saurornitholestes robustus, n. sp. - Robert M. Sullivan
saurornitholestes robustus, n. sp. - Robert M. Sullivan
saurornitholestes robustus, n. sp. - Robert M. Sullivan
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Lucas, S. G. and <strong>Sullivan</strong>, R.M., eds., 2006, Late Cretaceous vertebrates from the Western Interior. New Mexico Museum of Natural History and Science Bulletin 35.<br />
253<br />
SAURORNITHOLESTES ROBUSTUS, N. SP. (THEROPODA: DROMAEOSAURIDAE) FROM<br />
THE UPPER CRETACEOUS KIRTLAND FORMATION (DE-NA-ZIN MEMBER),<br />
SAN JUAN BASIN, NEW MEXICO<br />
ROBERT M. SULLIVAN<br />
Section of Paleontology and Geology, The State Museum of Pennsylvania, 300 North Street, Harrisburg, PA 17120-0024<br />
Abstract—Saurornitholestes <strong>robustus</strong>, n. <strong>sp</strong>., is named based on new frontal material from the upper part of the<br />
Kirtland Formation (De-na-zin Member), San Juan Basin, New Mexico. The frontal is larger, and unusually thick,<br />
compared to <strong>sp</strong>ecimens of Saurornitholestes langstoni. All <strong>sp</strong>ecimens of Saurornitholestes previously collected from<br />
the San Juan Basin, and referred to the Judithian taxon S. langstoni, are now provisionally referred to this new<br />
<strong>sp</strong>ecies. A single tooth and a left second pedal ungual are referred to cf. Saurornitholestes <strong>robustus</strong>. The recognition<br />
of a new <strong>sp</strong>ecies of Saurornitholestes from the Upper Cretaceous strata of the San Juan Basin, further supports the<br />
distinct temporal position of the upper Fruitland and Kirtland vertebrate faunas, and is consistent in the recognition of<br />
a Kirtlandian land-vertebrate “age” fauna.<br />
INTRODUCTION<br />
Ongoing fieldwork in the Upper Cretaceous strata of the San Juan<br />
Basin, New Mexico, continues to add to our knowledge of dinosaur and<br />
other fossil vertebrates during the Kirtlandian time interval (<strong>Sullivan</strong> and<br />
Lucas, 2003, 2006). Previously, <strong>Sullivan</strong> and Lucas (2000) reported on a<br />
water-worn left frontal of a small theropod dinosaur that compared favorably<br />
to <strong>sp</strong>ecimens of Saurornitholestes langstoni, with the exception that<br />
the anterior part of the New Mexico <strong>sp</strong>ecimen was slightly more constricted<br />
anteriorly. In the summer of 2005, a larger, more robust, left frontal, referable<br />
to the genus Saurornitholestes, was recovered from the upper part of<br />
the Kirtland Formation (De-na-zin Member) in Alamo Wash. This new<br />
frontal forces a reconsideration of the identity of the previously reported<br />
<strong>sp</strong>ecimen (SMP VP-1270). Here, I report on this new <strong>sp</strong>ecimen (SMP VP-<br />
1955), and designate a new <strong>sp</strong>ecies of Saurornitholestes, S. <strong>robustus</strong>, that<br />
adds to our knowledge of Late Cretaceous North American velociraptorines<br />
and <strong>sp</strong>ecifically the Kirtlandian vertebrate fauna. In this paper, AMNH =<br />
American Museum of Natural History (New York); GIN (IGM) = Mongolian<br />
Institute of Geology (Ulan Bataar); SMP = The State Museum of Pennsylvania,<br />
Harrisburg; and TMP = Royal Tyrrell Museum of Palaeontology,<br />
Drumheller.<br />
SYSTEMATIC PALEONTOLOGY<br />
DINOSAURIA<br />
SAURISCHIA Seeley, 1888<br />
THEROPODA Marsh, 1881<br />
DROMAEOSAURIDAE Matthew and Brown, 1922<br />
Velociraptorinae Barsbold, 1983<br />
SAURORNITHOLESTES Sues, 1978<br />
Type <strong>sp</strong>ecies— Saurornitholestes langstoni Sues, 1978.<br />
Comments— Previously, Saurornitholestes has been considered<br />
to a monotypic taxon known from skeletal material from the Dinosaur<br />
Park Formation of Alberta, Two Medicine Formation of Montana and the<br />
Kirtland Formation of New Mexico (Currie, 2005; <strong>Sullivan</strong> and Lucas,<br />
