S M I T H S O N I A N C O N T R I B U T I O N S T O P A L E O B I O L O G Y ? N U M B E R 6 2 
Fossil Birds from the 
Oligocene Jebel Qatrani Formation, 
Fayum Province, Egypt 
D. Tab Rasmussen, 
Storrs L. Olson, 
and 
Elwyn L. Simons 
SMITHSONIAN INSTITUTION PRESS 
Washington, D.C. 
1987 
A B S T R A C T 
Rasmussen, D. Tab, Storrs L. Olson, and Elwyn L. Simons. Fossil Birds from the 
Oligocene Jebel Qatrani Formation, Fayum Province, Egypt. Smithsonian Contributions 
to Paleobiology, number 62, 20 pages, 15 figures, 1986.?^Fossils from fluvial deposits 
of early Oligocene age in Egypt document the earliest known diverse avifauna from Africa, 
comprising at least 13 families and 18 species. Included are the oldest fossil records of 
the Musophagidae (turacos), Pandionidae (ospreys), Jacanidae (jacanas), and Balaenicipi-
tidae (shoebilled storks). Other families represented are the Accipitridae (hawks and 
eagles), Rallidae (rails), Gruidae (cranes), Phoenicopteridae (flamingos), Ardeidae 
(herons), Ciconiidae (storks), and Phalacrocoracidae (cormorants). A highly distinctive 
rostrum is described as a new family, Xenerodiopidae, probably most closely related to 
herons. A humerus lacking the distal end is tentatively referred to the same family. Two 
new genera and three species of large to very large jacanas are described from the distal 
ends of tarsometatarsi. This Oligocene avifauna resembles that of modern tropical African 
assemblages. The habitat preferences ofthe constituent species of birds indicate a tropical, 
swampy, vegetation-choked, fresh-water environment at the time of deposition. 
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's 
annual report, Smittisonian Year. SERIES COVER DESIGN: The trilobite Phacops rana Green. 
Library of Congress Cataloging in Publication Data 
Rasmussen, D. Tab 
Fossil birds from the Oligocene Jebel Qatrani Formation, Fayum Province, Egypt 
(Smithsonian contributions to paleobiology ; no. 62) 
Bibliography : p. 
SupL of Docs, no.: SI 1.30:62 
1. Birds, Fossil. 2. Paleontology?Oligocene. Paleontology?^Egypt?Fayum (Province). 
I. Olson, Storrs L. II. Simons, Elwyn L. III. Title. IV. Series 
QE701.S56 no. 62 560 s 87-9782 [QE8711 [568'.0962'2 
Contents 
Page 
Introduction 1 
Acknowledgments 2 
Systematic Paleontology 3 
Order and Family Indeterminate 3 
Eremopezus eocaenus Andrews, 1904 3 
Stromeria fajumensis Lambrecht, 1929 3 
Order CUCULIFORMES 3 
Family MUSOPHAGIDAE 3 
Genus and Species Indeterminate, aff. Crinifer 3 
Order "FALCONIFORMES" 5 
Family AcciprrRiDAE 5 
Genus and Species Indeterminate, aff. Haliaeetus 5 
Family PANDIONIDAE 5 
Genus and Species Indeterminate, aff. Pandion 5 
?Pandionidae, Genus and Species Indeterminate 6 
Order GRUIFORMES 6 
Family RALLIDAE 6 
Genus and Species Indeterminate 6 
Family GRUIDAE 6 
Genus and Species Indeterminate 6 
Order CHARADRHFORMES 7 
Family JACANIDAE 7 
Nupharanassa, new genus 7 
Nupharanassa bulotorum, new species 7 
Nupharanassa tolutaria, new species 8 
Janipes, new genus 8 
Janipes nymphaeobates, new species 8 
Family PHOENICOPTERIDAE 9 
Genus Indeterminate, aff. Palaelodus, species 1 9 
Genus Indeterminate, species 2 10 
Order "QcoNnFORMEs" 11 
Family XENERODIOPIDAE, new family 12 
Xenerodiops, new genus 12 
Xenerodiops mycter, new species 12 
Family ARDEIDAE 13 
Nycticorax sp 13 
Genus and Species Indeterminate 14 
Family CICONIIDAE 15 
Palaeoephippiorhynchus dietrichi Lambrecht, 1930 15 
IPalaeoephippiorhynchus dietrichi Lambrecht, 1930 15 
Family BALAENicipmDAE 15 
Goliathia andrewsi Lambrecht, 1930 15 
Order I*ELECANIFORMES 16 
111 
IV SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 
Family PHALACROCORACIDAE 16 
Genus and Species Indeterminate 16 
Discussion 17 
Literature Cited 19 
Fossil Birds from the Oligocene 
Jebel Qatrani Formation, 
Fayum Province, Egypt 
D. Tab Rasmussen, Storrs L. Olson, 
and Elwyn L. Simons 
Introduction 
Fossils from the Ohgocene Jebel Qatrani Formation of 
Fayum Province, Egypt, provide one of the earhest and most 
complete records of the continental Tertiary flora and fauna 
of Africa. The Fayum is best known for its early anthropoid 
primates, but in addition has yielded 13 other orders of 
mammals, including the only marsupial known from Africa, 
turtles, crocodiles, snakes, lizards, sharks and skates, lungfish, 
bony fishes, and a variety of plants and trace fossils (Bown et 
al., 1982; Simons and Bown, 1984; Gebo and Rasmussen, 
1985). 
Despite this diversity of fossil forms, very litde has been 
known of the avifauna. Previously, only four species of birds 
had been described from the Fayum, all of which were named 
over 50 years ago (Andrews, 1904; Lambrecht, 1929, 1930). 
Rich (1974) mentioned, but did not describe, a fifth avian 
specimen, evidendy a humerus belonging to the Pelecani-
formes. In 1962, a large tarsometatarsus was recovered from 
the lower part of the Jebel Qatrani Formation. Moustafa (1974) 
suggested that this might belong to Eremopezus but it was 
never described and has since been lost Apart from these few 
specimens from the Fayum, the only other Paleogene fossil 
bird hitherto reported from Africa is Gigantornis eaglesomei 
Andrews, based on a sternum from the Eocene of Ni?
geria belonging to the gigantic pseudo-toothed pelecaniform 
birds of the marine family Pelagornithidae (Olson, 1985). 
Fossils from the Fayum are usually poorly mineraUzed and 
D. Tab Rasmussen, Duke University Primate Center, Duke University, Durham, 
North Carolina 27706. Storrs L. Olson, Department of Vertebrate Zoology, National 
Museum of Natural History, Smithsonian Institution, Washington, DC. 20560. 
Elwyn L Simons, Duke University Primate Center and Departments of Anatomy 
and Anthropology, Duke University, Durham, North Carolina 27706. 
quickly disintegrate when exposed to surface weathering. In 
recent years, however, methods have been developed for the 
efficient recovery of small and delicate fossils from the Jebel 
Qatrani sandstones (Simons, 1967). This has led to the recovery 
of a number of specimens of birds, mainly of smaller species 
than those known previously, that would never have survived 
normal erosional exposure on the surface. Detailed study of the 
intrafamiUal taxonomic relationships of most of these fossils 
has not been attempted, as each of the species is represented 
by only a few fragmentary specimens. Instead, emphasis is 
placed on the zoogeographical and paleoecological signif?
icance of the fossU avifauna. 
The 340 m thick Jebel Qatrani Formation consists mainly 
of variegated sandstones and mudstones deposited as point 
bars and overbank deposits by several meandering streams or 
rivers (Bown and Kraus, 1987). These sediments are exposed 
in a series of escarpments forming the northern rim of the 
Fayum depression located southwest of Cairo (Figure 1). The 
Jebel Qatrani Formation conformably overlies the predomi?
nantly marine sediments of the late Eocene Qasr el Sagha 
Formation, and is capped by the Widan el Faras Basalt which 
has been dated at 31?1 my (Fleagle et al., 1986; see Figure 2), 
thus providing a minimum age for the very top of the formation 
(latest early or earliest middle Oligocene). The highest fossil 
bird and primate localities in the Jebel Qatrani Formation are 
100 m below this basalt layer and are separated from it by a 
major erosional unconformity, making it almost certain that 
none of the fossils are younger than early Oligocene. 
Two main fossil-producing levels occur in the Jebel Qatrani 
sediments, one near the top of the formation in what is known 
as the upper sequence, the other near the middle of the lower 
sequence (Bown and Kraus, 1987; these are the "Upper and 
Lower Fossil Wood Zones" of previous authors). The upper 
and lower sequences are separated by a cliff-forming marker 
SMiniSONLAN CONTRIBUTIONS TO PALEOBIOLOGY 
? E ST E RN 
D E S E R T 
.,^ '^ *' %*^^ 
/ 
.^^J-
? V ^ 
i ? * * . v> ' 
r^-^'^1-'-
^"?^t" 
"^ 
o?'""'l,,/' 
??.??'? * 
.'-; 
*' 
f - * ' ? * 
. a 
Widan el Faras Basalt (31.0 \ 1.0 m.y.) 
