Transcriber Note: Text emphasis denoted as _Italics_ and =Bold=.
The Cretaceous Birds
of New Jersey
STORRS L. OLSON
and
DAVID C. PARRIS
SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY • NUMBER 63
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SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY · NUMBER 63
The Cretaceous Birds of New Jersey
Storrs L. Olson and David C. Parris
[Illustration]
SMITHSONIAN INSTITUTION PRESS
Washington, D.C.
1987
ABSTRACT
Olson, Storrs L., and David C. Parris. The Cretaceous Birds of New
Jersey. Smithsonian Contributions to Paleobiology, number 63, 22
pages, 11 figures, 1987.--This is a revision of the fossil birds from
Late Cretaceous (Maastrichtian; Hornerstown and Navesink formations)
deposits in New Jersey. Material of previously named taxa, described
over a century ago, is augmented by more recently collected specimens
from a new locality at the Inversand Company marl pits near Sewell,
Gloucester County. With about 8 genera and 9 species, this is the most
diverse Cretaceous avifauna yet known. Most species belong to a group
of primitive Charadriiformes resembling in limb morphology the fossil
family Presbyornithidae and the living family Burhinidae. These are
tentatively referred to the “form family” Graculavidae Fürbringer,
1888, with its provisional synonyms Palaeotringinae Wetmore, 1940;
Telmatornithidae Cracraft, 1972, and Laornithidae Cracraft, 1972. The
species included are: _Graculavus velox_ Marsh, 1872; _Telmatornis
priscus_ Marsh, 1870 (synonyms: _Telmatornis affinis_ Marsh, 1870;
_Graculavus pumilis_ Marsh, 1872; _Palaeotringa vetus_ Marsh, 1870);
_Anatalavis rex_ (Shufeldt, 1915); _Laornis edvardsianus_ Marsh, 1870;
_Palaeotringa littoralis_ Marsh, 1870; _P. vagans_ Marsh, 1872; and
an undescribed genus and species probably different from any of the
preceding. _Anatalavis_ is proposed as a new genus for Telmatornis rex
Shufeldt, 1915. A new family, genus, and species (Tytthostonychidae,
_Tytthostonyx glauconiticus_) is proposed for a humerus showing
similarities to the Pelecaniformes and Procellariiformes and
tentatively referred to the latter, along with an ulna of a much
smaller species. The species in this fauna appear to be part of the
modern radiation of neognathous birds, but none can be referred to
modern families.
Official publication date is handstamped in a limited number of initial
copies and is recorded in the Institution's annual report, _Smithsonian
Year_. Series cover design: The trilobite _Phacops rana_ Green. /X
Library of Congress Cataloging-in-Publication Data Olson, Storrs
L. The cretaceous birds of New Jersey. (Smithsonian contributions
to paleobiology; no. 63) Bibliography: p. 1 Birds Fossil. 2.
Paleontology--Cretaceous. 3. Paleontology--New Jersey. I. Parris, David
C. II. Title. III. Series. QE701.S56 no. 63 560 s 86-29837 [QE871]
[568’.09749] X/
Contents
Page
Introduction 1
Acknowledgments 1
The Fossil Localities and Their Stratigraphy 1
Order Charadriiformes 4
“Form Family” Graculavidae Fürbringer, 1888 4
Genus _Graculavus_ Marsh, 1872 4
_Graculavus velox_ Marsh, 1872 4
_Graculavus velox?_ 6
Genus _Telmatornis_ Marsh, 1870 6
_Telmatornis priscus_ Marsh, 1870 6
Genus _Anatalavis_, new genus 11
_Anatalavis rex_ (Shufeldt, 1915), new combination 11
Genus _Laornis_ Marsh, 1870 12
_Laornis edvardsianus_ Marsh, 1870 12
Genus _Palaeotringa_ Marsh, 1870 12
_Palaeotringa littoralis_ Marsh, 1870 12
_Palaeotringa littoralis?_ 14
_Palaeotringa vagans_ Marsh, 1872 14
Graculavidae, Genus and Species Indeterminate 14
Order Procellariiformes? 14
Family Tytthostonychidae, new family 16
Genus _Tytthostonyx_, new genus 16
_Tytthostonyx glauconiticus_, new species 16
Family and Genus Indeterminate 16
Aves, incertae sedis 19
Discussion 19
Appendix 20
Literature Cited 21
The Cretaceous Birds of New Jersey
_Storrs L. Olson and David C. Parris_[1]
[Footnote 1: _Storrs L. Olson, Department of Vertebrate Zoology,
National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560. David C. Parris, New Jersey State Museum, 205
West State Street, Trenton, New Jersey 08625-0530._]
Introduction
Fossils of Cretaceous birds are scarce and usually difficult
to interpret. The better known forms such as _Hesperornis_ and
_Ichthyornis_ belong to strange and archaic groups having little or
nothing to do with the modern avian radiation. The only areas that have
yielded Cretaceous birds of essentially modern aspect in sufficient
quantities to be regarded as avifaunal assemblages are the inland
deposits of the Lance Formation and strata of similar age in Wyoming
(Brodkorb, 1963a) and the marine deposits of New Jersey. Of these, the
assemblage from New Jersey is the more diverse.
Fossil birds were described from the Cretaceous greensands of southern
New Jersey over a century ago by Marsh (1870, 1872). These have been
carried, largely uncritically, in lists and compilations ever since
(e.g. Hay, 1902; Lambrecht, 1933; Rapp, 1943; Miller, 1955; Brodkorb,
1963b, 1967). Although some of these specimens were subsequently
re-examined and their status altered (Shufeldt, 1915; Cracraft, 1972,
1973), there has been no modern comprehensive revision of all of the
avian taxa that have been named from the Cretaceous of New Jersey. In
recent years, additional fossil birds have been recovered from these
deposits that add further to our knowledge of late Mesozoic avifaunas,
making a review of this material all the more desirable.
