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THE PLEISTOCENE AVIFAUNA OF THE TALARA TAR SEEPS,
NORTHWESTERN PERU
By
KENNETH EUGENE CAMPBELL, JR.
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY
THE UNIVERSITY OF FLORIDA
1973
ACKNOWLEDGMENTS
I would like to thank the members of my supervisory committee,
Dr. Pierce Brodkorb, Dr. David Johnston, and Dr. Richard Edwards, for
their help and encouragement during the progress of this research and
the preparation of this dissertation. I would especially like to
thank the chairman of my supervisory committee, Dr. Pierce Brodkorb,
for his guidance and critical evaluation of my research and the pre-
paration of this manuscript. His continued interest in my progress
has been an invaluable contribution to this study.
I wish to thank the collector of the fossil material, Dr. A.
Gordon Edmund of the Royal Ontario Museum, Ontario, Canada, for the
opportunity to study the collection. Professor Gustav Orc6s of the
Institute Polyt6chnico, Quito, Ecuador, kindly permitted me to study
the fossil material from La Carolina, Ecuador, and extended many
courtesies to me while I visited his country. Peruvian officials,
particularly Ing. Eduardo Izquierdo C., were most cooperative in
allowing me to collect comparative material. Sr. Max Cerro provided
a welcome base of operations during field work in northwestern Peru.
The following individuals and institutions generously loaned
many specimens for comparative purposes: Dr. Dean Amadon, American
Museum cf Natural History; Dr. N. Philip Ashmole, Yale Peabody
Museum; Dr. Robert M. Mengel, Natural History Museum, University of
Kansas; Dr. Robert W. Storer, Museum of Zoology, University of Mich-
igan; Dr. Glen Woolfenden, University of South Florida; and Dr.
Richard Zusi, United States National Museum.
Special acknowledgment is due the Organization for Tropical
Studies, Inc., for providing field research funds under OTS Pilot
Research Grant N69-23. The Frank M. Chapman Fund, American Museum
of Natural History, and Sigma Xi also provided field research funds.
Without their support this study would be very incomplete.
Special acknowledgment is also due my wife, Terrie, for her
enduring patience and enthusiasm.
To all of these people and many more, I am deeply grateful.
TABLE OF CONTENTS
Page
AKNOWLEDGMENTS .................................................. ii
LIST OF TABLES................................................... v
LIST OF FIGURES................................................ vii
ABSTRACT .................................. ...................... viii
INTRODUCTION..................................................... 1
MATERIALS AND METHODS ........................................... 5
GEOLOGY......................................................... 6
SYSTEMATIC LIST................................................. 10
ECOLOGY.......................................................... 188
Recent Avifauna............................................. 189
Paleoecology............................................... 190
CLIMATOLOGY OF NORTHWESTERN PERU................................ 194
Neoclimatology............................................. 194
Atmospheric Circulation ............................... 194
Oceanic Circulation.................................. 195
Climatological Effects of the Atmospheric and
Oceanic Circulations............................. 197
El Nifno............................................... 199
Paleoclimatology........................................... 202
DISCUSSION AND CONCLUSIONS ...................................... 205
The Equatorial Arid Fauna.................................. 205
Origin of the Equatorial Arid Fauna ................... 205
Speciation Within the Equatorial Arid Fauna........... 207
Dating Some Extinct Species................................ 208
Comparison of Avifaunas of Talara Tar Seeps and
Rancho La Brea ........................................ 209
Dating the Marine Terraces................................. 210
Correlation of Talara Tar Seeps with La Carolina
of Ecuador............................................ 211
LITERATURE CITED ................................................ 213
APPENDIX......................................................... 218
BIOGRAPHICAL SKETCH ............................................. 227
iv
LIST OF TABLES
Table Page
1 List of non-passerine species known from the Talara
Tar Seeps, with number of elements and least number
of individuals of each indicated......................... 11
2 Measurements of the scapula, humerus, radius, femur,
tibiotarsus, and tarsometatarsus of Eudocimus n. sp.
and Recent E. albus, and of the scapula, tibiotarsus,
and tarsometatarsus of E. albus from the Talara Tar
Seeps (in mm) ................... ...................... 30
3 Measurements of the scapula, coracoid, humerus, radius,
ulna, carpometacarpus, femur, tibiotarsus, and tarso-
'metatarsus of Tadorninae, n. gen. n. sp. and Lophonetta
specularioides (in mm)................................... 42
4 Measurements of the scapula, coracoid, humerus, radius,
ulna, carpometacarpus, femur, tibiotarsus, and tarso-
metatarsus of Anas n. sp. 1 (excluding femur), Anas
n. sp. 2, Anas n. sp. 3 (coracoid, humerus, tibio-
tarsus, and tarsometatarsus only), A. leucophrys,
A. brasiliensis, A. cyanoptera, and A. crecca carolin-
ensis (in mm) ............................................ 58
5 Measurements of the scapula, coracoid, humerus, radius,
ulna, carpometacarpus, femur, tibiotarsus, and tarso-
metatarsus of Gymnogyps californianus, Recent Vultur
gryphus, V. gryphus from the Talara Tar Seeps (exclud-
ing scapula, humerus, and carpometacarpus), Gymnogyps
n. sp. (excluding carpometacarpus and femur), and
Vulturidae, n. gen. n. sp. (excluding scapula) (in mm)... 81
6 Measurements of the scapula, coracoid, humerus, radius,
ulna, carpometacarpus, femur, tibiotarsus, and tarso-
metatarsus of Geranoaetus n. sp., Recent G. melano-
leucus, G. melanoleucus from the Talara Tar Seeps
(excluding scapula), Harpyhaliaetus solitaria; and
of the coracoid, carpometacarpus, tibiotarsus, and
tarsometatarsus of Buteoninae?, n. gen. n. sp. (in mm)... 100
7 Measurements of the scapula, coracoid, humerus, radius,
ulna, carpometacarpus, femur, tibiotarsus, and tarso-
metatarsus of Milvago n. sp., M. chimachima, and M.
chimango (in m) .............. .... ... ................... 129
Table Page
8 Measurements of the humerus, coracoid, and femur of
Belonopterus chilensis, B. resplendens, and Hoplox-
ypterus cayanus; of the humerus and femur of Belon-
opterus n. sp.; and of the coracoid of Vanellinae,
n. gen. n. sp. (in mm)................................... 139
9 Measurements of the coracoid and tarsometatarsus of
Tringa n. sp., Recent T. solitaria, and T. solitaria
from the Talara Tar Seeps (in mm)......................... 149
10 Measurements of the femur of Steganopus n. sp.,
Recent S. tricolor, and S. tricolor from the Talara
Tar Seeps (in mm)....................................... 166
11 Measurements of the humerus of Thinocorus n. sp.,
Recent T. rumicivorus, and T. rumicivorus from the
Talara Tar Seeps (in mm) ................................. 171
12 Measurements of the coracoid and carpometacarpus of
Caprimulgus n. sp., C. cayennensis, C. longirostris,
and C. parvulus (in mm) .................................. 187
LIST OF ILLUSTRATIONS
Plate Page
I View of the Talara Tar Seeps, northwestern Peru,
looking north........................................... 4
II Two views of the Talara Tar Seeps........................ 9
III Holotypes of Eudocimus n. sp., Theristicus n. sp.,
Syrigma n. sp., Tadorninae, n. gen. n. sp., and
Anas n. sp. 1........................................... 220
IV Holotypes of Vulturidae, n. gen. n. sp., Sarcoramphus?
n. sp., Anas n. sp. 2, Anas n. sp. 3, and Milvago n. sp. 222
V Holotypes of Gymnogyps n. sp., Belonopterus n. sp.,
Tringa n. sp., and Buteoninae?, n. gen. n. sp ........... 224
VI Holotypes of Geranoaetus n. sp., Vanellinae, n. gen.
n. sp., Scolopacidae, n. gen. n. sp., Thinocorus n. sp.,
Steganopus n. sp., and Caprimulgus n. sp................ 226
Abstract of Dissertation Presented to the
Graduate Council of the University of Florida in Partial
Fulfillment of the Requirements for the Degree of Doctor of Philosophy
THE PLEISTOCENE AVIFAUNA OF THE TALARA TAR SEEPS,
NORTHWESTERN PERU
By
Kenneth Eugene Campbell, Jr.
June, 1973
Chairman: Dr. Pierce Brodkorb
Major Department: Department of Zoology
The Talara Tar Seeps are a series of fossil-bearing deposits
located in northwestern Peru. They have been dated at approximately
14,000 years B.P. and lie directly on a marine terrace thought to be
Sangamon in age.
The avifauna is known from 6200 elements representing a minimum
of 742 individuals of 13 orders, 24 families, 66 genera, and 88 species.
Five new genera and 20 new species are described. The following
families are represented, with the number of identified species for
each (numbers of new species in parenthesis): Tinamidae, 1; Podici-
pedidae, 2; Phalacrocoracidae, 2; Ardeidae, 5 (1); Ciconiidae, 2;
Plataleidae, 5 (2); Anatidae, 9 (4); Vulturidae, 6 (3); Accipitridae,
7 (2); Falconidae, 6 (1); Cracidae, 1; Rallidae, 1; Charadriidae,
7 (2); Scolopacidae, 13 (2); Phalaropodidae, 3 (1); Burhinidae, 1;
Jacanidae, 1; Thinocoridae, 3 (1); Laridae, 2; Columbidae, 4;
Psittacidae, 1; Tytonidae, 1; Strigidae, 3; Caprimulgidae, 2 (1).
Diagnostic osteological characters are given in all cases where
necessary to justify species identification.
The coast of northern Peru where the Talara Tar Seeps are located
is desert today. Very rare unpredictable rains occur during the
summer, and a continuous cloud cover with no precipitation prevails
during the winter. The paleoavifauna indicates that during the Wis-
consin glaciation northwestern Peru was a savanna woodland or a
savanna with extensive riparian forests. During the summer months
regular heavy monsoon rains occurred, followed by a winter dry season
with continuous cloud cover similar to the present winter season.
The explanation of the Pleistocene summer rains is found in the
higher zonal index resulting from increased polar activity during the
periods of glaciation. During this time the intertropical front was
located at or south of the equator. This allowed northern hemisphere
winds to cross the equator and become northwesterly winds that covered
the normally cold coastal waters with warm equatorial water, establish-
ing the necessary conditions for rainfall on land.
The two oldest marine terraces in the region, the Mancora and
Talara Tablazos, with molluscan faunas indicating a cool water environ-
ment, are similar in age and were formed during the Sangamon inter-
glacial. At this time the intertropical front was located well north
of the equator. The similarity of the molluscan fauna of the Lobitos
Tablazo to that of the modern beach indicates a rather recent age,
probably late or post-Wisconsin.
Most of the Equatorial Arid Fauna of Chapman is thought to have
originated during the glacial periods when the intertropical front was
probably located near the equator. Climatic conditions then existed
that restricted rainfall along the western coast of Colombia. The
drier coastal areas provided an avenue for the passage of arid and semi-
arid species from Central America south to the coast of Ecuador and
Peru.
Numerous distinctive subspecies of the Equatorial Arid Fauna
have evolved in southwestern Ecuador and northwestern Peru since the
Wisconsin. Divergence between populations of species of the Equatorial
Arid Fauna occurring in the coastal areas and those found in the
Maranon Valley of Peru is thought to have taken place during glacial
periods. Increased vegetation on the western slopes of the Andes
Mountains during the glacial periods would have effectively isolated
the two drier habitats.
Many of the extinct species probably evolved after the beginning
of the Wisconsin glaciation and became extinct at the end, thus
having a lifespan of approximately 50,000 years.
The Carolinian faunas of southwestern Ecuador are thought to be
either very early Wisconsin or Late Wisconsin based on their similar-
ities with the Talaran faunas and postulated climatic events.
The similarly aged Rancho La Brea avifauna includes 98 non-
passerine species, compared with the 88 non-passerine species found
in the Talara fauna. Sixteen species occur in both localities.
Twenty-three per cent of the non-passerine species of the Talara
avifauna are extinct, in contrast with 17 per cent of the non-
passerine species from Rancho La Brea.
INTRODUCTION
During January and February 1958, Dr. A. Gordon Edmund and Mr.
R. R. Hornell of the Royal Ontario Museum, Toronto, Canada, recovered
an extensive sample of plant, invertebrate, and vertebrate remains
from numerous surface tar seeps in the La Brea-Pariflas oil field, near
Talara, Peru (PI. I). These deposits are known as the Talara Tar
Seeps to avoid confusion with the similar deposits at Rancho La Brea,
California. This report is the first on the avian remains of the
deposit and is concerned with the non-passerine portion of the avifauna.
Additional work needs to be completed before the passerine species are
satisfactorily identified.
The Talara Tar Seeps are located just west of the Pan American
Highway, kilometer 1125, approximately 6 miles west of the Amotape
Mountains and 20 miles due east of the Pacific coast at Punta Parinas,
the westernmost point of South America. The site is marked on maps as
La Brea, but only a cemetery remains of the town. The tar seeps were
first worked by the Indians, who used the material in their road
construction program, and later the Spaniards made extensive use of
the pitch, developing it into a considerable economic resource. The
first oil well in South America was drilled at the site in 1862.
Most of the fossil bones are well preserved, and in numerous
cases broken specimens could be pieced back together, even though
the pieces may have been separated at the time of collection. The
species represented by the most elements is Anas bahamensis, of which
2
there are 2864 bones from a minimum of 243 individuals. Several
species are represented by only one element. Individual specimens
range in size from complete humeri of condors to those of swallows.
Numerous skulls of various species are also present.
Plate I
View of the Talara Tar Seeps, northwestern Peru, looking north.
:._I,;~~~L'~.- L_;~-
MATERIALS AND METHODS
The fossil specimens were studied under loan from the Royal
Ontario Museum, Ontario, Canada. At the termination of this study the
specimens will be cataloged and housed in that institution.
Except in a few instances only skeletons prepared by maceration
were used for the osteological diagnosis. Skeletons that have been
cleaned only by bugging are not free of all tendinal material and are
unsatisfactory for exact osteological description.
Osteological terminology was taken primarily from Howard (1929).
Additional anatomical works used were those of Fisher (1946), Fisher
and Goodman (1955), and Owre (1967).
All measurements were taken with vernier dial calipers accurate
to 0.05 mm. The following abbreviations are used in the table and
text: M, mean; N, number of specimens measured; OR, observed range.
GEOLOGY
The geology of northwestern Peru has been discussed by Bosworth
(1922), Iddings and Olsson (1928), Travis (1953), and Lemon and
Churcher (1961). Additional information is presented by Sheppard
(1937). The following is a brief summary of the geology of the
Talara region based on these works.
The major oil-producing area of Peru is known as the La Brea-
Parihas oil field, which straddles the 4th parallel. The oil field
occupies the westernmost part of the South American continent, and
lies between the Pacific Ocean on the west and the Amotape Mountains,
a small outlier of the Andes Mountains, to the east. The Amotape
Mountains and basement rocks of the coastal area consist of Penn-
sylvanian quartzites, graywackes, argillites, and shales. Directly
upon these lie a series of Cretaceous rocks in excess of 9000 feet
in thickness. The Cretaceous sediments consist of a lower series
of limestone rocks and an upper series of shales, sandstones, and
conglomerates.
The main oil-producing rocks are Eocene in age, and consist of
approximately 15,000 feet of marine elastic sediments. Shales, sands,
and a few conglomerates are present in these sediments that originated
during the orogenic uplift of the Andes Mountains directly to the east.
These sediments are highly faulted and of complex structure as a
result of subsequent orogenic movement.