FIGURE 1.?Map of Egypt showing the position of the field camp (asterisk) and the 
Jebel Qatrani Escarpment in relation lo the Fayum Depression and Cairo. The Jebel 
Qatrani Escarpment is composed of the upper 100 m of the Jebel Qatrani Formation 
and a resistant overlying basalt. North of the escarpment lies the Miocene Kashab 
Formation, immediately to the south are exposures of the Oligocene Jebel Qatrani 
Formation, and farther south is the marine Eocene Qasr el Sagha Formation. Most 
fossil-producing quarries occur within 5 km of the camp location. Modified from 
Bown etal. (1982). 
bed called the Barite Sandstone (Figure 2). 
The four previously described bird specimens from the 
Fayum were collected during the first decade of this century 
from what were then called the "fluvio-marine beds" (the 
fluvial Jebel Qatrani). The early expeditions collected nearly 
exclusively in the lower sequence, from which all four bird 
specimens came. When the first of these, Eremopezus eocaenus 
Andrews (1904), was described, the "fluvio-marine beds" were 
considered to be of late Eocene age, hence the specific name. 
Andrews' (1906) list of the vertebrate fauna associated with 
Eremopezus clearly places it in the lower Oligocene, however. 
Since 1960, Yale and Duke University expeditions led by 
Simons, in cooperation with the Egyptian Geological Survey 
and General Petroleum Company, have collected extensively 
throughout the suatigraphic range of the Jebel Qatrani 
Formation. The great majority of the bird bones, as well as ihc 
small mammals, come from upper sequence quarries (Figure 
2). 
Fossils were collected at each of the quarries by surface 
collecting, excavation, or a special wind-quarrying technique, 
described elsewhere (Simons, 1967), which is responsible for 
the recovery of all new bird specimens described here. The 
fossils were compared with recent and fossil skeletal material 
in the National Museum of Natural History, Smithsonian 
P 250-
Oasr el Sagha Formation (Eocene) 
FIGURE 2.?Simplifiexi section of the Jcbcl Qatrani Formation showing the 
straiigraphic positions of quarries that have produced birds. TTie sediments lying 
above the Barite Sandstone constitute the upper sequence, those below constitute 
the lower sequence. TTie vast majority of bird specimens have come from Quarries 
I and M of the upper sequence. Quarry V has not yet produced any diagnostic bird 
elements. Quarries E and A have each produced a single diagnostic bone (of a jacanid 
and accipitrid, respectively). Modified from Bown and Kraus (1987). 
Institution (USNM), and the Zoology DeparUncnt, Duke 
University. Specimens are cataloged in the collections of the 
Cairo Geological Museum (CGM) and the Duke University 
Primate Center (DPC). The sequence of taxa generally follows 
that in Olson (1985). 
AcKNowi.EDGMENTs.?We thank Richard Kay, Daniel Gcbo, 
and Asenath Bernhardt for comments and discussion; Priihijit 
Chatrath and Frederick V. Grady for preparing specimens; 
Louise Roth and Joseph Bailey for assistance with the 
ostcological collection of the Duke University Zoology 
Department; and the Egyptian Geological Survey and General 
Petroleum Company of Egypt for assistance in field operations. 
Specimens described here were collected by P.S. Chatrath, 
J.F. Fleagle, D.L. Gcbo, A.G. Ibrahim, R.F. Kay, A.C. 
McKenna, R.M. Madden, and Rasmussen and Simons. 
Photographs in Figures 9 and 11 arc by Victor E. Krant/, 
Smithsonian Institution. We acknowledge National Science 
Foundation grants BNS-80-16206 and BNS-81-14295 and 
Smith.sonian Foreign Currency grant 809479 to Simons. Wc 
are grateful to Jonathan Becker, Scott L. Wing, and Ronald 
NUMBER 62 
G. Wolff for comments on the manuscript, to Richard L. Zusi 
for remarks on the cranial adaptations of Xenerodiops, and 
George Steyskal for assistance with classical languages. 
Systematic Paleontology 
Order and Family Indeterminate 
Eremopezus eocaenus Andrews, 1904 
HOLOTYPE.?Distal end of left tibiotarsus, British Museum 
(Natural History) A843. 
TYPE LOCALITY AND HORIZON.?North of Birket Qarun, 
lower sequence, Jebel Qatrani Formation. 
REMARKS.?Andrews (1904, 1906) compared this rhea-sized 
tibiotarsus with a variety of ratites and concluded diat it 
combined features of several of them, while at the same time 
it differed widely from all other birds, ratite or carinate. He 
declined to assign the specimen to any higher level taxon 
except "Order Ratitae" (Andrews, 1906:258). With consider?
able doubt, he also referred a pedal phalanx recovered from the 
same pit to this species. This was "remarkable for die breadth 
and depUi of its proximal end compared with the distal" 
(Andrews, 1906:260). 
Andrews always expressed great uncertainty as to the 
relationships oi Eremopezus. Rothschild (1911) placed Eremo?
pezus in its own subfamily, Eremopezinae, of the Aepyorni-
thidae, the elephantbirds of Madagascar. This was presumably 
because of his behef that it was a "general type intermediate 
between Aepyornis and Struthio" (Rodischild, 1911:149). The 
name Eremopezinae is thus attributable to Rothschild (1911), 
rather than to Lambrecht (1933) as usually cited (e.g., 
Brodkorb, 1963). Lambrecht (1933) also included Eremopezus 
in its own subfamily in the Aepyornithidae on the basis of 
deep ligamental pits on the lateral and medial sides of die 
condyles. See additional remarks on this species below. 
Stromeria fajumensis Lambrecht, 1929 
HOLOTYPE.?A broken and much worn distal end of a right 
tarsometatarsus lacking all but the bases of the trochleae; 
specimen in Munich. 
TYPE LOCALITY AND HORIZON.?Nordi of Dimeh, Fayum, 
Jebel Qatrani Formation. 
REMARKS.?^The spelling fajumensis was used consistently 
in the original publication (Lambrecht, 1929), the emendation 
to fayumensis (Lambrecht, 1933) being unjustified. Lambrecht 
assigned this specimen to the Aepyornithidae on the basis of 
a projecting crest on the plantar surface of die shaft, a feature 
he considered to be shared with the similarly-sized Mullerornis 
betsilei Milne-Edwards and Grandidier. He found the similarity 
compelling enough to suggest that Stromeria represents a direct 
ancestor of Malagasy aepyorniUiids. We feel that such 
conclusions are not justified by the extremely poor nature of 
the specimen. Judging from Lambrecht's (1929) photographs, 
the plantar crest actually differs considerably in shape and 
proportions from diat of Mullerornis. 
At no point did Lambrecht (1929, 1933) ever state why 
Eremopezus eocaenus and Stromeria fajumensis, known only 
from noncomparable elements, should belong to different 
species, genera, and subfamiUes. Perhaps this was because the 
specimen of Eremopezus was judged to be markedly different 
from typical aepyornithids, whereas that of Stromeria was not. 
The distal widdi of the tibiotarsus oi Eremopezus was given 
as 48 mm (Andrews, 1904) and that of die tarsometatarsus of 
Stromeria as 55 mm (Lambrecht, 1929). In modern cursorial 
ratites such as Rhea, the distal ends of the tarsometatarsus and 
tibiotarsus are about equal in width, but in other ratites the 
distal end of the tarsometatarsus may be more expanded. Thus 
in the Aepyornithidae the distal width of the tibiotarsus is 
93-95% of the distal widdi of the tibiotarsus (data from 
Andrews, 1904; Milne-Edwards and Grandidier, 1869), and in 
moas 70-80% (data from Oliver, 1949). Therefore, with the 
widdi ofthe tibiotarsus oi Eremopezus being 87% of die width 
of the tarsometatarsus of Stromeria, it is entirely conceivable 
that both may be referable to a single genus or species, 
particularly given the likelihood of individual and sexual 
differences in size. 
In summary, there is no compelling evidence to suggest that 
more than one species of ratite-like bird is represented in the 
Fayum deposits, a point previously noted by Moustafa (1974), 
and the material is insufficient to determine with confidence 
anything about the relationships of the bird. As Andrews 
(1904:170) himself stated, widi his characteristic insight: 
''Eremopezus may be merely another instance of retrogressive 
modification leading to loss of flight and increase of bulk in a 
group of Carinate birds, such as has occurred in the case of die 
Stereornithes, die Gastornidies, and probably in most of die 
early so-called Ratite birds." 
Ratites are usually associated with open grassland and 
savanna, but since diis is not true of cassowaries or moas, the 
supposed ratite of the Fayum tells us nodiing about die 
environment in which it Uved. 
