a very large frac- 
tion of the total complement. The relatively few exceptions are sum- 
marized in footnotes to the table, and some of them are discussed 
briefly in the text. Included in the spinasternal musculature are 
(a) transverse spinal muscles; (b) spinal muscles of the preceding 
or succeeding furca; (c) spinal muscles of the preceding or succeed- 
ing coxa; (d) muscles stretched between successive spinae; and (e) 
spinabdominal muscles. Each of these groups is discussed under the 
corresponding subheading below. 

a. Transverse spinal muscles—The transverse muscles of cock- 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK 5 


TABLE 1.—Spinasternal muscles of cockroaches 


In this composite list, muscles that appear on the same horizontal line are 
considered serially homologous. Question marks indicate uncertain homologies. 
Symbols such as M4o, Cro3 refer to the numbers given the corresponding mus- 
cles in P. australasiae by Maki (1938) and in P. americana by Carbonell (1947). 
Each of the 19 species investigated here has all muscles shown in the table, 
except as stated in the notes below or in the text. For muscles without spinaster- 
nal attachments, see table 2. 


Footnote 
number First spina Second spina “Third spina” 
Bae os Isps-rils 2sps-2ils --- 
TDs ee Isps-epse 2sps-epss — 
M4o; Cro3 M74; Cr49 
Deh asa Isps-fus - “3sps’’-fus 
MS; Cog M41 M105 (partim) 
Bardens se —- 2sps-fis — 
Cror 
meas del Isps-fua 2sps-fus ? “Osps’-sira 
aia Gros M73; Cr5z Mro4; Cig1? 
Rieraterayeite: s _ 2SPS-S1rA4 _ 
C189 
Citak. st. Isps-C%1 25ps-CH2 ? fus-cxs post. rot. (partim) 
M24, 25; Co8 M56; C134 M88, 89; C171 
Jig stevatateye ISpS-CH2 2SPS-CH3 — 
C105 C173 
Br yates 9) Isps-2sps 2SpS-3Sps ? “3sps’-ventr. diaphr. 
M39; C106 M72; Cr52 M105 (partim) 


1Cryptocercus has both 1,a and 1,b; other species 1,b only. The abdominal transverse 
muscle, $;;4-Syz4 (M112, is a serial homolog. 

2 Muscle a2sps-fuz is lacking; Maki’s record (M4r) is probably an error of transcription. 
(See text.) 

8 No comment. 

4 Carbonell (1947, p. 20) describes muscle 19r in P. americana as follows: ‘‘Oblique ven- 
tral muscle. . . . From the base of the sternal arm to the anterior edge of the first [sic!] 
abdominal sternum.” 

5 Abdominal insertion wholly or partly on s,;;4 in Eurycotis, Macropanesthia, Neostylopyga, 
and Periplaneta brunnea. (See text.) 

6 Several species have two definitive muscles, Isps-cx1, one of which is probably equivalent 
to the eps,-fu,X of other species. (See figures and text.) In listing two furcal posterior 
rotators of the third coxa, Maki (1938) suggests that one of them may be serially homologous 
with the spinal posterior rotators of the other legs. (See text for discussion.) 

7 Blaberus, Diploptera, Cryptocercus, Leucophaea, Macropanesthia, Nauphoeta, and Pyc- 
noscelus lack Isps-cx,; the last five genera also lack 2sps-cxry. 

8 The fibers from “3sps’’ to the ventral diaphragm were not found in several species, but 
were possibly destroyed during dissection in some of these. 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


roaches are regularly present in the first two thoracic intersegments 
but absent in the third, where their failure to appear is no doubt re- 
lated to the near obliteration of the ventral region of the first abdomi- 
nal segment, reduction of which is a very general and probably very 
early feature of pterygote reorganization. However, the transverse 
muscles are represented in the abdomen by the muscular attachments 
of the ventral diaphragm on the anterior lateral angles of the second 
abdominal sternum. A striking development of these abdominal 
fibers is seen in the series Pycnoscelus, Diploptera, Nauphoeta, 
Leucophaea, in which last they attain the status of a powerful trans- 
verse muscle, Sya-Syra (figs. 1, 8, 11, 12: 34). The function of such 
a muscle is unknown. Curiously, Blaberus, Cryptocercus, and Macro- 
panesthia (figs. 5, 10, 17), which share other peculiarities of the four 
genera just mentioned, do not show any tendency toward hypertrophy 
of the transverse muscle of the ventral diaphragm, and in this respect 
are more like the other species included in this study. The lateral 
suspensions of the ventral diaphragm are not evident in the abdomi- 
nal intersegments posterior to the first; and in general the ventral 
diaphragm of cockroaches is much less extensive than that of some 
other insects, e.g., phasmids and the acridid Orthoptera. 

Some authors have listed as transverse muscles structures such as 
the ligament fus-fus, whose affinities are, however, with the spina- 
furcal muscles. 

b. Spinafurcal muscles —Cockroaches all have the muscle rsps-fuy 
and “3sps”-fus, the latter represented, as a result of loss of the third 
spina, by fibrous ligaments that often appear as a single transverse 
band, fus-fus. A corresponding 2sps-fug does not occur in any blattid 
I have examined, and I believe Maki’s record of this muscle (1938, 
fig. 6, No. 41) in P. australasiae must rest on an error of transcrip- 
tion, since all cockroaches have another, larger muscle, 2sps-fu,, that 
is omitted from his figure and description. 

Miall and Denny’s (1886) statement that the muscle 2sps-fu, is 
inserted on the base of the first leg in B. orientalis is misleading, for 
the connections in B. orientalis (fig. 2: 16) are identical with those 
of other cockroaches ; but the description reflects Miall and Denny’s 
awareness of a structural difference between the prothoracic sternal 
arm and those of other segments, a distinction that seems to have 
escaped comment by most others who have investigated the muscula- 
ture of cockroaches. (See section 3, below.) 

The muscles rsps-fug and 2sps-fus are also universally present in 
blattids as is their possible serial homolog, “3sps’’-sya, which is here 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK fe 


discussed under the spinabdominal muscles. (See e, this section, 
below). 

c. Spinacoxal muscles——The first and second spinae both carry 
posterior rotators (or remotors) of the preceding coxa (Isps-c+1, 
2sps-c%2) in all blattids examined. A corresponding muscle of the 
third spina is absent as such, but may be represented, as already sug- 
gested by Maki (1938), in the muscle fus-c1s post. rot., which fre- 
quently shows signs of a dual composition. In some specimens, a few 
of the fibers of fus-cas post. rot. appear to be continuous with those 
of the ligament fis-fus. Only in larval Dytiscus (Speyer, 1922), in 
Grylloblatta (Walker, 1938, 1943), in Zootermopsis (fig. 6: 29), and 
in the larvae of Cybister and Corydalus has a distinct muscle, 3sps- 
c#s3, been found; and in these species the muscle fus-crs post. rot., 
which is also present, seems to be a single band. 