2000). Other reports, based solely on teeth, have also been made by Baszio<br />
(1997a,b), Currie et al. (1990), Peng et al. (2001), Rowe et al. (1992),<br />
Sankey (1998, 2003) and Sankey et al. (2002, 2005). The referral of <strong>sp</strong>ecimens<br />
to cf. Saurornitholestes langstoni from the Horseshoe Canyon and<br />
Scollard formations by Baszio (1997b) should be viewed with skepticism.<br />
Baszio (1997b), himself, noted that isolated teeth from the Scollard, Lance<br />
and Frenchman formations may actually belong to another theropod. In<br />
fact, the newly named Atrociraptor from the Horseshoe Canyon Formation<br />
of Alberta, has a skull that is very different from Saurornitholestes<br />
(e.g., a shorter and deeper face), but its teeth are almost indistinguishable<br />
from Saurornitholestes (Currie and Varricchio, 2004). For the present, it<br />
seems that the genus Saurornitholestes is restricted to the Late Campanian<br />
(Judithian and Kirtlandian).<br />
Saurornitholestes <strong>robustus</strong>, n. <strong>sp</strong>.<br />
Holotype— SMP VP-1955, nearly complete left frontal.<br />
Horizon and Type Locality— De-na–zin Member of the Kirtland<br />
Formation. The type locality is SMP loc. no. 388 (Alamo Wash [north fork]);<br />
UTM data is on file and available to qualified researchers.<br />
Age— late Kirtlandian (73 Ma; see <strong>Sullivan</strong> and Lucas, 2006).<br />
Diagnosis— A <strong>sp</strong>ecies of Saurornitholestes distinguished from<br />
Saurornitholestes langstoni by its larger and more robust frontal<br />
(twice as thick).<br />
Description— The holotype (SMP VP-1955, Fig. 1) is a nearly<br />
complete left frontal. There is some damage behind the orbital region toward<br />
the posterior process and there is also some minor damage along the<br />
anterior-most part of the orbital adjacent to the lacrimal facet. The dorsal<br />
and lateral (orbital) surfaces are weathered and <strong>sp</strong>lintered, giving the frontal<br />
a rather rough appearance. The ventral and medial surfaces are largely<br />
unweathered. De<strong>sp</strong>ite the weathered and less than complete condition, the<br />
frontal is adequate for diagnostic purposes.<br />
The frontal measures (anteroposteriorly along the midline) 62 mm<br />
in length. Maximum width cannot be established with certainty because of<br />
the damage along the lateral portion of the postorbital, but I estimate it at<br />
about 30 mm.<br />
Dorsally, the frontal is relatively flat, rising slightly toward the rim<br />
of the orbital border. There is a slight depression on the dorsal surface above<br />
where the confluence of the posterior margin of the olfactory bulb and the<br />
anterior margin of the cerebral hemi<strong>sp</strong>here impression occur (on the ventral<br />
side). The area of the frontal-nasal contact is damaged, both medially<br />
and laterally. The lacrimal contact surface is intact, and is well-developed<br />
on the anterolateral side of the frontal, but it is slightly damaged along its<br />
lateroposterior-most portion at the front of the orbital rim.<br />
Ventrally, the olfactory bulb and cerebral hemi<strong>sp</strong>here impressions<br />
are present, although the former is weakly developed. Anteriorly, the frontal<br />
is relatively thick and presumably had a strong contact with the nasal.<br />
The olfactory bulb impression is less pronounced than that of SMP VP-<br />
1270 (<strong>Sullivan</strong> and Lucas, 2000), but that may be due to the fact that SMP<br />
VP-1270 is water-worn. The bridge between the olfactory bulb impression<br />
and the cerebral hemi<strong>sp</strong>here impression is not as constricted as in SMP<br />
VP-1270, again probably for the same reasons. The cerebral hemi<strong>sp</strong>here<br />
impression is well-defined, bordered laterally by the prominent descending<br />
orbital border. The frontoparietal articulation surface is weakly developed<br />
and slightly damaged.
