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SMITHSONIAN 
MISCELLANEOUS COLLECTIONS 



VOL. 131 






r^y^^'^rv'^^'vril 






'every man is a valuable member of society vvuo, by his observations, researches, 

AND experiments, PROCURES KNOWLEDGE FOR MEN" JAMES SMITHSON 



(Publication 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) 



CONTENTS 

1. Abbot, C. G. Leading operations of the Smithsonian Astro- 

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

2. Peterson, Mendel L. The last cruise of H.M.S. "Loo." 55 

pp., 17 pis., 3. figs. Nov. 23, 1955. (Publ. 4224.) 

3. Fairchild, G. B. Synonymical notes on neotropical flies of the 

family Tabanidae (Diptera). 38 pp. Jan. 11, 1956. (Publ. 
4225.) 

4. Cooper, G. Arthur. New Cretaceous Brachiopoda from Ari- 

zona. 18 pp., 4 pis. Dec. 21, 1955. (Publ. 4227.) 

5. 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.) 

6. Gazin, C. Lewis. Paleocene mammalian faunas of the Bison 

Basin in south-central Wyoming. 57 pp., 16 pis., 2 figs. Feb. 
28, 1956. (Publ. 4229.) 

7. Gazin, C. Lewis. The upper Paleocene Mammalia from the 

Almy formation in western Wyoming. 18 pp., 2 pis. July 31, 
1956. (Publ. 4252.) 

8. 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 pis., i fig. Oct. 30, 1956. (Publ. 4257.) 

9. Kilham, Lawrence. Breeding and other habits of casqued 

hornbills (Bycanistes sitbcylindricus) . 45 pp., 6 pis., 2 figs. 
Nov. 8, 1956. (Publ. 4259.) 

10. Snodgrass, R. E. Crustacean metamorphosis. 78 pp., 28 figs. 

Oct. 17, 1956. (Publ. 4260.) 

11. Chadwick, L. E. The ventral intersegmental thoracic muscles of 

cockroaches. 30 pp., 18 figs. Jan. 15, 1957. (Publ. 4261.) 



(v) 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 1 



Eoebling :f unb 



LEADING OPERATIONS OF THE 

SMITHSONIAN ASTROPHYSICAL 

OBSERVATORY, 1895 to 1955 



By 

C. G. ABBOT 

Research Associate, Smithsonian Institution 




(PUBUCATION 4222) 



CITY OF WASHINGTON 

PUBLISHED BY THE SMITHSONIAN INSTITUTION 

SEPTEMBER 22, 1955 



P. 



BiLTIMORB, IIS., n. B. A. 



LEADING OPERATIONS OF THE SMITHSONIAN 

ASTROPHYSICAL OBSERVATORY, 

1895 TO 1955 

By C. G. Abbot 
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 i. — Improved 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. 

2. Galvanometer. Theory investi- Astrophys. Journ., vol. 18, No. i, 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 
steadiness. 

4. Silver-disk pyrheliometer invented. Smithsonian Misc. Coll., vol. 56, No. 

About 100 copies have been 19, 191 1. 



1 Hereafter referred to simply as "Annals." 

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL 131, NO. 1 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



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

5. Water- flow and water-stir stand- 

ard pyrheliometers invented and 
used. The world's scale of solar 
radiation measurements rests on 
them. 2 

6. 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. 

7. 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. 

8. Balloon recording pyrheliometer 

invented and used at high alti- 
tudes. 

9. Two-mirror coelostat invented. 

10. Slide-rule extrapolator invented. 

Constantly used in long-method 
solar observing. 

11. High-power lamp and other de- 

vices prepared by F. E. Fowle 
for researches on deep infrared 
spectrum. 

12. Highly sensitive radiometer in- 

vented for measuring energy 
spectra of stars. 

13. A prism of nearly normal wave- 

length dispersion invented. 



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

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

16. A multiple rotating-sector dia- 



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

p. 105, 1954. 
Smithsonian Misc. Coll., vol. 95, No. 

23, 1937; vol. Ill, 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. no. No. II, 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, 2II, 1908. 
Annals, vol. 4, pp. 84-86, 1922. 



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



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

1929. 
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. 



2 A. P.O. modified form of Angstrom pyrheliometer is used in daily observa- 
tions. See .'Xnnals, 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 on public exhibition in 
Langley case (see above). 



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



Past 2. — Various itwentions, mainly for military use in World Wars I and II 



Variable-speed power-transmis- 
sion mechanism, Claim i, 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." 

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

Self-propelled rotating projectile 
for smooth-bore guns. Combina- 
tion with smooth-bore ordnance. 



U. S. Patent No. 893416 of July 
1908. 



14, 



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, 
1921. 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



8. 

9- 
10. 



Gyroscopic navigation instrument. 
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. 1501^ 
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 i, 1941. 
Patent No. 2460482, February i, 1949. 



SECTION B 
Part i. — Researches 



I. Bolometric map of infrared solar 
spectrum. 



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



Annals, vol. I, pp. 5-204, 1900; vol. 5, 

p. 54. 1932. 
Smithsonian Misc. Coll., vol. 82, No. i, 

1929. 
-Annals, vol. i, pp. 219-237, 253-262, 

1900. 



NO. I ASTROPHYSICAL OBSERVATORY, 1895 TO I955 — ABBOT 



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

3. Structure of water-vapor bands 

wi and wj. 

4. Total solar eclipses, 1900, 1901, 

1908, 1918, 1919. 



5. Theory of sensitive galvanometer. 



6. "The cheapest form of light." 

7. Solar-constant and solar -distribu- 

tion work, begun in 1902. 

8. Mount Wilson expeditions, begun 

1905. 

9. Theory of atmospheric transmis- 

sion. 

10. Methods for measuring the solar 

constant. 

11. Transmission of the spectrobo- 

lometer. 

12. Pyrheliometry. 

13. Details of solar-constant observ- 

ing. 

14. Sources of error in solar-constant 

work. 

15. Solar-contant results of stations 

compared. 



16. Normal solar-energy curves. Pre- 

ferred determination. 

17. Sun's temperature. 

18. Atmospheric transmission, many 

stations, sea level up to 14,000 
feet altitude. 

19. Theory of vacuum bolometer, cor- 

rected later. 



Annals, vol. i, 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. 29, 31, 34, 35, 

1922. 
Smithsonian Misc. Coll., vol. 69, No. 9, 

1 91 9. 
Annals, vol. I, pp. 244-252, 1900. 
Astrophys. Journ., vol. 18, No. i, July 

1903. 
Annals, vol. 2, p. S, 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, 1 17-124, 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. 21-29, 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, I10-I12, 
1908; vol. 3, pp. 104-I13, 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. 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



20. Infrared and ultraviolet correc- Annals, vol. 5, PP- 103-110, 1932. 

tions for solar-constant work. 

21. Solar variation : Astrophys. Journ., vol. 19, p. 305, June 

1904. 

a. First suspected. Annals, vol. 2, pp. 98-103, II7-I79, 

1908. 

b. Clayton's contributions. Annals, vol. 4, PP- 36, 185, 367-374 

1922. 

c. From solar-constant work Annals, vol. 5, pp. 246-269, 1932. 

1 920- 1 930. 



d. Short up and down trends 

and ( I ) temperatures, 
(2) ionosphere. 

e. Accompanying (i) hurri- 

canes, (2) magnetic 
storms, 
f. Accompanying sunspots. 



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

(i) Smithsonian Misc. Coll., vol. no, 
No. I, and (2) No. 6, 1948. 



Smithsonian Misc. Coll., vol. no, No. 

6, 1948. 
Annals, vol. 7, pp. 165-168, 1954. 
(i) Smithsonian Misc. Coll., vol. 104, 

^>o. 3, 1944; vol. 116, No. 4, 1951 ; 

(2) vol. Ill, No. 13, 1949. 
h. Long periodic and weather. 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- 

23. Defense of our solar-constant Annals, vol. 4, pp. 3.-'3-366, 1922. 

value (Abbot, Fowle, Aldrich). 

24. Brightness of the night sky. Astron. Journ., vol. 27, No. 3, pp. 17- 

24, June 20, 1 911. 

25. Direct and scattered radiation of Astron. Journ. vol. 28, No. 16, pp. 129- 

sun and stars. 135, March 1914. 

26. Tower telescope on Mount Wilson Annals, vol. 4, pp. 217-257, 1922. 

and solar-drift curves. Smithsonian Misc. Coll., vol. 78, No. 5, 

1926. 

27. Nature of the sun's sharp Scientia, vol. 19, pp. 171-181, March 



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



22. A large family of periodic varia 
tions: 

a. In the sun. 

b. In the weather. 



boundary. 



28. Volcanoes and climate. 

29. Summary of the work of the As- 

trophysical Observatory, 1890- 
1 920. 

30. Radiometer measurements of stel- 

lar-energj' spectra. 



1916. 
(See also Abbot, C. G., "The Sun," 

1911.) 
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. 293-311, 

1929. 



NO. I ASTROPHYSICAL OBSERVATORY, 1 895 TO 1 955 — ABBOT 



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. 

22. Daily solar-constant values, 1920- 
1952, with 1 0-day and monthly 
means. 

2Z. Convenient table for solar-constant 
tabulations. lo-day and monthly 
mean excesses over 1.900 in 
hundredths percentages of 1.94. 
Thus 1.950 becomes 

I.950-I-900X 100 = 2.58. 
1.94 

Similarly 1.940 becomes 2.06. 
This difference, 0.52, is 0.53 per- 
cent of mean solar constant. 
Note. — 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, 1954, 



Annals, vol. 5, pp. 177-182, 1932; vol. 

6, pp. 85-162, 169-175, 1942; vol. 7, 

pp. 26-94, 1954. 
Smithsonian Misc. Coll., vol. 117, No. 

10, pp. 20-24, 1952; vol. 128, No. 4 

(table i), 1955. 



Part 2. — Work of specialists 



L. B. Aldrich : 

a. The melikeron, an approxi- 

mately black-body pyra- 
nometer. 

b. Reflecting power of clouds, 

and earth's albedo. 

c. Eclipse expedition, June 

1 91 8. 

d. A study of body radiaticHi. 

e. Sun spots 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. 
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. 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



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. 

3. 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, 1929. 
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. 1-13, February 27, 1937. 



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

5, pp. i-io, August 14, 1953. 
Annals, vol. 7, pp. 99-104, 1954. 

.\nnals, vol. 7, pp. 144-164, 1954. 



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



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



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SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 2 



THE LAST CRUISE OF H.M.S. "LOO" 

(With 17 Plates) 



By 
MENDEL L. PETERSON 

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



(Publication 4224) 



CITY OF WASHINGTON 

PUBLISHED BY THE SMITHSONIAN INSTITUTION 

NOVEMBER 23, 1955 



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



THE LAST CRUISE OF H.M.S. "LOO" 

By MENDEL L. PETERSON 

Curator of Naval History 

U. S. National Museum 

Smithsonian Institution 

(With 17 Plates) 

In the spring of 195 1 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 (pi. 2, fig. i ) 
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 



^ 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 (pi. 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 (pi. 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 3^ 
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 (pi. 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 (pi. 6). In this lot was found a 6-pound 
shot with an arrow on it, which was immediately identified as the 
broad arrow (pi. 7). This was the first indication of the nationality 
of the ship, as this syml)ol 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 ^-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 i, 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 pis. 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 (pis. 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 (pis. 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 (pi. 15) 

^ Later a ehain 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 (pi. i6). 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 17 14 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 Looc 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 Ijegan March 15. The calendar year was 1744. (See Appen- 
dix B for UUing's letter.) 



NO. 2 LAST CRUISE OF II. 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 compleatcd 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. 

W: IC: GL:« 
By T.C. 

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. 



^ 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. 

^ 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 Spanisli 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. 

<* Initials 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. IS, Public Record Office.) 

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



6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

She was most probably launched in 1706^ since she first appears 
in the Navy List under the date April i, 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 17 10 she was attached to the 
Dunkirke Squadron and in the fall of that year was on convoy duty to 
the eastern countries.^- In April 171 1 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 171 1 was refitted and again sent to duty 
in British waters, meeting ships from Virginia and convoying them 
to British ports." During the winter of 171 2-13 she transported 
troops to Bayonne and returned with prisoners of war ^^ and then 
again served in the Channel, cruising against smugglers.^*' In the fall 
of 1 7 14 she was sent convoy to Port Mahon in the Mediterranean and 
on return was paid ofif and laid up.^^ 

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.^^ 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 

* 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). 

9 Navy List, Jan. i, 1707-Dec. 31, 1709, ADM 8/10, Public Record Office. 
" ADM 8/10. 

" Ibid. 

1- Ibid. 

" Ibid. 

"ADM 8/10 and 8/1 1. 

"ADM 8/11. 

10 ADM 8/12. 

17 Ibid. 

18 ADM 8/13. 
" Ibid. 

20 Ibid. ' 



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 patroUing 
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." 22 

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.-^ 



21 Ibid. 

22 Ibid. 

23 ADM 8/17 and 8/18. 

24 ADM 8/19 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. lago, 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. lago. 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. I3I 

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 -^ expected at 
St. Sebastian." ^^ 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, I743.-'' 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 LIunnington, 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. 

30 Admiralty In-Letters, ADM i, vol. 2625, pt. 3, No. 146. 

31 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. 

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

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

3'* ADM I, 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 hooks "to the 30 June, 
1742." ^^ Five days later admiralty orders "ahout 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,^^ 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. 

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

2^ Ibid., 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 I, vol. 233, extracts.) *T 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." 

^^ A drink distilled from rum. 

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



lO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Another order dated the same day directed Utting to have his ship 
"vichialled to four months only ^'•* for a f orreign 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 9-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 II orders were issued to the commanding officers of the 
Rye, 20 guns ; Flamborough, 20 guns, and the sloop Spy, 8 carriage 
gims 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- 
ment." " 



s» Thus rescinding the order of June 14, wliich had directed the loading of a 
6-months' supply of food. 

*o 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. 

<i ADM I, vol. 2625, pt. 3, No. 418. 

^~ ADM 2, vol. 60, p. 79. 

43 Ibid., 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 CaroHna, 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. 



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



12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 tlic 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 I3 

you shall reasonably suspect to have Wool ^^ 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. 

W. J.C. G. L. 

By 
T.C. 
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." *^ 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." *^ 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 

*" 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. 

48 ADM I, vol. 2625, pt. 3, No. 419. 

*'' Ibid., No. 420. 

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

4° Ibid., 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 i8th July, 1743. 

W. J.C. B. 

By 
T.C. 
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 j'e 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 ^o 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 yc summer months ^^ ; 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 20th 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 ^2 can go 12 months at first. There 
is an exceeding good careening place at Port Royall ^^ which can be made ours 

""> Cape Florida. 

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

^^ 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. 

^3 South Carolina. 



NO. 2 LAST CRUISE OF H.M.S. LOO — PETERSON 1 5 

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.-""'^ 

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.^^ 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 ofif Charleston 
Bar. In Charleston he found the sloop Spy ready for sea, the Rye 
"cleaned and almost fitt for sea," the Flamhorough "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 



■'■'•* ADM I, vol. 2625, pt. 3, No. 421. 

s5 Ibid. 

^^ Loo's General Muster Book, ADM 36, Ser. I, vol. 1823. 

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

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

^^ Ibid., 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 I, vol. 60, No. 435. 



l6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 . . ." ^^ Utting was mistaken, however, in his estimate of the 
damage, for closer examination revealed extensive damage to the mast 



61 Ibid. 

e- Ibid. 

83 ADM I, vol. 60, No. 436. 

61 Ibid. 

«'' Ibid. 

60 Ibid. 

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. 

6«ADM I, vol. 2625, No. 438. 

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

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

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



NO. 2 LAST CRUISE OF H.M.S. "lOO" PETERSON 1"] 

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 i8 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] ''^ 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 i8th day of November 1743. 

Ashby Utting 
To Captain Newnham of his 
Majesties Sloop Spy. 

" ADM I, vol. 2625, No. 426. 

^3 ADM 2, vol. 2625, No. 426. 

^* 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 
i8oo's. 



l8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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." **- 



^5 ADM I, 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. 

" ADM I, vol. 2625, No. 433. 
" Ibid 

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

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

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

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



NO. 2 LAST CRUISE OF II. M.S. LOO — rETERSON I9 

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." *^ 

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 

83 ADM I, vol. 2625, No. 427. 

^■* 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.) 

^^ ADM I, vol. 2625, No. 429. 

80 ADM I, vol. 2625, No. 424. 

^"^ ADM I, vol. 2625, No. 424. 



20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 "Irish 
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 Loo set sail with her charge it was 6 p.m. and growing dark. 
The "Pan of Matanzas" (fig. i), a fliat-topped mountain behind 
Matanzas Bay on the coast of Cuba, bore south by east at a distance 
of i8 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,°^ 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. 

80 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 I, vol. 5283.) 

"0 Utting's letter of February 15, 1744. (ADM i, vol. 2625. (See Appendix 
B.)) 

81 Bishop's deposition at the court martial. (ADM i, 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 
visibiHty not over a quarter of a mile.^- 

At 12: 30 a.m. and again at i : oo 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 oflf the wind 
the head sails were caught across wind and the ship struck the reef 



1\ 



hart . Uk> Pkii de Miitaii^'an 




I\kiUM.-iIjiii^aii 



Fig. I. — 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 oflf 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 

82 Deposition of John Randolph, master's mate, at the court martial of Cap- 
tain Utting. (ADM i, vol. 5283.) 

*2 Bishop's deposition. (ADM i, vol. 5283.) 
^* Utting's letter of February 15, 1744. 
85 Bishop's deposition. (ADM i. vol. 5283.) 
8° Utting's letter of February 15, 1744. 



22 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



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 




Fig. 2. — Map of the Florida Straits, showing the course of the last cruise of 
the Loo. I, 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 i^ 
cables ^^ length long and ^ broad which lay on the edge of the Bank 
of the IMartiers 3 leagues "* without them." ^^ (See fig. 2.) 

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



^'' A cable was 600 feet. 

8^ A league was 3 nautical miles. 

shutting'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 lo 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 oflf 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." ^°^ 

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 

'00 Randell's deposition. 
101 Ibid. 

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

lo^ Utting's letter of February 15, 1744. 
lo-" Ibid 



24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 — 60 in the altered longboat, 10 in the yawl,^°® 184 "in 
the little Sloop not 30 tuns" ^" and 20 in the captain's barge. Utting 
sent the sloop, the longboat, and the yawl 3 or 4 miles oflfshore 
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." ^°* 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 Ibid. 

100 y\ heavy double-ended rowboat. 

107 Utting's letter of February 15, 1744. 
"8 Ibid. 

108 Ibid. 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 CaroHna even for St. Augustine (if I could fetch nowhere 
else) rather than all be drowned which Doe assure you had very 
little other prospect." ^^^ 

The fair weather continued and the overloaded sloop arrived in 
Port Royal harbor (pi. 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." ^^^ 

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 vcssell bound from the Havannah to St. Augustine witli 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, tliat 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 

^10 The Gulf Stream would carry him northward. 

1^1 Utting's letter of February 15, 1744. 

1^2 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.^13 

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 ofificers 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,^'^^ 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 tlie 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 gaily s (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.^i" 

^13 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 i, 
vol. 2625, No. 455.) 

^1* 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. 

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

ii" 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.^^^ 

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." ^^* 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." ^^^ 

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 

%. 3-) 

Six days later. May 31, the court of 12 captains sat on board the 
ship-of-the-line Sandzvich 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." ^-^ 

On June 6 the Lords of the Adiniralty 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 Ibid. 

118 Ibid. 

119 Ibid. 

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




1 



i^M. ^.<-/// «^*«r/^^ /tf''* i«yt«.- ^/t »/. J<C« 
oD'<n/ -Attn /R/ ^' : /J^^^ ^tffi _^»,,i^ 

(ffTP ^coi / ^^jr^c-^-t^it^ ^f^/£^/ f..//2^/j^rw 

. ^'\^; 3— Letter of Capt. Asliby Utting to the Secretary of the Admiralty report- 
ing his arrival in England May 24, 1744, to stand trial by court martial for the 
loss of the Loo. (Photostat from the Public Record Office, London.) 
28 



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

pounds 4 shillings ii pence due them after deductions of 1,121 pounds 
4 shillings 1 1 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.^^^ 

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.^22 

On July 7, 1744, a commission was issued giving Utting command 
of the Gosport}^^ 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 10, 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." ^^^ 

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 
1 5th 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.^-" Utting's wish to return to South 

^^'^ Loo's pay list dated Aug. 10, 1744, Admiralty Ships Pay Books, Treasurers 
Series I, ADM 33, No. 352. 
122 Admiralty In-Letters, ADM i, vol. 2267. 
1-3 AD 6/16, Commission and Warrant Book, 1743-1745, p. 335. 

124 First Lord of the Admiralty. 

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

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



30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 ofif 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." ^-' Today the name "Looe" is peri:>etuated 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-Lctters, ADM 2, vol. 205, p. 344. 



APPENDIX A 

EXTRACTS FROM THE NAVY LIST RELATING TO THE "LOO" 



Period 
April I, 1707-May 31, 1707 
June I, 1707- June 30, 1707 
July I, 1707-September 30, 1707 
October i, 1707-October 31, 1707 
November i, 1707-November 30, 1707 

December i, 1707-January 31, 1708 

February i, 1708-March 31, 1708 
April I, 1708-April 30, 1708 
May I, 1708-July 31, 1708 
August I, 1708-February 28, 1709 
March i, 1709-JuIy 31, 1709 
August I, 1709-September 30, 1709 

October i, 1709-October 31, 1709 
November i, 1709-Novembcr 30, 1709 



December i, 1709-December 31, 1709 

January i, 1710-January 31, 171 o 
February i, I7i(>-February 30, 1710 

March i, 1710-March 31, T710 

April I, 1710-April 30, 1710 
May I, 1710-May 31, 1710 



June I, I7ia-Junc 30, 1710 



July I, 1710-July 31, 1710 

August I, 1710-Scptenibcr 30, 1710 
October i, 1712-October 31, 1712 
November i, 1710-December 31, 1710 
January i, 1711-January 31, 171 1 



Duty 

"Longreach taking in Guns" 

"Going to Archangell" 

"Gon to Archangell" 

"Arch-Angell" 

"Returned with the Russia ships to 
Grimsby" 

"Sheerness (fitting for Newfound- 
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" ^ 

". . . 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" 



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



31 



32 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



Period 
February i, 1711-February 28, 171 1 
March i, 1711-March 31, 1711 
April I, 1711-April 30, 171 1 

May I, 1711-May 31, 1711 
June I, 1711-June 30, 1 71 1 
July I, 1711-September 30, 171 1 
October i, 1711-Octobcr 31, 1711 
November i, 1711-November 30, 1711 
December i, 1711-December 31, 171 1 
January i, 1712-March 4, 17 12 
March 5, 1712-June 30, 1712 

July I, I7i2-July3i, 1712 

August I, 1712-September 30, 1712 

October i, 1712-October 3i, 1712 



Duty 

"Spithead, ordered to the Downes" 

"In Holland to return to the Downes" 

"In 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 of? the Orcades for the 
homeward bound Virginia ships" 

"Nore, ordered to the Downes" 



November i, 1712-November 30, 1712 "Downs, ordered to Bayonne with the 

Transports" 



December i, 1712-Decembcr 31, 1712 
January i, 1713-January 31, 1713 
February i, 1713-February 28, 1713 
March i, 1713-March 31, 1713 
April I, 1713-May 31, 1713 
June I, 1713-Junc 30, 1713 

July I, 1713-July 31, 1713 

August I, 1713-Scpteniber 30, 1713 

October i, 1713-October 31, 1713 

November i, 1713-December 31, 1713 



"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" ^ 



-Ibid., January i, 1710-December 31, 1711, ADM 8/11. 

s Ibid., 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 



LAST CRUISE OF H.M.S. LOO — PETERSON 



33 



Period 
January i, 1714-March 31, 1714 

April I, 1714-April 30, 1714 

May I, 1714-June 30, 1714 

July I, 1714-September 30, 1714 

October i, 1714-October 31, 1714 
November i, 1714-December 31, 1714 
January i, 1715-January 31, 171 5 
February i, 1715-March 14, 1717 
April I, 1717-October 31, 1717 

November i, 1717-December 31, 1717 

January i, 1718-March 25, 1718 
March 26, 1718-March 31, 17 19 

April I, 1719-April 30, 171 9 

May I, 1719-December 31, 1720 
January i, 1721-April 30, 1721 
May I, 1721-January 31, 1722 
February i, 1722-May 31, 1722 
June I, 1722-August 31, 1722 
September i, 1722-December 31, 1725 
January i, 1726-January 10, 1728 
January 11, 1728-March 31, 1728 
April I, 1728-April 30, 1728 
May I, 1728-July 31, 1728 
August I, 1728-December 31, 1728 
January i, 1729-January 31, 1729 
February i, 1729-April i, 1729 
April 2, 1729-May 31, 1729 

June I, 1729-July 31, 1729 
August I, 1729-August 31, 1729 
September i, 1729-September 30, 1729 
October i, 1729-December 31, 1729 
January i, i73(>-March 31, 1730 
April I, 1730-April 30, 1730 



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" ■* 

"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" ^ 

"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" ^ 

"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" 



* Ibid., January i, 1714-December i, 1714, ADM 8/13. 
5 Ibid., January i, I7is-April 30, 1721, ADM 8/14. 
8 Ibid., May i, 1721-December 31, 1725, ADM 8/15. 
^ Ibid., January i, 1726-December 31, 1728, ADM 8/16. 



34 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



Period 
May I, 1730-May 31, 1730 
June I, 1730-June 30, 1730 

July I, 1730-July 31, 1730 

August I, 1730-October 31, 1730 

November i, 1730-January 31, 1731 

February i, 1731-May 31, 1731 
June I, 1731-July 31, 1731 

August I, 1731-August 31, 1731 

September i, 1 73 i-November 30, 1731 

December i, 1731-December 31, 173 1 



January i, 1732-January 31, 1732 
February i, 1732-February 28, 1732 
March i, 1732-March 31, 1732 
April I, 1732-April 30, 1732 

May I, 1732-May 31, 1732 
June I, 1732-Dccember 31, 1734 
January i, 1735-May 4, 1735 
May 5, 1735-June 30, 1735 

July I, 1735-July 31, 1735 
August I, 1735-August 31, 1735 
September i, 1735-October 31, 1735 
November i, 1735-April 30, 1737 
May I, 1737-May 31, 1737 
June I, 1737-December 31, 1739 
January i, 1740-December 31, 1741 
January i, 1742-January 3, 1742 
January 4, 1742-April 30, 1742 



May I, 1742-May 31, 1742 
June I, 1742-Junc 30, 1742 



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 refitting for Channel 

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) 11 
(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 Ibid., January i, i72C)-December 31, 1731, ADM 8/17. 
»Ibid., January i, 1732-December 31, 1734, ADM 8/18. 
1" Ibid., January i, 1735-Decemhcr 31, 1739, ADM 8/19 and 8/20. 
"Ibid., January t, I74(>-Decembcr 31, 1741, ADM 8/21. 



NO. 2 



LAST CRUISE OF H.M.S, LOO — PETERSON 



35 



July I, 1742-August 31, 1742 

September i, 1742-September 30, 1742 

October i, 1742-October 31, 1742 
November i, 1742-November 30, 1742 
December i, 1742-December 31, 1742 
January i, 1743-March 31, 1743 

April I, 1743-April 30, 1743 
May I, 1743-May 31, 1743 

June I, 1743-June 30, 1743 

July I, 1743-July 31, 1743 

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



"Cruizing 50 leagues off Capte Finis- 
terre" 

"Plymouth refitting for Channel Serv- 
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 serv- 
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.i2 The Loo had been lost 
February S, 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 Ibid., January i, 1742-May i, 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. 

1 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 liaving 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 - 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 i 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 of? so 
far as to haul the head sails,'* when the Captain " came and told me the tiller 

^ A 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. i). 

3 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 ) . 

■* A vessel is said to be " at stays" when heading into the wind in tacking. 
° 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. 
8 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 ^^ 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 decks ^- 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 

■^ "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. Ti, 
London, 1781). 

8 The cal)le was 200 yards or one-tenth of a nautical mile. 

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

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

^2 To cut openings in the decks. 



38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 ^^ 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,!-* jq i,^ ^^jjg yauP'' 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 FewelP*' 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. 
1 ' The commanding officers boat. 
15 A heavy double-ended row boat. 
1" 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 oftkers 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. v.fas on board the Snow when cast away but I should have been very 
happy to have found Captain Hardy ^'^ 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/^ has deter- 
mined me to stay here till some opportunity ofifers 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 otlficer 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.^^ 

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

^'' 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. 
1^ 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 


Quartermaster 




*Acleane, W. 


Able Bodied Seaman 




Aiken, James 


Able Bodied Seaman 




Allen, Thomas 


Able Bodied Seaman 




Allman, John 


Landsman 


Deserted September 2, 1742 


Anderson, James 


Quarters Servant 




Angelo, Rogero 


Able Bodied Seaman 




*Arthur, James 


Able Bodied Seaman 




♦Atkinson, Thomas 


Able Bodied Seaman 




Bagster, John 


Alaster 




*Baker, William 


Able Bodied Seaman 




Balderson, William 


Able Bodied Seaman 


Deserted October 14, 1743, in 




and Quartermaster's 


South Carolina 




Mate 




Ball, William 


Surgeon's Mate 




Balls, Benjamin 


Able Bodied Seaman 




Banke, John 


Quarter Gunner 




Baptista, John 


Able Bodied Seaman 


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


Barnes, William 


Able Bodied Seaman 




Barsey, Richard 


Landsman 


Deserted March 30, 1743, at 
Plymouth, England 


Barsey, Thomas 


2d Master's Mate 




♦Bartlctt, Joseph 


Able Bodied Seaman 




♦Basham, Charles 


Gunner's Mate and 
Quartermaster 




Bates, John 


Able Bodied Seaman 





40 



NO. 2 



LAST CRUISE OF H.M.S. LOO 



-PETERSON 



41 



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 
*Buglcss, Stephen 
Bull, John 

Bullman, William 
Burdock, John 
Burns, Patrick 



Able Bodied Seaman 

Able Bodied Seaman 
1st 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 Seaman 

Master 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

and Coxswain 
Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 

Able Bodied Seaman 
Servant 

Able Bodied Seaman 
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 



Deserted November 27, 1743, 

at Port Royal, S. C. 
Deserted September 2, 1742 



Deserted December 19, 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 VOL. I3I 



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, 1743, 
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 



NO. 2 



LAST CRUISE OF H.M.S. LOO 



-PETERSON 



43 



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 
1st 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 



Deserted September 2, 1742 



Deserted October 13, 1743, in 
South Carolina 



Deserted August 11, 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. I3I 



*Duncan, William 

Dunn, George 
*Dunn, Matthew 

Dunstar, James 

Dyar, Anthony 
*Dyer, Darby 
Eades, Thomas 

*Easton, Thomas 
*Ecigecombe, William 

Ellis, Benjamin 
*Ellory, Robert 

Ervin, Adam 

Evans, George 

Fanson, Andrew 
*Farmer, William 

Parrel, James 
Ferguson, Robert 

Field, John 
*Fisher, William 
Fitzgerald, Morris 
Fitzpatrick, Henry 

Fletcher, John 
*Fling, Timothy 

Forrest, Richard 

Forsith, William 
*Forster, Randal 

Fortiene, Joseph 

Eraser, Daniel 

Frost, George 

*Fullmore, Henry 
Gaily, Thomas 
Gibson, George 
Gilbert, Thomas 

*Gilmore, Arthur 

Gilmore, John 
*Gold, William 

Goldsmith, John 
*Good, John 

Goodsides, Abraham 

Gordon, James 

Gordon, Robert 



Able Bodied Seaman 
Able Bodied Seaman 
Surgeon's Mate 
Able Bodied Seaman 

Able Bodied Seaman 
Able Bodied Seaman 
Landsman 

Able Bodied Seaman 
Able Bodied Seaman 
Master's Servant 
Landsman 

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 

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

Able Bodied Seaman 

Sailmaker and Mid- 
shipman 
Boatswain 
Master's Servant 
Able Bodied Seaman 
Able Bodied Seaman 

Quarter Gunner 
Able Bodied Seaman 
Able Bodied Seaman 
Ordinary Seaman 
Surgeon's 2d Mate 

Midshipman 
Midshipman 



Deserted October 5, 1743, at 
New York 



Deserted September 2, 1742, 
at Plymouth, England 



Deserted December 12, 1742, 
at Portsmouth, England 



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

Deserted November 27, 1743, 
at Port Royal, S. C. 



Deserted April 16, 1743, at 
Plymouth, England 



Deserted April 13, 1743 



Deserted October 8, 1742 



NO, 2 



LAST CRUISE OF H.M.S. LOO — PETERSON 



45 



Gorman, David 
Graham, Matthew 
*Graham, Samuel 
Grant, Henry 

*Green, Henry 

Gregory, Jo. 

Gresham, Charles 

Grossier, John 

Grun, Nicholas 
*Hallct, 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 
♦Higgcnson, James 
♦Higginson, John 
♦Highmas, Thomas 
♦Hinds, Michael 
♦Hogg, Peter 

HoUiday, 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 



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 MISCELLANEOUS COLLECTIONS VOL. I3I 



Horseman, Arthur 

Hughes, John 

*HulI, 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 
*Kno\vlcs, 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 
1st Lieutenant's Serv- 
ant 
Quarter Gunner 



Died March 24, 1742, at 

Plymouth, England 
Deserted December 12, 1742, 

at Portsmouth, England 



Deserted August 11, 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 



LAST CRUISE OF H.M.S. LOO — PETERSON 



47 



*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 

*McNcal, William 
Meachem, James 



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

Able Bodied Seaman 

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

and Midshipman 
Ordinary Seaman 
Able Bodied Seaman 

Ordinary Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Quartermaster 
1st Lieutenant's Serv- 
ant and Able Bodied 
Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 

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



Deserted April i6, 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 VOL. I3I 



Mellan, John 


Boatswain's Mate 


Meridith, John 


Ordinary Seaman 


Mignam, George 


Surgeon 


*Miller, James 


Able Bodied Seaman 


♦Miller, Nicholas 


Able Bodied Seaman 


Miller, Stephen 


Boatswain's Servant 


Mills, Nathan 




♦Millsom, James 


Able Bodied Seaman 


Mitchell, R. 


Ordinary Seaman 


*Molineaux, John 


Ordinary Seaman and 




Able Bodied Sea- 




man 


Moore, Samuel 


Yeoman 


More, Thomas 


Able Bodied Seaman 


Morran, William 


Able Bodied Seaman 


Morrison, John 


Able Bodied Seaman 


Morrow, Hugh 


Captain's Servant 


*Mortimer, Robert 


Able Bodied Seaman 


Murphy, Francis 


Able Bodied Seaman 


*Murphy, James 


Able Bodied Seaman 


♦Murphy, John 




Murphy, Patrick 


Able Bodied Seaman 


Murphy, Samuel 


Able Bodied Seaman 


♦Nagan, James 


Ordinary Seaman 


*Nellson, Robert 


Able Bodied Seaman 


♦Newgent, Patrick 


Able Bodied Seaman 


*Ne\vson, Richard 


Able Bodied Seaman 


Nobbs, Stephen 


Landsman 


Norman, Edward 


Ordinary Seaman 



Deserted April 13, 1743 



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



♦Nuikle, Robert 
Ogburnc, John 
Oliver, Richard 
*Orr, Archibald 
*Osbourne, William 
Parker, Peter 
Parr, William 

*Parrott, Samuel 
Patrick, Richard 
Patten, Hugh 

Pearse, Safnuel 



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 September 2, 1742 



Deserted September 10, 1742, 
at Plymouth, England 



Deserted August 11, 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 



NO. 2 



LAST CRUISE OF H.M.S. LOO 



-PETERSON 



49 



*Pearson, Thomas 



Ordinary Seaman and 
Able Bodied Sea- 
man 



Pegan, Roger 


Able 


Bodied 


Seaman 


Pegan, Thomas 


Able 


Bodied 


Seaman 


Pelican, John 


Able 


Bodied 


Seaman 


Pemell, Thomas 


Able 


Bodied 


Seaman 


♦Peters, William 


Able 


Bodied 


Seaman 




and 


Midshipman 


Phallcm, Edmund 


Able 


Bodied 


Seaman 


*Phillips, Thomas 


Cook 






*Phonix, Philip 


Able 


Bodied 


Seaman 


♦Pickering, Michael 


Clerk 






♦Plantain, Charles 


Able 


Bodied 


Seaman 


Poole, Thomas 


Pilot 






Potterfield, George 


Able 


Bodied 


Seaman 


Powell, Philip 


Able 


Bodied 


Seaman 



Pownswell, Edward Able Bodied Seaman 



♦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 



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 
1st 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 



Deserted November 27, 1743, 
at Port Royal, S. C. 

Deserted March 17, 1742, at 
Plymouth, England 



Deserted March 30, 1743, nt 

Plymouth, England 
Deserted December 12, 1742, 

at Portsmouth, England 



Deserted October 6, 1743, at 
New York 



50 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



Risden, Joseph 
Roach, Henry 

Roberts, David 

Roberts, Edward 

Roberts, John 
Roberts, John 
Roberts, Joseph 
♦Roberts, WilHam 
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 
*SaIter, 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 
1st Captain's Servant 
Able Bodied Seaman 
Able Bodied Seaman 
Able Bodied Seaman 
Gunner 
Landsman 

Able Bodied Seaman 
1st Lieutenant's Serv- 
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 Servant 

and Able Bodied 

Seaman 



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 



LAST CRUISE OF H.M.S. LOO 



-PETERSON 



51 



Softley, Robert 

Somerwel, Joseph 
Southard, Thomas 



Able Bodied Seaman 

Able Bodied Seaman 
Able Bodied Seaman 



Spare, Thomas 


Able Bodied Seaman 


Sparks, Joseph 


Able Bodied Seaman 


♦Spinks, Stephen 




Spry, Nicholas 


Able Bodied Seaman 


Stanford, Richard 




Stapleton, A. 


Quartermaster's Mate 


Stephenson, James 


Landsman 


*Steuart, Neil 


Surgeon 


Stevenson, Henry 


Able Bodied Seaman 


Stewart, Alexander 


Able Bodied Seaman 


*Stiveash, Stiven 


Able Bodied Seaman 


*Stoneham, John 


Able Bodied Seaman 


Stradder, Forbel 


Landsman 


Stroud, John 


Boatswain 


♦Sullivan, Thomas 


Ordinary Seaman 


Swain, William 


3d Lieutenant's Serv- 




ant and Captain's 




Servant 


*Sweeny, Daniel 


Able Bodied Seaman 


*Swickman, Thomas 


Landsman 


Swift, Theodore 


Able Bodied Seaman 


*Tabler, Thomas 


Captain's Servant and 




Able Bodied Sea- 




man 


♦Taylor, John 


Carpenter's Servant 




and Able Bodied 




Seaman 


♦Taylor, Jonathan 


Able Bodied Seaman 


♦Taylor, William 


Captain's Servant and 




Ordinary Seaman 


Thatcher, John 


Quartermaster 


♦Thompson, Richard 


Pilot 


Thoyer, Peter 


Able Bodied Seaman 


♦Tipper, John 


Able Bodied Seaman 


Tobyn, George 


Able Bodied Seaman 


Todd, Alexander 


Able Bodied Seaman 


♦Torginton, Joseph 


Carpenter's Servant 


Treacey, William 


Able Bodied Seaman 



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 VOL. I31 



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

*Trunker, William 
Tunis, Michael 
Turford, Thomas 

*Utting, Ashby 
Vincent, Aaron 
*Vivian, John 
nVadlin, Richard 
*Walkcr, James 
Walker, John 

Walker, Peter 

WalHs, 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 
♦Willmot, Theodore 

Wills, J. 

Wilson, George 

Wilson, Lawrence 
Wise, John 

Woodgate, William 
♦Wotton, William 
Wright, William 

♦Yeates, James 
♦Young, John 
Younger, John 



Able Bodied 
Able Bodied 
Able Bodied 
Able Bodied 
3d Lieutenant' 

ant 
Able Bodied 
Able Bodied 
Carpenter 



Seaman 
Seaman 
Seaman 
Seaman 

s Serv- 

Seaman 
Seaman 



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 

1st Lieutenant's Serv- 
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 



NO. 2 



LAST CRUISE OF II. M.S. LOO 



-PETERSON 



53 



MARINE LIST 



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

*BaII, 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 



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 VOL. I3I 

♦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 
♦Woodeson, John 

Woodley, Nicholas 
Colonel Wynyard's Regiment 

Aldridge, William 

Chappel, Samuel 

Qayton, Joseph 

Hancock, Justinian 

Hope, Richard 
♦Kent, Benjamin 
♦Korgett, John 

Parker, Daniel 
♦Stokes, William 
♦Trowes, Richard 
♦Wright, Samuel 



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 1743 i 

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 

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



APPENDIX E 

COMMANDING OFFICERS OF THE "LOO" 

1. Harland, Robert April 4, 1707-October i, 1709 

2. Herbert, Jona October i, 1709- ? 

3. Walrund, Beau 1709- 1715 

4. Splain, Tim April I7i7( ?)-December 1717 

5. Protherie, George April 1718- 1720 

6. Scott, — (Captain) October 1720- 1722 

7. Waterhouse, Tho. (Commodore) February 1727/28-October 1728 

8. Lee, FitzHcnry October 22, 1728-April 2, 1729 

9. Berkeley, William April 2, 1729- ? 

10. Best, John May 5, 1735-May 1737 

11. Northesk, Earl of January 4, 1741/42-September 1742 

12. Dent, Digby September 17, 1742-April 1 1, 1743 

13. Utting, Ashby April 1 1, 1 743-February 5, 1743/44 

1 Based on the Navy List, 1707-1744. 



55 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 2, PL. 2 





I. Copper cciins recovered from tlie wreck site (if the L(U) in the summer of 
1950. Left to right: Spanish 8 maravedi circa lOoo. Spanish 8 maravedi circa 
1C40, and Swedish \ ore dated 1720. This last coin was important evidence 
in dating the wreck. Lent to the V. S. National Aluseuni hy Dr. and Mrs. 
George Crile, Jr. 



,^ 




2. Small (Jncen .\nne pewter teapot fmni the wreck of tlic Loo. Gift 
to the National Mnseum from K. .\. Link. The pot was damaged hy fire 
when the Loo was hurncd hy her captain after heing wrecked. 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 2, PL. 3 






'• ' I, 




SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 2, PL. 4 




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VOL. 131, NO. 2, PL. 6 




K"^. 



A basket of coral-sand-encrusted shot coming u]i from the l)ottoni. 
(Photoyraph by Dr. George Crilc, Jr.) 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 2, PL. 7 




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SMITHSONIAM MI^rELLANEOUS COLLECTIONS 



VOL. 131, NO. 2, PL. 13 




K. A. Link riyging the main boom of the Blue Hcroii to raise a i-ton iron barrel from 
the wreck site. (Photograph by Mrs. George Crile, Jr.) 



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 Kertesz (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 Museum 
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 Museum d'Histoire Naturelle in 
Paris. The Neotropical species of Tabanidae described by Francis 
Walker between 1848 and i860, 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. I3I 

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 Museum 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 Memoires de la 
Societe Royale des Sciences, de I'Agriculture et des Arts de Lille, and 
in a separately published form put out by Librairie Encyclopedique de 
Roret, Paris. There is no difference in the text, but the pagination is 
different, and in some cases the reprint is of an earHer 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 Dipteres Nouveaux" published in 1892 in Mem. 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. pniinosus 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 w^as 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 abiciis Wlk. 1848, List, i : 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 albiscittellatus Macq. 1850, Dipt. Exot., Suppl. 4:34. Mexico = /<?m- 

caspis Wied. (Osten Sacken, 1878; Krober, 1934; Fairchild, 1941). Perhaps 

in Nearctic coll. 
Tabanus albiviitatns Macq. 1834, Hist. Nat. Dipt., i : 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 argentijrons Wlk. 1848, List, 1:186. Loc? B. M. 
Pangonia bicolor Macq. 1850, Dipt. Exot., Suppl. 4:27. Mtx'xco ^= Esenbeckia 

semijlava Wied. (Bellardi, 1859; Krober, 1934). Homonym of P. bicolor 

Macq. 1846. New Holland. 
Hacmafopota biznttata Macq. 1834, Hist. Nat. Dipt., i : 212. Amer. Merid. = 

Diachlorus bivittatus Wied. 1828 (Krober, 1934). From description, a Dia- 

chlorus, though not credited to Wiedemann in original description. 
Tabamis bivittatus Macq. 1845, Dipt. Exot., Suppl. i : 35. Brazil. Coll. Spinola. 
Tabanus bonarietisis Macq. 1838, Dipt. Exot., i (i):i42. Buenos Aires. Coll. 

Serville. Krober (1934) places in Agclanius with acupuuclatus Rond. 1868, 

as synonym. . 

Tabanus brcvivilta Wlk. 1850, Newman's Zoologist, 8, App., p. xcvi. Loc? 

B. M. Homonym of T. brcvivilta Wlk. 1848. Australia. 
Tabamis castancus Macq. 1834, Hist. Nat. Dipt., i : 198. Cayenne. The descrip- 
tion indicates probable synonymy with T. unicolor Wied. 
Tabanus castaneoventris Macq. 1838, Dipt. Exot., i (i) : 152. Loc? Mus. Paris. 
Tabanus chrysolcucus Wlk. 1854, List, 5, Suppl. i : 327. Brazil. B. M. 
Tabanus dcspectus 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 Krober. 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. tacniola P. de B. 

Tahamis flammans Wlk. 1848, List, i: 153. Loc? B. M. 

Tabanus flaznbarbis Macq. 1845, Dipt. Exot., Suppl. i : 169. Cayenne. Coll. 
Spinola. Krober (i930h) 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. i : 36. Colombia. Coll. de 
Breme. 

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

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

Tabanus fullo Wlk. 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 julvifasciatiis Macq. 1834, Hist. Nat. Dipt., 1:206. Loc? Coll. 
Percheron. 

Tabanus julviger Wlk. 1850, Dipt. Saund., i : 65. Loc? B. M. 

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

1859. 
Tabanus gigas Macq. 1834, Hist. Nat. Dipt., 1:200. Loc? A homonym of 

T. gigas Herbst 1787. 
Pangonia hcbes Wlk. 1848, List, i : 137. Loc? B. M. Ricardo (1901) says type 

lost, but see Oldroyd (1954). Probably Australian. 
Tabanus hispidus Wlk. 1850, Dipt. Saund., i : 63. Loc? B. M. 
Tabanus honcstus Wlk. 1850, Dipt. Saund., 1:64. Loc? B. M. Lutz (1907) 

lists as possible synonym of fnscofasciatus 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 lagenafcrus Macq. 1838, Dipt. Exot., i (i) : 148. Loc? Mus. Paris. 
Tabanus longifrons Krober 1930, Dipt. Pat. S. Chile, S (2) : 152. Chile. B. M. 

Apparently not returned by Kruber. 
Tabanus longipcnnis Macq. 1834, Hist. Nat. Dipt., i : 201. Loc? Coll. Percheron. 
Chrysops lugubris Macq. 1846, Dipt. Exot., Suppl. i : 44. Brazil. Coll. Robyns, 

Bruxelles. The description suggests possibility that this was not a Chrysops. 
Tabanus macuUpcnnis Macq. 1834, Hist. Nat. Dipt., 1 : 198. Brazil. Coll. Serville. 

A homonym of T. macuUpcnnis Wied. 1828, and Brulle 1832. Not same as 

macuUpcnnis Macq. 1846. Not listed by Krober (i934)- 
Tabanus macuUpcnnis Macq. 1846, Dipt. Exot., Suppl. i : 34. Brazil. Coll. 

