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

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.

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 9d

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.