Palaeogeography, Palaeoclimatology, Palaeoecology 230 (2006) 243#250 www.elsevier.com/locate/palaeo

Dinosaur sanctuary on the Chatham Islands, Southwest Pacific: First record of theropods from the K#T boundary Jeffrey D. Stilwell a,b,*, Christopher P. Consoli a, Rupert Sutherland c, Steven Salisbury d, Thomas H. Rich e, Patricia A . Vickers-Rich a, Philip J. Currie f, Graeme J. Wilson c a School of Geosciences, M onash University, Clayton Victoria 3800 A ustralia b Centre for E volutionary R esearch, A ustralian Museum, 6 College St., Sydney N SWA ustralia c Institute of Geological and Nuclear Sciences, P . O. Box 30-368, L ower H utt, New Zealand d School of Life S ciences, The University of Queensland, Brisbane Qld 4072 A ustralia e Museum Victoria, P . O. Box 666E, M elbourne Victoria 3001 A ustralia f Royal Tyrrell M useum of Palaeontology, D rumheller, Alberta TOJ OYO Canada Received 3 March 2005; received in revised form 21 July 2005; accepted 27 July 2005

Abstract

Cretaceous#Tertiary (K#T) boundary (ca. 65 Ma) sections on a Southwest Pacific island containing were unknown until March 2003 w hen theropod b ones w ere recovered from the Takatika Grit on the r emote Chatham Islands (latitude 448 S, longitude 1768 W), along the Chatham Rise. Tectonic and p alaeontologic evidence support the eastward extension of a ca. 900 km land b ridge that connected the islands to what is now New Zealand p rior to the K#T b oundary. The Chathams terrestrial fauna inhabited coastal, temperate environments along a low-lying, narrow, crustal extension ofthe New Zealand subcontinent, characterised by a tectonically dynamic, volcanic landscape w ith eroding hills (horsts) adjacent t o flood p lains and deltas, all sediments accumulating in grabens. This finger-like tract was b lanketed with a and clubmoss () dominated forest. The Chatham Islands region would h ave, along with New Zealand, provided a island sanctuary after separating from the margin ca. 80 Ma. D 2005 Elsevier B.V. All r ights reserved.

Keywords: #Tertiary b oundary; Dinosaur; ; Palaeobiogeography; Chatham Islands; New Zealand

1. Introduction Non-avian, terrestrial dinosaurs from the Mesozoic have not been p reviously reported from any oceanic islands in the Southwest Pacific, nor have any b irds. Terrestrial and marine deposits w ith latest Cretaceous

* Corresponding author. Tel.: +61 3 9905 1642; fax: +61 3 9905 4903. E-mail address: [email protected] (J.D. Stilwell).

003 1-0182/$ - see front matter D 2005 Elsevier B.V. All r ights r eserved. doi: 10.1016/j.palaeo.2005.07.017 dinosaur remains are, in fact, rare in b oth hemispheres and are p articularly so in the south. Only a handful of localities are known to yield depauperate latest Creta- ceous faunas in presumed marine deposits from Haw- ke#s Bay in N orth Island, New Zealand (Molnar, 198 1; Scarlett and Molnar, 1984; Wiffen and Molnar, 1988; Molnar and Wiffen, 1994; Wiffen, 1996; Long, 1998); southern Patagonia (see Coria, 1999; Luna et al., 2003); and the James Ross Island group (Gasparini et al., 1987; Hooker et al., 1991; Case et al., 2000; Rich et al., 1999) (James, Seymour and Vega islands) along the tip of A ntarctic Peninsula. This report of latest Creta-