In spite of the relative diversity of the New Jersey Cretaceous
avifauna, the total number of specimens is still small. The decline
of the glauconite greensand industry and the difficulty of recovering
small fossils have contributed to this paucity of specimens. The
glauconite industry is now confined to a single operation, the
Inversand Company in Sewell, Mantua Township, Gloucester County, New
Jersey. Fortunately, the late owner of the company, Mr. Churchill
Hungerford, Jr., generously allowed fossils to be recovered on his
property by the New Jersey State Museum, which houses most of the newly
discovered specimens, the Academy of Natural Sciences of Philadelphia
being the repository of the rest. Another specimen came from a locality
in Upper Freehold Township, Monmouth County, New Jersey and was donated
to the New Jersey State Museum by Gerard R. Case.
Acknowledgments.--We gratefully acknowledge the late Churchill
Hungerford, Jr., for permitting fossil material to be recovered from
his property by the New Jersey State Museum (NJSM). We are much
indebted to John H. Ostrom, Peabody Museum of Natural History, Yale
University (YPM), and Gay Vostreys and Charles Smart of the Academy
of Natural Sciences of Philadelphia (ANSP) for their patience in
lending types and other material from their collections for a very
extended period. Pat V. Rich, Monash University, assisted Parris in
the early stages of this study. Comparative material of _Presbyornis_
was obtained from the collection of the University of California
Museum of Paleontology (UCMP), the University of Wyoming (UW), and the
National Museum of Natural History, Smithsonian Institution (USNM).
The photographs are by Victor E. Krantz, Smithsonian Institution. For
valuable comments on the manuscript we are grateful to Donald Baird,
Princeton University, and Jonathan Becker, Smithsonian Institution.
=The Fossil Localities and Their Stratigraphy=
The extensive deposits of Cretaceous age in eastern North America
have been widely studied for over 150 years. These generally poorly
consolidated sediments have provided valuable resources, notably
glauconite, fire clay, and chalk. As the publications by Morton (1829),
Vanuxem (1829), Conrad (1869), and other early authors showed, the
sediments are also quite fossiliferous.
In the eastern United States, significant Cretaceous deposits occur
from New Jersey to Texas (Figure 1), with extensive outcrop and
subsurface records in both Atlantic and Gulf coastal plains. The
surface distribution and correlations were first summarized by
Stephenson et al. (1942). Subsequent works by various authorities
have refined, but not substantially altered his views of outcrop
stratigraphy. Petroleum exploration has encouraged more recent restudy
of the subsurface stratigraphy, notably along the east coast (Minard et
al., 1974; Perry et al., 1975; Petters, 1976).
[Illustration: Figure 1.--Distribution of Cretaceous rocks in the
eastern United States. Arrow indicates New Jersey. (Modified after
Moore, 1958, fig. 15.2).]
In New Jersey, the latest Cretaceous deposits are remarkably rich
in glauconite, especially the Navesink and Hornerstown formations.
Besides providing a local industry in agricultural fertilizers, the
glauconite greensands, locally called “marl,” yielded many specimens to
the fiercely competitive vertebrate paleontologists of the nineteenth
century. Preservation of vertebrate fossils in a glauconite deposit may
be excellent, apparently due to autochthonous formation of the mineral
and the probable quiescence of the depositional environment. The
Hornerstown Formation, for example, contains few grains of terrigenous
origin and little evidence of disturbance by water currents.
Such depositional environments were apparently favorable for the
preservation of small and delicate bones. The accumulation of sediment
occurred during a period of marine transgression with the shoreline not
far to the northwest but at sufficient distance to prevent deposition
of terrigenous material.
During their great rivalry, E.D. Cope and O.C. Marsh sought greensand
fossils vigorously. Marsh, however, obtained all of the Cretaceous
birds (Marsh, 1870, 1872), largely due to efforts of marl pit owner
J.G. Meirs. Although in the years subsequent to Marsh's original
descriptions of the New Jersey birds from the Navesink and Hornerstown
formations there was some confusion regarding their probable age
(Wetmore, 1930), this was later definitely established as Cretaceous
by Baird (1967), who attributed the specimens to the Navesink and
Hornerstown formations.
The summary of Petters (1976) represents current ideas of the
Cretaceous stratigraphy of New Jersey. Baird's (1967) discussion is
consistent with Petters's view that the Hornerstown Formation is
regarded as partly Cretaceous and partly Tertiary. Some authors have
used the term New Egypt Formation instead of Navesink in more southerly
outcrops.
Cretaceous birds have been recovered from three geographically
distinct localities in New Jersey (Figure 2). With the exception of
_Laornis_, all of the specimens described by Marsh (1870, 1872) came
from Upper Freehold Township, Monmouth County, in the area including
the settlements of Hornerstown, Arneytown, and Cream Ridge. The Meirs
family operated a number of pits in this area and it is no longer
possible to ascertain the exact provenance of specimens labelled only
as being from Hornerstown. These could have come either from the
basal Hornerstown Formation or the underlying Navesink Formation,
both of which are Maastrichtian in age. Baird (1967:261) ascertained
that the holotype of _Palaeotringa vetus_, from “friable green marl
near Arneytown” was from the lower (i.e., Cretaceous) part of the
Hornerstown Formation. The holotypes of _Telmatornis priscus_ and _T.
affinis_, from the Cream Ridge Marl Company pits, on the other hand,
are from the Navesink Formation. A more recently collected specimen
from this area is the proximal end of an ulna (NJSM 11900) collected
by Gerard R. Case from “marl piles near junction of Rtes. 537 and
539 in Upper Freehold Twp., Monmouth County, near Hornerstown.” This
definitely came from the Hornerstown Formation but it cannot be said
whether from the Cretaceous or Paleocene sediments included therein.
