[Figure 4] b,d,f,h
Graculavus velox Marsh, 1872:363.
Limosavis velox (Marsh).—Lambrecht, 1933:546.
Holotype.—Proximal end of left humerus, YPM 855.
Locality and Horizon.—From Hornerstown, Upper Freehold Township, Monmouth County, New Jersey; collected by J.G. Meirs; Late Cretaceous (Maastrichtian), either basal Hornerstown Formation or Navesink Formation.
Measurements (in mm).—Proximal end of humerus, YPM 855: proximal width through dorsal and ventral tubercles 21.1, depth through bicipital surface and tuberculum ventrale 11.6, depth of head 5.7.
Figure 4.—Proximal ends of left humeri of Graculavus velox and related birds: a, Esacus magnirostris (Burhinidae), USNM 19649; b,d,f,h, Graculavus velox, holotype, YPM 855; c,e,g, i, Presbyornis sp., UCMP 126205. a-c, anconal view; d,e, anconal view with distal portion tilted upwards; f,g, palmar view; h,i, proximal view. All figures × 2; specimens coated with ammonium chloride to enhance detail.
Comparisons.—Marsh (1872) originally described this as a species of cormorant (Phalacrocoracidae, Pelecaniformes) and included the species G. pumilis Marsh, 1872, also from New Jersey, and G. anceps Marsh, 1872, from the Late Cretaceous of Kansas, in the same genus. Marsh (1880) later referred G. anceps to the genus Ichthyornis, where it has remained. Shufeldt (1915:17-19) went into considerable detail to show that the species of Graculavus, particularly G. velox, were not cormorants, instead being limicoline shorebirds with similarities to the Burhinidae, Haematopodidae, and Charadriidae. Accordingly, Lambrecht (1933:540, 546) placed these taxa among the charadriiform birds, but rather inexplicably listed velox under Shufeldt's substitute name Limosavis in the suborder Laro-Limicolae, while retaining pumilis in the genus Graculavus in the suborder Limicolae. Brodkorb (1963b:249) ignored Shufeldt's assessment of relationships and placed G. velox and G. pumilis in the Phalacrocoracidae, subfamily Graculavinae. Cracraft (1972) did not examine the specimens attributed to Graculavus in his consideration of the relationships of Telmatornis.
We have synonymized Graculavus pumilis Marsh, 1872, with Telmatornis priscus Marsh, 1870, and discuss below the characters by which Graculavus (restricted to G. velox) may be separated from Telmatornis. Shufeldt (1915) has already presented adequate evidence that Graculavus is not a cormorant and is instead a charadriiform. The following combination of characters of the proximal end of the humerus is shared by Graculavus and Presbyornis and distinguishes these genera from other Charadriiformes: (1) lack of a distinct lanceolate scar for M. coracobrachialis cranialis; (2) lack of a distinctly excavated second (dorsal) tricipital fossa; (3) presence of a distinct tumescence in the proximoventral portion of the tricipital fossa; scars for (4) M. scapulohumeralis caudalis and (5) M. scapulohumeralis cranialis very large and distinct; (6) attachment of M. latissimus dorsi cranialis a well-defined, raised protuberance situated dorsal to the median ridge of the shaft; (7) tuberculum dorsale well defined, distinctly pointed. In most of the preceding characters that it preserves, the single proximal end of humerus referred to Telmatornis (the holotype of G. pumilis) agrees with Graculavus and Presbyornis.