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.

**Genus Telmatornis Marsh, 1870**

Type-Species.—Telmatornis priscus Marsh, 1870, by subsequent designation (Hay, 1902:528).

Included Species.—Type species only.

Telmatornis priscus Marsh, 1870

Figures 5b-j, 6c,e,g, 7a,d,g,j,n

Telmatornis priscus Marsh, 1870:210. Telmatornis affinis Marsh, 1870:211. Graculavus pumilis Marsh, 1872:364. ?Palaeotringa vetus Marsh, 1870:209.

Holotype.—Distal end of left humerus ([Figure 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 5f,g]), 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 5i]) and several fragments of shafts of long bones apparently associated (holotypical material of Graculavus pumilis Marsh, 1872); collected near Hornerstown, New Jersey, by J.G. Meirs; probably from the basal Hornerstown Formation, Maastrichtian, Late Cretaceous.

Distal end of left tibiotarsus ([Figure 7n]), 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 7d,g,j]), NJSM 11853; collected 27 March 1975 by David C. Parris from the main fossiliferous layer of the Inversand Company marl pit.

Figure 5.—Wing elements of Burhinus and Telmatornis. a, Burhinus vermiculatus (USNM 488870), proximal end of right humerus, anconal view, b-d, Telmatornis priscus (holotype of Graculavus pumilis, YPM 850), proximal end of right humerus (b, anconal view; c, palmar view; d, proximal view), e,h, T. priscus (holotype, YPM 840), distal end of left humerus (e, anconal view; h, palmar view), f,g, T. priscus (holotype of Telmatornis affinis, YPM 845), distal end of right humerus (f, aconal view; g, palmar view), i, T. priscus (associated with YPM 850), distal end of left carpometacarpus, dorsal view; j, T. priscus (NJSM 11900), proximal end of right ulna. (All figures x 2; specimens coated with ammonium chloride to enhance detail.)

Figure 6.—Humeri of Anatalavis, new genus, and Telmatornis. a, Anatalavis rex (holotype, YPM 902), right humerus, palmar view; × 1.5. b,d,f, A. rex, (YPM 948), left humerus (b, palmar view, × 1.5; d, enlarged, anconal view, × 2; f, enlarged, palmar view, × 2). c,e,g, Telmatornis priscus, (ANSP 15360), left humerus (c, palmar view, × 1.5; e, enlarged, anconal view, × 2; g, enlarged, palmar view, × 2); h, Burhinus vermiculatus (USNM 430630), left humerus, palmar view, × 2. (Specimens coated with ammonium chloride to enhance detail.)

Figure 7.—Hindlimb elements. a,b, Right pedal phalanx 1 of digit II (a, Telmatornis priscus, ANSP 15541; b, Presbyornis sp., USNM uncatalogued; part of associated foot), c-k, Distal end of left tarsometatarsus, anterior, posterior, and distal views, respectively (c,f,i, Presbyornis sp., UCMP 126178; d,g,j, T. priscus, NJSM 11853; e,h,k, Burhinus vermiculalus, USNM 488870). l-n, Distal portions of left tibiotarsi (l, Palaeotringa littoralis, holotype, YPM 830; m, P. vagans, holotype, YPM 835; n, T. priscus, holotype of P. vetus, ANSP 13361). (All figures × 2; specimens coated with ammonium chloride to enhance detail.)

Right pedal phalanx 1 of digit II ([Figure 7a]), ANSP 15541; collected in 1972 by Richard White at the Inversand Company marl pit.

Proximal end of right ulna ([Figure 5j), NJSM 1]1900; collected 14 July 1978 from spoil piles near junction of Routes 537 and 539, near Hornerstown, Upper Freehold Township, Monmouth County, New Jersey, by Gerard R. Case; presumably from the Hornerstown Formation but whether from Cretaceous or Tertiary sediments is not known.

