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

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 avifaima 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
Fiirbringer, 1888, with its provisional synonyms Palaeotringinae Wetmore, 1940; Telmatorni-
thidae Cracraft, 1972, and Laornithidae Cracraft, 1972. The species included are: Graculavus
velox Marsh, 1872; Telmatorius priscus Marsh, 1870 (synonyms: Telmatornis c^inis Marsh,
1870; Graculavus pumilus 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 PUBQCATIGN DATE is handstamped in a limited number of initial copies and is recorded in the Institution'
annual report, Smithsonian Year. SERIES COVER DESIGN: The trilobite Phacops rana Green.
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. IH. Series.
QE701.S56 no. 63 560 s 86-29837 [QE871] [568'.09749]

Contents
Page
Introduction	1
Acknowledgments	1
The Fossil Localities and Their Stratigraphy	1
Order CHARADRIIFORMES	4
"Form Family" GRACULAVIDAE Fiirbringer, 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
111



The Cretaceous Birds of New Jersey
Storrs L. Olson and David C. Parris

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 Town-
ship, 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
Storrs L. Olson. Department of Vertebrate Zoology, National Museum of Natural
History, Smittisonian Institution, Washington, D.C. 20560. David C. Parris, New
Jersey State Museum. 205 West State Street, Trenton, New Jersey 08625-0530.

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
Chtirchill 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 valu-
able 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 fossiiiferous.
  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 explora-
  
SMITHSONIAN CONTKIBUTIONS TO PALEOBIOLOGY


Cretaceous rocks known or inferred
to be present beneatli surface
concealed by post - Cretaceous
Cretaceous rocks (predominantly
sedinnentary) at surface


I-'IGI;RK 1.—Distribution of Oclaccou.s rocks in the eastern United States. Arrow indicates New Jersey. (Mcidified after
Moore, 1958. fig. 15.2).

tion has encouraged more recent restudy of the subsurface
suatigraphy, notably along the cast coast (Minard ct al., 1974;
Perry et al., 1975; Petters, 1976).
  In New Jersey, the latest Cretaceous deposits arc 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 compclitive 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 di.slurbancc 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
u-ansgression with the shoreline not i-ar 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 mari pit owner J.G. Mcirs. 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 (WeLmorc, 1930), this
was later definitely established as Cretaceous by Baird (1967),
who atu-ibuted 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)
di.scussion is consistent with Pellers's view that the Horners-
town Formation is regarded as piu-lly 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 geographi-
cally distinct localities in New Jersey (Figure 2). With the
exception oi Laornis, all ofthe specimens described by Marsh
(1870, 1872) came from Upper Freehold Township, Monmouth
  
NUMBER 63


FIGURE 2.—^Localities in southern New Jersey of the main fossiiiferous deposits
that have yielded Cretaceous birds. (The bold line demarcates the inner and outer
coastal plain physiographic provinces; B = Birmingham; H = Hornerstown; S =
Sewell.)
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 die holotype of Palaeotringa
vetus, from "friable green marl near Arneytown" was from the
lower (i.e.. Cretaceous) part of die 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.
  The second general locality is near Birmingham, Burlington
County, where the type oi 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 nodiing to be added to Baird's (1967) conclusion that
this specimen is latest Cretaceous in age.
FIGURE 3.—Stratigraphic diagram of the Inversand Company marl pit at Sewell,
Gloucester County, New Jersey.
  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, aldiough this information is
preserved in records sufficient in number and distribution to
assm^e that it will not be lost The Inversand specimens came
from the main fossiiiferous layer within the basal portion of
the Hornerstown Formation (Figure 3). This layer is of late
MaasU-ichuan 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


SMITHSONIAN CONTRBUTIONS TO PALEOBIOLOGY

           Order CHARADRIIFORMES
"Form Family" GRACULAVIDAE Fiirbringer, 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 die New Jersey deposits
belong wiUi what Olson (1985) has termed die "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. Odier apparent descendants include ibises
(Plataleidae) and the ducks and geese of the order Anseri-
formes. The latter are linked widi the "transitional Charadrii-
formes" through the Paleocene and Eocene genus Presbyornis.
which is known from abundant material from widely scattered
areas of Uie 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 dian to any modern group of birds
except the Btu-hinidae. 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 simdar 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 widi the "transitional Charadriiformes"
for which family-group names are available include: Gracula-
vinae Fiirbringer, 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 die same category (Olson and
Feduccia, 1980a) include: Apatornithidae Furbringer, 1888;
Cimolopterygidae Brodkorb, 1963a; Torotigidae Brodkorb,
1963a; and Lonchodytidae Brodkorb, 1963a.
  Tertiary taxa that may possibly be related to die "transitional
Charadriiformes" and diat have been used as the basis of
family-group names are: Presbyornithidae Wetmore, 1926
(Nautilornidiinae Weunore, 1926, and Telmabatidae Howard,
1955, are definitely synonyms); Scaniornidiidae Lambrecht,
1933; and Dakotornidiidae Erickson, 1975.
  