Spinal promotors of the mesocoxa and metacoxa (Isps-c%», 2sps- 
cs) also occur frequently in cockroaches, as they do in other primi- 
tive forms. However, the mesocoxal promotor is absent in Blaberus, 
Diploptera, Leucophaea, Macropanesthia, Nauphoeta, and Pycnosce- 
lus; and the last four of these genera also lack the metathoracic 
homolog. Both spinal promotors are likewise missing in Cryptocercus, 
which shares to some extent the tendency of these genera toward 
hypertrophy of the transverse muscles of the first spina although it 
differs markedly from them in certain other respects. In some other 
cockroaches, e.g., in Blattella, the spinal promotors, though present, 
are weak. Thus, the trend toward obliteration of these muscles, which 
has gone far among higher orders of insects, is evident even among 
the Blattariae. 

d. Spinaspinal muscles—The muscles rsps-2sps and 2sps-“3sps” 
were found in all the species studied here, although they are at times 
weakly developed and easily overlooked; this is particularly true of 
2sps-“3sps.” The first of these muscles is present also in most Or- 
thoptera (sensu stricto) and Mantodea, but not in other orders with 
the possible exception of Isoptera, where it was recorded by Fuller 
(1924, fig. 9, muscle n) for Termes latericius Hav. Since this muscle 
does not occur in other termites studied by Maki (1938) and the 
writer, it may be that Fuller misjudged the posterior attachment of 
his muscle n, which perhaps represents rsps-fu,, a muscle missing 
from Fuller’s account but present in all Isoptera studied by others. 

The muscle 2sps-“3sps” has so far been recorded only from blat- 
tids, where its general occurrence may be taken as one more indica- 
tion of primitive structure. What is probably a vestige of this muscle 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


has been found in the immature mantid, Tenodera sinensis Sauss. 
(fig. 7 +29). 

The spinaspinal muscles are the only portion of the longitudinal 
ventral musculature that has obviously retained its primary relation- 
ships in blattids. Both attachments are still unmistakeably interseg- 
mental. Like the other somatic muscles, the spinaspinal muscles are 
paired bilaterally ; but the right and left bands of rsps-2sps are often 
so closely appressed in the midline that they seem like a single ele- 
ment, and they have been so described by some authors. The pos- 
terior insertions of 2sps-“3sps” are usually well separated on the 
ligament ftts-fus (e.g., fig. 9: 23), and in some instances may even 
seem to be on fis at the site of attachment of the ligament. Care must 
be taken, therefore, not to confuse them with the usually better de- 
veloped spinafurcal muscles, 2sps-fus, from which they are morpho- 
logically distinct. ! 

The ligament fus-fus also serves, in several cockroaches, as a base 
of attachment for paired muscular strands that course posteriorly to 
join the meshwork of contractile fibers and membrane that consti- 
tutes the ventral diaphragm (figs. 2-5, 8, 11, 18: 30). These strands, 
“3sps’’-ventr. diaphr., may be serial homologs of the muscles /sps- 
2sps, 2sps-“3sps.” I did not succeed in finding these delicate strands 
in all species, but could not be sure, in the cases where they seemed 
absent, that I had not destroyed them. 

In pterygote insects, there is no homolog of the spinaspinal muscles 
anterior to Isps; but Maki (1938) has recorded muscles that are 
probably homologous in the prothorax of some Apterygota. 

e. Spinabdominal muscles—The spinabdominal muscles of cock- 
roaches include only 2sps-sy4, “3sps’-srra and “3sps’-ventr. diaphr. 
The last two have already been mentioned in this section, b and d, 
above. They arise on the ligament fus-fus, and not on the base of 
fus as some have stated. The muscle 2sps-syr4 is characteristic of 
blattids, and is present in all of them I have seen, though it is weak 
in Leucophaea. Elsewhere, it has been recorded only from Gryl- 
loblatta (Walker, 1938, No. z1zb). It is interesting as an example 
of a muscle more than one segment in length, a type that is of infre- 
quent occurrence in pterygote insects. In Macropanesthia, Peri- 
planeta, Eurycotis, and Neostylopyga the abdominal insertion of some 
or all the fibers is actually on syzz4. Apparently this modification 
may occur readily because 2sps-sy4 is ordinarily inserted along 
the antecosta of syy4 dorsal to the usual longitudinal bands, syz4- 
Sua, With which 2sps-s;z4 is more or less continuous. Dissolution of 
the integumental attachment at s774 adds one segment of muscle 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK 9 


to the length of 2sps-syz4; and this step, to judge by various instances 
observed, leads to an intervening stage in which the now floating 
muscle is still divided by a transverse septum at the original level of 
attachment on sya (figs. 3, 17: 21). Subsequently, all signs of the 
septum are lost. Reduction of the ventral region of the first abdomi- 
nal segment has doubtless contributed to developments of this nature, 
which are not confined to cockroaches or to the particular muscle 
in question (cf. fig. II: 32). 


2. THE INTERSEGMENTAL LATEROSTERNITES 


Intersegmental laterosternite (ils) is a term here introduced for 
sites of muscle attachment that lie at the lateral extremities of the 
ventral intersegmental folds. Such sites are believed to have been 
characteristic features of the anatomy of early arthropods in all in- 
tersegments. In existing forms, extensive modification of the original 
relationships is the rule, as will be seen below; nevertheless, recogni- 
tion of the presence and nature of these sites is helpful in understand- 
ing the manner in which the ventral musculature and associated 
structures have evolved. 

Primitively, the musculature of the ils included (1) the transverse 
muscles, which, with or without interruption by a median spina, 
stretched between the two ils of the same intersegment; (2) the out- 
ermost bands of the ventral longitudinal body musculature; (3) cer- 
tain dorsoventral muscles; and probably (4) other muscles of vari- 
ous types, some of which will be noted below. However, the original 
muscular relationships of the i/s are still not fully understood; and 
this fact, together with their varied fate in different lines of descent 
and in different parts of the body, presents the comparative morphol- 
ogist with many perplexing problems. Hence, it is not surprising that 
structures equivalent to the ils as here defined have been overlooked 
by some workers and variously named in different situations by 
others. Several have referred to them as “intersegmental pleurites,” 
a term which is unsatisfactory both because of the obvious sternal 
nature of the structures in question and because use of the name 
“pleurite” in reference to intersegmental elements is self-contradic- 
tory. Crampton (1926) avoided these objections by employing the 
term “furcilla.”” Unfortunately, this usage of “furcilla” seems likely 
to cause confusion, because the name had been applied in various 
other senses by earlier workers and because it suggests a nonexistent 
affinity with the segmental sternal apophyses, or “furcae” (fu). For 
these reasons, we have substituted the more accurately descriptive 
designation “is,” 


IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I1 


The principal primary muscular relationships of the ils are tolera- 
bly well preserved in the typical abdominal intersegment, granted that 
a secondary extension of sclerotization has here merged the in- 
tersegment indistinguishably with the following segmental sternal 
plate, of which the former intersegment now forms the antecosta 
(Snodgrass, 1929). The ils are here represented in the anterolateral 
angles of the definitive abdominal sterna, which in many insects dis- 
play the muscular relationships outlined above (Ford, 1923; Maki, 
1938). 