[Illustration: Figure 2.--Localities in southern New Jersey of the
main fossiliferous deposits that have yielded Cretaceous birds. (The
bold line demarcates the inner and outer coastal plain physiographic
provinces; B = Birmingham; H = Hornerstown; S = Sewell.)]
The second general locality is near Birmingham, Burlington County,
where the type of _Laornis edvardsianus_ was obtained from “greensand
of the upper, Cretaceous marl bed ... in the pits of the Pemberton Marl
Company” (Marsh, 1870:208). There is nothing to be added to Baird's
(1967) conclusion that this specimen is latest Cretaceous in age.
The third locality, and that yielding most of the recently obtained
specimens, is the Inversand Company marl pit, located near Sewell,
Gloucester County. In accordance with the wishes of the Inversand
Company, the precise locality of this pit will not be disclosed,
although this information is preserved in records sufficient in number
and distribution to assure that it will not be lost. The Inversand
specimens came from the main fossiliferous layer within the basal
portion of the Hornerstown Formation (Figure 3). This layer is of late
Maastrichtian age (latest Cretaceous), on the basis of invertebrate
fossils, including three genera of ammonites, and a substantial
vertebrate fauna, including mosasaurs (see Appendix). It is probable
that the upper part of the Hornerstown Formation within the pit is of
Paleocene age, as it is known to be elsewhere, but most paleontologists
believe the basal portion to be Cretaceous in age (Gaffney, 1975; Koch
and Olsson, 1977). One avian specimen is from an unknown level in the
pit.
[Illustration: Figure 3.--Stratigraphic diagram of the Inversand
Company marl pit at Sewell, Gloucester County, New Jersey.]
=Order Charadriiformes=
=“Form Family” Graculavidae Fürbringer, 1888=
Type Genus.--Graculavus Marsh, 1872.
Included Genera.--_Graculavus_ Marsh, 1872; _Telmatornis_ Marsh, 1870;
_Anatalavis_, new genus; _Laornis_ Marsh, 1870; _Palaeotringa_ Marsh,
1870; and an additional unnamed genus.
Remarks.--Most of the birds from the New Jersey deposits belong with
what Olson (1985) has termed the “transitional Charadriiformes,” a
group that seemingly tends to connect the Gruiformes and the more
typical Charadriiformes. The only living family in this group that
has traditionally been considered charadriiform is the Burhinidae,
the thick-knees or stone curlews. Other apparent descendants include
ibises (Plataleidae) and the ducks and geese of the order Anseriformes.
The latter are linked with the “transitional Charadriiformes” through
the Paleocene and Eocene genus _Presbyornis_, which is known from
abundant material from widely scattered areas of the world (Olson and
Feduccia, 1980b; Olson, 1985). _Presbyornis_ combines a long-legged
shorebird-like body with the head of a duck. The fragmentary Cretaceous
fossils from New Jersey, all of which are postcranial, usually show
more similarity to _Presbyornis_ than to any modern group of birds
except the Burhinidae. Therefore, our comparisons have been made
chiefly with these two groups.
With the fragmentary material at hand it is difficult, well nigh
impossible, to make hard and fast taxonomic judgments concerning the
number of species, genera, or families represented. Birds with very
similar wing or leg elements could have had completely different
feeding adaptations and could represent ancestral forms leading to
different modern groups not considered to be closely related. For
example, without the skull, _Presbyornis_ could not be determined as
having anything to do with the Anseriformes (Olson and Feduccia, 1980b:
12-13).
Late Cretaceous fossil birds of modern aspect have been described in
a variety of genera, most of which have been used as the basis for
family-group names. Taxa from New Jersey that appear to belong with
the “transitional Charadriiformes” for which family-group names are
available include: Graculavinae Fürbringer, 1888; Palaeotringinae
Wetmore, 1940; Telmatornithidae Cracraft, 1972; and Laornithidae
Cracraft, 1973.
Taxa from Upper Cretaceous deposits in western North America that
appear to fall in the same category (Olson and Feduccia, 1980a)
include: Apatornithidae Fürbringer, 1888; Cimolopterygidae Brodkorb,
1963a; Torotigidae Brodkorb, 1963a; and Lonchodytidae Brodkorb, 1963a.
Tertiary taxa that may possibly be related to the “transitional
Charadriiformes” and that have been used as the basis of family-group
names are: Presbyornithidae Wetmore, 1926 (Nautilornithinae Wetmore,
1926, and Telmabatidae Howard, 1955, are definitely synonyms);
Scaniornithidae Lambrecht, 1933; and Dakotornithidae Erickson, 1975.
Doubtless there are others that we have overlooked. How many families
are actually represented here and what their interrelationships may
be is purely a matter of conjecture in the absence of better fossil
material. Because the entire skeleton of _Presbyornis_ is known, the
familial name Presbyornithidae may justifiably be retained and used for
that genus.
In the case of the Cretaceous birds under consideration here, we
have decided for the time being to adopt a version of paleobotanical
convention in recognizing a “form family” Graculavidae, which implies a
general similarity in morphology of the constituent taxa, although the
material available is simply not sufficient for determining phylogeny
or key adaptations.
=Genus Graculavus Marsh, 1872=
_Limosavis_ Shufeldt, 1915:19.
Type-Species.--_Graculavus velox_ Marsh 1872, by subsequent designation
(Hay, 1902).
Included Species.--Type species only.
Remarks.--_Limosavis_ Shufeldt, 1915, substitute name for _Graculavus_,
considered inappropriate; not used in direct combination with any
specific name when originally proposed.
=_Graculavus velox_ Marsh, 1872=
Figure 4 _b,d,f,h_
_Graculavus velox_ Marsh, 1872:363.