Spinola. A homonym of macuUpcnnis Wied. 1828, and Macq. 1834, but a 

different species, probably a synonym of Tabanus (Hyboviitra) qimdripunctata 

and the genotype of Dasyphyrta End. (Krober 1934)- 

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

Tabanus microccrus Wlk. 1848, List, i: 150. Loc? B. M. 

I 



NO. 3 NEOTROPICAL FLIES, TABANIDAE — FAIRCHILD 7 

Tabanus nigripalpis Macq. 1845, Dipt. Exot., Suppl. i : 40, pi. 4, figs. 7-8. New 

Grenada. Coll. Bigot. Placed by Krober (i93ii, 1934) in Catachlorops, near 

nijescens Fab. A specimen, headless, in Mus. Paris labeled as rujescens 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 olivaceivcntris Macq. 1847, Dipt. Exot., Suppl. 2: 18. Brazil, Para. Coll. 

de Villiers. 
Tabanus opulenUis Wlk. 1848, List, i : 148. Loc? B. M. 
Tabanus ontatifrons 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 

Krober (1934) as a possible synonym of saussurei Bell. In Nearctic Coll. at 

Mus. Paris, where seen by Philip (i9S4b). 
Tabanus planus Wlk. 1850, Dipt. Saund., 1:61. Loc? B. M. 
Pangonia prasiniventris Macq. 1846, Dipt. Exot., Suppl. i : 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 Wlk. 1850, Dipt. Saund., i : 36. Brazil, Rio de Janeiro. Lutz 

(igoy) =: occidentalis Wied. Listed as (Neotabaniis) by Krober (1934), but 

he saw no specimens. 
Tabanus redactus Wlk. 1850, Dipt. Saund., i : 66. Loc? B. M. 
Tabanus ruber Macq. 1845, Dipt. Exot., Suppl. i : 42. Mexico. Coll. Guerin. 

A homonym of T. ruber Thunb. 1827. Renamed subruber by Bellardi, 1859. 

May be among Nearctic species in Mus. Paris. 
Pangonia rufa Macq. 1838, Dipt. Exot., 1(1): no. Lima (Peru). Coll. Serville. 

There are specimens determined by Bigot and Krober in B. M. 
Tabanus scutellatus Macq. 1839, Dipt. Exot, i (2) : 186. Loc? Mus. Paris. 

Krober (1934) claims to have seen the type and says ii=z macula Macq., but 

I was unable to find it and doubt the synonymy. In any case scutellatus has six 

years' priority. 
Tabanus secundus Wlk. 1848, List, i : 180. Loc? B. M. 
Tabanus subsenex Wlk. 1850, Dipt. Saund., i : 38. S. America. B. M. Lutz 

(1907) = triangulunt Wied. 
Tabanus surinamensis Macq. 1838, Dipt. Exot., 1 (i) : 132. Surinam. Coll. Ser- 
ville. The description suggests T. nebulosus de Geer. Krober (1934) placed in 

Tabanus with a query. 
Pangonia translucens Macq. 1845, Dipt. Exot., Suppl. i : 26. Brazil. Coll. Guerin. 

The species has long been placed in Esenbeckia. (Krober, 1934; Fairchild, 

I942d). 
Tabanus trifarius Macq. 1838, Dipt. Exot., i (i):i44. 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 "7". erythrogaster 

Colomb." and also as "T. variventris J. Macq. 2e. Suppl. Colombie." The 



8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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

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

Tabanus viridiflavtis Wlk. 1850, Newman's Zoologist, 5, App., p. Ixvi. Brazil. 
B. M. Lutz (igo7) = 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., i (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 ]\Iacquart 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 jrontalis Macq. 1838. i 6 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 scutcllatus Macq. 1839. 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 Museum 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 
Fidenu and Melpia of the catalog. Scaptia s.s. includes Osca and Cal- 
liosca. Scaptia subgenus Pseudoscione includes Listriosca, Listrapha, 
Parosca, Listraphella, and probably Lilaenaoi the catalog. Stenotabanus 
includes all the small Tabaniis-like species with bare subepaulets and at 
least some of the species placed in Stypommia 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 Hemichrysops 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- 



lO 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 Wlk. 1850, Dipt. Saund., 1:34. (Tabanus) B. M.(H) = *rubigini- 

pennis Macq. 1845, Krober (i934, 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.). 
*alb'idicollis Macq. 1850, Dipt. Exot., Suppl. 4:32. (Tabanus) B. M.(H) = 

T. importunus Wied., Krober (1934). Fairchild (i942f). 
*albidocinctus Big. 1892, 5:686. (Tabanus) B. M. = Stenotabanus. Krober 

(1934) as Leiuotabanns. 
♦albifasciata Macq. 1845, Dipt. Exot., Suppl. i : 28. (Pangonia) B. M. = Scione. 

Krober (1934) has it albojasciata in error. 
*albifrons Macq. 1838, Dipt. Exot., i(i):lo8. (Pangonia) M. P.(H) = 

(Pseudoscione). Krober (1934) as Lilaeina Borg. Genotype. 
*albipecttts Big. 1892, 5:611. (Mycieromyia) B. M. Krober (1933b) = Fidena 

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

(I929d). 
*albithorax Macq. 1838, Dipt. Exot., i (i):i07. (Pangonia) M. P. = Scaptia. 

Krober (1930k, 1934) as Osca. 
*alboater Wlk. 1850, Zoologist, 8, App., p. Ixvii. (Tabanus) B. M.(H) = 

*T. atricornis Big. 1892. =: T. albibarbis Wied., Krober (1932b). *T. angusti- 

fron^ Macq. 1847, *T. settlor 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. Angusiijrons 

has clear wings and slightly different antennae and palps. 
*albohirtus Wlk. 1857, Trans. Linn. Soc. London, 17:338. (Tabanus) B. M. = 

Dasybasis. Krober (i93oi, 1934) as Agelanius. 
*albomaculatus Wlk. 1854, List, 5:207. (Tabanus) B. M.(H)=*T. discifer 

Wlk. 1850. Krober (1931c) as Gymnochcla discifer; (1934) as Chelommia 

discifer. 
*alboniarginata 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 (igo7) =*Dichelacera marmorata Big., Lutz (1907). 
*albovarius Wlk. 1854, List, 5:206. (Tabanus) B. M.(H)=*r. unicinctus 

Wlk. 1857 = *Leuco tabanus leuconotum Fchld. 1941 (N.S.). 
*albovitlatus Krob. 1930, Dipt. Pat. S. Chile, 5, fasc. 2:146. (Therioplectes) 

B. M. = *Dasybasis scutulatus Krob. 1930. (N.S.) Krober (1934) as 

Sciladyn us. 
*alci3 Will. 1896, 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 

*alteripenms Wlk. i860, Trans. Ent. Soc. London, 5: 275. (Tabanus) B. M.(H) 

= T. caliginosus Bell. 1859, Hine (1925). Fairchild (i953) as (Philipo- 

tabanus). Philip (1954). 
♦altivagus O. S. 1886, Biol. Cent.-Amer., Dipt., 1:45. (Chrysops) B. M. 
*anwbilis Wlk. 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.) 
*angustijrons 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. luciduhts Beq. 1940 in part. = T. lu- 

cidulus Fchld. 1951. 
♦angustus Macq. 1838, Dipt. Exot., i (i) : 136. (Tabanus) M. P.(H)=*r. 

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, 1:198. (Chrysops) B. M.(H) = Diachlorus 

ferrugatus Fab. 1805. 
*ataenia Macq. 1838, Dipt. Exot, i (i) : 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) = *r. alboater Wlk. (q. v.) 

= ? T. albibarbis Wied. Not T. atricornis Meig. 1838. 
*atrifcra Wlk. i860, Trans. Ent. Soc. London, 5:272. (Pangonia) B. M. = 

Pangonia haustellata (Fab.) 1781. Palearctic. Philip (1954). 
attenuatus Wlk. 1848, List, i: 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. 
*auribarbis Macq. 1847, Dipt. Exot., Suppl. 3:12. (Tabanus) B. M.(H) = 

*Dasychela macula Macq. 1845. Krober (1934, 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, Krober (1934). 
*aurisquammatus Big. 1892, 5:665. (Atylotus) B. M. = Atylotus fulvus Meig. 

Palearctic. (N.S.) Lutz (1907) = T. tw/co/or Wied. 
*auroguttata Krob. 1930, Zool. Anz., 90(3-4) : 71. (Chrysops) B. M. Bequaert 

(1944). A distinct species from C. auroguttata pallidijemorata Krob. 
♦aurora Macq. 1838, Dipt. Exot., 1(1): 142. (Tabanus) M. P. Not T. aurora 

of Lutz (1914, 1918), Bequaert (1926), or Krober (1929c). Near T. jerreus 

Wlk. 1848 and T. impressus Wied. 1828. 
*austeni Krob. 1930, Zool. Anz., 86(11-12) : 294. (Tabanus) B. M. Not Tabanus 
(Ochrops) austeni Szil. (1915, Ent. Mitt. Berlin, 4:100). For Tabanus 



12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

(Phaeotabatiiis) aiisteni Krob. I hereby propose the name Tabanus sannio, 
nom. nov. 

*badia Wlk. 1848, List, i : 132. (Pangonia) B. M. = Fidena venosa Wied. 1828, 
Krober (i930g) as Sackenimyia venosa; (1934) as Melpia. 

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

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

*basalis Wlk. 1848, List, i : 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 
Krober (1934)- 

*basi-rufus Wlk. 1850, Dipt. Saund., i : 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. induus Wlk. 1850. ^ *T. bitinctus Wlk. 1857. = ? T. mar gin en evr is 
Macq. 1855. Type not found, but specimens det. Bigot in B. M. agree with 
basizntta. (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). Krober (1934). Not Tabanus tristis Fab. 1798. 
*bifascies Wlk. 1848, List, i: 191. (Dichelacera) B. M.(H). Barretto (1949b) 

as Rhamphis End. 
*bifenestratU3 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. Krober (1934). Fairchild (1942b) as 

Bellardia. 
*bitincfus 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. =r 

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) r= ? rubrithorax Macq. 1838. 
♦callosus Macq. 1848, Dipt. Exot., Suppl. 3:11. (Tabanus) B. M.(H) = 

? T. trizittatus Fab. Not T. callosus Fairchild 1942c. Type lacks antennae 

and palps and further series may show intergrading with other named forms, 

as in lineola 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 prcsbiter Rond. 1863. Krober (1929a) as Styponimia. 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.(n) zn Dicladocera. Krober 

(1931c) as var. of Gyvinochela satanica Big.; (1934) as syn. of Chclovmiia 



NO. 3 NEOTROPICAL FLIES, TABANIDAE — FAIRCHILD 13 

satanica Big. Not Chelommia castanca Barr. 1949a. Subepaulets bare. Type 

agrees quite well with a specimen det. Fairchild as Dicladocera imicolor Lutz 

from the description. Barretto (1948) as syn. of Amphichlorops satanica Big. 

= ? Amphichlorops jerruginea Barr. 1948. 
*casianea Sure. 1919, Mes. Arc. Merid. fiquat. Amer. du Sud, 10:222. {Ere- 

phopsis) M. P. = Fidena castaneiventris Krob. 1934, nom. nov. Not Fidcna 

castanca (Perty) 1830. 
*caustica O. S. 1886, Biol. Centr.-Amer., Dipt, i : 44- {Pangonia) B. M. = 

Esenbeckia tviedemanni Bell., Krober (1934)- Philip (i954a) as a distinct 

species. 
*chilensis Macq. 1838, Dipt. Exot, 1(1) : 145- (Tabanus) M. P.(H) = Dasy- 

basis. Krober (1934) as Agclatmis. 
*chionostigma O. S. 1886, Biol. Centr.-Amer., Dipt., i : 54- (Tabanus) B. M. == 

Stibasoma, Fairchild (1940b). Labels on type do not agree with original 

description. 
*cinerascens Big. 1892, 5:610. (Mycferomyia) B. M. Krober (i93of). 
*cingulifcr Wlk. 1857, Trans. Ent. Soc. London, 4(5) : 123. (Tabanus) B. M. = 

Leucotabanus exaestuans Linn. 1767. 
*claripemiis Big. 1892, 5:675. (Atylotus) B. M.(H) = Tabanus hookcri 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 Chclotahamis 

fuscus Wied. 
*colombensis Macq. 1846, Dipt. Exot., Suppl. i : 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. (Mclpia) B. M.(H)=: 

Fidena, Krober (1934); Fairchild (1951a). 
*commixtus Wlk. i860, Trans. Ent. Soc. London, 5:273. (Tabanus) B. M.(H) 

=:*Tabanus maya Bequaert 1932. Fairchild (1942c). Krober (1934) as syn. 

of T. lincola Fab. (N.S.) 
*communis Krob. 1931, Stett. Ent. Zeitg., 92: 282. (Agelanius) B. M. = *T. fre- 

quens Krob. 1934, nom. nov. Not T. communis Krob. 1930. = Dasybasis, 

with bare subepaulets. 
*compactus Wlk. 1854, List, 5, Suppl. i : 222. (Tabanus) B. M.(H) z= Stibasoma 

fulvohirtum Wied. 1828, Osten Sacken (1886). 
*comprchcnsa Wlk. 1850, Dipt. Saund., i : li. (Pangonia) B. M.(H) = Elaphella 

cervus Wied. 1828, Ricardo (1904). 
*con finis Wlk. 1848, List, 1:160. (Tabanus) B. M. = ? T. aurilineatus Sch.- 

Stekh. 1926, Oriental. Not T. con finis Zett. 1840. Lutz (1907) = T. tacniotcs 

Wied. Krober (1933a) as (Neotabanus). Not Neotropical, in my opinion. 
*confligcns Wlk. 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., i : 49. E. India. = T. cinerarius 

Wied. 1828, Krober (1931c, 1934) as Chelommia or Gynmochcla. 
*conica Big. 1857, Ann. Ent. Soc. France, ser. 3, 5: 278. (Pangonia) B. M.(II) 

= Mycteromyia Philippi, genotype. Krober (i93of). 
♦consequa Wlk. 1850, Zoologist, 8, App., p. cxxi. (Tabanus) B. M.(H) = 
? T. lineola var. carncus Bell., Fairchild (1942c). = *7". globulicallosns 
Krob. 1931. =: ? T. dorsiger var. pallidefemorata Krob. 1929. = ? T. ochro- 



14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

philus Lutz 1914. This is the small pale form of lUieola 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 Wlk. 1848, List, i : 198. (Chrysops) B. M. = Diachlorus ferrugatus 

Fab., Ricardo (1904). 
*cornuta Wlk. 1857, Trans. Linn. Soc. London, 17(3): 337. (Pangotiia) 

B. M.(H) =Rhabdotylus planiventris (Wied.) 1828, Lutz (1907). Krober 

(1934) as Amphichlorops. 
*corone O. S. 1886, Biol. Centr.-Amer., Dipt., 1:51. {Tabamis) B. M.(H). 
*cribellum O. S. 1886, Biol. Centr.-Amer., Dipt., 1:52. (Tabanus) B. M. ^ 

Stenotabanus, Stone (1938); Philip (1950). 
*cyanami Wlk. 1848, List, i : 208. (Hadrus) B. M. = Selasoma tibiale Fab., 

Ricardo (1904). 
*depressa Macq. 1837, Ann. Soc. Ent. France, 6:429; 1838, Dipt. Exot., 1: iii. 

(Pangonia) B. M. and M. P.(H) = Scaptia lata Guerin 1830. Walker 

(1850), genotype of Osca. Rondani (1863), genotype of Diatomineura. 
*dcrwatus 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. (Tabamis) B. M.(H) = 

*T. nuntius Wlk. 1854. = *T. untznttatus Macq. 1855. = T. sallei Bell. 1859. 

= T. angustivitta, Fchld. 1942. = *T. (Agelanhis) 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 Macrocorvnis. 
♦discifer Wlk. 1850, Dipt. Saund., 1:35. (Tabanus) B. M.(H)=*r. albo- 

viaculat us Wlk. 1854. Krober (1931c) as Gymnochda; (1934) as Chclommia. 
*discifer Big. 1892, 5:684. (Tabanus) B. M.(H)=3*T. desertus Wlk. 1850. 

Not *T. discifer Wlk. 1850. (N.S.) 
*diversipcnnis Wlk. 1848, List, 1:165. (Tabanus) B. M.(H) = *Esenbeckia 

fascipennis Macq. 1838, Krober (1934). 
*diversipcs Macq. 1848, Dipt. Exot., Suppl. 3:13. (Diabasis) B. M. = Dia- 
chlorus bicinctus Fab. 1805. Lutz (1913) ; Krober (1928b). 
*dives Wlk. 1848, List, 1:166. (Tabanus) B. M.(H) = *Stibasoma flavi- 

ventre Macq. 1847, Lutz (1915). Krober (1934) as var. of julvohirtum 

Wied. 
*dominicanus Krob. 1931, 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 Par osca; (1934) as Listrapha. 
♦dorsovittatus Macq. 1855, Dipt. Exot., Suppl. 5:30. (Tabanus) B. M.(H). 

z=zT. 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, Biul. Centr.-Amer., Dipt., 1:49. (Tabanus) B. M.(H) = 

(Philipotabanus) subgenotype. Fairchild (i942f). 



NO. 3 NEOTROPICAL FLIES, TABANIDAE — FAIRCHILD I5 

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

*clongatHS Macq. 1845, Dipt. Exot., Suppl. 1:38. (Tabamis) 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. 

endcrleini Krob. 1931, Zool. Anz., 94(9-10) : 252. (Esenbeckia) B. M. The 
holotype c? is in Berlin. The specimens in B. M. det. Krober, i d* and 2 $, do 
not agree with Krober'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. Merid. fiquat. Amer. du Sud, 10:219. 
(Scione) M. P.(H). Krober (1930J). 

*erebus O. S. 1886, Biol. Centr.-Amer., Dipt., 1:50. (Tabanus) B. M.(H) = 
Astigmatophthalmus satamis Krob. 1931, Stone (1938). Krober (1934) as 
syn. of T. alteripennis Wlk. Fairchild ( I942f ) 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 ^/j'/o/M.y o-yf/jra^MJ Big. 1892, 5 : 661. Krober (1934) as 
syn. of T. monochroma Wied. 

*erythrocephalus Big. 1892, 5:668. (Atylotiis) B. M.(H) =:Bolbodimyia, 
Krober (i93oh). Stone (1954). 

*erythronotata Big. 1892, 5:612. (Mycteromyia) B. M.(H) = Fidena. Lutz 
(1909) as Bombylopsis. Krober (1934) as Melpia. 

*eittaeniatns 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. 

*excclsHS Sure. 1919, Ales. Arc. Merid. fiquat. Amer. 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 Sure, indicating 
an intention on Surcouf's part to change his homonym; excelsior is hereby 
proposed for excelsus Sure. 1919, not Ricardo 1913. Krober (1934) as 
? Stypommia. Bequaert and Renjifo (1946) as Agelanius. = Dasybasis. 

*exeuns Wlk. 1850, Dipt. Saund., i : 12. (Pangonia) B. M. = ? (Pseudoscione) 
molesta Wied. 1828. Near seminigra Ric. Enderlein (1925) as Melpia geno- 
type. Not Melpia Wlk. Krober (1930k) as syn. of Parosca molesta Wied.; 
(1934) as Listrapha molesta Wied. 

*fallax Macq. 1847, Dipt. Exot., Suppl. 2:20. (Tabamis) B. M. Not T. fallax 
Macq. 184s, Africa. Krober (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 straiglitened out. 

*fasciata Macq. 1834, Hist. Nat. Ins. Dipt., i: 194; 1838, Dipt. Exot., 1(1) : 107. 
(Pangonia) M. P. Not P. jasciata Latr. 181 1, Egypt. Lutz (1907) as syn. 
of Esenbeckia esenbeckii Wied. 1830. There are 10 $ 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). 



l6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Osten Sacken (1886), Ricardo (1904), Krober (1934) as syn. of D. cerzn- 
coniis Fab. (N.S.) 

*fascipennis Macq. 1845, Dipt. Exot., Suppl. 1:35. (Tabanus) B. M.(H) = 
(Philipotabanus) Fairchild (i942f). 

*fascipennis Macq. 1838, Dipt. Exot., 1(1): no. {Pangonia) M. P.(H) = 
*Esenbeckia diver si pennis Wlk. 1848, Krober (1934). 

*jascipennis Krob. 1930, Zool. Anz., 88(9-10) 1237. (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 jascipennis Macq. 1845. The new specific name of 
vecordis is hereby proposed and the species may be known as Tabanus 
(Hemichrysops) vecordis, nom. nov. 

*feuestratus Alacq. 1838, Dipt. Exot., i(i):i39. (Tabanus) M. P.(H) = 
Pachyschelomyia notopleuralis Barr. 1950. Brethes (1910) and Krober 
(1934) as Siibasoma. 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. feuestratus Fab. 
1794. (N.S.) 

*fenestrata Macq. 1845, Dipt. Exot., Suppl. 1:26. (Pangonia) B. M. = 
(Pseudoscione). Lutz (1909) as Diatomineura. Krober (1930k) as Par osca; 
(1934) as Listraplia. Very close to seminigra Ric. and longipcnnis Ric. 

♦ferreus Wlk. 1848, List, 1:151. (Tabanus) B. AL(H). Lutz (1907) = ? ? 
D. rufipennis Macq. Krober (1932b) as Chclotabanus jcrreus. Very close to 
T. impressus Wied. 1828, but darker. 

*ferrifer Wlk. 1850, Dipt. Saund., 1:30. (Tabaniis) B. M. (H) = Tabanus 
nebulosus de Geer, Philip (1952). Fairchild (i942f). 

*jcrruginea Macq. 1839, Dipt. Exot., 1(2): 295. (Pangonia) M. P.(H) = 
*Esenbeckia illota Will., Fairchild (i942d). Not Tanyglossa ferrugin^a 
Latr. 1809. The type is like the form treated by Fairchild (i942d) as E. i. 
enderleini 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 *jacti- 
lator Fchld. and jidz'ivcntris Macq. 

*fiIiolus Will. 1901, Biol. Centr.-Amer., Dipt., i, 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 (1932a). 

♦flavinotum Krob. 1934. Rev. Ent., 4(3): 309. (Tabanus) B. AL(H). Nom. 
nov. pro T. nigriflaz'us 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-Vike. 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 T. flafipcnnis Ric. 1914, from the 
Moluccas. 



NO. 3 NEOTROPICAL FLIES, TABANIDAE — FAIRCHILD I7 

*flaviventris Macq. 1847, Dipt. Exot, Suppl. 3:90. (Tabanus) B. M.(H) = 
Stibasoma, Lutz (1915) with dives Wlk. 1848, as syn. = *T. dives Wlk. 
1848; = St. euglossa Lutz 1915 (fig. only, no description) ; = *St. stilbium 
Fchld. 1953 ; = 5"^ mallophoroides, J. Beq. 1944, not Walker 1857; = ? St. 
siilfurotaeitiatum 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 c? 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. (Tabamis) 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). 
Krober (1930a) as Stypovimisa. Subepaulets setose. 

*fulvoscricea Krob. 1931, Zool. Anz., 95(1-2): 26. (Fidena) B. M.(H) = 
*Scione rufescens (Ric.) 1900; = *Scione aiireopygia Fchld. 1942. (N.S.) 

*fumifera Wlk. 1854, List, 5, Suppl. i : 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. 

*iur Will. 1901, Biol. Centr.-Amer., Dipt., i, Suppl. : 261. (Tabamts) 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 occUata by Enderlein and festiva by Hine. See Fair- 
child (i9Sia) (N.S.). Not listed by Krober (1934). 

♦funinculus Will. 1901, Biol. Centr.-Amer., Dipt., i, Suppl. : 260. (Tabanus) 
B. M.(H). 

*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 intcreuns (Wlk.) 1856 (N.S.). Lutz (1907) noted it was 
probably Neotropical. Krober (1934) not listed. Oldroyd (1954). 

*juscipcnuis Macq. 1847, Dipt. Exot., Suppl. 2: 14. (Dichelacera) B. M.(H) = 
Catachlorops psoloptera (Wied.) 1828. Bequaert (1924) genotype of 
Catachlorops. Krober (1934, I939) ; Barretto (1946). Type agrees with 
specimen of psoloptera det. Barretto. 

*juscipennis Macq. 1838, Dipt. Exot., i(i) : 156. (Diabasis) M. P. =: Leptapha 
fumata (Wied.) 1821 (N.S.). 

♦fuscofasciatus Macq. 1838, Dipt. Exot., 1(1): 140. (Tabanus) M. P.(ri) = 
*Tabanus hilarii Macq. 1839. 

*fuscus Ric. 1902, Ann. Mag. Nat. Hist., ser. 7, 9:431. (Erephrosis) B. M. 
Lutz (1907) and Krober (1933a) as syn. of Fidena imnthemi Wied. Differs 
from imnthemi in B. M. in broader frons, stouter antennae, wholly pollinose 
clypeus, darker legs and white-haired pleura. Close also to obscuripes Krob. 



l8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

but differs in broader palps, frons, and antennae, and in abdominal coloring, 
which is as in isnnthcmi. 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. = 7". lineola var. carneus Bell, 
small form (N.S.). 

*grandis Ric. 1904, Ann. Mag. Nat. Hist., ser. 7, 14:371. (Dichelacera) 
B. M.(H). Krober (i934). 

*giiiterasi 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 ? in Berlin, but the specimen now at B. M. is a $ and 
there is now no c? in B. M. 

*guyanensis Macq. 1845, Dipt. Exot., Suppl. 1:41. (Tabanus) B. M.(H) = 
? Tabanus flaznbarbis Macq., Dipt. Exot., Suppl. i : 41. = T. flavibarbis of 
Krober (1929c, i93oh) and Bequaert (1926). The type of flavibarbis could 
not be found. The description of gnyancnsis precedes on the same page 
(N.S.). 

*haemagogus Will. 1901, Biol. Centr.-Amer., Dipt., i, 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 poUinosity, brown callus and not strongly bicolored fore tibiae. Speci- 
mens in B. M. from Br. Guiana det. luchwsa by Krober are the same, but 
*luctuosa Macq. is a different species. See Barretto (1946). 

*hemiptera Sure. 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 Sure, macula Macq., and minos Schin., and will go in Dasychela. 
I doubt its being Australian. Not listed by Krober (1934). 

*hilarii Macq. 1839, Dipt. Exot., 1(2) : 185. (Tabanus) M. P.(H) = *T. fusco- 
fasciatus Macq. 1838/= ? T. acer Brethes 1910. (N.S.) 

*hinmdus Wlk. 1850, Zoologist, 8, App., p. cxxii. (Dichelacera) B. M.(H). 
= *Dichelacera marginata Macq. 1847, Ricardo (1904). Krober (1934) 
as syn. of marginata Macq. 

♦hirtipalpus Big. 1892, 5:619. (Diatomineura) B. M.(H) = Mycteromyia 
z= *Mycteromyia edzvardsi Krob. 1930. Krober (i93of) 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., i, Suppl. : 254. (Pangonia) 
B. M.(H) = *Escnbcckia jcrruginca (Macq.) Fairchild 1942. Philip 

(1954a). 
♦immaculata Macq. 1838, Dipt. Exot., 1(1): 115. (Dichelacera) M. P.(H). 
= Amphichlorops angustifrons Krob. 1932 = ? Amphichlorops ferruginea 
Barr. 1948. Lutz and Neiva (1914). Krober (i934), 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 juscipcnnis Macq. The type of immaculata 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. (Rhinotriclista) B. M. 
= Scione, Krober (1934). 

*imponens Wlk. 1857, Trans. Ent. Soc. London, 4: 122. (Tabanus) B. M.(H) 
i=z Tabanus olivaceiventris Macq. 1847 (N.S.) = *Atylotus pulverulenius 
Big. 1892. Krober (1929a) as (Lophotabanns) with pulverulentits as syn. 
Bequaert (1926). 

*importiinus 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. 

*incertiis Macq. 1838, Dipt. Exot., 1(1) : 151. (Tabanus) M. P.(H) = Tabanus 
nebulosus de Geer 1776 = *T. ferrifer Wlk. 1850. Not 7". palpalis var. 
inccrtus Szil. 1926, East Indies. Blanchard's (1852) reference of specimens 
from Chile repeated by Krober (1934) is certainly an error. (N.S.) 

♦incipiens Wlk. i860, Trans. Ent. Soc. London, 5:275. (Tabanus) B. M. = 
Stenotabanus, Krober (i93oh, 1934). Type headless, subepaulets bare. Near 
St. macidijrons 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. fiilviceps, Krober 1925, Bequaert 1940. 
= ? C. julviceps Wlk. 1845. =C aurofasciata Krob. 1926. (N.S.) Not 
C. julviceps Lutz 1909. Not C. incisa Fairchild 1942. There are 3 $ cotypes; 
the one bearing Macquart's hand label has been selected and labeled as lecto- 
type. The others are different species. 

Hncisuralis Macq. 1847, Dipt. Exot., Suppl. 2: 12. (Pangonia) B. M. = ? Fi- 
dena opaca (Brethes) 1910 =: Fidena albibarba End. 1925, p. 293, not 
Melpia atiribarba 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. c? $; 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 

*incoitsl^icmis Wlk. 1848, List, 1:171. (Tabaniis) B. M. — Chlorotabanus 
inanis Fab., Krober (1930c). 

*indecisus Big. 1892, 5:666. (Atylotus) B. M.(H) = Tabanus = *Atylotus 
simplex Big. 1892, p. 667. Not *Tabanus simplex Wlk. 1850. Krober (1934) 
as Tabanus. (N.S.) 
infuscatipennis Macq. MS., Sure. 1919, Mes. Arc Merid. fiquat. Amer. du Sud, 
10(2) : 230. (Tabanus) M. P. i $ 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. Krober (1934) does not list. 

*inornatus Wlk. 1848, List, I : 199. (Clirysops) B. M.(H) ^ Diachlorus 
bivittatus (Wied.) 1828, Ricardo (1901). Krober (1934). 

*innotescens Wlk. 1854, List, 5, Suppl. 1327. (Tabanus) B. AL(H) = Cryp- 
totylus pallidipalpis Stone (1944) = Tabanus aurora, Bequaert (1926) and 
Krober (1929c). Not *T. aurora Macq. (N.S.) 

Hnlereuns Wlk. 1856, Dipt. Saund., i, pt. 5:450. (Tabanus) B. M.(H) = 
*Catachlorops fuscinevris (Macq.) 1839. Krober (1934). Barretto (1946). 
(N.S.) 

*interruptus Macq. 1838, Dipt. Exot., i : 156. (Diabasis) M. P. =1 Diachlorus 
immaculatus (Wied.) 1828, Lutz (1913). Krober (1928b). 

*jamaicensis Newst. 1909, 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 Diatomincura. Enderlein (1922, 1925, 
1929) as Listrapha genotype. Krober (i93of) 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. Krober (1934) species incert. scdis. (N.S.) 

*lativitta Wlk. 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 (i93of) as Parosca: (1934) as 
Listrapha. 

*limlnilus Big. 1892, 5:642. (Thcrioplcctes) B. M.(H) =: *Dichelacera uni- 
fasciata Macq. 1838, Brethes (1910) ; Krober (1934); Barretto (1949b). 

*limbithorax ALicq. 1855, Dipt. Exot., Suppl. 5:22. (Pangonia) B. M. = 
Scaptia. Krober (i93of) as Parosca; (1934) as Listrapha. Ferguson (1924) 
concluded on the basis of comparison by Austen with type of limbithorax 
that nivcointtata Ferg. and Henry was a synonym and hence the species 
Australian. Although Krober (i93of, 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., scr. 6, 20:21. (Tabanus mexi- 
canus var.) B. M.(H) = *Cryptotylus longiappendiculatus Macq. 1855. 



NO. 3 NEOTROPICAL FLIES, TABANIDAE FAIRCIIILD 21 

Knab (1916) as syn. of luteoflaims Bell. Not Cryptotyliis limomis 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- 
bantts. 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 d* is a different species, unknown to me. 

*lhndits Wlk. 1848, List, i: 162. (Tabamis) B. M.(H) = Tabanus importunus 
Wied. 1828. Krober (1934) as syn. of T. viduiis Wlk. (N.S.) 

*longiappendiculatus Macq. 1855, Dipt. Exot., Suppl. 5:32. (Tabanus) 
B. M.(H) = Cryptotylus. = Tabanus luteoflavus Bell. 1859. = *T. pnrns 
Wlk. i860. = *T. mexicamis var. limonus Towns. 1897. = T. pallidus Krob. 
1930. ^ T. pallidiilus Krob. 1934. Krober (1934) as (Macrocoriniis) . 
(N.S.) 

*longipalpis Macq. 1848, Dipt. Exot., Suppl. 3:9. (Pangonia) B. M. = 
Histriosilvius Krober i93od genotype; redescribes and figures type. Lutz 
(1909) as Esenbcckia; Ricardo (1900a) as Diatomineura; Enderlein (1925) 
as Protosih'iiis. 

*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. Krober (1933b) as longirostris. = ? Erephopsis brevistria 
Lutz 1909. The type also agrees with specimens in B. M. det. aurifasciata 
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. i : 322, not *basalis Wlk. 1848. 
Ricardo (1900a) says Walker's second basalis 1854 not same as his first. 
Krober'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. = *Tabamis obliqmis 
Wlk. 1850. Not T. luciduliis, 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: (i) 
*T. lucidulus Wlk. 1848 = *T. obliquus Wlk. 1850. = T. lucidulus Krob. 
1930. (2) T. townsendi Johns. =z *T. angustifrons Towns, not Macq. = 
*7". lucidulus Be(juacrt in part 1940. = *T. lucidulus Fchld. 1951. (3) 
T. obumbratus Beq. 1940 == *T. lucidulus 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, Krober (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 luctuosa. His nigripennis, from description and figures, is 
composite, the description agreeing fairly well with luctuosa, 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 (i934)- 

♦lutzi Sure. 1921, Gen. Inseet., Taban. p. 54; 1923, Ann. Ent. Soc. France, 
91(3) : 242. (Stigmatophthalmus) M. P. = Dasychela. Close to *D. riveti 
Sure. Krober (1940) as Dicladocera (Stigmatophthalmus ) . St. altivagus 
Lutz, the genotype, is quite different. Not listed by Krober (i934)- 

*macroceratus Big. 1892, 5:687. (Tabanus) B. M.(H) = *Pseudacanthocera 
sylveirii (Macq.) 1838. Krober (1934)- 

*macrodonta Macq. 1839, Dipt. Exot., 1(2): 183. (Tabaniis) M. P.(H) = 
Psalidia furcata (Wied.) 1828, Lutz (1907)- 

♦macula Macq. 1845, Dipt. Exot., Suppl. i : 43- (Tabanus) B. M.(H) = 
♦Dasychela aiiribarbis Macq. 1847. Krober (1940) as Dicladocera, with 
auribarbis, argyrophorns Schin. 1868 and scutellatus Macq. 1839 as syno- 
nyms. Bequaert and Renjifo (1946) as Dasychela with auribarbis Macq., 
argyrophortis Schin. and submacula Wlk. 1850. The type of scutellatus Macq. 
could not be found, but I doubt its identity with mactda. The type of *sub- 
tnacida Wlk. is somewhat different though closely related. 

*maculifrons Krob. 1931, Stett. Ent. Zeitg., 92:277. (Agelanins {Archiplatius)) 
B. M.(H) = *Dasybasis maculiceps (Krob.) 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 *fidviventris Macq. (q.v.). 

*macnlipcnms 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, Brulle 1832, Macquart 1834, or 
Macquart 1846. 

♦maculiventris Macq. 1850, Dipt. Exot., Suppl. 4:33- (Tabanus) M. P.(H) 
= Dasybasis. Krober (1934) as Tabanus, with rubromaculatus Blanch. 1852 
as synonym. The type is labeled "rubromarginatus Gay Chili," probably a 
lapsus for rubromacidatus, as well as with its published name, so that all three 
names refer to same specimens. 

♦maletectus Big. 1892, 5:664. (Atylotus) B. M. Krober (1934) as Tabanus. 
The subepaulets are bare, labella fleshy, eyes bare, antenrtae 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 (i944)- Lutz 
(1915) as Stibasoma. Krober (1934) as Stibasoma. Dyridophorum has less 
yellow on dorsum of abdomen but otherwise same. It is possible that jestivum 
Wied. and panamcnse Curr. are but races or color forms of the same species. 

*mamjcstus Wlk. 1850, Dipt. Saund., i : 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. E.xot., Suppl. 3:172. (Tabanus) B. M. = 
Tabanus. Not Tabanus margituitus Wlk. 1848 (November) Australia. 
Ricardo's (1901) transference of Walker's species to Silvius makes it also 
a homonym of marginatus Macq. 1838. Krober (i934) confused with Sihnus 
marginatus Macq. 1838. The type is fragmentary and probably unidentifiable 
with certainty. 



NO. 3 NEOTROPICAL FLIES, TABANIDAE — FAIRCHILD 23 

*marg{natus Macq. 1838, Dipt. Exot., 1(1), pi. 19, fig. i. (Silvius) M. P. = 
*Pseudacanthocera slyveirii (Macq.) 1838, i, c.p. 155. Not Silwus 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. 

*marmflrata Big. 1892, 5:634. (Dichclacera) 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 atirea 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 (i93oi, 1934) as {Agelanius). 

*miscra O. S. 1886, Biol. Centr.-Amer., Dipt., i : 47- (Diclisa) B. M.(H) = 
Scione aurulans (Wied.) 1828, Hine (1925). Fairchild (i942d). 

*missionum Macq. 1839, Dipt. Exot., 1(2): 186. (Tabanus) M. P. = Dasy- 
basis. Lutz et al. (1918) as Neotabanus. 

*modesfics Krob. 1931, Stett. Ent. Zeitg., 92(1-2) 1293. (Tabanus (Agelanius)) 
B. M. = Dasybasis modestinus Krob. 1934, nom. nov. Not T. modestus 
Wied. 1828. 

*montium Sure. 1919, Mes. Arc Merid. fiquat. Amer. du Sud, 10:229. (Ta- 
banus) M. P. = Dasybasis. Bequaert and Renjifo (1946) as Agelanius. 
Near osornoi Beq. and excelsus Sure. 

*multijascia Wlk. 1850, Dipt. Saund., 1:68. (Dichelacera) B. M.(H) = Di- 
chelacera cervicomis (Fab.) 1805; Ricardo (1904). 

*nana Wlk. 1850, Dipt. Saund., i : 11. (Pangonia) B. M. = Pseudelaphella. 
Genotype, Krober (i930e). 

*neglectus Will. 1901, Biol. Centr.-Amer., Dipt., i, 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., i (4) : 4-9. (Dasyrhamphis) B. M. 
Specimens so det. by Krober 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. arc 
conspecific and distinct from neo-submacula det. Krober. 

*nigra Krob. 1931, Rev. Ent., I (3): 290. (Rhamphidommia) 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, pi. XCVII, 
fig. 2; 1838, Voy. Coquille, Zool., 11:288. (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. I3I 

venosa (Wied.) 1821 by Krober (i930g, 1934). The palpi are very small and 
deeply grooved. = Fidena. 

*nigrithorax Krob. 1930, Zool. Anz., 90(3-4) : 75. (Diachlonis) B. M. A ? in 
B. M. from Br. Guiana is close but lacks median black on third tergite. It 
stands over the box label podagricns Fab. 

*nigriventris Krob. 1931, Zool. Anz., 94(9-10) : 254. (Escnbeckia) B. M.(H). 

*mgrhnttata Macq. 1850, Dipt. Exot., Suppl. 4:23. (Paiigonia) M. P. = 
Fidena marginalis (Wied.) 1830, Lutz (1907, 1909) ; Krober (1933b, 1934). 
The synonymy is probable but not certain as there are several similar species 
here. 

*nigrohirta Wlk. 1848, List, 1 : 132. (Pangonia) B. M. = Fidena venosa 
(Wied.). Krober (i930g). 

*nitens Big. 1892, 5:609. (Myctcromyia) B. M. (H) := Fidena. Krober 
(1933b) redescribes, but his description and figures inconsistent and probably 
composite. Fairchild (1951a) genotype of Bornbylopsis Lutz and lonopsis 
Lutz. 

*nivalis Wlk. 1850, Dipt. Saund., 1:71. (Scepsis) B. M.(H) genotype. Sur- 
couf (1921); Krober (1928a). 

*notabilis Wlk. 1850, Dipt. Saund., i : 18. (Pangonia) B. M. = Esenbeckia, 
Krober (1932a) with injramaciilata Lutz as synonym. If notabilis and iu- 
jramaculata 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). 

*nitiitius Wlk. 1854, List, 5, Suppl. 1:207. (Tahaiius) B. M.(H) = *Tabanus 
desertus Wlk. 1850. (N.S.) Krober (1934) says type lost. Philip (1952) 
as synonym of obsolctus 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. (Tabmius) B. M. = *Tabanus lu- 
cidulus Wlk. 1848. 

*obscurehiria Krob. 1930, Zool. Anz., 86 (ii-i2):284. (Tabanus (Phacota- 
bamis) aphanoptera var.) B. M. Krober (1934) as var. obscuripilis, nom. nov. 
Not T. obscnrchirta Ric. 1908. = Phaeotabanus obscuripilis Krob. 

*obscitriventris Krob. 1930, Zool. Anz., 87(1-2): 6. (Tahauus (Matroconnus)) 
B. M. =: *Tabanus (Lophotabanus) albocirculus Hine 1907. Krober (1934) 
as obscurigaster, nom. nov. Not T. obscuriventris Krober 1929. (N.S.) 

♦ocellus Wlk. 1848, List, i : 143. (Pangonia) B. M. = Dasychela. Ricardo 
(1900a) noted it was a tabanine. Krober (i93oh) 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. ochraccns Krober 1928 
which = ? *D. augitslifrons 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.) 

*ocul(ittts P.ig. 1892, 5:606. (Chrysops) B. M. = Chrysops molesta Wied. 1828, 
Ricardo 1901 ; Krober (1926, 1934). 



NO. 3 NEOTROPICAL FLIES, TABANIDAE — FAIRCHILD 25 

*oculus Wlk. 1848, List, 1:157. (Tahamis) B. M.(H) = Tabanus (Lopho- 
tabanus). Krober (1934) as Bcllardia. 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. Krober (1929, i.e.) lists as synonym of Lopho- 

tabanus druyvcsteijni Szil., a synonym of nebulosus de Geer. Fairchild 

(i942f) as synonym of ferrifer Wlk. The species is, in my opinion, a darker 

and smaller race of nebulosus. 

*pachycephahim Big. 1892, 5 : 636. (Stibasoma) B. M. = *Stibasoma chio- 
nostigma O. S. 1886. Fairchild (1940b). 

♦pachjrpalpus Big. 1892, 5:631. {Dichelacera) B. M.(H) =. Tabanus. Near 
bigoti Bell, and validus Hine. Same as rtifipcnnis Macq. 1846 (Dipt. Exot., 
Suppl. 1 141), specimen in B. M., but different from *rufipcnnis Macq. 1838, 
types in Paris. 

*pallidefemorata Krob. 1930, Zool. Anz., 90:72. (Chrysops auroguttafa van) 
B. M. = Chrysops pallidefemorata, Bequaert (1944) ; Pechuman (1937). 

*palHdctinctus Krob. 1930, Zool. Anz., 86(11-12) : 297. (Tabanus (Phaeota- 
bantis)) B. M.(H) = Tabanus (Philipotabanus) caliginosus Bell., Fair- 
child (1953). 

*parallelus Wlk. 1848, List, 1:187. (Tabanus) B. M. = Stenotabanus, 
Bequaert (1940) with *T. alene Towns, as synonym. 

*patellicornis Krob. 1930, Zool. Anz., 88(11-12) : 307. (Pseudelaphella) 
B. M.(H). 

*pavida Will. 1901, Biol. Centr.-Amer., Dipt., i, Suppl. :253. (Pangnnia) 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 Dicladoccra. The subepaulets are setose and the species belongs 
with hirtitibia Wlk. in the group treated as Chclommia 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 Chelonimia. Barretto (1949a) 
as Chelonimia. Bequaert and Renjifo (1946) as Dichelacera (Psalidia). 

*piceo-hirta Wlk. 1848, List, i : 132. (Pangonia) B. M. =: Fidena venosa Wied. 

*pictipennis Macq. 1834, Hist. Nat., Dipt., 1:199. (Tabanus). 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 
Kruber (1934). 

*pictipeniiis 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. 193 1. 



26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

♦pictus Krob. 1930, Zool. Anz., 86(11-12) : 264. (Eutabanus) B. M. Subepaulets 

bare. Nearest Stenotabamis but distinct. Bequaert (1939). 
♦plangens Wlk. 1854, List, 5, Suppl. i: 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., i : 69. (Dichelacera) B. M. = Cata- 

chlorops. Closest to haltcratus 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- 

tatiis Latr. Krober (1930c) as Macrocormiis, redescribes. Near strigimacu- 

his Fchld. 1942. 
*pntinosus Krob. 1931, Stett. Ent. Zeitg., 92(2) : 276. {Tabanus (Agclanius)) 

B. M.(H) = Dasybasis. Krober (1934) as pruinivitta, nom. nov. Not 

T. pruinostis Big. 1892, or Hine 1900, or Surcouf 1906, or Krober 1929. 
*ptibescens Macq. 1847, Dipt. Exot., Suppl. 2:20. (Tabanus) B. M. The type 

is in very bad condition, probably composite and not certainly Neotropical. 

Not T. pubescens Strom 1768 or Walker 1854. 
*pulchra Will. 1901, Biol. Centr.-Amer., Dipt., i, Suppl. : 263. (Dichelacera) 

B. M.(H) = Dichelacera salvadorcnsis Lutz 1915- (N.S.) 
*pulverulentiis Big. 1892, 5:665. (Atylotus) B. M.(H) = Tabanus olivacei- 

ventris Macq. 1847. (N.S.) = *Tabanus imponens Wlk., Krober (1929a). 
*pumiloides Will. 1901, Biol. Centr.-Amer., Dipt., i, Suppl. : 260. (Tabanus) 

B. M. = Stenotabanus. Fairchild (1953). 
*punctipennis Macq. 1838, Dipt. Exot., 1(2): 185. (Tabanus) M. P.(H) = 

Tabanus (Hybomitra) quadripunctatus Fab. Fairchild (i942f). Not 

T. punctipcnnis Macq. 1847 (Nearctic), or Stypommisa punctipennis End. 

1925- 
*piirus Wlk. i860, Trans. Ent. Soc. London, 5:274. (Tabanus) B. M.(H) = 

♦Cryptotylus longiappendiculatus Macq. 1855. Hine (1925) as synonym of 

luteoflaims Bell. (N.S.) 
*pyrausta O. S. 1886, Biol. Centr.-Amer., Dipt, i : 43- (Pangonia) B. M.(H) 

= Fidena rhinophora Bell. 1859, Fairchild (1953). 
*quadrimaculatus Macq. 1845, Dipt. Exot., Suppl. 1:39. (Tabanus) B. M. = 

Dichelacera. Krober (1932a) as Gymnochela; says poecilopfera Schiner 

same; (1934) as Amphichlorops. Close to *tcstacca Macq., and *alcis Will. 
*reinburgi Sure. 1919, Mes. .A.rc Merid. £quat. Amer. du Sud, 10:233. (Esen- 

beckia) M. P. 
*rcpanda Wlk. 1848, List, 1:190. (Dichelacera) B. M.(H) = ♦Dichelacera 

testacea Macq. 1846, Krober (1934). 
*reticulatus Krob. 1930, Zool. Anz., 86(11-12) : 298. (Tabanus (Phacota- 

banus)) B. M.(H) = (Philipotabanus), Fairchild (i942f). 
*riveti Sure. 1919, Mes. Arc. Merid. Iiquat. Amer. du Sud, 10:226. (Tabanus) 

M. P. = Dasychela. Krober (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 *lrigonophorus Macq. but probably now indeterminable. 
*rubescc}is Big. 1892, 5:663. (Atylotus) B. M. = Tabanus campestris Brethes 

191 1, nom. nov. Krober (1933a) redescribes. Subepaulets setose. Not rubes- 

cens Macq. 1838, Bellardi 1859. 