244 J.D. Stilwell et al. / Palaeogeography, P alaeoclimatology, P alaeoecology 230 (2006) 243$250 northwestern corner (see arrow). ceous dinosaurs in the Takatika Grit on the r emote Chatham Islands (Fig. 1) indicates t hat n on-avian ther- opods and other groups did exist on an island sanctuary for millions of y ears following the separation of New Zealand and the islands from the Gondwana margin. The Chatham Islands today are the eroded remnants of isolated oceanic isles that consist of a main island, Chatham (Rekohu in local Moriori language), and smal- ler island, Pitt (Rangiauria), located 22 km t o the southeast along with much smaller volcanic islets, reefs and rocks (Campbell et al., 1993). These islands are emergent, spectacular, low-lying, and steeply cliffed edifices at the eastern end of the Chatham Rise, a linear feature extending due east more than 1000 km from Canterbury Province of South Island, New Zealand. A major submarine extension of the New Zealand micro- continent, the Chatham Rise encompasses some 200,000 km ( and Herzer, 1993). Fortunately, the islands preserve an important Cretaceous to Recent record for this p art of the Pacific, w hich enables a detailed reconstruction of its geologic history over the last 100 Ma. The p refix for v ertebrate material GNS is an abbre- viation for the Institute of Geological and Nuclear Sciences, Lower Hutt, New Zealand. 2. Geology

The geologic history of the Chatham Islands extends back into the late with exposed Torlesse ter- rane basement rocks equivalent to those on the New Zealand mainland (Hay et al., 1970; Campbell et al., 1993), consisting of greywacke, argillite and associated metamorphics, including the Chatham Schist. Resting unconformably on basement rocks is a thick sequence of Upper Cretaceous#Paleogene aged transgressive p ack- ets, derived from a p rincipal divergent tectonic p hase reflecting the final fragmentation of the Gondwana. This extensional regime resulted in substantial, east#west trending half-grabens and volcanism in the Bounty Trough#Chatham Rise region (Campbell et al., 1993). The Takatika Grit is a distinctive, thin, fossiliferous rock u nit (maximum thickness 10 m; Fig. 2.), a glau- conitic quartzofeldspathic grit made u p of sandstone and siltstones with a volcanogenic component, resting nonconformably on basement schist (Hay et al., 1970; Campbell et al., 1993). It hosts several b one horizons (Nodular Phosphorite-Bone Package (NPB)) spanning 2 km of wave-cut platform (between 43.7508S, 176.6678W and 43.7438S, 176.6838W) and p reserves skeletal elements, mainly of marine reptiles. Based on associated content, the unit is lower-Hau- with associated (note: position of catalogued bones) and phosphorite nodules (see Legend).

J.D. Stilwell et al. / Palaeogeography, P alaeoclimatology, P alaeoecology 230 (2006) 243#250 245 murian to mid-Teurian in age; the microfauna is dom- inated b y the Early Haumurian , corre- lated to the Satyrodinium haumuriense Zone (Wilson, 1982; Wilson et al., 2005) (Haumurian#Teurian Stage, mid- to early ). The grit may rep- resent, in p art, an allochthonous accumulation of reworked u ppermost Cretaceous sediments and fossils that were deposited p ost K#T b oundary, in the Danian. However, the mixed latest Cretaceous#earliest Tertiary age of associated indicate that the K#T boundary may b e present b elow the exposed bone-rich beds. Vertebrate remains in the marine sediments include a theropod centrum (Fig. 3 A#B), the proximal p edal phalanx II-1 or III-1 ofa theropod (Fig. 3 F#G), as well as the probable theropod proximal head ofa tibia (Fig. 3 D# E) (all three elements from N PB). In slightly younger sediments, Greensand Package II, a theropod manual phalanx (Fig. 3 K#O) and manual ungual (Fig. 3 H#J) have b een recovered. These latter two b ones w ere col- lected stratigraphically above the major Cretaceous fau- nal assemblage in greensands together with rich assemblages clearly of early-to mid-Teur- ian (Danian) age. Interestingly, these bones exhibit no signs of reworking. The latest Cretaceous theropod frag- ments were evidently transported into the shallow ma- rine environment from adjacent terrestrial habitats and display differential p reservation and wear. The b ones are associated with authigenic p hosphorite nodules of p eb- ble to boulder size and some of the nodules actually contain b one.