_Limosavis velox_ (Marsh).--Lambrecht, 1933:546.
Holotype.--Proximal end of left humerus, YPM 855.
Locality and Horizon.--From Hornerstown, Upper Freehold Township,
Monmouth County, New Jersey; collected by J.G. Meirs; Late Cretaceous
(Maastrichtian), either basal Hornerstown Formation or Navesink
Formation.
Measurements (in mm).--Proximal end of humerus, YPM 855: proximal width
through dorsal and ventral tubercles 21.1, depth through bicipital
surface and tuberculum ventrale 11.6, depth of head 5.7.
[Illustration: Figure 4.--Proximal ends of left humeri of _Graculavus
velox_ and related birds: _a_, _Esacus magnirostris_ (Burhinidae), USNM
19649; _b,d,f,h_, _Graculavus velox_, holotype, YPM 855; _c,e,g, i_,
_Presbyornis_ sp., UCMP 126205. _a-c_, anconal view; _d,e_, anconal
view with distal portion tilted upwards; _f,g_, palmar view; _h,i_,
proximal view. All figures × 2; specimens coated with ammonium chloride
to enhance detail.]
Comparisons.--Marsh (1872) originally described this as a species of
cormorant (Phalacrocoracidae, Pelecaniformes) and included the species
_G. pumilis_ Marsh, 1872, also from New Jersey, and _G. anceps_ Marsh,
1872, from the Late Cretaceous of Kansas, in the same genus. Marsh
(1880) later referred _G. anceps_ to the genus _Ichthyornis_, where it
has remained. Shufeldt (1915:17-19) went into considerable detail to
show that the species of _Graculavus_, particularly _G. velox_, were
not cormorants, instead being limicoline shorebirds with similarities
to the Burhinidae, Haematopodidae, and Charadriidae. Accordingly,
Lambrecht (1933:540, 546) placed these taxa among the charadriiform
birds, but rather inexplicably listed velox under Shufeldt's substitute
name _Limosavis_ in the suborder Laro-Limicolae, while retaining
_pumilis_ in the genus _Graculavus_ in the suborder Limicolae.
Brodkorb (1963b:249) ignored Shufeldt's assessment of relationships
and placed _G. velox_ and _G. pumilis_ in the Phalacrocoracidae,
subfamily Graculavinae. Cracraft (1972) did not examine the specimens
attributed to _Graculavus_ in his consideration of the relationships of
_Telmatornis_.
We have synonymized _Graculavus pumilis_ Marsh, 1872, with
_Telmatornis priscus_ Marsh, 1870, and discuss below the characters
by which _Graculavus_ (restricted to _G. velox_) may be separated
from _Telmatornis_. Shufeldt (1915) has already presented adequate
evidence that _Graculavus_ is not a cormorant and is instead a
charadriiform. The following combination of characters of the proximal
end of the humerus is shared by _Graculavus_ and _Presbyornis_ and
distinguishes these genera from other Charadriiformes: (1) lack of
a distinct lanceolate scar for M. coracobrachialis cranialis; (2)
lack of a distinctly excavated second (dorsal) tricipital fossa; (3)
presence of a distinct tumescence in the proximoventral portion of the
tricipital fossa; scars for (4) M. scapulohumeralis caudalis and (5)
M. scapulohumeralis cranialis very large and distinct; (6) attachment
of M. latissimus dorsi cranialis a well-defined, raised protuberance
situated dorsal to the median ridge of the shaft; (7) tuberculum
dorsale well defined, distinctly pointed. In most of the preceding
characters that it preserves, the single proximal end of humerus
referred to _Telmatornis_ (the holotype of _G. pumilis_) agrees with
_Graculavus_ and _Presbyornis_.
Among living families, the Burhinidae are the most similar to
_Graculavus_; both agree in characters 1, 2, 4, and 7, with certain
species of _Burhinus_ also having characters 3 and 6 present but less
developed. _Graculavus_ differs from Burhinus mainly in having (8) the
head not as deep and bulbous; (9) distance from head to tuberculum
dorsale greater; (10) tuberculum dorsale smaller, much less projecting;
(11) tuberculum ventrale in ventral view more elongate; and (12) scar
on tuberculum ventrale for M. coracobrachialis caudalis much larger and
more distinct.
_Graculavus_ is very similar to _Presbyornis_, agreeing with that
genus in characters 8 and 10 but differing in characters 11 and 12
and in (13) having the head more deeply undercut. _Presbyornis_ is
intermediate between _Graculavus_ and the _Burhinidae_ in character 9.
_Graculavus velox_ was a fairly large bird, being approximately the
size of _Presbyornis_ cf. _pervetus_ and somewhat larger than the large
living burhinid _Esacus magnirostris_.
=Graculavus velox?=
Figure 9_d_
Referred Material.--Abraded right carpometacarpus consisting mainly of
the major metacarpal, NJSM 11854.
Locality and Horizon.--Collected from the main fossiliferous layer of
the Inversand Company marl pit, Sewell, Gloucester County, New Jersey;
Hornerstown Formation, latest Cretaceous (Maastrichtian); collected 25
February 1976 by David C. Parris.
Measurements (in mm).--Length 51.0.
Comparisons.--Nothing can be said about this very poor specimen except
that it came from a bird with a carpometacarpus slightly larger than
that of a modern specimen of the burhinid _Esacus magnirostris_.
Because _Graculavus velox_ is the only bird yet known in the New Jersey
fossil fauna that was of this same size, the present specimen may
possibly be referable to that species.
=Genus _Telmatornis_ Marsh, 1870=
Type-Species.--_Telmatornis priscus_ Marsh, 1870, by subsequent
designation (Hay, 1902:528).
Included Species.--Type species only.