NO. 3 NEOTROPICAL FLIES, TABANIDAE — FAIRCHILD 27 

*rnbidiis Macq. 1847, Dipt. Exot., Suppl. 2: 19. (Tabanus) B. M. Not T. rti- 
bidiis Wied. 1821. Krober (1934) as T. rubricosa, nom. nov. Not T. rubri- 
cosa Wulp. 1881. Type headless and excessively dirty, probably now inde- 
terminable. Near *fallax Macq. and albibarbis Wied. 

*rubiginipennis Alacq. 1845, Dipt. Exot., Suppl. 1:39. (Tabanus) B. M.(H) 
1= *Tabanus adustus Wlk. 1850. Krober (1940) as Didadocera. 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 fidminca 
Hine, but with bicolored fore tibiae and widely open cell R5. 

*rubribarbis Big. 1892, 5:630. (Atylotiis) B. M.(H) = Dichelacera (Psa- 
lidia) furcata (Wied.) 1828, Krober (1932b). 

*rubrinotatiis Big. 1892, 5:676. (Atylotus) B. M.(H) = *Tabanus guyanen- 
sis Macq. 1845. Krober (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 stibmaryinata Lutz 1915. Rondani (1850) transfers to Tabanus and 
changes name to brasiliensis, nom. nov. Lutz (1907) := ? D. januarii var. 
Krober (1934) ■= januarii Wied. A pale form with reduced wing markings 
and broad frons. 

*nifescens Ric. 1900, Ann. Mag. Nat. Hist., ser. 7, 6:294. (Erephrosis) 
B. M.(H). The holotype is in Budapest; this is a paralype. = *Scione 
aureopygia Fchld. 1942. = *Fidcna 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. = Didado- 
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. 

*rufivcntris Macq. 1838, Dipt. Exot., 1(1) : 145. (Tabanus) B. M. = *Tabanus 
bifloccus Hine 1925. Not T. rufiventris Fab. 1805 or Macquart 1845. Krober 
(1934) does not list. Bequaert (1940) = ? T. hookeri Knab. 



28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 

*ntfivcnlris 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) indioriim Beq. and Renjifo 1946, nom. nov. pro rufi- 
ventris Macq. 1845. (N.S.) 

*rnfohirta Wlk. 1848, List, 1:131. (Pangonia) B. M.(H) = Fidena venosa 
(Wied.), Krober (i930g, 1934). 

*rufopilosus Big. 1892, 5:620. (Veprius) B. M. Ricardo (1901) as Silvins. 
Krober (i93Gd, 1934) lists under Vepriiis with a query. Hind tibial spurs, 
bare subepaulet and subcosta, fleshy labella, and pilose holoptic eyes. 

*nifus Krob. 1931, Stett. Ent. Zeitg., 92:287. (Archiplaliits) B. M.(H) == 
*Tabanus desertus Wlk. Not T. rufits Scop. 1763 or Palisot de Beauvois 
1809. Krober (1934) as T. (Agelanius) ruficolor, nom. nov. (N.S.) 

*satanica Big. 1892, 5:632. (Dichelacera) B. M. =1 Dicladocera. Krober 
(1931c) as Gymnochela with castanea Big. as syn. ; (1934) as Chelommia. 
The subepaulets are bare and the species congeneric with unicolor Lutz and 
*rufipennis Macq. It also agrees well with the description of T. 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. E.xot., Suppl. 2:15. {Dichelacera) B. M.(H). 
Hine (1917). Krober (1934). Nearest to *nmrginata 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 riifescens Fab. 

*scutulatU3 Krob. 1930, Dipt. Pat. S. Chile, 5(2) : 143. (Tlierioplectes) 
B. M.(H) = Dasybasis. = *Tabanus (Therioplectes) albovittatns 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 Krober (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 scminigra Ric. Close to 
*fencstrata Macq. and *Iongipentiis Ric. 

*scmisordidus Wlk. 1854, List, 5:208. {Tabanus) B. M.(H) = Tabanus 
importunus Wied., Krober (1929c, 1934). 

*semiviridis Ric. 1900, Ann. Mag. Nat. Hist., ser. 7, 8: 181. {Pangonia) 
B. M.(H) = Esenbeckia prasiniventris Macq., Bequaert and Renjifo 
(1946). Krober (1932a, 1934). Described as from Barengo, Old Castile, 
Spain, but the original label is indecipherable and might have been "Vene- 
zuela." 

*senior Wlk. 1850, Dipt. Saund., i : 67. {Tabanus) B. M.(H) = Tabanus albi- 
barbis Wied. 1824. (N.S.) Bodkin and Cleare (1916). Krober (1934) not 
listed. 

*shannoni Krub. 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 uinbraticolus, which is entirely distinct. 



NO. 3 NEOTROPICAL FLIES, TABANIDAE — FAIRCHILD 29 

^simplex Big. 1892, 5:667. (Tabamis) B. M.(H) = *Tabanus indecisus 
(Big.) 1892. Not *'T. simplex W\k. (N.S.) Krober (1934) = ? T. {Neo- 
tabaiiHs) signativentris Brethes. There are three cotypes, the lectotype being 
the one with the Bigot name label. The other two are different species. 

*sparsa Wlk. 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., 92:299. (Tabauus) B. M.(H)=: 
*Tabanus (Philipotabanus) grassator Fchld. 1953. Type headless. My 
specimen becomes allotype. (N.S.) 

*subfascipc)uiis 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. (Tabaims) B. M.(H) =z 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. (Tabanns) B. M. = Esenbeckia, Krober 
(1932a, 1934). Walker (1849) = Pangonia fiiscipennis Wied. var. Ricardo 
(1900) := Pangonia, a distinct species. Close to *Esenbeckia notabilis Wlk., 
the proboscis rather heavy, incompletely sclerotized. 

*sulphurexis Macq. 1847, Dipt. Exot., Suppl. 2:19. (Tabanns) B. M.(H) = 
Chlorotabanus inanis (Fab.), Krober (1934). Lutz (1907) as a pale form 
of mexicanus L. 

*tabanipcnnis Macq. 1848, Dipt. Exot., i(i):io8. {Pangonia) M. P. The 
lectotype = Fidena castanea Party 1833 of Krober (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 Wlk. 1850, Newman's Zoologist, 8, App., p. Ixv. (Tabanns) B. M.(H) 
= Tabanns cinerarins Wied. 1828. Not T. tenens Wlk. 1850, Dipt. Saund., 
1:49. Walker (1854) changes name to confligens. Krober (1934) as 
Chelommia. T. cinerarins was proposed as a new name for T. glancus Wied. 

1819 (Zool. Alag., 1(3) :42) thought to be preoccupied by T. glancus 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. 1819. 

*tenuirostris Wlk. i860. Trans. Ent. Soc. London, 5:272. (Pangonia) B. M. = 
Esenbeckia flavohirta Bell. 1859. Krober (1934) as Ricardoa flavohirta. 

*tenuistria Wlk. 1848, List, 1:143. (Pangonia) B. M. = Fidena. Krober 
(ig30g) 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.) 

*terminns Wlk. 1848, List, i: 160. (Tabanns) B. M. = ? Tabanus sorbillans 
Wied. Type is a d* and seems to match sorbillans fairly well. Krober (1933a) 
as a valid species of (Neotabanns). (N.S.) 



30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

♦testacea Macq. 1846, Dipt. Exot., Suppl. 1:29. (Dichelacera) B. M.(H). 
Krdber (1934) with repanda as queried synonym = *Dichelacera repanda 
Wlk. 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, Krciber (1932a) with umbra Wlk. 1850 as synonym = 
*E. umbra Wlk. 

♦testaceomaculatus Macq. 1838, Dipt. Exot., 1(1) : 144. (Tabanus) M. P.(H) 
= Dasybasis, Krober (i93oi) as (Agelanius). = ? *D. trigonophorus 
Macq. 1838, p. 185. 

*testaceus Macq. 1838, Dipt. Exot., 1(1): 137. (Tabamis) M. P. Closest to 
*T. indecisiis Big. but differs. Krober (1930c) claims he saw type in London. 
Not T. testaccus 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 T. 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, Krober (i93ii, 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 c? Stenotabanus, but not further identifiable in present 
state of knowledge of this group. It is not the c? of *callosus Macq. as sug- 
gested by Krober (1934). 

♦trigonophorus Macq. 1838, Dipt. Exot., 1(2) : 185. (Tabamis) M. P. = Dasy- 
basis. Very close to *testaceomaculatus Macq. 1838, p. 144, structurally, but 
both types very denuded and certainty impossible. 

*tristi3 Big. 1892, 5 : 621. (Dasybasis) B. M. = ? Protodasyapha, Krober 
(1930k). A (S. Eyes densely pubescent, holoptic, facets demarcated. Ocelli 
present. Subcpaulet bare, hind tibiae spurred, labella fleshy, antennae subu- 
late, Pangonia-Wkt. 

*tritus Wlk. 1857, Trans. Linn. Soc. London, 17(3) : 338. (Tabanus) B. AL = 
Dasybasis. Krober (i93oi, 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., i: 19. (Pangonia) B. M.(H) = *Esenbeckia 
testaceiventris Macq. 1848, Krober (1934). 

*unicinctus Wlk. 1857, Trans. Ent. Soc. London, 4: 122. (Tabamis) B. M. = 
♦Leucotabanus albovarius (Wlk.) 1854. (N.S.) = *Lcucotabanus leu- 
conotum Fclild. 1941. Not T. utiicinctus 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 *aIbovarius 
Wlk. 

*unicolor Macq. 1845, Dipt. Exot, Suppl. 1:27. (Pangonia) B. M.(H) = 
Fidena castanea Perty 1830, Krober (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 3I 

*unifasciata Macq. 1838, Dipt. Exot., 1(1): 119. (Dichelacera) M. P.(H). 

Barretto (1949b) with *limbata Big., trigonotaenia Lutz and soror Krob. as 

syns. 
*unipunctatus Big. 1892, 5:663. (Atylotus) B. M.(H) = *Tabanus (Lopho- 

tabanus) piraticus Fchld. 1942. (N.S.) Philip (1952) = T. jilamensis 

Hine. Not T. (L.) unipitnctatus, Krob. 1929. Jilamensis Hine appears to be 

distinct. See Fairchild 1942a. 
*uniznttatus Macq. 1855, Dipt. Exot., Suppl. 5:30. (Tabanus) B. M.(H) = 

♦Tabanus desertus Wlk. 1850. (N.S.) 
*vaciUans Wlk. 1850, Dipt. Saund., 1:70. (^Dichelacera) B. M.(H) = ? Cata- 

chlorops capreolus Wied. 1828. (N.S.) Krober (1934) as synonym of potator 

Wied, 1828. Not vacillans, Barr. 1946. The type keys out to capreolus in 

Barretto's key (1946). Lutz (1907) = /'Otofor 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 ? Diatomincura. Krober {i()2,o'k) =^ Calliosca schoenemanii End. 

genotype. 
*rariprs Wlk. 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, Krober (1934) = Amphiclorops. Fairchild (1942b) = 

Stibasoma. Very close to *Rhab. viridiventris Macq. 
♦venosus Big. 1892, 5:685. (Tabanus) B. M.(H) = *Stenotabanus maculi- 

pennis (Krob.) 1929. (N.S.) Krober (1930a) as Stypommia, but his 

venosus not same species as the type. St. pequeniensis Fchld. 1942 also close. 
*viduus Wlk. 1850, Newman's Zoologist, 8, App., p. Ixviii. (Tabanus) B. M.(H) 

= *Tabanus basivitta Wlk. 1850. (N.S.) Krober (i93oh) = r. (Lopho- 

tabanus) lividus Wlk. 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 Lisirapha. 
*viridiventris Macq. 1838, Dipt. Exot., 1(1): 141. (Tabanus) M. P.(H) = 

Rhabdotylus. Close to venenatus O. S. and planiventris Wied. Carrera and 

Lane (i945). 

SUMMARY 

A study of the type specimens of Neotropical Tabanidae preserved 
in the British Museum of Natural History in London and the Museum 
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. I3I 

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 i8i 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: no 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 ty[)e. 
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. 

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NO. 3 NEOTROPICAL FLIES, TABANIDAE — FAIRCHILD 33 

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34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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NO. 3 NEOTROPICAL FLIES, TABANIDAE FAIRCHILD 35 

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Zool. Anz., vol. 83, Nos. 1-4, pp. 47-63; Nos. 5-8, pp. ns-i37- 
1929b. Die Ausbeute der deutschen Chaco — Expedition 1925 — 26. XIII. 

Tabanidae. Konowia, vol. 8, No. 2, pp. 174-193. 2 pis. 
1929c. 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. 
I929d. Die Stenotabaninae und die Lepiselaginae Sudamerikas. Encycl. 

Entom., ser. B, 11, Diptera, vol. 5, No. 3, pp. 101-154, 97 figs. 
1930a. Nachtrage zu den kleinen Gattungen der sudamerikanischen Ta- 
banini. Zool. Anz., vol. 86, Nos. 9-10, pp. 248-265, 8 figs. 
1930b. Die Tabanidenuntergattung Phaeotabanus Lutz. Zool. Anz., vol. 86, 

Nos. 11-12, pp. 273-300. 
1930C. Die Untergattungen Macrocornns Lutz und Chlorotabanus Lutz. 

Zool. Anz., vol. 87, Nos. 1-2, pp. 1-18, 13 figs. 
i93od. Die Tabaniden subfamilie Silviinae der neotropischen Region. Zool. 

Anz., vol. 88, Nos. 9-10, pp. 225-239, 11 figs. 
19306. Die Pityocerini (Tabanidae) der neotropischen Region. Zool. Anz., 

vol. 88, Nos. 11-12, pp. 305-312. 
l93of. Die Tribus Pangoniini der neotropischen Region. Zool. Anz., vol. 89, 

Nos. 7-10, pp. 211-228. 
I930g. Die Tabanidengattung Sackenimyia Big. Zool. Anz., vol. 90, Nos. 

1-2, pp. 1-12, 6 figs. 
i93oh. Neue Tabaniden und Zusatze zu bereits beschreibenen. Zool. Anz., 

vol. 90, Nos. 3-4, pp. 69-86, 21 figs. 
i93oi. Tabanidae. Diptera of Patagonia and South Chile, pt. 5, fasc. 2, 

pp. 106-161, 2 pis. British Museum. 
1930J. Die sudamerikanischen Arten der Gattung Scionc Wlk. (=: Rhino- 

triclista End.). Stett. Ent. Zeitg., vol. 91, No. 2, pp. 140-174. 
1930k. Die Pelecorhynchinae und Melpiinae Sudamerikas. Mitt. Zool. 

Staatinst. Zool. Mus. Hamburg, vol. 44, pp. 149-196, 33 figs. 
1931a. Neue sud- und mittelamerikanische Arten der Dipterengattung 

Tabanus L. Stett. Ent. Zeitg., vol. 92, No. 2, pp. 275-305. 
1931b. Neue neotropische Tabaniden aus den Unterfamilien Bellardiinae 

und Tabaninae. Rev. Ent., vol. i. No. 4, pp. 400-417, 18 figs. 
1931C. Die Tabanus — Untergattung Gymnochela End. Zool. Anz., vol. 96, 

Nos. 1-2, pp. 49-61, 9 figs. 
I93id. Neue Arten der Gattung Fidena Wlk. Zool. Anz., vol. 95, Nos. 1-2, 

pp. 17-37, 19 figs. 
193x6. Neue Arten aus dem Genus Esenbcckia Rond. Zool. Anz., vol. 94, 

Nos. 9-10, pp. 245-257, 7 figs. 
I93if. Die Tabanus — Gruppen Straba End. und Poecilosoma Lutz ( = 

Hybostraba und Hybopelma End.) der neotropischen Region. Zool. 

Anz., vol. 94, Nos. 3-4, pp. 67-89, 20 figs. 
I93ig. Die kleinen Gattungen der Dichelacerinae End. aus der sud- 
amerikanischen Region. Rev. Ent., vol. i. No. 3, pp. 282-298, 

II figs. 



36 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



I93lh. 



19311. 



1932a. 



1932b. 



1933a. 



1933b. 



1934- 



1939- 

1940. 

LuTZ, A. 
1907. 



Dreizehn neue neotropische Tabanusarten. Konowia, vol. 10, No. 4, 

pp. 291-300. 
Neue sudamerikanische Tabaniden des ungarischen National-Mu- 
seums und einiger aiiderer Institute. Ann. Mus. Nat. Hungarici, 

vol. 27, pp. 329-350, 19 figs. 
Das Genus Esenbeckia Rondani und die Gymnochela — Untergattung 

Amphichlorops Lutz. Rev. Ent., vol. 2, No. i, pp. 52-90, ^2 figs. 
Die Tabaniden — Subfamilie Bellardiinae End. der neotropischen 

Region. Rev. Ent., vol. 2, No. 2, pp. 289-302, 6 figs. 
Das Subgenus Neotabanus der Tabanidengattung Tabanus s. lat. 

Rev. Ent., vol. 3, No. 3, pp. 32i7-i(>7- 
Die neotropischen Arten der Tabanidengattung Fideiia Wlk. Arch. 

Naturg., N. F., vol. 2, No. 2, pp. 231-284. 
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- 

Das Tabanidengenus Catachlorops Lutz. Verhoflf. Deutsch. Kol. Mus. 

Bremen, vol. 2, pp. 211-232, 4 pis. 
Das Tabanidengenus Dicladocera Lutz. Verhoff. Deutsch. Kol. Mus. 

Bremen, vol. 3, No. i, pp. 58-92, 3 pis. 



1909. 



1913- 



1915- 



Lutz, A 
1918 



Bemerkungcn iiber die Nomenclatur und Bestimmung der Brasilian- 

ischen Tabaniden. Zentralbl. Bakteriol., vol. 44, No. i, pp. 137-144. 

Tabaniden Brasiliens und einiger Nachbarstaaten. Zool. Jahrb., 

Suppl., vol. 10, No. 4, pp. 619-692, 3 pis. 
Tabanidas do Brasil e alguns estados visinhos. Mem. Inst. Oswaldo 

Cruz, vol. 5, No. 2, pp. 142-191, 2 pis. 
Tabanidas do Brasil e de alguns estados visinhos. Segunda mcmoria. 
Mem. Inst. Oswaldo Cruz, vol. 7, No. i, pp. 51-119, 3 pls. 
Lutz, A., and Neiva, A. 

1914. As Tabanidae do Estado do Rio de Janeiro. Mem. Inst. Oswaldo 
Cruz, vol. 6, No. 2, pp. 69-80. 
Araujo, H. C. de Souza, and Fonseca Filho, O. da. 
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. de Oliveira. 

1935. Sobre algumas novas especies de motucas do genero Esenbeckia Rond. 
Mem. Inst. Oswaldo Cruz, vol. 30, No. 3, pp. 543-562. 
Mackerras, I. M. 

1954. The classification and distribution of Tabanidae I. General review. 
Australian Journ. Zool., vol. 2, No. 3, pp. 431-454, 10 figs. 
Macquart, J. 

1834. Histoire naturelle des insectes dipteres. In Suites a Buffon, vol. I, 

pp. 186-216. 
1838-1855. Dipteres exotiques nouveaux ou peu connus. Mem. Soc. Roy. 
Sci., Agr., Arts, Lille. Contemporaneously published as a separate 
work by Librairie Encylopedique 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, pis. I-XXV (1838, vol. I, pt. I, pp. 1-221, pis. I- 
XXV) ; 1839, pp. 121-323, pis. I-XIV (1838, vol. I, pt. 2, pp. 1-207, 
pis. I-XIV); 1841, pp. 283-413, pis. I-XXI (1840, vol. 2, pt. I, 
pp. 1-135, pis. I-XXI) ; 1842, pp. 63-200, pis. I-XXII (1842, vol. 2, 
pt. 2, pp. 1-140, pis. I-XXII) ; 1843, pp. 162-460, pis. I-XXXVI 
(1843, vol. 2, pt. 3, pp. 1-304, pis. I-XXXVI) ; 1845, pp. 133-364, 
pis. I-XX (1846, Suppl., pp. 1-238, pis. I-XX) ; 1847, pp. 21-210, 
pis. I-VI (1847, 2" Suppl., pp. 1-104. pis. I-VI) ; 1848, pp. 161-237, 
pis. I-VII (1848, Suite de 2' Suppl. [or third Supplement], pp. 
1-77, pis. I-VII) ; 1850, pp. 309-379, pis. I-XIV, and 1851, pp. 134- 
209, pis. XV-XXVIII (185 1, 4'' Suppl., pp. 1-336, pis. I-XXVII) ; 
1855, pp. 25-156, pis. I-VII (1855, s" Suppl, pp. 1-136, pis. I-VII). 
Oldroyd, 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 Misc. Coll. vol. 16, No. 270, xlvi -|- 276 pp. 

1886. Biologia Centrali-Americana, Insecta, Diptera, i (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. 
Philip, 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. Midi. 
Nat., vol. 27, No. 2, pp. 257-324. 

1950. New North American Tabanidae II. Ann. Ent. Soc. Amer., vol. 43, 
No. I, pp. 1 15-122. 

1952. The Linnaean and Degeerian species of American Tabanidae. Ann. 
Ent. Soc. Amer., vol. 45, No. 2, pp. 310-314. 

19543. 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. 

RiCARDO, 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, I pi. 
1900b. Description of five new species of Pangoninae from South America. 

Ann. Mag. Nat. Hist., ser. 7, vol. 6, pp. 291-294. 
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 

RONDANI, C. 

1850. Osservazioni sopra alquante specie di esapodi ditteri del Museo Tori- 

nense. Nuov. 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 urn die Erde, etc. Zoologische 
Teil, Diptcra, pp. i-vi, 1-388, 4 pis. 
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, N0.2, 
pp. 248-254. 
SURCOUF, J. M. R. 

1919. Dipteres. Brachyceres piqueurs (Tabanidae). Mes. Arc de Merid- 
ian Equatorial en Amerique du Sud, vol. 10, No. 2, pp. 217-233. 
1921. Genera insectorum, fasc. 175, Diptera, Tabanidae, pp. 1-182, 5 pis. 
Brussels. 
SZIL.\DY, Z. 

1926. New and Old World horseflies. Biol. Hungarica, vol. i, 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. Ft. i, pp. 1-299 (1848); pt. 5, pp. 1-330, i pi. 

(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. i, pp. 1-75. 2 pis. 
1850b. Characters of undescribed Diptera in the British Museum. New- 
man's Zoologist, vol. 8, Appendix, art. i, pp. Ixv-lxxii, art. 5, 

pp. xcv-xcix, art. 25, pp. cxxi-cxxii. 
Wiedemann, C. R. W. 
1821. Diptera exotica. 
1828-1830. Ausscreuropaische Zweiflugelige Insekten, vol. i, pp. i-xxxii, 

1-608, 7 pis. (1828) ; vol. 2, pp. i-xii, 1-684, 5 P's. (1830). 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 4 



Cljarlesi S. anb jWarp "^aux OTalcott 
jResiearcl) Jf unb 



NEW CRETACEOUS BRACHIOPODA 
FROM ARIZONA 

(With 4 Plates) 



By 
G. ARTHUR COOPER 

United States National Museum 
Smithsonian Institution 






^Ml^ 



(PUBUCATION 4227) 



CITY OF WASHINGTON 

PUBLISHED BY THE SMITHSONIAN INSTITUTION 

DECEMBER 21, 1955 



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



CfjarlefiJ B. anb JHarp ^aux Malcott 3^tieavtf) jFunb 

NEW CRETACEOUS BRACHIOPODA 
FROM ARIZONA 

By G. ARTHUR COOPER 
U. S. National Museum 
Smithsonian Institution 

(With Four Plates) 

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 
NW:|:SW^NE:^ 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. I3I 

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, TerebrateUa 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 miiralensis, 
and large forms of Lunatia? sp. often occur. The massive limestone 
is usually replete with Radiolites? 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 3 A, figures 1-3 

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, 

34. 

Types.— UoXoiy^e U.S.N.M. No. 124192. 

Horizon and locality. — Mural limestone, from a small hill 300 yards 
east of U. S. Highway 80, NW^SWiNEi 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 lA, figures 1-17 

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 

Holotype U.S.N.M. No. 124193a 14.0 

Paratype U.S.N.M. No. 124193b 1 1.8 



Width 


Thickness 


13.6 


9.0 


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. I24i86a-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, NWiSW^NEi 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 
rhynchonellids 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. amcricana. 

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 iB, 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 9.4 

Paratype U.S.N.M. No. 124194a 16.9 13.9 12.5 8.8 

" 124194c 20.3 17.4 17.4 11.6 

" I24i94d 54 4-7 3-9 2.7 

" " " I24i94e 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, 124194c, d, 124195, 124195a, 124196c; 
unfigured paratypes U.S.N.M. Nos. 124194a, e, 124196a, b, 124205, 
I 242 I 8. 

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. vcspertina and 
according to Sahni's figures (1929, pi, 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 iB, 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. 124195a. 
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 lateral 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. 124200a) 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. 124201a) 
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 



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. 


. lO.I 


8.8 


12.3 


8.7 


5.2 


Paratype U.S.N.M. 


No. 


124197b. 


. 131 


ii.i 


13.5 


10.4 


8.7 


" " 




124206a. 


. 12.2 


9.6 


14.0 


10.8 


6.5 


« « 




124206b. 


• 13.4 


10.6 


14.0 


10.7 


1-^ 


« « 




124206c. 


. 11.5 


9.0 


130 


10.9 


6.4 


<< <i 




i242o6d. 


• 7-2 


5.3 


7-5 


? 


4.2 


« (( 




i242o6e. 


. S-i 


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, 124201a, b, 124220; 
measured paratypes U.S.N.M, Nos. 124206 a-e; unfigured paratypes 
U.S.N.M. Nos. 124197c, 124200a, b. 

Horizon mid locality. — Mural limestone, from a small hill 300 yards 
east of U. S. Highway 80, NW^SWiNEi sec. 36, T. 23 S., 
R. 25 E., Bisbee Quadrangle, Arizona ; Rancho Nuevo, 3 miles east of 
Santa Rosalia, Sonora, Mexico, 

Discussion. — This species is characterized by having a low and in- 
distinct fold and sulcus and differs from all other described species of 
Genimarcula in this respect. This is the first report of the genus in 
North America. Gemvwrcula arisonensis 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. Genimarcula mcnardi 
(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. 



10 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 
to 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 OrnithcUa 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 i, 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 



12 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. I24202d). 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 (pi. 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. I24202g) 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. i24i9oh). 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 I3 

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 f ev^ 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 42 mm. the expanded end of the septum splits laterally and the 
ring enlarges, the septum having a deep reentrant 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 '].(i 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 



Length 

Holotype U.S.N. M. No. 124191 15. i 

Paratype U.S.N.M. No. 124189 13.3 

" " " 124190a 11.4 

" " " 124190b 12.4 



Types. — Holotype U.S.N.M. No. 124191 ; figured paratypes 
U.S.N.M. Nos. 124190a, c, e, f, h-j, 124202a, d-i ; measured paratypes 
U.S.N.M. Nos. 124189, 124190a, b; described but unfigured paratypes 
U.S.N.M. No. 124190k; unfigured paratypes U.S.N.M. Nos. I24i90d, 
g, 124202b. 

Horizon and locality. — Mural limestone, from a small hill 300 yards 
east of U. S. Highway 80, NWiSW^NEi sec. 36, T. 23 S., 



brachial 
length 


Width 


Thickness 


12.4 


13-9 


9-7 


II.O 


12.2 


8.1 


9.6 


10.7 


6.0 


10.4 


11.7 


7-5 



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 "Waldheiniia" 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 
Bosquet, J. 

1859. Monographic des brachiopodes fossiles du Terrain Cretace superieur 
du Duche de Limbourg. Haarlem. 
Cobban, W. A., and Reeside, 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. 
1 01 1 -1044. 
Davidson, 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 i, Sup- 
plement to the Recent, Tertiary and Cretaceous species, pp. 17-72. 
Deslongchamps, E. E. 

18S4. Etudes critiques sur des brachiopodes nouveaux ou peu connus. Art. 8. 
Note sur les modifications a apporter a la classification des 
Terebratulidae. 
Elliott, G. F. 

1947. The development of a British aptian brachiopod. Proc. Geol. Assoc, 
vol. 58, pt. 2, pp. 144-159- 
MuiR-Wooi). 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 I5 

Ransome, 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. 
Thomson, J. A. 

1927. Brachiopod morphology and genera (Tertiary and Recent). New 
Zealand Board of Sci. and Art, Manual 7. 

EXPLANATION OF PLATES 

Plate i 

Page 

A. Cyclothyris americana Cooper, new species 3 

1-5, Anterior, posterior, brachial, side, and pedicle views, respec- 
tively, X I, of holotype U.S.N.M. No. 124193a. 6, Brachial 
view of the holotype, X 2, showing elevated rim on deltidial 
plates around the foramen. 7, Beak of young pedicle valve, 
X 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. I24i86d. 10, 17, Cardinalia seen from the anterior and 
tilted, X 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. I24i86e. 13, Interior of the brachial vdve 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, NWiSW^NEi sec. 36, T. 23 S., 
R. 25 E., Bisbee Quadrangle, Cochise County, Arizona. 

B. Rectithyris vespertina Cooper, new species 4 

18-22, Pedicle, posterior, brachial, side, and anterior views, re- 
spectively, of a large specimen, X i, paratyi>e U.S.N.M. No. 
124194c. 23-27, Pedicle, posterior, anterior, brachial, and side 
views, respectively, X i. of holotype U.S.N.M. No. 124194b. 
28, Brachial view, X i, of an immature specimen, paratype 
U.S.N.M. No. I24i94d. 29, Brachial view of another paratype, 
X 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 I and X 2, respectively. 32, Interior of a brachial 
valve showing hinge plate, socket ridges, crural bases, and 
indistinct adductor scars, X 3. paratype U.S.N.M. No. 124188. 
33, Interior of a young brachial valve, X 3i showing a loop in 
an advanced stage, paratype U.S.N.M. No. 124196c. 34, In- 



l6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Page 
terior view of an adult brachial valve shovi^ing fully matured 
loop, X 2, paratype U.S.N.M. No. 124195. 35, The preceding 
loop enlarged, X 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. Gemviarcida arizonensis Cooper, new species 7 

1-5, Anterior, posterior, side, brachial, and pedicle views, re- 
spectively, of a complete specimen, X i, paratype U.S.N.AI. 
No. 124197b. 6-10, Brachial, side, posterior, anterior, and pedi- 
cle views, respectively, X 2, of holotype U.S.N.M. No. 124197a. 
II, Complete specimen tilted to show interarea and deltidium 
(= united deltidial plates), X 3. paratype U.S.N.M. No. 
I24i97d. 12, View of the interarea and disjunct deltidial plates 
of a young specimen, X 3, paratype U.S.N.M. No. I24i97g. 
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. I24i97h. 14, Im- 
mature pedicle valve, X 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. i2^igyt, showing cardi- 
nalia. 16, Interior of a brachial valve younger than the preced- 
ing. X 4, showing cardinalia, paratype U.S.N.M. No. I24i97f. 

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. 1241971. 

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. 124197J. Mural limestone, from a 
small hill 300 yards east of U. S. Highway 80, NWiSWi- 
NEi sec. 36, T. 23 S., R. 25 E., Bisbee Quadrangle, Cochise 
County, Arizona. 



NO. 4 CRETACEOUS BRACHIOPODA, ARIZONA — COOPER 1 7 

Page 
25-28, Interior, posterior, side tilted, and interior posteriorly 
tilted views, resijectively, of an adult specimen showing 
terebrataliform loop, X 2, paratype U.S.N.M. No. 124220. 
Mural limestone, Rancho Nuevo, 3 miles east of Santa Ro- 
salia, Sonora, Mexico. 

B. Gemviarciila mcnardi (Lamarck) 10 

29, 30, Brachial and side views of a complete specimen, X ii 
for comparison with Gemmarcula arisonensis, 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. :i2, 2i2, 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, X 2, for comparison 
with G. arisonensis, counterpart of pedicle valve shown by 
figure 31. Cretaceous (Green Sand), Le Mans, Sarthe, France. 

Plate 3 

A. Craniscu^ hesperin^ Cooper, new species 2 

I, 2, Brachial and side views, X i, 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, NWiSW^NEi sec. 36, T. 23 S., 
R. 25 E., Bisbee Quadrangle, Cochise County, Arizona. 

B. Psilothyris occidentalis Cooper, new species 11 

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 i, paratype U.S.N.M. No. 124190a. 11, 
Side view of a specimen broken to show descending branches 
of the long dalliniform loop and part of the wide ascending 
branch, X 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. I24202e, 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. I24202h. 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. I24202g. 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, X 4, paratype U.S.N.M. No. 
I24202i. 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 



l8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Page 
of tlie loop, X 4. paratype U.S.N.M. No. 124202!. 19, A 
slightly larger specimen than the preceding in dalliniform 
stage showing descending branches and remnantal septal at- 
tachments, X 4, paratype U.S.N.M. No. i24i9oh. 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, X 4, showing adductor scars 
indistinctly, paratype U.S.N.M. No. I24i90e. 22, Cardinalia 
of an adult brachial valve, X 4, showing crural processes 
and hinge plate with upturned edge, paratype U.S.N.M. No. 
1241901. 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. i24i9of. Horizon and locality same as above. 

C. Psilothyris tamarinda ( Sowerby ) 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 occidcntalis Cooper, new species n 

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 X 14 and X 10, 
respectively, paratype U.S.N.M. No. I24202g. Mural lime- 
stone, a small hill 300 yards east of U. S. Highway 80, NWi- 
SWiNEi sec. 36, T. 2^ S., R. 25 E., Bisbee Quadrangle, 
Cochise County, Arizona. 

B. Gonmarcula arizoncnsis Cooper, new species 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 i/, paratype U.S.N.M. No. 124199. 
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. 

Abhrcz>iations 

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.) 



LTHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131. NO. 4. PL. I 




Cyclothyris and Rectithyris 

(see explanation at end of text.) 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131. NO. 4. PL 




Gemmarcula 
(see explanation at end of text.) 



THSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131. NO. 4. PL. 3 




- J 



23 






// 



21 



22 



^&!^ 





.''^" 




■>v»- 



24 




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 



Cljarlesi ©. anb ifWarp IrJaux OTalcott 
l^esiearcl) Jf unb 



A CHECK-LIST OF THE FOSSIL AND 

PREHISTORIC BIRDS OF NORTH 

AMERICA AND THE 

WEST INDIES 



By 
ALEXANDER WETMORE 

Research Associate, Smithsonian Institution 




(Publication 4228) 



CITY OF WASHINGTON 

PUBLISHED BY THE SMITHSONIAN INSTITUTION 

JANUARY 25, 1956 



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



Cftacles! ©. anb iHarp *^aux lEalcott B^ctfcarcft jFunb 

A CHECK-LIST OF THE FOSSIL AND PRE- 
HISTORIC BIRDS OF NORTH AMERICA 
AND THE WEST INDIES 

By ALEXANDER WETMORE 
Research Associate, Smithsonian Institution 

The present check-list is an ampHfication 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 Mexico, with the addition of Baja 
California. It has seemed desirable now to include also the records, 
comparatively few in number, for Mexico 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. I3I 

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 
Itchtucknce 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 Ichthyornis have been associated with the 
Hcsperornis group in a superorder (Odontognathae) of the Neorni- 
thes, characterized by the possession of teeth. The skull of Ichthyor- 
nis always has presented an anomaly in that the teeth were in sockets 
instead of in grooves as in Hcsperornis. 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 Hcsperornis 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 
Hcsperornis, Ichthyornis, 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 
Hcsperornis 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. 



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 Megapaloclodiis 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 ambiguiis 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 1 1 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 sedis. 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. I3I 

Class AVES : Birds 

Subclass NEORNITHES: True Birds 

Superorder ODONTOGNATHAE: New World Toothed Birds 

Order HESPERORNITHIFORMES : Hesperornithes 

Family HESPERORNITHIDAE : Hesperornithes 

Genus HESPERORNIS Marsh 

Hesperornis Marsh, Amer. Journ. Sci., ser. 3, vol. 3, 1872, p. 360. Type, by 
monotypy, Hesperornis regalis Marsh. 

Hesperornis crassipes (Marsh) 

Lestornis crassipes Marsh, Amer. Journ. Sci., ser. 3, vol. 11, 1876, p. 509. 

Upper Cretaceous (Niobrara formation) : Western Kansas. 

Hesperornis montana Shufeldt 

Hesperornis montana Shufeldt, Auk, vol. 32, No. 3, July 1915, p. 293, pi. 18, 
figs. 4, 6, 8, 10, 12. 

Upper Cretaceous (Claggett formation) : i mile above mouth of 
Dog Creek, Fergus County, Montana. 

Hesperornis regalis Marsh 
Hesperornis regalis Marsh, 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 Marsh 1 
Hesperornis gracilis Marsh, Amer. Journ. Sci., ser. 3, vol. 11, 1876, p. 510. 

Upper Cretaceous (Niobrara formation) : Near Smoky Hill River, 
western Kansas. 

Genus CONIORNIS Marsha 

Coniornis Marsh, Amer. Journ. Sci., ser. 3, vol. 45, 1893, p. 82. Type, by 
monotypy, Coniornis alius Marsh. 

Coniornis altus Marsh 
Coniornis altus Marsh, 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 Marsh, Amer. Journ. Sci., ser. 3, vol. 14, 1877, p. 86. Type, by 
monotypy, Baptornis advenus Marsh. 

Baptornis advenus Marsh 

Baptornis advenus Marsh, Amcr. 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 : Ichthyornis and Allies 

Order ICHTHYORNITHIFORMES : Ichthyornis and Allies 

Family ICHTHYORNITHIDAE : Ichthyornithes 

Genus ICHTHYORNIS Marsh 

Ichthyornis Marsh, Amer. Journ. Sci., ser. 3, vol. 4, November 1872, p. 344. 
Type, by monotypy, Ichthyornis dispar Marsh. 

Ichthyornis agilis (Marsh) 

Craculavus 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 AIarsh, 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 Marsh, Amer. Journ. Sci., ser. 3, vol. 4, 1872, p. 344. 

Upper Cretaceous (Niobrara formation) : Near Solomon River, 
Kansas. 

Ichthyornis lentus (Marsh) 

Graculavus lentus Marsh, Amer. Journ. Sci., ser. 3, vol. 14, 1877, p. 253. 

Upper Cretaceous : Near McKinney, Texas. 

2 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. I3I 

Ichthyornis tener Marsh 
Ichthyornis tcner Marsh, Odontornithes, 1880, pp. 151, 198, pi. 30, fig. 8. 

Upper Cretaceous (Niobrara formation) : Wallace County, Kansas. 

Ichthyornis validus Marsh 

Ichthyornis validus Marsh, Odontornithes, 1880, pp. 147, 153, 198, pi. 30, figs. 
11-14. 

Upper Cretaceous (Niobrara formation) : Near Solomon River, 
Kansas. 

Ichthyornis victor Marsh 

Ichthyornis victor Marsh, 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 : Apatornitiies 

Genus APATORNIS Marsh 

Apatornis Marsh, Amer. Journ. Sci., ser. 3, vol. 5, Jan. 21, 1873, p. 162. Type, 
by monotypy, Ichthyornis celer Marsh. 

Apatornis celer (Marsh) 

Ichthyornis celer Marsh, Amer. Journ. Sci., ser. 3, vol. 5, 1873, p. 74. 

Upper Cretaceous (Niobrara formation) : Butte Creek, Logan 
County, near Smoky Hill River, Kansas. 

Supcrorder NEOGNATHAE: Typical Birds 

Order CAENAGNATHIFORMES : Caenagnatiius 

Family CAENAGNATHIDAE: Caenagnatiius 

Genus CAENAGNATHUS Sternberg 

Caenagnathus Stf.knrerg, Journ. Pal., vol. 14, January 1940, p. 81. Type, by 
original designation, Caenagnathus collinsi Sternberg. 

Caenagnathus collinsi Sternberg * 

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. 

* 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 Brunnich, Orn. Borealis, 1764, p. 38. 

Modern form reported from late Pleistocene (Palos Verdes sand) ; 
Newport Bay, Orange County, California. 

Gavia arctica (Linnaeus) : Arctic Loon 

Colymbus arciicus Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 135. 

Modern form reported from late Pleistocene (Palos Verdes sand) : 
San Pedro, Los Angeles County, California. 

Gavia concinna Wetmore 

Gavia concinna Wetmore, Journ. Morph., vol. 66, No. i, Jan. 2, 1940, p. 25, 
figs. 1-4. 

Pliocene (Etchegoin formation) : Sweetwater Canyon (type lo- 
cality), 5^ 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 Brodkorb 

Gavia howardae Brodkorb, Condor, vol. 55, No. 4, July 20, 1953, p. 212, fig. iB. 

Pliocene (Bone Valley formation) : Pierce (type locality) and 
Brewster, Polk County, Florida. 

Subfamily GAVIELLINAE: Gaviella 

Genua 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. I3I 

Gaviella pusilla (Shufeldt) 

Gavia pusilla Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, Febru- 
ary 1 91 5, p. 70, pi. 13, fig. 106. 

Probably from Oligocene (White River formation) : near Lusk, 

Wyoming.^ 

Order COLYMBIFORMES : Grebes 

Family COLYMBIDAE: Grebes 

Genus COLYMBUS Linnaeus 

Colymhus Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 135. Type, by subse- 
quent designation, Colymhus cristatus Linnaeus (Baird, Brewer, and 
Ridgway, 1884). 

Subgenus DYTES Kaup 

Dytes Kaup, Skizz. Entw.-Gesch. Eur. Thierw., 1829, p. 41. Type, by subse- 
quent designation, Dytes cormitus Kaup =: Colymhus auritus Linnaeus 
(Gray, 1842). 

Colymbus auritus Linnaeus: Horned Grebe 
Colymhus auritus Linnaeus, Syst. Nat., ed. 10, vol. i, 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 Hablizl: Eared Grebe 

Colymhus caspicus Hablizl, 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 Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, 
February 191 5, p. 54. 

? Oligocene (John Day) : Lower Willow Creek, Baker County, 
Oregon. 



^ See Wetmore, A., Journ. Morph., vol. 66, Jan. 2, 1940, p. 30. 
° Specimens from Fossil Lake, Oregon, formerly included under this species 
have been found by Hildegarde Howard to represent Colymhus caspicus and 

Podilymbus podiccps. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 9 

Colymbus parvus Shufeldt 
Colymbiis parvus Shufeldt, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 6, 
July 9, 1913, p. 136, pi. 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 Brodkorb, Ann. Mag. Nat. Hist., ser. 12, vol. 6, December 1953, 
p. 953, I fig. Type, by original designation, Pliodytes lanquisti Brodkorb. 

Pliodytes lanquisti Brodkorb 
Pliodytes lanquisti Brodkorb, Ann. Mag. Nat. Hist., ser. 12, vol. 6, December 

1953, P- 953, I fig- 
Pliocene (Bone Valley formation) : Near Brewster, Polk County, 

Florida. 

Genus AECHMOPHORUS Coues 

^chmophorus CouES, Proc. Acad. Nat. Sci. Philadelphia, vol. 14, No. 5, April- 
May (Aug. i), 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. Miller, Univ. California Publ., Bull. Dept. Geol., 
vol. 6, No. 4, Feb. 4, 191 1, p. 83, figs. 1-3. 

Late Pleistocene: Fossil Lake (type locality), Oregon;^ Palos 
Verdes sand, New^port Bay, Orange County, Playa del Rey, San 
Pedro, Los Angeles County, and near Manix, San Bernardino County, 
California. 



'' Includes 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. 



10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Genus PODILYMBUS Lesson 

PodilymbtiS Lesson, Traite d'Orn., livr. 8, June 11, 1831, p. 595. Type, by 
monotypy, Podiccps caroliiicnsis Latham = Colyinbits podiceps Linnaeus, 

Podilymbus podiceps (Linnaeus) : Pied-billed Grebe ^ 

Colymbus Podiceps Linnaeus, Syst. Nat., ed. 10, vol. i, 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, Mexico. 

Order PROCELLARIIFORMES : Albatrosses, Shearwaters, 
Petrels, and Allies 

Family DIOMEDEIDAE: Albatrosses 

Genus DIOMEDEA Linnaeus 

Diomedia Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 132. Type, by subse- 
quent designation, Diomedca cxnlans Linnaeus (Gray, 1840). 

Diomedea albatrus Pallas : Short-tailed Albatross 

Diomedca albatrus Pallas, Spic. Zool., vol. i, fasc. 5, 17C9, p. 28. 

Modern form reported from late Pleistocene (Palos Verdes sand) : 
Newport Bay, Orange County, Playa del Rey, Los Angeles County, 
California. 

Diomedea anglica Lvdekker 

Diomedea anglica Lvdekker, 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° 

Puffimts Brisson, Orn., 1760, vol. i, p. 56; vol. 6, p. 130. Type, by tautonymy, 
Puffinus Brisson z= Proccllaria puffinus Briinnich. 



^ Podilymbus magnus Shufeldt, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 
6, July 9, 1913, p. 136, pi. 38, figs. 439-440, 449, has been identified as F. podiceps 
by Wetmore, California Acad. Sci., vol. 23, Dec. 30, 1937, pp. 198-199. 

° Described by Lydckker from the Upper Pliocene at Foxhall, Suffolk, Eng- 
land. Recorded from Florida by Wetmore, Proc. New England Zool. Club, 
vol. 22, June 23, 1943, pp. 66-67, pl- 12, figs. 10-15. 

^° Puffinus parz'us Shufeldt, Ibis, October 1916, p. 632, from Recent deposits 
in the bone caves of Bermuda is considered a synonym of Puffinus Ihcrminieri. 
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 
PuflSnus griseus (Gmelin) : Sooty Shearwater 

Proccllaria grisca Gmelin, Syst. Nat., vol. i, 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. 

PufiSnus puflSnus (Brunnich) : Common Shearwater 

Proccllaria 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, Calif ornia.^^ 

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. Miller, Carnegie Inst. Washington Publ. 349, 
August 1925, p. Ill, pis. I, 2, 7a. 

Middle Miocene (Temblor formation, Turrit ella ocoyaiia zone) : 
Lompoc (type locality). Miocene (Monterey shale) : Lomita and 
San Pedro breakwater, San Pedro, California. 

Puffinus kanakoffi Howard ^^ 

Puffinus kanakoffi Howard, Carnegie Inst. Washington Publ. 584, June 22, 
1949, p. 187, pi. 2, figs. 3, 5. 

Middle Pliocene (San Diego formation) : Washington Boulevard 
Freeway, San Diego, California. 

Puffinus felthami Howard ^^ 

Puffinus felthami Howard, Carnegie Inst. Washington Publ. 584, June 22, 
1949, p. 194, pi. 2, figs. 4, 6. 

Late Lower Pliocene: 3 miles north of Corona del Mar, Orange 
County, California. 



^^ The California records refer to Puffiinus puffinus opisthomelas Coues, for- 
merly listed as a separate species. 
^2 Subgeneric allocation provisional. 



12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I3I 

Subgenus ARDENNA Reichenbach 

Ardenna Reichenbach, Avium Syst. Nat., 1852 (1853), p. iv. Type, by 
monotypy, Procellaria major Faber = P. gravis O'Reilly. 

Puflfinus conradi Marsh 
Puffinus conradi Marsh, Amer. Journ. Sci., ser. 2, vol. 49, 1870, p. 212. 

Middle Miocene (Calvert formation) : Maryland. 



Genus FULMARUS Stephens 

Fuhmrns Stephens, in Shaw, Gen. Z06I., vol. 13, pt. i, 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 jiircata Gmelin. 

Oceanodroma hubbsi Miller 

Oceanodroma hubbsi L. H. Miller, Condor, vol. 53, No. 2, Mar. 27, 1951, p. 78, 
fig. I. 