In the intersegments that follow each of the three thoracic seg- 
ments, the situation is rarely so transparent. One gains the impres- 
sion that, even in the most primitive forms that have come to hand, 
the musculature of the ils has already been subject to extensive shift- 
ing and reduction, while in more recent insects only a few scattered 
remnants suggest the original role of the ils as attachment sites for 
part of the longitudinal body muscles. Moreover, where the trans- 
verse muscles have been preserved, their lateral attachments now 
usually appear to be on segmental parts. In addition, we find a few 
muscles that originate on the thoracic ils or on their present equiva- 
lents, whose insertions are segmental and which have no counterparts 
in the legless abdomen. 

Equally difficult to analyze, because of the extremely varied skeletal 
and muscular relationships that exist in different groups, is the situa- 
tion in the cervical intersegment. Here one must be content for the 
present with the assurance that the former Oils are usually somehow 
represented, most often as part of one or more of the definitive cervi- 
cal sclerites. 

These problems are well illustrated in the cockroaches, in which 
the musculature of the is, though rich in comparison with that of 
more recently differentiated orders, can only be considered vestigial 
in relation to the inferred ancestral condition. 

Ventral muscles of cockroaches that appear to belong to the ils 
complex include (a) transverse muscles; (b) cruciate coxal and furcal 
muscles; (c) certain other furcal muscles; and (d) spinasternal 
muscles of the abdominal ils. (See tables 1 and 2.) 

a. Transverse muscles —The transverse muscles of the thorax ordi- 
narily have a median attachment on the spina, and have therefore 
been discussed under section I,a, above. The nature of their lateral 
attachments remains to be considered. As already noted, abdominal 
relationships support the view that the lateral attachments of the 
transverse muscles are morphologically intersegmental, i.e., on the 
ils. Comparative evidence from other arthropods and the scanty em- 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK II 


bryological data on insects (Heymons, 1895 ; Roonwal, 1937) justify 
this inference. In the postembryonic cockroach, however, the defini- 
tive connection in the thorax is usually with the anterior margin of 
the succeeding episternum, and the muscles are therefore to be desig- 
nated as Isps-eps,, 2sps-epss, even though Maki (1938, p. 58) de- 
scribes the attachment as “‘on the small sclerite before the [mes] epi- 
sternum” in P. australasiae. If this were all the evidence available, 
one would conclude that in blattids the rils and 2ils had fused with 


TABLE 2.—Ventral intersegmental muscles of cockroaches: 
muscles without spinasternal attachments 


Symbols such as M6, C55 refer to the numbers given the corresponding mus- 
cles in P. australasiae by Maki (1938) and in P. americana by Carbonell (1947). 
For muscles with spinasternal attachments, see table 1. 


Footnote 
number Muscle type 
I....Cruciate muscles Icv-C#1X €pse-Cx1X € pSe-furX 
Co7 C102 
2....Postcoxal ligaments Tils-fus 2ils-fus 3ils-fus 
3....Furcal muscles fus-tent. fus-2cv 
M6; C55 M7; C84 
fis-fue (2bands) 
M38; Croo 
fus-fus (2bands) 
M71; Cr48 


fu-sua (usually 3 bands) 
M103; C192, 193 


1The 3 cruciate muscles are not serially homologous. For discussion of shifts in origin 
of eps,-cx,X and eps,-fu,X, see text sections 2,a and 2,b. The three muscles or their equiva- 
lents are present in all species examined. 

2 The three postcoxal ligaments are serially homologous. For variations in their occurrence, 
see text section 2,c. 

8 The furcal muscles are probably all serially homologous, at least in a broad sense. For 
variations in the abdominal insertion of fus-s;74, see text section 3. Carbonell (1947) records 
the abdominal attachment of muscles number 192 as on s,, in P. americana, All the furcal 
muscles listed are present in all species investigated. 


eps, and epss, respectively. The arrangement of the cruciate muscles 
of the profurca and procoxa in P. americana and in Cryptocercus 
(see below) is also consistent with this interpretation, 
However, the full story is not that simple, for Cryptocercus pos- 
esses not only muscles Isps-eps,, 2sps-epss that are clearly homol- 
ogous with those of other blattids, but has in addition fibrous liga- 
ments (figs. 10, 13: 4, 17) that run from the spinae to small sclerites 
in the intersegmental membrane well in advance of the episternal 
margin. These transverse ligaments have evidently been derived from 
former muscles, and their lateral attachment sites not only occupy the 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


position of true ils but also carry the dorsoventral muscles typical of 
these structures. 

Furthermore, careful dissection of most cockroaches discloses a 
second series of straplike ligaments, also of muscular derivation, that 
run from the postcoxal membranes (i.e., from the intersegments) 
to the furcal arms of the respective preceding segments. These liga- 
ments, here designated sils-fuz, etc., are inserted on the furcal arms 
near the seat of the furcophragmatal muscle (see figs. 2, 10: 13, 
24,931). 

It appears, then, that in the thoracic intersegments of cockroaches 
the former i/s may now be represented by as many as three seemingly 
distinct sites: (1) the following episternum; (2) the original is; 
and (3) the origin of the postcoxal ligament. How this separation 
came about is by no means obvious. 

As already stated, the often straplike but still fibrous transverse 
ligaments rsps-rils, etc., are evidently derived from former muscles, 
and are even now represented in whole or in part by muscles in some 
species. However, again in the light of relationships found in other 
primitive groups (e.g., Dytiscus larva (Speyer, 1922), Corydalus 
larva, etc.), these transverse muscles seem to have served also (after 
loss of their contractile nature ?) as suspensory ligaments for a por- 
tion of the longitudinal ventral intersegmental musculature. Vestiges 
of this or an analogous arrangement are still present in the first 
thoracic intersegment of some cockroaches. 

In Cryptocercus, which in this respect is the most primitive blattid 
I have seen, both the ligament rsps-rils and the muscle Isps-eps2 are 
present and are, laterally, quite distinct (fig. 13: 4, 5). However, the 
mesal portion of zsps-zils, incidentally still composed of contractile 
tissue, is so confluent with the adjacent fibers of rsps-eps, that a 
separation of the two muscles in this region is hardly possible. Thus, 
the anterior portion of the muscle Isps-eps, could be described as 
“ligament-eps,.” Similarly, it is difficult to specify the origin of the 
cruciate profurcal muscle eps,fu,X (8), for part of its fibers arise on 
eps while the more ventral ones, not visible in figure 13, originate on 
the ligament rsps-zils, from which they run with the others to the in- 
sertion on the contralateral furcal apodeme fi. 

Variations of these relationships are exemplified in a number of 
other genera, viz, Periplaneta, Neostylopyga, Eurycotis, Blatta, and 
Blattella. In none of these is the peripheral attachment of the liga- 
ment rsps-rils preserved; but the central portion of the ligament is 
present and extends anterolaterally from the spina as a noncontractile 
septum on which fibers from eps, are attached and from which origi- 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK 13 


nate muscles (“epss”-fu,X, “eps,”-cv,X) that insert on the contra- 
lateral profurca and procoxa. These conditions in P. brunnea are 
illustrated in figure 14. Here it will be noted that the origin of the 
cruciate furcal muscle (8) is more central than that of the cruciate 
coxal muscle (9). In P. australasiae (fig. 15: 8) the more dorsal 
bands of the furcal muscle originate so near the midline that they 
appear to arise from the spina; and they are so recorded in Maki’s 
(1938) description. 