3. Dinosaurian elements

GNS CD 578: a partial centrum, Theropoda; (Fig. 3 A#B). Measurements: Width (max)= 89.3 mm. Locality and horizon: N orthwest Chatham Island, New Zealand; Tioriori Group (Cretaceous#Tertiary); Taka- tika Grit (Lower Haumurian#Lower Danian); NPB in association with theropod dinosaur and marine reptile remains and p hosphorites nodules. Description: Theropod centrum with no trace of a neural arch or transverse p rocesses. Its p latycoelous articular surface could b e anterior or p osterior. The centrum is constricted at the anteroposterior midpoint as compared with its diameter at the articular end. There are no traces of a p neumatic fossa or p neumatic foram- ina p reserved on lateral side of the centrum. The cen- trum is hollow but no internal p artitions are p reserved, but given the state of preservation of this specimen, it is unlikely that such internal p artitions w ould have b een retained. Remarks: The centrum was from a medium-sized the- ropod about 4 m long and 3 m high, but its position along the vertebral column and orientation is difficult to determine due to its incomplete nature. The p latycoe- lous articular surface and constriction of the midpoint indicates that it is not an axial element of a marine reptile and the theropod caudal vertebra reported from Mangahouanga Stream, Hawke$s Bay, New Zealand bears no resemblance to the Chatham Island centrum (Molnar, 198 1). GNS CD 579: manual ungual, Theropoda; (Fig. 3 H#J). Measurements: Length= 15.25 mm, width=4.5 mm, height= 6.6 mm. Locality and horizon: N orthwest Chatham Island, New Zealand; Tioriori Group (Cretaceous#Tertiary); Taka- tika Grit (Lower Haumurian#Lower Danian); Green- sand Package II in association w ith shark teeth and Danian microfossils. Description: A theropod m anual u ngual lacking the tip and distal-most portion of the dorsal surface; it is mediolaterally compressed, slightly asymmetric, and gently recurved. The weak flexor tubercle appears to be distally displaced. The p halangeal articular surface has a gently raised, p almodosally aligned ridge, sug- gestive of a ginglymoid j oint. Proximal to the flexor tubercle, the p roximal-most third of the p almar surface is flat in lateral/medial aspect. There are b road, shallow grooves for claw sheaths that approach the dorsal sur- face p roximally. The groove on the assumed lateral surface is much more distinctive and distally ap- proaches the dorsal surface slightly more than the lat- eral side. Remarks: The manual u ngual is comparatively smaller than most theropod claws but has several diagnostic features characteristic of theropods. It is likely to b e non-avian because it is not strongly recurved, charac- teristic of b irds and unguals of dromaeosaurs and ovir- aptorosaurs are generally larger and much more curved, lacking. The mediolateral flattening and presence of a flexor tubercle implies that this is not a pes ungual. The distal displacement of the weak flexor tubercle and lack of curvature is r eminiscent of some ornithomimosaur- ids, such as Ornithomimus edmontnensis from the upper Campanian#lower Judith River Formation of Alberta (Sternberg, 1933), and Harpymi- mus okadnikovi from the A ptian#Albian of Mongolia (Barsbold and Perle, 1984). The presence of an ornitho- mimosaurid in the Early Cretaceous of southern Victo- ria has b een suggested b y Rich and Vickers-Rich (1994), b ased on the discovery of two femora, which they referred to Timimus hermani. The slight curvature, 246 J.D. Stilwell et al. / Palaeogeography, P alaeoclimatology, P alaeoecology 230 (2006) 243#250