Doubtiess diere are odiers that we have overlooked. How
many families are actually represented here and what dieir
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 die case of die Cretaceous birds under consideration here,
we have decided for the time being to adopt a version of
paleobotanical convention in recognizing a "form famdy"
Graculavidae, which implies a general similarity in morphol-
ogy of the constituent taxa, aldiough the material available is
simply not sufficient for determining phylogeny or key
adaptations.
Genus Graculavus Marsh, 1872
Limasavis Shufeldt, 1915:19.
  TYPE-SPEQES.—Graculavus velox Marsh 1872, by subse-
quent 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 ve/ox Marsh, 1872
FIGURE Ab,df,h
Graculavus ve/ox Marsh, 1872:363.
Limosavis velox (Marsh).—Lambrecht, 1933:546.
HOLOTYPE.—Proximal end of left humerus, YPM 855.
  LocALriY AND HoRizoN.—From Hornerstown, Upper Free-
hold Township, Monmouth County, New Jersey; collected by
J.G. Meirs; Late Cretaceous (Maastrichtian), eidier basal
Hornerstown Formation or Navesink Formation.
  MEASLTREMENTS (in mm).—Proximal end of humerus, YPM
855: proximal widdi through dorsal and ventral tubercles 21.1,
depth through bicipital surface and tuberculum ventrale 11.6,
depth of head 5.7.
  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 similar-
FlGURE 4.—Proximal ends of left humeri of Graculavus velox and related birds: a.
Esacus magnirostris (Burhinidae), USNM 19649; b4f,K 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 x 2; specimens coated with ammonium chloride to enhance detail.

NUMBER 63


SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

ities to the Burhinidae, Haematopodidae, and Charadriidae.
Accordingly, Lambrecht (1933:540, 546) placed these taxa
among die charadriiform birds, but rather inexplicably listed
velox under Shufeldt's substitute name Limosavis in die
suborder Laro-Limicolae, while rexaining 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 relation-
ships 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
distinctiy 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,
distinctiy pointed. In most of the preceding characters that it
preserves, the single proximal end of humerus referred to
Telmatornis (the holotype of G. pumilus) agrees widi
Graculavus and Presbyornis.
  Among living famdies, die 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 die head more deeply undercut.
Presbyornis is intermediate between Graculavus and the
Burhinidae in character 9.
   Graculavus velox was a fairly large bird, being approxi-
mately the size of Presbyornis cf. pervetus and somewhat
larger than the large living burhinid Esacus magnirostris.
Graculavus velox?
FIGURE 9d
REFERRED MATERIAL.—Abraded right carpometacarpus con-

sisting mainly ofthe major metacarpal, NJSM 11854.
  LOCALITY AND HORIZON.—Collected from the main fos-
siiiferous 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 widi a carpometa-
carpus slightiy larger than that of a modern specimen of the
burhinid Esacus magnirostris. Because Graculavus velox is the
only bird yet known in die 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 subse-
quent designation (Hay, 1902:528).
INCLUDED SPECIES.—^Type species only.
Telmatornis priscus Marsh, 1870
FIGURES 5b-j. 6c.e,g. la,d,gj.n
Telmatornis priscus Marsh, 1870:210.
Telmatornis affinis Marsh, 1870:211.
Graculavus pumilis Marsh, 1872:364.
?Palaeotringa vetus Mars^, 1870:209.
  HOLOTYPE.—Distal end of left humerus (Figure 5e.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/^), YPM 845 (holotype of Telmatornis affinis Marsh 1870);
same data as holotype of T. priscus.
  Proximal end of right humerus (Figure 5b-d), YPM 850,
with distal end of right carpometacarpus (Figure 5/) and several
fragments of shafts of long bones apparentiy associated
(holotypical material of Graculavus pumilis Marsh, 1872);
collected near Hornerstown, New Jersey, by J.G. Meirs;
probably from the basal Hornerstown Formation, Maastrich-
tian, Late Cretaceous.
  Distal end of left tibiotarsus (Figure In), 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 6c,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 Id.gj), NJSM
11853; collected 27 March 1975 by David C. Parris from die
main fossiiiferous layer of the Inversand Company marl pit.
  

NUMBER 63

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 pumilus, 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), fg, 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.)

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

I-IGL'RE 6.—Humeri of Anatalavis, new genus, and TelrruiU-<rms. a, Anatalavis rex (holotype, YPM 902), right humerus,
palmar view; x 1.5. b,d,f, A. rex, (YPM 948), left humerus {h. palmar view, x 1.5; ci, enlarged, anconal view, x 2;/,
enlarged, palmar view, x 2). c.e.g, Telmatornis priscus, (ANSP 15360), left humerus (c, palmar view, x 1,5; e, enhu-gcd,
anconal view, x 2; g, enlarged, palmar view, x 2); h Burhinus vermiculatus (USNM 430630), left humerus, palmar
view, X 2. (Specimens coaled with ammonium chloride lo enhance detail.)


NUMBER 63

FIGURE 7.—Hindlimb elements. a,b, Right pedal phalanx I of digit 11 {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 (cf i, Presbyornis sp., UCMP 126178; d, g.j.T priscus, NJSM 11853; e. h k, Burhinus
vermiculatus, USNM 488870). l-n, Distal portions of left tibiotarsi (/, Palaeotringa littoralis. holotype, YPM 830; m,
P. vagans, holotype, YPM 835; n, T. priscus, holotype of P. vetus. ANSP 13361). (All figures X 2; specimens coated
with ammonium chloride to enhance detail.