The contrasting arrangement of the corresponding muscles in P. 
americana is apparent in figure 16. In this species, there is no visible 
remnant of the ligament rsps-rils, and the cruciate muscles (8, 9) 
originate far laterally, on the anterior margin of eps, as they were de- 
scribed by Carbonell (1947). Hence the structure of P. australasiae 
and P. americana is superficially quite distinct. Access to intervening 
forms, such as P. brunnea, etc., shows, however, that in P. australasiae 
and P. americana we are merely confronted with rather extreme vari- 
ations in the arrangement of morphologically identical elements. 

Cryptocercus is unique among the cockroaches studied in that the 
most dorsal fibers of the transverse muscle of the first intersegment, 
Isps-epS2, continue across the body without attachment on the spina 
(fig. 13: 5). This development is almost certainly secondary, for 
the more ventral fibers of this muscle have the usual spinal con- 
nection. 

The several variants we have seen in the cruciate muscles are such 
that all of them could have been derived, by gradual transition, from 
any one chosen as a starting point. There is also at present no bar 
to the alternative assumption that any or all of them might suddenly 
have arisen de novo from each other or from an unknown basic 
pattern as a result of gene mutation or recombination. Therefore, a 
decision as to which of the existing configurations portrays the most 
primitive condition is not warranted on the basis of the evidence so 
far presented. Although Cryptocercus shows some very primitive 
features in the first thoracic intersegment, it is even here less primi- 
tive in other respects than certain other cockroaches, and should not 
be regarded as the prototype for the arrangement of the cruciate 
muscles unless independent confirmation can be produced. Other data 
that bear on this question are cited in this section, b, below. 

At the present time, it also does not seem possible to decide whether 
muscles such as Isps-rils and Isps-eps, are fundamentally distinct, or 
whether the episternal branch is no more than a hypertrophied off- 
shoot from an originally single transverse band. Both elements are 
present simultaneously in a few other insects, not all of which are 


I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I3I 


closely related to the cockroaches. An example is shown in figure 6: 
4, 5. There is some indication also, in various other primitive groups, 
that there existed muscles of the type rils-eps,; if so, these too may 
have contributed to the arrangements now seen in Dlattids, for they 
could conceivably account for the lateral portions of the cruciate 
muscles that run in some species from eps, to the transverse ligament 
1sps-tils or to the septum that has replaced it. 

b. Cruciate coxal muscles—Mention of the cruciate coxal muscle, 
epSe-cx,X (9), has been made in the preceding section. This muscle 
is inserted together with the spinal posterior rotator Isps-cx; (7), 
from which it is morphologically distinct. However, species such as 
P. australasiae, in which the origin of “eps,”’-c7,X is far mesad, could 
properly be described as having two definitive spinacoxal posterior 
rotators, as was done by Maki (1938, fig. 6, Nos. 24, 25). In most 
instances these two muscles may still be distinguished by the fact 
that the true spinal muscle originates along the side of the spina ven- 
tral to the other spinal musculature, whereas the muscle equivalent 
to epSs-cx,X has a more dorsal origin, anterior to the transverse 
muscle Isps-epss. Yet the distinction is not always clear ; and in some 
blattids one or the other of these two muscles may even have been 
lost. 

Serial homologs of the muscle epsy-cx,;X do not occur in cock- 
roaches so far as is known, but homologs with origins on the ils are 
found in all three thoracic segments of larval Dytiscus (Speyer, 
1922) and in larval Cybister. In larval Corydalus, which lacks such 
muscles in the first intersegment, cruciate posterior rotators of the 
second and third coxae originate on the corresponding furcal arms. 
This shift in origin is easily understood from the fact that the furcal 
arms are here fused with the succeeding is, evidently, as judged by 
conditions still found in some other Megaloptera such as Sialis spp., 
in consequence of sclerotization along the line of the postcoxal liga- 
ments 2ils-fus, 3ils-fus. One infers from the position of the cruciate 
coxal muscles in these primitive forms that the attachments of the 
cruciate muscles of cockroaches on Iils or eps, are more likely the 
primary ones than any of the other variants observed in blattids. If 
so, Cryptocercus and, for some strange reason, P. americana but not 
its congeners have more nearly preserved the original condition. 

Cruciate promotors of the first coxa have been described from a 
number of orders, and are apparently present in a much reduced state 
in all cockroaches, although on account of their delicacy they have 
escaped the notice of myologists. In the adult insect, which is the 
stage usually chosen for dissection, they are extremely slender and 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK I5 


transparent. They are more easily seen, though not immediately 
recognizable as muscles, in the nymph, where they were first dis- 
covered by Scharrer (1948) as the bearers of the prothoracic glands. 
The glandular tissue, which encases the tenuous contractile filament 
and thus renders it more visible (fig. 7: 3), degenerates soon after 
metamorphosis, but the muscular core persists throughout life. The 
origin is near the anterior end of the first cervical sclerite, rcv, which 
for this and other reasons is to be regarded as incorporating the cervi- 
cal ils; and the insertion is on the proximal margin of the contralateral 
coxa vera, just laterad of the coxotrochantinal articulation. Cor- 
responding cruciate promotors of the second and third coxae have not 
been identified in any pterygote insect, but there is a possibility that 
they are represented in the usual spinal promotors Isps-cx9, 2sps-ci3. 

c. Lateral furcal intersegmental muscles—In cockroaches furcal 
muscles whose origins are on the i/s or on their present equivalents in- 
clude only the cruciate muscles eps -fu,X of the first intersegment ; 
and the three postcoxal ligaments rils-fuz, etc. The cruciate muscle 
has been discussed in the preceding sections. 

The postcoxal ligaments are often frail and transparent, and there- 
fore easily overlooked in dissection ; and they dissolve rapidly in alkali. 
These characteristics no doubt explain why the ligaments have not 
received more attention from morphologists, for they are quite fre- 
quently present in primitive insects. (See fig. 6: 13, 24, 31, and fig. 7: 
24, as well as the figures of cockroaches. ) 

By a process that has many analogies in the evolution of the ptery- 
gote thorax, the postcoxal ligaments have often been replaced, in 
phylogeny, by apodemal growths, a course of development that cul- 
minates in a firm skeletal union between the furcal arm and the suc- 
ceeding i/s. Such unions constitute, or at least contribute to, the post- 
coxal bridges, whose interpretation has interested several previous 
students of insect morphology. 

Cockroaches, however, show little or no indication of the trend to- 
ward formation of a postcoxal bridge by sclerotization along the line 
of this former muscle. Only in Blaberus, of the blattids I have seen, 
is the distal end of the ligament 2i/s-fu, converted into a stiff, well- 
sclerotized apodeme ; whereas the usual course of evolution of a post- 
coxal bridge in other Ptergygota seems, contrariwise, to have been 
via sclerotization from the furcal attachment outward. 

