clearly has the overall “graculavid” morphology but shows sufficient
differences from the humeri of _Telmatornis_ or _Anatalavis_ to warrant
its generic separation from them. In size it is about equal to the
modern form _Burhinus bistriatus vocifer_ and hence would be compatible
with _P. littoralis_. It differs from _Telmatornis_, _Anatalavis_, or
_Presbyornis_, and is more similar to _Burhinus_ in having (1) the
brachial depression wider, shallower, and more proximally situated.
Although 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 others. In distal view the specimen is more similar
to _Presbyornis_ than to the other Cretaceous humeri, although (3) the
olecranal fossa is shallower. If this specimen is correctly referred
to _Palaeotringa_, it shows that 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 7_m_
_Palaeotringa vagans_ Marsh, 1872:365.
Holotype.--Fragmented distal two-thirds of a left tibiotarsus lacking
the external condyle and the anterior portion of the internal condyle,
YPM 835.
Locality and Horizon.--From Hornerstown, Upper Freehold Township,
Monmouth County, New Jersey; collected by J.G. Meirs; Late Cretaceous
(Maastrichtian), “about ten feet below the surface of the 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 than _P. vetus_ (= _Telmatornis
priscus_). It differs from the latter and agrees with _P. littoralis_
in having the distal tendinal opening of a flattened oval shape, rather
than decidedly rounded. If we have correctly referred _P. vetus_ to
_Telmatornis priscus_, then it is certain that neither of the other
two species of _Palaeotringa_ can be referred to _Telmatornis_. In _P.
vagans_ the tendinal groove appears to be much narrower and the 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 9_b,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.
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 15
August 1972 by David C. Parris.
Measurements (in mm).--Humerus: distal width 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 still. In size this bird would have approximated the
modern flamingo _Phoeniconaias minor_, which it somewhat resembles in
morphology as well. The humerus is not greatly different from that
of other Graculavidae in general aspect but is distinct in having a
larger, much deeper, and more proximally situated brachial depression.
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 Procellariiformes?=
Among the newly collected material from the Inversand pit is a singular
avian humerus that cannot be assigned to the Graculavidae or to
any other 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.
[Illustration: Figure 9.--Miscellaneous elements, _a_, _Palaeotringa
littoralis?_ (NJSM 11303), distal end of left humerus, palmar view;
_b_, Graculavidae, genus and species indeterminate (NJSM 11302),
distal end of left humerus, palmar view; _c_, proximal end of
radius associated with _b_; _d_, _Graculavus velox?_ (NJSM 11854),
right carpometacarpus; _e,f_, Procellariiformes?, genus and species
indeterminate (ANSP 15713), 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_, × 2; _e,f,g_, × 5; specimens
coated with ammonium chloride to enhance detail.)]
The most distinctive features of this specimen are the deep brachial
depression and the incipient ectepicondylar spur, thus calling to mind
both the Lari (Charadriiformes) and the Procellariiformes among modern
birds. Among the Pelecaniformes it also bears a resemblance to the
Phaethontidae and especially to the 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 other Charadriiformes 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
developed than in any of the Pelecaniformes but not as well developed
as in the Procellariiformes. The apparently 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 the 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 the 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, 11
Holotype.--Right humerus lacking the ventral tubercle, portions of the
pectoral crest, and other parts of the proximal end, where partially
reconstructed, NJSM 11341.
Locality and Horizon.--Main fossiliferous layer of the 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 reconstructed, 110; width
and depth of shaft at midpoint 7.0 × 5.6; distal width 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 the Procellariiformes and
Pelecaniformes has been previously suggested (Sibley and Ahlquist,
1972:70; Olson, 1985:142), and among the pelecaniform taxa most often
mentioned as being procellariiform-like are the Fregatidae. It is
tempting to regard the humerus of _Tytthostonyx_ as being similar
to that possessed by the ancestral stock that gave rise to the
Procellariiformes. Its similarities also to the Eocene frigatebird
_Limnofregata_ would thus be seen as corroborating the primitiveness
of the Fregatidae within the 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 ectepicondylar spur and deep brachial
depression could be interpreted as tending in that direction.