10

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY



  Right pedal phalanx 1 of digit II (Figure 7a), ANSP 15541;
collected in 1972 by Richard White at the Inversand Company
marl piL
  Proximal end of right ulna (Figure 5;), NJSM 11900;
collected 14 July 1978 from spod piles near junction of Routes
537 and 539, near Hornerstown, Upper Freehold Township,
Monmouth County, New Jersey, by Gerard R. Case; presum-
ably from die Hornerstown Formation but whedier from
Cretaceous or Tertiary sediments is not known.
  Miller (1955) lists an additional specimen from near
Arneytown under die name Palaeotringa vetus (YPM 2808).
This was cataloged in 1937 as "part of a tibia" of "Eocene"
age but die 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; depdi dirough 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 widdi at midpoint (ANSP 15360 only)
4.7.
  Proximal end of humerus YPM 850: proximal widdi through
dorsal and ventral tubercles 13.1; depdi through bicipital
surface and ventral condyle 7.5, depth of head approximately
3.5.
  Proximal end of ulna NJSM 11900: depdi through dorsal
cotyla 7.0.
  Distal end of carpometacarpus YPM 840: depth at distal end
5.3; shaft widdi 2.9.
  Distal end of tibiotarsus ANSP 13361: shaft widdi 3.5,
approximate depth through medial condyle 6.9.
  Distal end of tarsometatarsus NJSM 11853: distal width
6.1-I-; shaft width 2.7.
Pedal phalanx 1 of digit II: lengdi 14.6; proximal width 3.0.
  COMPARISONS.—This is evidendy the most abundant bird
in the New Jersey Cretaceous deposits. Hidierto 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) Usted the genus under die
Rallidae. Shufeldt (1915:26) considered that Telmatornis was
not a heron but might be related either to rad-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 die genus in the family Rallidae,
subfamily Rallinae, without comment. Cracraft (1972) estab-
lished that Telmatornis did not belong in the Rallidae but was
instead very similar to the Burhinidae. He synonymized T.

affinis widi T. priscus and created a new family, Telmatorni-
thidae, 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 diat it preserves much more of die shaft
(Figure 6c), are indeed very simdar to the humeri in the
Burhinidae. In size diey are closely comparable to the small
living species Burhinus vermiculatus (cf. Figure 6g,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 slightiy 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 die humerus of Telmatornis is similar
to diat 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 distinctiy
demarcated but (4) more protrudent in lateral or medial view.
  The proximal end of humerus (YPM 850) diat is die 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 die
opposite end of the same bone as the holotype of T. affinis
(Figure 5b,cf,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 die 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 simdar to the humerus
in the Burhinidae but differs from that family and agrees widi
Graculavus in characters 8, 9, and 10 (p. 6). It differs further
from the Burhinidae in having the area for die attachment of
M. scapulohumeralis caudaUs 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 widi 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;), is
from a bird the size of Burhinus vermiculatus and not too
dissimilar to it except diat 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 5i) is slightiy
larger and more robust than in Burhinus vermiculatus, but not
  
NUMBER 63

11



so much as to be incompatible widi T. priscus. Compared to
Burhinus (1) die symphysial area is deeper and (2) the articular
surface for die 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 In) is similar in size to the comparable element
in Burhinus vermiculatus, though with a relatively more
slender shaft, and hence is from a bird die size of T. priscus,
being smaller tiian any of the odier 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 die type of Palaeotringa vetus
"apparendy 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 (Charadrii-
formes: Scolopacidae).
  The distal portion of tarsometatarsus NJSM 11853 (Figure
Td.g.j) is unfortunately quite abraded. It is from a small
charadriiform and has a shaft widdi about die 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 die New Jersey
Cretaceous deposits, then it is a very instructive specimen, for
it differs much more from Burhinus dian does the humerus of
Telmatornis. NJSM 11853 differs from die 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 die 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 red-acted as in
Presbyornis.
  Pedal phalanx ANSP 15541 (Figure 7a) is from a bird the
size of T priscus. This specimen is much longer and more
slender dian phalanx 1 of digit II in Burhinus vermiculatus but
has aknost exactiy the shape and proportions of die same
element in Presbyornis (Figure lb), although being much
smaller. Aldiough its assignment to Telmatornis is very
tentative, die lengdi of this element seems to indicate a wading
bird as opposed to one widi the terrestrially adapted shorter
toes of the Burhinidae.
Genus Anatalavis, new genus
TYPE-SPEQES.—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, widi (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 6a,b,df
Telmatornis rex Shufeldt, 1915:27, fig. 101.
  HOLOTYPE.—Right humerus lacking proximal end, YPM
902 (Figure 6a).
  LocALFTY AND HoRizoN.—From Hornerstown, Upper Free-
hold Township, Monmoudi County, New Jersey; collected by
W. Ross in 1878; probably Late Cretaceous (Maastrichtian),
basal Hornerstown Formation.
  REFERRED SPEQMEN.—Paratypical left humerus lacking
proximal end, YPM 948 (Figure 6b,df). From Hornerstown,
Upper Freehold Township, Monmouth County, New Jersey;
coUected by J.G. Meirs in 1869; probably Late Cretaceous
(Maastrichtian), basal Hornerstown Formation.
  MEASUREMENTS (in mm).—Humeri (YPM 902, YPM 948,
respectively): distal widdi 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 widdi 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 diat "except for
its decidedly larger size, T. rex does not differ from T priscus
in any significant features." In fairness to these audiors, 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 diat die 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
radier duck-like, except for the more expanded distal end. It is
still quite short and stout even for a duck.
  