Early, or perhaps Middle, Pleistocene to Recent marine deposits
overlie the Eocene rocks and consist of well stratified marls, lime-
stones, calcareous sands, coquinas, and pebble beds. These deposits
occupy three terraces, named the Mancora Tablazo (highest and oldest),
the Talara Tablazo (middle terrace and not much younger than the Mancora
Tablazo), and the Lobitos Tablazo (lowest and youngest). The terraces
represent periods of marine encroachment followed by orogenic uplift,
which was greater in the north than in the south, giving the terraces
a southward tilt. The deposits are thickest in the north and thin
southward, ranging only up to 50 or 80 feet in thickness. The Talara
Tar Seeps rest on the Mancora Tablazo. Recent deposits consist of
large alluvial breccia fans from the Amotape Mountains, valley
terraces, sand dunes, and Recent marine deposits.
The bone-bearing deposits occur as lenses or horizontal beds,
a few to several feet in thickness and of varying lengths (Pl. II,
Fig. A). The lenses closely resemble sand or gravel deposits that
form in normal stream deposition, or they may represent pools of tar
that filled with sands or gravels. Probably both types of deposits
are represented, for although the area is now desert, dry stream
beds and extensive gulleying indicate that the area was consider-
ably wetter during the Wisconsin glaciation. Oil and water currently
escaping from abandoned oil wells attract both vertebrate and inverte-
brate animals and act as active traps (P1. II,Fig. B). Windblown sand
usually forms a crust over most of the recent seeps. Persistent large
pools of pitch occur only in areas where man has excavated the natural
asphalt for road material or museum collections.
Plate II
Two views of the Talara Tar Seeps
Fig. A. Close-up view of the deposit, showing lensing type of
deposition and abundance of fossil material.
Fig. B. View of Phalacrocorax olivaceus recently entombed in
surface layer of tar.
Fig. A
Fig. B
SYSTEMATIC LIST
The following systematic list details the non-passerine species
known from the Talara Tar Seeps, as summarized in Table 1. Nomencla-
ture used is primarily that of Brodkorb (1963, 1964, 1967, 1971).
Except where otherwise noted species distributions are taken from
Meyer de Schauensee (1966).
In some instances specimens are thought to represent immature
individuals. This conclusion is based on the pitted appearance of the
surface of the bone and incomplete ossification in the articular areas.
Order Tinamiformes (Huxley)
Family Tinamidae Gray
Genus Crypturellus Brabourne & Chubb
Crypturellus cf. transfasciatus (Sclater & Salvin)
Pale-browed Tinamou
Material. Proximal end of 1 right humerus.
Remarks. At the present time C. transfasciatus occurs from the
equator in western Ecuador south to northwestern Peru in Tumbes and
Piura. It is the only species of tinamou found in the dry deciduous
forests that surround the semi-arid and arid regions of southwestern
Ecuador and coastal Peru. The fossil specimen corresponds in size
with C. transfasciatus, but as no skeletal material was available for
comparison the specimen is only tentatively identified.
Table 1. List of non-passerine species known from the Talara Tar
Seeps, with number of elements and least number of individuals of
each indicated.
Number of Least Number
Elements of Individuals
Tinamiformes
Tinamidae
Crypturellus cf. transfasciatus 1 1
Podicipediformes
Podicipedidae
Podiceps dominicus 4 2
Podilymbus podiceps 4 2
Pelecaniformes
Phalacrocoracidae
Phalacrocorax olivaceus 133 15
P. bougainvillii 1 1
Ardeiformes
Ardeidae
Ardea cocoi 6 2
Casmerodius albus 11 4
Egretta thula 7 2
E. caerulea 1 1
Syrigma n. sp. 1 1
Nycticorax nycticorax 53 10
Ciconiidae
Jabiru mycteria 47 5
Mycteria americana 1 1
Plataleidae
Theristicus n. sp. 1 1
Eudocimus n. sp. 9 2
E. albus 10 3
Ajaia 'ajia 1 1
Anseriformes
Anatidae
Dendrocygna autumnalis 299 33
Tadorninae, n. gen. n. sp. 112 10
Chloephaga melanoptera 91 7
Cairina moschata 33 4
Anas n. sp. 1 13 2
Anas n. sp. 2 29 5
Anas n. sp. 3 5 1
A. bahamensis 2864 243
Nomonyx dominicus 1 1
Table 1. Continued.
Number of Least Number
Elements of Individuals
Accipitriformes
Vulturidae
Vulturidae, n. gen. n. sp. 28 4
Gymnogyps n. sp. 30 4
Vultur gryphus 17 3
Sarcoramphus? n. sp. 3 1
Coragyps cf. atratus 72 6
Cathartes aura 26 3
Accipitridae
Buteoninae?, n. gen. n. sp. 8 2
Geranoaetus n. sp. 52 7
G. melanoleucus 31 4
Buteo polyosoma 30 4
Buteo sp. 1 4 2
Buteo sp. 2 10 4
Parabuteo unicinctus 30 4
Falconidae
Falco peregrinus 32 3
F. femoralis 7 2
F. sparverius 47 6
Falco sp. 6 1
Polyborus plancus 293 27
Milvago n. sp. 160 17
Galliformes
Cracidae
Penelope cf. purpurascens 4 1
Ralliformes
Rallidae
Porzana carolina 3 1
Charadriiformes
Charadriidae
Vanellinae, n. gen. n. sp. 1 1
Belonopterus n. sp. 3 2
Pluvialis dominica 32 6
Squatarola squatarola 3 1
Charadrius vociferous 7 2
C. collaris 3 1
C. semipalmatus 1 1
Scolopacidae
Tringa solitaria 21 5
Tringa n. sp. 2 1
Totanus flavipes 69 17
T. melanoleucus 11 2
Actitis macularia 1 1
Catoptrophorus semipalmatus 16 4
Table 1. Continued.
Number of Least Number
Elements of Individuals
Erolia minutilla 28 12
E. melanotos 220 57
Ereunetes mauri 16 3
Micropalama himantopus 1 1
Arenaria interpres 2 1
Numenius cf. borealis 1 1
Scolopacidae, n. gen. n. sp. 3 1
Phalaropodidae
Lobipes lobatus 2 2
Steganopus tricolor 65 11
Steganopus n. sp. 1 1
Jacanidae
Jacana spinosa 1 1
Burhinidae
Burhinus superciliaris 29 4
Thinocoridae
Thinocorus n. sp. 1 1
T. rumicivorus 1 1
cf. Attagis sp. 1 1
Laridae
Larus atricilla 31 8
L. pixcan 95 17
Columbiformes
Columbidae
Zenaida auriculata 102 13
Z. asiatica 7 2
Columbina talpacoti 3 2
C. cruziana 578 68
Psittaciformes
Psittacidae
Forpus coelestis 1 1
-Strigiformes
Tytonidae
Tyto alba 83 8
Str gi-ae
Bubo virginianus 101 11
Speotyto cunicularia 6 1
Asio lammeus 12 2
Caprimulgiformes
Caprimulgidae
Chordeiles acutipennis 6 1
Caprimulgus n. sp. 2 1
Total 6200 742
Order Podicipediformes (Furbringer)
Family Podicipedidae (Bonaparte)
Genus Podiceps Latham
Podiceps dominicus (Linnaeus)
Least Grebe
Material. One complete right coracoid, 1 nearly complete left
femur, proximal end of 1 left femur, proximal end of 1 left tarsometa-
tarsus. The 4 specimens represent a minimum of 2 individuals.
Characters. Easily identified by their small size, the fossil
specimens display characters well within the range of variation
found in Recent specimens.
Remarks. The present range of P. dominicus includes northwestern
Peru.
Genus Podilymbus Lesson
Podilymbus podiceps (Linnaeus)
Pied-billed Grebe
Material. Proximal end of 1 right humerus, 1 nearly complete
right femur, proximal end of 1 right femur, 1 complete left tarsometa-
tarsus. The 4 specimens represent a minimum of 2 individuals.
Characters. The fossil specimens vary within acceptable limits
from Recent specimens examined.
Remarks. The present range of P. podiceps includes northwestern
Peru, and I would expect to find this grebe on almost any body of
fresh-water in coastal Peru, although not in large numbers.
Order Pelecaniformes Sharpe
Suborder Sulae Sharpe
Family Phalacrocoracidae (Bonaparte)
Genus Phalacrocorax Brisson
Phalacrocorax olivaceus (Humboldt)
Olivaceous Cormorant
Material. Two maxillae, distal halves of 2 right and 2 left
mandibles, 5 right and 2 left scapulae, 5 complete right and 11
complete left coracoids, humeral ends of 1 right and 3 left coracoids,
sternal ends of 2 right and 4 left coracoids, 2 complete left humeri,
proximal ends of 2 right and 4 left humeri, distal ends of 7 right and
6 left humeri, proximal ends of 3 right and 2 left ulnae, distal ends
of 5 right and 2 left ulnae, proximal ends of 5 right and 2 left radii,
distal ends of 4 right and 1 left radius, 3 complete right and 1
complete left carpometacarpus, proximal ends of 1 right and 1 left
carpometacarpus, 4 complete right and 4 complete left femora, proximal
end of 1 left femur, distal ends of 1 right and 1 left femur, 2
complete right and 1 complete left tibiotarsus, proximal ends of 1
right and 2 left tibiotarsi, distal ends of 1 right and 3 left tibio-
tarsi, 8 complete right and 8 complete left tarsometatarsi, proximal
ends of 1 right and 2 left tarsometatarsi, distal ends of 3 left
tarsometatarsi. The 133 specimens represent a minimum of 15
individuals.
Characters. With the exception of certain cranial elements,
elements of P. olivaceus are generally smaller than elements of P.
gaimardi, although size ranges of the two species overlap. They are
much smaller, especially more slender, than elements of P. bougainvillii.
P. olivaceus differs from P. bougainvillii and P. gaimardi by
having premaxillary and mandible with (1) size large (small in P.
bougainvillii and P. gaimardi).
Scapula with (1) glenoid facet distinctly triangular (more
rounded in P. bougainvillii and P. gaimardi); (2) acromion wide
and moderately long (wide and long in P. bougainvillii, narrow and
moderately long in P. gaimardi); (3) protrusion on lateral surface
of shaft distal to glenoid facet round and prominent (elongated and
only slightly raised, if any, in P. bougainvillii and P. gaimardi).
Coracoid with (1) shaft very slender (very robust in P. bougain-
villii, slender in P. gaimardi); (2) bicipital attachment narrow and
deeply concave (wide and moderately to slightly concave in P. bougain-
villii and P. gaimardi); (3) procoracoid short and rounded (moderate
to long, stouter, and more angular in P. bougainvillii and P. gaimardi);
(4) attachment of Lig. humero-coracoideum anterius superius deep
(superficial in P. bougainvillii and P. gaimardi); (5) humeral end
narrow (wide in P. bougainvillii and P. gaimardi).
Humerus with (1) pneumatic fossa moderately excavated (deeply
excavated in P. bougainvillii and P. gaimardi); (2) deltoid crest
small, smoothly rimmed (larger and locally angular in P. bougainvillii
and P. gaimardi); (3) attachment of M. extensor metacarpi radialis
inset in shallow depression (attachment located on raised platform in
P. bougainvillii; attachment inset in depression in P. gaimardi, but
bordered distally by large ectepicondylar prominence); (4) attachment
of anterior articular ligament with moderate slope (lesser slope in
P. bougainvillii, greater slope in P. gaimardi).
Ulna with (1) internal cotyla narrow (wider in P. bougainvillii
and P. gaimardi); (2) external cotyla wide and moderately long (long
and narrow in P. bougainvillii, short and narrow in P. gaimardi);
(3) external condyle of moderate width and length (narrow and long in
P. bougainvillii, moderately wide and short in P. gaimardi); (4)
carpal tuberosity moderately large (larger in P. bougainvillii and
P. gaimardi).
Radius with (1) insertion of M. biceps moderately deep (very deep
in P. bougainvillii and P. gaimardi); (2) projection extending from
lateral end of carpal facet sharp and pointed (only slightly developed
in P. bougainvillii and P. gaimardi).
Carpometacarpus with (1) external rim of carpal trochlea small
(much wider in P. bougainvillii, slightly wider in P. gaimardi); (2)
process of metacarpal I narrow (broad in P. bougainvillii and P.
gaimardi); (3) pollical facet wide (narrower in P. bougainvillii and
P. gaimardi).
Femur with (1) iliac facet small (larger in P. bougainvillii and
P. gaimardi, even though the femur is shorter in P. gaimardi); (2)
internal condyle slightly angular distally (more rounded, and rotated
posteriorly, in P. bougainvillii and P. gaimardi); (4) shaft slender
(much heavier in P. bougainvillii and P. gaimardi).
Tibiotarsus with (1) outer cnemial crest wide and short (wide
and long in P. bougainvillii, narrow and short in P. gaimardi); (2)
shaft slightly to moderately concave distal to internal articular
surface (deeply concave in P. bougainvillii, slightly to moderately
concave in P. gaimardi); (3) internal condyle short and moderately
wide (long and wide in P. bougainvillii, long and narrow in P.
gaimardi); (4) size very small.
Tarsometatarsus with (1) internal cotyla short, narrow, and
moderately concave (longer, wider, and more concave in P. bougain-
villii and P. gaimardi); (2) inner extensor groove wide (narrow in P.
bougainvillii and P. gaimardi); (3) external trochlea narrow (wide in
P. bougainvillii and P. gaimardi); (4) distal foramen moderately
sized (smaller in P. bougainvillii, much larger in P. gaimardi).
Remarks. P. olivaceus is generally distributed throughout
South America, and is common in coastal lagoons, marshes, lakes,
and rivers.
Phalacrocorax bougainvillii (Lesson)
Guanay Cormorant
Material. Distal end of 1 right humerus.
Remarks. P. bougainvillii is a marine species that breeds on
offshore islands and comes ashore only rarely, and then usually only
during times of distress (Murphy, 1936). The population of this
species numbers in the millions, and it undergoes wide fluctuations.
At times when the population crashes the beaches are literally
covered with corpses. This happens whenever the warm current known
as El Nino appears. The single specimen in the collection can be
regarded as a rare straggler.
Order Ardeiformes (Wagler)
Suborder Ardeae Wagler
Family Ardeidae Vigors
Genus Ardea Linnaeus
Ardea cocoi Linnaeus
White-necked Heron
Material. One right scapula, proximal ends of 1 right and 1
left carpometacarpus, distal ends of 2 left carpometacarpi, distal end
of 1 right tarsometatarsus. The 6 specimens represent a minimum of
2 individuals.
Characters. As A. cocoi is the largest living South American
heron its elements are readily recognized on size.
Remarks. Generally distributed over all of South America, A.
cocoi is found in coastal marshes and lagoons in Peru, although only
in small numbers.
Genus Casmerodius Gloger
Casmerodius albus (Linnaeus)
Common Egret
Material. Two complete right and 2 complete left coracoids,
humeral ends of 2 right coracoids, 1 right scapula, 1 complete left
carpometacarpus, proximal end of 1 left carpometacarpus, distal end
of 1 right tibiotarsus, distal end of 1 right tarsometatarsus. The
11 elements represent a minimum of 4 individuals.
Characters. Intermediate in size between Ardea and Tigrisoma,
Casmerodius is easily distinguished on size.
Remarks. Generally distributed over all of South America,
C. albus is found commonly in coastal marshes and lagoons of Peru.
Genus Egretta T. Forster
Egretta thula (Molina)
Snowy Egret
Material. Two right scapulae, 2 complete right coracoids,
proximall end of 1 left humerus, 1 complete right femur, proximal
end of 1 right femur. The 7 specimens represent a minimum of 2
individuals.