In fact, the general impression left by the blattids is that their 
tendency is toward obliteration of these ligaments, and this tendency 
is increasingly manifest as one passes from the prothorax to the 
metathorax. All the cockroaches studied possess a fairly strong and 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


short rils-fu,; and a longer and more slender 2i/s-fug was found in 
all but Pycnoscelus. The presence of 3ils-fus was definitely ascer- 
tained only in the genera Periplaneta (4 species), Eurycotis, Blatta, 
and Cryptocercus. Failure to find a structure of this delicate nature 
is, of course, no proof of its absence ; but the observations cited never- 
theless do indicate quite well the tendency for these ligaments to 
weaken in the more posterior segments. The genera where their 
presence in the metathorax is most doubtful (Leucophaea, Nauphoeta, 
etc.) are notably those judged to have a more specialized muscular 
pattern on the basis of other criteria; and several of these are large 
insects, where such a structure, if present, should be relatively easy 
to find. 

The origins of the postcoxal ligaments of cockroaches, though 
clearly intersegmental, are at sites anterior and dorsal to the small 
sclerites indentified as the true ils by their reception in Cryptocercus 
of the transverse ligaments and of the usual dorsoventral muscles. 
In some other insects, the two sites are closer together or even indis- 
tinguishable, and I can offer no explanation for their separation in 
blattids. 

d. Spinasternal muscles of the abdominal ils—Only two such mus- 
cles have been found in cockroaches, namely 2sps-syz4 and “3sps’- 
Sua. The usual abdominal attachments for both are near the antero- 
lateral angle of the second sternum, somewhat anterior and ventral 
to the suspension of the ventral diaphragm. As explained above, this 
region of the definitive sternite is believed equivalent morphologi- 
cally to the thoracic ils. 

Identification of this attachment site with the ils renders dubious 
the homology, indicated as possible in table 1, of “3sps”-syr4 with the 
spinafurcal muscles rsps-fue, 2sps-fus; for it is very unlikely that 
the ils have contributed to the furcal structures of cockroaches. (See 
this section, c, above. ) 

The muscle from the second spina 2sps-syz4 clearly has no serial 
homolog in blattids. It is ordinarily inserted on sy74 somewhat mesad 
and ventrad of “3sps”-sjz4, and is thus two full segments in length. 
The variant attached on s;;74 has been discussed in section I,e. 


3. THE FURCAE 


The consensus of morphologists has been that the furcae (fu) of 
higher insects have been produced, in phylogeny, by the approxima- 
tion in the ventral midline of paired segmental sternal apophyses 
(Weber, 1928; Snodgrass, 1929). The resulting Y-shaped structure 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK wy. 


consists of the infolded furcabasis and the laterally extended furcal 
arms. In cockroaches, as in other primitive forms, right and left 
apophyses remain separate. For this reason, purists avoid applying 
the term “furca” to them, but for convenience we shall continue to 
refer to them as the furcae or furcal arms, with which they are 
homologous. Despite the seemingly incontestable segmental nature 
of these apophyses, they nevertheless carry a large fraction of the 
surviving longitudinal ventral intersegmental musculature in all ptery- 
gote insects. This situation poses a contradiction, long recognized and 
accepted by students of thoracic structure, that has never been satis- 
factorily resolved (cf. Snodgrass, 1929). 

Weber (1928) surmised that the present furcal intersegmental mus- 
cles had been derived from spinasternal muscles. He proposed that, 
as the furcal arms were gradually elevated in phylogeny, they inter- 
cepted the spinasternal muscles, which thereupon acquired furcal at- 
tachments and lost their primary connections with the spinae. This 
hypothesis, which regards the furcal muscles as replacements for the 
spinasternal muscles, is clearly untenable in the face of the presence 
of the usual complement of furcal intersegmental muscles in all those 
primitive forms, such as larval Dytiscus (Speyer, 1922) or Cybister, 
larval Corydalus, and the cockroaches, which still retain an extensive 
spinasternal musculature, including (in the blattids) both spinaspinal 
muscles Isps-2sps, 2sps-“3sps.’ Conceivably, the rising furcal arms 
could have intercepted some of the more lateral bands of the primary 
ventral longitudinal intersegmental muscles, for instance those at- 
tached on or below the transverse ligaments, Isps-rils, etc. ; but even 
this modification of Weber’s hypothesis is unconvincing in the absence 
of any known situation in insects where interception of a muscle by 
a skeletal element has led to the development of an attachment be- 
tween the two. Moreover, the data of Carpentier, Barlet, and others 
(see Barlet, 1954, for references) show that the essential features of 
the furcal complex exist in the Apterygota, which also possess an 
extensive array of muscles homologous with the spinasternal muscles 
of higher forms. Any notion that the furcal longitudinal muscles have 
arisen in the Pterygota through transfer of muscles from some other 
category must therefore be abandoned. How then can they be ac- 
counted for? 

An answer may be approached, we believe, through realization that 
the principal endoskeletal structures of insects and other arthropods 
have all developed as the result of sclerotization along the course of 
former muscles, and that the present sternal arms are of this nature. 
Although the genesis of certain endoskeletal structures lies so far in 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


the past that it will probably be impossible forever to document the 
details of the process in these instances, there are many other cases 
where the course of evolution can be deduced with reasonable cer- 
tainty from comparisons of existing forms. The complex endosternal 
structures of the Apterygota furnish a number of examples, for, as 
described by the Belgian authors cited above, many endosternal ele- 
ments that are ligamentous in one species or group are still repre- 
sented by functional muscles or by apparently degenerate muscles in 
others. In the opinion of the present writer, yet other parts of the 
endosternum that are invariably ligamentous in the apterygote species 
so far studied are homologous with muscles, such as the transverse 
muscles, that persist as contractile elements in some primitive Ptery- 
gota and as ligaments in others. Another clearcut set of examples of 
the replacement of muscles by endoskeletal structures is found in the 
later history of the sternal arms themselves, for instance in the de- 
velopment of the furcopleural fusion, which has occurred inde- 
pendently in numerous lines of descent. Here the process can be fol- 
lowed in some detail through several series of intermediate stages 
provided by existing forms. 

As a generalization we offer the hypothesis that all such endoskele- 
tal developments owe their inception to other structural or functional 
changes that have limited freedom of movement at the insertions of 
certain muscles. These muscles, deprived of their original effective- 
ness as contractile organs, are doomed to disappear unless they happen 
to retain some value in the role of static supports or braces. Further- 
more, the organism evidently finds it more economical to construct 
the braces it requires from other than contractile tissue, which cannot 
resist compression and which can maintain tension only through a 
continuous expenditure of energy, so that replacement of bracing mus- 
cles or tensors by noncontractile ligaments or by stiffer sclerotized 
apodemes is the usual evolutionary pattern. In our view, structures 
originating in this manner constitute the primary endoskeletal rudi- 
ments. Once established, these may be variously molded in later evo- 
lution in accordance with the mechanical requirements they are called 
upon to fill; and in the course of such modification their original 
derivation from muscles may be almost wholly obscured. 