Upper Miocene (Capistrano formation ") : About i mile south of 
Capistrano Beach, Orange County, California. 

Order PELECANIFORMES : Tropicbirds, Pelicans, Frigatebirds, 

and Allies 

Suborder PELECANI : Pelicans, Boobies, Cormorants, and Darters 

Superfamily PELECANOIDEA : Pelicans and Allies 

Family PELECANIDAE: Pelicans 

Genus PELECANUS Linnaeus 

Pelecanus Linnaeus, Syst. Nat., ed. 10, vol. i, 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 I3 

Subgenus CYRTOPELICANUS Reichenbach 

Cyrtopclicanns Reiciienbach, Avium Syst. Nat., 1852 (1853), p. vii. Type, 
by original designation, Pckcamis trachyrhynchns Latham = P. crythro- 
rhy licit OS Gmelin. 

Pelecanus erythrorhynchos Gmelin : White Pelican 

Pdccanus erythrorhynchos Gmelin, Syst. Nat., vol. i, pt. 2, 1789, 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 Wetmore, Smithsonian Misc. Coll., vol. 87, No. 20, Dec. 
27, 1933, p. 3, figs. 1-2. 

Upper Pliocene (Hagerman lake beds) : Near Hagerman, Idaho. 



Subgenus LEPTOPELICANUS Reichenbach 

Leptopclicanus 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. i, 1766, p. 215. 

Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 



Family CYPHORNITHIDAE: Cyphornithes 

Genus CYPHORNIS Cope 

Cyphornis Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 9, May 31, 
1894, p. 449. Type, by monotypy, Cyphornis tnagnus Cope. 

Cyphornis magnus Cope 

Cyphornis magnus Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 9, 
May 31, 1894, p. 451. 

Middle Oligocene : Carmanah Point, Vancouver Island, British 
Columbia. 

Genus PALAEOCHENOIDES Shufeldt 

Palaeochenoides Shufeldt, Geol. Mag., n.s. 4, vol. 3, August 1916, p. 347. 
Type, by monotypy, Palaeochendides mioceanus Shufeldt. 



14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 

Palaeochenoides mioceanus Shufeldt 

Palaeochenoides mioceanus Shufeldt, Geol. Mag., n.s. 4, vol. 3, August 1916, 
p. 347, pl. IS- 

Miocene (Hawthorn formation) : Near source of Stono River, 
South Carolina. 



Superfamily SULOIDEA: Boobies, Cormorants, Darters, and Allies 

Family SULIDAE: Boobies and Gannets 

Genus SULA Brisson 

Sula Brisson, Orn., 1760, vol. i, p. 60; vol. 6, p. 494. Type, by tautonymy, 
Sula Brisson = Pelccanus piscator Linnaeus. 

Subgenus SULA Brisson 

Sula stocktoni Miller 

Sula stocktoni L. H. Miller, Publ. Univ. California at Los Angeles Biol. 
Sci., vol. I, No. 5, Mar. 12, 1935, p. 75, fig. 2. 

Middle Miocene (Monterey shale) : Near Lomita, Los Angeles 
County, California. 

Sula willetti Miller 

Siila willetti L. H. Miller, Carnegie Inst. Washington Publ. 349, August 1925, 
p. 112, pis. 3, 8, fig. I. 

Middle Miocene (Temblor formation, Turritella ocoyana zone) : 
Lompoc, Santa Barbara County, California, 

Sula guano Brodkorb 

Sula guano Brodkorb, Florida Geol, Surv. Rep. Invest. No. 14, November 
1955, P- 9, figs. 2, 5, 8. 

Pliocene (Bone Valley formation) : Near Brewster, Polk County, 
Florida. 

Sula pbosphata Brodkorb 

Sula phosphata Brodkorb, Florida Geol. Surv. Rep. Invest. No. 14, November 
1955, p. II, figs. 3> 6, 9- 

Pliocene (Bone Valley formation) : Near Brewster, Polk County, 
Florida. 

Subgenus MICROSULA Wetmore 

Microsula Wetmore, Proc. U. S. Nat. Mus., vol. 85, Jan. 14, 1938, p. 25. Type, 
by original designation, Sula (Microsula) az'ita Wetmore. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 1 5 

Sula avita Wetmore 

Sula avita Wetmore, 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. Miller, Carnegie Inst. Washington Publ. 349, August 1925, 
p. 114. Type, by monotypy, Miosula media Miller. 

Miosula media Miller 

Miosula media L. H. Miller, 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 Howard 

Miosula recentior Howard, Carnegie Inst. Washington Publ. 584, June 22, 
1949, p. 190, pl. 2, figs. i-2a. 

Middle Pliocene (San Diego formation) : Curlew Street, opposite 
Ostego Drive, San Diego, California. 



Genus MORUS Vieillot 

Morns Vieillot, Analyse, April 1816, p. 63. Type, by monotypy, Pelecanus 
bassanus Linnaeus. 

Morus loxostyla (Cope)i* 

Sula loxostyla Cope, Trans. Amer. Philos. Soc, n.s., vol. 14, December 1870, 
p. 236, fig. S3. 

Miocene: Calvert County (type locality), Maryland; New Jersey. 

Morus vagabundus Wetmore 

Moris vagabundus Wetmore, 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. 

^* 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. 



l6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Moms lompocana (Miller) 

Sula lompocana L. H. Miller, Carnegie Inst. Washington Publ. 349, August 
1925, p. 114, pis. 4, 7b, 9- 

Middle Miocene (Temblor formation, Turritella ocoyana zone) : 
Lompoc, Santa Barbara County, California. 

Morus peninsularis Brodkorb 

Mortis peninsularis Brodkorb, Florida Geol. Surv. Rep. Invest. No. 14, No- 
vember 195s, p. 8, figs. I, 4, 7. 

Pliocene (Bone Valley formation) : Near Brewster, Polk County, 
Florida. 

Morus reyana Howard 
Moris reyana Howard, 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 is 

Graculavus Marsh, Amer. Journ. Set., scr. 3, vol. 3, 1872, p. 363. Type, by 
subsequent designation, Graculavus vclox Marsh (Hay, 1902). 

Graculavus pumilus Marsh 

Graculavus pumilus Marsh, Amer. Journ. Sci., ser. 3, vol. 3, 1872, p. 364. 

Paleocene (Hornerstown marl) : Hornerstown, New Jersey. 

Graculavus velox Marsh 

Graculavus velox Marsh, Amer. Journ. Sci., ser. 3, vol. 3, 1872, p. 363. 
Paleocene (Hornerstown marl) : Hornerstown, New Jersey. 

Genus PHALACROCORAX Brisson lo 

Phalacrocorax Brisson, Orn., 1760, vol. i, p. 60; vol. 6, p. 511. Type, by 
tautonymy, Phalacrocorax Brisson = Pelecanus carbo Linnaeus. 

Phalacrocorax wetmorei Brodkorb 

Pltalaerocorax ivetmorei Brodkorb, Florida Geol. Surv. Rep. Invest. No. 14, 
November 1955, p. 12, figs. 10, 11. 

Pliocene (Bone Valley formation) : Near Brewster, Polk County, 
Florida. 



^^ Limosavis Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, February 
1915, p. 19, proposed as a new genus for Graculavus vclox Marsh, is a synonym 
of Graculavus Marsh, as both names are based on the same species. 

1" No subgenera are recognized in recent studies of the cormorants. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE I7 

Phalacrocorax auritus (Lesson) : Double-crested Cormorant 
Carbo auritus Lesson, Traite 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 (Brandt) : Brandt's Cormorant 

Carbo penicillatus Brandt, Bull. Sci. Acad. Imp. Sci. St.-Petersbourg, 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. Miller, Condor, vol. 31, No. 4, July 15, 1929, 
p. 167, figs. 58-59. 

Upper Miocene (Modclo formation) : Calabasas, Los Angeles 
County, California. 

Phalacrocorax idahensis (Marsh) 
Graculus idahensis Marsh, Amer. Journ. Sci., ser. 2, vol. 49, 1870, p. 216. 

Pliocene: Castle Creek; Ovi^yhee County (type locality), Idaho; 
Pliocene (Bone Valley formation) : Near Brewster, Polk County, 
Florida. Upper Pliocene (Hagerman lake beds) : Near Hagerman, 
Idaho. 

Phalacrocorax macropus (Cope) 

Graculus macropus Cope, Bull. Geol. Gcogr. Surv. Terr., vol. 4, No. 2, 1878, 
p. 386. 

Late Pleistocene: Fossil Lake, Oregon." 

Phalacrocorax marinavis Shufeldt 

Phalacrocorax marinazns Shufeldt, Trans. Connecticut Acad. Sci., vol. 19, 
February 1915, p. 56, pi. 14, figs. 114, 116-118, 122. 

? Oligocene (John Day) : Willow Creek, Oregon. 



^■^ Shufeldt, Auk, 1915, pp. 485-488, has identified material from the Miocene of 
Montana as this si)ecies, but examination of the specimen reveals that this is 
in error. 



l8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Phalacrocorax mediterraneus Shufeldt 

Phalacrocorax mediterraneus Shufeldt, Trans. Connecticut Acad. Arts Sci., 
vol. 19, February 1915, p. 58, pi. 15, fig. 138. 

Lower Oligocene (Chadron formation) : Gerry's Ranch, Weld 
County, Colorado. 

Phalacrocorax rogersi Howard 

Phalacrocorax rogersi Howard, 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 Howard 

Phalacrocorax kennelli Howard, Carnegie Inst. Washington Publ. 584, June 
22, 1949, p. 188, pi. 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. i, p. 60; vol. 6, p. 476. Type, by tautonymy 
and monotypy, Anhinga Brisson = Plotiis anhinga Linnaeus. 

Aiihinga anhinga (Linnaeus) : Anhinga 

Plotus Anhinga Linnaeus, Syst. Nat., cd. 12, vol. i, 1766, p. 218. 

Modern form reported from Pleistocene (Melbourne bone bed) : 
Melbourne (stratum 2), Florida. 

Order CICONHFORMES: Herons, Storks, and Allies 

Suborder ARDEAE: Herons, Bitterns, and Allies 

Family ARDEIDAE : Herons and Bitterns 

Subfamily ARDEINAE: Herons and Egrets 

Genus ARDEA Linnaeus is 

Ardea Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 141. Type, by subsequent 
designation, Ardea cincrea Linnaeus (Gray, 1840). 



18 Ardea sellardsi Shufeldt, Journ. Geol., January-February (January) 1917, 
p. 19, described from Vcro (stratum 3), Florida, proves to be based on tiie 
tibiotarsus of Meleagris gallopavo. See Wetniore, Smithsonian Misc. Coll., 
vol. 85, No. 2, Apr. 13, 1931, p. 32. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE IQ 

Ardea herodias Linnaeus : Great Blue Heron 
Ardca Herodias Linnaeus, Syst. Nat., ed. lo, vol. i, 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 Brodkorb 
Ardea polkensis Brodkorb, Florida Geol. Surv. Rep. Invest. No, 14, November 
I9S5, p. 17, figs. 13, 14, 15- 

Pliocene (Bone Valley formation) : Near Brewster, Polk County, 
Florida. 

Genus CASMERODIUS Gloger 

Casmcrodius Gloger, Hand- und Hilfsbuch Naturg., 1842 (1841), p. 412. 
Type, by subsequent designation, Ardca egretta Gmelin (Salvadori, 1882). 

Casmerodius albus (Linnaeus) : Common Egret 

Ardca alba Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 144. 

Modern form reported from Pleistocene : Melbourne, Seminole 
Field, Pinellas County, and Venice, Florida. Late Pleistocene : Rancho 
La Brea, Los Angeles, California; Banos de Ciego Montero, Santa 
Clara Province, Cuba. 

Genus LEUCOPHOYX Sharpe 

Leucophoyx Sharpe, Bull. Brit. Orn. Club, vol. 3, Apr. 30, 1894, p. xxxix. 
Type, by original designation and monotypy, Ardea candidissima Gmelin == 
Ardea thnla Molina. 

Leucophoyx thula (Molina) : Snowy Egret 

Ardca Thida Molina, Sagg. Stor. Nat. Chili, 1782, p. 235. 

Modern form reported from Pleistocene: Bradenton, Florida, 

Genus HYDRANASSA Baird 

Hydranassa Baird, in Baird, Cassin, and Lawrence, Rep. Expl. Surv, R. R. 
Pac, vol. 9, 1858, p. 660. Type, by original designation, Ardca hidoviciana 
Wilson =: Egretta ritficollis Gosse. 

Hydranassa tricolor (Muller) : Tricolored Heron 

Ardea tricolor P. L. S. Muller, Natursyst. Suppl., 1776, p. iii. 

Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 

^8 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. I3I 

Genus FLORIDA Baird 

Florida Baird, 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. i, 1758, p. 143. 

Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 

Genus BUTORIDES Blyth 

Butorides Blyth, 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. i, 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 injausius Forster = /Irtira nyclicorax Lin- 
naeus. 

Nycticorax nycticorax (Linnaeus) : Black-crowned Night Heron 

Ardea Nycticorax Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 142. 

Modern form reported from Pleistocene : San Josecito Cave, Aram- 
berri, Nuevo Leon, Mexico ; Bradenton, and Itchtucknee River, Flor- 
ida. Late Pleistocene: McKittrick, Kern County, and Rancho La 
Brea, Los Angeles, California. 

Genus NYCTANASSA Stejneger 

Nyctamssa 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. i, 1758, p. 143. 

Modern form reported from Pleistocene : Seminole Field, Pinellas 
County, Florida.^" 

-0 Larus vera Shufeldt, Journ. Geol., 1917, p. 18, from stratum 3 of Vero, 
Florida, is Nyctanassa z'iolacca, according to Wetmore, Smithsonian Misc. Coll., 
vol. 85, No. 2, 1931, pp. 3, II, and 16. Cooke, Florida Geol. Surv., Geol. Bull. 29, 
I945i 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 
1915, P- 39- Type, by monotypy, Eoceornis ardetta Shufeldt. 

Eoceornis ardetta Shufeldt 

Eoceornis ardetta Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, 
February 1915, p. 39, pi. 13, fig. 102. 

Eocene (Bridger formation) : Henry's Fork, Wyoming. 

Subfamily BOTAURINAE: Bitterns 

Genus IXOBRYCHUS Billberg 

Ixohrychus Billberg, Syn. Faunae Scand., vol. i, 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. i, pt. 2, 1789, p. 645. 

Modern form reported from late Pleistocene : Banos 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 stcllaris Linnaeus (Gray, 1840). 

Botaurus lentiginosus (Rackett) : American Bittern 

Ardea lentiginosa Rackf.tt, 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 
1915, P- 33- Type, by monotypy, Botauroides parvus Shufeldt. 

Botauroides parvus Shufeldt 

Botauroides parz'us Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, 
February 1915, p. 33. 

Eocene ( ? Bridger formation) : "Spanish John Meadow," Wy- 
oming. 



'^^ Ardea paloccidcntalis 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. I3I 

Suborder CICONIAE: Storks, Ibises, and Spoonbills 

Superfamily CICONIOIDEA: Storks and Wood Ibises 

Family CICONIIDAE: Storks and Jabirus 

Subfamily CICONIINAE: Storks 

Genus CICONIA Brisson 

Ciconia Brisson, Orn., 1760, vol. i, p. 48; vol. 5, p. 361. Type, by tautonymy, 
Ciconia = Ardea ciconia Linnaeus. 

Ciconia maltha Miller 

Ciconia maltha L. H. Miller, Univ. California Publ., Bull. Dept. Geol., vol. 5, 
No. 30, Aug. 5, 1910, p. 440, figs. 1-7. 

Upper Pliocene (Hagerman lake beds) : Barbour Ranch, Snake 
River, Idaho. Pleistocene: American Falls, Idaho; Vero (stratum 2), 
Melbourne (stratum 2), Itchtucknee River, 6^ 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; Banos de Ciego Montero, Santa Clara Province, 
Culia.-2 

Subfamily MYCTERIINAE: Wood Ibises 

Genus MYCTERIA Linnaeus 

Myctcria Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 140. Type, by monotypy, 
Mycteria americana Linnaeus. 

Mycteria wetmorei Howard -^ 

Mycteria zvetmorci Howard, Condor, vol. 37, Sept. 15, 1935, p. 253, fig. 47. 

Late Pleistocene: Rancho La Brea, Los Angeles, California. 

Superfamily THRESKIORNITHOIDEA : Ibises 

Family THRESKIORNITHIDAE: Ibises and Spoonbills 

Subfamily THRESKIORNITHINAE: Ibises 

Genus PLEGADIS Kaup 

Plegadis Kaup, Skizz. Entw.-Gcs. Eur. Thierw., 1829, p. 82. Type, by mono- 
typy, Tantalus falcinellus Linnaeus. 



22 Records formerly listed as Jabiru myctcria (Lichtenstein) have all been 
assigned to the present species by Hildegarde Howard, in Carnegie Inst. Wash- 
ington Publ. 530, Jan. 19, 1942, p. 202. Jabiru zvcillsi Sellards, therefore, be- 
comes a synonym of Ciconia maltha. 

-3 Replaces Myctcria americana as listed in Check-list of North American 
Birds, ed. 4, 1931, P- 4i6. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 23 

Plegadis chihi (Vieillot) : White-faced Ibis 

Numenius chihi Vieillot, Nouv. Diet. Hist. Nat., nouv. ed., vol. 8, March 
1817, p. 303. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, California. 

Genus EUDOCIMUS Wagler 

Eudocimus Wagler, 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. i, 1758, p. 145. 

Modern form reported from Pleistocene : Seminole Field, Pinellas 
County, and Haile, Alachua County, Florida. 

Subfamily PLATALEINAE : Spoonbills 

Genus AJAIA Reichenbach 

Ajaia Reichenbach, Avium Syst. Nat., 1852 (1853), p. xvi. Type, by original 
designation, Ajaia rosea Reichenbach = Platalca ajaja Linnaeus. 

Ajaia ajaja (Linnaeus) : Roseate Spoonbill 

Platalca Ajaja Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 140. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, California. 

Suborder PHOENICOPTERI : Flamingos 

Family PHOENICOPTERIDAE: Flamingos 

Genus PHOENICOPTERUS Linnaeus 

Phoenicopterus Linnaeus, Syst. Nat., ed. 10, vol. i, 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 Howard 

Phoenicopterus jninutus Howard, Geol. Surv. Prof. Pap. 264-J, June 1955, 
p. 202, pi. 50. 

Late Pleistocene ; Manix lake beds, near Manix, San Bernardino 
County, California. 



24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Phoenicopterus stocki Miller 
Phoenicopterus stocki L. H. Miller, Wilson Bull., vol. 56, No. 2, June 1944, 
p. 77, figs. I, 2. 

Pliocene (Rincon) : Chihuahua, Mexico. 

Phoenicopterus floridanus Brodkorb 

Phoenicopterus floridanus Brodkorb, 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 : Paloelodus and Allies 

Genus MEGAPALOELODUS Miller 

Megapaloelodus A. H. Miller, 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. Miller, Univ. California Publ., Bull. Dept. 
Geol. Sci., vol. 27, No. 4, June 22, 1944, p. 86, fig. i. 

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. Miller and L. V. Compton, Condor, vol. 41, No. 4, July 15, 
1939. P- 153- Type, by original designation, Paranyroca magna Miller and 
Compton. 

Paranyroca magna Miller and Compton 

Paranyroca magna A. H. Miller 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 

Cygmis Bechstein, Orn. Taschenb. Deutschl., vol. 2, 1803, p. 404, footnote. 
Type, by tautonymy, Anas cygnus Linnaeus. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 2$ 

Subgenus STHENELIDES Stejneger 

Stheitclides Stejneger, Stand. Nat. Hist., vol. 4, 1885, p. 143. Type, by 
monotypy, Anas mclancoripha Molina. 

Cygnus paloregonus (Cope) 2* 

Cygnus paloregonus Cope, 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 Wagler, Isis von Oken, 1832, col. 1234. Type, by subsequent designa- 
tion, Cygnus nmsicus Bechstein ^ /4Ma.y cygnus Linnaeus (Gray, 1840). 



Subgenus OLOR Wagler 
Olor columbianus (Ord) : Whistling Swan 

Anas columbianus Ord, 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. i, July 1908, p. 3. Type, by mono- 
typy, Cygnus bticcinator 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 Scopoli, Annus I, Historico-Naturalis, 1769, p. 67. Type, by subse- 
quent designation. Anas bernicia Linnaeus (Bannister, 1870). 



2* Subgeneric allocation tentative. 

2^ Specimens named Cygnus matthczvi and Anscr 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. I3I 

Branta canadensis (Linnaeus) : Canada Goose 
Anas canadensis Linnaeus, Syst. Nat., ed. 10, vol. i, 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 (Richardson) : Richardson's Goose 

Anscr Hutchinsii Richardson, 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. i, 1758, p. 124. 
Modern form reported from Pleistocene : Fossil Lake, Oregon. 

Branta esmeralda Burt 

Branta esmeralda Burt, Univ. California Publ., Bull. Dept. Geol. Sci., vol. 18, 
No. 6, Mar. 19, 1929, p. 222, pi. 20. 

Upper Miocene (Esmeralda formation) : Fish Lake Valley, Es- 
meralda County, Nevada. 

Branta howardae Miller 

Branta hoivardae L. H. Miller, Condor, vol. 32, No. 4, July 15, 1930, p. 208, 
fig. 74- 

Lower Pliocene (Ricardo formation) : Mojave Desert area, Kern 
County, CaUfornia. 

Branta dickeyi Miller 

Branta dickeyi L. H. Miller, Condor, vol. 26, No. 5, Sept. 15, 1924, p. 179, 
fig. 46. 

Upper Pliocene: Dry Creek, Malheur County, Oregon. Late 
Pleistocene: McKittrick, California. 



'^ Specimens from Fossil Lake range in size from modern B. c. 7ninima to B. c. 
canadensis. 
'" Recorded as Branta canadensis canadensis. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 2/ 

Branta hypsibata (Cope)^^ 
Anscr hypsibaius Cope, 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, pi. IS, fig. 17. 

Late Pleistocene : Fossil Lake, Oregon. 

Genus ANABERNICULA Ross 20 

Aiuibcniicula Ross, Trans. San Diego Soc. Nat. Hist., vol. 8, No. 15, Aug. 24, 
1935, P- 107. Type, by monotypy, Anahernicula 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 IS, 1930, p. 92. Tyi>e, by original designation, Presbychen abavus 
Wetmore. 

Presbychen abavus Wetmore 

Presbychen abavus Wetmore, Proc. California Acad. Sci., ser. 4, vol. 19, 
No. 8, July IS, 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. i, p. 58; vol. 6, p. 261. Type, by tautonymy, 
Anser domesiica Brisson = /ina.j 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 hypcrborea. 

2^ Possibly representative of a distinct subfamily. See Howard, Carnegie Inst. 
Washington Publ. 551, Jan. 25, 1946, pp. 172-173. 

^° See Howard, Condor, 1936, p. 35. 



28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Anser albifrons (Scopoli) : White-fronted Goose 
Branta albijrons Scopoli, Annus I, Historico-Naturalis, 1769, p. 69. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon ; 
Rancho La Brea, Los Angeles, and San Pedro ^^ (Palos Verdes sand), 
Los Angeles Comity, California. 

Genus CHEN Boie 

Chen Boie, Isis von Oken, vol. 10, Heft 5, 1822, col. 563. Type, by monotypy, 
Anscr hyperborcus Pallas. 

Chen hyperborea (Pallas) : Snow Goose 

Anscr hyperborcus Pau:-as, Spic. Zool., vol. i, fasc. 6, 1769, p. 25. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon ; 
Rancho La Brea, Los Angeles, and McKittrick, California.^^ 

Chen rossii (Cassin) : 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 ^3 

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: Treeducks 

Genus DENDROCYGNA Svsrainson 

Dcndrocygna Swainson, Class. Birds, vol. 2, July i, 1837, p. 365. Type, by 
subsequent designation, Anas arcuata Horsfield (Gray, 1840). 

Dendrocygna eversa Wetmore 

Doidrocygna cvcrsa Wetmore, Proc. U. S. Nat. Mus., vol. 64, art. 5, Jan. 
15, 1924, p. 3, figs. 1-2. 

Upper Pliocene (Blancan) : Near Benson, Arizona. 



3^ Specimen with size of the subspecies frontalis. 

32 Chen caeritlcsccns 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. i, 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. Miller, 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. Miller, 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: Surface-feeding Ducks 

Genus ANAS Linnaeus 

Anas Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 122. Type, by subsequent 
designation, Anas boschas Linnaeus =::^. platyrhynchos Linnaeus (Les- 
son, 1828). 

Anas platyrhynchos Linnaeus: Mallard 
Anas platyrhynchos Linnaeus, Syst. Nat., ed. 10, vol. i, 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 
A7ias obscura rubripes Brewster, Auk, vol. 19, No. 2, April 1902, p. 184. 

Modern form reported from Pleistocene : Itchtucknee River, 
Florida. 

Anas fulvigula Ridgway : Mottled Duck 

Anas obscura var. ftdvigula Ridgway, 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. i, 1758, p. 125. 

Modern form reported from late Pleistocene: McKittrick and 
Rancho La Brea, Los Angeles, California.''* 

^* Listed erroneously in Check-list of North American Birds, ed. 4, 1931, p. 421, 
from Itchtucknee River, Florida. 



30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Anas acuta Linnaeus: Pintail 
Anas acuta Linnaeus, Syst. Nat., ed. 10, vol. i, 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. i, 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 Wetmore, Univ. Kansas Sci. Bull., vol. 30, pt. i. No. 9, 
May 15, 1944, P- 92, figs. 1-3- 

Upper Pliocene (Rexroad formation) : Meade County, Kansas 
(type locality) ; 2 miles south of Benson, Arizona. 

Anas cyanoptera Vieillot: Cinnamon Teal 

Anas cyanoptera Vieillot, Nouv. Diet. Hist. Nat., nouv. ed., vol. 5, December 
1816, p. 104. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon ; 
McKittrick, Kern County, California. 

Anas Integra (Miller). 

Querqucdula Integra A. H. Miller, 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 pcnelope Lin- 
naeus (Eyton, 1838). 



•'•'• There are also records for the Upper Miocene or lower Pliocene of Cedar 
Mountain, Nevada, by L. H. Miller, Univ. 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. i, 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. i, 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 Boie, Isis von Oken, vol. 10, Heft 5, 1822, col. 564. Type, by 
nionotypy, Anas clypeata Linnaeus. 

Spatula clypeata (Linnaeus) : Shoveler 
Anas clypeata Linnaeus, Syst. Nat., ed. 10, vol. i, 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: Diving Ducks" 

Genus AYTHYA Boie 

Ay thy a Boie, Tageb. Reise Norwegcn, 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 coUaris (Donovan) : Ring-necked Duck 

Anas collaris Donovan, Brit. Birds, vol. 6, 1809, pi. 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. 

^'^ Polysiicta stelleri, Bucephala islandica, and Ilistrionicns histrionicus re- 
ported from Fossil Lake by Sliufeldt were wrongly identified and are eliminated 
from the list. See Howard, Carnegie List. Washington Publ. 551, Jan. 25, 1946, 
p. 176. 



32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Aythya valisineria (Wilson) : Canvasback 
Anas valisineria Wilson, Amer. Orn., vol. 8, 1814, p. 103, pi. 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 Baird, in Baird, Cassin, and Lawrence, Rep. Expl. Surv. R. R. Pac, 
vol. 9, 1858, pp. xxiii, L, 787, 788, 795. Type, by original designation, 
Atms albcola Linnaeus. 

Bucephala albeola (Linnaeus) : Bufiflehead 
Anas Albcola Linnaeus, Syst. Nat., ed. 10, vol. i, 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 Brodkorb 

Bucephala ossivallis Brodkorb, Florida Geol. Surv. Rep. Invest. No. 14, 
November 1955, p. 18, figs. 16, 17. 

Pliocene (Bone Valley formation) : Near Brewster, Polk County, 
Florida. 

Genus CLANGULA Leach 

Clangula Leach, in Ross, Voy. Discovery, 1819, app., p. XLViii. Type, by 
monotj'py, Clangula glacialis Linnaeus = Anas hyemalis Linnaeus. 

Clangula hyemalis (Linnaeus) : Oldsquaw 

Anas hyemalis Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 126. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon. 

Genus MELANITTA Boie 

Melanitta Boie, Isis von Okeii, 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 (Bonaparte) : 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. i, 1758, p. 125. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon ; 
San Pedro (Palos Verdes sand), Los Angeles County, California. 

Genus CHENDYTES Miller 

Chendytcs L. H. Miller, Condor, vol. 27, No. 4, July 15, 1925, P- I45- Type, 
by monotypy, Chendytes lawi Miller. 

Chendytes lawi Miller 

Chendytes lazvi L. H. Miller, 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 Howard 

Chendytes milleri H. Howard, Condor, vol. 57, No. 3, May 25, 1955, p. 137, 
fig. I a, d, e, f, g, i, fig. 2 b, c, fig. 3. 

Early Pleistocene : San Nicolas Island, California. 

Subfamily OXYURINAE: Ruddy and Masked Ducks 

Genus OXYURA Bonaparte 

Oxyura Bonaparte, Ann. Lye. 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. i, pt. 2, 1789, 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 Wetmore, Journ. Pal., vol. 12, No. 3, May 1938, p. 280. Type, by 
original designation, Eonessa attaticula Wetmore. 



34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Eonessa anaticula Wetmose 

Eonessa anaticula Wetmore, Journ. Pal., vol. 12, No. 3, May 1938, p. 280, 
figs. 1-5. 

Eocene (Uinta C horizon) : My ton Pocket, Utah. 

Subfamily MERGINAE: Mergansers 

Genus LOPHODYTES Reichenbach 

Lophodytcs Reichenbach, Avium Syst. Nat., 1852 (1853), p. ix. Type, by 
original designation, Mergus cticiillatus Linnaeus. 

Lophodytes cucullatus (Linnaeus) : Hooded Merganser ^9 
Mergus cucullatus Linnaeus, Syst. Nat., ed. 10, vol. i, 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)^^ 

Qnerquedula floridana Siiufeldt, 9th Ann. Rep. Florida State Geol. Surv., 
1917, P- 36, pi. I, 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. i, 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. i, 1758, p. 129. 

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. i, 1758, p. 129. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon. 



3" 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. 

<o See Wetmore, Condor, vol. 57, No. 3, 1955, p. 189. 

•*i Formerly recorded as Mergus serrator; see Wetmore, Wilson Bull., 1948, 
p. 240. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 35 

Order FALCONIFORMES : Vultures, Hawks, and Falcons 

Suborder CATHARTAE : New World Vultures 

Superfamily NEOCATHARTOIDEA : Neocathartes 

Family NEOCATHARTIDAE : Neocathartes 

Genus NEOCATHARTES Wetmore 

Neocathartes Wetmore, Auk, vol. 67, No. 2, April 1950, p. 235. Type, by 
original designation, Eocathartes grallator Wetmore. 

Neocathartes grallator (Wetmore) 

Eocathartes grallator Wetmore, Ann. Carnegie Mus., vol. 30, May 24, 1944, 
p. 58, pis. i-s, figs. i-io. 

Upper Eocene (Upper Washakie beds) : Sand wash one-half mile 
north of Dobe Town Road crossing, Sweetwater County, Wyoming. 

Superfamily CATHARTOIDEA : New World Vultures 

Family CATHARTIDAE : New World Vultures 

Genus CATHARTES Illiger 

Cathartes Illiger, Prodromus, 181 1, p. 236. Type, by subsequent designation, 
Vultur aura Linnaeus (Vigors, 1825). 

Cathartes aura (Linnaeus) : Turkey Vulture ^2 

Vultur aura Linnaeus, Syst. Nat., ed. 10, vol. i, 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 Geoffrey 

Coragyps Geoffroy Ms in Le Maout, Hist. Nat. Oiseaux, 1853, p. 66. Type, 
by monotypy, Vultur urubu Vieillot = Vultur atratus Bechstein. 



*2 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. 

*3 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. I3I 

Coragyps atratus (Bechstein) : Black Vulture 

Vultur atratus Bechstein, in John Latham's allg. Ucbers Vogel, Bd. i, Anh., 
1793, p. 655. 

Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, and cavern deposits near Lecanto, Florida. Quaternary 
(probably Recent) : Rocky Arroyo, New Mexico. 

Corag3rps occidentalis (Miller)** 

Catharista occidenialis L. H. Miller, 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 Wetmore, 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 Wetmore, Proc. Colorado Mus. Nat. Hist., vol. 7, No. 2, 
July 15, 1927, p. 3, figs. 1-4. 

Lower Oligocene (Chadron formation) : Florsetail Creek, Weld 
County, Colorado. 

Genus PALAEOGYPS Wetmore 

Palaeogyps Wetmore, Proc. Colorado Mus. Nat. Hist., vol. 7, No. 2, July 
15, 1927, p. 5. Type, by monotypy, Palaeogyps prudronius 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. 

♦* Coragyps shastcnsis (Miller) is a synonym according to Miller, Condor, 
1941, pp. 140-141. 

*^ 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 37 

Genus GYMNOGYPS Lesson 

Gymnogyps Lesson, Echo du Monde Savant, ser. 2, vol. 6, Dec. 8, 1842, col. 
1037. Type, by monotypy, Vitltur californianus Shaw. 

Gymnogyps amplus Miller ^^ 

Gymnogyps amplus L. H. Miller, 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, Niievo 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. Miller and H. Howard, 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. Miller, Univ. California Publ., Bull. Dept, 
Geol., vol. 6, No. i, Nov. 28, 1910, p. 11, figs. 3a, ^b. 

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 Dum^ril 

Sarcoramphtis Dumeril, Z06I. Anal., 1806, p. 32. Type, by subsequent designa- 
tion, Vultur papa Linnaeus (Vigors, 1825). 

Sarcoramphus kernense (Miller) 

Vultur kernensis L. H. Miller, Condor, vol. 22, Mar. 18, 1931, p. 70, fig. 16, 

Pliocene: Pozo Creek, Kern River Divide, Kern County, about 9 
miles northeast of Bakersfield, California. 

Family TERATORNITHIDAE : Teratornithes 

Genus TERATORNIS Miller 

Teratornis, L. H. Miller, Univ. California Publ., Bull. Dept. Geol., vol. 5, 
No. 21, Sept. 10, 1909, p. 307. Type, by monotypy, Teratornis merriami 
Miller. 



*8 Fisher, Pacific Science, vol. i. 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. I3I 

Teratomis merriami Miller 

Teratornis merriami L. H. Miller, 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),*^ Los Angeles, McKittrick, 
Kern County, and Carpinteria, Santa Barbara County, California. 

Teratornis incredibilis Howard 

Teratornis incredibilis Howard, Bull. Southern California Acad. Sci., vol. 51, 
pt. 2, 1952, p. 51, pi. 10, figs. 1-2. 

Quaternary (probably late Pleistocene) : Smith Creek Cave, 34 
miles north of Baker, White Pine County, Nevada. 

Genus CATHARTORNIS Miller ^s 

Cathartornis L. H. AIiller, Univ. California Publ., Bull. Dept. Geol., vol. 6, 
No. I, Nov. 28, 1910, p. 14. Type, by monotypy, Cathartornis gracilis 
Miller. 

Cathartornis gracilis Miller 

Cathartornis gracilis L. H. Miller, Univ. California Publ., Bull. Dept. Geol., 
vol. 6, No. I, Nov. 28, 1 910, p. 14, figs. 40, 4b. 

Late Pleistocene : Rancho La Brea, Los Angeles, California. 

Suborder FALCONES: Secretary-birds, Hawks, and Falcons 
Superfamily FALCONOIDEA: Hawks, Falcons, and Allies 

Family ACCIPITRIDAE: Hawks, Old World Vultures, and 

Harriers 

Subfamily AEGYPHNAE : Old World Vultures 

Genus PALAEOBORUS Coues 

Palaeoborus Coues, Key North Amer. Birds, ed. 2, 1884, p. 822. Type, by 
original designation, Catharlcs ximbrosus Cope. 

Palaeoborus umbrosus (Cope)4o 

Cathartes umbrosus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 26, Oct. 20, 

1874, p. 151- 
Pliocene : North of Pojauque, New Mexico. 



4^ 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)- 

43 Allocated 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. 

4» 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 hoivardae Wetmore, 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. Miller 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. Miller, Univ. California Publ., Bull. Dept. Geol., vol. 9, No. 9, 
Mar. 10, 1916, p. 108. Type, by monotypy, Neogyps errans Miller. 

Neogyps errans Miller 

Neogyps errans L. H. Miller, Univ. California Publ., Bull. Dept. Geol., vol. 9, 
No. 9, Mar. 10, 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. Miller, 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 atnericanus L. H. Miller, 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, P- 344, fig- I. 

Lower Pliocene : Big Spring Canyon, 15 miles southwest of Martin, 
Bennett County, South Dakota. 



''" 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). 

^1 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). 



40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Neophrontops vetustus Wetmore 

Ncophrontops 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: White-tailed Kites 

Genus ELANUS Savigny 

Elanus Savigny, Descr. figypte, vol. i, 1809, pp. 69, 97. Type, by monotypy, 
Elami-s caesius Savigny ;= Falco caeruleus Desfontaines. 

Elanus leucurus (Vieillot) : White-tailed Kite 

Milvus leucunis Vieillot, Nouv. Diet. Hist. Nat., nouv. ed., vol. 20, May 181 8, 

p. 563 [errore=:S56]. 

Modern form reported from Pleistocene: San Josecito Cave, 
Aramberri, Nuevo Leon. Late Pleistocene : Rancho La Brea, Los 
Angeles, California. 

Subfamily MILVINAE: True Kites 

Genus PROICTINIA Shufeldt 

Proictinia Shufeldt, Bull. Anicr. Mus. Nat. Hist., vol. 32, art. 16, Aug. 4, 
1913, P- 301. Type, by monotypy, Proictinia gilmorei Shufeldt. 

Proictinia effera Wetmore 

Proictinia effera Wetmore, 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, pi. 55, fig. 27. 

Lower Pliocene (Ogallala formation) : Long Island, Phillips 
County, Kansas. 

Subfamily ACCIPITRINAE: Bird Hawks 

Genus ACCIPITER Brisson 

Accipiter Brisson, Orn., 1760, vol. i, p. 28; vol. 6, p. 310. Type, by tautonymy, 
Accipiter Brisson = Fa/co nisus Linnaeus. 



NO, 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 4I 

Accipiter gentilis (Linnaeus) : Goshawk 

Falco gentilis Linnaeus, Syst. Nat, ed. 10, vol. i, 1758, p. 89. 

Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, and Rancho La Brea, Los Angeles, California. 

Accipiter striatus velox (Wilson) : Sharp-shinned Hawk 
Falco velox Wilson, Amer. Orn., vol. s, 1812, p. 116, pi. 45, fig. i. 

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. i, pi. 10, fig. i. 

Modern form reported from late Pleistocene : McKittrick, Kern 
County, Carpinteria, Santa Barbara County, and Rancho La Brea, 
Los Angeles, California. 

Subfamily BUTEONINAE: Buzzards and Eagles 

Genus BUTEO Lac^pMe 

Butco Lacepede, Tabl. Ois., 1799, p. 4. Type, by tautonymy, Falco bnlco Lin- 
naeus. 

Buteo jamaicensis (Gmelin) : Red-tailed Hawk 

Falco jamaicensis Gmelin, Syst. Nat., vol. i, pt. i, 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. i, pt. i, 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 (Vieillot) : Broad-winged Hawk 

Sparvius platypterus Vieillot, Tabl. Encycl. Meth. 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 
Bntco szvai)isoiii 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 (Pontoppidan) : Rough-legged Hawk 
Falco lagopus Pontoppidan, 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. i, pt. i, May 1844, pi. 6. 

Modern form reported from late Pleistocene: Hawver Cave, El- 
dorado County, Rancho La Brea, Los Angeles, Carpinteria, Santa 
Barbara County, and McKittrick, Kern County, California. 

Buteo fuscescens (Vieillot) : Buzzard Eagle 

Spisa'etus fuscescens Vieillot, Nouv. Diet. Hist. Nat., nouv. ed., vol. 32, Sep- 
tember 1819, p. 55. 

Modern form " reported from late Pleistocene : Banos de Ciego 
Montero, Santa Clara Province, Cuba. 

Buteo antecursor Wetmore 

Buteo antecursor Wetmore, Bull. Mus. Comp. Zool., vol. 75, October 1933, 
p. 298, figs. 1-5. 

Oligocene (Brule formation) : Near Torrington, Goshen County, 
Wyoming. 

Buteo granger! Wetmore and Case 

Buteo grangeri Wetmore and Case, Contr. Mus. Pal. Univ. Michigan, vol. 4, 
No. 8, Jan. 15, 1934, p. 129, i pi. 

Middle Oligocene (Brule formation, Oreodon beds) : Big Badlands 
of Pass Creek, Washabaugh County, South Dakota. 

Buteo fluviaticus Miller and Sibley 

Buteo fluviaticus A. H. Miller and C. G. Sibley, Condor, vol. 44, No. i, 
Jan. IS, 1942, p. 39, fig. 12. 

Middle Oligocene (Brule formation, Oreodon beds) : Owl Creek, 
6 miles east of Carr, Weld County, Colorado. 

°2 Formerly called Buteo melanoleucu^ (Vieillot). The modern range extends 
from the mountains of Venezuela and Colombia, south through Ecuador and 
Peru to Chile, and from southeastern Brazil and Paraguay to Tierra del Fuego. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 43 

Buteo typhoius Wetmore 

Biiteo 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) 

Gcranoa'ctus ales Wetmore, Ann. Carnegie Mus., vol. 16, No. 4, Apr. 10, 1926, 
p. 403, pi. 38, figs. 1-5. 

Lower Miocene (Lower Harrison beds) : Quarry No. 2, Agate 
Springs Fossil Quarries, Sioux County, Nebraska. 

Buteo contortus (Wetmore) 

Geranoactus contortus Wetmore, 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 Marsh, Amer. Journ. Sci., ser. 3, vol. 2, August 1871, p. 125. 

Lower Pliocene (Upper Snake Creek beds) : Loup Fork River, 
Nebraska. 

Buteo conterminus (Wetmore) 

Geranoactus conterminus Wetmore, 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 

Parahutco Ridgway, 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, pi. 313. 

Modern form reported from Pleistocene: San Josecito Cave, 
Aramberri, Nuevo Leon, 



^3 Possibly early Pliocene ; cf. Cook, H. J., and Cook, M. C, Nebraska Geol. 
Surv., Paper No. 5, I933, P- 42. 



44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Genus CALOHIERAX Wetmore 

Calohlcrax Wetmore, Bull. Mus. Comp. Z06I., vol. 80, No. 12, October 1937, 
p. 428. Type, by original designation, Calohierax quadrattis Wetmore. 

Calohierax quadratus Wetmore 

Calohierax quadratus Wetmore, Bull. Mus. Comp. Z06I., vol. 80, No. 12, 
October 1937, p. 429, figs. 1-3. 

Recent (extinct) : °* Cave deposits on Great Exuma Island, Ba- 
hama Islands. 

Genus MIOHIERAX Howard 

Miohierax Howard, Condor, vol. 46, No. 5, Sept. 27, 1944, p. 236. Type, by 
original designation, Miohierax stocki Howard. 

Miohierax stocki Howard 

Miohierax stocki Howard, 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 " 

Hypomorphnus Cabanis, Arch. Naturg., vol. 10, Bd. i, 1844, p. 263. Type, by 
original designation, Falco urubitinga Linnaeus, 

Hypomorphnus enectus (Wetmore) 

Urubitinga cnccta 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 (SnuFELDT)f'^ 
Aquila sodalis Shufeldt, Amer. Nat., vol. 25, No. 297, September 1891, p. 821. 

Late Pleistocene: Fossil Lake, Oregon. 

Genus TITANOHIERAX Wetmore 

Tilanohicrax Wetmore, Bull. Mus. Comp. Zool., vol. 80, No. 12, October 
I937i P- 430. Type, by original designation, Tilanohicrax gloveralleni 
Wetmore. 



** Included here since it has not been found in living form, being known only 
from its bones. 

^* For the use of Hypomorphnus to replace Urubitinga see Peters, Check-list 
of the birds of tiie world, vol. i, 1931, p. 244. 

''^ 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 

Titanohicrax gloveralleni Wetmore, Bull. Mus. Comp. Zool., vol. 80, No. 12, 
October 1937, p. 431, figs. 4-9. 

Recent (extinct) : " Cave deposits on Great Exuma Island, Bahama 
Islands. 

Genus BUTEOGALLUS Lesson 

Buteogallns Lesson, Traile d'Orn., Hvr. 2, 1830, p. 83. Type, by monotypy, 
Buteogallns cathartoides Lesson = Fa/co aequinoctialis Gmelin. 

Buteogallus milleri (Howard) ^^ 

Uruhitinga milleri Howard, Carnegie Inst. Washington Publ. 429, October 
1932, p. 25, pi. 2, figs. 3-3a, pi. 3, fig. 2. 

Late Pleistocene : Hawver Cave, Eldorado County, California. 

Buteogallus fragilis (Miller) ^^ 

Gemnoaetiis fragilis L. H. Miller, 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. Miller, Condor, vol. 30, No. 4, July 16, 1928, p. 255. 
Type, by original designation, Morphnus daggetti Miller. 

Wetmoregyps daggetti (Miller) 
Morphnus daggetti L. H. Miller, 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, Diet. Sci. Nat., vol. i, Suppl., October 1816, p. 88. Type, 
by subsequent designation, Falco giiianensis Daudin (Chubb, 1916). 



^^ Included here since it has not been found in living form, being known only 
from its bones. 

^8 Referred to this genus by Howard, Carnegie Inst. Washington Publ. 551, 
Jan. 25, 1946, p. 177. 

^^ Recorded fflso 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). 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. 



46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Morphnus woodward! Miller 

Morphniis ivoodivardi L. H. Miller, Univ. California Publ., Bull. Dept. Gcol., 
vol. 6, No. 12, Oct. 9, 1911, p. 312, figs. 3a, 3b. 

Late Pleistocene : Rancho La Brea, Los Angeles, California.^*' 

Genus SPIZAfiTUS Vieillot 

Spisactus Vieillot, Analyse, 181 6, p. 24. Type, by subsequent designation, 
Falco oniatus Daudin (Gray, 1840). 

Spizaetus grinnelli (Miller) ^i 

Geranoa'ettis grinnelli L. H. Millek, Univ. California Publ., Bull. Dcpt. Geol., 
vol. 6, No. 12, Oct. 9, 191 1, 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. 

Spizaetus willetti Howard 
Spizaetus willetti Howard, Condor, vol. Z7, 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. 

Spizaetus pliogryps (Siiufeldt) 

Aqiiila pliogryps Shufeldt, Amer. Nat., vol. 25, No. 297, September 1891, 
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 ataznis 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. 



<"> 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 Spicactus by Howard, Carnegie Inst. Washington Publ. 429, 
1932, pp. 33-44- 

62 Placed in Spisactus 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 ^s 

Aquila Brisson, Orn., 1760, vol. i, pp. 28, 419. Type, by tautonymy, Aquila 
Brisson = Falco chrysa'ctus Linnaeus. 

Aquila chrysaetos (Linnaeus) : Golden Eagle 
Falco Chrysaetos Linnaeus, Syst. Nat., ed. 10, vol. i, 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 

Haliacetus Savigny, Descr. figypte, Ois., vol. i, 1809, pp. 68, 85. Type, by 
monotypy, Haliacetus nisus Savigny = Falco albicilla Linnaeus. 

Haliaeetus leucocephalus (Linnaeus) : Bald Eagle 
Falco leucocephalus Linnaeus, Syst. Nat., ed. 12, vol. i, 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 : Palaeoplancus 

Genus PALAEOPLANCUS Wetmore 

Palaeoplancus Wetmore, Smithsonian Misc. Coll., vol. 87, No. 19, Dec. 26, 
1933. P- I- Type, by original designation, Palaeoplancus st ember gi 
Wetmore. 