Additionally, the following specimens are identified to genus
only: 1 complete right and 2 complete left carpometacarpi, distal
ends of 3 left tibiotarsi, proximal end of 1 right tarsometatarsus,
distal ends of 3 right and 5 left tarsometatarsi.
Characters. The three species of Egretta, E. thula, E.
(Hydranassa) tricolor, and E. (Florida) caerulea, are extremely
difficult to separate osteologically. Characters used to identify the
fossil specimens of E. thula are given below, but intraspecific
variation is so great that even these characters may prove to be
invalid if sufficient series are examined.
Scapula with (1) thickness between acromion and coracoidal
articulation moderate, but variable (similar in E. caerulea, slightly
thicker in E. tricolor); (2) length of acromion base moderate (less
in E. caerulea, greater in H. tricolor).
Coracoid with (1) curvature of coracohumeral surface slight to
moderate as traced by central groove (similar in E. caerulea, very
slight in E. tricolor); (2) angle between internal edge of furcular
facet and triosseal canal slightly more than 90 (much larger in E.
caerulea, and the area is well rounded; slightly larger in E. tricolor).
Humerus with (1) distal end of attachment of M. proscapulo-
humeralis surrounded by attachment of M. triceps, external head (not
surrounded in E. caerulea, roughly similar in E. tricolor); (2)
external attachment of M. supracoracoideus short and deep (short
and shallow in E. caerulea, long and shallow in E. tricolor).
Carpometacarpus with (1) intermuscular line between pollical
facet and pisiform process lying near pollical facet (similar in E.
caerulea, positioned posteriorly in E. tricolor). No characters
were found that reliably separated the carpometacarpi of E. thula
from those of E. caerulea. The three fossil specimens could thus be
separated from E. tricolor, but assignment to either E. thula or E.
caerulea was not possible.
Femur with (1) iliac facet narrow antero-posteriorly (narrow to
moderate width in E. caerulea, broader in E. tricolor); (2) external
condyle flat dorsally (similar in E. caerulea, extending farther
dorsally in E. tricolor); (3) distal end broad (narrow in E. caerulea,
broad in E. tricolor).
Tibiotarsus with (1) rotular crest low (low in E. caerulea,
higher in E. tricolor); (2) external condyle slightly undercut
dorso-proximally (not undercut in E. caerulea; slightly, or not,
undercut in E. tricolor); As no characters were found that could
reliably be used to separate the tibiotarsi of E. thula and E.
tricolor, the specimens are simply referred to the genus.
Tarsometatarsus. No reliable characters were found that could
be considered valid for separating tarsometatarsi of the three
species of Egretta. For this reason the specimens are simply referred
to the genus.
Remarks. Generally distributed over almost all of South America,
E. thula is common in coastal lagoons and marshes of Peru.
Egretta caerulea (Linnaeus)
Little Blue Heron
Material. One complete left coracoid.
Remarks. Generally distributed over almost all of South America,
E. caerulea is found in coastal lagoons and marshes of Peru, but not
in such large numbers as E. thula.
Genus Syrigma Ridgway
Syrigma n. sp.
Holotype. Complete left coracoid. (PI. III, Fig. C)
Diagnosis. Agrees with Syrigma and differs from all other genera
of South American herons by having (1) head broad and slightly peaked
in medial view (subangular to subrounded); (2) brachial tuberosity
prominent and angular; (3) triosseal canal very deep adjacent to glenoid
facet; (4) attachments of M. coracobrachialis anterior and Lig. humero-
coracoideum anterius superius deep, elongated, and notched; with center
of latter lying parallel, and roughly in line with, anterior edge of
shaft; (5) protrusion internal to glenoid facet large, bordering ventral
side of triosseal canal; (6) external side of shaft with slight ridge
running ventrally from middle of head.
Differs from Syrigma sibilatrix (Temminck) by having (1)
coracohumeral surface short (displaces dorsal-most point of head
posteriorly); (2) furcular facet larger; (3) brachial tuberosity
larger; (4) proximal end with medial side flatter (locally more
convex and concave in S. sibilatrix); (5) protrusion internal to
glenoid facet smaller; (6) head thicker between attachment of Lig.
humero-coracoideum anterius superius and anterior furcular facet; (7)
glenoid facet rotated posteriorly, lengthening distance from external
edge of glenoid facet and anterior edge of shaft; (8) external sternal
facet much higher. Measurements of the holotype, with those of one
male specimen of S. sibilatrix in parenthesis, are as follows: length,
42.3 (41.6); head to scapular facet, 13.3 (13.5); proximal depth, 9.5
(9.2); least depth of shaft, 4.2 (4.3).
Remarks. The genus Syrigma is represented by one living
species in South America, S. sibilatrix, and it occurs only east of
the Andes. The presence of a second, extinct species west of the
Andes suggests that perhaps the climate west of the Peruvian Andes
was more equitable during the late Pleistocene than it is today.
Syrigma n. sp. is the first paleospecies of the genus to be
described.
Genus Nycticorax T. Forster
Nycticorax nycticorax (Linnaeus)
Black-crowned Night Heron
Material. Middle third of 1 left mandible, 1 right and 3 left
quadrates, 4 right and 3 left scapulae, 1 complete right coracoid,
humeral ends of 9 right and 4 left coracoids, proximal end of 1 left
humerus, proximal end of 1 right ulna, distal ends of 2 right and 3
left ulnae, distal end of 1 left radius, proximal end of 1 left
carpometacarpus, 1 complete right femur, proximal end of 1 right and 1
left femur, distal ends of 3 right and 5 left tibiotarsi, 1 complete
right and 1 complete left tarsometatarsus, proximal end of 1 right
tarsometatarsus, distal ends of 1 right and 3 left tarsometatarsi.
The 53 specimens represent a minimum of 10 individuals.
Characters. All elements of Nycticorax examined are separable
from those of Egretta by size, those of Nycticorax being either much
longer, shorter, wider, or stouter than corresponding elements of
Egretta. Nycticorax differs from Nyctanassa Stejneger and Tigrisoma
Swainson by having scapula with (1) coracoidal articulation large
(large and subrounded in Tigrisoma, smaller in Nyctanassa); (2)
glenoid facet oval (broad oval in Tigrisoma, narrow oval in Nyctanassa);
.(3) acromion deflected moderately externally (less in Tigrisoma,
more in Nyctanassa).
Coracoid with (1) triosseal canal moderately inset (slightly
inset, if any, in Trigrisoma and Nyctanassa); (2) glenoid facet large
(much smaller in Tigrisoma, of similar size in Nyctanassa); (3)
dorsal surface of head notched by deep concavity in medial view
(rounded in Tigrisoma, notched and more angular in Nyctanassa);
(4) anterior end of head rotated moderately externally (rotated less
in Tigrisoma, much more in Nyctanassa).
Humerus with (1) attachment of M. proscapulohumeralis elongated
and moderately inset (oval and inset only slightly in Tigrisoma and
Nyctanassa); (2) pneumatic fossa very shallow (moderately deep in
Tigrisoma and Nyctanassa); (3) dorsal edge of attachment of external
head of triceps curving away from capital shaft ridge distal to
attachment of M. latissimus dorsi posterioris (extends farther
proximally on capital shaft ridge in Tigrisoma and Nyctanassa).
Ulna with (1) olecranon narrow (broad in Tigrisoma, similar in
Nyctanassa); (2) external cotyla large, with small ridge leading
distally (smaller, without ridge in Tigrisoma and Nyctanassa); (3)
external condyles long (short in Tigrisoma, moderate in Nyctanassa);
(4) carpal tuberosity moderately long (long in Tigrisoma, short in
Nyctanassa).
Radius with (1) palmar surface of distal end moderately excavated
(slightly excavated in Tigrisoma, greatly excavated in Nyctanassa);
(2) tendinal groove broad and deep (narrower and not as deep in
Tigrisoma and Nyctanassa); (3) carpal facet broad (narrow in Tigrisoma
and Nyctanassa).
Carpometacarpus with (1) internal ligamental attachment of
pisiform process and ligamental attachment of ulnare confluent
(separated in Tigrisoma and Nyctanassa); (2) corner on distal end
of internal rim of carpal trochlea small (large in Tigrisoma, absent
in Nyctanassa); (3) carpal trochlea moderately wide (wider in
Tigrisoma, similar in Nyctanassa); (4) process of metacarpal I
slender (stouter in Tigrisoma and Nyctanassa).
Femur with (1) proximal lateral face curving medially (curvature
slightly less in Tigrisoma, much less in Nyctanassa); (2) concavity
distal to iliac facet posteriorly deep (moderate concavity in Tigrisoma,
slight concavity in Nyctanassa); (3) posterior intermuscular line
curving medially to base of internal condyle (lies more in center of
shaft and does not curve in Tigrisoma, does not curve as far internally
in Nyctanassa); (4) gap between external condyle and origin of M.
gastrocnemius, pars externa, wide (ridge present between the two in
Tigrisoma, slight gap in Nyctanassa).
Tibiotarsus with (1) internal articular surface large (similar
in Tigrisoma, smaller in Nyctanassa); (2) inner cnemial crest short
and thick (long and of moderate width, tapering distally, in
Tigrisoma and Nyctanassa); (3) ridge leading to postero-most corner of
internal articular surface absent (present in Tigrisoma, absent in
Nyctanassa); (4) internal condyle undercut dorsally (not undercut
in Tigrisoma and Nyctanassa); (5) internal condyle flaring extensively
internally (much less in Tigrisoma and Nyctanassa).
Tarsometatarsus with (1) shaft large (more slender in Tigrisoma,
much more slender in Nyctanassa); (2) cotylae and trochleae wide
(narrow in Tigrisoma and Nyctanassa); (3) internal edge to shaft
angular rounded in Tigrisoma and Nyctanassa).
Remarks. N. nycticorax is generally distributed throughout
South America, including coastal rivers, swamps, and lagoons of Peru.
On the basis of osteological comparisons listed above I believe
Nycticorax is generically distinct from Nyctanassa.
Suborder Plataleae Newton
Family Plataleidae Bonaparte
Subfamily Threskiornithinae (Richmond)
Genus Theristicus Wiagler
Theristicus n. sp.
Holotype. Complete left coracoid. (P1. III, Fig. B)
Diagnosis. Agrees with Theristicus and differs from all other
genera of South American ibises (with possible exception of Harpiprion
Wagler and Cercibis Wagler which were not available for comparison), by
having (1) head long and subangular in medial view; (2) attachment of M.
coracobrachialis anterior moderately deep, long, with a narrow posterior
section and a much wider anterior section; (3) attachment of Lig.
humero-coracoideum anterius superius shallow and confluent with anterior
section of attachment of M. coracobrachialis anterior; (4) coracohumeral
surface broad, of almost uniform width; (5) head very wide, with external
and internal sides roughly parallel; (6) brachial tuberosity large, well
rounded; (7) glenoid facet large, with widest area immediately dorsal
to scapular facet; (8) shaft with anterior edge immediately dorsal to
sternal facet flattened and rotated slightly internally; (9) anterior
external sternal facet with large protrusion posteriorly.
Differs from Theristicus caudatus (Boddaert) by having (1)
coracohumeral surface much shorter, giving a more rounded head with
lower profile in medial view; (2) attachments of M. coracobrachialis
anterior and Lig. humero-coracoideum anterius superius shallower; (3)
procoracoid deeply concave internally; (4) shaft with anterior edge
immediately dorsal to sternal facet more flattened and rotated less
internally; (5) external sternal facet and external distal angle much
shorter; (6) concavity between external sternal facet and flattened
anterior edge of shaft shallow. Measurements of the holotype, with
those of three male specimens of T. caudatus in parenthesis are as
follows: length, 50.5 (52.0-54.3; mean, 52.8); head to scapular facet,
19.9 (20.8-21.1; mean, 20.9); proximal depth, 12.9 (13.7-14.1; mean,
13.8); least depth of shaft, 8.5 (9.0-9.1; mean, 9.1).
Referred Material. Fragmentary proximal end of 1 right humerus.
Characters. Agrees with Theristicus and differs from all other
examined genera of South American ibises by having (1) pneumatic fossa
shallow; (2) internal tuberosity short; (3) median crest very prominent.
Differs from Theristicus caudatus by having (1) internal tuberosity
shorter; (2) attachment of M. coracobrachialis posterior rotated
toward axis of shaft; (3) median crest more prominent; (4) attachment
of Lig. humero-coracoideum anterius inferius positioned more anconally.
The specimen is too broken to yield accurate measurements.
Remarks. Theristicus is represented by one living species in
South America, T. caudatus, that occurs over almost the entire
continent, including cultivated areas of coastal Peru (Koepcke, 1970).
No significant differences were seen between specimens of T. c. caudatus
and T. c. branickii examined. Although no specimens of T. c. melanopis
were available, I doubt that it would vary significantly from T. c.
caudatus or T. c. branickii.
Theristicus n. sp. is the first described paleospecies of the
genus.
Genus Eudocimus Wagler
Eudocimus n. sp.
Holotype. Complete left tarsometatarsus. (PI. III, Fig. A)
Paratype. Distal end of 1 left tarsometatarsus.
Diagnosis. Agrees with Eudocimus and differs from all other
genera of South American ibises (with possible exception of Harpiprion
and Cercibis which were unavailable for comparison) by having (1)
shaft long, moderately stout; (2) cotylae, trochleae, and hypotarsus
moderately compressed.
Differs from Eudocimus albus Wagler by having (1) shaft with
channeling of anterior face very shallow, deepest on external side,
with proximal ligamental attachment raised to level of sides of shaft
(shaft with anterior face deeply channeled, deepest on internal side,
with proximal ligamental attachment lying below level of sides of
shaft in E. albus); (2) intercotylar prominence larger, rounder,
and rotated dorsally; (3) proximal end wider and distal end narrower
(when compared with specimen of E. albus of same, or greater, length);
(4) cotylae slightly longer and wider; (5) groove between intercotylar
prominence and external cotylae distinct; (6) hypotarsal ridges
extend farther distally and forming "V"; (7) external side of hypo-
tarsus and shaft anterior to internal side of hypotarsus more excavated;
(8) attachment of external ligament strong, positioned posteriorly;
(9) convexity at anterior end of internal trochlea slightly developed;
(10) depth of middle trochlea less; (11) external trochlea with
external face large, extending less posteriorly.
For measurements see Table 2. Measurements of the holotype
are as follows: length, 100.8; proximal width, 13.7; distal width,
12.7; least width of shaft, 5.3; width of middle trochlea, 4.9.
Referred material. One right scapula, distal end of 1 left
humetus, proximal end of 2 radii, 1 complete left femur, proximal
end of 1 right femur, 1 complete right tibiotarsus (immature), distal
end of 1 right tibiotarsus. The 10 specimens, including holotype and
paratype, represent a minimum of 2 individuals.
Characters. Scapula with (1) distal end of glenoid facet
slightly elevated above surface of shaft, and rotated ventrally only
slightly (more elevated and rotated ventrally in E. albus).
Humerus with (1) attachment of anterior articular ligament
broader, extending less distally; (2) attachment of M. pronator
brevis located farther distally; (3) internal condyle projecting
less distally, more palmarly; (4) intercondylar groove shallower.
Radius with (1) capital tuberosity larger; (2) ligamental
papillae smaller; (3) bicipital attachment a very deep pit; (4)
ulnar facet larger.
Femur with (1) anterior intermuscular line straight, lying near
external side of shaft (curved and positioned medially in E. albus);
(2) iliac facet shorter; (3) concavity distal to posterior edge of
iliac facet shallower; (4) trochanter projecting less anteriorly; (5)
internal flexure of shaft proximal to internal condyle greater; (6)
ridge leading to posterior corner of internal condyle much smaller,
with more curving edge; (7) external condyle smaller, less rounded.