Returning to the narrower problem of the nature of the sternal arms 
and their longitudinal musculature, we may point out that the arms 
are represented in the Apterygota by ligamentous straps that con- 
nect the thoracic endosterna, which are mainly intersegmental in char- 
acter, with the respective preceding segmental sternal regions (refer- 
ences in Barlet, 1954). In these insects the endosterna provide the 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK 19 


attachment sites for almost all the ventral musculature, including, of 
course, the usual longitudinal intersegmental muscles. We have al- 
ready indicated our belief that the entire endosternal complex, which 
is ligamentous in consistency, is of muscular derivation ; and we sug- 
gest here that the sternal ligaments are merely another example of 
transformed muscles. We may then regard the endosternum schemati- 
cally as a point of junction of various intersegmental muscles, among 
which are the usual longitudinal bands and a muscle to the preceding 
segmental sternum. 

The configuration thus summarized is, however, exactly what is 
seen in the musculature associated with the sternal arms of pterygote 
insects. True, the number of elements that impinge upon this focus 
is less than in the Apterygota; but those elements that do occur in 
the Pterygota all have their counterparts in muscles that are attached 
on the apterygote endosternum in proximity to the attachments of 
the sternal ligaments, or in similarly directed portions of the endo- 
sternum itself. Only the fact that the sternal arm of Pterygota is 
usually a heavily sclerotized ingrowth of the ventral body wall gives 
the superficial impression of a fundamental difference between the 
two subclasses. 

In cockroaches, even this distinction breaks down; for in the 
prothorax of blattids the paired furcal pits do not give rise at once 
to sclerotized apodemes, as they do in the mesothorax and metathorax, 
or in the prothorax of most Pterygota. Instead, there extends inward 
from the pit a flexible, fibrous ligament that connects with the apex 
of a sclerotic bar whose other end articulates firmly with the pleural 
apodeme. Upon this bar, at or near its junction with the sternal liga- 
ment, are inserted the usual muscles of the furcal complex. 

On the basis of these facts and the considerations outlined above, 
we suggest that the sternal arms of pterygote insects represent muscles 
that formerly ran from the segmental sternal region posteriorly to a 
common junction of various other intersegmental muscles, including 
the forerunners of the present longitudinal furcal muscles. In the 
course of evolution, the sternal muscles were replaced first by non- 
contractile ligaments, a condition still manifest in the Apterygota and 
in the prothorax of blattids, and finally by sclerotized apodemes, the 
form in which they now appear in the pterothorax of cockroaches and 
in all three thoracic segments of most winged insects. These changes 
in the sternojunctional muscle have not altered the morphological rela- 
tionships of the other muscles attached at the junction, which may 
still be regarded as an intersegmental locus in the morphological sense. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


There is thus no problem of explaining a shift of their attachments 
to a segmental site, for no shift has occurred. 

Apart from the structure of the profurca, with its connotations for 
the evolution of the furcal structures of pterygote insects in general, 
there is little that is remarkable about the sternal apophyses and 
their musculature in cockroaches that has not already been touched 
on in preceding sections. The ventral furcal intersegmental muscles 
found in the Blattariae may be classified as (a) spinafurcal muscles; 
(b) furcal muscles from the ils; and (c) furcofurcal muscles, includ- 
ing muscles with furcal origins and insertions in the head, neck, or 
abdomen. If the suggestion offered above is correct, that the definitive 
furcal apophysis is partly of intersegmental nature, a number of other 
muscles with furcal attachments, such as those of the appendage, may 
also be primarily intersegmental. However, further work is needed 
on the details of such relationships, and it seems best to leave them 
for future consideration. 

Except for muscles with spinasternal attachments (table 1), the 
ventral furcal intersegmental muscles of cockroaches are listed in 
table 2. 

Furcal muscles from the spinae and i/s have been discussed above, 
particularly under sections 1,b, 2,a, and 2,c. Like these muscles, the 
furcofurcal muscles of blattids can be homologized in considerable 
detail with those of comparable location in other insects. Readily dis- 
tinguished in most cockroaches are a usually slender mesal band and 
a more massive lateral band of both fw,-fu2 and fus-fus. Components 
probably homologous with each of these bands can also be identified 
in many blattids in the muscle fi3-sr74, which often includes an addi- 
tional more ventral group of fibers. These subdivisions, ordinarily 
lumped together in descriptions, seem to possess a fair degree of 
constancy in a number of insect orders, and may be of significance in 
future more detailed comparative studies. 

The fact that the furcabdominal muscles are inserted on the second 
(never on the first) abdominal sternum, is what would be expected 
if the furcal attachment is really intersegmental, as has been argued 
above. Morphologically, these muscles still run from the third 
thoracic intersegment to the first abdominal intersegment, and have 
neither lost nor shifted their original attachment sites. However, just 
as the muscle 2sps-syz4 has become 2sps-syzz4 in some species (sec- 
tion 1,e), so is a portion of fts-sy4 found, at times, as fus- 
Sra, aS a result of an analogous development (figs. 1, 5, 8, II, 12, 


w7ose) 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK 21 


The profurca bears two anteriorly directed longitudinal ventral 
intersegmental muscles. The stronger, consisting of two or more 
bands and serially homologous with the muscles fu;-ftte, fue-fus, 
passes into the head to insert on the tentorium. This muscle, fu,-tent 
(figs. 1-18: r), is commonly considered to be more than one segment 
in length (Snodgrass, 1935, p. 159). The weaker, usually a thin, flat 
strap of somewhat degenerate appearance, has a more ventral origin 
on fu; (figs. 4, 5, 9: 2) and is inserted on the mesal lobe of the 
ipselateral second cervical sclerite. Possibly this site should be re- 
ferred to the Oils, which are certainly included in the first cervical 
sclerites, rcv, of which the 2cv may be merely subdivisions (Cramp- 
ton, 1926) ; but this question cannot be settled until the constitution 
and muscular connections of the various cervical structures of insects 
are better understood. 


GENERAL DISCUSSION 


Reinvestigation of the ventral intersegmental muscles of the three 
cockroaches previously studied by others has shown each of the 
earlier accounts deficient in some respects. The defects are mostly 
errors of omission. Thus, none of the earlier investigators noted the 
muscle Icv-c+,X or the three postcoxal ligaments Z1/s-fu;, etc. That 
they did not is understandable, for these are hardly muscles in a func- 
tional sense, even though the cervicocoxal “muscle” does have a con- 
tractile core a few microns in diameter. Carbonell’s (1947) exclusion 
of the ligament fis-fus3(=3sps-fus) and of the bands “3sps’’-ventr. 
diaphr. from his description is likely to have been on similar grounds ; 
for in most other details his depiction of P. americana is accurate 
and complete. However, unless Maki’s (1938) specimens of 
P. australasiae differed radically from those of this species available 
to the writer, one must reject Maki’s assertions that the muscles 
Isps-fuy, Isps-cx%2, 2sps-fuy, 2sps-cx3, and 2sps-sy4 are absent, and 
that a muscle, 2ps-fu2, unknown in any other cockroach, is present. 
Miall and Denny (1886) stated explicitly that they had not given a 
complete account of the muscles of B. orientalis, so that there is no 
cause for surprise in the fact that this species has numerous muscles 
unmentioned in their description. 