Miller (1955) lists an additional specimen from near Arneytown under the name Palaeotringa vetus (YPM 2808). This was cataloged in 1937 as "part of a tibia" of "Eocene" age but the specimen cannot now be located in the Yale collections and its age and identity must be considered very doubtful.

Measurements (in mm).—Distal ends of humeri (YPM 840, YPM 845, ANSP 15360, respectively): distal width 10.9, 10.1, 11.3; depth through dorsal condyle 5.7, 5.2, 5.5; width of shaft at proximal extent of brachial fossa 6.3, 5.5,6.4; length from distal end of pectoral crest to ventral condyle (ANSP 15360 only) 45.1; shaft width at midpoint (ANSP 15360 only) 4.7.

Proximal end of humerus YPM 850: proximal width through dorsal and ventral tubercles 13.1; depth through bicipital surface and ventral condyle 7.5, depth of head approximately 3.5.

Proximal end of ulna NJSM 11900: depth through dorsal cotyla 7.0.

Distal end of carpometacarpus YPM 840: depth at distal end 5.3; shaft width 2.9.

Distal end of tibiotarsus ANSP 13361: shaft width 3.5, approximate depth through medial condyle 6.9.

Distal end of tarsometatarsus NJSM 11853: distal width 6.1+; shaft width 2.7.

Pedal phalanx 1 of digit II: length 14.6; proximal width 3.0.

Comparisons.—This is evidently the most abundant bird in the New Jersey Cretaceous deposits. Hitherto it had been known only from the two distal ends of humeri that are the holotypes of Telmatornis priscus and T. affinis. Marsh (1870) did not clearly place Telmatornis with any living family but mentioned species of Rallidae, Scolopacidae, and Ardeidae in his comparisons. Hay (1902:528) listed the genus under the Rallidae. Shufeldt (1915:26) considered that Telmatornis was not a heron but might be related either to rail-like or charadriiform birds, the material, according to him, being insufficient for positive determination. He (1915:27) also described a larger species, Telmatornis rex, which we have removed to a new genus. Lambrecht (1933:489) maintained Telmatornis as a genus incertae sedis in his order Ralliformes. Brodkorb (1967) placed the genus in the family Rallidae, subfamily Rallinae, without comment. Cracraft (1972) established that Telmatornis did not belong in the Rallidae but was instead very similar to the Burhinidae. He synonymized T. affinis with T. priscus and created a new family, Telmatornithidae, for T. priscus and T. rex.

We concur in synonymizing T. affinis with T. priscus. The holotypes and the new specimen of humerus (ANSP 15360), which is instructive in that it preserves much more of the shaft ([Figure 6c]), are indeed very similar to the humeri in the Burhinidae. In size they are closely comparable to the small living species Burhinus vermiculatus (cf. [Figure 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 slightly more distally located; in distal view (3) the ventral condyle smaller and less rounded; and (4) the dorsal tricipital groove shallower.

The distal portion of the humerus of Telmatornis is similar to that in Presbyornis but differs in having (1) the dorsal condyle decidedly more elongate; (2) olecranal fossa much shallower; (3) ventral epicondyle in ventral view less distinctly demarcated but (4) more protrudent in lateral or medial view.

The proximal end of humerus (YPM 850) that is the holotype of Graculavus pumilis was considered by Shufeldt (1915:19) definitely to be from a limicoline charadriiform. It is from a bird exactly the size of Telmatornis priscus and its coloration and preservation would not be incompatible with its being the opposite end of the same bone as the holotype of T. affinis ([Figure 5b,c,f,g]). The following differences between the holotypical humeri of G. velox and _"G." pumilis establish that these belong to different genera: (1) in velox the area dorsal to the ventral tubercle and distal to the head is much more excavated, undercutting the head; (2) the dorsal tubercle is more pronounced; (3) there is a distinct excavation distomedial to the ventral tubercle, lacking in pumilis; (4) the ventral tubercle in ventral view is much more produced in velox than in pumilis.