Remarks. E. ruber differs more from E. albus and Eudocimus n. sp.
than the latter two do from each other, thus a detailed comparison
with E. ruber is not considered necessary.
Eudocimus n. sp. is the first paleospecies of the genus
described from the New World. Its presence in the same deposit as
E. albus, which does not quite reach so far south as the fossil
site today, poses the interesting question of whether the two species
were sympatric or merely had seasonally overlapping ranges. Neither
of the two species occur in very large numbers in the collection.
Eudocimus albus (Linnaeus)
White Ibis
Material. One complete left coracoid, humeral ends of 2 left
coracoids, distal end of 1 right ulna, 1 complete right carpometa-
carpus, distal ends of 2 left tibiotarsi, proximal end of 1 right
Table 2. Measurements of the scapula, humerus, radius, femur, tibiotarsus, and tarsometatarsus
of Eudocimus n. sp. and Recent E. albus, and of the scapula, tibiotarsus, and tarsometatarsus of
E. albus from the Talara Tar Seeps (in mm).
Eudocimus n. sp.
11.9
1
3.5
Eudocimus albus
Talara Tar Seeps
11.0
1
3.7
17.2
1
8.9
1
5.6-5.9
5.8
2
Scapula
Proximal
Height
Proximal
Width
Humerus
Distal
Width
Depth of
External
Condyle
Radius
Proximal
Width
Recent
9.3-11.8
10.9
5
3.1-4.1
3.7
14.6-17.8
16.3
5
7.9-9.9
8.9
5
4.3-5.4
4.9
5
Table 2. Continued.
Eudocimus n. sp.
Eudocimus albus
Talara Tar Seeps
59.0
1
11.9-12.2
12.1
2
12.6
1
141.0
10.6-11.0
10.8
2
9.5-10.4
9.9
2
Femur
Length
Proximal
Width
Recent
Distal
Width
Least
Width of
Shaft
Tibiotarsus
Length
Distal
Width
56.1-67.1
62.9
S
10.8-13.5
13.2
5
11.1-13.8
12.4
S
5.0-6.5
102.3-141.8
127.3
5
8.5-10.6
9.8
5
Table 2. Continued.
Eudocimus n. sp.
Least
Width of
Shaft
Depth of
Internal
Condyle
Tarsometatarsus
Length
Proximal
Width
Distal
Width
Least
Width of
Shaft
Width of
Middle
Trochlea
Eudocimus albus
Talara Tar Seeps
5.2-5.4
5.3
2
11.3-11.7
11.5
2
10.0-11.8
10.9
2
100.8
13.7
1
12.7
1
12.4
1
11.2
1
4.6-4.9
4.8
2
Recent
4.4-5.4
5.0
5
9.2-12.1
10.8
5
73.0-103.8
92.7
5
10.2-12.8
11.8
5
10.4-13.3
12.1
5
4.6-5.4
4.9
5
4.0-5.0
4.6
5
tarsometatarsus, distal end of 1 right tarsometatarsus. The 9
specimens represent a minimum of 3 individuals.
Remarks. E. albus occurs today in the northernmost part of
coastal Peru, western Ecuador, Colombia, and Venezuela.
Subfamily Plataleinae Bonaparte
Genus Ajaia Reichenbach
Ajaa ajaja (Linnaeus)
Roseate Spoonbill
Material. Proximal end of 1 right carpometacarpus.
Characters. Carpometacarpus with (1) metacarpal I long and
slender; (2) posterior carpal fossa elongated and very deep; (3)
pollical facet moderately wide; (4) size large.
Remarks. A. ajaja is widespread in South America, and has
been recorded from northwestern Peru.
Suborder Ciconiae Bonaparte
Family Ciconiidae (Gray)
Subfamily Ciconiinae Gray
Genus Jabiru Hellmayr
Jabiru mycteria (Lichtenstein)
Jabiru
Material. One cranium, 1 sternal fragment, proximal ends of 2
left scapulae, symphyseal area of 1 furculum, portions of 3 right and
4 left coracoids, distal ends of 1 right and 1 left humerus, proximal
ends of 2 left ulnae, distal ends of 2 right and 2 left ulnae, distal
ends of 2 right and 1 left radius, proximal end of 1 left carpometa-
carpus, distal ends of 2 right carpometacarpi, 5 right and 1 left
carpal digits, proximal end of 1 left tibiotarsus, distal ends of 3
right and 4 left tibiotarsi, proximal ends of 1 right and 2 left
tarsometatarsi, distal ends of 3 right and 1 left tarsometatarsi.
The 47 elements represent a minimum of 5 individuals.
Remarks. J. myctera has been recorded on the coast of southern
Peru, but only as isolated individuals, and these are regarded as
trans-Andean vagrants. The specimens represented here may also
represent trans-Andean vagrants, or they may represent individuals
that moved southward from the moister coastal regions of southern
Ecuador. Although there are no Recent records of Jabiru in Ecuador,
I see no reason why it may not have been there in the past, or why
it may not be there now.
Subfamily Mycteriinae American Ornithologists' Union
Genus Mycteria Linnaeus
Mycteria americana Linnaeus
Wood Stork
Material. Proximal end of 1 carpometacarpus.
Remarks. Although M. americana has been recorded from north-
western Peru, it has not been recorded from the remainder of coastal
Peru. The scarcity of M. americana may indicate that there were no
large wet areas near the fossil site. The single specimen must
represent a very rare vagrant.
Order Anseriformes Wagler
Suborder Anseres Wagler
Family Anatidae Vigors
Subfamily Dendrocygninae Reichenbach
Genus Dendrocygna Swainson
Dendrocygna autumnalis (Linnaeus)
Black-bellied Tree Duck
Material. Portions of 2 crania, 1 frontal, 4 right and 7 left
quadrates, 1 right and 2 left articulars, 1 right and 5 left man-
dibular fragments, 15 right and 14 left scapulae, 13 complete right
and 21 complete left coracoids, humeral ends of 5 right and 4 left
coracoids, sternal ends of 2 left coracoids, 2 complete right and 2
complete left humeri, proximal ends of 14 right and 12 left humeri,
distal ends of 17 right and 12 left humeri, 2 complete right and 1
complete left ulna, proximal ends of 6 right and 6 left ulnae,
distal ends of 8 right and 7 left ulnae, 1 complete left radius,
proximal ends of 5 right and 4 left radii, distal ends of 9 right and
8 left radii, 12 complete right and 10 complete left carpometa-
carpi, proximal ends of 7 right and 7 left carpometacarpi, 7 complete
right and 4 complete left femora, proximal ends of 4 right and 5
left femora, distal ends of 1 right and 1 left femora, distal ends
of 14 right and 4 left tibiotarsi, 1 complete right tarsometatarsus,
proximal end of 1 right and 1 left tarsometatarsus, distal ends of
4 right and 5 left tarsometatarsi. The 299 specimens represent a
minimum of 33 individuals.
Characters. The larger size of D. autumnalis serves to distin-
guish its elements from those of D. viduata (Linnaeus). Although D.
bicolor (Vieillot) is usually smaller than D. autumnalis, the two
species overlap in size. Characters of D. autumnalis used to separate
it from D. bicolor are listed below. For the scapula, radius and ulna,
elements not utilized by Woolfenden (1961), generic characters are
also given.
Scapula of Dondrocygna with (1) glenoid facet long and narrow,
lacking an antero-ventral projection. Scapula of D. autumnalis
with (1) acromion longer; (2) glenoid facet longer and more elevated.
Coracoid with (1) angle between furcular facet and edge of
triosseal canal much smaller; (2) brachial tuberosity less rounded;
(3) head more rounded in medial view; (4) attachment of Lig. humero-
coracoideum anterius superius larger.
Humerus with (1) proximal and distal ends much wider; (2)
deltoid crest longer and more flaring; (3) internal tuberosity much
larger; (4) ectepicondylar prominence more rounded; (5) attachment of
anterior articular ligament higher; (6) entepicondyle much larger.
Ulna of Dendrocygna with (1) external cotyla large; (2)
bicipital attachment large and prominent; (3) carpal tuberosity very
large. Ulna of D. autumnalis with (1) olecranon much thicker; (2)
internal cotyla larger; (3) distal end of external cotyla more rounded;
(4) external condyle longer; (5) carpal tuberosity larger.
Radius of Dendrocygna with (1) shaft immediately posterior to
head very narrow; (2) ulnar facet narrow and U-shaped; (3) capital
tuberosity large; (4) carpal facet narrow, with distinct anconal
projection centrally located. Radius of D. autumnalis with (1)
capital tuberosity smaller, but more prominent shaft flaring more
abruptly to meet a wider distal end.
Carpometacarpus with (1) process of metacarpal I larger; (2)
lobe at end of external rim of carpal trochlea more prominent; (3)
area of fusion between metacarpal II and metacarpal III longer;
(4) internal rim of carpal trochlea larger.
Femur with (1) head rotated dorsally, and concavity between
iliac facet and head greater; (2) iliac facet larger and more
irregularly shaped; (3) trochanter larger and more prominent; (4)
distal end wider; (5) ligamental attachment at base of fibular condyle
more prominent; (6) condyles larger; (7) rotular groove deeper.
Tibiotarsus with (1) shaft reasonably straight and stout (in
D. viduata and D. bicolor there is a greater internal concavity and
distal third of shaft has larger posteriorly directed convexity);
(2) inner cnemial crest larger; (3) condyles not reflected anteriorly.
Tarsometatarsus with (1) intercotylar prominence less prominent;
(2) cotylae large; (3) external edge of hypotarsus longer, positioned
proximally; (4) external trochlea slightly narrower, but middle and
internal trochleae wider; (5) internal trochlea projecting less
distally.
Remarks. D. autumnalis is recorded in western Ecuador south
to the southernmost coastal province (Chapman, 1926), and occasionally
on the coast of Peru (Koepcke, 1970). However, it was not recorded
by Marchant (1958) on the Santa Elena Peninsula of Ecuador. The
presence of this species may indicate an abundance of vegetation
surrounding the fossil site because of the species preference for
quiet waters surrounded by forests (Delacour, 1954).
Subfamily Tadorninae Reichenbach
"Humerus with (1) fairly prominent capital shaft ridge directed
toward external tuberosity; (2) area of origin of external head of
triceps relatively narrow; (3) deltoid crest relatively large and
flaring, and in lateral view more rounded, or if an abrupt bend
occurs, it lies more posteriad; (4) deltoid crest extending farther
distad; (5) head rotated so that external tuberosity is higher."
(Woolfenden, 1961, p. 108)
Tadorninae, n. gen.
Holotype. Tadorninae, n. gen. n. sp.
Diagnosis. Humerus resembles that of Lophonetta Riley and
differs from Chloephaga Eyton and Neochen Salvadori by having (1)
head more rounded in anconal view; (2) external tuberosity reduced
and elongated; (3) external head of triceps only slightly, if any,
undercutting head; (4) deltoid crest shorter and less flaring; (5)
capital groove rotated internally, reducing size of internal tuberosity
and increasing size of head; (6) attachment of anterior articular
ligament less elevated; (7) shaft flaring less to meet ectepicondyle;
(8) internal condyle elongated, less bulbous.
IIumerus differs from that of Lophonetta by having (1) attachment
of M. supracoracoideus on external tuberosity proportionately more
elongated; (2) capital groove narrower and straighter; (3) deltoid
crest shorter and less flaring,.with sharp angle at its midpoint;
(4) bicipital crest shorter and more rounded; (5) pneumatic fossa
completely rimmed by heavy bone, such that distal edge is elevated
above floor of bicipital crest; (6) pneumatic fossa with highest
point located under tip of internal tuberosity (under median crest in
Lophonetta); (7) shaft with greater palmar flexure in distal quarter,
such that line drawn along anterior edge of shaft would almost bisect
distal-most point of ectepicondyle (bisects anterior edge of ectepicon-
dyle in Lophonetta); (8) ectepicondylar prominence smaller, with
narrow ridge along proximal portion; (9) ectepicondyle less rounded
and more V-shaped, extending farther distally; (10) attachment of
anterior articular ligament wider than it is long (longer than wide
in Lophonetta), less elevated, and located farther distally; (11)
entepicondyle more prominent and rounded, not as large, with larger
entepicondylar prominence and external curvature to anconal side;
(12) condyles more rounded, giving deeper and narrower intercondylar
groove; (13) external condyle shorter, retaining internal flexure
found in Lophonetta. but lacking distinct ridge leading internally
from tip of condyle; (14) shaft widening more gradually toward distal
end.
Tadorninae, n. gen. n. sp.
Holotype. Complete right humerus. (P1. III, Fig. D)
Paratypes. One complete right and 3 complete left humeri,
proximal ends of 6 right and 5 left humeri, distal ends of 2 right
and 7 left humeri.
Diagnosis. As for genus. For measurements see Table 3.
Measurements of the holotype are as follows: length, 68.6; proximal
width, 14.7; distal width, 11.0; least width of shaft, 5.2.
Referred material. Three right and 4 left scapulae, 8 complete
right and 7 complete left coracoids, humeral ends of 2 right and 1 left
coracoid, 4 complete right and 3 complete left ulnae, proximal ends of
5 right and 3 left ulnae, distal ends of 5 right and 3 left ulnae,
2 complete left radii, proximal ends of 3 left radii, distal end of
1 right radius, 3 complete right and 2 complete left carpometacarpi,
proximal ends of 2 right and 6 left carpometacarpi, 2 complete right
and 2 complete left femora, proximal end of 1 right femur, 1 complete
left tibiotarsus, distal ends of 4 right tibiotarsi, 4 complete right
and 5 complete left tarsometatarsi, proximal end of 1 left tarso-
metatarsus. The 112 specimens, including holotype and paratypes,
represent a minimum of 10 individuals.
Characters. All elements of Tadorninae, n. gen. n. sp. agree
with the humerus in resembling Lophonetta more than Chloephaga or
Neochen. Tadorninae, n. gen. n. sp. differs from Lophonetta
specularioides by having scapula with (1) coracoidal articulation
proportionately more prominent, confluent with glenoid facet; (2)
intermuscular line between attachments of M. rhomboidius superficialis
and M. subscapularis better developed; (3) glenoid facet rotated such
that it faces more anteriorly and distal end is displaced dorsally.
Coracoid with (1) head projecting less anteriorly; (2) head
with width much greater; (3) glenoid facet flaring less laterally;
(4) scapular facet larger.
Ulna with (1) attachment of anterior articular ligament more
oval and less elongated; (2) external cotyla shorter but wider; (3)
external condyle longer but less prominent; (4) carpal tuberosity
more prominent.
Radius with (1) capital tuberosity more prominent, and shaft
more constricted immediately distal to it; (2) carpal facet less
curved; (3) shaft with internal flexure at distal end.
Carpometacarpus with (1) attachment of M. flexor carpi ulnaris
brevis deeper; (2) external scapho-lunar ligamental attachment small,
lying proximal to ridge extending from external cuneiform ligamental
attachment to proximal fornix, and lying very close to external
cuneiform ligamental attachment (the large size and posterior position
of this attachment in other genera of the subfamily Tadorninae is
listed as a subfamilial character by Woolfenden (1961)); (3) lobe
at end of external rim of carpal trochlea, beginning at notch in rim,
very long and prominent; (4) internal rim of carpal trochlea more
prominent, giving more rounded proximal end in internal view; (5)
internal ligamental fossa deeper; (6) area of fusion of metacarpal I
and metacarpal II much shorter both proximally and distally.