Palaeoplancus sternbergi Wetmore 

Palaeoplancus sternbergi Wetmore, Smithsonian Misc. Coll., vol. 87, No. 19, 
Dec. 26, 1933, p. 12, figs. 1-19. 

Middle Oligocene (Brule formation, Upper Oreodon beds) : East 
side of Plum Creek, Niobrara County, Wyoming. 

^3 Aquila fero.r Shufeldt proves to be a mammal. See Wetmore, Amer. Mus. 
Nov., No. 680, Dec. 4, 1933, pp. 1-2. 

"* Howard, Auk, vol. 64, April 1947, pp. 287-291, 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 Lac^p^de 

Circus Lacepede, Tabl. Ois., 1799, p. 4. Type, by subsequent designation, Falco 
acruginosus Linnaeus (Lesson, 1828). 

Circus cyaneus (Linnaeus) : Marsh Hawk 
Falco cyaneus Linnaeus, Syst. Nat., ed. 12, vol. i, 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 Savigny, Descr. figypte, Ois., vol. i, 1809, pp. 69, 96. Type, by 
monotypy, Pandion fluznalis Savigny ^ Falco haliaetus Linnaeus. 

Pandion haliaetus Linnaeus : Osprey 
Falco Haliaetus Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 91. 

Modern form reported from Pleistocene: Melbourne (stratum 2), 
and Itchtucknee River, Florida. 



Family FALCONIDAE: Caracaras and Falcons 

Subfamily CARACARINAE: Caracaras 

Genus CARACARA Merrem 

Caracara Merrem, 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 (Howard) 

Polybonts prelutosus Howard, Carnegie Inst. Washington Publ. 487, July 7, 
1938, p. 226, pis. 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.*'^ 



^'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 (Howard) 

Polyboriis prelutosus grinnelli Howard, Condor, vol. 42, No. i, 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, pi. 2, figs. 5, 6. 

Recent (extinct) : ®® Cave deposits in Cueva Torano, near Utuado, 
Puerto Rico. 

Subfamily FALCONINAE: Falcons 

Genus FALCO Linnaeus 

Falco Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 88. Type, by subsequent 
designation, Falco subbuteo Linnaeus (A. O. U. Comm., 1886). 

Subgenus HIEROFALCO Cuvier 

Hierofalco Cuvier, Regne Animal, vol. i, 1817 (Dec. 7, 1816), p. 312. Type, 
by monotypy, Falco candicans Gmelin. 

Falco mexicanus Schlegel: Prairie Falcon 
Falco mexicanus Schlegel, Abh. Geb. Zool. 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 su'arthi L. H. Miller, Condor, vol. 29, No. 3, May 15, 1927, p. 152, 
fig. 54- 

Late Pleistocene: McKittrick, California. 
Falco oregonus Howard 

Falco oregonus H. Howard, Carnegie List. Washington Publ. 551, Jan. 25, 
1946, p. 178, pi. I, figs. 2, 3. 

Late Pleistocene : Fossil Lake, Oregon. 

Subgenus RHYNCHODON Nitzsch 

Rhynchodon Nitzsch, Obs. Avium Art. Carot. Comm., 1829, p. 20. Type, by 
subsequent designation, Falco peregrinus Tunstall (A. O. U. Comm., 
1886). 



•^^ Licluded here since it has not been found in living form, being known only 
from bones. 



50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Falco peregrinus Tunstall: Peregrine Falcon 
Falco Peregrinus Tunstall, Orn. Brit., 1771, p. 1. 

Modern form reported from Late Pleistocene : Potter Creek Cave, 
Shasta County, McKittrick, and Rancho La Brea, Los Angeles, 
California. 

Subgenus TINNUNCULUS Vieillot 

Tinmincnhis Vieillot, Ois. Amer. Sept., vol. i, 1807, p. 39. Type, by subse- 
quent designation, Falco columbarhis Linnaeus (Walden, 1872). 

Falco columbarius Linnaeus : Pigeon Hawk 

Falco columbarius Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 90. 

Modern form reported from late Pleistocene: McKittrick, and 
Rancho La Brea, Los Angeles, California. 

Falco ramenta Wetmore 

Falco ramenta Wetmore, Proc. U. S. Nat. Mus., vol. 84, Nov. 3, 1936, p. 75, 
fig. 14. 

Miocene (Sheep Creek formation) : Dawes County, Nebraska. 

Subgenus CERCHNEIS Bole 

Cerchncis Boie, 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. i, 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: Megapodes, Curassows, Pheasants, 
and HoATZiNS 

Suborder GALLI : Megapodes, Curassows, Grouse, and Pheasants 

Superfamily CRACOIDEA : Megapodes, Curassows, and Guans 

Family GALLINULOIDIDAE: Gallinuloides 

Genus GALLINULOIDES Eastman 

Gallinuloides Eastman, Geol. Mag., February 1900, p. 54. Type, by mono- 
typy, Gallinuloides wyomingcnsis Eastman. 



NO, 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 5 1 

Gallinuloides wyomingensis Eastman 

Gallinuloides wyomingetisis Eastman, Geol. Mag., n. s., vol. 7, pt. 4, No. 2, 
February 1900, p. 54, pi. 4. 

Middle Eocene (Green River formation): Fossil (type locality), 
and Henry's Fork, Wyoming, 

Family CRACIDAE: Curassows, Guans, and Chaciialacas 

Genus ORTALIS Merrem 

Ortalida (accusative case) = Ortalis (nominative) Merrem, Avium Rar. 
Icones et Descrip., vol. 2, 1786, p. 40. Type, by original designation, 
Phasiamis motmot Linnaeus. 

Ortalis phengites Wetmore 

Ortalis phengites Wetmore, Bull. Amer. Mus, Nat. Hist., vol, 48, art. 12, 
Dec. 3, 1923, p. 487, figs, 1-2. 

Lov^er Pliocene (Upper Snake Creek beds) : South of Agate, Sioux 
County, Nebraska, 

Ortalis tantala Wetmore 

Ortalis tantala Wetmore, Condor, vol, 35, No. 2, Mar. 15, 1933, p. 64, figs. 
10-14, 

Lower Miocene (Lower Harrison beds) : Carnegie Hill, Sioux 
County, Nebraska. 

Ortalis poUicaris MnxER 

Ortalis pollicaris A, H, Miller, 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 Brodkorb, Wilson Bull., vol. 66, No. 3, September (Oct. 29), 
1954. P- i8o- Type, by original designation, Boreortalis laesslci Brodkorb. 

Boreortalis laesslei Brodkorb 

Boreortalis laesslei Brodkorb, Wilson Bull., vol. 66, No. 3, September (Oct. 
29), 1954, P- 182, fig, I (on p. 181). 

Lower Miocene (Hawthorn formation) : Thomas Farm, 8 miles 
north of Bell, Gilchrist County, Florida. 



52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Superfamily PHASIANOIDEA : Grouse, Quails, Pheasants, and Turkeys 

Family TETRAONIDAE: Grouse and Ptarmigans 

Genus DENDRAGAPUS Elliot 

Dendragaptis Elliot, Proc. Acad. Nat. Sci. Philadelphia, vol. 16, No. 1, 
January-February (April 22,), 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 (SnuFELDT)^^ 
Pedioccctes lucasi Siiufeldt, Auk, vol. 8, No. 4, October 1891, p. 367. 
Late Pleistocene : Fossil Lake, Oregon. 

Dendragapus nanus (Siiufeldt) ^''^ 
Pedioccctes nanus Shufeldt, Amer. Nat., vol. 25, No. 297, September 1891, 
p. 821. 

Late Pleistocene : Fossil Lake, Oregon. 

Genus BONASA Stephens 

Bonasa Stephens, in Shaw, Gen. Zool., vol. 9, pt. 2, 1819, p. 298. 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. i, 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 c^ 

Tytnpanuchus Gloger, Hand- und Hilfsbuch Naturg., 1842 (pp. 1-450, 1841), 
p. 396. Type, by monotypy, Tetrao cupido Linnaeus. 



6T 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, 191 3, 
p. 299, pi. 55, figs. 18-20, pi. 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 Tympamichus 
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 luUi Shufeldt 

Tympamichus lulli Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, 
February 1915, p. 69, pi. 12, fig. 90. 

? Pleistocene : '^° Hornerstown, New Jersey, 

Tympanuchus stirtoni Miller 

Tympamichus stirtoni A. H. Miller, Univ. California Pub!., 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 Baird, Rep. Expl. and Surv. R. R. Pac, vol. 9, 1858, pp. xxi, xliv. 
Type, by monotypy, Tetrao phasianellns Linnaeus. 

Pedioecetes phasianellus (Linnaeus) : Sharp-tailed Grouse 
Tetrao Phasianellus Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 160. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon. 

Genus CENTROCERCUS Swainson 

Centrocerais Swainson, in Swainson and Richardson, Fauna Bor.-Amer., 
vol. 2, 1831 (1832), pp. 358, 496. 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 Shufeldt, Amer. Nat., vol. 25, No. 297, September 1891, p. 821, 
Type, by monotypy, Palaeotetrix cjilli Shufeldt. 



^° Cited in the original description as "Post-Pliocene.' 



54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Palaeotetrix gilli Shufeldt 

Palaeotctrix gilli Shufeldt, Amer. Nat., vol. 25, No. 297, September 1891, 
p. 821. 

Late Pleistocene : Fossil Lake, Oregon. 

Genus PALAEOPHASIANUS Shufeldt 

Palaeophasiamis Shufeldt, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 16, 
Aug. 4, 1913, p. 291. Type, by monotypy, Palaeophasiamis meleagroides 
Shufeldt. 

Palaeophasianus meleagroides Siiuff.ldt 

Palaeophasiamis meleagroides Shufeldt, Bull. Amer. Mus. Nat. Hist., vol. 
32, art. 16, Aug. 4, 1913, p. 291, pi. 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: Quails, Pheasants, and Peacocks 
Subfamily ODONTOPHORINAE : American Quails 

Genus COLINUS Goldfuss 

Colimis Goldfuss, Handb. Zool., vol. 2, 1820, p. 220. Type, by monotypy, 
Perdix mcxicamis, Caille de la Louisiane, PI. Enl. 149= Tetrao virgini- 
anus Linnaeus. 

Colinus virginianus (Linnaeus) : Bobwhite 

Tetrao virginianus Linnaeus, Syst. Nat., ed. 10, vol. i, 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. i. No. 9, 
May 15, 1944. P- 96, figs. 4-8. 

Upper Pliocene (Rexroad fauna) : Meade County, Kansas, 

? Colinus eatoni Shufeldt ''^ 

Colimis eatoni Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, Febru- 
ary 1915, p. 70, pi. I3> fig- 103. 

Geologic age uncertain : Western Kansas, 



^1 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 califoniicits Shaw (Gray, 1840). 

Lophortyx calif ornicus (Shaw) : California Quail 

Tetrao califoniicus Shaw, in Sliaw and Nodder, Nat. Misc. vol. 9, 1798, text 
to pi. 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 Baird, Rep. Expl. and Surv. R. R. Pac, vol. 9, 1858, pp. xxi, xlv, 
638, 642. Type, by original designation, Ortyx picta Douglas. 

Oreortyx pictus (Douglas) : Mountain Quail 
Ortyx picta Douglas, Trans. Linn. Soc. London, vol. 16, pt. i, 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. Miller, 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. Miller, 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 Gould, Monogr. Odontophoridae, pt. i, 1844, pi. and text. Type, by 
monotypy, Ortyx massena Lesson ^ Ortyx montesumae Vigors. 

Cyrtonyx montezumae (Vigors) : Harlequin Quail 

Ortyx Montesumae Vigors, Zool. Journ., vol. 5, June 1830, p. 275. 

Modern form reported from Pleistocene : San Josecito Cave, Aram- 
berri, Nuevo Leon. 



56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Cyrtonyx cooki Wetmore 

Cyrtonyx cooki Wetmore, Condor, vol. 36, No. i, Jan. 15, 1934, p. 30, fig. 5. 

Upper Miocene (Upper Sheep Creek beds) : 17 miles south of 
Agate, Sioux County, Nebraska. 

Cyrtonyx tedfordi Miller ''^ 

Cyrtonyx tedfordi L. H. Miller, Condor, vol. 54, No. 5, Sept. 22, 1952, p. 298, 
fig. 2. 

Upper Miocene (Barstow formation) : Lake bed horizon, near 
Barstow, California. 

Subfamily PHASIANINAE: Old World Partridges and 
Pheasants 

Genus PHASIANUS Linnaeus 

Phasianus Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 158. Type, by tau- 
tonymy, Phasianus colchicus Linnaeus. 

Phasianus alfhildae Siiufeldt^^ 

Phasianus alfhildae Siiufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, 
February 191S, p. 7i- 

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)^* 

Phasianus mioceanus Siiufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, 
February 1915, p. 60, pi. 13, figs. 94, 96. 

Miocene : Chimney Rock and Scott's Bluff, Nebraska. 

■^2 Allocation in this genus tentative. 

^3 Allocation of this species to the Old World genus Phasianus follows the 
usage of the original describer, and is subject to verification. 

■f* Given as "Parilina" in the original place of publication, through an error 
in reading the field label. 

''^ Described from fragmentar>' 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 

Mcleagris Linnaeus, Syst. Nat., ed. lo, vol. i, 1758, p. 156. Type, by tau- 
tonymy, Meleagris gallopavo Linnaeus. 

Meleagris gallopavo Linnaeus : Turkey "^^ 
Meleagris Gallopavo Linnaeus, Syst. Nat., ed. 10, vol. i, 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 Marsh 

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 Marsh, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, p. 261. 

Pleistocene : Monmouth County, New Jersey. 

Meleagris richmondi Siiufeldt 

Meleagris richmondi Siiufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, 
February 1915, p. 67, pi. 2, fig. 19. 

Pleistocene: Near Mission San Jose, Alameda County, California. 

Meleagris superba Cope 

Meleagris superbus Cope, Trans. Amer. Philos. Soc, n.s., vol. 14, pt. i, 
December 1870, p. 239. 

Pleistocene: Monmouth County (type locality), and Manalapan,^^ 
New Jersey ; Frankstown and Port Kennedy caves, Pennsylvania. 

''^ Ardea sellardsi Shufeldt, 9th Ann. Rep. Florida State Geol. Surv., 1917, 
p. 38, pi. 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, 
pp. 10- 1 1, 32-33. Tiie deposit is now considered to be of Recent age. See Cooke, 
Florida Geol. Surv. Geol. Bull. 29, 1945, pp. 306-307. 

'^'^ Possibly Upper Pliocene. 

''^ Type locality of Meleagris alius Marsh, Amer. Journ. Sci., ser. 3, vol. 4, 
1872, p. 260, which is a synonym. 



58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Meleagris tridens Wetmore 
Meleagris tridetus Wetmore, 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. Miller, 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. Miller, Univ. California Publ., Bull. Dept. Geol., vol. 9, 
No. 9, Mar. 10, 1916, p. 96. Type, by monotypy, Pavo calijornicus Miller. 

Parapavo calif omicus (Miller) 

Pavo californicus L. H. Miller, Univ. California Publ., Bull. Dept. Geol., 
vol. 5, No. 19, Aug. 14, 1909, 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 ),"'•' Los Angeles, California. 

Order GRUIFORMES : Cranes, Rails, and Allies 
Suborder GRUES : Cranes, Limpkins, Trumpeters, and Rails 
Superfamily GRUOIDEA : Cranes, Limpkins, and Trumpeters 

Family GERANOIDIDAE : Geranoides 
Genus GERANOIDES Wetmore 

Geranoides Wetmore, Condor, vol. 35, No. 3, May 15, 1933, p. 11 5- Type, 
by original designation, Geranoides jepseni Wetmore. 

Geranoides jepseni Wetmore 

Geranoides jepseni Wetmore, Condor, vol. 35, No. 3, May 15, 1933, p. 115. 
fig. 22. 

Lower Eocene (Gray Bull member) : South Elk Creek, Bighorn 
County, Wyoming. 

'" 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 
oklahomacnsis 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 so 

Alctoniis Marsh, Amer. Journ. Sci., scr. 3, vol. 14, October 1872, p. 256. 
Type, by subsequent designation, Aletornis nobilis Marsh (Hay, 1902). 

Aletomis bellus Marsh si 

Aletornis bellus Marsh, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, 
p. 258. 

Eocene (Bridger formation) : Grizzly Buttes, Wyoming. 

Aletornis gracilis Marsh ^^ 

Aletornis gracilis Marsh, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, 
p. 258. 

Eocene (Bridger formation) : Henry's Fork, Wyoming. 
Aletornis nobilis Marsh ^2 
Aletornis nobilis Marsh, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, p. 256. 
Eocene (Bridger formation) : Grizzly Buttes, Wyoming. 

Aletornis pemix Marsh 
Aletornis pemix Marsh, 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. 



s" Allocation in the subfamily Gruinae provisional. 

51 Considered by Shufcldt, Trans. Connecticut Acad. Arts Sci., vol. 19, Febru- 
ary 191 5, pp. 32, 76, as possibly a species of Scolopacidae. 

52 Marsh in his original proiwsal of the genus Aletornis included in it five 
si)ecies without selecting a type. From the five in question Hay, U. S. Geol. 
Surv., Bull. 179, 1902, p. 527, designated Aletomis 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. yy) Grtis 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 Aletomis. 



60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Fulicaletornis venustus (Marsh) ^^ 

Aletornis venustus Marsh, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, 
p. 257- 

Eocene (Bridger formation) : Henry's Fork, Wyoming. 

Genus PARAGRUS Lambrecht 

Paragrus Lambrecht, Handb. Palaeorn., 1933, p. 520. Type, by monotypy, 
Gallinuloides prentici Lootnis. 

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, lo miles west of Otto, 
Wyoming. 

Genus GRUS Pallas 

Grus Pallas, Misc. Zool., 1766, p. 66. Type, by tautonymy, Ardca grtis Lin- 
naeus. 

Grus americana (Linnaeus) : Whooping Crane 
Ardca americana Linnaeus, Syst. Nat., cd. 10, vol. i, 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 s* 
Ardea canadcyisis Linnaeus, Syst. Nat., ed. 10, vol. i, 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, y6, placed this species in the genus 
Fulica, the principal basis for Lambrecht's action in proposing Fulicaletornis. 

^* 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 haydcni 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 6l 

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 
Gnts proavus Marsh, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, p. 261. 

Pleistocene: Monmouth County, New Jersey. 

Grus nannodes Wetmore and Martin 

Grus nannodes Wetmore and Martin, Condor, vol. 32, No. i, 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. Miller and C. G. Sibley, 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 ^s 

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 : Limpkins 
Genus ARAMUS Vieillot 

Aramus Vieillot, Analyse, 181 6, p. 58. Type, by monptypy, Courliri Buf- 
fon = Ardea scolopacea Gnielin. 

Aramus guarauna Linnaeus: Limpkin 
Scolopax Guarauna Linnaeus, Syst. Nat., cd. 12, vol. i, 1766, p. 242. 

Modern form reported from Pleistocene : Seminole Field, Pinellas 
County, and Itchtucknee River, Florida. 

Genua BADISTORNIS Wetmore 

Badistornis Wetmore, Journ. Morph., vol. 66, Jan. 2, 1940, p. 30. Type, by 
original designation, Badistornis aramus Wetmore. 



8^ Generic allocation doubtful. See footnote under Aletomis nobilis (p. 59). 



62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Badistornis aramus Wetmore 

Badistornis aramus Wetmore, Journ. Morph., vol. 66, Jan. 2, 1940, p. 30, 
figs. 7-10. 

OHgocene (Metamynodon zone, Brule formation) : 35 miles south- 
west of Scenic, South Dakota. 



Genus ARAMORNIS Wetmore 

Aramornis Wetmore, Amer. Mus. Nov., No. 211, Mar. ii, 1926, p. i. Type, 
by original designation, Aramornis longurio Wetmore. 

Aramornis longurio Wetmore 

Aramornis longurio Wetmore, 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. loi, No. 14, May 11, 1942, 
p. I. Type, by monotypy, Gnotornis araviiellus Wetmore. 

Gnotornis aramiellus Wetmore 

Gnotornis aramiellus Wetmore, Smithsonian Misc. Coll., vol. loi, No. 14, 
May II, 1942, p. I, figs. 1-4. 

Upper OHgocene (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: Rails 

Family RALLIDAE: Rails, Gallinules, and Coots 

Subfamily RALLINAE: Rails 

Genus TELMATORNIS Marsh s^ 

Telmatornis Marsh, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, p. 210. 
Type, by subsequent designation, Telmatornis priscus Marsh (Hay, 1902). 

Telmatornis affinis Marsu 

Telmatornis affinis Marsh, Amer. Journ. Sci., ser. 2, vol. 49, Alarch 1870, 
p. 211. 

Paleocene (Hornerstown marl) : Hornerstown, New Jersey. 



s'^ Allocation in the subfamily Rallinae provisional. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 63 

Telmatornis priscus Marsh 

Tchnatornis 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 1915, p. 27, pi. 13, fig. lOI. 

Paleocene (Hornerstown marl) : Hornerstown, New Jersey. 

Genus PALAEORALLUS Wetmore 

Palacorallus Wetmore, Condor, vol. 2>i> No. 3, May 15, 1931, p. 108. Type, 
by original designation, Palacorallus troxclli Wetmore. 

Palacorallus troxelli Wetmore 

Palacorallus troxclli Wetmore, Condor, vol. 2i2>, No. 3, Alay 15, 1931, p. 108, 
figs. 26-29. 

Lower Eocene (Wasatch formation) : Northwest of Little Tatman 
Mountain, near Btirlington, Wyoming. 

Genus CRECCOIDES Shufeldt 

Crcccoides Shufeldt, Proc. Amer. Pliilos. Soc, vol. 30, Apr. 14, 1892, p. 125. 
Type, by monotypy, Creccoidcs osbornii Shufeldt. 

Creccoides osbornii Shufeldt 

Creccoides osbornii Shufeldt, 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. Miller, Univ. California Publ. Z06I., vol. 47, Mar. 6, 1942, 
p. 43. Type, by monotypy, Epirallus natator Miller. 

Epirallus natator Miller 

Epirallus natator L. H. AIiller, Univ. California Publ. Zool., vol. 43, Mar. 6, 
1942, p. 43, fig. I a. 

Pleistocene: San Josecito Cave, Aramberri, Nuevo Leon. 

Genus RALLUS Linnaeus 

Rallus Linnaeus, Syst. Nat., cd. 10, vol. i, 1758, p. 153. Type, by subsequent 
designation, Rallus aquaticus Linnaeus (Fleming, 1821). 

Rallus elegans Audubon : King Rail 
Rallus elegans Audubon, Birds Amer. (folio), vol. 3, 1834, pi. 203. 



64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, and Itchtucknee River, Florida, 

Rallus longirostris Boddaert: Clapper Rail 

Rallus longirostris Boddaert, Table Planch. Enlum., 1783, p. 52. 

Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 

Rallus limicola Vieillot : Virginia Rail 

Rallus limicola Vieillot, Nouv. Diet. Hist. Nat., nouv. ed., vol. 28, May 1819, 
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. i, No. 9, 
May 15, 1944, p. 99, figs. 9-19. 

Upper Pliocene (Rexroad fauna) : Meade County, Kansas. 

Genus PORZANA Vieillot 

Porsana Vieillot, Analyse, 1816, p. 61. Type, by monotypy and tautonymy, 
Marouette Buff on = Rallus porsana Linnaeus. 

Porzana Carolina (Linnaeus) : Sora 
Rallus carolinus Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 153. 

Pleistocene : Near Reddick, Marion County, Florida. 

Porzana auffenbergi Brodkorb 

Porsana auffenbergi Brodkorb, 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 vielanophaius Vieillot. 

Laterallus guti Brodkorb 

Laterallus guti Brodkoru, Wilson Bull., vol. 64, No. 2, June 16, 1952, p. 80, 
fig. I. ' 

Pleistocene : i 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, Fulica 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 

Ncsotrochis Wetmore, Proc. U. S. Nat. Mus., vol. 54, Nov. 21, 1918, p. 516. 
Type, by original designation, Nesotrochis dcbooyi Wetmore. 

Nesotrochis debooyi Wetmore 

Nesotrochis debooyi Wetmore, Proc. U. S. Nat. Mus., vol. 54, Nov. 21, 1918, 
p. 516, pi. 82. 

Recent (extinct) :^^ Archeological sites on St. Thomas ^^ and St. 
Croix, Virgin Islands ; and at Barrio Canas, near Ponce ; cavern 
deposits in Cueva Clara and Cueva San Miguel, near Morovis ; Cueva 
Torano, and a cave on Hacienda Jobo, near Utuado, Puerto Rico. 

Subfamily GALLINULINAE: Gallinules 

Genus PORPHYRULA Blyth 

Porphyriila Blyth, Cat. Birds Mus. Asiat. Soc, 1849 (1852), p. 283. Type, 
by monotypy, P. chloronotus Blyth = Porphyria alleni Thomson. 

Porphyrula martinica (Linnaeus) : Purple Gallinule 
Fulica martinica Linnaeus, Syst. Nat., ed. 12, vol. i, 1766, p. 259. 

Modern form reported from Pleistocene : Haile, Alachua County, 
Florida. 

Genus GALLINULA Brisson 

GaUinula Brisson, Orn., 1760, vol. i, 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. i, 1758, p. 152. 

Modern form reported from Upper Pliocene (Hagerman lake 
beds) : Near Hagerman, Idaho. From Pleistocene : ''° Seminole Field, 

S8 Included here as it has not been found in living form, being known only 
from bones. Possibly the species lived until Spanish colonial times. 

^^ Type locality a kitchen midden at Magen's Bay, on the north coast of 
St. Thomas. 

^0 Reported from Pleistocene at Haile, Alachua County, Florida, on basis of a 



66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Pinellas County, and Itchtucknee River, Florida. Late Pleistocene : 
Banos de Ciego Montero, Cuba. 

Genus PALAEOCREX Wetmore 01 

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 prhmts 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: Coots 

Genus FULICA Linnaeus 

Fidica Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 152. Type, by subsequent 
designation, Fulica atra Linnaeus (Gray, 1840). 

Fulica americana Gmelin : American Coot 
Fulica americana Gmelin, Syst. Nat., vol. r, pt. 2, 1789, 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. i, March (Apr. 22), 

1953, P- 50. 
81 Subfamily allocation provisional. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 67 

Fulica minor Shufeldt''^ 
Fulica minor Siiufeldt, Amer. Nat., vol. 25, No. 297, September 1891, p. 820. 
Late Pleistocene: Fossil Lake, Oregon. 

Suborder CARIAMAE: Cariamas and Allies 

Family BATHORNITHIDAE : Bathornithes 

Genus BATHORNIS Wetmore 

Bathornis Wetmore, Proc. Colorado Mus. Nat. Hist., vol. 7, No. 2, July 15, 
1927, p. II. 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. II, 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 Wetmore, Bull. Mus. Comp. Zool., vol. 75, October 1933, 
p. 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. Zool., vol. 75, October 1933, 
p. 310, figs. IS-19. 

Upper Oligocene (Brule formation) : Near Torrington, Goshen 
County, W^yoming. 

Bathornis geographicus Wetmore 

Bathornis geographicus Wetmore, Smithsonian Misc. Coll., vol. loi, No. 14, 
May II, 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 : Diatrymas 

Family DIATRYMIDAE : Diatrymas 

Genus BARORNIS Marsh 

Barornis Marsh, Amer. Journ. Sci., ser. 3, vol. 48, 1894, p. 344. Type, by 
monotypy, Barornis regens Marsh. 

Barornis regens Marsh ^^ 

Barornis regens Marsh, Amer. Journ. Sci., ser. 3, vol. 48, October 1894, p. 344, 
text fig. 

Eocene: Squankum, Monmouth County, New Jersey. 

Genus DIATRYMA Cope 

Diatryma Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 28, sign. 2, April 18, 
1876, p. II. Type, by monotypy, Diatryma gigantea Cope. 

Diatryma ajax Shufeldt 

Diatryma ajax Shufeldt, Bull. Amer. Mus. Nat. Hist., vol. 2>^, art. 16, 
Aug. 4, 1913, p. 287, pi. 52, figs. 4-5, pi. 53, figs. 8-10, pi. 54, figs. 13-14. 

Lower Eocene (Wasatch formation) : 3 (type locahty) and 5 miles 
southeast of mouth of Pat O'Hara Creek, Clark's Fork Basin, Wy- 
oming. 

Diatryma giganteum Cope 

Diatryma gigantea Cope, 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 steini Matthew and Granger, Bull. Amer. Mus. Nat. Hist., vol. 
37, art. II, May 28, 1917, p. 322, pis. 20-33. 

Lower Eocene (Wasatch, Gray Bull member) : South Elk Creek, 
Bighorn Basin, Wyoming. 

Genus OMORHAMPHUS Sinclair 

Omorhamphus Sinclair, Proc. Amer. Philos. Soc, vol. 67, 1928, p. 51. Type, 
by monotypy, Omorhamphus storchii Sinclair. 



^3 Considered a species of Diatryma by Shufeldt, Trans. Connecticut Acad. 
Arts Sci., vol. 19, February 1915, pp. 37-38. 

^* 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 

Oinorhamphus storchii Sinclair, Proc. Amer. Philos. Soc, vol. 67, 1928, p. 52, 
pis. 1-2, figs. 1-3. 

Lower Eocene (Lower Gray Bull horizon, Lower Wasatch) : i^ 
miles southeast of Dorsey Creek, about 2 miles south of Otto-Basin 
Road, Big Horn County, Wyoming. 



Order CHARADRIIFORMES : Shorebirds, Gulls, and Auks 

Suborder CHARADRII : Shorebirds 

Superfamily CHARADRIOIDEA : Plovers, Sandpipers, and Allies 

Family RHEGMINORNITHIDAE: Rhegminornis 

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, pi. II, figs. 1-5. 

Lower Miocene (Tampa limestone) : °^ Thomas Farm, 8 miles north 
of Bell, Gilchrist County, Florida. 

Family HAEMATOPODIDAE : Oystercatchers 

Genus PARACTIORNIS Wetmore 

Paractiornis Wetmore, 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. 



^^ Cooke, Florida Geol. Surv., Geol. Bull. 29, 1945, pp. 1 19-120, 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. 



70 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Genus PALOSTRALEGUS Brodkorb 

Palostralcgus Brodkorb, Florida Geol. Surv. Rep. Invest. No. 14, November 
1955. P- 19- Type, by original designation, Palostralcgus sulcatiis Brod- 
korb. 

Palostralegus sulcatus Brodkorb 

Palostralcgus sulcatus Brodkorb, Florida Geol. Surv. Rep. Invest. No. 14, 
November 1955, p. 20, fig. 18. 

Pliocene (Bone Valley formation) : Near Brewster, Polk County, 
Florida. 



Family CHARADRIIDAE : Plovers, Turnstones, and Surfbirds 

Subfamily CHARADRIINAE: Plovers 

Genus CHARADRIUS Linnaeus 

Charadr'ms Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 150. Type, by tau- 
tonymy, Charadrius hiaticula Linnaeus. 

Charadrius sheppardianus Cope 

Charadrius sheppardianus Cope, Bull. Geol. Geogr. Surv. Terr., vol. 6, No. i, 
Feb. II, 1 881, p. 83. 

OHgocene (Florissant lake beds) : Florissant, Colorado."" 

Charadrius vocif erus Linnaeus : Killdeer 

Charadrius vocijerus Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 150. 

Modern form reported from late Pleistocene : McKittrick, Kern 
County, and Rancho La Brca, Los Angeles, California. 

Genus EUPODA Brandt 

Eiipoda J. F. Brandt, in Tchihatchev, Voy. Sci. Altai Orient., 1845, p. 444. 
Type, by monotypy, Charadrius asialicus Pallas. 

Eupoda montana (Townsend) : Mountain Plover 

Charadrius montatius J. K. Townsend, Journ. Acad. Nat. Sci. Philadelphia, 
vol. 7, pt. 2, Nov. 21, 1837, p. 192. 

Modern form reported from late Pleistocene: McKittrick, Kern 
County, California. 



8" 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 7I 

Genus SQUATAROLA Cuvier 

Squatarola Cuvier, Rcgne Animal, vol. i, 1817 (Dec. 7, 1816), p. 467. Type, 
by tautonyniy, Tringa squatarola Linnaeus. 

Squatarola squatarola (Linnaeus) : Black-bellied Plover 

Tringa Squatarola Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 149. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, California. 

Genus LIMICOLAVIS Shuf eldt ^^ 

Limicolavis Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, February 
iQiSi P- 55- Type, by monotypy, Limicolavis pluvianella Shufeldt. 

Limicolavis pluvianella Shufeldt 

Limicolavis pluvianella Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 
19, February 1915, p. 55, pl- I5, fig- 129. 

? Oligocene : Lower Willow Creek, Oregon. 



Family SCOLOPACIDAE : Woodcock, Snipes, and Sandpipers 

Subfamily PALAEOTRINGINAE : Palaeotringas 

Genus PALAEOTRINGA Marsh 

Palaeotringa Marsh, Anier. Journ. Sci., ser. 2, vol. 49, March 1870, p. 208. 
Type, by subsequent designation, Palaeotringa littoralis Marsh (Hay, 
1902). 

Palaeotringa littoralis Marsh ^^ 

Palaeotringa littoralis Marsh, Amer. Journ. Sci., ser. 2, vol. 49, Alarch 1870, 
p. 208. 

Paleocene (Hornerstown marl) : Hornerstown, New Jersey. 

Palaeotringa vagans Marsh 
Palaeotringa vagans Marsh, Amer. Journ. Sci., ser. 3, vol. 3, May 1872, p. 365. 

Paleocene (Hornerstown marl) : Hornerstown, New Jersey. 

Palaeotringa vetus Marsh 

Palaeotringa vetus Marsh, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, 
p. 209. 

Paleocene (Hornerstown marl) : Arneytown, New Jersey. 



^^ Family relationship uncertain. 

88 Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, February 1915, pp. 23, 
J7, pl. 6, fig. 35, believes this to be a gull, but this is open to question. 



72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Subfamily SCOLOPACINAE: Woodcock and Snipes 

Genus CAPELLA Frenzel 

Capella Frenzel, Beschr. Vogel und Eyer Wittenberg, 1801, p. 58. Type, by 
monotypy, Scolopax coelcstis Frenzel =^ Scolopax gallinago Linnaeus. 

Capella gallinago (Linnaeus): Common Snipe »® 
Scolopax GaUinago Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 147. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, California. 

Capella anthonyi (Wetmore) 

Gallinago anthonyi Wetmore, Proc. Biol. Soc. Washington, vol. 33, Dec. 30, 
1920, p. 78, pi. 2, figs. I, 2. 

Recent (extinct) : ^ Cave deposits in Cueva Catedral (type lo- 
cality) and Cueva Clara, near Morovis, Puerto Rico. 

Subfamily TRINGINAE: Curlews, Yellowlegs, and Allies 

Genus NUMENIUS Brisson 

Numenius Brisson, Orn., 1760, vol. i, p. 48; vol. 5, p. 311. Type, by tau- 
tonymy, Numenius Brisson = Scolopax arquata Linnaeus. 

Numenius americanus Beciistein: Long-billed Curlew 

Numenius americanus Bechstein, in Latham, AUgcm. Uebers. Vogel, vol. 4, 
pt. 2, 1812, p. 432. 

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. i, 1758, p. 146. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, California. 

09 Capella delicata (Ord), Wilson's snipe, of the previous list. 
1 Included here as it has not been found in living form, being known only 
from bones. 
^Phaeopus hudsonicus (Latham), Hudsonian curlew of the previous list. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 73 

Genus PALNUMENIUS Miller 

Pahmmcnhis L. Miller, Univ. California Publ. Zool., vol. 43, Mar. 6, 1942, 
p. 45. Type, by monotypy, Palmimenms victima Miller. 

Palnumenius victima Miller 

Palnumenius Tnctima L. Miller, Univ. California Publ. Zoo!., vol. 43, Mar. 6, 
1942, p. 45, fig. lb. 

Pleistocene : San Josecito Cave, Aramberri, Nuevo Leon. 



Genus BARTRAMIA Lesson 

Bartramia Lesson, Traite d'Orn., livr. 7, Apr. 9, 1831, p. 553. Type, by 
monotypy, Bartramia laticanda Lessons 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 Bechstein, Orn. Taschenb. Deutschland, vol. 2, 1803, p. 282. Type, 
by tautonymy, Totanus maculatus Bechstein ^ Scolopax totanus Linnaeus. 

Totanus melanoleucus (Gmelin) : Greater Yellowlegs 

Scolopax melanolcuca Gmelin, Syst. Nat., vol. i, pt. 2, 1789, p. 659. 

Modern form reported from Pleistocene: Fossil Lake, Oregon; 
Rancho La Brea, Los Angeles, and McKittrick, Kern County, Cali- 
fornia, 

Subfamily CALIDRIINAE : Sandpipers, Godwits, and Allies 

Genus CALIDRIS Merrem 

Calidris pads Merrem, Lit. Zeitung, vol. 2, No. 168, June 8, 1804, col. 542. 
Type, by tautonymy, Tringa calidris Gmelin = Tringa canntus Linnaeus. 

Calidris pacis Brodkorb 

Calidris pads Brodkorb, 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. I3I 

Genus EROLIA Vieillot 

Erolia Vieillot, Analyse, 1816, p. 55. Type, by monotypy, Erolia variegata 
Vieillot = Scolopax tcstacca Pallas. 

Erolia penepusilla Brodkorb 

Erolia penepusilla Brodkorb, 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. i, 1758, p. 149. 

Modern form reported from late Pleistocene: McKittrick, Kern 
County, California. 

Genus LIMNODROMUS Wied 

Lhnnodromus Wied, Beitr. Naturg. Brasil, vol. 4, Abt. 2, 1833, p. 716. Type, 
by monotypy, Scolopax noveboracctisis Gmelin = Scolopax grisea Gmelin. 

Limnodromus griseus (Gmelin) : Dowitcher 
Scolopax grisea Gmelin, Syst. Nat., vol. i, pt. 2, 1789, p. 658. 

Modern form reported late Pleistocene : McKittrick, Kern County, 
and Rancho La Brea, Los Angeles, California. 

Genus MICROPALAMA Baird 

Micropalama Baird, Rep. Expl. and Surv. R. R. Pac., vol. 9, 1858, pp. xxii, 
xlvii, 714, 726. Type, by monotypy, Tringa himantoptis Bonaparte. 

Micropalama hesternus Wetmore 

Micropalama hesternus Wetmore, Proc. U. S. Nat. Alus., vol. 64, art. 5, Jan. 
15, 1924, p. n, figs. 6-7. 

Upper Pliocene (lilancan) : 2 miles south of Benson, Arizona. 

Genus LIMOSA Brisson 

Limosa Brisson, Orn., 1760, vol. i, p. 48; vol. 5, p. 261. Type, by tautonymy, 
Limosa Brisson = Scolopax limosa Linnaeus. 

Limosa vanrossemi Miller 

Limosa varvrossemi L. H. Miller, Carnegie Inst. Washington Publ. 349, 
August 1925, p. 116, pi. 6. 

Middle Miocene (Temblor, Turritella ocoyana zone) : Lompoc, 
California. 



NO, 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 75 

Family RECURVIROSTRIDAE : Ibis-bills, Avocets, and Stilts 

Subfamily RECURVIROSTRINAE: Avocets and Stilts 

Genus RECURVIROSTRA Linnaeus 

Recurvirostra Linnaeus, Syst. Nat., ed. lo, vol. i, 1758, p. 151. Type, by 
monotypy, Recurvirostra avosetta Linnaeus. 

Recurvirostra americana Gmelin : Avocet 
Recurvirostra americana Gmelin, Syst. Nat., vol. i, pt. 2, 1789, p. 693. 

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. i, 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. Muller, Natursyst., Suppl., 1776, p. 117. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon. 

Family PRESBYORNITHIDAE : Presbyornithes 

Genus PRESBYORNIS Wetmore 

Presbyornis Wetmore, 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, pi. 37, figs. 10-20. 

Eocene (Lower Green River formation) : White River, Utah, 2 
miles from Colorado State line. 

Family PHALAROPODIDAE: Piialaropes 

Genus LOBIPES Cuvier 

Lobipes Cuvier, Regne Animal, vol. i, 1817 (Dec. 7, 1816), p. 495. Type, by 
original designation, Tringa hyperborea Linnaeus = rn'»<70 lobata 
Linnaeus. 

Lobipes lobatus (Linnaeus) : Northern Phalarope 

Tringa lobata Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 148. 

Modern form reported from late Pleistocene : Fossil I-ake, Oregon. 



76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Suborder LARI : Skuas, Gulls, Terns, and Skimmers 

Family STERCORARIIDAE: Jaegers and Skuas 

Genus STERCORARIUS Brisson 

Stcrcorarius Brisson, Orn., 1760, vol. i, p. 56; vol. 6, p. 149. Type, by tau- 
tonymy, Stercorarius Brisson = Larus parasiticus Linnaeus. 

Stercorarius shufeldti Howard 

Stcrcorarius shufeldti H. Howard, Carnegie Inst. Washington Publ. 551, 
Jan. 25, 1946, p. 184, pi. 2, figs. I, 2. 

Late Pleistocene : Fossil Lake, Oregon.' 

Family LARIDAE: Gulls and Terns 

Subfamily LARINAE: Gulls 

Genus LARUS Linnaeus '' 

Larus Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 136. Type, by subsequent 
designation, Lams 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 calif ornicus Lawrence : California Gull 
Larus Californicus Lawrence, Ann. Lye. Nat. Hist. New York, vol. 6, 1854, 

p. 79- 
Modern form reported from late Pleistocene : Fossil Lake, Oregon. 

Larus Philadelphia (Ord) : Bonaparte's Gull 
Sterna Philadelphia Ord, in Guthrie, Geogr., 2d Amer. ed., 1815, p. 319. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon.^ 

Larus oregonus Shufeldt 
Larus oregonus Shufeldt, Amer. Nat., vol. 25, No. 297, September 1891, 
p. 820. 

Late Pleistocene: Fossil Lake, Oregon. 



3 The type of Stcrcorarius shufeldti originally was identified by Shufeldt as 
Larus argentatus, this specimen being the basis for the record of the herring 
gull from Fossil Lake. 

•* Larus vcro Shufeldt, Journ. Geol., January-February 1917, p. 18, has been 
identified by Wetmore as Nycianassa znolacca Linnaeus (Smithsonian Misc. Coll., 
vol. 85, No. 2, Apr. 13, 1931, p. 16). 

5 Records of Xema sabiui from Fossil Lake, so far as identified, refer to Larus 
Philadelphia. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 'J'^ 

Larus pristinus Shufeldt^ 

Larus pristinus Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, Febru- 
ary 191S, p. 54, pi. 14, fig. 112. 

? OHgocene (John Day) : Willow Creek, Oregon. 

Larus robustus Shufeldt 

Larus robustus Shufeldt, Amer. Nat., vol. 25, No. 297, September 1891, 
p. 819. 

Late Pleistocene: Fossil Lake, Oregon. 

Larus elmorei Brodkorb 

Larus elmorei Brodkorb, 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 ^ 

Gavioia A. H. Miller and C. G. Sibley, 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. Miller and C. G. Sibley, Auk, vol. 58, No. 4, Octo- 
ber 1 941, p. 563, fig. I. 

Late Upper Miocene (Barstovian, Niobrara River zone) : Niobrara 
Game Preserve, Cherry County, Nebraska. 

Subfamily STERNLNAE: Terns 

Genus STERNA Linnaeus 

Sterna Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 137. Type, by tautonymy. 
Sterna hirundo Linnaeus. 

Subgenus STERNA Linnaeus 
Sterna forsteri Nuttall: Forster's Tern 
Sterna forsteri Nuttall, 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. i. No. 8, Feb. 21, 1822, 
p. 3, col. 5. Type, by monotypy. Sterna melanops Rafinesque = Sterna 
surinatnensis Gmelin. 



^ Generic assignation in original description tentative. 
^ Allocation to subfamily provisional. 



78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I3I 

Chlidonias niger (Linnaeus) : Black Tern 
Sterna nigra Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 137. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon, 

Suborder ALCAE: Auks 

Family ALCIDAE: Auks, Murres, and Puffins 

Subfamily NAUTILORNITHINAE : Nautilornithes 

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 Wetmore, Ann. Carnegie Mus., vol. 16, Apr. 10, 1926, p. 392, 
pi. 36, figs. 1-8. 

Eocene (Lower Green River formation) : White River, Utah, 2 
miles from Colorado State line. 

Nautilornis proavitus Wetmore 

Nautilornis proavitus Wetmore, Ann. Carnegie Mus., vol. 16, Apr. 10, 1926, 
P- 394, Pl- 36, fig. 9- 

Eocene (Lower Green River formation) : White River, Utah, 2 
miles from Colorado State Hne. 

Genus HYDROTHERIKORNIS Miller 

Hydrotherikornis A. H. Miller, 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. Miller, Univ. California Publ., Bull. Dept. 
Geol. Sci., vol. 20, No. 3, Apr. 21, 1931, p. 24, fig. i. 

Upper Eocene (Arago series) : Sunset Bay, near Coos Bay, Coos 
County, Oregon. 

Subfamily ALCINAE: Auks and Murres 

Genus AUSTRALCA Brodkorb 

Austratca Brodkokb, 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 Brodkorb 

Australca grandis Brodkorb, 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. i, p. 52; vol. 6, p. 70. Type, by tautonymy, 
Uria Brisson ^ Colymbus aalgc Pontoppidan. 

Uria aalge (Pontoppidan) : Common Murre 

Colymbus aalge Pontoppidan, Danske Atlas, vol. i, 1763, p. 621, pi. 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 mccltmgi Wetmore. 

Miocepphus mcclungi Wetmore 

Miocepphus mcclungi Wetmore, Journ. Morph., vol. 66, Jan. 2, 1940, p. 35, 
figs. 11-14. 

Miocene (Calvert formation, zone 12) : Near the mouth of Parker 
Creek, Calvert County, Maryland.^ 

Genus BRACHYRAMPHUS Brandt 

Brachyramphus M. Brandt, Bull. Sci. Acad. Imp. Sci. St.-Petersbourg, vol. 2, 
No. 22, Mar. 19, 1837, col. 346. Type, by subsequent designation, Colymbus 
marmoratus Gniclin (Gray, 1840). 

^ Two records. 



80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Brachyramphus pliocenum Howard 

Brachyramphus pliocenus Howard, Carnegie Inst. Washington Publ. 584, 
June 22, 1949, p. 191. 

Middle Pliocene (San Diego formation) : Washington Boulevard 
Freeway, San Diego, California. 

Genus SYNTHLIBORAMPHUS Brandt 

Synthlihoramphus M. Brandt, Bull. Sci. Acad. Imp. Sci. St.-Petersbourg, 
vol. 2, No. 22, Mar. 19, 1837, col. 347. Type, by subsequent designation, 
Alca antiqua Gmelin (Gray, 1840). 

Ssmthliboramphus antiquum (Gmelin) : Ancient Murrelet 

Alca antiqua Gmeun, Syst. Nat., vol. i, pt. 2, 1789, p. 554. 

Modern form reported from late Pleistocene (Palos Verdes sand) : 
San Pedro, California. 

Genus PTYCHORAMPHUS Brandt 

Ptychoramphiis M. Brandt, Bull. Sci. Acad. Imp. Sci. St.-Petersbourg, 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. Lye. Nat. Hist. New York, vol. 2, 1828, p. 427. 
Type, by monotypy, Cerorhinca occidentalis Bonaparte = ^/co monocerata 
Pallas. 

Cerorhinca dubia Miller 

Cerorhinca dubia L. H. Miller, Carnegie Inst. Washington Publ. 349, August 
1925, p. IIS, pi. 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 51 

Mancalla californiensis Lucas 
Mancalla californiensis Lucas, Science, n.s., vol. 13, Mar. 15, 1901, p. 428.^ 

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 diccjcnse L. H. Miller, Trans. San Diego Soc. Nat. Hist., vol. 8, 
Dec. IS, 1937, p. 2,7^, 2 figs. 