Order CUCULIFORMES 
Family MusoPHAcroAE 
Genus and Species Indeterminate, aff. Crinifer 
FIGURES 3, 4 
MATERL?LL EXAMINED.?CGM 42836, distal end of left 
tarsometatarsus (Figure 3); CGM 42837, distal end of right 
humerus (Figure 4). 
LOCALITY AND HORIZON.?Bodi specimens are from Quarry 
M, upper sequence. 
MEASLIREMENTS.?^Tarsometatarsus CGM42836: distal width. 
SMiraSONLAN C O N T R I B U T I O N S TO PALEOBIOLOGY 
METRIC 1 
nil nil nil 
2 
III! 
FIGURE 3.?Anterior view of distal end of left tarsometatarsus (CGM 42836) of a 
turaco (Musophagidae) resembling Crinifer. 
9.4 mm; width and depdi of shaft, 4.8x3.2 mm. Humerus CGM 
42837: distal width, 14.0 mm; depth of external condyle, 6.9 
mm; widdi and depdi of shaft at midpoint, 5.8x5.0 mm. 
COMPARISONS.?The tarsometatarsus lacks part of the inner 
trochlea, and the middle trochlea is somewhat abraded but 
preserves a deep trochlear groove (Figure 3). The specimen 
lacks the unusual configuration of the outer trochlea seen in 
Musophaga and to a lesser extent in Tauraco and Corythaeola. 
The distal foramen is small and discrete, as in Crinifer and 
unlike the double opening, with the distal foramen partially 
separated from the intertrochanteric foramen, observed in 
Musophaga. The condition in Corythaixoides shows an 
approach to diat of Musophaga. The fossil is indistinguishable 
from specimens of the extant genus Crinifer except that it is 
larger than in any modern musophagids except Corythaeola. 
The humerus is slightly abraded at the distal end but is intact 
as far as die pectoral crest (Figure 4). The entepicondylar 
process is large, extending distally well beyond the condyles, 
the brachial depression is small, and the shaft is distinctiy 
curved. The modern genera of musophagids are not readily 
distinguished from one another on the basis of characters of 
the humerus. The fossil is somewhat larger dian in any of die 
extant forms except Corythaeola and hence is of a size 
FIGURE 4.?Palmar view of distal end of right humerus (CGM 42837) of a turaco 
(Musophagidae) resembling Crinifer. 
compatible with being from the same species as the 
tarsometatarsus. 
REMARKS.?^Previously, fossils of Musophagidae had been 
reported from the late Oligocene of Bavaria and die early 
Miocene of France and Kenya (Olson, 1985). Thus the Fayum 
fossils provide the earliest record for the family. That they 
cannot be distinguished generically from extant forms indicates 
that the humerus and tarsometatarsus of at least some 
musophagids have probably not evolved appreciably since die 
Oligocene. The greater resemblance of the fossils to Crinifer 
may only reflect the fact that diis genus appears to be the most 
primitive of modern musophagids in tarsal morphology. 
The modern species of Crinifer tend to prefer open parkland 
and woodland as opposed to the mesic, densely canopied forest 
frequented by most musophagids. Musophagids are among the 
NUMBER 62 
most primitive living birds (Olson, 1985), and as Stegmann 
(1978) has noted, the young are slow in developing and use 
die wings, which have claws on die alular and major digits, to 
clamber about in trees. All modern musophagids are strictly 
arboreal and frugivorous, and die Fayum birds may thus have 
relied to some extent on die same food sources as the 
contemporaneous primates. 
Order "FALCONIFORMES" 
Family ACCIPITRIDAE 
Genus and Species Indeterminate, aff. Haliaeetus 
MATERIAL EXAMINED.?DPC 1652, distal end of left 
tarsometatarsus. 
LOCALTTY AND HORIZON.?Quarry A, lower sequence. 
MEASUREMENTS.?Distal widdi, 23.9 mm; distance between 
distal foramen and external intertrochlear space, 4.6 mm. 
COMPARISONS.?This specimen comes from a large accipi?
trid similar in size and morphology to modern eagles of the 
genus Haliaeetus. The distal foramen is large and situated far 
distally, and the trochleae are widely spaced, in both of which 
features the specimen differs markedly from die Old World 
vultures (Gypaetinae or Aegypiinae auct.). The internal 
intertrochlear space is wider than in any of the modern genera 
of Accipitridae examined. There is a distinct muscle scar on 
the ridge between the facet for metatarsal I and die inner 
trochlea. This is variably present in some extant genera of 
Accipitridae, being especially noticeable in Gypohierax. The 
strong buttressing along the lateral edges of the shaft that is 
characteristic of Aquila and buteonine hawks, is absent in the 
fossil. The overall similarity of the fossd is greatest to modern 
kites and eagles of the genera Milvus and Haliaeetus, which 
are closely related in any case (Olson, 1982), and to the more 
distandy related African pahn-nut vulture, Gypohierax. 
REMARKS.?^The intrafamilial systematics of modern Accipi?
tridae is still very poorly understood and it is hazardous to 
attempt to determine die closest relationships of isolated 
fossils of hawks and eagles. The overall morphology of die 
Fayum fossil suggests that it could be from a fishing eagle 
allied with Haliaeetus, die modern forms of which are hunting 
and scavenging birds that frequent bodies of fresh water and 
seacoasts. During deposition of the lower sequence, where this 
specimen was found, the Fayum was situated much closer to 
the seacoast dian during deposition of die upper sequence 
(Bown and Kraus, 1987). Similarities to Gypohierax may 
reflect shared primitive characters. The monotypic living genus 
Gypohierax is specialized in its food habits, subsisting mainly 
on nuts of die oil palm. It may be die sole remnant of a group 
that was perhaps more diverse and widespread in die past. For 
example, the early Miocene accipitrid Palaeohierax from 
France has been noted by several audiors as being closely 
related to Gypohierax (see Olson, 1985). 
( ^ ^ ^ 
METRIC 1 
1 II 
2 
nil 
FIGURE 5.?Palmar view of distal end of right humerus (DPC 3082) of an osprey 
(Pandionidae) resembling Pandion. 
The Accipitridae are fairly common in the fossil record, 
which extends at least as far back as the late Eocene or early 
Oligocene. The present specimen constitutes die first Paleo?
gene record of die family for Africa. 
Family FANDIONTOAE 
Genus and Species Indeterminate, aff. Pandion 
FIGURE 5 
MATERIAL EXAMINED.?DPC 3082, distal end of right 
humerus (Figure 5). 
LOCALITY AND HORIZON.?Quarry M, upper sequence. 
MEASLIREMENTS.?Distal widdi, 16.9 mm; depth of external 
condyle, 10.2 mm. 
COMPARISONS.?The skeleton in die Pandionidae differs 
markedly from diat in the Accipitridae, die humerus being of 
a rather generalized waterbird type that is more similar in some 
respects to diat in certain Pelecaniformes than to the 
SMITHSONEAN CONTRIBUTIONS TO PALEOBIOLOGY 
Accipitridae. The Fayum fossU is most similar to the humerus 
in Pandion and die Phaediontidae, differing from die latter in 
the shallower, less pronounced olecranal fossa and in details 
of shape and position of muscle scars (Figure 5). The fossil is 
essentially identical to die extant species of Pandion except 
for its smaller size, relatively narrower breadth across die 
brachial depression, and die presence of a small, deep pit 
between die internal condyle and die attachments of the 
anterior articular ligament. 
REMARKS.?^Fossil ospreys, all referable to the extant genus 
Pandion, have been identified from the middle Miocene of 
California, and the late Miocene and early Pliocene of Rorida 
and North Carolina (Warter, 1976; Becker, 1985; Olson, 1985). 
The Fayum record is thus the earliest for the family and die 
first fossU record for Africa. The living osprey is practically 
cosmopoUtan in distribution, although in Africa it occurs 
ahnost entirely as a migrant, with only scattered nesting records 
(Chorley, 1939; Snow, 1978). The osprey subsists almost 
entirely on fish captured by diving, usually from a considerable 
height, and because of its diet is necessarily restricted to aquatic 
and marine environments. If the habits of the fossd form were 
similar, it would mean diat a certain amount of open, relatively 
shallow water was present near the site of deposition. 
?PANDIONIDAE, Genus and Species Indeterminate 
MATERIAL EXAMINED.?DPC 1929, damaged right carpome-
tacarpus. 
LOCALITY AND HORIZON.?Quarry I, upper sequence. 
MEASUREMENTS.?^Lengdi, 85.0 mm; widdi of trochlea, 9.7 
mm; breaddi of distal articular surface, approximately 10.3 
mm. 
COMPARISONS.?The very short distal symphysis and the 
broad, ciu-ved, distal base of the minor metacarpal indicate that 
this specimen is from a raptorial bird. The proximal base of the 
minor metacarpal is not as offset ventrally from the carpal 
trochlea as in the Accipitridae, and the facet for digit II is a 
strong crest rather dian a weak protuberance as in the 
Accipitridae. In both respects the fossil is more similar to the 
Pandionidae. The specimen is in die size range of the modern 
osprey, Pandion haliaetus, but is slightiy more robust and may 
have come from a species larger than the preceding one. 