Femur with (1) neck more constricted; (2) area between anterior
corner of trochanter and head more concave but similarly curved,
not angular; (3) ridge on posterior surface of shaft leading to
external condyle more prominent; (4) fibular condyle similar in
shape, but less produced proximally; (5) external condyle well
rounded where joined by fibular condyle (pointed in L. specularioides);
(6) pit for M. tibialis anticus positioned more distally; (7) ridge
on shaft leading distally to internal condyle, as traced by inter-
muscular line, less developed; (8) lip of internal condyle larger.
Tibiotarsus with (1) rotular crest forming straight line
(curved in L. specularioides); (2) outer cnemial crest thickened on
distal side; (3) intermuscular line curving internally proximally
to approach attachment of M. flexor digitorum longus (curves less
internally in L. specularioides); (4) external ligamental attachment
elliptical (similar in L. specularioides), but positioned more
proximally.
Tarsometatarsus with (1) anterior lip of internal cotyla more
undercut; (2) intercotylar prominence with greater anterior, but
smaller dorsal component; (3) posterior border of internal cotyla
with more distinct lip; (4) trochleae very spread (similar in L.
specularioides), but internal and external trochleae turned slightly
more medially; (5) external rim of external trochlea less elongated;
(6) internal side of middle trochlea lacking prominent internally
directed ridge.
Remarks. Tadorninae, n. gen. n. sp. is the smallest represent-
ative of the subfamily known to date, and it is considerably smaller
than related living South American species. Agreement of humeral
Table 3. Measurements of the scapula, coracoid, humerus, radius,
ulna, carpometacarpus, femur, tibiotarsus, and tarsometatarsus of
Tadorninae, n. gen, n. sp. and Lophonetta specularioides (in mm).
Tadorninae
n. gen. n. sp.
Scapula
Proximal
Height
Proximal OR
Width M
N
Coracoid
Length
Head to
Scapular
Facet
Proximal OR
Depth M
N
Least
Depth of
Shaft
Length of
Sternal
Facet
Humerus
Length
Proximal OR
Width M
N
Distal OR
Width M
N
8.2-9.0
8.6
7
2.5-2.8
2.6
7
34.3-36.4
35.4
14
11.8-13.0
12.3
18
6.2-7.6
6.9
18
3.5-4.0
3.8
15
13.4-14.9
14.2
4
61.9-68.6
65.0
4
13.9-14.7
14.5
16
9.9-11.0
10.5
14
Lophonetta
specularioides
11.9-12.9
12.5
3
3.6-3.8
3.7
3
45.4-51.0
49.1
3
16.7-19.2
18.2
3
9.5-10.1
9.8
3
5.1-5.7
5.5
3
17.7-20.3
19.4
3
95.3-101.8
99.5
3
19.0-22.4
21.1
3
14.5-15.7
15.3
3
Table 3. Continued.
Tadorninae
n. gen. n. sp.
Least
Width of
Shaft
Radius
Length
Proximal OR
Width M
N
Distal
Width
Ulna
Length
Proximal OR
Width M
N
Distal OR
Depth M
N
Least
Width of
Shaft
Carpometacarpus
Length OR
M
N
Height OR
Through M
Metacarpal I N
4.6-5.2
4.9
11
51.5-52.9
52.2
2
3.5-3.9
3.6
5
4.8-5.2
5.0
3
54.7-58.0
56.3
4
7.1-7.7
7.4
11
6.3-7.6
7.0
14
3.4-4.0
3.7
7
38.4-41.2
40.3
5
9.2-9.9
9.5
12
Lophonetta
specularioides
6.1-6.5
6.3
3
82.6-86.8
85.1
3
4.9-5.2
5.0
3
6.1-7.3
6.9
3
89.3-94.7
92.1
3
10.0-11.8
10.9
3
9.5-10.8
10.2
3
4.8-5.6
5.1
3
57.1-60.7
59.1
3
12.9-15.0
14.2
3
Table 3. Continued.
Tadorninae
n. gen. n. sp.
Lophonetta
specularioides
Proximal OR
Width M
N
Least
Width of
Shaft
Length of
Distal
Fornix
Femur
Length
Proximal OR
Width M
N
Distal OR
Width M
N
Least
Width of
Shaft
Tibiotarsus
Length
Proximal OR
Width M
N
Distal
Width
4.1-4.6
4.4
12
3.3-4.6
3.6
5
4.9-5.3
5.2
5
36.9-37.3
37.1
3
7.5-8.0
7.7
5
8.0
8.0
3
3.2-3.3
3.2
5
5.9-6.7
6.4
3
3.9-4.8
4.5
3
8.9-9.1
9.0
3
52.6-59.1
56.1
3
9.9-11.2
10.5
3
11.7-12.9
12.5
3
4.8-5.8
5.3
3
85.7-95.4
92.1
3
9.6-10.4
10.0
3
9.4-10.4
10.0
3
6.5-6.9
6.6
5
Table 3. Continued.
Tadorninae
n. gen. n. sp.
Least
Width of
Shaft
Depth of
Internal
Condyle
Tarsometatarsus
Length OR
M
N
Proximal OR
Width M
N
Distal
Width
Least
Width of
Shaft
Width of
Middle
Trochlea
7.2-7.3
7.3
5
33.6-38.5
35.3
7
6.8-7.4
7.1
9
7.2-7.8
7.5
7
3.5-3.8
3.6
10
2.8-3.3
3.1
9
Lophonetta
specularioides
4.3-4.7
4.4
3
10.5-11.0
10.8
3
51.0-56.9
54.3
3
10.3-10.9
10.7
3
10.4-11.0
10.8
3
4.0-4.5
4.3
3
4.6-5.0
4.8
3
characters with those of the subfamily as listed by Woolfenden (1961)
indicate that it is placed in the correct subfamily, despite its
small size and variance with a few other subfamilial characters in
other elements.
Genus Chloephaga Eyton
Chloephaga melanoptera (Eyton)
Andean Goose
Material. One right quadrate, 1 right and 2 left scapulae, 4
complete right and 2 complete left coracoids, humeral ends of 1 right
and 3 left coracoids, 1 complete left humerus, proximal ends of 1
right and 4 left humeri, distal ends of 1 right and 5 left humeri,
1 complete right ulna, proximal end of 1 right ulna, distal ends
of 5 right and 4 left ulnae, proximal end of 1 right and 1 left
radius, distal ends of 2 right and 1 left radius, 4 complete right
and 1 complete left carpometacarpus, proximal ends of 4 right and 2
left carpometacarpi, 1 complete pelvis, 2 complete right and 1
complete left femur, proximal ends of 2 right and 4 left femora,
distal ends of 4 right and 4 left femora, proximal end of 2 left
tibiotarsi, distal ends of 3 right and 6 left tibiotarsi, 1 complete
left tarsometatarsus, proximal ends of 2 right and 2 left tarsometatarsi,
distal ends of 2 right and 1 left tarsometatarsus. The 91 specimens
represent a minimum of 7 individuals.
Characters. The large size of Chloephaga readily separates it
from other genera of the subfamily Tadorninae found in South America,
and indeed, in size of individuals it is the largest genus of the family
in South America. As the fossil specimens did not differ from the C.
melanoptera available for comparison, and C. melanoptera is the only
species of the genus occurring in Peru today, the material is referred
to that species.
Remarks. C. melanoptera is primarily resident in the high Andes,
living up to the snow line. It does, however, migrate to lower
altitudes during the non-breeding season. It has not been recorded
in the Andes closer than 100 miles south of the fossil site (Koepcke,
1970). Because this species feeds almost exclusively on grass and is
seldom seen on water (Delacour, 1954), the presence of an abundant
ground cover of vegetation near the fossil site would seem to be more
likely to attract this species than a body of water surrounded by
sparse vegetation.
A number of the specimens appear to represent immature individuals.
Subfamily Anatinae (Vigor)
"Humerus with (1) capital shaft ridge obsolete; (2) pneumatic
fossa opening, containing bony struts; (3) ectepicondyle typically
equal or subequal in height to entepicondyle." (Woolfonden, 1961,
p. 109)
Genus Cairina Fleming
Cairina moschata (Linnaeus)
Muscovy Duck
Material. One right articular, 4 left scapulae, 2 complete
right coracoids, humeral end of 1 right coracoid, proximal ends of
2 right humeri, distal ends of 2 right and 1 left humerus, 2 complete
right ulnae, proximal end of 1 left ulna, distal ends of 1 right and 3
left ulnae, proximal ends of 1 right and 1 left radius, distal ends
of 1 right and 1 left radius, 1 complete left carpometacarpus,
proximal ends of 1 right and 1 left carpometacarpus, distal end of
2 right tibiotarsi, 1 complete left tarsometatarsus, proximal end of
1 left tarsometatarsus, distal end of 2 left tarsometatarsi. The 33
specimens represent a minimum of 4 individuals.
Characters. Cairina is easily separated from Sarkidiornis, the
most closely related genus present in South America.
Remarks. C. moschata, a forest duck, occurs on the arid coast
of Peru only by accident today. It has been recorded only once, near
Lima. Marchant (1958) did not report it from the arid Santa Elena
Peninsula of Ecuador. Chapman (1926) reported this species near the
coast only 20 miles north of the Peru-Ecuador border, and less than
120 miles from the fossil site, but in a completely different type of
habitat.
The sedentary habits of this species would seem to indicate that
either its range extended much farther south than today, increasing
the number of stragglers, or that it was in residence. If Cairina was
in residence its preference for water surrounded by forests would
indicate a considerable amount of vegetation in the area.
Genus Anas Linnaeus
Anas n. sp. 1
Holotype. Complete left humerus (internal tuberosity and
bicipital crest broken). (P1. III, Fig. E)
Diagnosis. Agrees with Anas and differs from other genera of
subfamily Anatinae by having those characters of the genus as
described by Woolfenden (1961).
Differs from living species of Anas by having (1) head of humerus
small, profile rounded in palmar view, and projecting more anconally
than distally (similar in Anas (Callonetta) leucophrys Vieillot; head
large, subrounded, and projecting more distally than anconally in
Anas (Amazonetta) brasiliensis Gmelin, A. crecca Linnaeus, and A.
cyanoptera (Vieillot)); (2) head without inflection (similar in A.
leucophrys, A. crecca, and A. cyanoptera; head inflected in A.
brasiliensis); (3) internal tuberosity broken, but appearing
prominent (similar in A. brasiliensis, A. crecca and A. cyanoptera;
reduced in A. leucophrys); (4) attachment of M. coracobrachialis
posterior forming short, broad shelf, not extending down median crest
(moderate to long, broad, usually not forming shelf, and usually ex-
tending down median crest in A. leucophrys, A. brasiliensis, A.
crecca, and A. cyanoptera); (5) median crest prominent (similar in A.
leucophrys and A. brasiliensis; slender to moderately prominent in
A. crecca and A. cyanoptera); (6) pneumatic fossa elongated, not
entering into internal tuberosity (moderately elongated, not entering
into internal tuberosity in A. brasiliensis; slightly elongated and
entering into internal tuberosity in A. leucophrys, A. crecca, and
A. cyanoptera); (7) deltoid crest not flaring, with depression at
distal end (similar in A. crecca; flaring, without depression in A.
leucophrys and A. brasiliensis; not flaring, without depression, and
reduced in size in A. cyanoptera); (8) attachment of M. pectoralis
superficialis rotated palmarly (similar in A. brasiliensis; not
rotated in A. leucophrys, A. crecca, and A. cyanopteral; (9) shaft
with moderate internal curvature immediately distal to bicipital
crest, giving moderately sigmoid shaft (similar in A. leucophrys;
sharp internal curvature in A. brasiliensis, giving strongly sigmoid
shaft; slight internal curvature in A. crecca and A. cyanoptera,
giving weakly sigmoid shaft); (10) impression of M. brachialis anticus
relatively deep, oriented at small angle to axis of shaft (deep in
A. brasiliensis; shallow in A. leucophrys, A. crecca, and A. cyanoptera;
oriented at large angle to axis of shaft in all); (11) attachment of
anterior articular ligament longer than wide (width roughly equal to
length in A. leucophrys, A. brasiliensis, A. crecca, and A. cyanoptera);
(12) entepicondyle rotated so internal side faces antero-distally
(rotated strongly anteriorly in A. leucophrys and A. brasiliensis;
rotated slightly to moderately anteriorly in A. cyanoptera; not
rotated in A. crecca); (13) entepicondyle rounded distally, not
produced (subrounded distally and very produced in A. leucophrys;
subangular distally and moderately produced in A. brasiliensis;
angular to subangular distally and moderately produced in A. crecca
and A. cyanoptera); (14) internal tip of external condyle broken
(undercut in A. leucophrys, A. brasiliensis, A. crecca, and A.
cyanoptera); (15) olecranol fossa long (similar in A. loucophrys and
A. cyanoptera; short in A. brasiliensis and A. crecca).
For measurements see Table 4. Measurements of the holotype are
as follows: length, 56.4; proximal width, 13; distal width, 9.2;
least width of shaft, 4.3.
Referred material. One right scapula, 2 complete right coracoids,
1 complete right and 1 complete left ulna, distal end of 1 right
ulna, proximal end of 1 left radius, distal end of 1 right radius,
proximal end of 1 right carpometacarpus, distal end of 1 right and 1
left tibiotarsus, 1 complete left tarsometatarsus. The 13 elements,
including holotype, represent a minimum of 2 individuals.
Characters. Scapula with (1) acromion of moderate length,
narrow, with strong anterior curvature (wide, of similar length and
curvature in A. leucophrys; short and wide, with very slight anterior
curvature in A. crecca; long and wide, with moderate anterior curvature
in A. cyanoptera); (2) glenoid facet of moderate length and narrow,
lacking strong projection to antero-ventral corner (similar, but long,
in A. leucophrys; long and wide, with prominent antero-ventral corner
in A. brasiliensis, A. crecca, and A. cyanoptera); (3) attachment of
Lig. furculo-scapulare dorsale oval, located dorsal to midline of shaft
(similar in A. leucophrys, A. brasiliensis, and A. crecca; elongated
and situated ventral to midline of shaft in A. cyanoptera).
Coracoid with (1) coracohumeral surface long, relatively wide,
not widening markedly anteriorly, with moderate curvature, and
positioned internally to edge of triosseal canal (short, narrow, not
widening markedly anteriorly, of moderate curvature, positioned
slightly internally in A. leucophrys; long, wide, widening markedly
anteriorly, of slight curvature, and positioned internally in A.
brasiliensis; long, narrow, moderate to marked widening anteriorly,
of slight to moderate curvature, not positioned internally in A.
crecca and A. cyanoptera); (2) attachment of Lig. humero-coracoideum
anterius superius slightly elevated (similar in A. leucophrys and A.
brasiliensis; not elevated in A. crecca and A. cyanoptera); (3)
scapular facet small and elongated (similar in A. leucophrys and A.
brasiliensis; large oval in A. crecca and A. cyanoptera).
Ulna with (1) distal lip of external cotyla broad, extending
moderately distally (similar in A. brasiliensis; internal side of
external cotyla extends distally much farther than external side in
A. leucophrys, A. crecca, and A. cyanoptera); (2) ridge leading to
shaft from carpal tuberosity large (similar in A. crecca; slight to
moderate ridge in A. leucophrys, A. brasiliensis, and A. cyanoptera).
Radius with no distinguishing characters other than size.