The holotype of G. pumilis is very similar to the humerus in the Burhinidae but differs from that family and agrees with Graculavus in characters 8, 9, and 10 ([p. 6]). It differs further from the Burhinidae in having the area for the attachment of M. scapulohumeralis caudalis extending farther distally in ventral view. It differs from Presbyornis mainly in lacking the excavation to and undercutting the head. Because pumilis is not congeneric with Graculavus velox and because of its size and similarities with the Burhinidae and Presbyornis, we have no hesitation about considering Graculavus pumilis Marsh, 1872, to be a junior subjective synonym of Telmatornis priscus Marsh, 1870.

The proximal end of an ulna, NJSM 11900 ([Figure 5j]), is from a bird the size of Burhinus vermiculatus and not too dissimilar to it except that the shaft is more robust in the fossil. The specimen is too imperfect to merit detailed study and is referred to Telmatornis priscus only on size and probability.

The very fragmentary distal end of carpometacarpus associated with the type of G. pumilis ([Figure 5i]) is slightly larger and more robust than in Burhinus vermiculatus, but not so much as to be incompatible with T. priscus. Compared to Burhinus (1) the symphysial area is deeper and (2) the articular surface for the major digit is proportionately larger, the specimen being somewhat more similar to the carpometacarpus in Presbyornis.

The three specimens of Palaeotringa Marsh from the Cretaceous of New Jersey are based on poorly preserved distal ends of tibiotarsi. The holotype of Palaeotringa vetus Marsh, 1870 ([Figure 7n]) is similar in size to the comparable element in Burhinus vermiculatus, though with a relatively more slender shaft, and hence is from a bird the size of T. priscus, being smaller than any of the other species of Palaeotringa. It is more similar to Presbyornis than to Burhinus. Because it is from a charadriiform the size of T. priscus, as first revisers we tentatively consider Palaeotringa vetus Marsh, 1870, to be a subjective synonym of Telmatornis priscus Marsh, 1870. The only alternative would be to consign it to Aves incertae sedis. It is of passing historical interest to recall Marsh's (1870:209) comment that the type of Palaeotringa vetus "apparently was the first fossil bird-bone discovered in this country," having been mentioned both by Morton (1834) and Harlan (1835) as belonging to the genus Scolopax (Charadriiformes: Scolopacidae).

The distal portion of tarsometatarsus NJSM 11853 ([Figure 7d,g,f]) is unfortunately quite abraded. It is from a small charadriiform and has a shaft width about the same as in Burhinus vermiculatus. If this fossil came from an individual of Telmatornis priscus, as we assume, T. priscus being the smallest and most abundant "graculavid" in the New Jersey Cretaceous deposits, then it is a very instructive specimen, for it differs much more from Burhinus than does the humerus of Telmatornis. NJSM 11853 differs from the Burhinidae and agrees with Presbyornis in having (1) the distal foramen proportionately large and oval, not very small and circular; (2) a large, well-developed scar for the hallux (hallux absent in Burhinidae); (3) external trochlea proximodistally more elongate. That which remains of the inner trochlea indicates that it was (1) somewhat more posteriorly retracted than in Burhinus but (2) not nearly as elevated and retracted as in Presbyornis.

Pedal phalanx ANSP 15541 ([Figure 7a]) is from a bird the size of T. priscus. This specimen is much longer and more slender than phalanx 1 of digit II in Burhinus vermiculatus but has almost exactly the shape and proportions of the same element in Presbyornis ([Figure 7b]), although being much smaller. Although its assignment to Telmatornis is very tentative, the length of this element seems to indicate a wading bird as opposed to one with the terrestrially adapted shorter toes of the Burhinidae.

**Genus Anatalavis, new genus**

Type-Species.—Telmatornis rex Shufeldt, 1915.

Included Species.—Type-species only.