Middle Pliocene (San Diego formation) : Market Street, near 
Euclid Avenue (type locality), and Mission Hills district, San Diego, 
California. 

Order COLUMBIFORMES : Sand-grouse, Pigeons, and Do\'es 

Suborder COLUMBAE: Pigeons and Doves 

Family COLUMBIDAE : Pigeons and Doves 

Subfamily COLUMBINAE : Pigeons and Doves 

Genus COLUMBA Linnaeus 

Columha Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 162. Type, by subsequent 
designation, Columba ocnas Linnaeus (Vigors, 1825). 

Columba fasciata Say : Band-tailed Pigeon 

Columha 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) 

Chlorocnas miciila 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, Columha carolincnsis Linnaeus. 



8 See also Lucas, Proc. U. S. Nat. Mus., vol. 24, Sept. 27, 1901, pp. 133-134, 
figs. I, 2. 



82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Zenaidura macroura (Linnaeus) : Mourning Dove 
Colmnba macroura Linnaeus, Syst. Nat., ed. 10, vol. i, 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. Tj^pe, by subsequent designation, Columba migratoria Linnaeus 
(Swainson, 1837). 

Ectopistes migratorius (Linnaeus) : Passenger Pigeon 

Columba migratoria Linnaeus, Syst. Nat., ed. 12, vol. i, 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 Gosse, 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) 

Orcopclcia larva Wetmore, Proc. Biol. Soc. Washington, vol. ^2), Dec. 30, 1920, 
p. 79, pi. 3, figs. 1-2. 

Recent (extinct) : ^° Cave deposits in Cueva Clara (type locality) 
and Cueva Catedral, near Morovis ; Cueva Torano, near Utuado ; 
kitchen middens near Mayagiicz, and at Barrio Canas, near Ponce, 
Puerto Rico. 

Order PSITTACIFORMES : Lories, Parrots, Parakeets, and Macaws 

Family PSITTACIDAE: Lories, Parrots, and Macaws 

Subfamily PSITTACINAE: Parakeets and Macaws 

Genus ARA Lac^p^de 

Ara Lacepede, Tableaux Ois., 1799, p. i, Type, by subsequent designation, 
Psittacus macao Linnaeus (Ridgway, 1916). 



1° 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 Bech STEIN : Cuban Macaw 

Ara tricolor Bechstein, in Latham, Allg. Ucbcrs. Vog., vol. 4, Th. i, 181 1, 
p. 64, pi. I. (Cuba.) 

Modern form recorded from late Pleistocene : Baiios de Ciego 
Montero, Santa Clara Province, Cuba. 

Ara autocthones Wetmore 

Ara autoctJwncs Wetmore, Journ. Agr. Univ. Puerto Rico, vol. 21, No. i, 
January 1937, p. 12, pi. i, figs. 8, 9. 

Recent (extinct) : ^^ Prehistoric kitchen midden deposits at Con- 
cordia, near Southwest Cape, St. Croix, Virgin Islands. 

Genus RHYNCHOPSITTA Bonaparte 

Rhynchopsitfa Bonaparte, Rev. et Mag. Zool., ser. 2, vol. 6, March 1854, 
p. 149. Type, by monotypy, Macrocercus pachyrhynchns Swainson. 

Rhynchopsitta pachyrhyncha (Swainson) : Thick-billed Parrot 

Macrocercus pachyrhynchus Swainson, Philos. Mag., n.s., vol. i, No. 6, June 
1827, p. 439. 

Modern form reported from Pleistocene : San Josecito Cave, Aram- 
berri, Nuevo Leon. 

Genus CONUROPSIS Salvadori 

Connropsis Salvadori, Cat. Birds Brit. Mus., vol. 20, 1891, pp. 146, 203. Tyi)e, 
by original designation, Psittacus carolinensis Linnaeus, 

Conuropsis fratercula Wetmore 

Connropsis jratercnla Wetmore, 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 : Plantain-eaters and Cuckoos 

Suborder CUCULI : Cuckoos, Roadrunners, and Anis 

Family CUCULIDAE : Cuckoos, Roadrunners, and Anis 

Subfamily NEOMORPHINAE: Ground Cuckoos 

Genus GEOCOCCYX Wagler 

Geococcyx Wagler, Isis von Oken, vol. 24, Heft 5, May 1831, col. 524. Type, 
by monotypy, Geococcyx varicyala Wagler = Saurothcra calif orniana 
Lesson. 



'^ Included here since it has not been found in living form, being known only 
from bones. 



84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I3I 

Geococcyx calif ornianus (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 Howard 

Geococcyx conklingi Howard, Condor, vol. 22, 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 : Owls ^^ 

Family PROTOSTRIGIDAE : Protostrix 

Genus PROTOSTRIX Wetmore 

Protostrix Wetmoue, Ainer. Mus. Nov., No. 680, Dec. 4, 1933, p. 3. Type, 
by original designation, Aquila lydekkeri Shufeldt. 

Protostrix lydekkeri (Shufeldt) 

Aquila lydekkeri Shufeldt, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 16, 
Aug. 4, 1913, P- 298. 

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 lo miles from Fort Bridger, Wyoming. 

Protostrix leptosteus ( Marsh )i* 
Bubo leptosteus Marsh, Amer. Journ. Sci., ser. 3, vol. 2, August 1871, p. 126. 

Eocene (Bridger formation) : Grizzly Buttes, near Fort Bridger, 
Wyoming. 

1- Possibly of Recent period. 

13 Aquila antiqua Shufeldt, type of the genus Mineri'a Shufeldt, formerly con- 
sidered an owl, proves to be a mammal. See Wetmore, Amer. Mus. Nov., No. 
680, Dec. 4, 1933. PP- I. 2. 

" See Wetmore, Condor, 1937, pp. 84-85. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 85 

Protostrix mimica Wetmore 

Protostrix mimica Wetmore, 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 Owls 

Subfamily TYTONINAE: Barn Owls 

Genus TYTO Billberg 

Tyto BuiBERG, Syn. Faunae Scand., vol. i, pt. 2, 1828, tab. A. Type, by 
monotypy, Strix flammea auct. = Strix alba Scopoli. 

Tyto alba (Scopoli): Barn Owl 

Strix alba Scopoli, Annus i, Historico-Naturalis, 1769, p. 21. 

Modern form reported from Pleistocene: Cavern deposits near 
Lecanto, Florida ; ^^ San Josecito Cave, Aramberri, Nuevo Leon. Late 
Pleistocene : Carpinteria, Santa Barbara County and Rancho La Brea, 
Los Angeles, California. 

Tyto cavatica Wetmore 

Tyto cavatica Wetmore, Proc. Biol. Soc. Washington, vol. 33, Dec. 30, 1920, 
p. 80, pi. 3, figs. 3-6, 

Recent (extinct) : " Cave deposits in Cueva Toraiio, 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 Wetmore, Bull. Mus. Comp. Zocil., vol. 80, No. 12, October 1937, 
p. 436, figs. 10-16. 

Recent (extinct) : " Cave deposits on Great Exuma Island, Ba- 
hama Islands. 



IS 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. 

1^ 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., 1769, p. 3. Type, by monotypy, Otus bakkamoetia 
Pennant. 

Otus asio (Linnaeus) : Screech Owl 

Strix asio Linnaeus, Syst. Nat., ed. 10, vol. i, 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) Hammeola Kaup, in Jardine, Contr. Orn., 1852 (1853), 
p. III. 

Modern form reported from Pleistocene : San Josecito Cave, Aram- 
berri, Nuevo Leon. Late Pleistocene : Samwel Cave,^^ 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 Dumgril 

Bubo DuMERiL, Zool. Analytique, 1806, p. 34. Type, by tautonymy, Strix 
bubo Linnaeus. 

Bubo virginianus (Gmelin) : Horned Owl 
Strix virginiaiia Gmelin, Syst. Nat., vol. i, pt. i, 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 Sinclair! Miller 

Bubo sinclairi L. H. Miller, Univ. California Publ., Bull. Dept. Geol., vol. 6, 
No. 16, Oct. 28, 191 1, p. 393, figs. 4-5. 

Late Pleistocene: Samwel and Potter Creek (type locality) caves, 
Shasta County, California. 



i'^ Recorded originally as Micropallas whitneyi. See Miller, L. H., Trans. San 
Diego Soc. Nat. Hist., vol. 7, No. 19, Mar. 31, 1933, pp. 209-210. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 87 

Genus GLAUCIDIUM Boie 

Glaucidium Boie, Isis von Oken, Bd. 2, 1826, col. 970. Type, by subsequent 
designation, Strix passerina Linnaeus (Gray, 1840). 

Glaucidium gnoma Wagler : 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 Gloger, Hand- und Hilfsbuch Naturg., 1842 (1841), p. 226. Type, 
by monotypy, Strix cuntcularia Molina. 

Speotyto cunicularia (Molina) : Burrowing Owl 
Strix Cnnicidaria Molina, 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 huhtda =: Strix huhula Daudin. 

Ciccaba virgata (Cassin) : 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. i, 1758, p. 92. Type, by tautonymy, 
Strix stridtda 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. 
ID, i860), p. 193. 

Modern form reported from Pleistocene : San Josecito Cave, Aram- 
berri, Nuevo Leon. 



88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Strix brea Howard 
Strix brea Howard, Condor, vol. 35, No. 2, Mar. 15, 1933, p. 66, fig. 15. 

Late Pleistocene: Rancho La Brea, Los Angeles, California. 

Strix dakota Miliar 

Strix dakota A. H. Miller, 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. i, p. 28. Type, by tautonymy, Asio Brisson = 
Strix otus Linnaeus. 

Asio otus (Linnaeus) : " Long-eared Owl 

Strix Otus Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 92. 

Modern form reported from late Pleistocene : Samwel Cave, Shasta 
County, McKittrick, Kern County, and Carpinteria, Santa Barbara 
County, California.^^ Pleistocene: San Josecito Cave, Aramberri, 
Nuevo Leon. 

Asio flammeus (Pontoppidan) : Short-eared Owl 
Strix flammea Pontoppidan, Danske Atlas, vol. i, 1763, p. 617, pi. 25. 

Modern form reported from late Pleistocene : Rancho La Brea, 
Los Angeles, California. 



Genus AEGOLIUS Kaup 

Aegolius Kaup, Skizz. Entw.-Gesch. Eur. Thierw., 1829, p. 34. Type, by 
monotypy, Strix tengmabni Gmelin = 6'/rt;r ju>tereus Linnaeus, 1758. 

Aegolius funereus (Linnaeus) : Boreal Owl 
Strix junerea Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 93. 

Modern form reported from Pleistocene : -° Shelter cave. Pyramid 
Peak, Organ Mountains, Dona Ana County, New Mexico. 

'^^ Asio wilsonianus (Lesson) of the preceding list. 

1" 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. i, pt. i, 1788, p. 296. 

Modern form reported from late Pleistocene: Rancho La Brea, 
Los Angeles, California. Pleistocene: San Josecito Cave, Aramberri, 
Nuevo Leon. 



Order CAPRIMULGIFORMES : Oilbirds, Goatsuckers, and Allies 

Suborder CAPRIMULGI : Goatsuckers, Potoos, and Frogmouths 

Family CAPRIMULGIDAE: Goatsuckers 

Subfamily CAPRIMULGINAE: Goatsuckers 

Genus PHALAENOPTILUS Ridgway 

Phalaenoptilus Ridgway, Proc. U. S. Nat. Mus., vol. 3, 1880, p. 5. Type, by 
original designation, Caprimiilgus nuttallii Audubon 

Phalaenoptilus nuttallii (Audubon) : Poor- will 

Caprimulgiis Nuttalli Audubon, 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 Woodpeckers 

Suborder PICI : Woodpeckers and Wrynecks 

Family PICIDAE: Woodpeckers, Wrynecks, and Piculets 

Subfamily PICINAE : Woodpeckers 

Genus COLAPTES Vigors 

Colaptes Vigors, Trans. Linn. Soc. London, vol. 14, pt. 3, 1826, p. 457. Type, 
by original designation, Cuciilus aiiratus Linnaeus. 

Colaptes cafer (Gmelin) : Red-shafted Flicker 
Pictts cafer Gmelin, Syst. Nat., vol. i, pt. i, 1788, p. 431. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon ; 
Samwel and Potter Creek caves, Shasta County, Hawver Cave, El- 
dorado County, McKittrick, Kern County, Carpinteria, Santa Barbara 
County, and Rancho La Brea, Los Angeles, California. 



90 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Colaptes chrysoides (Malherbe) : Gilded Flicker 

Gcopicus (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 

Dryocoptts Boie, Isis von Oken, Bd. 2, 1826, col. 977. Type, by monotypy, 
Pious martins Linnaeus. 

Dryocopus pileatus (Linnaeus) : Pileated Woodpecker 
Picns pileatus Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 113. 

Modern form reported from Pleistocene : Cave deposits of Tennes- 
see. Late Pleistocene : Rancho La Brea, Los Angeles, California. 

Genus ASYNDESMUS Coues 

Asyndesmns Coues, Proc. Acad. Nat. Sci. Philadelphia, vol. 17, No. i, 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 : Perching Birds 

Suborder PASSERES: Song Birds 

Family ALAUDIDAE: Larks 

Genus EREMOPHILA Brehm 

Eremophila Brehm, Isis, vol. 21, pts. 3-4, 1828, p. 322. Type, by subsequent 
designation, Alauda alpestris Linnaeus (Sharpe, 1890). 

Eremophila alpestris (Linnaeus) : Homed Lark 

Alauda alpestris Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 166. 

Modern form reported from late Pleistocene: McKittrick and 
Rancho La Brea, Los Angeles, California. 

Family PALAEOSPIZIDAE: Palaeospiza 

Genus PALAEOSPIZA Allen 

Palaeospiza Allen, Bull. Geol. Geogr. Surv. Terr., vol. 4, No. 2, May 3, 1878, 
p. 443. Type, by monotypy, Palaeospiza hella Allen. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 9I 

Palaeospiza bella Allen 

Palaeospica bella Allen, Bull. Geol. Geogr. Surv. Terr., vol. 4, No. 2, May 3, 
1878, p. 443, pi. I, figs. 1-2. 

OUgocene (Florissant lake beds) : ^^ Florissant, Colorado. 

Family HIRUNDINIDAE: Swallows 

Genus PETROCHELIDON Cabanis 

Pctrochelidon Cabanis, Mus. Hein., vol. i, October (after Oct. 23), 1851, 
p. 47. Type, by subsequent designation, Hirundo melanogaster Swainson 
(Gray, 1855). 

Pctrochelidon pyrrhonota (Vieillot) : Cliflf Swallow 

Hirundo pyrrhonota Vieillot, Nouv. Diet. Hist. Nat., nouv. ed., vol. 14, Sep- 
tember 1817, p. 519. 

Modern form reported from late Pleistocene: McKittrick, Cali- 
fornia. 

Family CORVIDAE: Jays, Magpies, and Crows 

Subfamily GARRULINAE : Jays and Magpies 

Genus CYANOCITTA Strickland 

Cyanocitta Strickland, Ann. Mag. Nat. Hist., ser. i, vol. 15, No. 98, April 
1845, p. 261. Type, by original designation, Corvtis cristatus Linnaeus. 

Cyanocitta stelleri (Gmelin) : Steller's Jay 
Corvus stelleri Gmelin, Syst. Nat., vol. i, pt. i, 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 

Aphclocoma Cabanis, Mus. Hein., vol. i, sign. 28, Oct. 15, 1851, p. 221. Tyi>e, 
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. i, pt. i, 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. 



92 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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. i, p. 30; vol. 2, p. 35. Type, by tautonymy, 
Pica Brisson^ Corvus pica Linnaeus. 

Pica nuttallii (Audubon) : Yellow-billed Magpie 
Corvus nuttallii Audubon, Birds Amer. (folio), vol. 4, 1836, pi. 362, fig. i. 

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 

Corviis Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 105. Type, by tautonymy, 
Corvus = Corvus corax Linnaeus. 

Corvus corax Linnaeus: Common Ravenna 

Corvus Corax Linnaf.us, Syst. Nat., ed. 10, vol. i, 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 Couch : White-necked Raven 

Corvus cryptoleucus Couch, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, No. 2, 
May 20, 1854, p. 66. 

Modern form reported from late Pleistocene : McKittrick and 
Rancho La lirea, Los Angeles, California. 

Corvus brachyrhynchos Breiim: Crow 

Corvus brachyrhynchos C. L. Brehm, Bcitr. Vogelkundc, 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 Corinis shufcldti Sharing is a synonym of C. corax. See Howard, Carnegie 
Inst. Washington Publ. 551, Jan. 25, 1946, p. 189. 
2* Record formerly given from Carpinteria refers to C. caurinus. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE 93 

Corvus caurinus Baird: Northwestern Crow 

Corims caurinus Baird, 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 Wilson : Fish Crow 
Corvus ossifragus Wilson, Amer. Orn., vol. 5, 1812, p. 27, pi. 27, 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, pi. 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 

Gyvmorhinus Wied, Reise Nord-Amer., vol. 2, 1841, p. 21. Type, by monotypy, 
Gymnorhimis cyanocephalus Wied. 

Gymnorhinus cyanocephalus Wied: Pifion Jay 
Gymnorhinus cyanocephalus Wied, 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: Nuthatches 

Subfamily SITTINAE: Typical Nuthatches 

Genus SITTA Linnaeus 

Sitta Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 115. Type, by monotypy, 
Sitla europaea Linnaeus. 

Sitta canadensis Linnaeus : Red-breasted Nuthatch 
Sitta canadensis Linnaeus, Syst. Nat., ed. 12, vol. i, 1766, pp. 176, 177. 

Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 

'5 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. I3I 

Sitta pygmaea Vigors : Pygmy Nuthatch 
Sitta pygmaea Vigors, in Zool. Beechey's Voy., 1839, p. 25, pi. 4, fig. 2. 

Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 

Family CHAMAEIDAE: V^ren-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, Panis 
jasciatus Gambel. 

Chamaea fascia ta (Gambel) : Wren-tit 

Parii^s jasciatus Gambkl, 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: Thrashers and Mockingbirds 

Genus TOXOSTOMA Wagler 

Toxostoma Wagler, Isis von Oken, vol. 24, Heft 5 (May) 1831, col. 528. 
Type, by monotypy, Toxostoma vcltila Wagler = Orpheus curvirostris 
Swainson. 

Toxostoma bendirei (Coues) : Bendire's Thrasher 
Harporhynchus betidirei Coues, 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 

Harpcs rcdiviva 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 

Oreoscoptcs Baird, 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: Thrushes 

Genus TURDUS Linnaeus 

Turdus Linnaeus, Syst. Nat., ed. lo, vol. i, 1758, p. 168. Type, by subsequent 
designation, Turdus viscivorns Linnaeus (Gray, 1840). 

Turdus migratorius Linnaeus: Robin 
Turdus migratorius Linnaeus, Syst. Nat., ed. 12, vol. i, 1766, p. 292. 

Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 

Genus SIALIA Swainson 

Sialia Svi'AINSON, Philos. Mag., n. s., vol. i, No. 5, May 1827, p. 369. Type, by 
monotypy, Sialia asurea Swainson = Motacilla sialis Linnaeus. 

Sialia mexicana Swainson : Western Bluebird 

Sialia mexicana Swainson, Fauna Bor.-Amer., vol. 2, 1831 (February, 1832), 
p. 202. 

Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 

Family BOMBYCILLIDAE : Waxwings 

Genus BOMBYCILLA Vieillot 

Bombycilla Vieillot, Hist. Nat. Ois. Amer. Sept., vol. i, 1807 (1808), p. 88. 
Type, by monotypy, Bombycilla cedrorum Vieillot. 

Bombycilla cedrorum Vieillot: Cedar Waxwing 

Bombycilla cedrorum Vieillot, Hist. Nat. Ois. Amer. Sept., vol. i, 1807 
(i8o8),p. 88, pi. 57. 

Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, and Rancho La Brea, Los Angeles, California. 

Family LANIIDAE: Shrikes 

Subfamily LANIINAE: Shrikes 

Genus LANIUS Linnaeus 

Lanius Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 93. Type, by subsequent 
designation, Lanius excubitor Linnaeus (Swainson, 1824). 



96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Lanius ludovicianus Linnaeus: Loggerhead Shrike 

Lanius ludovicianus Linnaeus, Syst. Nat., ed. 12, vol. i, 1766, p. 134. 

Modern form reported from late Pleistocene: McKittrick, Kern 
County, and Rancho La Brea, Los Angeles, California. 

Family ICTERIDAE : Meadowlarks, Blackbirds, and Troupials 

Genus STURNELLA Vieillot 

Sturnella Vieillot, Analyse, 1816, p. 34. Type, by monotypy, Stourne, ou 
Merle a fer-a-cheval Buff on = Alauda magna Linnaeus. 

Sturnella neglecta Audubon : Western Meadowlark 

Sturnella neglecta Audubon, Birds Amer., octavo ed., vol. 7, 1844, p. 339, 
pi. 489. 

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 AGELAITJS Vieillot 

Agelaius Vieillot, Analyse, 181 6, p. Z2i- Type, by subsequent designation, 
Troupiale commandeur Buff on = Ono/M.s phoeniceus Linnaeus (Gray, 
1840). 

Agelaius phoeniceus (Linnaeus) : Red-winged Blackbird 

Oriolus phoeniceus Linnaeus, Syst. Nat., ed. 12, vol. i, 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 20 

Psarocolius cyanocephalus Wagler, Isis von Oken, vol. 22, Heft 7 (July), 
1829, col. 758. 

Modern form reported from late Pleistocene : Fossil Lake, Oregon ; 
McPherson County, Kansas (Kentuck locality). 



28 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. 399, 
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 docs 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. Miller, Univ. California Publ., Bull. Dept. 
Geol. Sci., vol. 19, No. i, Dec. 21, 1929, p. 14, pi. i, figs. /, h. 

Late Pleistocene : Rancho La Brea, Los Angeles, California. 

Genus CASSIDIX Lesson 

Cassidix Lesson, Traite d'Orn., livr. 6, Feb. i, 1831, p. 433. Type, by sub- 
sequent designation, Cassidix mexicanus Lesson = Corvus mexicantis 
Gmelin (Gray, 1840). 

Cassidix mexicanus (Gmelin) : Boat-tailed Grackle 

Corvus mexicanus Gmelin, Syst. Nat., vol. i, pt. i, 1788, p. 375. 

Modern form reported from Pleistocene: Seminole Field, Pinellas 
County, Florida. 

Genus QUISCALUS Vieillot 

Quiscaliis Vieillot, Analyse, 1816, p. 36. Type, by subsequent designation, 
Gracula quiscula Linnaeus (Gray, 1840). 

Quiscalus quiscula (Linnaeus) : Grackle 
Graada Quiscula Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 109. 

Modern form reported from Pleistocene : Seminole Field, Pinellas 
County, Florida. 

Genus PYELORHAMPHUS Miller 

Pyelorhamphus A. H. Miller, Auk, vol. 49, No. i, January 1932, p. 39. Type, 
by original designation, Pyelorhamphus molothroides Miller. 

Pyelorhamphus molothroides Miller 

Pyelorhamphus molothroides A. H. Miller, Auk, vol. 49, No. l, January 
1932, p. 39, Pl- 4- 

Quaternary ( ? Pleistocene) : " Shelter Cave, Pyramid Peak, Organ 
Mountains, Dona Ana County, New Mexico. 

Genus PANDANARIS Miller 

Pandanaris A. H. Miller, Condor, vol. 49, No. i, Feb. 6, 1947, p. 22. Type, 
by original designation, Pandanaris convexa A. H. Miller. 

Pandanaris convexa Miller 

Pandanaris convexa A. H. Miller, Condor, vol. 49, No. i, Feb. 6, 1947, p. 22, 
fig. 4 a-d. 

Late Pleistocene: Pit "A," Rancho La Brea, Los Angeles, Cali- 
fornia. 



27 The deposits in which this extinct species was found are possibly of Recent 
age. 



98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Family FRINGILLIDAE : Grosbeaks, Finches, Sparrows, and 

Buntings 

Subfamily RICHMONDENINAE: Cardinals and Allies 

Genus PHEUCTICUS Reichenbach 

Pheucticus Reichenbach, Av. Syst. Nat., June i, 1850, pi. 78. Type, by sub- 
sequent designation, Pitylus aureovcniris Lafresnaye and d'Orbigny (Gray, 
1855). 
Pheucticus melanocephalus (Swainson) : Black-headed Grosbeak 

Guiraca melanocephala Swainson, Philos. Mag., n. s., vol. i, No. 6, June 
1827, p. 438. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, California. 

Subfamily CARDUELINAE : Purple Finches, Goldfinches, and 

Allies 

Genus HESPERIPHONA Bonaparte 

Hesperiphona Bonaparte, Consp. Gen. Avium, vol. i, 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. Lye. Nat. Hist. New York, vol. i, pt. 2, 
1825, p. 220. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, California. 

Genus CARPODACUS Kaup 

Carpodacus Kaup, Skizz. Entw.-Gesch. Eur. Thierw., 1829, p. 161. Type, by 
subsequent designation, Loxia rosea Pallas (Gray, 1842). 

Subgenus BURRICA Ridgway 

Burrica Ridgway, Man. North Amer. Birds, 1887, p. 390. Type, by original 
designation, Fringilla tnexicana Miiller. 

Carpodacus mexicanus (Muller) : House Finch 

Fringilla wcxicaua P. L. S. Muller, 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 SPINTJS Koch 

Spinus Koch, Syst. Baier. Zool., vol. i, 1816, p. 233. Type, by tautonymy, 
Fringilla spinus Linnaeus. 

Spinus pinus (Wilson) : Pine Siskin 
Fringilla pinus Wilson, Amer. Orn., vol. 2, 1810, p. 133, pi. 17, fig. i. 

Modern form reported from Pleistocene: Carpinteria and Rancho 
La Brea, Los Angeles, California. 

Spinus tristis (Linnaeus) : American Goldfinch 
Fringilla tristis Linnaeus, Syst. Nat., ed. 10, vol. i, 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. i, 1758, p. 171. Type, by subsequent 
designation, Loxia curvirostra Linnaeus (Gray, 1840). 

Lozia curvirostra Linnaeus : Red Crossbill 
Loxia Curvirostra Linnaeus, Syst. Nat., ed. 10, vol. i, 1758, p. 171. 

Modern form reported from late Pleistocene: Carpinteria, Santa 
Barbara County, California. 



Subfamily EMBERIZINAE : Sparrows and Buntings 
Genus PALAEOSTRUTHUS Wetmore 

Palaeostruthus Wetmore, Bull. Mus. Comp. Zool., vol. 67, May 1925, p. 192. 
Type, by original designation, Palacospisa hatcheri Shufeldt. 

Palaeostruthus hatcheri (Shufeldt) 

Palaeospiza hatcheri Shufeldt, Bull. Amer. Mus. Nat. Hist., vol. 32, art. 16, 
Aug. 4, 1913, p. 301, pi. 55, fig- 28. 

Middle Pliocene : Near Long Island, Kansas. 

Genus PIPILO Vieillot 

Pipilo Vieillot, Analyse, 181 6, p. 32. Type, by monotypy, Pinson aux yeux 
rouges Buff on = Fringilla erythrophthalma Linnaeus. 

Pipilo maculatus Swainson : Spotted Towhee 
Pipilo tnaculata Swainson, Philos. Mag., n. s., vol. i, 1827, p. 434. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, and Carpinteria, California. 



lOO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Pipilo fuscu3 SwAiNSON : Brown Towhee 

Pipilo fiisca SwAiNSON, Philos. Mag., n. s., vol. i, 1827, p. 434. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, and Carpinteria, California. 

Pipilo angelensis Dawson 
Pipilo angelettsis Dawson, Condor, vol. 50, No. 2, Mar. 16, 1948, p. 39, fig. 16. 

Late Pleistocene : Rancho La Brea, Los Angeles, California. 

Genus CALAMOSPIZA Bonaparte 

Calamospisa 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. i, 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. i. No. 6, June 1827, p. 435. 
Type, by monotypy, Ammodramus bimacxdatus Swainson. 

Ammodramus savannarum (Gmexin) : Grasshopper Sparrow 
Fringilla savannarum Gmelin, Syst. Nat., vol. i, 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 

Pooecetcs Baird, in Baird, Cassin, and Lawrence, Rep. Expl. Surv. R. R. Pac, 
vol. 9, 1858, pp. XX, XXXIX. Type, by monotypy, Fringilla graminca 
Gmelin. 

Pooecetes gramineus (Gmelin) : Vesper Sparrow 

Fringilla graminea Gmelin, Syst. Nat., vol. i, pt. 2, 1789, p. 922. 

Modern form reported from Pleistocene : Rancho La Brea, Los 
Angeles, California. 

Genus CHONDESTES Swainson 

Chondcstes Swainson, Philos. Mag., n. s., vol. i, No. 6, June 1827, p. 435. 
Type, by monotypy, Chondcstes strigatus Swainson. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE lOI 

Chondestes grammacus (Say) : Lark Sparrow 
Fringilla grammaca Say, in Long, Exped. Rocky Mts., vol. i, 1823, p. 139. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, California, 

Genus AMPHISPIZA Coues 

Amphispiza Coues, Birds Northwest, 1874, p. 234. Type, by original designa- 
tion, Emberisa bilineata Cassin. 

Amphispiza bilineata (Cassin) : Black-throated Sparrow 

Emberiza bilineata Cassin, Proc. Acad. Nat. Sci. Philadelphia, vol. 5, No. 5, 
September-October (Dec. 7), 1850, p. 104, pi. 3. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, California. 

Amphispiza belli (Cassin) : Bell's Sparrow 

Emberiza Belli Cassin, Proc. Acad. Nat. Sci. Philadelphia, vol. 5, No. 5, 
September-October (Dec. 7), 1850, p. 104, pi. 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, pi. 120, fig. I. 

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, 183 1 (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. I3I 

Genus PASSERELLA Swainson 

Passerella Swainson, Nat. Hist, and Class. Birds, vol. 2, July i, 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. ^7, 
pi. ID. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, and Carpinteria, California, 



Genus MELOSPIZA Baird 

Melospisa Baird, 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 (Wilson) : Song Sparrow 
Fringilla melodia Wilson, Amer. Orn., vol. 2, 1810, p. 125, pi. 16, fig. 4. 

Modern form reported from late Pleistocene : Rancho La Brea, Los 
Angeles, California. 

INCERTAE SEDIS 
Genus CIMOLOPTERYX Marsh =8 

Cimolopteryx Marsh, Amer. Journ. Sci., ser. 3, vol. 38, 1889, p. 83, foot- 
note. Type, by monotypy, Cimolopteryx rarus Marsh. 

Cimolopteryx rarus Marsh 

Cimolopteryx rarus Marsh, Amer. Journ. Sci., ser. 3, vol. 38, July 1889, 
p. 83, footnote. 

Upper Cretaceous (Lance formation) : Niobrara County, Wyoming. 

Cimolopteryx retusus Marsh 

Cimolopteryx retusus Marsh, 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 diflferent 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 IO3 

Genus EOPTERYX Meyer 

Eopteryx Meyer, Ber. Senckenberg. Nat. Ges. Frankfurt am Main, 1887, 
p. 14. Type, by monotypy, Eopteryx mississippiejisis Meyer. 

Eopteryx mississippiensis Meyer -^ 

Eopteryx mississippiensis Meyer, Ber. Senckenberg. Nat. Ges. Frankfurt am 
Main, 1887, p. 14, pi. 2, figs. 22a-22c. 

Eocene : Jackson, Mississippi. 

(Genus uncertain) 
Falco falconellus Shufeldt so 
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 ^i 

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 Shufeldt, Journ. Geol., vol. 21, October-November (Nov. i), 1913, 
p. 644. Type, by monotypy, Hebe schucherti Shufeldt. 

Hebe schucherti Shufeldt ^2 

Hebe schucherti Shufeldt, Journ. Geol., vol. 21, October-November (Nov. i), 
1913, p. 644, fig. 10, a, b. 

Eocene : ^^ 5 miles west of Green River, Wyoming. 



28 Described from a fragmentary vertebra. 

3° Not a falcon ; relationships doubtful. See Wetmore, A., Proc. U. S. Nat. 
Mus., vol. 84, Nov. 3, 1936, pp. 77-78. 

^1 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. Z06I., vol. 67, May 1925, p. 184. Possibly of Oligo- 
cene age. 

32 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. 

33 From data furnished by Dr. M. R. Thorpe, of the Peabody Museum, Yale 
University. 



104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Genus IGNOTORNIS Mehl 

Ignotornis Mehl, Amer. Journ. Sci., ser. 5, vol. 21, May 1931, p. 443. Type, 
by monotypy, Ignotornis mccomielli Mehl. 

Ignotornis mcconnelli Mehl ^4 

Ignotornis mcconnelli Mehl, Amer. Journ. Sci., ser. 5, vol. 21, May 1931, 
p. 444, fig. I. 

Cretaceous (Dakota sandstone) : About i^ miles northwest of 
Golden, Colorado. 

Genus LAOPTERYX Marsh 

Laopteryx Marsh, Amer. Journ. Sci., ser. 3, vol. 21, April 1881, p. 341. Type, 
by monotypy, Laopteryx prisctis Marsh. 

Laopteryx priscus Marsh 35 

Laopteryx priscits Marsh, 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 Marsh ^s 

Laornis edvardsianus Marsh, Amer. Journ. Sci., ser. 2, vol. 49, March 1870, 
p. 206. 

Paleocene (Homerstown 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 ^~ 

Palaeonornis Struthionoides Emmons, Amer. Geol., pt. 6, 1857, p. 148, fig. 114. 

? Triassic : Anson County, North Carolina. 



3' Described from fossil impressions of 4-toed footprints, apparently with webs 
connecting the three anterior toes. 

35 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. 

38 Doubtfully related to Anseriformes. Lambrecht, Handb. Palaeorn., 1933, 
pp. 526-527, has placed it uncertainly after the Aramidae. 

37^ Affinity doubtful : possibly not avian. 



NO. 5 CHECK-LIST OF FOSSIL BIRDS — WETMORE IO5 

Genus UINTORNIS Marsh 

Uintoniis Marsh, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, p. 259. 
Type, by monotypy, Uintornis lucaris Marsh. 

Uintomis lucaris Marsh ^s 

Uintoniis lucaris Marsh, Amer. Journ. Sci., ser. 3, vol. 4, October 1872, p. 259. 

Eocene (Bridger formation) : Near Henry's Fork, Wyoming. 

Genus YALAVIS Shufeldt 

YaJavis Shufeldt, Journ. Geol., vol. 21, October-November (Nov. i), 1913, 
p. 649. Type, by monotypy, Yalavis tenuipes Shufeldt. 

Yalavis tenuipes Shufeldt ^9 

Yalavis tenuipes Shufeldt, Journ. Geol., vol. 21, October-November (Nov. i), 
1913, p. 649, figs. lie and 12c. 

Geologic age and locality of occurrence not known. 



38 According to Shufeldt, Trans. Connecticut Acad. Arts Sci., vol. 19, February 
191 5> PP- 50-52, 77, pi. 6, fig. 42, this species is of uncertain affinity, and is not a 
woodpecker as suggested by Marsh. 

29 Said in the original description to be a passeriform bird of uncertain affinity. 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 6 



Cftarlesi ©, anb JHarp ^aux SKHalcott 
3Res!earcf) Jf unb 



PALEOCENE MAMMALIAN FAUNAS OF THE 

BISON BASIN IN SOUTH-CENTRAL 

WYOMING 

(With 16 Plates) 

By 
C. LEWIS GAZIN 

Curator, Division of Vertebrate Paleontology 

United States National Museum 

Smithsonian Institution 




(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 

Introduction i 

Acknowledgments 2 

History of investigation 2 

Occurrence and preservation of material 3 

The Bison basin faunas 4 

Environment and relationships between the Bison basin faunas 7 

Age and correlation of the faunas lo 

Systematic description of vertebrate remains I2 

Reptilia 12 

Sauria 12 

Anguidae 12 

Mammalia 12 

Multitubcrculata 12 

Ptilodontidae 12 

Marsupialia 14 

Didelphidae 14 

Insectivora 15 

Leptictidae 15 

Pantolestidae 17 

Primates 19 

Plesiadapidae 19 

Carnivora 25 

Arctocyonidae 25 

Mesonychidae 35 

Miacidae 35 

Condylarthra 36 

Hyopsodontidae 36 

Phenacodontidae 42 

Pantodonta 47 

Coryphodontidae 47 

References 51 

Explanation of plates 54 



ILLUSTRATIONS 

Plates 

(All plates following page 58.) 

1. Mullituberculates and insectivores from the Bison basin Paleocene. 

2. Primates and marsupials from the Bison basin Paleocene. 

3. Pronothodcctes from the Bison basin Paleocene. 

4. Plcsiadapis from the Bison basin Paleocene. 

5. Triccntcs and Chriacus from the Bison basin Paleocene. 

6. Thryptacodon from the Bison basin Paleocene. 

7. Clacnodon from the Bison basin Paleocene. 

8. Litomylus and Protoselcnc? from the Bison basin Paleocene. 

9. Haplaletes and Gidleyina from the Bison basin Paleocene. 

10. Phcnacodusl from the Bison basin Paleocene. 

11. Condylarths and Titanoidcs from the Bison basin Paleocene. 

12. Caenolambda from the Bison basin Paleocene. 

13. Caenolambda from the Bison basin Paleocene. 

14. Caenolambda from the Bison basin Paleocene. 

15. South rim of Bison basin showing fossil localities. 

16. Two fossil localities in the Bison basin. 



Figures 

Page 

1. Map of western Wyoming and portions of adjacent States 5 

2. Histogram of length of Mj of Claenodon 33 



IV 



Cijarletf ■©. anli iHarp ^aux ffiSalcott 3^cfi£carcf) jFunb 

PALEOCENE MAMMALIAN FAUNAS OF THE 

BISON BASIN IN SOUTH-CENTRAL 

WYOMING 

By C. lewis GAZIN 

Curator, Dknsion of Vertebrate Paleontology 

United States National Museum 

Stnithsonian Institution 

(With 16 Pi-ates) 

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. I3I 

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 1 1 were 
made by Lawrence B. Isham ; those of Caenolambda patter soni 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 1952 ^ 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 
Bison 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 

1 A very cursory examination of these beds was made by the writer in 1951, 
accompanied by C. L. 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 pis. 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 
pattersoni, 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. I3I 

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 




Fig. I. — Map of western Wyoming and portions of adjacent States showing Eocene 
sedimentary basins, with nearby Paleocene fossil localities numbered as follows: 
I, 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. Geo!. Surv. Monogr. 
55, figs. 9. 49. 1929. 



6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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. 



Reptilia : 

Sauria : ^ 

Anguid, undct 6 

Mammalia : 

Multituberculata : 

Cf. Ptilodus montanus Douglass i 

Cf. Ectypodiis niusculus Matthew and Granger. ... i 

Cf . Ectypodus ha::cni Jepsen i 

Cf. Anconodon russclli ( Simpson) i 

Marsupialia : 

Peradectcs clegans Matthew and Granger i 

Peradectes paidi, new species 2 

Insectivora : 

Diacodon pcarcei, new species I 

Disonalveus brozimi, new genus and species 4 

Primates : 

Pronothodcctcs, cf. mattJtezin Gidley 4 

Pronothodcctes simpsoui, new species 

Pronothodectes, cf. simpsoni, new species 8 

Plesiadapis, cf. jodinatus Jepsen 

Plesiadapis jepseni, new si)ecies 11* 

Plesiadapis, cf. jepseni, new species 

Carnivora : 

Tricentcs freinoutcnsis, new species 11 

Chriacus, near C. pehidots (Cope) 3 

Chriacus, sp. ( small ) I 

Thryptacodon, cf. aiistralis Simpson 

Thryptacodon demari, new species 

Thryptacodon, cf. demari, new species 2 

Thryptacodon belli, new species 11 



.s o 

^ 1 



NO. 6 



PALEOCENE FAUNAS OF BISON BASIN — GAZIN 



"O bo m 

Mammalia — continued "^ "§ fe 

Carnivora — continued ^ 

Claenodon, z{. procyonoldcs (Matthew) 3 

Claenodon, ci. montanensis (Gidley) 5 

Claenodon, cf. jerox (Cope) 5 • • 3 

Claenodon acrogenius, new species 6 I 

Dissacus, sp 2 

Didymictis, near D. tenuis Simpson i 

Condylarthra : 

Promioclacnus pipiringosi, new species I 

Cf . Promioclaenus pipiringosi, new species 2 

Promioclaemis ? sp I 

Litomylus scaphicus, new species 2 i 

Litomyliis scapliiscns, new species i 2 

Haplaletes pelicatus, new species 4 

Haplaletes serior, new species 

Protoselene? novissimus, new species 2 

Litolestes lacunatus, new species 

Cf. Litolestes lacunattis, new species i 

Gidleyina ivyoiningensis, new species lO 

Gidleyina, cf. Wyoming ensis, new species 5 • • • • 3 

Phetiacodns? bisonensis, new species 28 2 

Phenacodus? sp. (large) I i 2 

Pantodonta : 

Titanoides primaevus Gidley I 

Caenolambda pattersoni, nevf genus and species. .. . I 

a. Caenolambda pattersoni, new genus and species, i i 
Pantodont, undet. (tooth frags.) 2 5 3 i 

* Two of these are from the vicinity of the saddle locality but approximately so feet 
higher. 



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 Tifa- 
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 perhajis 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 T. aiis- 
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, T. 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 
jcrox 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. I3I 

tween the two faunas ; however, a single specimen of Haplaletcs en- 
countered at the Titanoidcs locality is of a surprisingly larger form 
than the Ilaplaletes represented at the saddle. On the other hand, a 
single specimen from the saddle referred tentatively to Litolestcs 
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 Phcnacodus? 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 Promioclacnus (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, Litolcstes, 
Gidleyina, and Phcnacodus may be regarded as Tiffanian in first ap- 
pearance, whereas Pronothodcctes, Tricentes, Clacnodon, Promioclac- 
nus, Litomylus, Haplaletcs, 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 Pronothodcctes simpsoni, 
Clacnodon acrogcnius, Litomylus scaphicus, and Haplaletcs 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 Gidlcyina ivyomingensis, as well as in the close approach 
that Phenacodus? bisonensis makes to Tctraclaenodon. 

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. fodinatiis 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 tlian 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. I3I 

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 
Pcltosaurtis has been recognized in horizons as early as Lance and 
Fort Union. Nevertheless Glyptosaurus and Xesfops 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 Torre jonian 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 I, figure i 

A relatively large ptilodont multituberculate is represented by a 
single incomplete left P4, U.S.N.M. No. 2o8yy. There is no certainty 



NO. 6 PALEOCENE FAUNAS OF BISON BASIN — GAZIN I3 

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 outhne, as well as in the spacing of the serrations, 
of which there were at least 13, the tooth rather closely resembles P4 
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. jnontanus. 



Cf. ECTYPODUS MUSCULUS Matthew and Granger, 1921 

Plate I, figure 3 

A small jaw fragment with Mi and the alveoli for M2, U. of Wyo. 
No. 1105, would appear to represent Ectypodus miisculus. 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. iiiuscnlus, 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 P4, U.S.N.M. 
No. 20878, compared below with Ectypodus hascni but is appreciably 
shorter than the 3.2 mm. cited by Jepsen for the length of Mi in 
E. haseni, although the cusp formula for this tooth is the same as that 
for E. mus cuius. 



Cf. ECTYPODUS HAZENI Jepsen, 1940 
Plate I, figure 2 

An isolated though complete P4, U.S.N.M. No. 20878, compares 
very closely to this tooth in the Silver Coulee Ectypodus haceni. 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 P4 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 P3, but the absence of other associated material pre- 
cludes certain generic identification. 



14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

Cf. ANCONODON RUSSELLI (Simpson), 1935 
Plate I, figure 4 

Anconodon russelli may be represented by a fragment of the right 
mandibular ramus showing the root portion of the incisor and P4, 
U. of Wyo. No. 1065. The preserved premolar is close in size to that 
referred above to Ectypodus haseni, but its profile is slightly more 
convex dorsally and has a long straight front edge more as in An- 
conodon. P4 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 Peradcctes clegans 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 Pcradectes 
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. paiili 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 PcradcctCS 

Peradectes elegans Peradectes pauli, n. sp. 

, ' >/ " > 

A.M.N.H. U.S.N.M. 

No. 17376 U. ofWyo. No. 20879 U.S.N.M. 

(type) No. 1 104 (type) No. 20880 

Ml, anteroposterior diameter 1.7S ... ... 1.6 

transverse diameter of trigonid 0.9 ... ... 0.7 

transverse diameter of talonid 0.95 ... ... 0.75 

M2, anteroposterior diameter 1.75 i-7 

transverse diameter of trigonid i.o i.o 

transverse diameter of talonid 1.0 1.0 

M3, 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 

Ml, anteroposterior diameter 1. 75 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- 



1 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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- 
cinerei (Jepsen) and Palaeictops pineycnsis (Gazin) from among 
the lower Eocene forms, and possibly also Palaeictops miniitus (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,3 new species 
Plate I, figure 6 

Type. — Left ramus of mandible with P3-M1, 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. 
P4 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 I7 

Discussion. — The closeness of the resemblance between D. con- 
cordiarcensis and D. pearcci 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 P4, 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 P4 
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 P2 and P3 in Prodiacodon 
puercensis and much better developed in species of Palaeictops is 
not seen on the P2 of D. concordiarcensis or the P3 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 Prodiaco- 
don and Palaeictops. 

Among the other early leptictids, Leptacodon has a less progressive 
P4 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 P4 with paraconid for- 
ward as in Diacodon and Palaeictops, though lower; however, the 
greatly elongated talonids exhibited in the molars and P4 immediately 
distinguish this genus. The abbreviation of the talonid on P4 and the 
lower molars of Xenacodon is suggestive of Diacodon, but P4 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 

OF Diacodon pearcei, u.s.n.m. no. 20970 

Pa, anteroposterior diameter : greatest transverse diameter 1.8:1.0 

P*, anteroposterior diameter : transverse diameter of trigonid 2.4:1.4 

Ml, anteroposterior diameter : transverse diameter of trigonid 2.2:1.6 

PANTOLESTIDAE 

BIS0NALVEUS,4 new genus 

Type. — Bisonalveiis broumi, new species. 

Generic characters. — Resembling Aphronorus, but P4 much smaller 



* Bison + alvcus, basin — for Bison basin. 



l8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 Mj and M2 in- 
distinct and less prominently projecting on M3, Entoconid on Mi and 
M2 more forward in position. Small cuspule on crest anterolingual 
to hypoconid on M3. 

Discussion. — The structure of P4 and the general form of the molars 
suggest an alliance of this form with Aphr ononis 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. 

P4 of Bisonalveiis 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 Mi and M2 than in Aphronorus. 
In M3 the entoconid and hypoconulid are closer together. An- 
terior to the hypoconid on M3 (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 I, figure 5 

Type.—Leh ramus of mandible with P4-M3, U.S.N.M. No. 20928. 
Horizon and locality. — Bison basin Tiffanian, saddle locality, south 



6 Named for Dr. Roland W. Brown. 



NO. 6 PALEOCENE FAUNAS OF BISON BASIN — GAZIN I9 

rim of Bison basin, sec. 28, T. 27 N., R. 95 W., Fremont County, 
Wyo. 

Specific characters. — In size of teeth Bisonalvens brozvni is dis- 
tinctly smaller than Aphronorus jraudator Simpson. The length of 
the lower molar series is about 13 percent shorter, whereas P4 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 M3 (No. 20929) and 
an isolated P4 (No. 20930). A jaw fragment with a much worn P4 
and Ml in the University of Wyoming collections (No. 1067) may 
also represent this species. 