254<br />
FIGURE 1. Saurornitholestes <strong>robustus</strong>, n. <strong>sp</strong>. SMP VP-1955 (holotype), nearly complete left frontal, from SMP loc. # 388, De-na-zin Member, Kirtland<br />
Formation. A, dorsal view (stereo pair); B, ventral view (stereo pair); C, orbital rim (lateral view) (dorsal direction is to the right); D, medial view of midline<br />
sutural surface (dorsal direction is to the left). Bar scale = 10 mm.
FIGURE 2. Cf. Saurornitholestes <strong>robustus</strong>, n. <strong>sp</strong>. SMP VP-1901 (referred<br />
<strong>sp</strong>ecimen), islolated tooth, from SMP loc. # 350, De-na-zin Member, Kirtland<br />
Formation. Bar Scale = 5 mm.<br />
Median suture thickness is 8 mm, measured at the confluence of<br />
the posterior margin of the olfactory bulb and the anterior margin of the<br />
cerebral hemi<strong>sp</strong>here impression. The maximum thickness of the frontal is<br />
where the posterior border descends ventrally. Here, the frontal measures<br />
10 mm thick, and is more than twice the thickness of the holotype (TMP<br />
74.10.5) in this region. The median suture surface is very rugose.<br />
The lateral surface of the orbital wall is weathered. No foramina are<br />
visible, due to the condition of the lateral surface.<br />
Comments— The massive nature of the holotype frontal (SMP VP-<br />
1955) clearly distinguishes it from the holotype of Saurornithololestes<br />
langstoni (TMP 74.10.5), leaving little doubt as to its <strong>sp</strong>ecific distinctiveness.<br />
The ratio of the length (measured along the midline) to the thickness<br />
(posterior part of the frontal) is 6:1 for S. <strong>robustus</strong> and 10:1 for S. langstoni.<br />
Moreover, the holotype of S. <strong>robustus</strong> differs from referred <strong>sp</strong>ecimens of<br />
Velociraptor mongoliensis (Barsbold and Osmólska, 1999), although they<br />
share a number of features (i.e., length/width ratio; shallow depression along<br />
mid-length of frontal; strong frontal nasal contact; lacrimal overlapping the<br />
frontal dorsally, etc.), which are herein considered characters of<br />
velociraptorines in general. Thus, there are sufficient reasons to retain both<br />
Saurornitholestes and Velociraptor as distinct genera.<br />
The frontal, once attributed to the <strong>sp</strong>ecies S. langstoni, has all the<br />
morphological attributes of the genus. These include: 1) triangular shape;<br />
2) posteroventral projection of the frontoparietal contact; 3) sigmoidally<br />
curved ridge running anterolaterally onto the frontal margin of the postorbital<br />
process; and 4) posterior concavity around the supratemporal fossae<br />
(Sues, 1978; Currie, 1987; <strong>Sullivan</strong> and Lucas, 2000). The <strong>sp</strong>ecific difference<br />
is the massive thickness of the frontal and somewhat larger size.<br />
Cf. Saurornitholestes <strong>robustus</strong><br />
Material— SMP VP-1901, tooth (Fig. 2), SMP loc #350; and SMP<br />
VP-1741, left second pedal ungual (Fig. 3), SMP loc. #361. Both <strong>sp</strong>ecimens<br />
were collected from the De-na-zin Member, Kirtland Formation, San<br />
Juan Basin, New Mexico<br />