When these few corrections have been made, it is seen that all 
species of cockroaches thus far studied have nearly identical com- 
plements of ventral intersegmental muscles. The spinasternal pro- 
motors of the coxae, Isps-cx, and 2sps-cx 3, are absent from some 
species and show signs of weakness in others. The metathoracic post- 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


coxal ligament 3is-fus is either lacking or very frail in several genera. 
Cryptocercus alone of the species investigated possesses muscles as 
yet found in no other blattid ; these are the semiligamentous transverse 
bands rsps-rils and 2sps-2ils, parts of which seem to have survived 
as septa in some other cockroaches. With these minor exceptions, the 
differences among the several species are merely variations in rela- 
tive size and proportions of the various muscles, or occasionally in 
their position. Such differences, some of them quite conspicuous, ob- 
viously indicate shifts in functional emphasis, although in most in- 
stances the details of their interpretation from this viewpoint are 
obscure. 

In contrast with most modern insects, the cockroaches enjoy a 
relatively rich ventral intersegmental musculature. Some authorities 
would be inclined, perhaps, to regard this as a consequence of 
secondary reduction of sclerotization in the ventral regions of the 
blattid thorax ; but acceptance of this view would make it very hard 
to account for the presence of homologous muscles in a number of 
other groups in which the thorax is extensively sclerotized. There 
is no muscle recorded in this paper for which either a direct or a serial 
homolog has not been found in at least one other order of pterygote 
insects, and most of them are known from several. Coupled with the 
fact that those other orders that display a similar degree of com- 
plexity in the ventral intersegmental musculature are the ones con- 
sidered highly primitive in various other respects, the evidence seems 
more consistently interpreted in the conclusion that the cockroaches 
also are primitive in this feature, and that the primitive state of the 
ventral intersegmental muscles was a complex one. As already indi- 
cated in the introduction, the various structural patterns preserved 
for our scrutiny among the more recently evolved orders of insects, 
which constitute a progressive series of specializations toward greater 
efficiency in flight, show that improvement in the flight mechanism 
has been accompanied regularly by reduction in the ventral inter- 
segmental thoracic musculature. These facts also favor the view that 
the possession of numerous discrete muscles in this category is a 
primitive characteristic. The results of a more comprehensive inquiry 
into these problems will be reported elsewhere. 


SUMMARY 


1. A comparative study was made of the ventral intersegmental 
musculature of the thorax in 19 species of cockroaches. The observa- 
tions produced a few corrections, mainly additions, to earlier de- 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK 23 


scriptions of 3 of these species. In general, there are very minor 
differences among the species in respect to the presence or absence 
of individual muscles in the category studied, although there are 
numerous differences, some conspicuous, in the relative size and pro- 
portions of the various muscles. 

2. The present ventral intersegmental thoracic muscles had their 
primary attachments on the spinae, on the intersegmental latero- 
sternites, or on the forerunners of the furcal apophyses. Cockroaches 
still have two typical spinae and definite vestiges of a third. They 
possess an extensive spinasternal musculature. Remnants of the 
musculature of the intersegmental laterosternites are present, but some 
of these muscles now have segmental attachments, and others are 
represented by noncontractile ligaments. The longitudinal furcal 
muscles are equivalent to those of other pterygote insects. 

3. Attention is called to the postcoxal ligaments that run between 
the furcal apophysis and the immediately following intersegmental 
laterosternite in each of the thoracic intersegments, and to the sig- 
nificance of these former muscles in the development of the postcoxal 
bridges of higher insects. 

4. Evidence and arguments are presented for the hypothesis that 
the furcal apophyses represent former muscles that have been replaced 
in phylogeny by sclerotized apodemes. It is suggested that one at- 
tachment of these muscles was on the segmental sternum, the other 
at a common intersegmental junction of several other muscles, among 
them elements of the longitudinal ventral group. Loss of movement 
at the sternal insertion led first to transformation of the sterno- 
junctional muscle into a fibrous ligament and eventually to the scle- 
rotization of the ligament. Analogous events have occurred fre- 
quently in the evolution of the pterygote thorax. The blattid 
prothorax exemplifies a stage in the evolution of the typical furcal 
apophysis when the postulated sternal muscle was still in a liga- 
mentous condition. The subsequent sclerotization of this ligament, 
which has occurred in the other thoracic segments of cockroaches and 
in all three segments of most Pterygota, in no way alters the morpho- 
logical relationships of the other muscles inserted upon its central 
end; morphologically, then, the present attachments of the longitudi- 
nal ventral muscles on the furcal arms are still intersegmental, and 
it is therefore unnecessary to invent mechanisms whereby they might 
have been shifted from an intersegmental to a segmental site of 
attachment. 

5. The ventral intersegmental thoracic musculature of cockroaches 
is rich in number of elements, compared to that of more recently 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 


evolved groups. However, direct or serial homologs of all the ele- 
ments occur in one or more other orders of winged insects. The most 
extensive complements of these muscles are found in those forms, such 
as larval dytiscids, Grylloblattodea, and Megaloptera, that are regarded 
as primitive on the basis of other criteria. It is concluded, therefore, 
that the blattids also are primitive in respect to the ventral interseg- 
mental muscles ; and that possession of a rich ventral intersegmental 
musculature was characteristic of the early Pterygota. 


ACKNOWLEDGMENT 


The writer is grateful to Dr. R. E. Snodgrass for helpful discussion 
of several questions considered in this paper. 


GLOSSARY OF ABBREVIATIONS 


CR aaah eines ehisaesie cervical sclerite 

Cae NG Soehion tue acis ls coxa 

OPS mal ccatate eihate Oehatere! tev .. episternum 

| (OIG A TiS CRS AIG ae oo furca, furcal arm, segmental sternal apophysis 

AUS aE Misia io tebe chet carats intersegmental laterosternite 

Dy vericic eisgesiite atelier phragma, or the primary dorsal intersegmental fold 
from which the phragma is derived 

DOSET Ob tole cisions Sees posterior rotator, a functional designation used to 
distinguish certain leg muscles 

SHER Saas sath eee segmental sternum 

SEP Een alate cistotoinioisietais tothe septum 

OS te aca chan pce mitt spinasternite or spina 

Pe ie ofetcistaciecraiete wiorslueteleie.s segmental tergum 

COME Ao aeicte eee a hee tentorium 

Vente APMP) Fs cece aes ventral diaphragm 

EN ee eer iat cule poor shat cruciate, used of a muscle whose origin and insertion 


are on opposite sides of the midline 


For the way in which these abbreviations are compounded into designations 
of muscles, see section on Method and Material in text. 