12

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY



Genus Laornis Marsh, 1870
TYPE-SPEQES.—Laornis edvardsianus Marsh, 1870, by
monotypy.
INCLUDED SPECIES.—^Type species only.
Laornis edvardsianus Marsh, 1870
FIGURE 8a,c,e
Laornis edvardsianus Marsh, 1870:206.
HOLOTYPE.—Distal end of right tibiotarsus, YPM 820.
  LocALFTY AND HoRizoN.—^From pits of the Pemberton Marl
Company at Birmingham, Burhngton County, New Jersey;
collected by J.C. GaskiU; Late Cretaceous (Maastiichtian),
basal Hornerstown Formation.
  MEASUREMENTS (in mm).—Distal end of tibiotarsus, YPM
820: distal width across condyles 22.6, depdi of external
condyle 19.3, depth of internal condyle 21.1, least shaft widdi
11.7, least shaft depdi 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 thai Laornis
"shows a sti^ong 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 bodi of these groups." In
its own nebulous way, diis assessment is concordant widi our
placement of Laornis in a charadriiform group diat was near
the ancestry of the Anseriformes. Doubtiess only on the
sti-ength of Marsh's comments. Cope (1869-1870:237) placed
Laornis in the "Lamellirosfres." 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 tibio-tarsus,
Turkey, Swan, Crane, and even odier groups all combined in
iL" Lambrecht (1933:526) included Laomw as a genus incertae
sedis in his "Telmatoformes," between die Aramidae and
Otididae.
  The type was restudied by Cracraft (1973:46) who put
Laornis in the Gruiformes and created a new family
(Laornidiidae) 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) l\s\&ALaornis under
Aves incertae sedis and guessed that it might be related to the
Pelecaniformes.
Except for the extreme difference in size, die 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
nutiient foramen situated in the groove for M. peroneus brevis
(Figure 8c). This is absent in Presbyornis but is present in both
of die tibiotarsi from die 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 diat graculavid-like species. The foramen
in the peroneus brevis groove may also be found in at least
some specimens of Stercorariidae, which is partiy what led
Olson (1974) to suggest a relationship between Laornis and
the Lari. Laornis appears to have been an extremely large
member of die "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,
2iRd Palaeotringa vagans Marsh, 1872.
Palaeotringa littoralis Marsh, 1870
FIGURE 11
Palaeotringa littoralis Marsh, 1870:208.
  HOLOTYPE.—Distal portion of left tibiotarsus lacking most
of the inner condyle, YPM 830.
  LocALFTY AND HoRizoN.—CoUectcd in the "middle marl
beds" by Nicolas Wain from his marl pits near Hornerstown,
New Jersey; Late Cretaceous (Maasu-ichtian), 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. Bodi have die 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 diis Uttie 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 loo large to be referable to
Telmatornis priscus.
  
NUMBER 63

13


FIGURE 8.—Distal end of right tibiotarsus of {a,c,e) Laornis edvardsianus, holotype, YPM 820, compared with {b,df)
the same element enlarged in Presbyornis sp., tJW BQ305: a,b, anterior views; c,d, lateral views (note large foramen
in peroneus brevis groove of Laornis); ef, distal views. {a,c,e, x 1.5, b,d/, x 4; specimens coated with ammonium
chloride to enhance detail.)