Comments— The tooth (SMP VP-1901) is nearly complete and is<br />
characteristically laterally compressed and strongly recurved. It has a FABL<br />
(fore-aft basal length) of 6.5 mm. The height of the tooth is 12 mm, mea-<br />
255<br />
FIGURE 3. Cf. Saurornitholestes <strong>robustus</strong>, n. <strong>sp</strong>. SMP VP-1741 (referred<br />
<strong>sp</strong>ecimen), left pes ungual (digit II), from SMP loc. # 361, De-na-zin Member,<br />
Kirtland Formation. A, medial view; B, lateral view. Bar Scale = 10 mm.<br />
sured from the base to the tip of the crown along the posterior carina; 15<br />
mm along the anterior carina. Denticles are present along both the anterior<br />
and posterior carinae, and they are slightly hooked toward the tip. Some of<br />
the denticles are damaged, e<strong>sp</strong>ecially along the anterior edge towards the<br />
base. The enamel veneer is largely missing, with only remnants adhering<br />
to the tooth in the vicinity of the denticles on both the anterior and posterior<br />
carinae. Although somewhat larger than most Saurornitholetes teeth, this<br />
tooth cannot be distinguished from those of Saurornitholestes (S. langstoni)<br />
as illustrated by Currie et al. (1990), Sankey et al. (2002) and Brinkman<br />
(2005).<br />
The second pedal ungual (SMP VP-1741) is incomplete both proximally<br />
and distally, making it difficult to accurately determine its total size.<br />
However, it measures 69 mm along the preserved outside curvature. It is<br />
laterally compressed and possesses a prominent lateral groove on each side.<br />
This groove is situated slightly higher on the left lateral side compared to<br />
the right side, thus it is from the left pes (Kirtland et al., 1993; Rahut and<br />
Werner, 1995; Norell and Makovicky, 1997). The pedal ungual is deeper<br />
than that of Velociraptor mongoliensis (IGM 100/985) described and illustrated<br />
by Norell and Makovicky (1997, fig. 6).<br />
DISCUSSION<br />
This is the second frontal of Saurornitholestes known from the San<br />
Juan Basin, New Mexico, and arguably it is the most important, as it documents<br />
features that serve to distinguish it from its presumed predecessor<br />
Saurornitholestes langstoni. The previous report of Saurornitholestes from<br />
the Kirtland Formation was also based on a left frontal from the San Juan<br />
Basin (<strong>Sullivan</strong> and Lucas, 2000). However, this <strong>sp</strong>ecimen (SMP VP-1270)<br />
was smaller, slightly water-worn, and in some re<strong>sp</strong>ects, similar to the holotype<br />
of S. langstoni. At the time of its report, some minor differences were<br />
noted, but these were relegated to individual variation (<strong>Sullivan</strong> and Lucas,<br />
2000). A re-evaluation of SMP VP-1270, compared to the holotype of S.<br />
langstoni (TMP 74.10.5), which are the same size, shows it to be more<br />
like the holotype of S. <strong>robustus</strong> in having a thicker frontal. A single tooth<br />
(SMP VP-1901) and a left second pedal ungual (SMP VP-1714) are both<br />
from the De-na-zin Member of the Kirtland Formation and clearly pertain<br />
to Saurornitholestes (Figs. 2, 3). They are provisionally referred to S.