MEASUREMENTS IN MILLIMETERS OF LOWER TEETH IN TYPE SPECIMEN 

OF Bisonalveus broimii, u.s.n.m. no. 20928 

Length of lower cheek tooth series, P4-M3, incl 9.5 

Length of lower molar series, M1-M3, incl 7.5 

Pj, anteroposterior diameter : greatest transverse diameter 2.2:1.5 

Ml anteroposterior diameter : transverse diameter of trigonid 2.5: 1.7 

Ma anteroposterior diameter : transverse diameter of trigonid 2.6:2.0 

M3 anteroposterior diameter : transverse diameter of trigonid 2.6:1.5 

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. fodinatiis, 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 jepseni and Plesiadapis cf. jodinatiis (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. I3I 

PRONOTHODECTES, cf. MATTHEWI Gidley, 1923 
Plate 2, figures i, 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 P4-M2 and part of M3, and a second 
specimen, U. of Wyo. No. 1062, exhibits P3-M1. 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 siinpsoni. 

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. jodinatus. 

MEASUREMENTS IN MILLIMETERS OF TEETH REFERRED TO 

Pronothodectes matthczin gidley 

U. of Wyo. 
No. 1099 

M\ anteroposterior diameter 2.5 

M", anteroposterior diameter 2.5 

U.S.N.M. U. of Wyo. 

No. 20758 No. 1062 

P3, anteroposterior diameter : transverse diameter 1.8:1.5 

P4, anteroposterior diameter : transverse diameter .... 2.1:1.7 2.2:1.8 

Ml, anteroposterior diameter : transverse diameter 

of talonid 2.4 : 2.0 2.5 : 2.1 

M2, anteroposterior diameter : transverse diameter 

of talonid 2.5 : 2.3 

Ma, anteroposterior diameter 3.2' 

» Approximate. 

PRONOTHODECTES SIMPSONI,« new species 

Plate 3 

Type. — Right ramus of mandible with P4-M3, 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, W^ sec. 28, T. 27 N., R. 95 W., Fremont County, 
Wyo. 

Specific characters. — Size nearly intermediate between Pronotho- 
dectes mattheivi and Plcsiadapis gidleyi, closer to the latter. Pi 
pressed close to incisor, posterior lower premolars and molar trigonids 
of more inflated appearance than in P. matthewi. 

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, Pi and P2 
being, of course, single-rooted teeth as in P. mattheivi. 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. mattheivi, in addition to 
its greater size, by exhibiting teeth of a more Plesiadapis-Vike 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 M2 and M3. The variant from the sad- 
dle locality overlaps in size range that represented at the ledge locality 
and is less obviously different from P. mattheivi 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, pi. 3, fig. 2) of P. cf. simpsoni with 
P4 to M3 from the saddle locality exhibits a disproportionately long 
M3 (buccally incomplete), but other specimens with M3 from the 
saddle show this tooth to be normally proportioned. Mi and Mo 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. I3I 

MEASUREMENTS IN MILLIMETERS OF LOWER TEETH IN SPECIMENS OF 

Pronothodectes simpsoni 

U.S.N.M. 

No. 207S4 U.S.N.M. U. of Wyo. 

(type) No. 20770 No. 1057 * 

Length of lower molar series 9.8 8.9 9.4 

P4, anteroposterior diameter : transverse 

diameter 2.5 : 1.9 2.2 : 2.0 

Ml, anteroposterior diameter : greatest 

transverse diameter 2.8 : 2.4 2.6 : 2.3 2.5 : 2.1 

Ma, anteroposterior diameter : greatest 

transverse diameter 3.1 : 2.8 2.9 : 2.5 2.7 : 2.5 

M3, anteroposterior diameter : greatest 

transverse diameter 4.3 : 2.5 3.6 : 2.2 4.4 :• . 

* Small variant but with large M3 

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 P4 to M3, and only 
one (U. of Wyo. No. 1085) of the specimens is a maxilla, exhibiting 
M2 and M3. 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. jodinatus 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. jodinatus 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 P2 had been 
present, although there was no evidence of a Pi. P2 is not present in 
the type or other observed materials of P. jodinatus from the Prince- 
ton Quarry but is present in P. gtdleyi 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 Plcsiadopis, CF. 

fodinatus jepsen 

U. of Wyo. 
No. 1 08 s 

M*, anteroposterior diameter : greatest transverse diameter 3.8:5.9 

M*, greatest transverse diameter 5.5 

U. of Wyo. U.S.N.M. U.S.N.M. 

No. 1082 No. 20783 No. 20784 

P4, anteroposterior diameter : greatest 

transverse diameter 2.6 : 2.2 

Ml, anteroposterior diameter : greatest 

transverse diameter 3.6 : 2.9 3.2 : 2.9 

M2, anteroposterior diameter : greatest 

transverse diameter 3.6 : 3.4 3-9 : 3-6 

Ms, anteroposterior diameter : greatest 

transverse diameter 5-5 : 3-5 

PLESIADAPIS JEPSENI/ new species 
Plate 4 

Type. — Left ramus of mandible with P4-M3, U.S.N.M. No. 20760. 

Horizon and locality. — Bison basin Tiffanian, ledge locality at south 
rim of Bison basin, W^ sec. 28, T. 27 N., R. 95 W., Fremont County, 
Wyo. 

Specific characters. — Close in size to Flesiadapis gidleyi and Plesi- 
adapis anceps. P2 almost always present and hypoconulid portion of 
talonid of M3 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 jepseni 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. 



7 Named for Dr. Glenn L. Jepsen in appreciation of his work on the 
Plesiadapidae. 



24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I3I 

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. jodinatus or the later P. dubius and 
P. rubeyi. 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 M3 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 P2 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' 12.4 

Length of upper molar series, M^-M* 8.5 

P', anteroposterior diameter : greatest transverse 

diameter 2.0 : 2.3 2.1 : 2.5 

P*, anteroposterior diameter : greatest transverse 

diameter 2.2 : 3.1 2.3 : 3.1 

AP, anteroposterior diameter : greatest transverse 

diameter 2.9 : 4.1 2.8 : 4.0 

M*, anteroposterior diameter : greatest transverse 

diameter 3-0 : 4.7 3.0 : 4.6 

M', anteroposterior diameter : greatest transverse 

diameter 2.9 : 3.9 

U.S.N.M. 

No. 20760 U.S.N.M. 

(type) No. 20586 

Length of cheek tooth series, Pi-Ms 12.6 

Length of lower molar series, Mi-AL 10.4 

Pa, anteroposterior diameter : greatest transverse 

diameter 2.4 : 1.9 

P«, anteroposterior diameter : greatest transverse 

diameter 2.4 : 2.8 2.5 : 2.4 



NO. 6 PALEOCENE FAUNAS OF BISON BASIN GAZIN 2$ 

Ml, anteroposterior diameter : greatest transverse 

diameter 3.0 : 3.0 3.2 : 3.0 

Mj, anteroposterior diameter : greatest transverse 

diameter 3-3 : 3-2 3-3 : 3-2 

M3, anteroposterior diameter : greatest transverse 

diameter 4.4 : 2.9 

Carnivora 

ARCTOCYONIDAE 

Creoclonts 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. 

OXYCLAENINAE 

TRICENTES FREMONTENSIS,s new species 

Plate 5, figure 4 

Typc.—Leh ramus of mandible with Mx-Mg, 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 suh- 
trigonus but anterior portion of lower dentition reduced, with trigonid 
of Ml narrow and premolars smaller. 

Discussion. — The above new specific name is proposed with some 
hesitancy, as the Torrejon species, Tricentes subtrigomts, 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 jremo}itensis. Neverthe- 
less, P3 and P4, 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. I3I 

the dozen or more other specimens of T. jremontensis at hand the 
relative narrowness of the anterior molars, particularly the trigonid 
of Ml, 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 Mi), though matched in certain individuals of 
T. subtrigonus, is rather less than the average in the Torrejon ma- 
terials observed. Other specimens of T. jremontensis appear to be 
outside the range of T. subtrigonus in this respect. Moreover, the 
paraconid on M2 and M3 is placed low on the trigonid of lower molars 
in T. jremontensis 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 M3. 

MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMENS OF 

Triccntcs jremontensis 

U.S.N.M. 
No. 20584 

Ps, anteroposterior diameter : transverse diameter 4.0 : 2.4 

P4, anteroposterior diameter : transverse diameter 4.5 : 2.9 

U.S.N.M. 

No. 20582 

(type) 

Ml, anteroposterior diameter : transverse diameter of trigonid 6.2:3.8 

Ml, transverse diameter of talonid 4.5 

Ma, anteroposterior diameter : greatest transverse diameter 6.4 : 4.9 

Ms, 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 P4, in one of the two specimens 
that retains this tooth, is distinctly better developed, as it is in small 
Spanoxyodon latruncidus. 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 2/ 

P4 in U.S.N.M. No. 20983 measures 6.1 mm. long by 3.7 mm. wide. 
Ml 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. pclvidens. 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. 21019, 
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 P4, Mi, and M3, 
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 mctaconid on P4 and the prominently 
isolated hypoconulid on M3 suggest possibly a closer relationship to 
Thryptacodon australis than to T. demari described as new in the 
following section. 

The trigonid of M3 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 M3 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. I3I 

T. antiqiius material exhibiting M3. A slightly more posteriorward 
position for the hypoconulid is indicated also for Mi, 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 

P4, anteroposterior diameter : greatest transverse diameter 5.1:2.9 

Ml, anteroposterior diameter : greatest transverse diameter 6.7 : 4.6 

M3, anteroposterior diameter : greatest transverse diameter 7-1:5.3 



THRYPTACODON DEMARI.o new species 
Plate 6, figures 2, 3 

Type.— Right ramus of mandible with P1-M3, U.S.N.M. No. 20985. 

Horizon and locality. — Bison basin Tiffanian, ledge locality at south 
rim of Bison basin, W^ sec. 28, T. 27 N., R. 95 W., Fremont County, 
Wyo. 

Specific characters. — Size a little smaller than Thryptacodon aus- 
tralis. P2 isolated by longer diastemata. Metaconid of P4 much bet- 
ter developed. Proportions of Mi about as in T. australis but pos- 
terior molars comparatively short and broad. Hypoconulid of M3 
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 Tififany 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. Mi is about the same size as in more nearly con- 
temporary T. australis, but Mo and M3 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 T. 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 M3 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. 
detnari, hence possibly not of diagnostic significance. 

Differences from T. antiquus, in addition to the development of the 
metaconid on P4, include a little less difference in width between M2 
and the trigonid of M3, with M3 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,^o new species 
Plate 6, figures i, 4 
Type. — Right ramus of mandible including M1-M3, 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. I3I 

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 dcmari and lower jaw 
much shallower. Upper and lower molars similar to those in T. de- 
mari, except that M3 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 danari 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 



M\ anteroposterior diameter buccally : transverse 

diameter 5.7 : 6.2 

M", anteroix5sterior diameter buccally : transverse 

diameter 6.5 ": 8.0 



T. demari 
U.S.N.M. 

No. 20984 


T. belli 
U.S.N.M. 
No. 20986 


57 : 6.2 




6.5 "18.0 


5.2:7.1 


U.S.N.M. 

No. 20985 

(type) 


U. of Wyo. 

No. 1045 

(type) 


5-2 : 2.7 




6.7 : 4-3 


5.6:3.8 


6.2 : 5.3 


5.4 : 4.7 


6.2:4.8 


5-3: 40 



P4, anteroposterior diameter : transverse diameter 
Ml, anteroposterior diameter : transverse diameter 
M2, anteroposterior diameter : transverse diameter 
M3, anteroposterior diameter : transverse diameter 
' 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 Clacnodon 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 TALEOCENE FAUNAS OF BISON BASIN — GAZIN 3I 

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 M2 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 Mg 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. aero genius, 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 
Mi-Ms, a jaw portion retaining only P2 and P3, U.S.N.M. No. 21007, 
and an isolated M3. 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, cf. procyonoides (matthew), u.s.n.m. no. 20630 

Length of lower molar series, Mi-Ma 24.0 

Mi, anteroposterior diameter : greatest transverse diameter 77:6.1 

Ma, anteroposterior diameter : greatest transverse diameter 8.0:6.7 

Ma, anteroposterior diameter : greatest transverse diameter 8.7 : 5.6 



32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 inchides 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 difificult its clear separa- 
tion from that represented by the materials referred to C. procyo- 
noides. 

MEASUREMENTS IN MILLIMETERS OF TEETH IN SPECIMEN OF 

Claenodon, of. montanensis (gidley), u.s.n.m. no. 20574 

M3, anteroposterior diameter : greatest transverse diameter 9-5": 7-5 

M3, 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. jerox. 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 M3 preserved rather than M2, 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 



NO. 6 



PALEOCENE FAUNAS OF BISON BASIN — GAZIN 



33 



3 



I 



C. oerogenius n.sp. 



C.cf. 
procyonoides 



C. cf montanensis C. cf. ferox 



7.5 8.0 8.5 9.0 9.5 10.0 105 1 1.0 115 12.0 12.5 13.0 135 14.0 14.5 15.0 



C. ferox 



a: 


6 


UJ 




CD 


5 


5 




-D 


4 


Z 






i 




2 

1 



Cproeyonofdss 



73 7.8 8.3 8.8 9.3 9.8 10.3 108 11.3 11.8 12.3 12.8 13.3 13.8 143 i4B 15.3 

Fig. 2. — Histogram of length of M^ 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. jerox (cope) 

U.S.N. M. 
No. 20797 

M*, anteroposterior diameter : transverse diameter* 10.7 : 15.9 

M', anteroposterior diameter : transverse diameter 8.2 : ii.i 

U.S.N.M. 
No. 20633 

Ma, anteroposterior diameter : transverse diameter 11.4: io-4 

Ms, anteroposterior diameter : transvcr.se diameter 12.7 : 9.1 

• Transverse diameter taken lingually to base of enamel. 



34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

CLAENODON ACROGENIUS," new species 
Plate 7, figures i, 6 

Type.— Right ramus of mandible with Pi, Mi, and M3, U.S.N.M. 
No. 20634. 

Horison 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. P2 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 P4. 

Although the size of the teeth in this material is within, or nearlj 
within, the upper limits of the size range for Claenodon ferox as 
recognized in the Torre j on 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. Pi is close to the canine and 
P2 is missing. A short distance posterior to Pi there is a depression 
that may have been an alveolus for a root of P2. 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 Pi and P3 is very mucli greater than would be 
required for a P2. 

The two molars preserved in this jaw are much worn and do not 
include Mo. 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 Clacnodon acrogcnius, 

U.S.N.M. NO. 20634 

Length of cheek tooth series, posterior margin of alveolus of canine to 

posterior margin of Ms 87.* 

Diastema between Pi and Ps 15.O 

1^ From Greek, akrogcneios =:^f/ith prominent chin. 



NO. 6 PALEOCENE FAUNAS OF BISON BASIN — GAZIN 35 

Depth of jaw beneath diastema between Pi and P3 31.O 

Pi, antcroi)osterior diameter : transverse diameter 3-5" : 2.2 

Ps, anteroposterior diameter at alveoH il.o 

Pi, anteroposterior diameter at alveoli 13.5 

Mi, anteroposterior diameter : transverse diameter 12.2 : 9.7 

Ms, anteroposterior diameter : transverse diameter 15.0' : 9.9 

" Approximate. 
* 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 P4-M2 in Dissactts. In size it is a trifle larger than Mg in D. nava- 
joviiis 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 M2 and the talonid of Mi 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 Mi has proportions almost the 
same as in the type of Didymictis tenuis from the Gidley quarry in 
the Crazy Mountain Fort Union. M2 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. M2 in No. 
1063 measures 2.6 mm. long by 1.4 mm. wide. The talonid of Mi is 
about 1.5 mm. wide. 



36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

CONDYLARTHRA 

HYOPSODONTIDAE 

PROMIOCLAENDS PIPIRINGOSI/^ new species 

Plate II, figures i, 2 

Type. — Right ramus of mandible with Mi and M2, U.S.N.M. 
No. 20571. 

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. — Close in size of molar teeth to Promioclaenus 
lemuroides (Matthew), but lower premolars noticeably smaller. Pre- 
molars simple and but slightly inflated. P2 and P3 without parastylid 
and without talonid cusps or crest. P4 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 Mi and M2 well 
defined. 

Discussion. — In addition to the type, a fragmentary jaw with P2-P4 
(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 Promioclaenus 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- 
loniiis of the Montana Fort Union. The lower premolars, if No. 21 021 
is correctly referred, are small and not so inflated as in P. lemuroides, 
though less slender than in P. aquilonius. Moreover, the anterior pre- 
molars are without parastylid or any talonid cusps. P4, 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 37 

P. acolytus and P. Icmuroides. 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 OP 

Promioclaenus pipiringosi 

U.S.N.M. 

No. 2 1 02 I 

P2, anteroposterior diameter : greatest transverse diameter 2.7 : 1.6 

Ps, anteroposterior diameter : greatest transverse diameter 3.3 : 2.1 

P4, anteroposterior diameter : greatest transverse diameter 3.8 : 2.8 

U.S.N.M. 

No. 20571 

(type) 

Ml, anteroposterior diameter : greatest transverse diameter 4.1 : 3.4 

M:, anteroposterior diameter : greatest transverse diameter 4.4 : 37 

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 M2 and part of the trigonid as 
well as the roots of M3. The form is close in size to Litaletes disjunc- 
tiis 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 M2, though somewhat worn, suggest a lower crowned tooth 
as in Promioclaenus, distinctly less inflated than in Mioclaemis. 

LITOMYLUS SCAPHICUS,!^ new species 
Plate 8, figures 2, 4 

Type. — Right ramus of mandible with M2 and M3, U.S.N.M. 
No. 21014. 

Horizon and locality. — Bison basin Tiflfanian, 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 



^^ Skaphikos, from Greek j/;o/>/joj = anything hollowed out as a basin, in allu- 
sion to the Bison basin. 



38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

scaphiciis, 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 P3, Mi and M2, U.S.N.M. 
No. 21010. 

Horizon and locality. — Bison basin Tiffanian, ledge locality at south 
rim of Bison basin, W^ 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. scaphiciis, P3 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 Mi and part of M2, 
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 Mj), 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 IMomylus 

L. scaphicus L. scaphiscus 

U.S.N.M. U.S.N.M. 

No. 21014 No. 21010 

(type) (type) 

Ps, anteroposterior diameter : greatest transverse 

diameter 3-7'-^'-(>5 

P4, anteroposterior diameter : greatest transverse 

diameter 3-8 * : — 

^* 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 

Ml, anteroposterior diameter : greatest transverse 

diameter 3.2 : 2.2 

Mi, anteroposterior diameter : greatest transverse 

diameter 3-6 '■ 3^- 3-3 '• ^-5 

Ms, anteroposterior diameter : greatest transverse 

diameter 3.8 : 2.6 

' Estimated. 

HAPLALETES PELICATUS,i5 new species 
Plate 9, figure i 

Type. — Left ramus of mandible with P3-M3, 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 P4 weaker. Paraconid weak on Mi, vestigial or absent 
on M2 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 P4 is not so prominent. Haplaletes 
is rather distinctive among condylarths, and the posterior molars, 
particularly M3, look surprisingly like these teeth in Thryptacodon. 
The first molars, however, bear little resemblance. 

Comparison of Haplaletes pelicatus with Haplaletes diniinutimis 
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 SERI0R,i6 new species 
Plate 9, figure 2 
Type. — Left ramus of mandible with Mj and M3, U. of Wyo. 
No. 1078. 

^'' From Greek, pelike = hasin, in allusion to the Bison basin. 
" Serior = later, with reference to its stratigraphic position. 



40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 M2 and M3 vestigial. Talonid basin shallow. Hypo- 
conulid of M3 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) 

Ps, anteroposterior diameter : greatest transverse 

diameter 2.6° : 1.6° 

P4, anteroposterior diameter : greatest transverse 

diameter 3-3 : 2.0 

Ml, anteroposterior diameter : greatest transverse 

diameter 30 : 2.3 

Ma, anteroposterior diameter : greatest transverse 

diameter 3.2 : 2.8 37 : 3-4 

M3, anteroposterior diameter : greatest transverse 

diameter 3-3:2.6 37: 3-1 

" Approximate. 

PROTOSELENE? NOVISSIMUS,i' new species 
Plate 8, figures i, 3 
Type.—Ltii ramus of mandible with Mo 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 Noi'issimns = 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 M2 and M3 
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 Mi (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. Mg in the type 
specimen measures 5.4 mm. long by 3.8 mm. wide, M3 is 5.5 by 
3.2 mm. 

The isolated Mi 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. Mi measures 5.6 mm. long by 3.6 mm. wide. 

LITOLESTES LACUNATUS.is new species 
Plate II, figures 3, 4 

Type. — Left ramus of mandible with P4 and Mi, 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 Liiolestcs 
notissimus in size of lower molars and well outside the range given by 
Simpson (1937a). Lower premolars relatively larger. P.i with small 
anterolingual parastylid, metaconid weak and close to primary cusp, 
and two small talonid cuspules very close together. Paraconid weak 
or absent on M2 and M3. Metaconid and protoconid about equal on 
Ml and M2, 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 

^8 Lacivnatus = hollowed out, with reference to the Bison basin. 



42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I3I 

species of this Tiffanian genus. The genotype Litolestcs ignotus Jep- 
sen is from the Silver Coulee horizon in the Polecat Bench series and 
L. nofissimiis from the Melville portion of the Crazy Mountain Fort 
Union. Litolcstes 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 Mi and M2 (U. of Wyo. No. 1083) 
and a jaw fragment with Ms and part of M2 (U. of Wyo. No. 1079). 
A lower jaw portion with P3, P4, and a much worn Mi (U. of Wyo. 
No. 1059) from the saddle locality may represent this species, but 
P4 is lower and wider and lacks any evidence of a metaconid ; more- 
over, the details of Mi are rather obscured by wear. A jaw fragment 
with only P4 (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 P4 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. 3 1 01 6 

(type) 

Ft, anteroposterior diameter : greatest transverse diameter 3.2 : 1.9 

Ml, anteroposterior diameter : greatest transverse diameter 2.7:2.0 

U. of Wyo. 
No. 1079 

Mj, transverse diameter of talonid 2.2 

Ms, anteroposterior diameter : greatest transverse diameter 2.2:1.6 

PHENACODONTIDAE 

GIDLEYINA WYOMINGENSIS, new species 
Plate 9, figures 3, 4 

Type. — Right ramus of mandible with P3-M1, U.S.N.M. No. 20790. 

Horizon and locality. — Bison basin TifTanian, locality at west end 
of Bison basin, N^ sec. 29, T. 27 N., R. 95 W., Fremont County, 
Wyo. 



NO, 6 PALEOCENE FAUNAS OF BISON BASIN — GAZIN 43 

Specific characters. — Size of P4 and molars close to that in Gid- 
leyina silherlingi. P3 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 zvyomingensis 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 Ecfocion. Comparison of the Tif- 
fanian materials with the genotype of Ecfocion, E. oshornianum, 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. 
P4 in both the Montana and Wyoming Gidleyina material would ap- 
pear distinctive when compared with Eocene material of Ecfocion, 
principally in that the trigonid is elongate in comparison with the 
talonid length, whereas in E. oshornianum 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. oshornianum the talonid of P4 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- 
herlingi is a synonym of G. montanensis. Simpson (1937b) called 



44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 gidleyi 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. gidleyi 
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. 2079s 

Length of upper molar series, M^-M' 20.1 

M\ anteroposterior diameter : greatest transverse diameter 7.4: 9.6* 

M*, anteroposterior diameter : greatest transverse diameter 7.2:10.4 

M', anteroposterior diameter : greatest transverse diameter 5.7: 8.4 

U.S.N.M. 

No. 20790 U.S.N.M. 

(type) No. 20793 

Pj, anteroposterior diameter : greatest transverse 

diameter 5-8 : 3.5 

P4, anteroposterior diameter : greatest transverse 

diameter 6.8 : 4.7 

Ml, anteroposterior diameter : greatest transverse 

diameter 6.5 : 5.2 6.6 : 5.2 

Mj, anteroposterior diameter : greatest transverse 

diameter 6.8 : 5.6 

Ma, transverse diameter of trigonid 4.8 

" Approximate. 

PHENACODUS? BISONENSIS,'" 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. 

1^ Named for the Bison basin. 



NO. 6 PALEOCENE FAUNAS OF BISON BASIN — GAZIN 45 

Specific characters. — Size very close to that of Phenacodus ahnien- 
sis, about intermediate between that of PJienacodus? grangeri and 
Phenacodus? rnattheivi 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 P4 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 P4 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. I3I 

to the primary cusp, and in this respect is certainly distinct on a specific 
level from the Clarkforkian materials, such as Phenacodiis 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 Tetraclacnodon 
puerccnsis. 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 

Phenacodiis ? 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 

diameter 9.4 : ii.o 

M'', anteroposterior diameter : greatest transverse 

diameter 9.4:12.8 

U.S.N.M. U.S.N.M. 

No. 20567 No. 20569 

Length of lower molar series 28.7 

Pa, anteroposterior diameter : greatest transverse 

diameter 7.7 : 4.6 

P4, anteroposterior diameter : greatest transverse 

diameter 8.8 : 5.7 

Ml, anteroposterior diameter : greatest transverse 

diameter 8.9:7.6 8.2:7.3 

M2, anteroposterior diameter : greatest transverse 

diameter 9-5 " : 74 9.6 : 8.2 

Ms, anteroposterior diameter : greatest transverse 

diameter 10.5:7.2 

" Approximate. 

PHENACODUS? sp. (large) 

Plate 10, figures 4, 5 

A fragmentary right mandibular ramus with Mg, U.S.N.M. 
No. 21025, ^^ 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. puerccnsis and P. primac- 
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. 
M3 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 

CORYPHODONTIDAE 

TITANOIDES PRIMAEVTJS Gidley, 1917 

Plate II, 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 Titanoidcs 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 Titanoidcs 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 

-^ Jepscn'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 seeu to belong 
beyond doubt to Titanoidcs primaevus. 



48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 AP 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 of upper molar series, M^-M' 66.8 

Ml, anteroposterior diameter : transverse diameter * 20.8 : 23.3 

M2, anteroposterior diameter : transverse diameter * 25.3 : 27.9 

Ms, anteroposterior diameter : greatest transverse diameter 20.4 : 34.0 

* Anteroposterior diameters of Mj and Ma taken across outer styles and transverse 
diameters perpendicular to line between outer styles. Anteroposterior diameter of M3 taken 
perpendicular to anterior face. 

CAEN0LAMBDA,2i new genus 

Type. — Caenolamhda 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 Barylamhda-Haplolamhda group, but with molars IVP 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 "Sparactolamhda" 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, -|- lambda, the Greek letter — named in 
analogy with Paiitolamhda, 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 Titatioides, 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 IVP 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 M^ 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 pattersoni 
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 difiference 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 PATTERS0NI,22 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 qiiinni 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 determinal)le 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. 2IO36, 

TYPE SPECIMEN OF CacHolambda pattersoni 

Length of skull from the anterior margin of premaxillae to posterior 

margin of occipital condyles 320. 

Length from anterior margin of canine alveolus to posterior margin 

of occipital condyles 300. 

Distance from posterior margin of palate at posterior narial aperture 

to posterior margin of occipital condyles 170. 

Width across postorbital processes no. 

Width across nasals about midway of length 24. 

Length of upper dentition, C (at alveolus) to M^, incl 135. 

Lengtli of upix;r check tooth series, P' to M^ incl 92. 

Lengtii of upper molar series, M^ to M' 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 13.5 : 23.5 

M\ anteroposterior diameter : transverse diameter 18.5 : 27.0 

M', anteroposterior diameter : transverse diameter 19.0 : 30.0 

M", anteroposterior diameter : transverse diameter 16.0 : — 

* Measurements are nearly all approximate 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 1 

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52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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NO. 6 PALEOCENE FAUNAS OF BISON BASIN — GAZIN 53 

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EXPLANATION OF PLATES 

Plate i 

multituberculates and insectivores from the bison basin paleocene 

Fig. I. Cf. Ptilodus montanus Douglass: Left P* (U.S.N.M. No. 20877), lateral 

view. Four times natural size. 
Fig. 2. Cf. Ectypodus hazcni Jepsen: Left P* (U.S.N.M. No. 20878), lateral 

view. Four times natural size. 
Fig. 3. Cf. Ectypodus vmscidus Matthew and Granger : Left ramus of mandible 

with Ml (U. of Wyo. No. 11 05), occlusal view. Four times natural size. 
Fig. 4. Cf. Anconodon russelli (Simpson) : Right ramus of mandible with P* 

(U. of Wyo. No. 1065), lateral view. Four times natural size. 
Fig. 5. Bisoiialveus hrozmi, 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. I, 2. Pronothodectes, cf. matthcim Gidley: i. 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. jodinatus Jepsen: Left ramus of mandible (U.S.N.M. 
No. 20784), lateral and occlusal views. Three times natural size. 

Figs. 4, 5. Peradcctcs 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. 1 104), lingual and occlusal views. Four times natural size. 

Plate 3 
pronothodectes from the bison basin paleocene 

Figs. I, 3. Pronothodectes simpsmn, new species : i. 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. Plesiadapis jepseni, new species: I, 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. I, 2. Chriacus, near C. pehndcns (Cope) : i, 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. Chriaciis, sp: Left M^? (U.S.N.M. No. 21019), occlusal view. Two and 

one-half times natural size. 
Fig. 4. Tricentes jremontensis, 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 

tiiryptacodon from the bison basin paleocene 

Fig. I. Thryptacodon belli, new species: Right ramus of mandible (U. of Wyo. 

No. 1045), type specimen, lateral and occlusal views. Twice natural size. 
Fig. 2. Thryptacodon, cf. dcmari, 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. I, 6. Claenodon acrogenius, new species: i. 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.^L 
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. I3I 

Plate 8 
litomylus and protoselene? from the bison basin paleocene 

Figs. I, 3. Protoselene? novissimus, new si)ecies: i, 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. I. Haplalctcs 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. Haplalctcs scrior, new species : Left ramus of mandible (U. of Wyo. 

No. 1078), type specimen, lateral and occlusal views. Four times natural 

size. 
Figs. 3, 4. Gidlcyina Ziryomingensis, 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. Phenacodtis? bisonoisis, new species: i. Right ramus of mandible 
(U.S.N.M. No. 20567), lateral and occlusal views, li times natural size; 
2, right maxilla (U.S.N.M. No. 20564), type specimen, occlusal view, li 
times natural size; 3, right maxilla (U.S.N.M. No. 20566), occlusal view, 
twice natural size. 

Figs. 4, 5. Pheuacodus? sp. (large) : 4, Right ramus of mandible (U.S.N.M. 
No. 21025), occlusal view, i^ 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. I, 2. Promioclaciius pipiringosi, new species : i. Right ramus of mandible 
(U.S.N.^L 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. Litolestcs lacunalus, new si>ecies : 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. Tiianoidcs Prittwci'iis Gidley : Riglit 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 15 

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). 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. ni. NO. 6. PL. 1 




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VOL. 131. NO. 6. PL. 2 






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Primates and Marsupials from the Bison basin Paleocene 

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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 



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CUAENODON FROM THE BISON BASIN PALEOCENE 

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SMITHSONIAN 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, PL. 13 




CAENOLAMBDA FROM THE BlSON BASIN PALEOCENE 

(SEE EXPLANATION AT END OF TEXT.) 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL 131. NO. 6. PU 14 




Caenolambda from the Bison Basin Paleocene 

(see explanation at end of text.) 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 




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VOL. 131. NO. 5. PL. 16 









Two Fossil localities in the Bison Basin 

(SEE EXPLANATION AT END OF TEXT.) 



SMITHSONIAN MISCKLl.ANEOUS COLLECTIONS 
VOLUME 131, NUMBER 7 



Cf^arlefi! ®. anl> iSlarp ¥^aux OTalcott 



THE UPPER PALEOCENE MAMMALIA 

FROM THE ALMY FORMATION IN 

WESTERN WYOMING 

(With 2 Plates) 



By 
C. LEWIS GAZIN 

Curator, Division of Vertebrate Paleontology 

United States National Museum 

Smithsonian Institution 




(PuBj-.'CAnoN 4252) 



CITY OK WASHINGTON 

PUBLISHED BY THi SMITHSONIAN INSTITUTION 

JULY 31, 1956 



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



CONTENTS 

Page 

Introduction and history of investigation I 

Geologic relations and occurrence of remains 2 

The Almy fauna 3 

Age and correlation of the fauna 3 

Systematic description of the mammalian remains 4 

Primates 4 

Plesiadapidae 4 

Carpolestidae 7 

Carnivora 8 

Arctocyonidae 8 

Mesonychidae 9 

Miacidae 9 

Condylarthra 10 

Phenacodontidae 10 

Dinocerata 16 

Uintatheriidae 16 

References 17 

Explanation of plates 18 

ILLUSTRATIONS 
Plates 



(Following p. 18) 

1. Primates, Anacodon? and Probathyopsis? from the Almy Paleocene. 

2. Condylarths from the Almy Paleocene. 



Cfjarlesf B. anb ittarp ^aux Maltott i^efiearcfj jFunb 

THE UPPER PALEOCENE MAMMALIA FROM 

THE ALMY FORMATION IN 

WESTERN WYOMING 

By C. lewis GAZIN 

Curator, Division of Vertebrate Paleontology 

United States National Museum 

Smithsonian Institution 

(With 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 Qarkforkian 
upper Paleocene age interpreted for the scant, earlier materials nov^ 
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 NW:iNE^ 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 Plcsiadapis 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. I3I 

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 
almieiisis, 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 : 

Plesiadapis rubeyi Gazin 2 

Plesiadapis cookei Jepsen 3 

Plesiadapis? pearcei, new species 2 

Carpolestidae : 

Carpolestes, cf . dubiiis Jepsen I 

Cabnivora : 

Arctocyonidae : 

Anacodon ? nexus, new species i 

Mesonychidae : 

Dissacus, sp 2 

Miacidae : 

Didymictis?, sp I 

CONDYLARTHRA : 

Phenacodontidae : 

Ectocion ralstonensis Granger 13 

Ectocion, cf. oshornianuin (Cope) 4 

Phenacodus almiensis Gazin 32 

Phenacodus primaevus Cope 10* 

DiNOCERATA : 

Uintatheriidae : 

Probathyopsis? , sp 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. I3I 

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 cookei, 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, Haplomyliis speirianns, 
and a species of Coryphodon. 

The further evidence given the distinctiveness of Qark 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 I, 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 P3-M2, but 
lacking the trigonid of Mi, and a left M^ tentatively referred to 
Plesiadapis riiheyi. 

P. rubeyi clearly belongs to the group of species that includes 
P. gidleyi, P. fodinahis, P. dubius, and probably P. cookei. 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. fodhiatus; however, in view of the distinctly small size 
of Ml, the anteroposteriorly shorter appearing summit of the trigonid 
of IM2, and the comparatively slender premolars showing an incipient 
metaconid on P4 (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 

Plcsiadapis rubeyi, u.s.n.m. no. 16696 

Ps, anteroposterior diameter : transverse diameter 2.8: 1.9 

P4, anteroposterior diameter : transverse diameter 2.9 : 2.2 

Ml, transverse diameter of talonid 2.7 

Ml, anteroposterior diameter : transverse diameter of talonid 3-7 '• 3-2 

PLESIADAPIS COOKEI Jepsen, 1930 

Plate I, 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 

1941 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 
M3, 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 SPECTMENS OF 

Plasiadapis cookei 

U.S.N.M. U.S.N.M. 

No. 16698 No. 2078s 

P», anteroposterior diameter : transverse diameter... 5-2: 3-7 

P«, anteroposterior diameter 5.3 

Ml, anteroposterior diameter : transverse diameter of 

talonid 6.1 : 5-6 

Mj, anteroposterior diameter : transverse diameter of 

talonid 6.4 : 5.5 6.4 : 6.3 

Ma, anteroposterior diameter : transverse diameter of 

trigonid lo.S" : . . . lo.o" : 6.2 

" Approximate. 

PLESIADAPIS? PEARCEI,2 new species 
Plate I, figure 9 

Type.— Right ramus of mandible (U.S.N.M. No. 20787), with Mi 
and M2. 

Horizon and locality. — Buckman Hollow Clarkforkian Paleocene 
on La Barge Creek, NW^NE:^ 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 Mi and M2 distinctly more posterolingual in position. 

Discussion. — The two lower jaw portions considered to represent 
this peculiarly distinctive form were at first allocated to P. rubeyi 
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. 



NO. 7 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 Mi the paraconid is farther forward. In 
M2, 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. jepsenl-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 

Ml, anteroposterior diameter 3.1 3,2 

Ml, transverse diameter of trigonid 2.1 2.0 

Ml, transverse diameter of talonid 2.5 2.5 

Ma, anteroposterior diameter 3.4 

Mj, transverse diameter of trigonid 2.4 

Mj, transverse diameter of talonid 2.8 

CARPOLESTIDAE 

CARPOLESTES, cf. DUBIUS Jepsen, 1930 

Plate I, figure 4 

A carpolestid P4 (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 haselae. 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 P4 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 P4 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,3 new species 

Plate I, figure i 

Type.—Leit ramus of mandible (U.S.N.M. No. 21282) with Mi 
and M2. 

Horizon and locality. — Buckman Hollow Clark forkian Paleocene 
on La Barge Creek, NW^NE^ 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- 
tanemis. Primary cusp pattern of lower molars better defined than 
in Anacodon ursidens or A. cidtridens, 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 Mi as 
well as M2. 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 

^ ncxtis (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 
Cluenodon montanensis. Nevertheless, the lovi^er 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 Aiiacodon? 
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. 

Ml in No. 21282 of Anacodon? nexus measures approximately 
9.5 mm. in length by 7.8 mm. across the talonid. Mo 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 P2 or P3, 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 Ml may well belong to a species of Didymictis, but is very 
much smaller than contemporary Didymictis protenus proteus from 



10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

the Clarkforkian of the Big Horn Basin. It is only a Httle 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 i 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 Pg-Ma 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 II 

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 Phenacodtis, 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 Mi. 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 Pi has but one root. 

The essential difference between Ectocion ralstonensis, as exempli- 
fied by No. 20736, and the Ectocion osborniamim 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 



12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

No. 20736, but it may be moderately developed on the cingulum of 
P* and sometimes P^ in E. oshornianum. 

The lower premolars appear relatively more slender in Ectocion 
ralstonensis than in E. oshornianum, 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. oshornianum ; also E. ral- 
stonensis exhibits a shallower mandible. 

Upper and lower molars of Ectocion ralstonensis are apparently not 
distinctive in comparison with E. oshornianum, 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. silherlingi 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. 



MEASXTREMENTS 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 margin of M' 49.7 

Length of upper cheek tooth series, P*-M', inclusive 41.8 

Length of upper premolar series, P*-P*, inclusive 24.0 

Length of upper molar series, M*-M^ inclusive 18.0 

C, anteroposterior diameter at alveolus : transverse diameter at alveolus. 4.1 : 2.8 

P', anteroposterior diameter : transverse diameter 3.0 : 1.5 

P^ anteroposterior diameter : transverse diameter 4.0 : 2.2 

P^ anteroposterior diameter : transverse diameter 5-7 : 5-S 

P*, anteroposterior diameter : transverse diameter 6.0 : 7.3 

M\ anteroposterior diameter : transverse diameter across anterior por- 
tion 6.2 : 8.5 

A'P, anteroposterior diameter : transverse diameter across anterior por- 
tion 6.2 : 9.1 

M^ anteroposterior diameter : greatest transverse diameter 4-9 : 7-S 

Length of lower cheek tooth series Pi (at alveolus )-M3, inclusive 42.8 

Length of lower premolar series. Pi (at alveolus) -P4, inclusive 23.3 

Length of lower molar series, M1-M3, inclusive 20.0 

P3, anteroposterior diameter : transverse diameter 5.5 : 3.2 

Pi, anteroposterior diameter : transverse diameter 6.7 : 4.1 

Ml, anteroposterior diameter : transverse diameter of talonid 6.5 : 5.1 

M2, anteroposterior diameter : transverse diameter of trigonid 6.5 : 5.4 

Ma, anteroposterior diameter : transverse diameter of trigonid 6.7 : 4.5 



NO. 7 MAMMALIA FROM THE ALMY FORMATION GAZIN I3 

ECTOCION, cf. OSBORNIANUM (Cope), 1882 

A right lower jaw fragment with P4-M2 (U.S.N.M. No. 20645) 
has teeth more robust than in the jaw belonging to the E. ralstonensis 
skull, and in addition P4 is more progressive with a better developed 
talonid basin and a large entoconid. P4, moreover, has a rather dis- 
tinctly developed paraconid or parastylid. This specimen is tentatively 
regarded as representing the Gray Bull species E. oshornianum. 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. oshorn- 
ianum, Simpson (1937b), in his treatment of the material, believed 
(except for rare E. parvus) that a single species was represented in 
which there w-as a shift in the mean size, the length of Mi 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 P4 in the larger 
Almy specimen. The character of P4 might likewise show marked 
variation within a species, but there are three small-toothed or 
E. ralstonensis specimens which have P4 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 

P4, anteroposterior diameter : transverse diameter of talonid 7.5 : 5.1 

Ml, anteroposterior diameter : transverse diameter of trigonid 6.8 : 5.9 

M2, anteroposterior diameter : transverse diameter of trigonid 7.1 : 6.5 

PHENACODUS ALMIENSIS Gazin, 1942 

Plate 2, figures 3 and 4 

A relatively small species of Phenacodus, P. ahniensis, 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 
wnth the canines and P^ to M^ in a scarcely worn state, together with 
certain limb and vertebral portions. Although collected in 194 1, it 
remains after six subsequent collecting trips the best specimen of this 
species extant. 



14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

P. ahnietisis is much smaller than Phettacodiis p. intermedius but 
appreciably larger than P. copei. It differs essentially from the Phena- 
codiis 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. ahniensis 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. ahniensis. Granger (191 5) noted that 
the metaconules were weak or absent in P. copei. These are appar- 
ently not reduced in P. ahniensis. Moreover, the upper premolars, 
strangely enough, appear more advanced than in P. copei. P^ has a 
well-defined and separate tritocone, described as weak in P. copei, 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. ahniensis is distinctly larger than 
P. mattheivi, 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 bisonensis. 

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 I5 

MEASUREMENTS OF UPPER TEETH IN SPECIMENS OF 

Phenacodus alniiensis 



Length of cheek tooth series, P*-M*, inclusive 

Length of molar series, M^-M", inclusive 

P^ anteroposterior diameter : transverse diameter . . 
P*, anteroposterior diameter : transverse diameter.. 
M\ anteroposterior diameter : transverse diameter * 
M^ anteroposterior diameter : transverse diameter * 
M^ anteroposterior diameter : transverse diameter * 

• Approximate. 

* Anteroposterior diameter of upper molars taken perpendicular to anterior margin and 
transverse diameter across anterior portion. 



U.S.N.M. 

No. 1 669 1 

Type 


U.S.N.M. 
No. 21286 


42.7" 


41.0" 


26.1" 


26.7 


8.2:7.4 


7.7": 8.6 


8.5:8.5 


7.9 : 9.2 


9.0: 10.0 


9.0: II.O' 


9-2: 1 1.3 


8.8:12.5' 


7.7 : 10.8 


8.7" : 10.5' 



PHENACODUS PRIMAEVUS Cope, 1873 
Plate 2, figure 5 

Tvi^o 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 P3 to M2, inclusive, and the other an incomplete lower molar. 
The teeth in No. 21287 ^-^e comparable in size to those in the Qark 
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 intermedins. 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- 
viedius. 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. intermedins 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. 



p. p.. cf. 
intermedins 
U.S.N.M. 
No. 20644 


P.p. 

prtmaevus 
U.S.N.M. 
No. 21287 


II-5 


12.7 


7.8 


10.2 


11.8 


I2.I 


9.8 


1 1.2 




I2.S' 




12.5 



16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

MEASUREMENTS IN MILLIMETERS OF CERTAIN LOWER TEETH IN JAWS OF 

Phenacodus 

p. ahnii^nsis 
U.S.N.M. 
No. 20643 

P4, anteroposterior diameter 9.6 

P4, transverse diameter of talonid 6.2 

Ml, anteroposterior diameter 9.1 

Ml, transverse diameter of talonid.... 7.4 

M2, anteroposterior diameter 

M2, transverse diameter of trigonid 

" Approximate. 

DINOCERATA 
UINTATHERIIDAE 

PROBATHYOPSIS?, sp. 
Plate I, 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 flssi- 
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 I7 

REFERENCES 
Cope, Edward 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. i. 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, pis. 1-7. 
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, pis. 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, pis. 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. 329- 
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, pis. i-io. 
Matthew, William D. 

1915a. A revision of the lower Eocene Wasatch and Wind River faunas. 
Part I. — Order Ferae (Carnivora), Suborder Creodonta. Bull. 
Amer. Mus. Nat. Hist., vol. 34, art. i, 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, pi. 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. lOO-III. 

ScHULTz, Alfred R. 

1 914. Geology and geography of a portion of Lincoln County, Wyoming. 
U. S. Geol. Surv., Bull. 543, pp. 1-141, figs. 1-8, pis. i-ii. 
Simpson, George G. 

1928. A new mammalian fauna from the Fort Union of southern Montana. 
Amer. Mus. Nov., No. 297, pp. 1-15, figs. 1-14. 



l8 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. 1-3. 

1935. The Tiffany fauna, upper Paleocene. 3. — Primates, Carnivora, Con- 

dylarthra, and Amblypoda. Amer. Mus. Nov., No. 817, pp. 1-28, 
figs. 1-14. 

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, 

pis. I-IO. 
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?, and probathyopsis ? from the almy paleocene 

Fig. I. Anacodon? nexus, new species: Mi and M2 (U.S.N.M. No. 21282), 

type specimen, occlusal view. Natural size. 
Figs. 2 and 3. Probathyopsisf, 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 : P* (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 riibcyi Gazin: Right ramus of mandible (U.S.N.M. No. 

16696), type specimen, occlusal and lingual views. Three times natural 

size. 

Plate 2 
condylakths from the almy paleocene 

Figs. I and 2. Ectocion ralstonensis Granger: i. 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 (Mi restored from right 
side). Natural size. 

Figs. 3 and 4. Phenacodus almicnsis 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 



■^x*.-*^ 



J 




A' 



^^^ 



vt 



> 



«t* 



■<fr- 





ai 



^^mB 



> 







10 



Primates. Anacodon?, and Probathyopsis? from the 

ALMY PALEOCENE 

(SEE EXPLANATION AT END OF TEXT.) 



SMITHSONIAN MISCELLAMEOUS COLLECTIONS 



VOL. 131, NO. 7, PL. 2 






CONDYLARTHS FROM THE AUMY PAUEOCENE 

(SEE EXPLANATION AT END OF TEXT.) 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 8 



Cjjarles; M. anb ilarp "^aux OTalcott 
l^esiearci) Jf unb 



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 
BADWATER AREA 

(With 3 Plates) 



By 
C. LEWIS GAZIN 

Curator, Division of Vertebrate Paleontology 

United States National Museum 

Smithsonian Institution 




(Publication 4257) 

CITY OF WASHINGTON 

PUBLISHED BY THE SMITHSONIAN INSTITUTION 

OCTOBER 30, 1956 



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



CONTENTS 

Page 

Introduction i 

Acknowledgments 2 

History of investigation 2 

Occurrence and preservation of material 3 

The Badwater fauna 4 

Correlation and age of the fauna 5 

Systematic description of the Mammalia g 

Alarsupialia ? 9 

Didelphidae ? 9 

Lagomorpha 9 

Leporidae 9 

Rodentia 10 

Carnivora 10 

Limnocyonidae 10 

Miacidae 10 

Condylarthra 11 

Hyopsodontidae 11 

Perissodactyla 12 

Equidae 12 

Brontotheriidae 13 

Chalicotheriidac 13 

Helaletidae 16 

Hyracodontidae 23 

Artiodactyla 23 

Dichobunidae 23 

Agriochoeridae 26 

Oromerycidae 30 

Leptomerycidae 30 

References 31 

Explanation of plates 34 

ILLUSTRATIONS 

Plates 

(All plates follow page 35.) 