Carpometacarpus with (1) process of metacarpal I low and of
moderate length (low, with distal edge sloping sharply proximally in
A. leucophrys; high and short in A. brasiliensis; of moderate height
and length in A. crecca and A. cyanoptera); (2) proximal edge of
metacarpal I sloping sharply proximally (similar in A. leucophrys
and A. cyanoptera; moderate slope in A. crecca; slight slope in A.
brasiliensis); (3) internal face of metacarpal I slightly excavated
(similar in A. crecca; moderately to deeply excavated in A. leucophrys,
A. brasiliensis, and A. cyanoptera); (4) proximal area of fusion of
metacarpal II and metacarpal III very long (long in A. leucophrys
and A. brasiliensis; short in A. crocca and A. cyanoptera); (5)
external rim of carpal trochlea rounded proximally (subangular to
angular in A. leucophrys, A. brasiliensis, A. crecca, and A. cyanoptera);
(6) area covered by external cuneiform ligament markedly convex (similar
in A. leucophrys; slightly convex in A. brasiliensis; flat in A.
crecca and A. cyanoptera); (7) posterior carpal fossa of essentially
uniform width (similar in A. leucophrys and A. brasiliensis; wide
distally, narrowing gradually proximally in A. crecca; wide prox-
imally and distally with central constriction in A. cyanoptera).
Tibiotarsus with (1) external face of distal end flat (similar
in A. leucophrys, A. brasiliensis, and A. crecca; flat or slightly
concave in A. cyanoptera); (2) internal condyle flaring medially
(not flaring in A. leucophrys, A. brasiliensis, A. crecca and A.
cyanoptera).
Tarsometatarsus with (1) intercotylar prominence broad (similar
in A. leucophrys, A. brasiliensis, A. crecca, and A. cyanoptera); (2)
shaft narrowing sharply immediately distal to internal cotyla
(similar in A. leucophrys, A. brasiliensis, and A. crecca; narrows
gradually in A. cyanoptera); (3) hypotarsus short (similar in
A. leucophrvs and A. crecca; long in A. brasiliensis and A. cyanoptera);
(4) anterior face of shaft straight in lateral view (similar in A.
cyanoptera; slight concavity in A. brasiliensis and A. crecca;
moderate concavity in A. leucophrysl; (S) internal trochlea projecting
distally without turning externally (similar in A. leucophrys, A.
brasiliensis, and A. cyanoptera; turned slightly externally in A.
crecca).
Remarks. Anas n. sp. 1 and the following two new species of
Anas are compared with A. leucophrys, A. brasiliensis, A. crecca,
and A. cyanoptera because the fossil species are all small and the
four living species represent the smallest forms of the genus.
Although A. crecca does not reach South America it was used rather
than A. discors, which does reach South America, because the two
species are extremely difficult to separate osteologically, and of
the two the former has smaller individuals.
The numbers under each element correspond in the comparisons of
all three of the new species.
Anas n. sp. 2
Holotype. Complete left humerus. (P1. IV, Fig. D)
Paratypes. One complete right and 1 complete left humerus,
proximal ends of 3 right and 1 left humerus, distal ends of 2 right
and 1 left humerus.
Diagnosis. Agrees with Anas and differs from other genera of
the subfamily Anatinae by having those characters of the genus as
described by Woolfenden (1961).
Differs from other species of Anas by having (1) head large,
profile subrounded in palmar view, and projecting more anconally than
distally; (2) head not inflected; (3) internal tuberosity reduced;
(4) attachment of M. coracobrachialis posterior moderate to long,
broad, forming slight shelf, extending down median crest; (5) median
crest prominent; (6) pneumatic fossa round, entering moderately into
internal tuberosity; (7) deltoid crest long, very flaring, without
depression at distal end; (8) attachment of M. pectoralis superficialis
not rotated palmarly; (9) shaft with sharp internal curvature
immediately distal to bicipital crest, giving strongly sigmoid shaft;
(10) impression of M. brachialis anticus extremely shallow, oriented
at large angle to axis of shaft; (11) attachment of anterior articular
ligament high and very small; (12) entepicondyle rotated moderately
anteriorly; (13) entepicondyle rounded distally, not produced; (14)
internal lip of external condyle not undercut, but with slight ridge
leading internally; (15) olecranol fossa short.
For measurements see Table 4. Measurements of the holotype are
as follows: length, 65.2; proximal width, 14.1; distal width, 9.7;
least width of shaft, 4.5.
Referred material. One left scapula, 2 complete right and 4
complete left coracoids, 1 complete right ulna, proximal ends of 2
left radii, 1 complete left carpometacarpus, proximal ends of 1 right
and 1 left carpometacarpus, proximal end and shaft of 1 right femur,
distal ends of 5 right and 1 left tibiotarsus, 1 complete left tarso-
metatarsus. The 29 specimens, including holotype and paratypes,
represent a minimum of 5 individuals.
Characters. Scapula with (1) acromion long and wide, with
moderate anterior curvature; (2) glenoid facet long and wide, with
antero-ventral corner not projecting; (3) attachment of Lig. furculo-
scapulare dorsale round, located on midline of shaft.
Coracoid with (1) coracohumeral surface wide and of moderate
length, with moderate to marked widening anteriorly, moderate to
marked curvature, and positioned slightly internally; (2) attachment
of Lig. humero-coracoideum anterius superius elevated; (3) scapular
facet large oval.
Ulna with (1) internal side of external cotyla extending much
farther distally than external side; (2) ridge leading from carpal
tuberosity to shaft very small.
Radius with no distinguishing characters other than size.
Carpometacarpus with (1) process of metacarpal I long and low;
(2) proximal edge of metacarpal I sloping slightly proximally; (3)
proximal area of fusion of metacarpal II and metacarpal III long, and
distal area of fusion short (distal area short in A. leucophrys; long
in A. brasiliensis and A. crecca; very long in A. cyanoptera); (4)
internal face of metacarpal I slightly excavated; (5) external rim
of carpal trochlea subangular proximally; (6) area covered by
external cuneiform ligament markedly convex; (7) posterior carpal
fossa wide distally, narrowing abruptly proximally.
Femur with (1) head large, projecting medially (small to
moderate sized head projecting medially in A. leucophrys, A. crecca,
and A. cyanoptera; very large head projecting equally dorsally and
medially in A. brasiliensis); (2) attachment of M. ilio-trochantericus
medius very long (long in A. leucophrys; of moderate length in A.
brasiliensis and A. crecca; short in A. cyanoptera); (3) iliac facet
curving sharply medially posteriorly (moderate curvature in A.
leucophrys and A. brasiliensis; slight if any curvature in A. crecca
and A. cyanoptera).
Tibiotarsus with (1) external face of distal end markedly
concave; (2) internal condyle not,flaring medially.
Tarsometatarsus with (1) intercotylar prominence broad; (2)
shaft narrowing sharply immediately posterior to internal cotyla;
(3) hypotarsus long; (4) anterior face of shaft straight in lateral
view; (5) internal trochlea moderately turned externally.
Anas n. sp. 3
Holotype. Complete left humerus. (PI. IV, Fig. E)
Paratypes. Proximal end of 1 right humerus, distal end of 1
right humerus.
Diagnosis. Agrees with Anas and differs from other genera of
the subfamily Anatinae by having those characters of the genus as
described by Woolfenden (1961).
Differs from other species of Anas by having (1) head large,
profile rounded in palmar view, and projecting more distally than
anconally; (2) head not inflected; (3) internal tuberosity reduced;
(4) attachment of M. coracobrachialis posterior short broad shelf, not
extending down median crest; (5) median crest moderately prominent;
(6) pneumatic fossa slightly elongated, entering only slightly into
internal tuberosity; (7) deltoid crest long, moderately flaring,
without depression at distal end; (8) attachment of M. pectoralis super-
ficialis not rotated palmarly; (9) shaft with moderate internal curva-
ture immediately distal to bicipital crest, giving moderately sigmoid
shaft; (10) impression of M. brachialis anticus shallow, oriented at
large angle to axis of shaft; (11) attachment of anterior articular
ligament as wide as it is long; (12) entepicondyle rotated moderately
anteriorly; (13) entepicondyle rounded distally, not produced, very
small; (14) internal tip of external condyle not undercut, but with
slight ridge present leading internally; (15) olecranol fossa short.
For measurements see Table 4. Measurements of the holotype are
as follows: length, 57.7; proximal width, 12.3; distal width, 9.1;
least width of shaft, 4.4.
Referred material. One complete left coracoid, distal end of 1
left tibiotarsus, 1 complete right tarsometatarsus. The 5 specimens,
excluding holotype but including paratypes, represent a minimum of
1 individual.
Characters. Coracoid with (1) coracohumerual surface short,
narrow, widening moderately anteriorly, of moderate curvature, and
not positioned internally; (2) attachment of Lig. humero-coracoideum
anterius superius elevated; (3) scapular facet large oval.
Tibiotarsus with (1) external face of distal end flat; (2)
internal condyle flaring medially.
Tarsometatarsus with (1) intercotylar prominence slender;
(2) shaft narrowing sharply immediately posterior to internal
cotyla; (3) hypotarsus short; (4) anterior face of shaft markedly
convex in lateral view; (5) internal trochlca strongly turned
externally.
Remarks. The holotype of Anas n. sp. 3 is from La Carolina,
Ecuador, not the Talara Tar Seeps. The Ecuadorian specimen was
chosen for the holotype because of its completeness and the lack of
a complete specimen in the Talara material. The paratypes and
all referred material are from the Talara Tar Seeps.
Anas bahamensis Linnaeus
Bahama Pintail
Material. Portions of 14 crania, 5 frontals, 9 right and 11
left quadrates, 3 right and 3 left articulars, 5 right and 7 left
Table 4. Measurements of the scapula, coracoid, humerus, radius,
ulna, carpometacarpus, femur, tibiotarsus, and tarsometatarsus of
Anas n. sp. 1 (excluding femur), Anas n. sp. 2, Anas n. sp. 3
(coracoid, humerus, tibiotarsus, and tarsometatarsus only), A.
leucophrys, A. brasiliensis, A. cyanoptera, and A. crecca
carolinensis (in mm).
Anas
n. sp. 1
Anas
n. sp. 2
Anas
n. sp. 3
Scapula
Proximal
Height
Proximal OR
Width M
N
Coracoid
Length
Head to
Scapular
Facet
Proximal
Depth
Least
Depth of
Shaft
Length of
Sternal
Facet
Humerus
Length
Proximal
Width
2.4
1
31.7
1
10.9
1
6.4
1
3.2-3.7
3.5
2
56.4
1
12.7
1
32.9-33.7
33.4
3
10.7-11.6
11.2
4
5.9-6.5
6.3
4
3.4-3.9
3.6
6
12.2-13.5
12.9
2
61.1-65.2
63.1
2
13.2-14.1
13.5
4
57.7
1
12.3-13.1
12.7
2
Table 4. Extended.
Anas Anas
leucophrys brasiliensis
7.1
1
2.0
1
32.8
1
11.2
1
6.6
1
4.3
1
12.4
1
59.8
1
13.5
1
9.5
1
3.0
1
41.1
1
13.8
1
8.3
1
4.7
1
16.4
1
74.8
1
17.0
1
Anas
cyanoptera
7.5-9.6
8.3
5
2.4-2.7
2.5
5
32.6-39.8
35.1
5
10.8-13.5
11.8
5
6.2-7.5
6.7
5
3.3-4.1
3.7
5
12.7-16.0
13.7
5
61.0-71.5
64.6
5
12.7-15.7
13.6
5
Anas
crecca
7.3-8.1
7.7
8
2.5-3.0
2.7
8
31.5-34.1
32.6
8
10.8-11.5
11.1
8
6.1-7.0
6.3
8
3.2-4.0
3.6
8
12.8-14.4
13.6
8
55.9-59.5
57.9
8
12.5-13.2
12.8
8
Table 4. Continued.
Anas
n. sp. 1
Distal
Width
Least
Width of
Shaft
Radius
Length
Proximal OR
Width M
N
Distal OR
Width M
N
Ulna
Length OR
M
N
Proximal OR
Width M
N
Distal
Depth
Least
Width of
Shaft
Carpometacarpus
Length
Anas
n. sp. 2
9.6-10.1
9.8
5
Anas
n. sp. 3
9.1-9.4
9.2
2
4.3-4.8
4.6
3.6-3.7
3.7
2
56.0
1
7.6
1
7.6
1
3.8
1
53.0-53.6
53.3
2
6.8
1
6.5-6.7
6.6
3
3.2-3.5
3.4
2
Table 4. Extended.
Anas Anas Anas Anas
leucophrys brasiliensis cyanoptera crecca
9.7 12.6 9.1-11.7 8.3-9.7
9.9 9.3
1 1 5 8
4.7 5.8 4.4-5.1 4.2-4.7
S- 4.6 4.4
1 1 5 8
50.7 61.5 46.8-56.7 44.2-46.6
50.5 45.8
1 1 5 8
3.3 4.3 3.1-4.0 3.1-3.3
3.4 3.2
1 1 5 8
4.6 5.7 3.9-5.3 4.2-4.5
4.6 4.3
1 1 5 8
54.7 66.3 50.6-61.1 48.3-51.1
54.4 50.2
1 1 5 8
6.8 8.9 6.6-8.5 6.2-6.9
7.1 6.6
1 1 5 8
6.9 8.6 6.7-8.2 6.6-7.0
7.1 6.8
1 1 5 8
3.4 4.4 3.3-3.9 3.3-3.6
3.5 3.4
1 1 5 8
37.6 46.0 36.0-43.9 34.5-37.1
38.9 36.2
1 1 5 8
Table 4. Continued.
Anas
n. sp. 1
Proximal OR
Width M
N
Height OR
Through M
Metacarpal I N
Least OR
Width of M
Shaft N
Length of OR
Distal M
Fornix N
Femur
Length OR
M
N
Proximal OR
Width M
N
Distal OR
Width M
N
Least
Width of
Shaft
Tibiotarsus
Length
Proximal OR
Width M
N
6.3-6.5 6.
6.5
6 1
Anas
n. sp. 3
Anas
n. sp. 2
4.3
4.3
3
8.9
8.9
2
3.0-3.3
3.1
2
Distal
Width
6.1-6.5
6.3
2
Table 4. Extended.
Anas
leucophrys
3.9
1
8.3
1
3.2
1
4.9
1 .
33.6
1
7.1
1
7.4
1
2.9
1
55.1
1
6.1
1
5.8
1
Anas
brasiliensis
4.9
1
11.0
1
3.8
1
7.2
1
43.4
1
9.3
1
9.1
1
3.4
1
67.7
1
7.4
1
7.5
1
Anas
cyanoptera
3.9-4.7
4.1
5
8.7-10.2
9.2
5
2.8-3.7
3.3
5
4.9-6.8
6.1
5
31.5-38.3
34.8
5
6.1-7.5
6.7
5
6.5-8.4
7.2
5
2.8-3.2
2.9
5
53.1-64.5
57.7
5
5.2-6.9
6.1
5
5.4-6.7
5.9
5
Anas
crecca
3.8-4.1
3.9
8
8.4-9.0
8.7
8
2.8-3.3
3.1
8
4.5-5.3
5.1
8
31.9-33.6
32.9
8
6.1-6.8
6.6
8
6.7-7.3
6.8
8
2.6-3.0
2.8
8
52.7-56.5
54.4
8
5.5-6.0
5.7
8
5.3-5.8
5.5
8
u
Table 4. Continued.
Anas
n. sp. 1
Least
Width of
Shaft
Depth of
Internal
Condyle
Tarsometatarsus
Length
Proximal
Width
Distal
Width
Least
Width of
Shaft
Width of
Middle
Trochlea
Anas
n. sp. 2
2.9-3.2
3.0
3
6.7-7.3
7.0
4
35.6
1
Anas
n. sp. 3
29.5
1
Table 4. Extended.
