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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 />

mongoliensis: American Museum Novitates, no. 3282, 28 p.<br />

Osborn, H.F., 1924, Three new Theropoda, Protoceratops zone, central Monogolia:<br />

American Museum Novitates, no. 144, 12 p.<br />

Peng, J., Russell, A.P., and Brinkman, D.B., 2001, Vertebrate microsite assemblages<br />

(exclusive of mammals) from the Foremost and Oldman formations of the Judith<br />

River Group (Campanian) of southeastern Alberta: An illustrated guide: The<br />

Provincial Museum of Alberta, Natural History Occasional Paper, no. 25, 54 p.<br />

Rauhut, O.W.M. and Werner, C., 1995, First record of the family Dromaeosauridae<br />

(Dinosauria: Theropoda) in the Cretaceous of Gondwana (Wadi Milk Formation,<br />

northern Sudan). Paläontologische Zeitschrift, v. 69, p. 475-489.<br />

Rowe, T., Cifelli, R.L., Lehman, T.M. and Weil, A., 1992, The Campanian Terlingua<br />

local fauna, with a summary of other vertebrates from the Aguja Formation,<br />

Trans-Pecos Texas: Journal of Vertebrate Paleontology, v. 12, p. 472-493. Sankey,<br />

J.T., 2003, New theropod and bird teeth from the Late Cretaceous (Maastrichtian)<br />

Hell Creek and Lance Formations (abs): Journal of Vertebrate Paleontology, v.<br />

23, p. 93A.<br />

Sankey, J.T., 2005, Late Cretaceous vertebrate paleoecology of the Big Bend National<br />

Park, Texas; in Braman, D.R., Therrien, F., Koppelhaus, E.B. and Taylor,<br />

W., Dinosaur Park Symposium, p. 89-106.<br />

Sankey, J.T., Brinkman, D.B., Guenther, M., and Currie, P.J., 2002, Small theropod<br />

and bird teeth from the Late Cretaceous (Late Campanian) Judith River Group,<br />

Alberta: Journal of Paleontology, v. 76, p. 751-763.<br />

Sues, H.-D., 1977, The skull of Velociraptor monogoliensis, a small Cretaceous<br />

theropod dinosaur from Mongolia: Paläontologische Zeitschrift, v. 51, p.173-<br />

184.<br />

Sues, H.-D., 1978, A new small theropod dinosaur from the Judith River Formation<br />

(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.

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