On the other hand, we must admit that we are dealing with only a
single bone and one of very great age at that, so that the 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 9_e,f_
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 depth 3.1, width and
depth of shaft near point of break 1.8 × 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 straight shaft of the ulna, the shape and depth of the
tendinal grooves, and the relatively well-developed scars for the
attachment of the secondaries. It differs from the Oceanitidae in
having the ventral lip of the external condyle much more rounded and
protrudent past the plane of the shaft, whereas the carpal tubercle in
dorsal view is markedly smaller. On this basis, the fossil certainly
could not be referred to the Oceanitidae and that it should be
associated with the Procellariiformes may be doubted as well.
[Illustration: 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, × 0.8;
_c,d_, stereophotographs of coated specimen in anconal and palmar
views, × 1.3.]
[Illustration: 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 × 2; specimens coated
with ammonium chloride to enhance detail.)]
=Aves, incertae sedis=
Figure 9_g_
Referred Material.--Distal end of left femur, NJSM 12119.
Locality and Horizon.--Inversand Company marl pit, Sewell, Gloucester
County, New Jersey; from processed spoil piles, precise stratum
unknown; collected 12 December 1981 by Cynthia Miller. Presumably
from the Hornerstown Formation but could be either Late Cretaceous or
Paleocene.
Measurements (in mm).--Distal width 4.3, distal depth 3.8.
Comparisons.--This is also from a very small bird, possibly the same
size as the species represented by the preceding ulna (ANSP 15713;
Figure 9_e,f_) 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 treated here are avian and of Mesozoic age, it is
almost certain that too much importance will be made of them by some
future authors. 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 fossils. Therefore, in an effort
to forestall overenthusiasm for these fragments we shall present our
own brief assessment of their significance.
Unlike most other Cretaceous birds, such as the Hesperornithiformes,
Ichthyornithiformes, 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 fossils
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
the incompleteness of the fossils. There are certain modern birds, for
example the 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 families are represented
in the fauna, although if the various members of the “form-family”
Graculavidae were better known there can scarcely be any doubt that
more than one family would be recognized in this group. Within
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
smallest 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 well. 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 the various members
of the Graculavidae were probably as divergent from one another as are
_Presbyornis_ and _Burhinus_, their similarities being almost certainly
due to the retention of primitive characters.
Including the two genera and species that show some similarities to the
Procellariiformes, along with the small indeterminate femur, the total
avifauna from the New Jersey greensands comprises 8 or 9 genera and 9
or 10 species. As far as can be determined, all of the birds 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 their fantastic and highly improbable conjectures as
to the mode of life of _Archaeopteryx_. Indeed, just the opposite is
probably the case (Olson, 1985), the lack of Late Cretaceous fossils of
truly terrestrial or arboreal birds most likely being due to sampling
bias.
Appendix
The nonavian megafauna of the main fossiliferous layer (Basal
Hornerstown Formation), at the Inversand Company marl pit, Sewell,
Gloucester County, New Jersey is listed below. Also found in the
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).
Brachiopoda
_Terebratulina atlantica_ (Morton)
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)
_Nuculana stephensoni_ Richards
_Etea delawarensis_ (Gabb)
Nautiloidea
_Eutrephoceras dekayi_ (Morton)
Ammonoidea
_Baculites ovatus_ Say
_Sphenodiscus lobatus_ (Tuomey)
_Pachydiscus_ (_Neodesmoceras_) sp.
Crustacea
cf. _Hoploparia_ sp.
Chondrichthyes
_Lamma appendiculata_ (Agassiz)
_Odontaspis cuspidata_ (Agassiz)
_Squalicorax pristodontus_ (Morton)
_Hexanchus_ sp.