=_Telmatornis priscus_ Marsh, 1870=
Figures 5_b-j_, 6_c,e,g_, 7_a,d,g,j,n_
_Telmatornis priscus_ Marsh, 1870:210.
_Telmatornis affinis_ Marsh, 1870:211.
_Graculavus pumilis_ Marsh, 1872:364.
_?Palaeotringa vetus_ Marsh, 1870:209.
Holotype.--Distal end of left humerus (Figure 5_e,h_), YPM 840;
collected in pits of the Cream Ridge Marl Company, near Hornerstown,
New Jersey by J.G. Meirs. Navesink Formation, Maastrichtian, Late
Cretaceous (Baird, 1967).
Referred Specimens.--Distal end of right humerus (Figure 5_f,g_),
YPM 845 (holotype of _Telmatornis affinis_ Marsh 1870); same data as
holotype of _T. priscus_.
Proximal end of right humerus (Figure 5_b-d_), YPM 850, with distal
end of right carpometacarpus (Figure 5_i_) and several fragments of
shafts of long bones apparently associated (holotypical material of
_Graculavus pumilis_ Marsh, 1872); collected near Hornerstown, New
Jersey, by J.G. Meirs; probably from the basal Hornerstown Formation,
Maastrichtian, Late Cretaceous.
Distal end of left tibiotarsus (Figure 7_n_), ANSP 13361 (holotype
of _Palaeotringa vetus_), collected near Arneytown, on the
Monmouth-Burlington county boundary, New Jersey; Basal Hornerstown
Formation, Maastrichtian, Late Cretaceous (Baird, 1967).
Left humerus lacking proximal end (Figure 6_c,e,g_), ANSP 15360;
collected in 1971 from the Inversand Company marl pit, Sewell,
Gloucester County, New Jersey, by Keith Madden. Basal Hornerstown
Formation, Maastrichtian, Late Cretaceous.
Distal end of left tarsometatarsus (Figure 7_d,g,j_), NJSM 11853;
collected 27 March 1975 by David C. Parris from the main fossiliferous
layer of the Inversand Company marl pit.
[Illustration: Figure 5.--Wing elements of _Burhinus_ and
_Telmatornis_. _a_, _Burhinus vermiculatus_ (USNM 488870), proximal end
of right humerus, anconal view, _b-d_, Telmatornis priscus (holotype
of _Graculavus pumilis_, YPM 850), proximal end of right humerus
(_b_, anconal view; _c_, palmar view; _d_, proximal view), _e,h_, _T.
priscus_ (holotype, YPM 840), distal end of left humerus (_e_, anconal
view; _h_, palmar view), _f,g_, _T. priscus_ (holotype of _Telmatornis
affinis_, YPM 845), distal end of right humerus (_f_, aconal view; _g_,
palmar view), _i_, _T. priscus_ (associated with YPM 850), distal end
of left carpometacarpus, dorsal view; _j_, _T. priscus_ (NJSM 11900),
proximal end of right ulna. (All figures x 2; specimens coated with
ammonium chloride to enhance detail.)]
[Illustration: Figure 6.--Humeri of _Anatalavis_, new genus, and
_Telmatornis_. _a_, _Anatalavis rex_ (holotype, YPM 902), right
humerus, palmar view; × 1.5. _b,d,f_, _A. rex_, (YPM 948), left
humerus (_b_, palmar view, × 1.5; _d_, enlarged, anconal view, × 2;
_f_, enlarged, palmar view, × 2). _c,e,g_, _Telmatornis priscus_,
(ANSP 15360), left humerus (_c_, palmar view, × 1.5; _e_, enlarged,
anconal view, × 2; _g_, enlarged, palmar view, × 2); _h_, _Burhinus
vermiculatus_ (USNM 430630), left humerus, palmar view, × 2. (Specimens
coated with ammonium chloride to enhance detail.)]
[Illustration: Figure 7.--Hindlimb elements. _a,b_, Right pedal
phalanx 1 of digit II (_a_, _Telmatornis priscus_, ANSP 15541; _b_,
_Presbyornis_ sp., USNM uncatalogued; part of associated foot), _c-k_,
Distal end of left tarsometatarsus, anterior, posterior, and distal
views, respectively (_c,f,i_, _Presbyornis_ sp., UCMP 126178; _d,g,j_,
_T. priscus_, NJSM 11853; _e,h,k_, _Burhinus vermiculalus_, USNM
488870). _l-n_, Distal portions of left tibiotarsi (_l_, _Palaeotringa
littoralis_, holotype, YPM 830; _m_, _P. vagans_, holotype, YPM 835;
_n_, _T. priscus_, holotype of _P. vetus_, ANSP 13361). (All figures ×
2; specimens coated with ammonium chloride to enhance detail.)]
Right pedal phalanx 1 of digit II (Figure 7_a_), ANSP 15541; collected
in 1972 by Richard White at the Inversand Company marl pit.
Proximal end of right ulna (Figure 5_j_), NJSM 11900; collected 14
July 1978 from spoil piles near junction of Routes 537 and 539, near
Hornerstown, Upper Freehold Township, Monmouth County, New Jersey, by
Gerard R. Case; presumably from the Hornerstown Formation but whether
from Cretaceous or Tertiary sediments is not known.
Miller (1955) lists an additional specimen from near Arneytown under
the name _Palaeotringa vetus_ (YPM 2808). This was cataloged in 1937
as “part of a tibia” of “Eocene” age but the specimen cannot now be
located in the Yale collections and its age and identity must be
considered very doubtful.
Measurements (in mm).--Distal ends of humeri (YPM 840, YPM 845, ANSP
15360, respectively): distal width 10.9, 10.1, 11.3; depth through
dorsal condyle 5.7, 5.2, 5.5; width of shaft at proximal extent of
brachial fossa 6.3, 5.5,6.4; length from distal end of pectoral crest
to ventral condyle (ANSP 15360 only) 45.1; shaft width at midpoint
(ANSP 15360 only) 4.7.