1. 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 

Page 

I. Suggested phylogenetic arrangement of North American tapiroids 16 



CfjarlefiS B. anb iHarp IJaux iiaUott J^esearcfj jFunb 

THE GEOLOGY AND VERTEBRATE PA- 
LEONTOLOGY OF UPPER EOCENE 
STRATA IN THE NORTHEASTERN 
PART OF THE WIND RIVER 
BASIN, WYOMING 

PART 2} 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 Badwatcr 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 I 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, early 
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 1-3 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 Ainynodon advcnus, 
Tchnatherium, cf. cidtridcns, 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 TelmatJierium, 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 
Bad water 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 2^ and 3^ miles almost due northwest of the site of the now 
abandoned Badwater Post Ofifice. 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 i 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 liadwater Creek, 
are about 20 miles due west and include the sites for the Malaquiferus 
and Eomoropus skulls, in the NW^ 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 Diplobimops and the others are of Protoreodon, Malaquiferiis, and 
Eomoropiis. 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 i 
of this paper. 

THE BADWATER FAUNA 

Marsupialia?: 
Didelphidae ? : 

Peratheriumf, sp. 
Lagomorpha: 
Leporidae : 

Mytonolagus zvyomingensis A. E. Wood 

RODENTIA * : 

Paramyidae : 

Rapamys?, sp. 

Sciuravus diihius A. E. Wood 

Paramyid indet. (large) 

Paramyid indet. (small) 
Eomyidae : 

Protadjidauntof, sp. 
Cricetidae : 

Cricetid indet. 
Carnivora : 

Limnocyonidae : 

Limnocyon?, sp. 
Miacidae : 

Miacis, cf. robustiis (Peterson) 
Condylartha : 

Hyopsodontidac : 

Hyopsodus, cf. nintensis Osborn 
Peris sodactyla : 
Equidae : 

Epihippus, cf. gracilis (Marsh) 

Epihippus, cf. pan-US Granger 
Brontotheriidae : 

Brontotheriid indet. 
Chalicotheriidae: 

Eomoropiis auarsius, new species 



NO. 8 MAMMALIAN FAUNA, BADVVATER AREA — GAZIN 

Helaletidae : 

Desmafotherimn zvoodi, new species 

Dilophodon, cf. Icotanus (Peterson) 
Hyracodontidae : 

Epitriplopusf, sp. 
Amynodontidae* : 

Amynodon advetius (Marsh) 
Artiodactyla : 
Dichobunidae : 

Pcntaccmylusf, sp. 

Apricitlus praeteritits, new genus and species 
Agriochoeridae : 

Protorcodon, cf. pctcrsoni Gazin 

Protoreodon, near P. pmniius (Marsh) 

Protoreodon pcarcei, new species 

Diplobunops, cf. inatthewi Peterson 
Oromerycidae : 

Malaquiferiis tourteloti Gazin 
Leptomerycidae : 

Leptotraguliis, cf. mcdius Peterson 

Leptorcodonf, 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 Protadjidanmo 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 di.stinct from that of Myton, including it ^ (Gazin, 
1955, chart I ) 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. 131 

Protadjidamno, only Epihippus, Amynodon, and Protoreodon are 
common to the two, and the genera Mytonolagus, Rapamys, Eomoro- 
pus, Epitriplopus, Poitacemylus, Diplobttnops, Leptotragulus, Lep- 
toreodon, and possibly Desmatotheriiim and Dilophodon are charac- 
teristic of the Uintan. Some of these ahnost 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 Sciiiravtis, 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- 
tadjidamno, 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. Protadjidamno 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. mutthemi 
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 7 

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. H, 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 Protapirits. 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 advcnus 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 Chasmothcroides, cf . intcrmcdius 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. I3I 

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 Ictops and is unlike that, for example, in M2 of Metacodon 
mellingeri where the hypoconulid is closely connected to the entoconid. 
For these reasons I have tentatively assigned this specimen to Pera- 
theriumf, sp. 

LAGOMORPHA 
LEPORIDAE 

MYTONOLAGUS WYOMINGENSIS Wood, 1949 

Plate I, figure i 

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^-]\P 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 M2 and particularly Mi 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. petersoni 
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. 
wyomingensis as perhaps more primitive than M. petersoni. 



NO. 8 MAMMALIAN FAUNA, BADWATER AREA GAZIN 



RODENTIA 

Description of the known Roclentia 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 I, figure 2 

A comparatively large miacid is represented by a lower jaw exhibit- 
ing the teeth P4 to Mg inclusive. Miacis would appear to be indicated 
by the distinctly basined form of the relatively small talonid of Mo. 
The talonid of Mi 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 M2, 
short and decidedly narrower than the trigonid. Mg 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 P4 and Mi as subequal in size 
so that P4 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 Mi and Mo likewise suggests M. 
robustus. 

The type of Miacis iiintensis Osborn (1895) fro"i Uinta B would 
appear from the scale of Osborn's illustration to be a little shorter 



10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

in length of cheek teeth and distinctly shallower jawed than the Bad- 
water form. Moreover, P4 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. Mx is relatively smaller, and M2 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 P4 is evidently characterized by two posterior cusps in ad- 
dition to the cingulum, 

MEASUREMENTS IN MILLIMETERS OF LOWER TEETH 

IN Miacis, cf. robiistiis, u.s.n.m. no. 21087 

P4, anteroposterior diameter : transverse diameter 10.5 : 5.4 

Ml, anteroposterior diameter : transverse diameter of trigonid 13.0 : 7.5 

M2, anteroposterior diameter : transverse diameter of trigonid 5.6 : 4.3 

CONDYLARTHRA 

HYOPSODONTIDAE 

HYOPSODUS, cf. UINTENSIS Osborn, 1902 

Plate I, figure 3 

A single Hyopsodus upper molar, U.S.N.M. No. 21089, """^.y well 
represent H. nintcnsis, 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 

3 Personal communication. 



NO. 8 MAMMALIAN FAUNA, BADWATER AREA — GAZIN II 

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 I, figure s 

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 Mo, 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 Mcsohippus. 
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 (Mi) in the type of Epihippus uintcnsis (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 (Duchesnchippus) intermedins. The V-shaped crests of 
the lower molar, however, are a little less acute than in the Duchesncan 
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 I, 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. 



12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

P" in the specimen compared with E. 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 Bad water 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. 24097. 

* Anarsios (Gr.), incongruous, strange — in allusion to the large and unex- 
pected canine. 



NO. 8 MAMMALIAN FAUNA, BADWATER AREA — GAZIN I3 

Horizon and locality. — Ilendiy Ranch member of Tepee Trail for- 
mation on Dry Creek, S\L\ sec. 9, near line between sees. 9 and 16, 
T. 39 N., R. 92 W., Wind River Basin, Wyo. 

Specific characters. — Teeth only slightly larger the& 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. yy) 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 i. 



14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 

by wear of Mi (compare height of cusps shown in lateral view, pi. 2, 
fig. I, with Osborn's fig. 3A on p. 267, 1913). 

Unlike the type of E. amarorum, the cranial portion of the E. 
anarsnis 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 

Eomoropiis anarsius, u.s.n.m, no. 21097 

Length of upper molar series, M'-M' inclusive, parallel to tooth row 54.4 

Ml, anteroposterior diameter perpendicular to anterior margin 14.9 

Ml, greatest transverse diameter across parastyle and protocone 16.7 

M2, anteroposterior diameter perpendicular to anterior margin 19.0 

M:, greatest transverse diameter across parastyle and protocone 23.5° 

M3, anteroposterior diameter perpendicular to anterior margin 19.6 

Ma, greatest transverse diameter across parastyle and protocone 25.0 

Length of preserved lower cheek tooth series, P3-M3, inclusive 78.0 

Length of lower premolars, P3-P4 24.0 

Length of lower molars, Mi- Ms 54-0 

P3, anteroposterior diameter : transverse diameter of talonid 12.8 : 7.8 

Pi, anteroposterior diameter: transverse diameter of talonid 11.2: 8.1 

Ml, anteroposterior diameter : transverse diameter of talonid 13.9 : 9.0 

Mj, 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. Eomoropiis 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. AP 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 AP and no significant differences are observed. Neverthe- 
less, the anteroexternal root for M- and AP 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 anncctcns, which Peterson (1919) 



NO. 8 MAMMALIAN FAUNA, BADWATER AREA — GAZIN I5 

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 E. anarsius are quite similar to those in E. 
amaronim and about the same size, although the anterior premolars 
may be relatively a little larger. E. amaronim includes all the lower 
cheek teeth from P2 to M3, inclusive. In No. 21097 only P2 of this 
series is missing, although the inner walls of Mo and M3 are not com- 
plete. As Osborn has shown, these teeth are much like those in 
Moropus; however, in the earlier form P2 is a relatively larger tooth 
and M3 retains a prominent hypoconulid. Moreover, as observed in 
occlusal view of the Dry Creek specimen, the crista obliqua in all the 
cheek teeth following P2 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. 

HELALETIDAE 

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 guyotii 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." 



i6 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 




liiEioct horiton unc«rla> 



Fig. I. — Suggested phylogenetic arrangement of North American Eocene 

tapiroids. 



NO. 8 MAMMALIAN FAUNA, BADVVATER AREA — GAZIN I7 

Scott's generic comparisons were made largely with Hyrachyus, 
from which it clearly differs. I am convinced, however, that a distinctly 
closer relationship to Hclaletes is indicated. The Badwater, Sage 
Creek, and Washakie materials together provide the evidence showing 
the sequence from Bridgerian Hclaletes through Uintan Desmato- 
therimn 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,e new species 
Plate 2, figure 4 

Type.— Right maxilla with P3-I\P (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 guyotii Scott or Desmatotherium kayi 
Hough. Upper premolars smaller than in these species but relative 
size intermediate between them and closer to D. guyotii. 

Discussion. — Four 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. zvoodi. Two of these, 
part of the type and two upper premolars (part of U.S.N.M. No. 
20202) were figured by Hough (1955, pi. 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. "guyotii" 
but with smaller premolars. The premolars of D. zvoodi are a little 
smaller than in D. kayi, but the ratio of their size to that of the molars 
is more nearly as in £>. guyotii. 

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 

" Named for H. E. Wood, II, in appreciation of his work on the Plyrachyidae. 



l8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

teeth a distinctly more molariform appearance. Resemblance is im- 
mediately seen to the earlier Hclaletcs nanus (genotype) which in a 
similar way is distinguished from Helaletes boops. Washakie D. 
guyotii 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, pi. 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. guyotii 
is comparable to Colodon oecidentalis 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. 
guyotii and only certain isolated teeth and tooth fragments are included 
in the materials of D. zvoodi. 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 IMa 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- 
clusive 33.3 

P\ anteroposterior diameter : greatest trans- 
verse diameter 7.5 : 6.0 

P", anteroposterior diameter 8.3 : — 

P^ anteroposterior diameter : greatest trans- 
verse diameter 9.0: 11.7 8.9: 11.8 

P*, anteroposterior diameter : greatest trans- 
verse diameter 8.8:12.4 9.2:12.7 9.1:12.6 



Length of upper molar series, parallel to 

tooth row 37.5 

Ml, anteroposterior diameter perpendicular 

to anterior margin 11 .0 

Ml, transverse diameter across parastyle and 

protocone 14.0 

M«, anteroposterior diameter perpendicular 

to anterior margin 12.9 

M2, transverse diameter across parastyle and 

protocone 15.8 

M3, anteroposterior diameter perpendicular 

to anterior margin 13.7 

M3, transverse diameter across parastyle and 

protocone 15.6 



II.O 

13.8 



U.S.N.M. 

No. 

20204 



12.8 

15.0 

13-0 
iS.i 



Colodon, and this may logically include Heptodon in the lower Eocene 
which, in addition to a much reduced Pi, 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. I3I 



DILOPHODON Scott, 1883 

Dilophodon was described by Scott (1883) in tbe same publication 
as Desmatotherium and, as in the case of D, giiyotii, the type of 
Dilophodon minusculiis 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. minuscidus 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 hoops, 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 I, 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 
P4," 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^-]VP 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 Ml and IMg. These teeth in No. 20207 ^r^ 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 minuscnhis, 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 Desinatotherium line is 
noted in the anterofx^sterior 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. 

Dilopliodon 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, P2 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 IVP, 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 boops, not divided as in the D esmatotherium-C olodon 
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 Hcptodon material would not be excluded as 
potentially ancestral. 



22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

The combination of characters seen in the upper dentition is highly- 
suggestive of Prota pirns 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 

D., cf. 

Icotanus 
U.S.N.M. 

No. 
21098 

Length of upper dentition, P'-M'' inclusive 39-0 ' 

Length of upper premolars, P^-P* inclusive, at alveoli 17.0* 

Length of upper molar series, M' (at alveolus) -M^ inclusive 23.2 

P', anteroposterior diameter : transverse diameter 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., cf. 

cuius Icotanus 

P.U. No. U.S.N.M. 

1 00 1 9 No. 

Type 20207 

Length of lower cheek tooth series, P2-M3 inclusive 46.6 41. i 

Length of lower premolar series, P2-P4 inclusive 18.7 16.2 

Length of lower molar series, M1-M3 inclusive 28.3 25.4 

P2, anteroposterior diameter : greatest transverse diam- 
eter 5-6 : 3-4 4-8 : 3-5 

P3, anteroposterior diameter : greatest transverse diam- 
eter 6.5 : 4-6 57 : 4-5 

Pi, anterioposterior diameter : greatest transverse diam- 
eter 6.8 : 5.5 6.1 : 5-2 

Ml, anteroposterior diameter : transverse diameter of tri- 

gonid 8.1:5.2 7-2: 5-2 

Mt, anteroposterior diameter : transverse diameter of tri- 

gonid 94: 5-9 8.8:6.1 

M3, anteroposterior diameter : transverse diameter of tri- 

gonid 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. Icotanus. 



NO. 8 MAMMALIAN FAUNA, BADWATER AREA GAZIN 23 

HYRACODONTIDAE 

EPITRIPLOPUS?, sp. 

Plate I, 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 Epitriplopiis. The Badwater teeth are closer in size to 
those in Epitriplopiis iiintense 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 Mcsamynodon mcdiiis. The lower tooth in No. 21099 measures 
16.2 mm. long by ii.i mm. wide. 

Dr. H. E. Wood concurs with me that, of the various possible alloca- 
tions which may be made of this material, Epitriplopiis is the most 
probable. 

ARTIODACTYLA 
DICHOBUNIDAE 

PANTACEMYLUS?, sp. 

A homacodont that may well be Pentacemyliis is represented by the 
posterior two-thirds of M3. 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 Bunomeryx. 

APRICULUS,^ new genus 

Type. — Apricuhis praeteritus, new species. 

Generic characters. — Simple conical cusps on upper molars as in 
Helohyiis, 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 Helohyiis. P* with single primary 
cusp and deuterocone. 

Discussion. — Apriciilus is almost certainly a descendant of Bridg- 
erian Helohyiis and belongs in the Helohyinae, but its trend has been 

'' Apriculns, diminutive of /Ipcr (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 Hngual position. This is particularly 
noticeable for IVP which, rather than having a triangular outline, has 
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 Perchoerns; 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 Apricidiis so far as known, but in the retention and 
marked development of the protoconule. 

APRICULUS PRAETERITUS,8 new species 
Plate 3, figure I 

Type.— Right maxilla with P^-M^* (teeth incomplete), U.S.N.M. 
No. 21 100. 

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-Wke 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 praetcritus 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 scries, P*-IVP inclusive, is 
found to be 30 mm. The molar series is about 24.5 mm. long. J\P 
is estimated to be about 8.7 (anteroposteriorly) by 10.6 mm. 

In size, upper molars of Apriculus praetcritus are only a little larger 
than the dichobunid tooth figured by Russell and Wickenden (1933). 



8 Praetcritus, overlooked or passed over — overlooked in recent review of col- 
lections for upi)cr 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 Helohyusf, 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 b}' 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 Mi, 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 Mi. 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 low^er 
molars of Archaeotherium. In a footnote Peterson called attention to 
the resemblance of P4 to that in Archaeotherium, but discredited such 
a relationship by the presence of a paraconid on Mi, 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 

" 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. 



26 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 Helohyiis 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 Lophiohyiis. 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}^ The type is the lower jaw portion described by Peterson, CM. 
No. 11912.^2 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. 

^^ Dyskritos (Gr.), hard to determine or doubtful, and choiros (Gr.), pig. 
^1 lapointensis, 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 2'] 

These teeth are all of about the size of those in Uintan P . puniilus. 
There is, moreover, no certain evidence that another species such as 
Protoreodon primus is not represented. 

PROTOREODON PEARCEI,i3 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, SW-j 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 
puniilus, very close in size to Diplohunops 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 puniilus 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. matthewi. 

The skull of Protoreodon pearcei differs rather noticeably from that 
of Diplobunops, cf. matthewi in the Badwater collection (see Gazin, 
1955, pis. 10-12) in the anterior extremity of the rostrum. Although 
the canines are actually larger and as far apart in P. pearcei, 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 deutcrocone 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 AP. 

The lower jaw is not so constricted through the symphysial portion 
as it is in the Diplobunops material figured by Scott (1945, pi. 5, fig. 

13 Named for Franklin L. Pearce, who found the type specimen. 



28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

2). Pi 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. peorcei are a little nar- 
rower than in D., cf. viatthewi. 

Proforeodon pearcei makes a close approach to Agriochoeriis 
antiqmis in size and in the presence of a short though distinct 
diastema between the upper canine and P^. There is, however, no 
diastema between Pi and P2. Moreover, the posterior premolars above 
and below are not nearly so progressive. Also, the protoconule, though 
very weak on ]\P, 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 Pi and P2 may represent P. pearcei 
rather than Diplobunops matthewi. 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, pi. 8, fig. 8) as "Mesagriochoerus, 
cf. primus" and the catalog number is incorrectly cited as "ciSzy." 

Measurements for the teeth in the type of P. pearcei are given with 
those for Diplobunops, cf. mattheivi 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 
a])undant 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 Plarry A. 
Tourtelot, one of these, U.S.N.M. No. 20303, has been previously 
figured (Gazin, 1955, pis. 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 pearcei and Diplobtinops, CF. matthezvi 

D., cf. matthezvi P. pearcei 

, * , U.S.N.M. 

U.S.N.M. U.S.N.M. No. 

No. No. 2030s 

20304 20303 Type 
Length of upper dentition, C (at alveolus) - 

M^ inclusive 96.5 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 39.5 40.0" 39.6 

C, anteroposterior diameter (at alveolus) : 

greatest transverse diameter 7.7:7.1 11.0:8.7 

P*, anteroposterior diameter : greatest trans- 
verse diameter 9.0 : 3.8 7.7 : 4.3 

P', anteroposterior diameter : greatest trans- 
verse diameter 9.6:5.9 10.3:6.4 11.1:6.5° 

P^ anteroposterior diameter : transverse di- 
ameter * 10.2:10.2 10.3:9.8 11.1:9.5 

P^ anteroposterior diameter : transverse di- 
ameter 10.1:12.3 10.0:12.9 10.2:12.5 

M\ anteroposterior diameter : transverse di- 
ameter* 12.8:15.1 12.0": — 12.5:13.5 

M', anteroposterior diameter : transverse di- 
ameter 13.8:16.6 13.8:17.5 13.7:16.6 

^r, anteroposterior diameter : transverse di- 
ameter 13.7:18.0 14.0": 19.5" 15.0:18.3 

Length of lower cheek tooth series. Pi (at 

alveolus) -M3, inclusive 82.5 

Length of lower premolar series, Pi (at 

alveolus)-P4, inclusive 43.2 38.3 

Length of lower molar series, M1-M3, in- 
clusive 44.0 

Pi, anteroposterior diameter (at alveolus) : 

greatest transverse diameter lo.o : 8.3 

Ps, anteroposterior diameter : greatest trans- 
verse diameter 10.0:5.3 9.5:4.6 

P3, anteroposterior diameter : greatest trans- 
verse diameter 11.0:7.0 10.8:6.2 

Pi, anteroposterior diameter : greatest trans- 
verse diameter 12.2:8.0 11.5:7.2 

Mi, anteroposterior diameter : transverse di- 
ameter of talonid 10.9 : 8.1 

Ms, anteroposterior diameter : transverse di- 
ameter of talonid 14.4:11.3'' i3-0'':9.3 

M3, anteroposterior diameter : transverse di- 
ameter of trigonid — : 10.7 20.3 : 10.4 

" 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 tootb, lingually to base of enamel or cingulum. 



30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 

toreodon pearcei 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 
Protorcodon 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, pi. 16), and need 
not be repeated here. An isolated oromerycid upper molar (U.S.N.M. 
No. 21 102) in the Badwater collection exhibits the same rectangular 
form, rugosity, posteriorly bifurcate protocone, and outstanding ribs 
on the outer cusps as in Malaquiferoiis tourteloti. It corresponds 
closely in form and size, and in the possession of a slightly outward- 
deflected metastyle to IVP. It differs in having a heavier, antero- 
posteriorly developed mesostyle. The shape of this tooth is entirely 
different, although structurally related to that in Oromcryx plicatus. 
Likewise, it would not be confused with the comparatively large 
Protylopusf 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 Eoinoropus. 

LEPTOMERYCIDAE 

LEPTOTRAGULUS, cf. MEDIUS Peterson, 1919 

Plate 3, figure S 

Two lower jaw portions, and probably several of the isolated molars, 
represent a form close or identical to the upper Uintan Lcptotragulus 



NO. 8 MAMMALIAN FAUNA, BADWATER AREA — GAZIN 3I 

medius. One of the jaw portions (U.S.N.M. No. 21 104) with P4 and 
Ml falls within the size range of the Myton material, but the other 
(U.S.N.M. No. 21 103), which has P3-M1, 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 Lcptotraguhis. 

The teeth in both Nos. 21 103 and 21 104 are significantly smaller 
than in the type of Leptotragiilus? 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 Lepfomeryx 
rather than Lcptotragulus. 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 diflferent in size from Lepto- 
traguhis 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 
Lepfomeryx. Nevertheless, as in Leptoreodon and unlike Leptomeryx, 
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 P4 are in most cases sharply separated and join forward with the 
protoconid and metaconid respectively. The second isolated P4 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. 

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in S.E. Wyoming, pp. 53-67. 
1953. Geology of the Badwater area. U. S. Geol. Surv. Oil and Gas. Invest. 
Map OAI124 (2 sheets). 
Wood, Albert E. 

1949. Small mammals from the uppermost Eocene (Duchesnean) near 

Badwater, Wyoming. Journ. Paleont., vol. 23, No. 5, pp. 556-565, 
figs. 1-24. 
Wood, Horace E., II. 

1934. Revision of the Hyrachyidae. Bull. Amer. Mus. Nat. Hist., vol. 67, 
pp. 181-295, figs. 1-51, pis. 20-24. 
Wood, 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, Jacob 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 

i-agomorph, carnivore, condylarth, and perissodactyls from 
the badwater upper eocene 

Fig. I. Mytonolagus wyom'mgensis A. E. Wood: Right maxilla (U.S.N.M. 

No. 21090), occlusal view of teeth. 4 x natural size. Badwater upper 

Eocene, Wind River Basin, Wyo. 
Fig. 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. 
Fig. 3. Hyopsodus, cf. uintensis Osborn: Right upper molar (U.S.N.M. No. 

21089), occlusal view. 4 x natural size. Badwater upper Eocene, Wind 

River Basin, Wyo. 
Fig. 4. Epitriplopus? , sp. : Left lower molar (U.S.N.M. No. 21099), occlusal 

view. Natural size. Badwater upper Eocene, Wind River Basin, Wyo. 
Fig. 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. 
Fig. 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. 
Figs. 7, 8. Dilophodon, cf. leotainis (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 

Figs. 1-3. Eomoropus anarsius, new species: i, 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. 

Fig. 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 

Fig. I. ^/TiCK/K.? /Tflf/rn'/HJ, new genus and species : Right maxilla (U.S.N.M. 
No. 21 100), type specimen, occlusal view. Twice natural size. Badwater 
upper Eocene, Wind River Basin, Wyo. 



NO. 8 MAMMALIAN FAUNA, BADWATER AREA — GAZIN 35 

Fig. 2. Malaquiferus tonrtcloti Gazin : Right upper molar (U.S.N.M. No. 

21 102), occlusal view. Twice natural size. Badwater upper Eocene, Wind 

River Basin, Wyo. 
Fig. 3. Leptoreodonf, sp. : Left P4 (U.S.N.M. No. 21105), occlusal view. 

Twice natural size. Badwater upper Eocene, Wind River Basin, Wyo. 
Fig. 4. Protoi-eodon, cf. pctcrsoni Gazin: Right maxilla (U.S.N.M. No. 21101), 

occlusal view. Twice natural size. Badwater upper Eocene, Wind River 

Basin, Wyo. 
Fig. 5. Leptotragulus, cf . mcdius Peterson : Right ramus of mandible 

(U.S.N.M. No. 21 103), occlusal and lateral views. Twice natural size and 

natural size, respectively. Badwater upper Eocene, Wind River Basin, Wyo. 
Fig. 6. Diplobnnops, cf. mafthewi 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. 
Figs. 7, 8. Protoreodon pcarcci, 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 (Pi reversed from left side). Natural size, Badwater upper Eocene, 

Wind River Basin, Wyo. 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 8, PL. 1 






J' " J 






'i y 





Lagomorph, carnivore. Condylarth, and Perissodactyls from 

THE BADWATER upper EOCENE 

(.See cxplanatinii nf jilates at end of text.) 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 8, PL. 2 



^m:. 



f 







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 




'i^* /. 



m 




ARTIODACTYLS FROM THE BADWATER UPPER EOCENE 

(See explanation of plates at end of text.) 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 131, NUMBER 9 



BREEDING AND OTHER HABITS OF CASQUED 
HORNBILLS (BYCANISTES SUBCYLINDRICUS) 

(With 6 Plates) 



By 
LAWRENCE KILHAM 

Bethesda, Md. 






(Publication 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 suhcylindricits (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. 

Acknozvledginents. — 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. 

L.K. 



CONTENTS 

Page 

Preface iii 

Introduction I 

Methods 2 

General habits of hornbills 4 

Nest construction 8 

Activities of nesting pairs 17 

Eggs, young, and nest openings 20 

Territory, and relations of hornbills with one another 24 

Relations with other birds 29 

Food 29 

Some anatomical features in relation to function 32 

Discussion of factors controlling hornbill populations 34 

Comparative studies of other hornbills 35 

Discussion of hornbill biology 40 

Summary 44 

References 45 



BREEDING AND OTHER HABITS OF CASQUED 
HORNBILLS (BYCANISTES SUBCYLINDRTCUS) 

By Lawrence Kilham 

Bethesda, Md. 

(With 6 Plates) 

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 MISCELLAMEOUS COLLECTIONS, VOL. 131, NO. 9 



2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 2^ miles long. An African 
ranger and his assistants who lived at the headquarters clearing (pi. i, 
fig. i) 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 (Psittaciis erithaciis) . 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 schiittii), run- 
ning squirrel-like along high branches, were quiet and difficult to find. 
Two other, smaller hornbills {Tockus alboterminatiis 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 {Stcpha- 
noaetus coronatus) . The harrier hawk (Polyboroides typiis), 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 melanoleiiciis) 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 {Ceuthmo chares aereiis), 
and Narina's trogon {Apaloderma narina). Among mammals, troops 
of redtail monkeys {Cercopitheciis 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 (i nest); shrill screaming of female hornbill from behind 
wall when her nest was approached by foreign hornbills (i nest) ; 
commotion of a group of hornbills surrounding an eagle (i nest). 
Two other methods involved observations of a male bird: When on 
a direct, purposeful flight into the forest (i nest) ; and following a 
bird after it picked up dirt from the ground (i 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 schweinfiirthii 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 
(pi. I, 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 (pi. 2, fig. i), 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. I3I 

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 
(pi. 2, fig. i) and a female (pi. 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 10 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 zvitJi sticks and bill zvhacking. — 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 I 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. I3I 

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 i 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 I 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 i. 
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 i, 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. 

Gatheriny dirt for bidlding at nest i. — I learned the most about wall 
building from a pair of hornbills in Mpanga Forest. Their hole was 



10 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



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SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 9, PL. 1 




I. HeadquarUTs clearing, Nfpanoa Research Poorest. 




rmtir^fm 



2. ^'ol^l.t; male liornl)!!! > (la\.> alter eiiier.:;iii,i; iruiu iil>i 5. 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 9, PL. 2 




1. ".M i>aii.ua," liaii(l-n.-arLd male li(irnl)ill lo moiith.s ot a^e. 




2. "Zika." liaiid-i-oared fcnialf lidrnhill lo iiuimli> of Hiiu. 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 9, PL. 3 




I. Li)cati()n ( sIkiwu hy arrow) of nest 13 in stub of broken limb 




Xcst 5 (location sbou n by arrows) in I 'il'Uuliiii<i tr(.-c in 
Jiotanical Gardens, luitebl)e. 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 9, PL. 4 




T. 'rcniiitc nuiund at lu-adquarUr^ clcarinj.;, Ali>anga KoL-arch Foix-^l, 
where female hornbill collected earth. 




2. l--nti-;iiuf ''I luM III, --hnwing ceiiieiit wall 



NO. 9 CASQUED HORNBILLS — KILHAM II 

ideally located. It was 70 feet up in the trunk of a huge tree {Celtis 
saoyouxii), 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 




Fig. I. — 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. I3I 



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. 




Fig. 2. — Nest i, 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 I3 

lined by soldiers, showed where the hornbill had collected earth. On 
October 27 the male again visited the stump. At 12 130 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 I 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 :o8 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 i. 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. I3I 

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 :30 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, i 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 zvall. — 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 i 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 I955> 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 i 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 



l6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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, i^ to 2 inches wide and 4 to 10 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 1 7 

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 lo. 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 schwcinfurthii) 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 (pi. 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 7 130 a.m. 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 



l8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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, i 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 1 1 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 9 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 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 9, PL. 5 







I, |-\'iiiaK- iKiiiiliill at nine ^lu.• \\a- rcniDved from nest lo, appruxiniately 
twD-tliirds of the wav tlir(Higli nesting period. 







z. "Mpanga" and "Zika" wluii aiii»r(i\inialel\ -' months of age. 



SMITHSONIAN MISCELLANEOUS COLLECTIONS 



VOL. 131, NO. 9, PL. 6 




I. "Mpanga" at f) niontlis of age. 




2. Young crowned liornbill (Tockiis alhotcrjiiiitatus). 
(Pliotograpli by Dr. Herbert Friedmann.) 



NO. 9 CASQUED HORNBILLS — KILHAM I9 

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 10 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. I3I 

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 lo 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 i 130 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 scaflfold 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 &gg 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 21 

It was January i, 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, 1 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 i. 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 i 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 ill had 



22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 i, 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 
400 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 i, 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, 11, 
12, and 13, table i) all opened between January i 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 iiiotJier 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 
(pi. I, 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. I3I 

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 i, figure 2, shows his appearance 2 days after 
leaving the nest. I estimated that he was 10 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 i until May 15, when we left, a pair of hornbills came 
every day, often remaining fur 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. H 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. I3I 

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 9, 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 10 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 2/ 

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 
15 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. I3I 

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 2g 

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 Canarium 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 HORNEILL NESTS 

Canarium schweinfurthii Engl. Aniiaris toxicaria (Pers.) Lesch. 

Pycnanthus angolensis (y\[tbfi.) 'E-x.dX. Chlorophora excelsa (Wehv.) Benth. 

Pseudospondias microcarpa (A. Rich.) Eugenia jatnbolana Lam. 

Engl. Dracaena steudneri Engl. 

Maesopsis eminii Engl. Ficus nalalcnsis ITochst. 

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 
{Lyhiiis bidentatus) was excavating in a tree when the hornbills flew 



NO. 9 CASQUED HORNBILLS — KILHAM 3I 

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 

Rhyparohia grandis (Sauss.) — large cockroach 

Long-horn cricket 

Tenebrionid beetle, Mctallonolus 

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 i 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. I3I 

from the ground. The hornbills made a clapping noise as they snapped 
at the passing termites and were at the game for over lo minutes. 
On May i 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 splendidns), 
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, i^ 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 lo 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 lamelligeriis). 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 
(pi. 5, fig. 2) of two young hornbills, male and female, at 7 to 8 
weeks of age. Plate 6, figure i, shows Mpanga's bill at 6 months of 
age and plate 2, figure i, 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. I3I 

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 
10 and 16 (table i) 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. Byconistes 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- 
w^ard. Searching the area, I found a leaning tree with a bole, 40 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 Tavcta, 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 i 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 [pi. 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. I3I 

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 [pi. 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 i 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 10 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 (Tockiis 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 scnegalensis) 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 alboterminattts) 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 wdth 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. I3I 

equally well to Bycanistes siibcylindricus. 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 G:gg 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 grufif 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 
w-rites 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 — KILIIAM 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 lo 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 casf|ued 
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. I3I 

fashion, the side of the end portion . . . beating against the surface, 
to which the moist dung is appHed." 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 ( i ) 
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- 
natiis, presumably hold for casqued hornbills. 

Ground hornbills. — I 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 zvoof woof with every stroke. The call note is a deep 
boom boovt, 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, "yoxi ahvays say that; boom, you ahvays 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. I3I 

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 suhcylindricus. 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 hrevis just after the 
female had emerged from her morning's work and about 10 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 i 
the pair tried for weeks to close the opening. Interest, however, began 
to fall oft' a week after coition, a situation that paralleled one de- 
scribed by Moreau (1936) in Usambara. Pairs of Bycanistes hrevis 
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 

(Aqtiila zvahlbergi) 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 II 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 stud3^ 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. I3I 

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 sitbcylin- 
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 w^ith 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 hombill 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 119 ±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 vicinit}' 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 hombills 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 

B.\NNERMAN, D. A. 

1953- The birds of West and Equatorial Africa. 2 vols., 1,526 pp., 144 figs., 
54 pis. 
Browx, Leslie H. 

1952. On the biology of the large birds of prey of the Embu district, Kenya 
Colony. Ibis, vol. 94, pp. 577-620. 
Mackworth-Praed, Cyril W., and Grant, C. H. B. 

1952. Birds of eastern and northeastern Africa, xxv + 836 pp., 53 col. pis., 
6 black-and-white pis., 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. i 
and 2, pp. 1-28. 
Ranger, G. 

1949a. Life of the crowned hornbill, Lophoceros suahelicus attstralts. 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. 



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 




(Publication 4260) 



CITY OF WASHINGTON 

PUBLISHED BY THE SMITHSONIAN INSTITUTION 

OCTOBER 17, 1956 



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



CONTENTS 

Page 

Introduction i 

I. Evolution of the arthropods 6 

II. The nauplius and the metanaupHus 9 

III. Examples of crustacean metamorphoses 13 

Branchiopoda 14 

Ostracoda 17 

Copepoda 20 

Cirripedia 35 

Isopoda 46 

Euphausiacea 52 

Decapoda 54 

Stomatopoda 62 

IV. Structure and evolution of arthropod appendages 65 

References 73 



CRUSTACEAN METAMORPHOSES 

By R. E. SNODGRASS 

Collaborator of the Smithsonian Institution and of the 
US. 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, 
w-hen 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 hcteromorphosis, 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. 10 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 
tliere 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 
wuU 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 orthomorphosis. 

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 tgg. 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 consequendy 
most parasites undergo metamorphic changes. jMany 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 



NO. 10 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 s)^mpathy 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. I3I 

ment ; in those having antennal glands they he 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 f oliaceous. 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 out on movable lateral lobes of the segments, the so-called 
parapodia, that serve for swimming and burrowing. 



NO. 10 



CRUSTACEAN METAMORPHOSES — SNODGRASS 



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.i A) the animals resembled a modern onychophoran, and are 




Fig. I. — 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 tasnianiae. 

I Ant, first antenna; 2Ant, second antenna; Mxpd, maxilliped; PIpds, pleo- 
pods; Prpds, pereiopods; Tel, telson; Urpd, uropod; II-XVIII, body segments. 



perhaps represented by such fossils as the Pre-Cambrian Xcnusion 
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. i 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, ""f ollows 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. 
I 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. i 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. 10 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 nauplins, 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 metanaiiplius, 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 (lAnt), typically biramous second antennae 
(2 Ant) and mandibles (Md), and a median eye of two or more parts. 



10 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



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 





rarl 



A 



B 



Fig. 2. — Nauplius and metanauplius of Apus cancriformis Bosc (from Claus, 

1873). 
A, nauplius, with first antennae (lAnt), second antennae (^Ant), and man- 
dibles (Md), rudiments of teloblastic appendages seen through cuticle. B, 
metanauplius, appendages of teloblastic segments (IbSegs) 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. 10 CRUSTACEAN METAMORrilOSES — SNODGRASS II 

Our chief interest in the nauplius is the question of its theoretical 
value in phylogeny. The naupHus has been Hkened 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 grozvth, 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. 2 A). 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 
(tbSegs). 



12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 
(^939) > 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 N eh alia. 

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. 10 CRUSTACEAN METAMORPHOSES — SNODGRASS I3 

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 Nehalia are 
essentially the same as those of Sollaud for the embryo of Leander 
and of Fransemeier for the larva of Arteniia. 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 Nehalia 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 Nehalia. 

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 



14 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 




Fig. 3. — Branchiopoda. Branchinecta occidcntalis 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, 2.9 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 (^Ant). 

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 



l6 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 1 1 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 (1-3) are commonly described as endites, while the large, 
so-called flabellum (<5) 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. 10 CRUSTACEAN METAMORPHOSES — SNODGRASS I7 

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 incongrucns, 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 Cytheridac and in Nesideidae, appendage 5 is a 
typical leg well separated from 4. If, therefore, appendage ./ is 
interpreted as the maxilla, appendages 5, 6, and 7 are thoracic legs, 
and Kesling (1951) says this is now the accepted interpretation of 



i8 



SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 



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. 




/^' A 






4- 5 6 T 7 



Fig. 4. — Ostracoda. Larval stages of Cyprinotus hicongnicns Ramdohr (from 

Schreiber, 1922). 
A, nauplius, with three pairs of appendages. B, second instar, with one pair 
of added appendages {4) and caudal furca (/). 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 



NO. 10 



CRUSTACEAN METAMORPHOSES — SNODGRASS 



19 



lAnt 




Fig. 5. — Ostracocla. 
A, Limnocy there inopinata (Baird), eighth instar (from Scheerer-Ostermycr, 
1940). B, Cypris tcstudinaria Sharpe, adult, left shell removed. C, Philomcdes 
globosa (Lilljeborg), adult, left shell removed. D, Cypris teshidinaria, 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. I3I 

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, ^) is represented in Philomedes (C, F, ^) 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 testiidinaria (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 10 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. 10 



CRUSTACEAN METAMORPHOSES — SNODGRASS 



21 



(H) 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' 



Fig. 6. — Copepoda. Calanus cristatus Kroyer, adult. 
I Ant, first antenna; 2Ant, second antenna; H, "head"; Md, mandible; iMx, 
first maxilla; 2Mx, second maxilla; Mxpd, maxilliped; VI, VII, XI, body 
segments. 



which, elaborated in their own bodies, is thus passed on as food for 
the larger animals. It w'ould 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, 8 A), 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 (Mxpd) and two pairs of legs (/L, 




Fig. 7. — Copepoda. Nauplius and two metanaupliar instars of Cyclops (outlines 
from Ziegelmayer, 1925). 
A, nauplius, ventral, showing three pairs of appendages and labrum {Lni). 
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. 10 



CRUSTACEAN METAMORPHOSES — SNODGRASS 



23 



The genus Calanus, a typical free-swimming marine copepod of 
the suborder Gymnoplca, hkewise develops from nauplius to adult 
by ordinary anamorphic growth without any metamorphic changes 




Fig. 8. — Copepoda: Podoplea. Developmental stages and adult of Cyclops. 
(A-F from Dietrich, 1915; G from Claus, 1863). 

A, Cyclops strenuiis 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. 

I Ant, first antenna; 2Ant, second antenna; iL, 2L, sL, legs; Md, mandible; 
iMx, sMx, first and second maxillae ; Mxpd, ma.xilliped. 



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. I31 

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 diflferent 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. 10 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 Cyniba- 
softia rigidum Thompson, described by Malaquin (1901) as Haemo- 
cera danae (Claparede), 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, R), and on the enlarged conical 
posterior part of the body it becomes armed with circles of spines 
directed forward. Tlie organs of the future adult now gradually de- 
velop within the cuticle of the larva (F), and the abdomen forms as 
a ventral flexure {Ah) of the posterior part of the body. At an early 
stage the first antennae are regenerated (F, G, lAnt) and eventually 
penetrate into the rostrum (I) 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 (I), the parasite becomes strongly 




Fig. 9. — Copepoda: Monstrillidae. Larval stages and adult of Haemocera 
danae (Claparede) parasitic in the polychaete SaJmacina dystcri Huxley (from 
Malaquin, 1901). 

A, free nauplius. B, after penetration into host. C, same, with nutritive ten- 
tacles (0- D, same, later stage. E, later stage, with cuticular envelope, rostrum 
{R), and spines. F, begimiing 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; lAnt, first antenna, ef, egg filaments, gSeg, genital segment, 
Ov, ovary. 

26 



NO. 10 CRUSTACEAN METAMORPHOSES — SNODGRASS 2.y 

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, Ov^ ; 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 (1921a), and a detailed account of the 
larval stages of Caligus ciirtus (O. F. Miiller) 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 chal'muis 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 nonf ceding, 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 chaHmus 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 




Fig. 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, Trcbius latijiircatus Wilson, adult female. F, Blakcanus cornigcr Wilson, 
adult female. G, Hacmobaphes 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. 10 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 chaHmus 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 C aligns cur His (fig. 10 D) 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 (191 5) 
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 amhloplitis (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, /). The young larva has two pairs of feath- 
ery swimming legs, and its maxillipeds (Mxpd) 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. I3I 



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 




Fig. II. — Copepoda : Lernaeopodidae. Developmental stages of fish parasites 
(from Wilson, 1910, 1915). 
A, Achtheres ambloplitis Kellicott, first copepodid, with filament (/) 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. ii 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 3I 

large second maxillae (C, 2Mx) are much thickened and securely 
grasp the base of the attachment filament (/) 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 (/) 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 
hranchialis we may draw on the work of Pedaschenko (1898), Scott 
(1901), Wilson (1917), Schuurmans-Stekhovcn (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 




Fig. 12. — Copepoda. Developmental stages of the fish parasite Lernaeoccra 
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. 
1 1.4 mm. I, adult egg-carrying female on cod, 40 mm. J, section of mature 
female. 

An, anus; AlCnl, alimentary canal; lAnt, first antenna; 2Ant, second an- 
tenna ; CmGld, cement gland ; es, egg string ; Mth, mouth ; 2Mx, second maxilla ; 
Mxpd, maxilliped; Odd, oviduct; Ov, ovary; Sphr, spermatophore. 



NO. 10 CRUSTACEAN METAMORPHOSES — SNODGRASS 33 

Stage, and called this stage a naupHus (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. I3I 

After attachment on the cod the head of the female undergoes a 
curious transformation. Large branching, hornHke processes grow 
out from it and sink into the host tissue as anchoring devices (fig. 
12 I). The proboscisHke 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 (I). The female in 
her final stage is said by Wilson (1917) to attain a length of 40 
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 
(Odd), 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 attaclv the vital inner organs. The worst of them are members 
of the genus Phrixocephaliis, 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. I3I 

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 niira 
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 ( lAnt) 
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 meiacrinicola (fig. 13 B). This species has the 
entire body enclosed in the shell, the abdomen being relatively short, 
but otherwise it is similar to 6*. mira. Okada finds well-dififerentiated 
males and females in 5". 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 
Synngoga 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. 10 



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 




m 


# ■ 


. ■ ■ \y 


i 


-^s>.,.. r,. , 


''^^l.::' 


£, 


^ -'".v. ^ '^ ' ' ■ 


^ 1 


■ ■- '' <4 


_';-j>' ■-■■-- 


- ■•' 



G 



Fig. 13. — Cirripedia: Ascothoracica. (A from Norman, 1913; B from Okada, 
1938; C, D, E from Heegaard, 195 1 ; F, G from Knipowitsch, 1890.) 

A, Sy^iagoga mira Norman, adult. B, Synagoga melacrintcola Okada. C, 
Ascothorax bulbosa Heegaard, shell of female with small male on top, internal 
parasite of ophiuroid. D, same, female. E, same, male. F, Dendrogastcr 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. I3I 

however, is a typical cirriped cypris, much resembling the adult of 
Synagoga (A). A species figured by Fisher (1911) as Dendrogaster 
arbuscuhis, found in a Calif ornian 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 ofT 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 
Runnstrom (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 (lAnt), 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 Rhisocephala. — In this suborder of parasitic cirripeds we en- 
counter the strangest metamorphic phenomena known in the whole 




Fig. 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 ebnrneus 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 Sacciilitui. 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 Sacculhia 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 Saccnlina, 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 Saccnlina (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 naupHus 
becomes a cypris larva (B) with a length of 0.20 mm. On leaving 




Fig. 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, lAnt) , 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 (TJi) 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. 10 CRUSTACEAN METAMORPHOSES — SNODGRASS 4I 

the original cypris (B). Soon a new cuticle is secreted on the surface 
of the sac (D, iCt) 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 (^Ct) 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. i6 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. I3I 




Fig. 16. — Cirripedia: Rhizocephala. Metamorphosis of Sacculina carci>ii Thomp- 
son (from Delage, 1884). 

A, nauplius. B, free-swimming cjpris stage ready to moult. C, cypris fixed 
by antenna at base of hair of crab, shell separated, thorax detached and thrown 
of? with internal tissues. D, larva still in shell has formed a new cuticle (iCt). 
E, shell being shed, larval body retracted witliin cuticle, with long, hollow "dart" 
(d) extended toward base of antenna. F, larva with a second inner cuticle 
i^Ct), the dart extended into antenna. G, larval body expanded, the dart has 
pierced the hair membrane of the crab. 

Ab, abdomen; lAnt, first antenna; iCt, outer cuticle; ^Ct, 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 {he). The figures at C and D, taken from G. Smith 
(1906), depict a species of Peltogastcr, 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) and a 
single nerve ganglion (Gng). 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. I3I 



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. 




Fig. 17. — Cirripedia ; Rhizocepliala. Internal and external parasitic stages. (A-D 
from G. Smith, 1906; E, F from Potts, 1915.) 

A, Sacculina neglecta attached on intestine of crab Inachus scorpio. B, same, 
later stage with root system developed. C, PeUogaster 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; op, external 
opening of brood cavity; Ov, ovary; r, nutritive roots; Tcs, 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. 10 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, I3I 

surface. Thompsonia, unlike Sacculhia, appears to do no specific 
damage to its host, so that it can continue its parasitic Hfe and indefi- 
nitely repeat its reproductive processes. The inoculation of the host 
by the free-swimming cypris has not been observed. 

The Thompsonia-mitsitd 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 Sacciilina 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 of!^er 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, 10 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 {iMxpd) 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 {iMx), supported by the paragnaths {Pgn) beneath them, 
puncture the integument. The much reduced second maxillae {2Mx) 
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, I3I 




Ps-n IMxpd IMx 2Mx Q- 



FiG. 18. — Isopoda: Gnathiidea. Paragnctha formica (Hesse) (from Monod, 

1926) . 

A, first frce-swiinming 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; iMx, first maxilla; sMx, second maxilla; 
Mxpd, 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 10 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. 18 D) 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. 



5© SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I3I 

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 Hfe as a parasite. 

Danalia ciirvata 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 sty li form 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 larv^al 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. 10 



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 




Fig. 19. — Isopoda: Epicaridea. Life history of Danalia airvata 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 ofif with the exuviae except a pair of small 
hooklike second pereiopods {^Prpd) 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 



$2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I31 

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