_Edaphodon stenobryus_ (Cope)
_Edaphodon mirificus_ Leidy
_Ischyodus_ cf. _I. thurmanni_ Pictet and Campiche
_Squatina_ 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. serrulalus_ Fowler
_Paralbula casei_ Estes
Chelonia
_Adocus beatus_ Leidy
_Osteopygis emarginatus_ Cope
_Taphrospys sulcatus_ (Leidy)
_Dollochelys atlantica_ (Zangerl)
Crocodilia
cf. _Procaimanoidea_ sp.
_Hyposaurus rogersii_ Owen
_Thoracosaurus_ sp.
_Bottosaurus harlani_ Meyer
_Diplocynodon_ sp.
Lacertilia
_Mosasaurus_ sp.
_Plioplatecarpus_ sp.
Literature Cited
Baird, Donald
1967. Age of Fossil Birds from the Greensands of New Jersey. _Auk_,
84:260-262.
Brodkorb, Pierce
1963a. Birds from the Upper Cretaceous of Wyoming. _Proceedings
of the XIIIth International Ornithological Congress_, pages
55-70, 10 figures.
1963b. Catalogue of Fossil Birds, Part 1 (Archaeopterygiformes
through Ardeiformes). _Bulletin of the Florida Slate Museum_,
_Biological Sciences_, 7(4):179-293.
1967. Catalogue of Fossil Birds, Part 3 (Ralliformes,
Ichthyornithiformes, Charadriiformes). _Bulletin of the
Florida Slate Museum_, _Biological Sciences_, 11(3):99-220.
1978. Catalogue of Fossil Birds, Part 5 (Passeriformes). _Bulletin
of the Florida Stale Museum_, _Biological Sciences_,
23(3):140-228.
Conrad, Timothy A.
1869. Notes on American Fossiliferous Strata. _American Journal of
Science_, series 2, 47:358-364.
Cope, Edward Drinker
1869-1870. Synopsis of the Extinct Batrachia, Reptilia and Aves of
North America. _Transactions of the American Philosophical
Society_ [for 1871], new series, 14:252 pages, 55 figures, 14
plates.
Cracraft, Joel
1972. A New Cretaceous Charadriiform Family. _Auk_, 89:36-46, 3
figures, 2 tables.
1973. Systematics and Evolution of the Gruiformes (Class Aves), 3:
Phylogeny of the Suborder Grues. _Bulletin of the American
Museum of Natural History_, 151:1-128, 51 figures, 49 tables.
Erickson, Bruce R.
1975. _Dakotornis cooperi_, a New Paleocene Bird from North Dakota.
_Scientific Publications of the Science Museum of Minnesota_,
new series, 3(1):7 pages, 3 figures.
Fürbringer, Max
1888. _Untersuchungen zur Morphologie und Systematik der
Vögel, zugleich ein Beitrag zur Anatomie der Stütz- und
Bewegungsorgane._ 2 volumes, 1751 pages, 30 plates. Amsterdam:
Von Holkema. [Issued as _Koninklijk Zoölogisch Genootschap
"Natura Artis Magistra," Bijdragen tot de Dierkunde_
(Amsterdam), 15.]
Gaffney, Eugene
1975. A Revision of the Side-necked Turtle _Taphrosphys sulcatus_
(Leidy) from the Cretaceous of New Jersey. _American Museum
Novitates_, 2751:24 pages.
Harlan, Richard
1835. _Medical and Physical Researches: or Original Memoirs
in Medicine, Surgery, Physiology, Geology, Zoology, and
Comparative Anatomy,_ xxxix + 9-653 pages, illustrated.
Philadelphia: L.R. Bailey.
Hay, Oliver P.
1902. Bibliography and Catalogue of the Fossil Vertebrata of North
America. _Bulletin of the United States Geological Survey_,
179:868 pages.