Order GRUIFORMES 
Family RALLIDAE 
Genus and Species Indeterminate 
MATERIAL EXAMINED.?DPC 3858, distal end of left 
tarsometatarsus. 
LOCALITY AND HORIZON.?Quarry M, upper sequence. 
MEASUREMENTS.?Distal width, 4.5 mm. 
COMPARISONS.?The posterior portions of die inner and 
outer trochleae have been lost through abrasion. These two 
trochleae are shorter than the middle one and are deflected 
posteriorly, but considerably less so than in the Charadrii-
formes or in Turnix. The middle trochlea has a distinct groove 
whereas the odier two trochleae are smoodi and rounded; the 
distal foramen is large, with a deep tendinal groove. In these 
features, and in size, this specimen resembles small rails of the 
genera Sarothrura, Coturnicops, and Laterallus, but the fossil 
is otherwise much too incomplete to be able to ascertain its 
relationships within the large and relatively homogeneous 
family Ralhdae. 
REMARKS.?Although the Rallidae may well have existed 
throughout most of the Tertiary, dieir record prior to the 
Oligocene is based on very imsatisfactory material that has not 
been certainly identified. The Fayum specimen is the earliest 
raU thus far reported from Africa. Although some modern 
species of rails occiu- in moist forests or in grasslands, most 
species inhabit well-vegetated marshes. 
Family GRUTOAE 
Genus and Species Indeterminate 
MATERIAL EXAMINED.?DPC 5330, distal end of left 
tarsometatarsus and portions of shaft. 
LocALFTY AND HoRizoN.?Quarry M, upper sequence. 
MEASimEMENTS.? D^istal widdi, 14.9 mm; depdi of middle 
trochlea, 6.4 mm; approximate shaft depth, 5.6 to 5.9 mm, and 
shaft width, 5.3 to 5.6 mm. 
COMPARISONS.?The trochleae are well preserved except for 
the proximo-ventral portion of the middle trochlea. The bone 
is broken and incomplete immediately proximal to the 
trochleae, but a distal section of shaft 80 mm in length is intact. 
The second and fourth trochleae are shorter than the middle 
one and are deflected posteriorly. The specimen resembles 
gruids (including Aramus) and differs from the somewhat 
similar Phoenicopteridae in die distinct protruding wing at the 
ventro-distal extremity of the second trochlea, the less deeply 
furrowed middle trochlea, and the shallower, hollowed ventral 
area immediately proximal to the trochleae. 
The specimen resembles Balearica and Anthropoides and 
differs from Aramus in the more elevated second trochlea, and 
in the lateral profile of the middle trochlea, which is slightiy 
compressed dorsoventrally rather than rounded. The tarsometa?
tarsus differs from all modern gruids in two details. The wing 
on the second trochlea is weaker and more distally produced, 
radier dian being distinctly hooked proximally; and the shaft 
is as deep or deeper dian it is wide. In size, the Fayum fossil 
is intermediate between the smaller Aramus and the larger 
Anthropoides virgo, although closer to Aramus. 
REMARKS.?The Fayum specimen is the earliest crane 
recorded from Africa. A diversity of small to medium-sized 
cranes resembling die crowned cranes of the modern African 
genus Balearica have been reported from Europe and North 
NUMBER 62 
America (Olson, 1985). The forms of Balearica are smalhsh 
cranes diat today occur in wet habitats across much of 
sub-Saharan Africa. They may forage in open grasslands, but 
nest only in heavily vegetated swamps where die cover is tiiick 
and high enough to hide die birds (Johnsgard, 1983). Balea?
rica, alone among modern cranes, frequendy roosts in trees. 
The diet includes a diversity of animal and vegetable foods. 
Order CHARADRHFORMES 
Family jACANroAE 
The distinctive lily-trotters or jacanas are represented by a 
diversity of species in die Fayum deposits, all of die elements 
recovered so far being die distal ends of tarsometatarsi, which 
are highly distinctive in having the huge distal foramen, broad 
tendinal groove, and flattened shaft unique to diis family. 
Modern jacanas are much oversplit at the generic level, widi 
the seven or eight species being placed in six genera. A more 
realistic classification is diat of Strauch (1976), who recognized 
only two genera based on differences in morphology and 
plumage of the wing. In Jacana (including Hydrophasianus) 
die wings are distincdy patterned, the radius is unmodified, 
and there is a strong carpal spur. In Metopidius (including 
Actophilornis, Microparra, and Irediparra), the wings are 
unpatterned, the carpal spur is lacking, and die radius is greatiy 
expanded distally into a unique broad blade. Fry (1983) has 
noted that Microparra capensis is apparendy a neotenic form 
of Actophilornis, and aldiough the radius is not gready 
expanded in this species, it may secondarily have reverted to 
the primitive condition. Fry's classification, in which all 
jacanas are lumped under the genus Jacana except for the 
monotypic genera Microparra and Hydrophasianus, is at odds 
with the preceding characters and Fry's own observations. 
Except for the modifications of the wing, there are no major 
differences in osteology between the various modern species 
ofjacanas. 
Nupharanassa, new genus 
TYPE SPECIES.?Nupharanassa bulotorum, new species. 
DLSIGNOSIS.?Differs from all modern Jacanidae in having 
the distal foramen of the tarsometatarsus much larger, not 
circular in outline but tapering proximally to produce a 
tear-drop shape; trochleae more nearly in die same proximo-
distal plane, die inner and outer trochleae being more distally 
produced relative to die middle trochlea; scar for hallux and 
the opposite lateral portion of die posterior surface of shaft 
more excavated; anterior portion of shaft lateral to distal 
foramen more expanded and flattened. 
ETYMOLOGY.?Greek nouphar (Latin nuphar), a wateriily, 
plus anassa, a queen. From die association of jacanas widi 
waterlilies, die large size of die fossil forms relative to modern 
jacanas, and in allusion to the giant wateriily Victoria regia, 
FIGURE 6.?Anterior (top row) and posterior (bottom row) views of distal ends of 
tarsometatarsi of Jacanidae: ad, right tarsometatarsus, holotype of Nupharanassa 
tolutaria, new genus and species (DP*C 2850); b,e, left tarsometatarsus, holotype of 
Janipes nymphaeolxites, new genus and species (CGM 42839); cf, right 
tarsometatarsus, holotype of Nupharanassa bulotorum, new genus and species (DPC 
3848). All to same scale. 
also named for a queen. The gender is feminine. 
Nupharanassa bulotorum, new species 
FIGURE 6C, f 
HOLOTYPE.-?DPC 3848, distal end of right tarsometatarsus 
(Figure 6c/). 
TYPE LOCALITY AND HORIZON.?Quarry M, Fayum Province, 
Egypt. Upper sequence of the Jebel Qatrani Formation, Lower 
Oligocene. Collected by R.H. Madden. 
MEASUREMENTS OF HOLOTYPE.?^Width at base of trochleae, 
11.4 mm; distal width, 12.2 mm; lengdi of distal foramen, 3.6 
mni. 
REFERRED SPEQMENS.?DPC 5327, distal end of left 
8 
SMITHSONIAN CONTEUBUTIONS TO PALEOBIOLOGY 
tarsometatarsus lacking the distal portions of the trochleae, 
from Quany M; DPC 5328, distal end of left tarsometatarsus 
lacking lateral portion of shaft and outer d-ochlea, also from 
Quarry M. 
DIAGNOSIS.?Much larger dian any previously known 
species of Jacanidae, being some 30 to 35 percent larger dian 
a male oi Metopidius indicus, die largest living species. 
ETYMOLOGY.?"Of large lotuses" from Latin, bu, large, and 
die genitive plural of lotus (from die Greek lotos). 
REMARKS.?The referred specimens are smaller than the 
holotype but probably fall within the size range of die species, 
given die pronounced sexual dimorphism in size in extant 
jacanas. In die holotype there is no osseous bridge defining the 
intertrochlear foramen, so diat diis opening is continuous widi 
die distal foramen. It is difficult to say whether die absence of 
diis bridge is normal or due to wear, diough there is no clear 
sign of breakage. By contrast, diere is a distinct intertrochlear 
foramen in die referred specimen, DPC 5327. 
Nupharanassa tolutaria, new species 
FIGURE 6a, d 
HOLOTYPE.?DPC 2580, distal end of right tarsometatarsus 
lacking most of die inner and outer trochleae and part of the 
middle trochlea (Figure 6a,(f). 
TYPE LOCALITY AND HORIZON.?Quarry E, Fayum Province, 
Egypt. Lower sequence of die Jebel Qatrani Formation, Lower 
Oligocene. Collected by R.F. Kay. 