Diagnosis.—Differs from Telmatornis and Presbyornis in (1) having the shaft very short, stout, and much more curved, both in dorsoventral and lateromedial views. Differs from Telmatornis and agrees with Presbyornis in (2) having the distal end in distal view deeper, with (3) a narrower and much deeper olecranal fossa. Also, (4) the brachial depression is smaller and narrower than in Telmatornis but not as deep, nor as proximally situated as in Presbyornis.

Etymology.—"Duck-winged bird," from Latin anas, duck, ala, wing, and avis, bird. The gender is feminine.

Anatalavis rex (Shufeldt, 1915), new combination

Figure 6a,b,dJ

Telmatornis rex Shufeldt, 1915:27, fig. 101.

Holotype.—Right humerus lacking proximal end, YPM 902 ([Figure 6a]).

Locality and Horizon.—From Hornerstown, Upper Freehold Township, Monmouth County, New Jersey; collected by W. Ross in 1878; probably Late Cretaceous (Maastrichtian), basal Hornerstown Formation.

Referred Specimen.—Paratypical left humerus lacking proximal end, YPM 948 ([Figure 6b,d,f]). From Hornerstown, Upper Freehold Township, Monmouth County, New Jersey; collected by J.G. Meirs in 1869; probably Late Cretaceous (Maastrichtian), basal Hornerstown Formation.

Measurements (in mm).—Humeri (YPM 902, YPM 948, respectively): distal width 13.6, 13.2; depth through dorsal condyle 7.3, 7.5; width of shaft at proximal extent of brachial fossa 7.2,7.5; length from distal end of pectoral crest to ventral condyle 49.1, 50.7; shaft width at midpoint 5.4, 5.6.

Remarks.—Shufeldt (1915:27) described this species in the same genus as T. priscus and T. affinis but correctly noted that the humerus "is a short one ... its sigmoid curve very pronounced." Cracraft (1972:41) considered that "except for its decidedly larger size, T. rex does not differ from T. priscus in any significant features." In fairness to these authors, it should be noted that the great differences between Anatalavis and Telmatornis are much more apparent in comparisons with the new humerus of T. priscus (ANSP 15360), which preserves much more of the shaft than the previously known specimens. Both Shufeldt and Cracraft considered YPM 948 to belong to the same species as the holotype of T. rex, and we concur.

The specimens of A. rex are not comparable with the type of Graculavus velox, which was from a larger bird. Anatalavis rex was a larger, heavier bird than Telmatornis priscus, with the humerus remarkably short and robust, so that the overall length of the humerus in A. rex would scarcely have exceeded that of T. priscus. Anatalavis must have been a bird of considerably different flight habits from Telmatornis or Presbyornis. The overall appearance of its humerus is in fact rather duck-like, except for the more expanded distal end. It is still quite short and stout even for a duck.

Genus Laornis Marsh, 1870

Type-Species.—Laornis edvardsianus Marsh, 1870, by monotypy.

Included Species.—Type species only.

Laornis edvardsianus Marsh, 1870

Figure 8a,c,e

Laornis edvardsianus Marsh, 1870:206.

Holotype.—Distal end of right tibiotarsus, YPM 820.

Locality and Horizon.—From pits of the Pemberton Marl Company at Birmingham, Burlington County, New Jersey; collected by J.C. Gaskill; Late Cretaceous (Maastrichtian), basal Hornerstown Formation.

Measurements (in mm).—Distal end of tibiotarsus, YPM 820: distal width across condyles 22.6, depth of external condyle 19.3, depth of internal condyle 21.1, least shaft width 11.7, least shaft depth 9.6.