REFERENCES 
BaRteT, J. 
1954. Morphologie du thorax de Lepisma saccharina L. (Aptérygote Thy- 
sanoure). II.—Musculature (2™° partie). Bull. Ann. Soc. Ent. 
Belgique, vol. 90, pp. 299-321. 
CARBONELL, C. S. 
1947. The thoracic muscles of the cockroach Periplaneta americana (L.). 
Smithsonian Misc. Coll., vol. 107, No. 2, 23 pp., 8 pls. 
Crampton, G. C. 
1926. A comparison of the neck and prothoracic sclerites throughout the 
orders of insects from the standpoint of phylogeny. Trans. Amer. 
Ent. Soc., vol. 52, pp. 199-248. 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK 25 


Forp, N. 
1923. A comparative study of the abdominal musculature of orthopteroid 
insects. Trans. Roy. Can. Inst., vol. 14, pp. 207-319, 17 pls. 
Futter, C. 
1924. The thorax and abdomen of winged termites. Union of South Africa 
Dept. Agr. Ent. Mem., vol. 2, pp. 49-78. 
Heymons, R. 
1895. Die Embryonalentwickelung von Dermapteren und Orthopteren unter 
besonderer Beriicksichtigung der Keimblatterbildung. viii + 134 pp., 
12 pls. Jena. 
MaAkgI, T. 
1938. Studies on the thoracic musculature of insects. Mem. Fac. Sci. Agr. 
Taihoku Imp. Univ., vol. 24, No. 1, 343 pp., 17 pls. 
Mratt, L. C., and Denny, A. 
1886. The structure and life history of the cockroach (Periplaneta ori- 
entalis). 224 pp. London. 
REHN, J. W. H. 
1951. Classification of the Blattaria as indicated by their wings (Orthop- 
tera). Mem. Amer. Ent. Soc., No. 14, 134 pp., 13 pls. 
Roonwat, M. L. 
1937. Studies on the embryology of the African migratory locust, Locusta 
migratoria migratorioides Reiche and Frm. II.—Organogeny. 
Philos. Trans. Roy. Soc. London, B, vol. 227, pp. 175-244. 
ScHARRER, B. 
1948. The prothoracic glands of Leucophaea maderae (Orthoptera). Biol. 
Bull., vol. 95, pp. 186-108. 
Snoperass, R. E. 
1929. The thoracic mechanism of a grasshopper, and its antecedents. Smith- 
sonian Misc. Coll., vol. 82, No. 2, 111 pp. 
1935. Principles of insect morphology. ix + 667 pp. New York. 
Speyer, W. 
1922. Die Muskulatur der Larve von Dytiscus marginalis. Ein Beitrag zur 
Morphologie des Insektenkérpers. Zeitschr. wiss. Zool., vol. 119, 
PP. 423-492. 
WALKER, E. M. 
1938. On the anatomy of Grylloblatta campodeiformis Walker. 3. Exoskele- 
ton and musculature of the neck and thorax. Ann. Ent. Soc. Amer., 
vol. 31, pp. 538-640. 
1943. On the anatomy of Grylloblatta campodeiformis Walker. 4. Exo- 
skeleton and musculature of the abdomen. Ann. Ent. Soc. Amer., 
vol. 36, pp. 681-706. 
Weter, H. 
1928. Die Gliederung der Sternopleuralregion des Lepidopterenthorax. 
Zeitschr. wiss. Zool., vol. 131, pp. 181-254. 


26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 


EXPLANATION OF FIGURES 


All muscles figured are numbered uniformly according to the list below. 
Instances in which the definitive attachments differ from those given in the list 
have been discussed fully in the text. 

The arrangement of ventral muscles in certain species is such that not all of 
them can be shown in a single drawing. However, all cockroaches studied have 
all muscles given in the list, whether the muscles appear in the figures or not, 
except as noted under tables 1 and 2 or in the text. The termite Zootermopis 
(fig. 6) and the mantid Tenodera (fig. 7) have only the ventral intersegmental 
muscles shown in the drawings. 

In a few instances different levels of dissection have been shown in different 
parts of the same figure; this does not imply an absence of the usual bilateral 
symmetry. To assist in orientation, some figures contain a few muscles that do 
not belong to the ventral intersegmental category. 

The scale indication represents I mm. 


NUMBERING OF MUSCLES IN FIGURES I-18 


Number Muscle Number Muscle Number Muscle 
1 fus-tent. 13 rils-fitx 25 fus-2ph 
2 fur-2cv 14 fiti-Iph 26 ftts-fuls 
3 Icv-cxiX 15 fits-fus 27 “3sps’-fus 
4 Isps-trils 16 2sps-fus 28 “3sps”-sira 
5 Isps-epss 17 2sps-2ils 29 fils-c2s post. rot.* 
6 Isps-fus 18 2sps-epss 30 “3sps’-ventr. diaphr. 
7 Isps-Cx1 IQ 2Sps-CH2 31 3tls-fus 
8 epse-fusX * 20 2sps-fus 32 fus-Srra * 
Q epse-cx1X * 21 2sps-sua * 33. fus-3ph (or -tra) 
10 Isps-fits 22 25SpS-CHs 34 Sura-Sura 
Il ISpPS-CHs 23 2SpS-3ZSps 35 Sua-Sua 
I2 Isps-2sps 24 «2ils-fus 


* Variants from these attachments are discussed in the text. 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK 








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hb SS 


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Wi ZA 

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4. Rage 
i 
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16- ea ‘thts 


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‘NY 734 


Fics. 1-4.—1, Leucophaea maderae (Fabr.), male. 2, Blatta orientalis L., 
male nymph. 3, Neostylopyga rhombifolia (Stoll), male. 4, Blattella vaga 


(Heb.), female. 


: 
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28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 





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Fics. 5-8—Blaberus craniifer (Burm.), nymph. 6, Zootermopsis angusticollis 
(Hagen), worker (Isoptera). 7, Tenodera sinensis Sauss.. nymph (Mantodea). 
The prothorax is shown on the left, the mesothorax and metathorax on the right. 
The portion of the prothorax posterior to the sternal arms, which does not carry any 
ventral intersegmental muscles, has been omitted. Glandular tissue that invests the 
muscles Icv-cv:X (3) is shown in solid black. 8, Nauphoeta cinerea (Oliv.), nymph. 


NO. II THORACIC MUSCLES OF COCKROACHES—CHADWICK 


29 






{NA Lin ~. 
ae ISN 41). 6+8 


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Fics. 9-12.—9, Supella supellectilium (Serv.), female. 10, Cryptocercus punc- 


tulatus Scud., female. 11, Pycnoscelus surinamensis (L.), female. 12, Diploptera 
dytiscoides (Serv.), nymph. 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. T3t 





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Fics. 13-18.—13, Cryptocercus punctulatus Scud., female, detail of first thoracic in- 
tersegment. 14, Periplaneta brunnea Burm., nymph, detail of first thoracic interseg- 
ment. 15, Periplaneta australasiae (L.), nymph, detail of first thoracic intersegment. 
16, Periplaneta americana (L.), nymph, detail of first thoracic intersegment. 17, 
Moree rhinocerus Sauss., female. 18, Parcoblatta pennsylvanica (DeGeer), 
nymph. 











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