Howard, Hildegarde
1955. A New Wading Bird from the Eocene of Patagonia. _American
Museum Novitates_, 1710:25 pages, 8 figures, 3 tables.
Koch, Robert C., and Richard K. Olsson
1977. Dinoflagellate and Planktonic Foraminiferal Biostratigraphy
of the Uppermost Cretaceous of New Jersey. _Journal of
Paleontology_, 51:480-491, 4 figures.
Lambrecht, Kalman
1933. _Handbuch der Palaeornithologie._ 1024 pages, 209 figures, 4
plates. Berlin: Gebrüder Borntraeger.
Marsh, O.C.
1870. Notice of Some Fossil Birds from the Cretaceous and Tertiary
Formations of the United States. _American Journal of
Science_, series 2, 49:205-217.
1872. Preliminary Description of _Hesperornis regalis_, with
Notices of Four Other New Species of Cretaceous Birds.
_American Journal of Science_, series 3, 3:360-365.
1880. Odontornithes: A Monograph on the Extinct Toothed Birds of
North America. _Report of the Geological Exploration of the
Fortieth Parallel_, volume 7: xv + 201 pages, 40 figures, 34
plates.
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.P., W.J. Perry, E.G.A. Weed, E.C. Rhodehamel, E.I. Robbins,
and R.B. Mixon
1974. Preliminary Report on Geology along Atlantic Coastal Margin
of Northeastern United States. _American Association of
Petroleum Geologists Bulletin_, 58(6):1169-1178, 7 figures.
Moore, Raymond C.
1958. _Introduction to Historical Geology._ 2nd edition, 656 pages,
593 figures. New York: McGraw Hill.
Morton, Samuel G.
1829. Description of the Fossil Shells Which Characterize the
Atlantic Secondary Formation of New Jersey and Delaware,
Including Four New Species. _Journal of the Academy of Natural
Sciences of Philadelphia_, series 1, 6:72-100.
1834. _Synopsis of the Organic Remains of the Cretaceous Group of
the United States._ 96 pages, 19 plates. Philadelphia: Key and
Biddle.
Olson, Storrs L.
1974. [Review of] Joel Cracraft. Systematics and Evolution of the
Gruiformes (Class Aves), 3: Phylogeny of the Suborder Grues.
_Auk_, 91(4):862-865.
1977. A Lower Eocene Frigatebird from the Green River Formation
of Wyoming (Pelecaniformes: Fregatidae). _Smithsonian
Contributions to Paleobiology_, 35:33 pages, 31 figures.
1981. The Museum Tradition in Ornithology--A Response to Ricklefs.
_Auk_, 98(1):193-195.
1985. The Fossil Record of Birds. In Donald S. Farner, James
R. King, and Kenneth C. Parkes, editors. _Avian Biology_,
8:79-238, 11 figures. New York: Academic Press.
Olson, Storrs L., and Alan Feduccia
1980a. Relationships and Evolution of Flamingos (Aves:
Phoenicopteridae). _Smithsonian Contributions to Zoology_,
316:73 pages, 40 figures, 2 tables.
1980b. Presbyornis and the Origin of the Anseriformes (Aves:
Charadriomorphae). _Smithsonian Contributions to Zoology_,
323:24 pages, 15 figures.
Perry, W.J., Jr, J.P. Minard, E.G.A. Weed, S.L. Robbins, and E.C.
Rhodehamel
1975. Stratigraphy of Atlantic Coastal Margin of United States
North of Cape Hatteras--Brief Survey. _American Association of
Petroleum Geologists Bulletin_, 59(9):1529-1548, 12 figures.
Petters, Sunday W.
1976. Upper Cretaceous Subsurface Stratigraphy of Atlantic Coastal
Plain of New Jersey. _American Association of Petroleum
Geologists Bulletin_, 60(1):87-107, 7 figures.
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.
_Transactions of the Connecticut Academy of Arts and
Sciences_, 19:1-109, 15 plates.
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., P.B. 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