14

SMrniSONL\N CONTRIBUTIONS TO PALEOBIOLOGY



Palaeotringa littoralis?
FIGURE 9a
  REFERRED MATERIAL.—Distal portion of a left humerus,
NJSM 11303.
  LOCALITY AND HORIZON.—Collected from die main fos-
sdiferous layer of the Inversand Company marl pit, Sewell,
Gloucester County, New Jersey; Hornerstown Formation, latest
Cretaceous (Maasfrichtian); collected 27 September 1972 by
David C. Parris.
  MEASUREMENTS (in mm).—Distal width 12.8, depth dirough
dorsal condyle 6.9, widdi of shaft at proximal extent of brachial
fossa 8.2.
  COMPARISONS.—This interesting specimen, although consid-
erably worn, clearly has the overall "graculavid" morphology
but shows sufficient differences from the humeri of Telma-
tornis or Anatalavis to wairant its generic separation from
them. In size it is about equal to die modern form Burhinus
bistriatus vocifer and hence would be compatible with P.
littoralis. It differs from Telmatornis, Anatalavis, or Pres-
byornis, and is more simdar to Burhinus in having (1) the
brachial depression wider, shallower, and more proximally
situated. Aldiough affected by wear, (2) the dorsal condyle is
nevertheless considerably smaller and not produced as far
proximally as in any of the preceding genera, although
Presbyornis is more similar in this respect than the odiers. In
distal view the specimen is more similar to Presbyornis than
to die odier Cretaceous humeri, aldiough (3) the olecranal fossa
is shallower. If this specimen is correctiy referred to
Palaeotringa, it shows diat genus to be distinct from any of
the others yet known in the fauna except possibly Graculavus,
for which the distal end of the humerus is unknown.
Palaeotringa vagans Marsh, 1872
                              FIGURE Im
Palaeotringa vagansMditsh, 1872:365.
  HOLOTYPE.—Fragmented distal two-diirds of a left tibio-
tarsus lacking the external condyle and the anterior portion of
the internal condyle, YPM 835.
  LOCALITY AND HORIZON.—From Hornerstown, Upper Free-
hold Township, Monmoudi County, New Jersey; collected by
J.G. Meirs; Late Cretaceous (Maasfrichtian), "about ten feet
below the surface of die marl" (Marsh, 1872:365), either basal
Hornerstown Formation or Navesink Formation.
  MEASUREMENTS (in mm).—Width of shaft just proximal to
external condyle 5.8.
  COMPARISONS.—This very unsatisfactory specimen comes
from a species smaller than P. littoralis and larger dian P.
vetus (= Telmatornis priscus). It differs from the latter and
agrees widi P. littoralis in having die distal tendinal opening
of a flattened oval shape, rather dian decidedly rounded. If we

have correctiy referred P. vetus to Telmatornis priscus, then it
is certain that neidier of the other two species of Palaeotringa
can be referred to Telmatornis. In P. vagans the tendinal groove
appears to be much narrower and die bridge much deeper than
in P. littoralis, but this is in part due to damage and possible
immaturity in the latter specimen, so it remains possible that
these species are in fact congeneric. The species P. vagans can
be retained as it is smaller than any of the other graculavids
in the fauna except T. priscus, from which it is generically
distinct
Graculavidae, Genus and Species Indeterminate
FIGURE 9b,c
  REFERRED MATERIAL.—Abraded distal end of left humerus
and associated proximal portion of humeral shaft, proximal
end of radius, and fragment of shaft of ulna, NJSM 11302.
  LocALnY AND HoRizoN.—Collected from the main fos-
sdiferous layer of the Inversand Company marl pit, Sewell,
Gloucester County, New Jersey; Hornerstown Formation, latest
Cretaceous (Maastrichtian); collected 15 August 1972 by
David C. Parris.
  MEASUREMENTS (in mm).—Humerus: distal widdi 19 mm,
depth through dorsal condyle 9.7, width of shaft at proximal
extent of brachial fossa 11.0; greatest proximal diameter of
radius 7.0.
  COMPARISONS.—The distal end of the humerus is the only
reasonably diagnostic element in this assortment and indicates
a large, robust species that would have exceeded in size any
of the others known in this Cretaceous avifauna except Laornis
edvardsianus, which was much larger stiU. In size this bird
would have approximated the modern flamingo Phoeniconaias
minor, which it somewhat resembles in morphology as well.
The humerus is not greatiy different from diat of other
Graculavidae in general aspect but is distinct in having a larger,
much deeper, and more proximally situated brachial depres-
sion. It represents a species distinct from any of the others yet
known in the fauna and is certainly generically distinct from
all except possibly Graculavus, for which comparable elements
are unknown.
Order FROCELLARm^oRMES?
  Among die newly collected material from the Inversand pit
is a singular avian humerus that cannot be assigned to the
Graculavidae or to any odier known family, fossil or modern.
Although it is generally inadvisable to name even Paleogene
birds on single elements, to say nothing of Cretaceous ones,
the specimen under consideration here is superior to any of the
other avian fossils yet collected from the Cretaceous of New
Jersey, both in preservation and in diagnostic qualities, and it
would seem incongruous to leave it innominate when
practically all the other fragments from the same deposits have
received names.
  

FIGURE 9.—.Miscellaneous elements, a, Palaeotringa littoralis? (NJSM 11303), distal end of left humerus, palmar view;
b, Graculavidae, genus and species indeterminate (NJS.M 11302), distal end of left humerus, palmar view; c, proximal
end of radius associated with b; d Graculavus veloxl (NJSM 11854), right carpometacarpus; ef Procellariiformes?,
genus and species indeterminate (ANSP 157l3),distal end of left ulna {e, external view;/ dorsal view); g, Aves, incertae
sedis (NJSM 12119), distal end of left femur, posterior view. {a,b,c.d x2; e.f g. x5; specimens coated with ammonium
chloride to enhance detail.)