MEASUREMENTS.?^Width at base of trochleae, 6.5 mm; 
length of distal foramen, 2.4 mm. 
DIAGNOSIS.?Although badly damaged, die specimen pre?
serves most of die characters distinguishing Nupharanassa 
from other jacanas. The species is much smaller dian N. 
bulotorum, falling within the size range of all but the smallest 
species of modern jacanas. 
ETYMOLOGY.?^Latin, tolutarius, trotting; from the modern 
name "hly-trotter," often applied to the jacanas. 
REMARKS.?This specimen is significant in that it comes 
from die lower sequence, about 150 m below die odier Fayum 
fossils of jacanas. Aldiough the mammalian faunas ofthe upper 
and lower sequences are broadly similar, few species overlap 
both levels, and in some cases genera or families present at 
one level are absent from die other (for example, die 
propliopithecid primates occur only in the upper quarries). The 
two levels thus represent two significantly different temporal 
and ecological assemblages. The entire 340 m of the Jebel 
Qatrani evidently represents a period of time on the order of 
8-10 million years, 150 m may span 3-4 million years or more, 
and the presence of a jacana in the lower sequence thus 
suggests diat ecological conditions suitable for jacanas 
probably persisted diroughout at least the major period of fossd 
deposition in die Jebel Qatrani Formation. 
Janipes, new genus 
TYPE SPECIES.?Janipes nymphaeobates, new species. 
DIAGNOSIS.?Differs from Nupharanassa and all living 
jacanas in the proportionately much smaller size of the distal 
foramen, the relatively broader and more flattened shaft, the 
shghtly wider intertrochanteric notches, and die more exca?
vated anterior and posterior portions of the lateral part of the 
shaft. 
ETYMOLOGY.?^"Foot with an arched passageway," from 
Latin janus, arched passageway, connecting vowel "i," plus 
pes, foot; for die distinctive shape of the large distal foramen 
as it passes through the bone. The gender is masculine. 
Janipes nymphaeobates, new species 
Figure 6b,e 
HOLOTYPE.?CGM 42839, distal end of left tarsometatarsus, 
with abraded trochleae (Figure 6b,e). 
TYPE LocALrrv AND HORIZON.?Quarry M, Fayum Province, 
Egypt. Upper sequence of the Jebel Qatrani Formation, Lower 
Oligocene. Collected by J.G. Fleagle. 
MEASUREMENTS.?^Width at base of trochleae, 9.2 mm; 
length of distal foramen, 1.9 mm. 
DIAGNOSIS.?This species exceeds all extant forms in size, 
but is smaller dian Nupharanassa bulotorum. 
ETYMOLOGY.?Greek, nymphaia (Latin stem nymphae- plus 
connecting vowel "o"), wateriily, and bates, a walker; again 
from the habits of jacanas of walking on wateriilies. 
REMARKS.?The only Tertiary member of the Jacanidae yet 
recognized is die much younger Jacana farrandi Olson from 
the late Miocene (latest Clarendonian and early Hemphillian) 
of Florida (Olson, 1976; J. Becker, personal communication). 
The Fayum specimens constitute the earliest record of die 
family and the first for Africa. Janipes exhibits derived features 
that indicate the Oligocene radiation of this family involved 
more dian just size differentiation. Neverdieless, the large size 
of both upper sequence taxa is remarkable. The only modern 
species of jacanas diat are sympatric are "Actophilornis" 
africanus and "Microparra" capensis in Africa, which differ 
greatiy in size, and Metopidius indicus and "Hydrophasianus" 
chirurgus in Asia, which belong to the two different groups 
of jacanas and also differ, diough less significandy, in size. 
The habitat requirements of jacanas are quite restricted. 
They are found exclusively in tropical regions on quiet bodies 
of fresh water widi lush floating vegetation, particularly 
Nymphaeaceae. Their toes are greatiy expanded as an 
adaptation for spreading dieir weight over a greater area to 
prevent sinking into the vegetation, which accounts for die 
extreme specialization of the tarsometatarsus. The Fayum 
jacanas are simUarly adapted, so similar habitats must have 
been present at the time of deposition. In addition, die much 
larger size of two of die fossil forms suggests that the 
NUMBER 62 
vegetation present was particularly dense or diat die species 
of Nymphaeaceae present were very large. 
The value of jacanas as paleoenvironmental indicators is of 
special interest widi regard to Nupharanassa tolutaria. which 
comes from die enigmatic Quarry E. This has yielded a higher 
proportion of shark teedi and skate plates dian other Fayum 
quarries. The small size and poor condition of odier fossds in 
parts of diis quarry appear to indicate diat diey have been 
reworked or transported some distance. The occurrence of a 
stricdy fresh-water species such as die jacana, along widi skates 
and sharks, which prefer brackish or salt water, tends to support 
this conclusion. 
Family PHOENICOPTERIDAE 
Genus Indeterminate, alT. Palaelodus, species 1 
FIGURES 7, Sa 
MATERIAL EXAMINED.?DPC 5326, abraded distal end of 
right tibiotarsus (Figure 7). DPC 2646, proximal and distal 
ends of a left carpometacarpus lacking die intermediate 
portions of die major and minor metacarpals (Figure Sa). DPC 
3083, proximal end of left carpometacarpus. 
LOCALPFY AND HORIZON.?Carpometacarpus DPC 3083, and 
tibiotarsus 5326, Quarry M; carpometacarpus DPC 2646, 
Quarry I; all upper sequence. 
MEASUREMENTS.?Carpometacarpus DPC 2646: proximal 
depdi, 11.8 mm; widdi of trochlea, 4.8 mm; distal depdi x 
widdi, 5.9 x 3.9 mm. Tibiotarsus DPC 5326: distal widdi 10.8 
mm; condylar depth, 10.4 mm (a minimum figure as die 
condylar edges are abraded). 
COMPARISONS.?The tibiotarsus, DPC 5326, has a deep, 
distinct anterior intercondylar fossa as in die Phoenicopteridae 
and Ciconiidae (Figure 7). Unlike the latter, the fossa is not 
round, but radier trapezoidal in shape, and the lateral part of 
the fossa extends deeply into die proximal part of the external 
condyle (see Olson and Feduccia, 1980), a feature not present 
in ciconiids, nor in other Charadriiformes. The distal opening 
of the tendinal foramen is separated from the intercondylar 
fossa by a distinct ridge of bone that extends medially to die 
internal condyle, a distinctive characteristic of flamingos. The 
anterior intercondylar tubercles, although abraded, appear to 
have been weak, and the peroneal grooves on the lateral part 
of the external condyle are absent In these details the Fayum 
specimen differs from modern flamingos and more closely 
matches specimens of Palaelodus. 
The Fayum tibiotarsus differs from that of Palaelodus 
primarily in the absence of a strong protuberance on the lateral 
surface of the external condyle, and the shghdy smaller size 
ofthe tendinal foramen. In size, the fossd closely matches die 
holotype oi Palaelodus gracilipes Milne-Edwards. 
One of the two carpometacarpi, DPC 3083, is poorly 
preserved, but so far as can be determined agrees in detail with 
METRIC 1 
FIGURE 7.?Anterior view of distal end of right tibiotarsus (DPC 5326) of a flamingo 
(Phoenicopteridae) resembling Palaelodus. The porous appearance of the condylar 
edges is due to abrasion that has removed the smooth outer surface. 
the Other carpometacarpus, DPC 2646, to which aU following 
comments pertain. The fossd resembles the carpometacarpus 
in die Phoenicopteridae (especially Palaelodus) and Podicipe-
didae in having a straight, splint-like minor metacarpal that 
runs closely parallel with the major metacarpal, leaving a 
narrow intermetacarpal space (Figure Sa,b). Odier features 
10 SMITHSONL\N CONTRIBUTIONS TO PALEOBIOLOGY 
METRIC 1 
FIGURE 8.?^Internal views of left carpometacarpi of fossil flamingos (Phoenicopte?
ridae): a, middle Oligocene specimen from the Fayum (DPC 2646); b, late Oligocene 
to early Miocene (Aquitanian) specimen of Palaelodus crassipes from France 
(USNM 256086). The shape of the alular metacarpal in the specimen of Palaelodus 
has been altered somewhat by severe abrasion. The length ofthe Fayum specimen 
is unknown, but was probably shorter than the specimen of Pcdaelodus, as indicated. 
that the fossd shares with flamingos and grebes include the 
facet for digit III, which is indistinct and proximal to that for 
digit II, and the long distal symphysis, which is not as extensive 
as in modern flamingos, but closely resembles die condition 
in Palaelodus. The Fayum specimen differs further from the 
Podicipedidae and resembles Palaelodus in having the dorsal 
process of the alular metacarpal much larger and shaped as in 
the Charadriiformes. It differs from all odier phoenicopterids 
in having a deep excavation on the internal side of the base of 
the alular metacarpal. The Fayum carpometacarpus is smaller 
than specimens of Palaelodus crassipes Milne-Edwards at 
USNM, and is about the size expected for P. gracilipes. Since 
the carpometacarpi and tibiotarsus come from the same 
stratigraphic level, and all are from a bird the size of P. 
gracilipes, it is likely that they represent a single species. 