Proximal end of humerus YPM 850: proximal width through dorsal and
ventral tubercles 13.1; depth through bicipital surface and ventral
condyle 7.5, depth of head approximately 3.5.
Proximal end of ulna NJSM 11900: depth through dorsal cotyla 7.0.
Distal end of carpometacarpus YPM 840: depth at distal end 5.3; shaft
width 2.9.
Distal end of tibiotarsus ANSP 13361: shaft width 3.5, approximate
depth through medial condyle 6.9.
Distal end of tarsometatarsus NJSM 11853: distal width 6.1+; shaft
width 2.7.
Pedal phalanx 1 of digit II: length 14.6; proximal width 3.0.
Comparisons.--This is evidently the most abundant bird in the
New Jersey Cretaceous deposits. Hitherto it had been known only
from the two distal ends of humeri that are the holotypes of
_Telmatornis priscus_ and _T. affinis_. Marsh (1870) did not clearly
place _Telmatornis_ with any living family but mentioned species
of Rallidae, Scolopacidae, and Ardeidae in his comparisons. Hay
(1902:528) listed the genus under the Rallidae. Shufeldt (1915:26)
considered that _Telmatornis_ was not a heron but might be related
either to rail-like or charadriiform birds, the material, according
to him, being insufficient for positive determination. He (1915:27)
also described a larger species, _Telmatornis rex_, which we have
removed to a new genus. Lambrecht (1933:489) maintained _Telmatornis_
as a genus incertae sedis in his order Ralliformes. Brodkorb (1967)
placed the genus in the family Rallidae, subfamily Rallinae, without
comment. Cracraft (1972) established that Telmatornis did not belong
in the Rallidae but was instead very similar to the Burhinidae. He
synonymized _T. affinis_ with _T. priscus_ and created a new family,
Telmatornithidae, for _T. priscus_ and _T. rex_.
We concur in synonymizing _T. affinis_ with _T. priscus_. The holotypes
and the new specimen of humerus (ANSP 15360), which is instructive in
that it preserves much more of the shaft (Figure 6_c_), are indeed
very similar to the humeri in the Burhinidae. In size they are closely
comparable to the small living species _Burhinus vermiculatus_ (cf.
Figure 6_g,h_). The fossils differ from _Burhinus_ in having (1) the
shaft less curved, both in dorsal and in lateral views; (2) brachial
depression shorter, wider, and slightly more distally located; in
distal view (3) the ventral condyle smaller and less rounded; and (4)
the dorsal tricipital groove shallower.
The distal portion of the humerus of _Telmatornis_ is similar to that
in _Presbyornis_ but differs in having (1) the dorsal condyle decidedly
more elongate; (2) olecranal fossa much shallower; (3) ventral
epicondyle in ventral view less distinctly demarcated but (4) more
protrudent in lateral or medial view.
The proximal end of humerus (YPM 850) that is the holotype of
_Graculavus pumilis_ was considered by Shufeldt (1915:19) definitely
to be from a limicoline charadriiform. It is from a bird exactly the
size of _Telmatornis priscus_ and its coloration and preservation would
not be incompatible with its being the opposite end of the same bone
as the holotype of _T. affinis_ (Figure 5_b,c,f,g_). The following
differences between the holotypical humeri of _G. velox_ and _“G.”
pumilis_ establish that these belong to different genera: (1) in
_velox_ the area dorsal to the ventral tubercle and distal to the head
is much more excavated, undercutting the head; (2) the dorsal tubercle
is more pronounced; (3) there is a distinct excavation distomedial to
the ventral tubercle, lacking in _pumilis_; (4) the ventral tubercle in
ventral view is much more produced in _velox_ than in _pumilis_.
The holotype of _G. pumilis_ is very similar to the humerus in the
Burhinidae but differs from that family and agrees with _Graculavus_
in characters 8, 9, and 10 (p. 6). It differs further from the
Burhinidae in having the area for the attachment of M. scapulohumeralis
caudalis extending farther distally in ventral view. It differs from
_Presbyornis_ mainly in lacking the excavation to and undercutting
the head. Because pumilis is not congeneric with _Graculavus velox_
and because of its size and similarities with the Burhinidae and
_Presbyornis_, we have no hesitation about considering Graculavus
pumilis Marsh, 1872, to be a junior subjective synonym of _Telmatornis
priscus_ Marsh, 1870.
The proximal end of an ulna, NJSM 11900 (Figure 5_j_), is from a bird
the size of _Burhinus vermiculatus_ and not too dissimilar to it
except that the shaft is more robust in the fossil. The specimen is
too imperfect to merit detailed study and is referred to Telmatornis
priscus only on size and probability.
The very fragmentary distal end of carpometacarpus associated with the
type of _G. pumilis_ (Figure 5_i_) is slightly larger and more robust
than in _Burhinus vermiculatus_, but not so much as to be incompatible
with _T. priscus_. Compared to _Burhinus_ (1) the symphysial area
is deeper and (2) the articular surface for the major digit is
proportionately larger, the specimen being somewhat more similar to the
carpometacarpus in _Presbyornis_.
The three specimens of _Palaeotringa_ Marsh from the Cretaceous of New
Jersey are based on poorly preserved distal ends of tibiotarsi. The
holotype of _Palaeotringa vetus_ Marsh, 1870 (Figure 7_n_) is similar
in size to the comparable element in _Burhinus vermiculatus_, though
with a relatively more slender shaft, and hence is from a bird the
size of _T. priscus_, being smaller than any of the other species of
_Palaeotringa_. It is more similar to _Presbyornis_ than to _Burhinus_.