16

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY



  The most distinctive features of this specimen are the deep
brachial depression and die incipient ectepicondylar spur, thus
calling to mind botii die Lari (Charadriiformes) and the
Procellariiformes among modern birds. Among die Pelecani-
formes it also bears a resemblance to the Phaethontidae and
especially to die Eocene frigatebird Limnofregata (Fregatidae)
(Olson, 1977).
Family TYTTHOSTONYCHIDAE, new family
TYPE GENUS.—Tytthostonyx, new genus.
INCLUDED GENERA.—Type genus only.
  DIAGNOSIS.—Differs from the Lari and odier Charadri-
iformes in (1) the low, narrow head; (2) the very large, long
pectoral crest; (3) the virtual absence of the incisura capitis or
any excavation for M. coracobrachialis cranialis; and (4) the
shallow, indistinct tricipital grooves. It agrees with the
Procellariiformes and differs from Phaethon and Limnofregata
in characters 2 and 4, and in the large, deeply excavated
brachial depression. The ectepicondylar spur is better devel-
oped than in any of die Pelecaniformes but not as well
developed as in the Procellariiformes. The apparendy very
broad pectoral crest extends much farther distally than in any
of the Procellariiformes or even in Limnofregata, to which the
fossil is somewhat more similar in this respect. Tytthostonyx
differs from any of die taxa compared in having the ventral
condyle very rounded, extending distally well past the dorsal
condyle.
Genus Tytthostonyx, new genus
TYPE-SPECIES.—Tytthostonyx glauconiticus, new species.
INCLUDED SPECIES.—Type species only.
DIAGNOSIS.—As for die family.
  ETYMOLOGY.—Greek, tytthos, little, plus stonyx, any sharp
point. The name is masculine in gender and refers to the small,
presumably rudimentary, ectepicondylar spur. It should not be
confused with the coleopteran genus Tytthonyx, based on onyx,
claw.
Tytthostonyx glauconiticus, new species
FIGURES 10, II
  HOLOTYPE.—Right humerus lacking die venu-al tubercle,
portions of die pectoral crest, and odier parts of the proximal
end, where partially reconsfructed, NJSM 11341.
  LocALm' AND HORIZON.—Main fossiiiferous layer of die
Inversand Company marl pit, Sewell, Gloucester County, New
Jersey; basal portion of the Hornerstown Formation, latest
Cretaceous (Maastrichtian); collected 11 October 1973 by
David C. Parris.
MEASUREMENTS OF HOLOTYPE (in mm).—Length as recon-

stincted, UO; widdi and depdi of shaft at midpoint 7.0 x 5.6;
distal widdi 14.8; depth through dorsal condyle 8.7.
  ETYMOLOGY.—^From Latin, glaucus (Greek, glaukos), bluish
green or gray, sea-colored, applied to greensands because of
their color, although appropriate because of their marine
origins as well; in reference to the holotype having been
recovered from beds of glauconite.
  REMARKS.—A possible relationship between die Procel-
lariiformes and Pelecaniformes has been previously suggested
(Sibley and Ahlquist, 1972:70; Olson, 1985:142), and among
the pelecaniform taxa most often mentioned as being
proceUariiform-hke are die Fregatidae. It is tempting to regard
the humerus of Tytthostonyx as being similar to that possessed
by die ancesfral stock that gave rise to the Procellariiformes.
Its similarities also to die Eocene frigatebird Limnofregata
would dius be seen as corroborating the primitiveness of die
Fregatidae within die Pelecaniformes. Whereas Tytthostonyx
definitely has not achieved the highly distinctive and
presumably derived morphology of the humerus of modern
Procellariiformes, the incipient development of the ectepicon-
dylar spur and deep brachial depression could be interpreted
as tending in that direction.
  On the other hand, we must admit diat we are dealing with
only a single bone and one of very great age at that, so that die
risk of overinterpreting the fossil is correspondingly great.
We can only discern the overall similarities of the specimen
and phylogenetic inferences can therefore be only tentative
at best.
Family and Genus Indeterminate
FIGURE 9ef
  REFERRED MATERIAL.—Distal portion of left ulna ANSP
15713.
  LOCALITY AND HORIZON.—Inversand Company marl pit,
near Sewell, Gloucester County, New Jersey; Hornerstown
Formation, Late Cretaceous (Maastrichtian); not found in situ,
collected on shelf formed by "blue bed"; collected 31 August
1977 by Richard S. White.
  MEASUREMENTS (in mm).—Distal width 2.6, distal depdi
3.1, widdi and depdi of shaft near point of break 1.8 x 1.9.
  COMPARISONS.—This specimen comes from a very small
bird. The only modern pelagic birds in this size range are the
storm-petrels of the family Oceanitidae and the fossil resembles
this family in the extremely sfraight shaft of the ulna, the shape
and depdi of die tendinal grooves, and die relatively
well-developed scars for die attachment of the secondaries. It
differs from the Oceanitidae in having the venfral lip of the
external condyle much more rounded and profrudent past die
plane of the shaft, whereas the carpal tubercle in dorsal view
is markedly smaller. On diis basis, die fossd certainly could
not be referred to the Oceanitidae and that it should be
associated widi die Procellariiformes may be doubted as well.
  
NUMBER 63

17


FIGURE 10.—Tytthostonyx glauconiticus, new genus and species (holotype, NJSM 11341), right humerus: a,b, anconal
and palmar views of uncoated specimen to show reconstructed areas, x 0.8; cd, stereophotographs of coated specimen
in anconal and palmar views, x 1.3.