REMARKS.?See account under following species. 
Genus Indeterminate, species 2 
MATERIAL EXAMINED.?^DPC 5513, fragment of left coracoid 
lacking sternal end, fmacular facet, and tip of procoracoid. 
LOCALTTY AND HORIZON.?Quarry M, upper sequence. 
MEASUREMENTS.?DPC 5513: medio-lateral width at narrow?
est point of shaft, 7.1 mm; length and width of glenoid facet, 
7.0 X 6.7 mm. 
COMPARISONS.?This specimen has the stout shaft with a 
marked "waist" and die large medially-projecting perforate 
procoracoid that distinguishes flamingos from storks and other 
large waterbirds. It agrees closely with modern flamingos and 
with the extinct genus Palaelodus as well. The procoracoid is 
slightly smaller and more gracde than in most specimens of 
Phoenicopterus spp., but resembles that in Palaelodus. The 
scapular facet is proportionately smaller dian in Phoenico?
pterus or Palaelodus. The fossil resembles modern flamingos 
in the absence of a strong ridge running down a short distance 
from the furcular facet on the postero-medial side of the head, 
a feature that is prominent in specimens of Palaelodus. The 
size of the Fayum coracoid matches that of Palaelodus 
crassipes and die living species Phoeniconaias minor. 
REMARKS.?^Flamingos are fairly common in the fossil 
record, with described species from North America, Etu^ope, 
Africa, and Australia (Olson and Feduccia, 1980; Olson, 1985). 
The Fayum specimens are the earliest recorded flamingos from 
Africa. They evidendy represent at least two species?a small 
one within the size range of Palaelodus gracilipes of the 
European early Miocene (Aquitanian); and a larger species in 
the range of P. crassipes. The morphology of the smaller 
specimens suggests a radier close affinity with Palaelodus, 
while the generic relationships of the larger species are not 
evident. 
Modern flamingos are normally gregarious birds that prefer 
open, shallow water, usually saline or alkaline, in which they 
filter feed for microorganisms. The genus Palaelodus has 
shorter tibiotarsi and shorter, more laterally compressed 
tarsometatarsi than die modern flamingos, possibly an adapta?
tion for swimming (Olson and Feduccia, 1980). Because die 
Fayum flamingos cannot be definitely referred to modern 
genera, dieir habitat requirements are uncertain, except diat 
NUMBER 62 11 
diey would not have occurred far from water. 
Order "CICONHFORMES" 
The traditional order Ciconiiformes appears to be a 
polyphyletic assemblage (see Olson, 1979,1985) and diere are 
considerable uncertainties siuTounding the relationships of die 
herons (Ardeidae) in particular. Even more uncertainty attaches 
to two additional fossil specimens that we have not been able 
to refer to any Uving family of birds. These, a rostrum and 
most of a humerus, are among the best preserved of the avian 
fossils from die Fayum (Figures 9-11). They share more 
simdarities widi herons than with any other group of birds, but 
are so distincdy different that a new famUy is warranted for 
them. 
The rostrum is very heavdy ossified, widi the ventral bars 
of die premaxiUae completely fused so diat the ventral surface 
of the rostrum is covered with bone as in the core 
Ciconiiformes, Ardeidae, and Pelecaniformes. The fossil 
rostrum differs from all of those groups except the Ardeidae 
in having large, open nostrils with narrow, well-defined nasal 
bars. The diagnosis therefore emphasizes the characters 
FIGURE 9.?Rostrum (holotype, DPC 3088) of Xenerodiops rrtycter, new genus and species: a, left lateral view; b, dorsal 
view; c, ventral view. About natural size, bar = 40 mm. 
12 SMrraSONL\N CONTRIBUTIONS TO PALEOBIOLOGY 
FIGURE 10.?^Drawing of the rostrum (holotype, DPC 3088) of Xenerodiops mycter, new genus and species, in left lateral 
view. About five-thirds natural size (total length of specimen, 88 mm). 
differentiating the new family from the Ardeidae. 
Family XENERODIOPIDAE, new family 
TYPE GENUS.?Xenerodiops, new genus. 
DIAGNOSIS.?The rostrum is heavy and decurved, with 
weU-defined lateral grooves that delimit a somewhat swollen 
terminus, quite unlike the straight, sharply pointed rostrum of 
herons. The dorsal bar is narrow and deep, and does not become 
expanded and flattened posteriorly as in die Ardeidae. It 
appears that die nasal septum may have been completely 
ossified (aldiough it is broken posteriorly so this cannot be 
determined with certainty), whereas in die Ardeidae diere is 
practically no indication of an ossified nasal sepmm. The nasal 
bars are thick and almost cylindrical, unlike the thin flattened 
nasal bars in the Ardeidae. The posterolateral corner of die 
premaxilla is a large, deep, posteriorly directed flange above 
which the quadratojugal bar articulates at some distance 
dorsally; in herons this is a slender process that runs under, 
and is in contact widi, the quadratojugal bar. The humerus is 
on first appraisal heron-like but more robust, with a more 
expanded and triangular pectoral crest and a more rounded 
bicipital crest that is much more distinctiy set off from die 
shaft. Unlike herons or any of the "core" Ciconiiformes, the 
tricipital fossa is excavated and not pneumatic, although there 
are a few small foramina around die proximal margin, as in the 
Phoenicopteridae. 
Xenerodiops, new genus 
TYPE SPECIES.?Xenerodiops mycter, new species. 
DUGNOSIS.?As for the family. 
ETYMOLOGY.?Formed from Greek xenos, stranger; erodios, 
a heron; and ops, countenance; from die very peculiar, though 
heron-ldce, appearance of the rostrum. The name is used as 
masculine in gender. 
Xenerodiops mycter, new species 
FIGURES 9-11 
HOLOTYPE.?DPC 3088, nearly complete rostrum lacking 
the dorsal portion of the descending process of the right nasal 
bar and the posterior portion of the dorsal bar and attached 
parts of die nasal septum; on the ventral surface, the posterior 
third of the right side is cracked and lacking some bone, and 
the posterior portion of the premaxilla has been compressed 
medially, but the left side is intact and appears undistorted 
(Figures 9, 10). 
TYPE LOCALTTY AND HORIZON.?Quarry M, Fayum Province, 
Egypt. Upper sequence of the Jebel Qatrani Formation, Lower 
Oligocene. Collected by Simons. 
MEASLIREMENTS OF THE HOLOTYPE.?Length of ventral chord 
from tip to posterior flange of premaxilla, 86.2 mm; least width 
of dorsal bar, 4.3 mm; greatest proximal width (measured from 
left side to midline and dien doubled) 26.2 mm; widdi at most 
swollen point of tip, 7.1 mm. 
PARATYPE.?DPC 2350, left humerus lacking distal end, 
from Quarry I, upper sequence, Jebel Qatrani Formation, 
Fayum district, Egypt (Figure 11). Collected by J.G. Fleagle. 
MEASUREMENTS OF PARATYPE.?^Estimated total lengdi, 120 
mm; width and depth of shaft at approximate midpoint, 8.8 
x6.5 mm; width through dorsal and ventral tubercles, 24.1 
mm; depth of head, 7.3 mm. 
DIAGNOSIS.?As for die famdy. The humerus indicates a 
species somewhat smaller than the smallest modern stork, 
Ciconia abdimii, and probably larger than Ardea alba and 
most modern herons except the larger species of Ardea. The 
proportions of the rostrum are so different from those of 
possible relatives that size comparisons are very difficult, but 
the width at die base would seem to indicate a species of about 
the same size as suggested by die humerus. 
ETYMOLOGY.?Greek mykter, nose, with regard to die most 
salient attributes of the species. The name is a masculine noun 
NUMBER 62 13 
FIGURE 11.?Left huunerus (paratype, DPC 2350) of Xenerodiops mycter, new genus and species: a, anccnal view; 
fepalmar view. About natural size, bar = 40 mm. 
in apposition. 
REMARKS.?The nostril in Xenerodiops was holorhinal and 
die skull highly kinetic. To judge by die remaining portions 
of the nasal bars, there was a well-developed naso-frontal 
hinge, probably more like that in storks or Cochlearius dian 
that in typical herons. Fimctionally, as well as morphologically, 
the bill in Xenerodiops was adapted for strong biting, and it 
appears intermediate between the pointed spearing device such 
as diat typical of herons, and the cylindrical, decurved bill of 
storks of the genus Mycteria (sensu lato), which is adapted for 
feeding tactilely in turbid or vegetation-choked water by 
grasping prey on contact with a "bill-snap" reflex (Kahl and 
Peacock, 1963). 