256<br />
<strong>robustus</strong> based on geographic and stratigraphic parsimony.<br />
Two <strong>sp</strong>ecies of Saurornitholestes are now known: S. langstoni and<br />
S. <strong>robustus</strong>. Currie (2005) cited three undescribed partial skeletons, two<br />
from the Dinosaur Park Formation of Alberta and one from the Two Medicine<br />
Formation of Montana.<br />
Although Saurornitholestes is considered to be similar to the Asian<br />
dromaeosaurid Velociraptor, recognition of two separate genera has been<br />
the consensus of most workers (Currie, 2005). Differences in the postcranial<br />
skeleton have already been documented by Norell and Makovicky<br />
(1997), and the deeper pedal ungual in Saurornitholestes <strong>robustus</strong>, if properly<br />
referred, further reinforces the argument for generic and <strong>sp</strong>ecific distinction.<br />
Although the frontals are not preserved in the holotype of<br />
Velociraptor mongoliensis (AMNH 6515) (Osborn, 1924; Sues, 1977),<br />
frontals are known from referred material (GIN 1000/24, 100/25) described<br />
in detail by Barsbold and Osmólska (1999). Comparison of the frontal material<br />
to that seen in a cast of skull of Velociraptor mongoliensis from a<br />
private collection, also verify the taxonomic distinction between the genera<br />
Velociraptor and Sauronitholestes.<br />
All <strong>sp</strong>ecimens of Saurornitholestes <strong>robustus</strong> come from the De-nazin<br />
Member of the Kirtland Formation, which is Kirtlandian age. The duration<br />
of the Kirtlandian is approximately 2 million years (74.8 to 72.8 Ma)<br />
based on recent correlations (see <strong>Sullivan</strong> and Lucas, 2006). The fossils<br />
from the De-na-zin Member can be more precisely dated at 73 Ma based<br />
on the 40 Ar/ 39 Ar dates of 73.04 Ma (Ash J) and 73.37 Ma (Ash H), pub-<br />
Barsbold, R. and Osmólska, H., 1999, The skull of Velociraptor (Theropoda) from<br />
the Late Cretaceous of Mongolia: Acta Palaeontologica Polonica, v. 44, p.189-<br />
219.<br />
Baszio, S., 1997a, Palaeoecology of dinosaur assemblages throughout the Late Cretaceous<br />
of South Alberta, Canada: Courier Forschungsinstitut Senckenberg, v.<br />
196, p. 1-31.<br />
Baszio, S., 1997b, Systematic Palaeontology isolated dinosaur teeth from the latest<br />
Cretaceous of south Alberta, Canada: Courier Forschungsinstitut Senckenberg,<br />
v. 196, p. 33-77.<br />
Brinkman, D., 2005, An illustrated guide to the vertebrate microfossils from the<br />
Dinosaur Park Formation: Unpublished guide, prepared for the Alberta Paleontological<br />
Society, Workshop on vertebrate microfossils (Jan. 26, 2002); revised<br />
April 3, 2005, 141 p.<br />
Currie, P.J., 1987, Theropods of the Judith River Formation of Dinosaur Provincial<br />
Park, Alberta, Canada; in Currie, P.J. and Koster, E.H., eds., Fourth Symposium<br />
on Mesozoic Terrestrial Ecosystems, Short Papers, v. 3: Drumheller, Tyrrell<br />
Museum of Palaeontology, p. 52-60.<br />
Currie, P.J., 2005, Theropoda, including birds; in Currie, P.J. and Koppelhus, E.B.,<br />
eds., Dinosaur Provincial Park: Bloomington and Indianapolis, Indiana University<br />
Press, p. 367-397.<br />
Currie, P.J., Rigby, J.K., Jr., and Sloan, R.E., 1990, Theropod teeth from the Judith<br />
River Formation of southern Alberta, Canada; in Carpenter, K. and Currie, P.J.,<br />
eds., Dinosaur systematics: Per<strong>sp</strong>ectives and approaches: Cambridge, Cambridge<br />
University Press, p.107-125.<br />
Currie, P.J., and Varricchio, D.J., 2004, A new dromaeosaurid from the Horseshoe<br />
Canyon Formation (Upper Cretaceous) of Alberta, Canada; in Currie, P.J.,<br />
Koppelhus, E.B., Shugar, M.A., and Wright, J.L., eds., 2004, Feathered Dragons.<br />