Anas
leucophrys
2.7
1
6.6
1
29.9
1
6.5
1
6.5
1
3.0
1
2.9
1
Anas
brasiliensis
3.5
1
8.9
1
38.4
1
7.8
1
7.7
1
3.9
1
3.7
1
Anas
cyanoptera
2.5-2.9
2.7
5
6.3-7.8
6.8
5
31.3-35.5
32.8
5
6.1-7.0
6.3
5
5.5-6.7
6.2
5
2.7-3.3
3.1
5
2.6-3.3
2.7
5
Anas
crecca
2.4-2.8
2.6
8
6.0-6.6
6.3
8
28.1-31.8
29.6
8
5.7-6.2
5.9
8
5.7-6.0
5.8
8
2.5-3.0
2.8
8
2.5-2.9
2.7
8
mandibular fragments, 122 right and 106 left scapulae, 172 complete
right and 128 complete left coracoids, humeral ends of 71 right and
87 left coracoids, 35 complete right and 34 complete left humeri,
proximal ends of 86 right and 104 left humeri, distal ends of 103
right and 110 left humeri, 71 complete right and 64 complete left
ulnae, proximal ends of 69 right and 61 left ulnae, distal ends of
87 right and 78 left ulnae, 14 complete right and 19 complete left
radii, proximal ends of 83 right and 61 left radii, distal ends of
52 right and 41 left radii, 135 complete right and 103 complete left
carpometacarpi, proximal ends of 73 right and 73 left carpometacarpi,
29 complete right and 21 complete left femora, proximal ends of 22
right and 25 left femora, distal ends of 11 right and 7 left femora,
5 complete right and 4 complete left tibiotarsi, proximal ends of 26
right and 15 left tibiotarsi, distal ends of 102 right and 91 left
tibiotarsi, 60 complete right and 65 complete left tarsometatarsi,
proximal ends of 17 right and 19 left tarsometatarsi, distal ends of
24 right and 21 left tarsometatarsi. The 2,864 specimens represent
a minimum of 243 individuals.
Characters. Only two species (A. bahamensis and A. discors
Linnaeus) of the subfamily Anatinae have been previously recorded
from northwestern Peru, although I have seen A. cyanoptera (subspecies
not determined) near Ciudad de Piura, Departemento de Piura (4 October
1971). A. bahamensis is easily separated on size from these two
species, including the largest subspecies of A. cyanoptera, A. c.
orinomus.
Anas flavirostris Vieillot, A. georgica Gmelin, A. puna Tshudi,
and A. platalea Vieillot also occur in Peru today, but not in the
northwestern section. A. bahamensis is readily separable from A. puna
and A. platalea by numerous osteological characters, and from smaller
A. flavirostris by size. Elements of A. bahamensis are usually smaller
than those of A. georgica, the most closely related species, but the
two species overlap in size. A. bahamensis differs from A. georgica
by having scapula with (1) concavity dorsal to proximal end of
glenoid facet deep; (2) acromion longer.
Coracoid with (1) coracohumeral surface narrower and shorter;
(2) head more rounded in medial view; (3) procoracoid projecting
farther dorsally.
Humerus with (1) capital groove narrower; (2) internal tuberosity
projecting less anconally; (3) attachment of Lig. humero-coracoideum
anterius inferius smaller and positioned anconally on internal
tuberosity; (4) concavity immediately external to attachment of M.
proscapulohumeralis much narrower; (5) ectepicondylar prominence
larger; (6) entepicondyle smaller; (7) external condyle narrower.
Ulna with (1) olecranon longer; (2) external cotyla smaller, with
distal projection smaller; (3) internal cotyla more concave.
Radius with (1) capital tuberosity smaller; (2) ligamental
prominence narrower, more distinct, and situated more laterally.
Carpometacarpus with (1) process of metacarpal I slightly longer
and more rectangular, lacking a sharp anterior curvature; (2)
posterior carpal fossa larger; (3) internal scapholunar ligamental
attachment on proximal end of metacarpal III usually positioned
proximally; (4) proximal area of fusion of metacarpal II and metacarpal
III shorter.
Femur with (1) head smaller; (2) iliac facet narrower; (3)
internal condyle not as deep; (4) external condyle not as deep,
more rounded, and extending farther proximally.
Tibiotarsus with (1) concavity between internal articular
surface and rotular crest shallower; (2) area between proximal
internal ligamental attachment and flexor attachment slightly
convex (very convex in A. georgica); (3) condyles less deep.
Tarsometatarsus with (1) cotylae smaller; (2) external ligamental
attachment positioned proximally; (3) trochleae less compressed;
(4) middle trochlea extending less proximally.
Remarks. A. bahamensis is widespread throughout tropical and
subtropical South America, and is capable of rapidly colonizing
impermanent bodies of water in large numbers, even in very arid areas
(Marchant, 1958). It is very characteristic of coastal lagoons and
where found is present in large numbers.
The large number of specimens of this species represents almost
a third of the collection of fossil birds from this site.
I believe material referred to Archeoquerquedula lambrechti
by Spillman (1942) is in actuality Anas bahamensis. Although I have
not seen his material, I do have considerable material from the
same site as that cited by Spillman and it agrees with Anas bahamensis.
Spillman did not describe any of the elements referred to Archeoquer-
quedula lambrechti, but he did figure a skull. From his figure it
is possible to determine that the length of the orbit is greater than
the distance from the posterior edge of the orbit to the parietal.
Of the species of Anas found in South America this condition is
found only in Anas bahamensis. In all other species the distance from
the posteior edge of the orbit to the parietal is greater than the
length of the orbit.
Subfamily Oxyurinae Phillips
Genus Nomonyx Ridgway
Nomonyx dominicus (Linnaeus)
Masked Duck
Material. One complete left humerus.
Remarks. N. dominicus has been recorded as far south in western
Peru as Lambeyeque, and Marchant (1958) reported it as scarce on the
arid Santa Elena Peninsula of Ecuador. It is sedentary in habits,
but strays great distances. It would appear from the singe specimen
that N. dominicus was not a regular resident at or near the fossil
site.
Discussion of Family Anatidae
The nine species of waterfowl found in the Talara Tar Seeps
avifauna represent almost 10 per cent of the non-passerine portion of
the avifauna and give a good indication of what the habitat was like
at the time of their fossilization. Although some of the species
are well known colonizers of temporary ponds (A. bahamensis) or
widely wandering species (Nomonyx dominicus), others indicate the
presence of abundant grasses (Chloephaga melonoptera) or forests
(Dendrocygna autumnalis and Cairina moschata).
The most interesting species are of course the four that are
extinct, and perhaps special noteshould be made that all of them are
of very small size. The relatively large number of elements repre-
senting each of the four species is large enough to remove any doubt
that they are indeed separate species, especially considering the
possibility of confusion in the three new small species of Anas.
The fact that the four extinct species are all small suggests
that although there was adequate water present there may have been
annual periods of stress resulting from dry periods. If there were
such annual periods of stress smaller sized individuals would allow
for a greater density of each species than larger individuals. Of
course there is the possibility that the four species were derived
from similarly sized ancestral species.
That the four species became extinct suggests that they were
endemic to at least northwestern Peru, and southwestern Ecuador in the
case of Anas n. sp. 3, and that there must have been a sufficient
period of stability for them to evolve as endemics. This would require
a long period of adequate water supply, which recently disappeared,
making the habitat uninhabitable. The speciation of the four species
may have occurred much as it is currently occurring in Anas cynopptera,
where one finds three allopatric subspecies of quite different
size ranges in northern South America.
The three new species of Anas tend to fill in some of the larger
gaps in osteological characters between Anas and Anas (Callonetta)
leucophrys and Anas (Amazonetta) brasiliensis noted by Woolfenden
(1961). For this reason I have considered the latter two species
and the three new species of the subfamily Anatinae as members of a
larger, more inclusive genus Anas rather than proposing new genera
for the new intermediate forms.
Order Accipitriformes (Vieillot)
Suborder Sarcoramphi (Ridgway)
Family Vulturidae (Illiger)
Vulturidae, n. gen.
Holotype. Vulturidae, n. gen. n. sp.
Diagnosis. Tarsometatarsus agrees with that of Vultur Linnaeus
and Gymnogyps Lesson by having (1) intercotylar prominence large; (2)
anterior metatarsal groove very deep with large attachment of M.
tibialis anticus located immediately distal to proximal foramina;
(3) hypotarsus flattened, roughly triangular in shape, with central
ridge leading distally; (4) shaft robust, rectangular, widening
proximally and distally to meet articular surfaces; (5) external and
internal trochleae much smaller than middle trochlea and turned
slightly toward midline of shaft; (6) size large.
Tarsometatarsus differs from that of Vultur and Gymnogyps by
having (1) intercotylar prominence very high and narrow (low and broad
in Vultur, high and of moderate width in Gymnogyps); (2) internal
cotyla very deep, narrow, with very high, almost straight internal
edge (deep, wide, internal edge moderately curved in Vultur; deep,
of moderate width, with internal edge rounded in Gymnogyps); (3)
external cotyla round, with articular surface extending onto external
anterior metatarsal ridge (very elongated with articular surface
restricted in Vultur, oval with articular surface restricted in
Gymnogyps); (4) hypotarsus narrow, deeply grooved, moderately notched
externally, with central ridge leading distally narrow (broad, very
deeply grooved, deeply notched externally, with central ridge
broad in Vultur; of moderate width, moderately grooved, moderately
notched externally, with central ridge broad in Gymnogyps); (5)
anterior metatarsal groove of moderate width and appears slightly
pneumatic proximal to attachment of M. tibialis anticus (wide and
highly pneumatic in Vultur, narrow and moderately pneumatic in
Gymnogyps); (6) attachment of M. tibialis anticus very large with
length equal to or exceeding length of anterior metatarsal groove
proximal to it (of moderate size, wider than long, much shorter than
proximal portion of anterior metatarsal groove in Vultur; of moderate
size, length greater than width, much shorter than proximal portion
of anterior metatarsal groove in Gynnogyps); (7) attachment of M.
extensor hallucis longus located approximately halfway between base of
anterior metatarsal groove and top of internal metatarsal ridge
(located near top of internal metatarsal ridge in Vultur and Gymnogyps);
(8) external anterior metatarsal ridge broad and rounded and internal
anterior metatarsal ridge of moderate width but with sharp proximal
edge (broad, rounded external and narrow, moderately rounded internal
anterior metatarsal ridge in Vultur; both internal and external
anterior metatarsal ridge broad and rounded in Gymnogyps); (9)
shaft moderately depressed external to hypotarsus (slightly, if any,
depressed in Vultur, moderately to deeply depressed in Gymnogyps);
(10) shaft transversely concave posteriorly at midpoint of shaft
(flat to rounded in Vultur; flat to slightly concave in Gymnogyps);
(11) shaft widens slightly to moderately to meet internal cotyla,
widens abruptly to meet external cotyla (widens abruptly to meet both
external and internal cotylae in Vultur, widens moderately to abruptly
to meet internal cotyla and abruptly to meet external cotyla in
Gymnogyps); (12) shaft widens much more to meet internal trochlea
than it does to meet external trochlea, placing middle trochlea off
center (widens equally to both internal and external trochleae,
placing middle trochlea near midline of shaft in Vultur; similar in
Gymnogyps); (13) internal trochlear surface small (large in Vultur;
moderate to large in Gymnogyps); (14) internal trochlea extending
distally to almost reach distal-most point of middle trochlea (does
not project as far distally in Vultur and Gymnogyps); (15) internal
trochlea with rounded posterior edge (posterior edge arrow-shaped,
directed posteriorly in Vultur; arrow-shaped, directed postero-
proximally in Gymnogyps); (16) middle trochlea short (moderate to
long in Vultur and Gymnogyps); (17) external trochlear surface large,
not delimited sharply antero-proximally (moderate, sharply delimited
antero-proximally in Vultur; moderate to large, moderately delimited
antero-proximally in Gymnogyps); (18) external trochlea with distinct
postero-proximal corner to posterior edge (rounded in Vultur; rounded
to prominent proximal projection in Gymnogyps); (19) external trochlear
notch moderately wide (very wide in Vultur; moderately wide in Gymnoyps).
Vulturidae, n. gen. n. sp.
Holotype. Complete right tarsometatarsus. (PI. IV, Figs. A & B)
Paratypes. Complete right tarsometatarsus and distal end of 1
right tarsometatarsus.
Diagnosis. As for genus. For measurements see Table 5.
Measurements of the holotype are as follows: length, 121.8; proximal
width, 25.4; distal width, 29.0; least width of shaft, 13.8; width of
middle trochlea, 10.9.
Referred material. Portions of 2 sterna, 1 complete left
coracoid, 2 left coracoids lacking head, humeral end of 1 right
coracoid, sternal end of 1 left coracoid, shaft and distal end of
1 left humerus, proximal end of 1 right humerus, distal end of 1
right humerus, proximal end of 1 left ulna, distal end of 1 left ulna,
proximal ends of 1 right and 1 left radius, distal ends of 1 right
and 2 left radii, proximal end of 1 right carpometacarpus, distal
end of 1 left tibiotarsus, proximal ends of 4 right femora, distal
ends of 2 right femora. The 28 specimens, including holotype and
paratypes, represent a minimum of 4 individuals. In addition the
following elements are tentatively referred to the species, but
cannot be assigned with certainty: 2 left articulars, 1 left quadrate,
portion of 1 cranium.
Characters. Sternum with (1) ventral lip of coracoidal sulcus
markedly concave along ventral edge, matching short external anterior
sternal facet of coracoid (not concave in Vultur and Gymnogyps); (2)
carinal apex situated very far distally (situated short distance
distally in Vultur, moderate distance in Gymnogyps); (3) carina
large (of moderate size in Vultur and Gymnogyps); (4) ventral
manubrial spine very broad (broad in Vultur, narrow in Gymnogyps).
Coracoid with (1) coracoidal fenestra lacking (present in Vultur
and Gymnogyps); (2) shaft leading distally from external corner of
glenoid facet forming large ridge (slight to moderate in Vultur, very
slight in Gymnogyps); (3) shaft very concave between procoracoid and
anterior sternal facet (not as concave in Vultur and Gymnogyps); (4)
external portion of anterior sternal facet short, distinctly set off
from base of shaft distally (long, not distinctly set off in Vultur
and Gymnogyps); (5) procoracoid directed internally along plane of
glenoid facet (directed more anteriorly in Vultur and Gymnogyps).
Humerus with (1) head narrow (wide in Vultur, narrow in Gymnogyps):
(2) capital shaft ridge dropping off sharply on both sides (shaft
more rounded in Vultur and Gymnogyps); (3) attachment of M. proscap-
ulohumeralis brevis elevated (not elevated in Vultur, elevated or
not elevated in Gymnogyps); (4) pectoral attachment very prominent
(less prominent in Vultur, more similar in Gymnogyps); (5) deltoid
crest flaring distally (not flaring as much in Vultur and Gymnogyps);
(6) shaft ventral to M. pectoralis superficialis not depressed
(depressed in Vultur, not depressed in Gymnogyps); (7) external
condyle wide and short (narrow and long in Vultur and Gymnogyps); (8)
internal condyle short (very long in Vultur, moderately long in Gym-
nogyps); (9) ectepicondylar prominence long, moderately produced prox-
imally (of moderate length, not produced distally, greatly produced
and angular proximally in Vultur; short, greatly produced and rounded
in Gymnogyps); (10) internal condyle moderately rotated anteriorly,
giving moderately flexed distal end (greatly rotated in Vultur,
moderately rotated in Gymnogyps); (11) impression of M. brachialis
anticus moderately deep (very deep in Vultur and Gymnogyps).