The humerus is assigned to diis species because it came from 
a bird of simdar size to the holotype and because it is heron-like 
but distinctiy different from all modern herons. Its refeiral can 
only be regarded as tentative at diis point. As widi the rostrum, 
the humerus of Xenerodiops could be viewed as being 
intermediate in structure, as die bicipital and pectoral crests are 
better developed dian in the Ardeidae and not as expanded as 
in die Ciconiidae. The phylogenetic significance of Xenero?
diops can be determined satisfactorily only with additional 
material. 
Family ARDEIDAE 
Nycticorax sp. 
FIGURE 12 
MATERIAL EXAMINED.?CGM 42840, nearly complete right 
tarsometatarsus lacking distal portions of the trochleae and 
most of die proximal articular siuface (Figure 12); DPC 3856, 
left coracoid lacking the sternal end. 
HORIZON AND LOCALTTY.?^Bodi specimens are from Quarry 
M, upper sequence. 
MEASUREMENTS.?^Tarsometatarsus CGM 42840: approxi?
mate length, 74 mm; width and depth at base of trochleae, 6.9 
X 3.0 mm. Coracoid DPC 3856: depth at glenoid facet, 7.3 mm. 
14 SMiraSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 
METRIC 1 
l l f 
FIGURE 12.?Anterior view of right tarsometatarsus (CGM 42840) of a night heron, 
Nycticorax sp. (Ardeidae). 
COMPARISONS.?The tarsometatarsus is relatively short and 
robust, with die trochleae short and widely spaced (Figure 12). 
Proximally, the medial side of the shaft is markedly thin, 
tapering to a sharp edge. In these respects the specimen differs 
from all modern genera of herons compared {Ardea, Syrigma, 
Pilherodius, Cochlearius, Tigrisoma, Botaurus) except Nycti?
corax and Egretta sacra. From die last it differs in die 
morphology of die medial flange. The specimen closely 
matches Nycticorax in all detads, falling well widiin the range 
of variation of USNM specimens of N. nycticorax. The long, 
narrow shaft of the fossd coracoid is also characteristic of 
Nycticorax (Olson, 1977) and it is likely therefore that diis 
specimen is referrable to the same species as the tarsometa?
tarsus. 
REMARKS.?Although we are hesitant to assert the existence 
of a modern genus of heron in the Oligocene of Egypt, diere 
is no basis for considering diese specimens as belonging to 
anything odier than a species of Nycticorax. If the unusual 
tarsometatarsal anatomy of diis genus is not primitive for the 
Ardeidae, it would indicate that the night herons separated from 
other members of the family at least 31 million years ago. This 
is the first Tertiary record of die genus. Night herons are 
crepuscidar or nocturnal in habits and feed in aquatic 
environments. 
Genus and Species Indeterminate 
FIGURE 13 
MATERIAL EXAMINED.?CGM 42838, rostrum lacking poste?
rior portions (Figure 13). CGM 42835, distal end of a left 
tarsometatarsus of a juvenile individual. DPC 3085, complete 
but weathered right pedal phalanx 1 of digit II. 
LOCALTTY AND HORIZON.?The rostrum is from Quarry I, 
upper sequence; the other two specimens are from Quarry M, 
upper sequence. 
MEASUREMENTS.?^Rostrum CGM 42838: width and depdi 
40 mm from tip, 11.2 x 6.8 mm. Tarsometatarsus CGM 42835: 
widdi and depth at base of trochleae, 8.8 x 3.7 mm. Phalanx 
DPC 3085: length, 28.3 mm; proximal widdi and depdi, 7.6 
x3.7 mm; distal width, 3.6 mm. 
COMPARISONS.?The rostrum (Figure 13) is the only 
reasonably diagnostic element among the above three spec?
imens, all of which are of a size compatible widi a 
medium-sized modern heron such as Ardea alba or Egretta 
rufescens. Because there is no good evidence to suggest that 
more than one species is represented in this material, all the 
specimens are considered together here, although there is 
likewise no proof that all come from a single species. 
The rostrum is long, narrow, and nearly straight. It is not 
unlike that in the extant genus Egretta, but is more compressed 
dorsoventrally, with a narrower median ridge, strong rostral 
grooves, and a more concave ventral surface. This combination 
of characters distinguishes it from any of die extant species of 
herons examined. 
The juvenile tarsometatarsus is almost completely undiag?
nostic except that die alignment of die trochleae in the same 
antero-posterior plane, and the small distal foramen and faint 
tendinal groove, make it recognizable as coming from a heron. 
The pedal phalanx has the broad, flat proximal end diat is quite 
characteristic of the basal phalanges in the Ardeidae. 
REMARKS.?^Although one woidd expect from their relatively 
NUMBER 62 15 
FIGURE 13.?^Dorsal view of rostrum (CGM 42838) of a medium-sized heron 
(Ardeidae). 
large size and generally aquatic habits that herons would have 
a reasonably good fossil record, they are actually quite scarce 
in Tertiary deposits, many previous records having been based 
on misidentified specimens (Brodkorb, 1980; Olson, 1985). 
There are scattered records of true herons beginning in the late 
Eocene or early Ohgocene of France and extending through 
die remainder of die Tertiary, aldiough die relationships of 
most of these fossils to living herons have not been firmly 
established. Hitherto, die earliest record for the Ardeidae in 
Africa was Zeltornis ginsburgi Balouet (1981), known from 
an incomplete coracoid from die early Miocene of Libya. The 
sample of fossil herons from the Fayum provides the earliest 
representation of the family for the continent. 
Family CICONITOAE 
Palaeoephippiorhynchus dietrichi Lambrecht, 1930 
HOLOTYPE.?A nearly complete rostrum and mandible with 
associated occipital region of cranium; specimen in Stuttgart. 
TYPE LOCALTTY AND HORIZON.?^North of Qasr Qarun, Jebel 
Qatrani Formation. 
REMARKS.?See following account. 
?Palaeoephippiorhynchus dietrichi Lambrecht, 1930 
FIGURE 14 
MATERIAL EXAMINED.?DPC 2347, distal end of right 
tibiotarsus. 
LOCALTTY AND HORIZON.?Quarry M, upper sequence. 
MEASLIREMENTS.?Intercondylar widdi, 17.9 mm; condylar 
depth, 24.2 mm. 
COMPARISONS.?The specimen is slightiy abraded and only 
a short portion of the shaft is present. It resembles die 
Ciconiidae and differs from the somewhat similar Balaenicipi-
tidae and Vulturidae in the deep, narrow tendinal groove, and 
the distinct intercondylar fossa. It differs from recent 
Ciconiidae in the weak intercondylar tubercle (not just due to 
abrasion), the relatively greater intercondylar width, the 
transversely furrowed supratendinal bridge, and the smaller 
distal tendinal opening. In size it would rank among die larger 
living storks such as Leptoptilos and Jabiru, and therefore 
would presumably have been about the size oi Palaeoephippio?
rhynchus dietrichi. That species was described as being most 
simdar in bdl morphology to the extant genus Ephippio-
rhynchus, whereas the fossU tibiotarsus shows no particular 
resemblance to that of any living genus. Thus it cannot 
certainly be established whether one or two species of storks 
are represented in the Fayum deposits. 
REMARKS.?Palaeoephippiorhynchus is the earliest certain 
ciconiid. Modern storks may either be highly aquatic or may 
be scavengers that frequent open grassland and savanna. 
Ephippiorhynchus is one of those that is very aquatic in its 
preferences, so if the similarities in feeding adaptations 
between it and Palaeoephippiorhynchus are any indication, die 
latter may have been also. 
Family BALAENiciPiTroAE 
Goliathia andrewsi Lambrecht, 1930 
HOLOTYPE.?Complete ulna, British Museum (Natural His?
tory), A883. 
16 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 
FIGURE 14.?Anterior view of distal end of right tibiotarsus (DPC 2347) of a large 
stork (Ciconiidae), possibly referable to Palaeoephippiorhynchus. 
formation that is definitely referable to that famdy. Because it 
is unlikely that there were two species of shoebilled storks of 
the same size present at the time of deposition, it is reasonable 
to assume that both specimens refer to the same species, or at 
least a single evolving lineage, as the holotype probably comes 
from a stiatigraphic level considerably lower than that of the 
referred tarsometatarsus. Whether Goliathia can actually be 
separated generically from Balaeniceps remains to be deter?
mined. 
The only previous fossil occurrence of a shoebdled stork is 
another distal end of a tarsometatarsus from the late Miocene 
of Tunisia (Rich, 1972). This specimen is smaller than in 
modern Balaeniceps or the Fayum tarsometatarsus. Rich 
considers the Tunisian form to be generically distinct from die 
modern one based on size and the morphology ofthe trochleae. 
The lack of trochlear surfaces on die Fayum specimen makes 
direct comparison impossible. 