18

sMrrasoNiAN CONTRIBUTIONS TO PALEOBIOLOGY


FIGURE 11.—Tytthostonyx glauconiticus, new genus and species (holotype, NJSM 11341), stereophotographs of distal
end of right humerus: a, anconal view; b, palmar view; c, ventral view; d, dorsal view; e, distal view. (All figures x 2;
specimens coated with ammonium chloride to enhance detail.)

NUMBER 63

19



Aves, incertae sedis
FIGURE 9^
  REFERRED MATERLU..—Distal end of left femur, NJSM
12119.
  LOCALITY AND HORIZON.—^Inversand Company marl pit,
Sewell, Gloucester County, New Jersey; from processed spoil
piles, precise sfratum unknown; collected 12 December 1981
by Cyndiia Miller. Presumably from die Hornerstown Forma-
tion but could be eidier Late Cretaceous or Paleocene.
MEASUREMENTS (in mm).—^Distal width 4.3, distal depdi 3.8.
  COMPARISONS.—^This is also from a very small bird, possibly
the same size as die species represented by the preceding ulna
(ANSP 15713; Figure 9ef) but probably somewhat larger. It
is characterized by an extremely well-developed tubercle for
the attachment of M. gastrocnemius lateralis. A perfunctory
perusal of modern taxa revealed nothing similar.
Discussion
  Because the specimens freated here are avian and of
Mesozoic age, it is almost certain that too much importance
will be made of them by some future audiors. Indeed, it will
probably be years before the literature can be expunged of the
records of presumed occurrences that arose from previous
misidentifications of these fossds. Therefore, in an effort to
forestall overendiusiasm for these fragments we shall present
our own brief assessment of their significance.
  Unlike most other Cretaceous birds, such as the Hesperorni-
thiformes, Ichdiyornidiiformes, and Enantiornithiformes, which
represent totally extinct lineages (Olson, 1985), the Cretaceous
birds of New Jersey are of essentially modern aspect. However,
there are no modern families of birds represented in the fauna.
The differences among the fossds suggest that at least two
orders are represented, but whether any or all of the species
can be placed in modern orders is more difficult to say. This
stems as much from the unsatisfactory state of the ordinal
classification of modern birds (Olson, 1981,1985), as from die

incompleteness of the fossils. There are certain modern birds,
for example die Burhinidae, with sufficient similarities to
some of the Cretaceous fossils that there would be no problem
with associating them in the same ordinal-level taxon, though
it would be more difficult to say which other modern families
should also be included.
  The material is too poor to state how many famdies are
represented in the fauna, although if the various members of
the "form-famdy" Graculavidae were better known there can
scarcely be any doubt diat more dian one family would be
recognized in this group. Widiin the Graculavidae from New
Jersey there appear to be six genera {Graculavus, Telmatornis,
Palaeotringa, Laornis. Anatalavis, and an unnamed genus).
These are diverse, ranging in size from the smaUest of the
modern Burhinidae to that of a large crane. The very short,
robust, curved humeri of Anatalavis indicate some diversity
in mode of flight as wed. The greatest similarity of most of
these forms is to the early Paleogene bird Presbyornis, and
then to the modern family Burhinidae. Because these two
groups are very different in their habits and feeding adaptations
we may expect that die various members of the Graculavidae
were probably as divergent from one anodier as are Presbyornis
and Burhinus, their similarities being almost certainly due to
the retention of primitive characters.
  Including the two genera and species diat show some
simdarities to the Procellariiformes, along with die small
indeterminate femur, the total avifauna from die New Jersey
greensands comprises 8 or 9 genera and 9 or 10 species. As far
as can be determined, all of the bfrds in this assemblage were
probably marine or littoral in habits. We certainly would not
interpret this as indicating that waterbirds are primitive and
that they gave rise to land birds, as suggested by Thulborn and
Hamley (1985) in dieir fantastic and highly improbable
conjectures as to the mode of life of Archaeopteryx. Indeed,
just the opposite is probably die case (Olson, 1985), the lack
of Late Cretaceous fossils of fruly terrestrial or arboreal birds
most likely being due to sampling bias.
  
Appendix
The nonavian megafauna of the main fossiiiferous layer (Basal Hornerstown Formation),
at the Inversand Company marl pit, Sewell, Gloucester County, New Jersey is listed
below. Also found in die deposits were numerous coprolites of sharks and crocodilians,
some amber, phosphatized wood, and a few seeds. Voucher specimens are in the
collections of the New Jersey State Museum, Academy of Natural Sciences of
Philadelphia, and Yale University (Princeton University collections).

BRACmOPODA
Terebratulina atlantica (Morion)
GASTROPODA
Gyrodes abyssinus (Morton)
Acteon cretacea Gabb
Anchura abrupta Conrad
Turbinella parva Gabb
Lunatia halli Gabb
Pyropsis trochiformis (Tuomey)
Volutoderma ovata Whitfield
Turbinella subconica Gabb
Turritella vertebroides Morton
PELECYPODA
Cardium tenuistriatum Whitfield
Glycymeris mortoni (Conrad)
Gryphaea convexa (Say)
Gervilliopsis ensiformis (Conrad)
Panopea decisa Conrad
Veniella conradi Morton
Crassatella vadosa Morton
Cucullaea vulgaris Morton
Lithophaga ripleyana Gabb
Xylophagella irregularis (Gabb)
NiAculana stephensoni Richards
Etea delawarensis (Gabb)
NAUTILOIDEA
Euirephoceras dekayi (Morton)
AMMONOIDEA
Baculites ovatus Say
Sphenodiscus lobatus (Tuomey)
Pachydiscus (Neodesmoceras) sp.
CRUSTACEA
cf. Hoploparia sp.