Comparisons.—The very large size of this specimen has undoubtedly been a factor in misleading those who have attempted to identify it, as it came from a bird the size of a swan or a large crane. The affinities of this fossil have long been questioned and the species has for most of its history been in limbo. Marsh (1870:207) concluded only that Laornis "shows a strong resemblance in several respects to the Lamellirostres [Anseriformes], and also to the Longipennes [Charadriiformes (Lari) and Procellariiformes], but differs essentially from the typical forms of both of these groups." In its own nebulous way, this assessment is concordant with our placement of Laornis in a charadriiform group that was near the ancestry of the Anseriformes. Doubtless only on the strength of Marsh's comments. Cope (1869-1870:237) placed Laornis in the "Lamellirostres." Hay (1902:531) included Laornis in the Anatidae. Shufeldt (1915:23) hardly clarified matters when he characterized Laornis as "at least one of the generalized types of waders," being a "remarkable type, which seems to have, judging from this piece of the tibiotarsus, Turkey, Swan, Crane, and even other groups all combined in it." Lambrecht (1933:526) included Laornis as a genus incertae sedis in his "Telmatoformes," between the Aramidae and Otididae.

The type was restudied by Cracraft (1973:46) who put Laornis in the Gruiformes and created a new family (Laornithidae) and superfamily (Laornithoidea) for it. He included it in his suborder Ralli, the only other member of which was the Rallidae. After preliminary comparisons, Olson (1974) ventured that Laornis belonged in the suborder Lari of the Charadriiformes. Brodkorb (1978:214) listed Laornis under Aves incertae sedis and guessed that it might be related to the Pelecaniformes.

Except for the extreme difference in size, the tibiotarsus of Laornis is in many respects similar to that of Presbyornis ([Figure 8]), especially in (1) the shape and position of the tubercle proximal to the external condyle; (2) the transverse pit in the intercondylar sulcus; and (3) the broad, shallow intercondylar sulcus as seen in distal view. It differs in a seemingly minor but quite characteristic feature, the large nutrient foramen situated in the groove for M. peroneus brevis ([Figure 8c]). This is absent in Presbyornis but is present in both of the tibiotarsi from the Cretaceous of New Jersey in which that portion of the bone is preserved (the holotypes of Palaeotringa littoralis and P. vagans), as well as in a tibiotarsus (Science Museum of Minnesota P75.22.25) from the type-locality of Dakotornis cooperi Erickson, 1975, that may be referable to that graculavid-like species. The foramen in the peroneus brevis groove may also be found in at least some specimens of Stercorariidae, which is partly what led Olson (1974) to suggest a relationship between Laornis and the Lari. Laornis appears to have been an extremely large member of the "transitional Charadriiformes," though where its relationships may lie within that group cannot be determined.

**Genus Palaeotringa Marsh, 1870**

Type-Species.—Palaeotringa littoralis Marsh, 1870; by subsequent designation (Hay, 1902:527).

Included Species.—Palaeotringa littoralis Marsh, 1870, and Palaeotringa vagans Marsh, 1872.

Palaeotringa littoralis Marsh, 1870

Figure 7l

Palaeotringa littoralis Marsh, 1870:208.

Holotype.—Distal portion of left tibiotarsus lacking most of the inner condyle, YPM 830.

Locality and Horizon.—Collected in the "middle marl beds" by Nicolas Wain from his marl pits near Hornerstown, New Jersey; Late Cretaceous (Maastrichtian), either basal Hornerstown Formation or Navesink Formation.

Measurements (in mm).—Depth through outer condyle 8.2; width of shaft just proximal to outer condyle 7.0.

Comparisons.—This specimen and that of P. vagans are too fragmentary for useful comparison. Both have the foramen in the groove for M. peroneus brevis, mentioned above. Their overall similarity to Presbyornis and to charadriiform birds in general justifies retaining them with the other "graculavids" but other than this little else can be said. In size, Palaeotringa littoralis would have been about equal to Burhinus bistriatus vocifer and smaller than Esacus magnirostris. Hence it would seem to be too small to belong to the same species as Graculavus velox and is definitely too large to be referable to Telmatornis priscus.

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

Palaeotringa littoralis?

Figure 9a

Referred Material.—Distal portion of a left humerus, NJSM 11303.

Measurements (in mm).—Distal width 12.8, depth through dorsal condyle 6.9, width of shaft at proximal extent of brachial fossa 8.2.

Comparisons.—This interesting specimen, although considerably worn, 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 7m

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.