The only extant species in this family, the shoebdl, B. rex, 
is confined to east-central Africa and inhabits fresh-water 
swamps that are thickly overgrown with vegetation, upon 
which it walks by means of its large feet. It is structurally and 
behaviorally highly adapted for capturing large prey in heavy 
vegetation, most commonly lungfish, catfish, and bichirs 
(Guillet, 1979). Lungfish and catfish are abundant in the 
Fayum fauna (Bown et al., 1982). The shoebill, along widi die 
jacanas, provides strong evidence for an environment of 
fresh-water swamp much overgrown with vegetation. 
Order PELECANIFORMES 
Family PHALACROCORACIDAE 
Genus and Species Indeterminate 
TYPE LOCALITY AND HORIZON.?Fayum Province, lower 
sequence, Jebel Qatrani Formation. 
REFERRED SPEQMEN.?DPC 2303, distal end of right 
tarsometatarsus lacking trochleae. 
LOCALTTY AND HORIZON OF REFERRED SPEQMEN.?Quarry 
M, upper sequence. 
MEASUREMENTS OF REFERRED SPECIMEN.?Width at level of 
distal foramen, 23 mm (thus matching that in a specimen of 
Balaeniceps rex in the Smithsonian collections). 
COMPARISONS.?The specimen shows die distinctive flat?
tened shaft of the Balaenicipitidae, with the remains of die 
trochleae in die same antero-posterior plane, and the distal 
foramen situated far distally in a deep, V-shaped tendinal 
groove. 
REMARKS.?^The holotypical ulna of Goliathia andrewsi was 
originally described as diat of a heron, but Brodkorb (1980) 
has presented cogent arguments for referring it to the 
Balaenicipitidae. The validity of tiiis assignment is corrobo?
rated by die discovery of an additional fossil from the same 
FIGURE 15 
MATERIAL EXAMINED.?DPC 5658, distal portion of a 
premaxilla. 
LOCALTTY AND HORIZON.?Quarry M, upper sequence. 
MEASUREMENTS.?Minimum height proximal to swollen 
terminus, 4.2 mm; width at same point, 4,5 mm. 
COMPARISONS.?The palatal surface of die bdl is straight 
along the proximal-distal axis, not recurved as in modern 
Phalacrocorax auritus, P. olivaceus, and some odiers. The bill 
broadens only gradually towards die base, unlike the robust, 
distinctiy tapering rostrum of P. atriceps and some odier 
modern species; nor is the bill distinctiy compressed dorsoven?
trally as in P. pelagicus. The terminus has a moderate to strong 
hook, which differs from the slight hook typical of many 
modern species. In size, proportions, and shape, die Fayum 
fossil compares best, out of 20 modern species examined, with 
P. bougainvillii, a medium to large marine cormorant widi a 
proportionately long, narrow bill. The proximal portion of the 
fossil specimen seems to show the rosfral groove opening into 
NUMBER 62 17 
METRIC 1 
FIGURE 15.?Right lateral view of rostrum (DPC 5658) of a cormorant (Phalacrocoracidae). 
a nostril, but this may be due to breakage. 
The rostral groove of the fossd, and the series of foramina 
that line the groove, are simdar to modern cormorants. In 
modern species the distal end of the groove curves strongly 
downward and empties gutter-like to the lateral palatal edge. 
In the Fayum specimen the rostral groove connects weakly 
widi a more distal and lower groove that ends abruptiy past the 
swollen terminus but does not continue to the lateral edge of 
the bid. In addition, the fossd preserves a pair of deep sulci 
miming antero-posteriorly across the dorsal surface of the 
most curved, swollen part of the terminus, which differs from 
the two shallow grooves diat enter small foramina well 
proximal to the hooked terminus in all modern cormorants. In 
both of these features?the distal configuration of the rosfral 
groove, and the superior sidci at the hooked terminus?the 
specimen differs from all modern cormorants but is nearly 
identical to modern species of Sulidae. The Fayum fossil differs 
completely from sulids in other aspects of gross morphology. 
REMARKS.?^The configuration of the grooves on the distal 
portion of the rostrum are best interpreted as primitive features 
reflecting a common ancestry with the Sulidae. The Fayum 
specimen must either predate, or be on a separate lineage from, 
the common ancestor of modern species of Phalacrocorax. 
The earliest known cormorant is from the Eo-Ohgocene 
Phosphorites du Quercy, and numerous specimens referred to 
Phalacrocorax have been described from die early Miocene 
onward in North America and Europe (Olson, 1985). The 
Fayum specimen is the earliest recorded cormorant from 
Africa. 
Modern cormorants are cosmopolitan in distribution, rang?
ing broadly over coastal and freshwater habitats. They dive in 
open, calm or flowing water for fish, crustaceans, and other 
aquatic animals. The Fayum cormorant, along with the ospreys 
and flamingos, suggests that in addition to the heavily 
overgrown swampy areas, there were open areas free of 
vegetation. 
Discussion 
The environment indicated by the Fayum bird assemblage 
differs from previous reconstructions in its emphasis on slow 
current and lentic conditions in association widi a great deal 
of emergent aquatic vegetation. Lidiologic characteristics of 
the Fayum quarries indicate that deposition of most bones 
occurred on point bars by active current There is no doubt, 
however, that swampy areas were developed along certain 
sections of river, along the river borders, or perhaps in 
extensive over-bank areas. A variety of mudstones found in the 
upper sequence, including parts of Quarries I and M, were 
perhaps deposited in such an environment (Bown and Kraus, 
1987). Vegetation-choked swamp may well have been die 
preferred habitat of some of the large, stocky-bodied mamma?
lian herbivores such as the anthracotheres and giant hyracoids, 
which are among the most common of the Fayum mammals. 
The fossil avifauna of the Fayum doubtiess has not sampled 
all of the habitats present in the area at the time of deposition, 
but is dominated by birds from a habitat that had not been 
clearly identified previously. The absence of certain groups 
commonly found in modern tropical African rivers such as 
ducks, ibises, shorebirds other dian Jacanidae and Phoenicopter-
18 SMiraSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 
idae, and certain land birds such as Coraciiformes, Galliformes, 
and Passeriformes, may be due eidier to sampling bias or to 
these taxa actually being absent in northern Africa in the early 
Oligocene. Ducks and passeriforms, for example, do not appear 
in European and North American deposits untd about the 
beginning of die Miocene. Rich (1974) noted the unexpected 
absence of anatids, galliforms, and passerines from the middle 
to late Miocene Beglia Formation of Tunisia, Odierwise, die 
Tunisian assemblage includes taxa now typical of sub-Saharan 
Africa. 
From a zoogeographic standpoint, die Fayum avifauna is 
extremely similar to one that would be expected along a 
fropical swampy river in central Africa today. The Balaenicipi?
tidae and Musophagidae are now restricted to Africa, although 
fossil musophagids are known from the Tertiary of Europe. 
The Fayum fossils belonging to the Pandionidae and Jacanidae 
are by far the earliest records for diese famdies, both of which 
are now widely distributed, the Jacanidae in the New and Old 
World tropics, and the Pandionidae worldwide. The Accipi?
tridae, Rallidae, Gruidae, Phoenicopteridae, Ciconiidae, Ar?
deidae, and Phalacrocoracidae are represented elsewhere by 
early Tertiary fossils from die New and Old World, and each 
is widely disdibuted today. The avian assemblage from the 
Fayum corroborates evidence from other localities in the 
Tertiary of North Africa (Brunet, 1971; Rich, 1972) suggesting 
that diere has been relative stabdity in some components of the 
African avifauna since at least the beginning of the Ohgocene. 
The principal change observed is the retreat of numerous taxa 
from the northern part of the continent doubtiess as a result of 
the geologically very recent development of the Sahara desert. 
All of the birds in the Fayum sample belong to living 
families except the supposed ratite and Xenerodiops. This is 
in contrast to the fossd mammals, which include numerous 
forms grossly different from modern African groups, such as 
anthracotheres, arsinoitheres, creodonts, ptolomaiids, giant 
hyracoids, parapidiecid and tarsiid primates, and marsupials. 
For this reason, the fossil birds are superior to mammals as 
indicators of past environmental conditions and permit 
paleoecological inferences to be made with some confidence. 
In Oligocene times, the Fayum contained a low swampy river, 
or series of rivers, overgrown in places widi papyrus, reeds, 
and floating vegetation such as Salvinia and water Idles (Wing 
and Tiffhey, 1982). Analysis of the fossd birds shows that the 
most closely analogous modern avifauna exists today only in 
a Umited area of Uganda, north and west of Lake Victoria 
(Olson and Rasmussen, 1986). This region presents several 
odier striking parallels widi die biota and environment known 
or inferred for the Fayum in the early Oligocene. Our evidence 
from fossil birds strongly supports die paleoenvironmental 
reconstruction of Bown et al. (1982), and is at odds widi die 
treeless, arid, sahelian habitat suggested for the Fayum by 
Kortlandt (1980). 
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