Because it is from a charadriiform the size of _T. priscus_, as first
revisers we tentatively consider _Palaeotringa vetus_ Marsh, 1870,
to be a subjective synonym of _Telmatornis priscus_ Marsh, 1870. The
only alternative would be to consign it to Aves incertae sedis. It is
of passing historical interest to recall Marsh's (1870:209) comment
that the type of _Palaeotringa vetus_ “apparently was the first fossil
bird-bone discovered in this country,” having been mentioned both by
Morton (1834) and Harlan (1835) as belonging to the genus _Scolopax_
(Charadriiformes: Scolopacidae).
The distal portion of tarsometatarsus NJSM 11853 (Figure 7_d,g,f_)
is unfortunately quite abraded. It is from a small charadriiform and
has a shaft width about the same as in _Burhinus vermiculatus_. If
this fossil came from an individual of _Telmatornis priscus_, as we
assume, _T. priscus_ being the smallest and most abundant “graculavid”
in the New Jersey Cretaceous deposits, then it is a very instructive
specimen, for it differs much more from Burhinus than does the humerus
of Telmatornis. NJSM 11853 differs from the Burhinidae and agrees with
_Presbyornis_ in having (1) the distal foramen proportionately large
and oval, not very small and circular; (2) a large, well-developed scar
for the hallux (hallux absent in Burhinidae); (3) external trochlea
proximodistally more elongate. That which remains of the inner trochlea
indicates that it was (1) somewhat more posteriorly retracted than
in _Burhinus_ but (2) not nearly as elevated and retracted as in
_Presbyornis_.
Pedal phalanx ANSP 15541 (Figure 7_a_) is from a bird the size of _T.
priscus_. This specimen is much longer and more slender than phalanx 1
of digit II in _Burhinus vermiculatus_ but has almost exactly the shape
and proportions of the same element in _Presbyornis_ (Figure 7_b_),
although being much smaller. Although its assignment to _Telmatornis_
is very tentative, the length of this element seems to indicate a
wading bird as opposed to one with the terrestrially adapted shorter
toes of the Burhinidae.
=Genus _Anatalavis_, new genus=
Type-Species.--Telmatornis rex Shufeldt, 1915.
Included Species.--Type-species only.
Diagnosis.--Differs from _Telmatornis_ and _Presbyornis_ in (1) having
the shaft very short, stout, and much more curved, both in dorsoventral
and lateromedial views. Differs from _Telmatornis_ and agrees with
_Presbyornis_ in (2) having the distal end in distal view deeper, with
(3) a narrower and much deeper olecranal fossa. Also, (4) the brachial
depression is smaller and narrower than in _Telmatornis_ but not as
deep, nor as proximally situated as in _Presbyornis_.
Etymology.--“Duck-winged bird,” from Latin _anas_, duck, _ala_, wing,
and _avis_, bird. The gender is feminine.
=_Anatalavis rex_ (Shufeldt, 1915), new combination=
Figure 6_a,b,d_J
Telmatornis rex Shufeldt, 1915:27, fig. 101.
Holotype.--Right humerus lacking proximal end, YPM 902 (Figure 6_a_).
Locality and Horizon.--From Hornerstown, Upper Freehold Township,
Monmouth County, New Jersey; collected by W. Ross in 1878; probably
Late Cretaceous (Maastrichtian), basal Hornerstown Formation.
Referred Specimen.--Paratypical left humerus lacking proximal end,
YPM 948 (Figure 6_b,d,f_). From Hornerstown, Upper Freehold Township,
Monmouth County, New Jersey; collected by J.G. Meirs in 1869; probably
Late Cretaceous (Maastrichtian), basal Hornerstown Formation.
Measurements (in mm).--Humeri (YPM 902, YPM 948, respectively): distal
width 13.6, 13.2; depth through dorsal condyle 7.3, 7.5; width of shaft
at proximal extent of brachial fossa 7.2,7.5; length from distal end of
pectoral crest to ventral condyle 49.1, 50.7; shaft width at midpoint
5.4, 5.6.
Remarks.--Shufeldt (1915:27) described this species in the same genus
as _T. priscus_ and _T. affinis_ but correctly noted that the humerus
“is a short one ... its sigmoid curve very pronounced.” Cracraft
(1972:41) considered that “except for its decidedly larger size, _T.
rex_ does not differ from _T. priscus_ in any significant features.”
In fairness to these authors, it should be noted that the great
differences between Anatalavis and Telmatornis are much more apparent
in comparisons with the new humerus of _T. priscus_ (ANSP 15360), which
preserves much more of the shaft than the previously known specimens.
Both Shufeldt and Cracraft considered YPM 948 to belong to the same
species as the holotype of _T. rex_, and we concur.
The specimens of _A. rex_ are not comparable with the type of
_Graculavus velox_, which was from a larger bird. _Anatalavis rex_
was a larger, heavier bird than _Telmatornis priscus_, with the
humerus remarkably short and robust, so that the overall length of the
humerus in _A. rex_ would scarcely have exceeded that of _T. priscus_.
_Anatalavis_ must have been a bird of considerably different flight
habits from _Telmatornis_ or _Presbyornis_. The overall appearance of
its humerus is in fact rather duck-like, except for the more expanded
distal end. It is still quite short and stout even for a duck.
=Genus _Laornis_ Marsh, 1870=
Type-Species.--_Laornis edvardsianus_ Marsh, 1870, by monotypy.
Included Species.--Type species only.
=_Laornis edvardsianus_ Marsh, 1870=
Figure 8_a,c,e_
_Laornis edvardsianus_ Marsh, 1870:206.
Holotype.--Distal end of right tibiotarsus, YPM 820.
Locality and Horizon.--From pits of the Pemberton Marl Company at
Birmingham, Burlington County, New Jersey; collected by J.C. Gaskill;
Late Cretaceous (Maastrichtian), basal Hornerstown Formation.