CHONDRICHTHYES
Lamna appendiculata (Agassiz)
Odontaspis cuspidata (Agassiz)
Squalicorax prislodontus (Morton)
Hexanchus sp.
Edaphodon stenobryus (Cope)
Edaphodon mirificus Leidy
Ischyodus cf. /. thurmanni Pictet and Campiche
Squalina sp.
Myliobatis cf. M. leidyi Hay
Ischyrhiza mira Leidy
Rhinoptera sp.
cf. Rhombodus levis Cappetta and Case
OSTEICHTHYES
Enchodus cf. E.ferox Leidy
Enchodus cf. E. serrulatus Fowler
Paralbula casei Estes
CHELONIA
Adocus beatus Leidy
Osteopygis emarginalus Cope
Taphrospys sulcatus (Leidy)
Dollochelys atlantica (Zangerl)
CROCODILIA
cf. Procaimanoidea sp.
Hyposaurus rogersii Owen
Thoracosaurus sp.
Botlosaurus harlani Meyer
Diplocynodon sp.
LACERTILIA
Mosasaurus sp.
Plioplatecarpus sp.

20

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Conrad, Timothy A.
1869.    Notes  on  American  Fossiiiferous  Strata. American Journal of
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Cope, Edward Drinker
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Cracraft, Joel
1972. A New Cretaceous Charadriiform Family. Auk, 89:36-46, 3 figures,
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Erickson, Bruce R.
1975.    Dakotornis cooperi, a New Paleocene Bird from North Dakota.
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Fiirbringer, Max
1888. Untersuchungen zur Morphologie und Systematik der Vogel,
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Gaffney, Eugene
1975.    A Revision of the Side-necked Turtle Taphrosphys sulcatus (Leidy)
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Harlan, Richard
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Hay, Oliver R
1902.    Bibliography  and Catalogue of the Fossil Vertebrata of North
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Howard, Hildegarde
1955.    A New Wading Bird from the Eocene of Patagonia. American
Museum Novitates. 1710: 25 pages, 8 figures, 3 ubles.
Koch, Robert C, and Richard K. Olsson
1977. Dinoflagellate and Planktonic Foraminiferal Biostratigraphy of the
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Lambrecht, Kalman
1933.    Handbuch der Palaeornithologie. 1024 pages, 209 figures, 4 plates.
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Marsh, O.C.
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1872.    Preliminary Description of Hesperornis regalis. with Notices of
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1880.    Odontornithes: A Monograph on the Extinct Toothed Birds of North
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Miller, Halsey W, Jr.
1955.    A Check-list of the Cretaceous and Tertiary Vertebrates of New
Jersey. Journal of Paleontology, 29(5):903-914.
Minard, J.R, W.J. Perry, E.G.A. Weed, E.C. Rhodehamel, E.L Robbins, and
R.B. Mixon
1974.    Preliminary Report on Geology along Atlantic Coastal Margin of
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Moore, Raymond C.
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Morton, Samuel G.
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Petters, Sunday W
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of New Jeney. American Association of Petroleum Geologists
Bulletin. 60(1):87-107, 7 figures.

21

22

SMITHSONIAN CONTRIBUTIONS TO BOTANY



Rapp, William F., Jr.
1943.    List of the Fossil Birds of New Jersey. Journal of Paleontology,
17(1):124.
Shufeldt, R.W.
1915. Fossil Birds in the Marsh Collection of Yale University. Trans-
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Sibley, Charles G., and Jon E. Ahlquist
1972.    A Comparative Study of the Egg White Proteins of Non-Passerine
Birds.  Peabody Museum  of Natural History,   Yale  University.
Bulletin, 39: 276 pages, 37 figures.
Stephenson, L.W., RB. King, W.H. Monroe, and R.W. Imlay
1942. Correlation of the Outcropping Cretaceous Formations of the
Atlantic and Gulf Coastal Plain and Trans-Pecos, Texas. Geological
Society of America Bulletin. 53:435-448, 1 plate.

Thulborn, Richard A., and Tim L. Hamley
1985. A New Palaeoecological Role for Archaeopteryx. In M.K. Hecht,
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Vanuxem, Lardner
1829.    Remark on the Characters and Classification of Certain Rock
Formations. American Journal of Science. 16:254-256.
Wetmore, Alexander
1926.    Fossil Birds from the Green River Deposits of Eastern Utah. Annals
ofthe Carnegie Museum. 16(3-4):391-402, plates 36-37.
1930.    The Age of the Supposed Cretaceous Birds from New Jersey. Auk.
47(2): 186-188.
1940.    A Check-list of the Fossil Birds of North America. Smithsonian
Miscellaneous Collections. 99(4): 1-81.

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