Ulna with (1) olecranon of moderate size (very large in Vultur,
small to moderate in Gymnogyps); (2) attachment of anterior articular
ligament small, not extending far distally (large, extending very
far distally in Vultur; large, extending far distally in Gymnogyps);
(3) area between external and internal condyles flat in external
view (internal condyle projects slightly farther distally in Vultur
and Gymnogyps); (4) carpal tuberosity large, joins shaft abruptly
(large in Vultur, small in Gymnogyps, merges more gently with shaft
in both).
Radius with (1) bicipital attachment large (small in Vultur and
Gymnogyps); (2) bicipital tubercle large (large in Vultur, very
small in Gymnogyps); (3) capital tuberosity bordered internally by
deep notch, pierced by large nutrient foramina (shallow in Vultur,
moderate in Gymnogyps, nutrient foramina small to moderate in both);
(4) external corner of distal end rounded (flattened so angle is
formed at external end of carpal facet in Vultur more rounded in
Gymnogps; ; (5) tendinal groove bordered externally by high ridge
(very slight in Vultur, moderate ridge in Gymnogyps); (6) distal end
with external edge curving sharply palmarly (almost straight in
Vultur and Gmnogs) .
Carpometacarpus with (1) process of metacarpal I low, curving
sharply proximally (high, with moderate proximal curvature in Vultur;
moderately high, with slight proximal curvature in Gymnogyps); (2)
external ligamental attachment a small deep pit bordered posteriorly
by raised area (of moderate size, lying in a pit, with no raised
area posteriorly in Vultur and Gymnogyps).
Femur with (1) trochanter of moderate size, subrounded (large,
well rounded in Vultur; of moderate size, subrounded in Gymnogyps);
(2) attachment of M. obturator externis deep (larger, not as deep,
positioned similarly in Vultur; similar size, not as deep, located
more posteriorly in Gymnogyps); (3) attachments of M. ilio-trochan-
tericus medius and M. iliacus confluent and with surface ribbed
(separate and not ribbed in Vultur and Gymnogyps); (4) concavity
immediately distal to iliac facet posteriorly deep, bordered extern-
ally by large protuberance (shallow, very small protuberance in
Vultur; very deep, moderate sized protuberance in Gymnogyps); (5)
distal end of internal condyle subrounded in medial view (more
rounded in Vultur; less rounded in Gymnogyps); (6) rotular groove
moderately undercut distally (deeply undercut in Vultur, moderately
undercut in Gymnogyps).
Tibiotarsus with (1) internal condyle short antero-posteriorly
(long in Vultur and Gymnogyps); (2) distal internal ligamental
attachment very deep (deep in Vultur, moderately deep in Gymnogyps).
Remarks. The tarsometatarsus is traditionally made the holotype
of fossil species when available, particularly in the case of
falconiform species, and the sum of many tarsometatarsal characters
of Vulturidae, n. gen. n. sp. establishes it as a distinctive genus
and species, although from the above diagnosis it is apparent that it
resembles Gymnogyps more than Vultur. Additional very apparent
large differences found in other elements strengthens this belief,
i.e., the shape of the sternum and the manner of articulation with
the coracoid, the absence of a coracoidal fenestra, and the confluent
ribbed attachments of M. ilio-trochantericus medius and M. iliacus
of the femur.
Genus Gymnogyps Lesson
Gymnogyps n. sp.
Holotype. Complete right tarsometatarsus. (P1. V, Figs. A & B)
Paratypes. One complete left tarsometatarsus, proximal ends of
2 right and 3 left tarsometatarsi, distal ends of 1 right and 3 left
tarsometatarsi.
Diagnosis. Agrees with Gymnogyps and differs from Vultur and
Vulturidae, n. gen. by having those characters of Gymnogyps listed in
description of Vulturidae, n. gen. n. sp.
Differs from Gymnogyps californianus (Shaw) and G. amplus L.
Miller (Collection of Pierce Brodkorb no. PB498, Reddick, Florida) by
having (1) intercotylar prominence with prominent concavity on
external side (slight to deep groove present in G. californianus and
G. amplus); (2) internal cotyla with moderately curved posterior
rim (moderately to sharply curved in G. californianus and G. amplus);
(3) hypotarsus extending internally to at least middle of internal
cotyla (does not extend as far internally in G. californianus and G.
amplus); (4) posterior projection of external cotyla large (small to
large in G. californianus and G. amplus); (5) ridge leading distally
from hypotarsus very prominent, extending distally for more than
third of length of bone, with deep depression of shaft on both sides
(moderately prominent, extending distally for less than third length
of shaft, with depression internally greater than that externally in
G. californianus and G. amplus); (6) anterior metatarsal groove with
marked concavity distal to attachment of M. tibialis anticus (no
noticable concavity to marked concavity in G. californianus and G.
amplus); (7) anterior metatarsal ridges well rounded and approximately
equal in size (external anterior metatarsal ridge larger and more
rounded than internal anterior metatarsal ridge in G. californianus and
G. amplus); (8) internal anterior metatarsal ridge continues distally
and forms a marked corner to shaft that ends at base of internal rim
of middle trochlea (is not traceable as a distinct corner in G. cali-
fornianus and G. amplus); (9) internal trochlea long, large,rotated
posteriorly, with longest part of postero-internal edge directed
postero-proximally (short, small to large, not rotated as much
posteriorly, with sharp to rounded postero-internal edge directed
more posteriorly than proximally in G. californianus and G. amplus);
(10) middle trochlea very long, lying parallel to main axis of shaft,
rims straight, and ending posteriorly in gradual taper (short, distal
end turned slightly inward from main axis of shaft, rims flaring and
terminating more abruptly posteriorly in G. californianus and G. amplus);
(11) external trochlea short, not extending as far distally as internal
trochlea, rotated slightly posteriorly, trochlear surface very long
anteriorly, with posterior edge well rounded and directed posteriorly
(long, extending as far distally as internal trochlea, rotated
slightly, if any, posteriorly, trochlear surface short, with posterior
edge directed posteriorly or proximally in G. californianus and
G. amplus.
For measurements see Table 5. Measurements of the holotype are
as follows: length, 122.0; proximal width, 26.9; distal width, 30.2;
least width of shaft, 13.5; width of middle trochlea, 11.1.
Referred material. Portion of 1 sternum, 1 left scapula, 1
complete right coracoid, 1 right coracoid lacking head, 1 complete
right and 1 complete left humerus, shaft and distal end of 1 right
humerus, proximal end of 1 left humerus, distal ends of 1 right and
2 left humeri, proximal ends of 1 right and 1 left ulna, distal ends
of 1 right and 2 left ulnae, distal end of 1 right radius, distal
ends of 2 left tibiotarsi. The 30 specimens, including holotype and
paratypes, represent a minimum of 4 individuals.
Characters. Sternum with (1) ventral manubrial spine narrow
(wider in G. californianus); (2) ventral lip of coracoidal sulcus
curved along ventral edge (less curvature in G. californianus,
straight in Vultur).
Scapula with (1) internal surface very concave anteriorly (less
in G. californianus, similar in Vultur); (2) distal end of dorsal
acromion edge greatly elevated above shaft (similar, but not quite
as sharply set off in G. californianus; moderately elevated, merging
gradually with shaft in Vultur).
Coracoid with (1) anterior sternal facet with small dorsal
slope (moderate in G. californianus, great in Vultur); (2) scapular
facet moderately elevated above shaft (slightly elevated in G.
californianus, greatly elevated in Vultur); (3) head narrow (similar
in G. californianus, wide in Vultur).
Humerus with (1) head narrow (slightly wider in G. californianus);
(2) attachment of M. proscapulohumeralis brevis elevated (not elevated
.in G. californianus); (3) capital groove divides base of median crest
and capital shaft ridge (does not extend as far distally in G.
californianus); (4) internal tuberosity moderately prominent
(very prominent in G. californianus); (5) deltoid crest small, with
anconal side very concave (larger, with anconal side less concave in
G. californianus); (6) shaft moderately robust (very robust in G.
californianus); (7) internal condyle small (large in G. californianus);
(8) external condyle moderately produced distally (very produced in
G. californianus); (9) cctepicondylar prominence short, distinctly
set off from shaft distally (long, not distinctly set off from shaft
in G. californianus); (10) entepicondyle moderately produced (more
produced in G. californianus); (11) impression of M. brachialis
anticus deep (very deep in G. californianus).
Ulna with (1) olecranon of moderate size (short, narrow, but
deep in G. californianus); (2) attachment of anterior articular
ligament long, elevated, and terminating abruptly (long, not elevated,
terminating gradually in G. californianus); (3) internal condyle
projecting slightly distal to external condyle (projects moderately
distal to external condyle in G. californianus); (4) carpal tuberosity
long (shorter in G. californianus).
Radius with (1) ulnar depression moderately deep (shallow in
G. californianus); (2) shaft widening very abruptly internally for
carpal facet (widens more gradually in G. californianus).
Tibiotarsus with (1) distal internal ligamental attachment
moderately deep (moderately deep in G. californianus, deep in Vultur);
(2) internal condyle not flaring internally at antero-proximal end
(similar in G. californianus, flares internally in Vultur).
Remarks. The appearance in the fossil record of a South American
Table 5. Measurements of the scapula, coracoid, humerus, radius,
ulna, carpometacarpus, femur, tibiotarsus, and tarsometatarsus of
Gymnogyps californianus, Recent Vultur gryphus, V. gryphus from the
Talara Tar Seeps (excluding scapula, humerus, and carpometacarpus),
Gymnogyps n. sp. (excluding carpometacarpus and femur), and
Vulturidac, n. gen. n. sp. (excluding scapula) (in mm).
Gymnogyps
californianus
27.2
1
8.1
1
102.6
1
46.9
1
Scapula
Proximal
Height
Proximal
Width
Coracoid
Length
Head to
Scapular
Facet
Proximal
Depth
Least
Depth of
Shaft
Length of
Sternal
Facet
Humerus
Length
Proximal
Width
Gymnogyps
n. sp.
16.4
1
38.6
1
273.0
1
55.7
1
96.2
1
43.2
1
25.5
1
14.8-18.1
16.4
2
256.0-265.0
260.5
2
52.5-54.3
53.4
2
Table 5. Extended.
Vultur gryphus
Recent Talara Tar Seeps
28.6-30.5
29.5
2
7.4-9.6
8.5
2
98.2-98.8 103.1
98.5
2 1
50.5-51.3 52.0
50.9
2 1
29.8-30.3 30.0
30.1
2 1
16.6-18.0 16.9
17.3
2 1
43.3-44.1 42.5
43.7
2 1
265.0-292.0
278.5
2
62.6-63.4
63.0
2
Vulturidae
n. gen. n. sp.
89.4
1
44.0-51.9
47.9
2
25.7-28.2
26.9
2
13.9-14.9
14.4
3
55.7
1
Table 5. Continued.
Distal
Width
Least
Width of
Shaft
Radius
Length
Proximal
Width
Distal
Width
Ulna
Length
Proximal
Width
Distal
Depth
Least
Width of
Shaft
Carpometacarpus
Length
Gymnogyps
n. sp.
46.4-49.0
47.6
5
18.2-19.6
19.1
3
29.3-31.9
30.6
2
23.6-25.3
24.5
2
Gymnogyps
californianus
51.1
1
23.0
1
304.0
1
14.1
1
22.8
1
319.0
1
33.5
1
24.7
1
13.5
1
139.0
1
Table 5. Extended.
Vultur gryphus
Recent Talara Tar Seeps
55.1-55.6
55.4
2
21.2-21.8
21.5
2
298.0-328.0
313.0
2
15.6-15.7
15.7
2
24.3-24.4
24.4
2
313.0-341.0
327.0
2
33.7-36.3
35.0
2
26.7-28.8
27.7
2
14.3-15.4
14.9
2
132.2-140.3
136.2
2
14.6
1
24.1
1
Vulturidae
n. gen. n. sp
48.5
1
13.2-13.4
13.3
2
22.7-24.3
23.4
3
33.5-34.0
33.8
2
Table 5. Continued.
Gymnogyps
n. sp.
Proximal
Width
Height
Through
Metacarpal I
Least
Width of
Shaft
Length of
Distal
Fornix
Femur
Length
Proximal
Width
Distal
Width
Least
Width of
Shaft
Tibiotarsus
Length
Proximal
Width
Distal
Width
Gymnogyps
californianus
13.2
1
14.4
1
137.7
1
32.3
1
214.0
1
28.7
1
24.2-25.3
24.8
2
Table 5. Extended.
Vultur gryphus
Recent Talara Tar Seeps
17.2-17.4
17.3
2
36.3-38.9
37.6
2
13.0-14.6
13.8
2
18.1-20.5
19.3
2
143.6-158.3
150.9
2
37.6-38.5 37.3
38.0
2 1
37.9-41.1 37.9
39.5
2 1
18.0-19.5
18.7
2
216.0-242.0 213.0
229.0
2 1
28.6-30.6 30.0-31.1
29.6 30.6
2 2
25.1-27.1 23.9-26.1
26.1 25.0
2 4
Vulturidae
n. gen. n. sp.
15.4
1
32.6
1
31.7-32.8
32.2
3
34.1-34.7
34.4.
2
24.1
I
87
Table 5. Continued.
Gymnogyps
n. sp.
Least
Width of
Shaft
Depth of
Internal
Condyle
Tarsometatarsus
Length
Proximal
Width
Distal
Width
Least
Width of
Shaft
Width of
Middle
Trochlea
Gymnogyps
californianus
14.7
1
22.8
1
118.5-122.0
120.3
2
25.8-29.9
27.0
7
29.5-34.0
31.3
4
13.4-13.5
13.5
2
10.6-12.1
11.3
6
117.8
1
26.7
1
29.8
1
14.0
1
Table 5. Extended.
Vultur gryphus
Recent Talara Tar Seeps
13.9-15.3 14.1
14.6
2 1
24.9-26.2 22.9-24.6
25.6 23.9
2 4
118.8-134.7
126.7
2
27.5-29.8 30.0
28.7
2 1
30.7-33.6
32.1
2
13.7-14.7
14.2
2
12.1-12.9
12.5
2
Vulturidae
n. gen. n. sp.
22.5
1
121.8-124.5
123.1
2
25.4-26.3
25.8
2
29.0-30.0
29.5
2
13.8
13.8
2
10.6-11.6
11.0
3
species of Gymnogyps poses the question of its relationships with the
G. amplus-G. californianus line of North America. The continent of
origin of the genus must remain uncertain until specimens of an
earlier period are found.
Genus Vultur Linnaeus
Vultur gryphus Linnaeus
Andean Condor
Material. One complete right coracoid, distal end of 1 right
humerus, proximal end of 1 right and 1 left ulna, distal end of
1 right ulna, proximal end of 1 left radius, distal end of 1 left
radius, proximal end of 1 left femur, distal ends of 3 left femora,
1 complete right tibiotarsus, proximal end of 1 right tibiotarsus,
distal ends of 2 right and 1 left tibiotarsus, proximal end of 1
left tarsometatarsus. The 17 specimens represent a minimum of 3
individuals.
Characters. See diagnosis for Vulturidae, n. gen. n. sp. and
Gymnogyps n. sp.
Remarks. V. gryphus is found almost throughout the Andes of
South America and often on the Pacific coast.
Discussion
In addition to the two species of living condors, seven large
extinct vulturid species have been named from the Pliocene and
Pleistocene, one each in the two living genera and the other five in
extinct genera. The possibility of one or more of these extinct
North American species being identical to an extinct form from the
Talara Tar Seeps requires a discussion of their characters.
In the subfamily Vulturinae [Illiger) two extinct monotypic
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