Measurements (in mm).--Distal end of tibiotarsus, YPM 820: distal width
across condyles 22.6, depth of external condyle 19.3, depth of internal
condyle 21.1, least shaft width 11.7, least shaft depth 9.6.
Comparisons.--The very large size of this specimen has undoubtedly
been a factor in misleading those who have attempted to identify
it, as it came from a bird the size of a swan or a large crane.
The affinities of this fossil have long been questioned and the
species has for most of its history been in limbo. Marsh (1870:207)
concluded only that _Laornis_ “shows a strong resemblance in several
respects to the _Lamellirostres_ [Anseriformes], and also to the
_Longipennes_ [Charadriiformes (Lari) and Procellariiformes], but
differs essentially from the typical forms of both of these groups.” In
its own nebulous way, this assessment is concordant with our placement
of _Laornis_ in a charadriiform group that was near the ancestry of
the Anseriformes. Doubtless only on the strength of Marsh's comments.
Cope (1869-1870:237) placed _Laornis_ in the “Lamellirostres.” Hay
(1902:531) included _Laornis_ in the Anatidae. Shufeldt (1915:23)
hardly clarified matters when he characterized _Laornis_ as “at least
one of the generalized types of waders,” being a “remarkable type,
which seems to have, judging from this piece of the tibiotarsus,
Turkey, Swan, Crane, and even other groups all combined in it.”
Lambrecht (1933:526) included _Laornis_ as a genus incertae sedis in
his “Telmatoformes,” between the Aramidae and Otididae.
The type was restudied by Cracraft (1973:46) who put _Laornis_ in the
Gruiformes and created a new family (Laornithidae) and superfamily
(Laornithoidea) for it. He included it in his suborder Ralli, the only
other member of which was the Rallidae. After preliminary comparisons,
Olson (1974) ventured that _Laornis_ belonged in the suborder Lari
of the Charadriiformes. Brodkorb (1978:214) listed _Laornis_ under
Aves incertae sedis and guessed that it might be related to the
Pelecaniformes.
Except for the extreme difference in size, the tibiotarsus of _Laornis_
is in many respects similar to that of _Presbyornis_ (Figure 8),
especially in (1) the shape and position of the tubercle proximal to
the external condyle; (2) the transverse pit in the intercondylar
sulcus; and (3) the broad, shallow intercondylar sulcus as seen in
distal view. It differs in a seemingly minor but quite characteristic
feature, the large nutrient foramen situated in the groove for M.
peroneus brevis (Figure 8_c_). This is absent in _Presbyornis_ but is
present in both of the tibiotarsi from the Cretaceous of New Jersey
in which that portion of the bone is preserved (the holotypes of
Palaeotringa littoralis and _P. vagans_), as well as in a tibiotarsus
(Science Museum of Minnesota P75.22.25) from the type-locality of
_Dakotornis cooperi_ Erickson, 1975, that may be referable to that
graculavid-like species. The foramen in the peroneus brevis groove
may also be found in at least some specimens of Stercorariidae, which
is partly what led Olson (1974) to suggest a relationship between
_Laornis_ and the Lari. _Laornis_ appears to have been an extremely
large member of the “transitional Charadriiformes,” though where its
relationships may lie within that group cannot be determined.
=Genus _Palaeotringa_ Marsh, 1870=
Type-Species.--_Palaeotringa littoralis_ Marsh, 1870; by subsequent
designation (Hay, 1902:527).
Included Species.--_Palaeotringa littoralis_ Marsh, 1870, and
_Palaeotringa vagans_ Marsh, 1872.
=_Palaeotringa littoralis_ Marsh, 1870=
Figure 7_l_
_Palaeotringa littoralis_ Marsh, 1870:208.
Holotype.--Distal portion of left tibiotarsus lacking most of the inner
condyle, YPM 830.
Locality and Horizon.--Collected in the “middle marl beds” by Nicolas
Wain from his marl pits near Hornerstown, New Jersey; Late Cretaceous
(Maastrichtian), either basal Hornerstown Formation or Navesink
Formation.
Measurements (in mm).--Depth through outer condyle 8.2; width of shaft
just proximal to outer condyle 7.0.
Comparisons.--This specimen and that of _P. vagans_ are too fragmentary
for useful comparison. Both have the foramen in the groove for
M. peroneus brevis, mentioned above. Their overall similarity to
_Presbyornis_ and to charadriiform birds in general justifies retaining
them with the other “graculavids” but other than this little else can
be said. In size, _Palaeotringa littoralis_ would have been about
equal to _Burhinus bistriatus vocifer_ and smaller than _Esacus
magnirostris_. Hence it would seem to be too small to belong to the
same species as _Graculavus velox_ and is definitely too large to be
referable to _Telmatornis priscus_.
[Illustration: Figure 8.--Distal end of right tibiotarsus of (_a,c,e_)
_Laornis edvardsianus_, holotype, YPM 820, compared with (_b,d,f_) the
same element enlarged in _Presbyornis_ sp., UW BQ305: _a,b_, anterior
views; _c,d_, lateral views (note large foramen in peroneus brevis
groove of _Laornis_); _e,f_, distal views. (_a,c,e_, × 1.5, _b,d,f_, ×
4; specimens coated with ammonium chloride to enhance detail.)]
=_Palaeotringa littoralis?_=
Figure 9_a_
Referred Material.--Distal portion of a left humerus, NJSM 11303.
Locality and Horizon.--Collected from the main fossiliferous layer of
the Inversand Company marl pit, Sewell, Gloucester County, New Jersey;
Hornerstown Formation, latest Cretaceous (Maastrichtian); collected 27
September 1972 by David C. Parris.
Measurements (in mm).--Distal width 12.8, depth through dorsal condyle
6.9, width of shaft at proximal extent of brachial fossa 8.2.