Studies on theTransition from Dinosaurs to Birds. Bloomington, Indiana<br />
University Press, p. 112-132.<br />
Eberth, D.A., 2005, The geology; in Currie, P.J. and Koppelhus, E.B., eds., Dinosaur<br />
Provincial Park: Bloomington and Indianapolis, Indiana University Press,<br />
p. 367-397.<br />
Fassett, J.E. and Steiner, M.B., 1997, Precise age of C33N-C32R magnetic-polarity<br />
reversal, San Juan Basin, New Mexico and Colorado: New Mexico Geological<br />
Society, Guidebook 48: 239-247.<br />
Kirkland, J.I., Gaston, R., and Burge, D., 1993, A large dromaeosaur (Theropoda)<br />
from the Lower Cretaceous of Eastern Utah: Hunteria, v. 2, 1-16.<br />
Norell, M.A. and Makovicky, P.J., 1999, Important features of the dromaeosaurid<br />
skeleton II: Information from newly collected <strong>sp</strong>ecimens of Velociraptor<br />
REFERENCES<br />
lished by Fassett and Steiner (1999), which bracket the upper and lower<br />
portions of the De-na-zin Member. By contrast, <strong>sp</strong>ecimens of<br />
Saurornitholestes (S. langstoni) from the Dinosaur Park and Two Medicine<br />
formations are considerably older, 75 Ma at the very least (Eberth,<br />
2005). It is not unreasonable to conclude that given the age difference, that<br />
the morphological differences between the <strong>sp</strong>ecies of Saurornitholestes<br />
are significant. De<strong>sp</strong>ite ongoing claims of endemic faunas and provincialism<br />
of Western Interior dinosaurs during the Late Cretaceous (see <strong>Sullivan</strong><br />
and Lucas, 2006, for citations), these differences are mostly due to temporal<br />
differences, not biogeographic ones. The cosmopolitan nature of the<br />
dinosaur faunas during the Late Cretaceous is borne out by the generic<br />
similarities of faunas from the north to the south. The coastal plain along<br />
the western margin of the Western Interior seaway provided a barrier free<br />
corridor for dinosaur di<strong>sp</strong>ersal.<br />
ACKNOWLEDGMENTS<br />
I thank James Gardner (Royal Tyrrell Museum of Palaeontology,<br />
Drumheller) and Phil J. Currie (University of Alberta, Edmonton) for access<br />
to comparative material; and Phil J. Currie and Don Brinkman (Royal<br />
Tyrrell Museum of Palaeontology, Drumheller) for discussions concerning<br />
Saurornitholestes. Don Brinkman and Spencer G. Lucas reviewed this<br />
contribution and I thank them for the comments and suggestions.<br />
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W., Dinosaur Park Symposium, p. 89-106.<br />
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and bird teeth from the Late Cretaceous (Late Campanian) Judith River Group,<br />
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theropod dinosaur from Mongolia: Paläontologische Zeitschrift, v. 51, p.173-<br />
184.<br />
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(Campanian) of Alberta: Zoological Journal of the Linnean Society, v. 62. p.<br />
381-400.<br />
<strong>Sullivan</strong>, R.M. and Lucas, S.G., 2000, First occurrence of Saurornitholestes<br />
(Theropoda: Dromaeosauridae) from the Upper Cretaceous of New Mexico:<br />
New Mexico Museum of Natural History and Science, Bulletin 17, p. 105-108.<br />
<strong>Sullivan</strong>, R.M. and Lucas, S.G., 2003, The Kirtlandian, a new land-vertebrate “age”<br />
for the Late Cretaceous of Western North America: New Mexico Geological<br />
Society, 54 th Field Conference, Guidebook, p. 369-377.<br />
<strong>Sullivan</strong>, R.M. and Lucas, S.G., 2006, The Kirtlandian land-vertebrate “age”—<br />
faunal composition, temporal position and biostratigraphic correlation in the<br />
nonmarine Upper Cretaceous of western North America: New Mexico Museum<br />
of Natural History and Science Bulletin, this volume.