Part 2

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 9_d_

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 5_b-j_, 6_c,e,g_, 7_a,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 5_e,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_f,g_), YPM 845 (holotype of _Telmatornis affinis_ Marsh 1870); same data as holotype of _T. priscus_.

Proximal end of right humerus (Figure 5_b-d_), YPM 850, with distal end of right carpometacarpus (Figure 5_i_) 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 7_n_), 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 6_c,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 7_d,g,j_), NJSM 11853; collected 27 March 1975 by David C. Parris from the main fossiliferous layer of the Inversand Company marl pit.

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

Proximal end of right ulna (Figure 5_j_), NJSM 11900; 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 6_c_), 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 6_g,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 5_b,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 5_j_), 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 5_i_) 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 7_n_) 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 7_d,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 7_a_) 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 7_b_), 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 6_a,b,d_J

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

Holotype.--Right humerus lacking proximal end, YPM 902 (Figure 6_a_).

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 6_b,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 8_a,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 8_c_). 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 7_l_

_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_.

=_Palaeotringa littoralis?_=

Figure 9_a_

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

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 27 September 1972 by David C. Parris.

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