Fig. 3. A#C Partial theropod centrum (GNS CD578), considerably constricted at the anterior midpoint in comparison with its diameter at the articular end; belongs to medium-sized theropod approximately 4 m long and 3 m high, probably distinct from the Mangahouanga Stream, New Zealand theropod. D#E Probable theropod proximal head of tibia (GNS CD 584; Width=93 mm). F#G Probable theropod proximal p edal phalanx II-1 or III-1 (GNS CD 583) from a large n eotheropodan theropod similar to Carnotaurus, possibly a b asal allosauroid, estimated to b e u p to 10 m long. H#J Theropod manual u ngual (GNS CD579) that is mediolaterally compressed and not recurved w ith w ell-defined lateral and medial grooves on the sides; note that the lateral groove on the medial surface merges closer t o the dorsal surface of the bone distally. K#O Manual phalanx (GNS CD580) identified as a proximal manual phalanx of a medium-sized theropod with a marked, inter-phalangeal articular surface. J.D. Stilwell et al. / Palaeogeography, P alaeoclimatology, P alaeoecology 230 (2006) 243#250 247 compressed n ature and distinct grooves determined that this specimen is most likely derived from a manus of a theropod dinosaur, and although a single p artially p re- served claw is not substantial enough for confirmation, it is most reminiscent of ornithomimosaurs. GNS CD 580: manual phalanx, Theropoda; (Fig. 3 K#O) Measurements: Length=33.6 mm, width (distal)= 9.05 mm, height (proximal)=9.8 mm, height (dis- tal)= 7.7 mm. Locality and horizon: N orthwest Chatham Island, New Zealand; Tioriori Group (Cretaceous#Tertiary); Takatika Grit (Lower Haumurian#Lower Danian); Greensand Package II, in association w ith fossil shark teeth and Danian microfossils. Description: Theropod manual phalanx, lacking some of the proximal articular surface, and the medial side. Overall, it is elongate, slender and is p almodosally compressed w ith proximal and distal expansion, the shaft tapering distally. The interphalangeal surface is smooth and deeply concave, and has an obtuse surface, which lacks a significant p almodosally aligned r idge; suggesting it is a proximal digit. The distal articular surface is well-developed with a deep central vertical groove formed by the condyles. These are relatively large and almost circular, being of similar width to the proximal articular surface, the lateral side is b etter preserved. Condyles have deep subrounded p its, most prominent on the lateral surface, where the pit deepens almost to the shaft. The asymmetry of this distal end suggests that it is not a central axis element but either in the medial or lateral row of digits. The presence of attachments for extensor ligaments rules it out as a metapodial. Remarks: The phalanx is comparable to those of many theropods (e.g. troodontids, oviraptorosaurids, ornitho- mimids), but its generalized nature p recludes an assign- ment to recognized genera. The morphology of the Chatham phalanx is also reminiscent of some pes p ha- langes of b irds. A volant b ird element was p reviously reported from the Takatika Grit deposit of the Chatham Islands. Fordyce (1991) briefly mentions an u nde- scribed, u nprepared, p neumatic, proximal end of a ra- dius of Early Paleocene age, and thus the p robability of an avifauna p opulation present on the Chatham Islands in the Earliest Cenozoic, and that the manual phalange is comparable to a b ird cannot b e dismissed. N everthe- less, the shape, size and deep p its on the medial and lateral sides of the distal end mark it as a medial or proximal manual phalanx of a modest-sized animal. Unsurprisingly, it does not resemble the -aged manual phalange from N orth Island, lacking a triangu- lar shaft and a distal articular face which is dorsally cylindrical (Molnar et al., 1998). The phalanx of either a t errestrial b ird or theropod dinosaur from the Late Cretaceous Mangahouanga Stream, N orth Island is comparable in size and distal articular morphology to the Chatham specimen. The shaft, however, is high and the proximal articular surface acutely angled, in con- trast to the compressed nature of the shaft and w ide angle of the interphalangeal articular surface of the Chatham manual p halange. GNS CD 583: proximal p edal phalanx II-1 or III-1, Theropoda; (Fig. 3 F#G) Measurements: Length= 185 mm, width= 102 mm. Locality and horizon: N orthwest Chatham Island, New Zealand; Tioriori Group (Cretaceous#Tertiary); Taka- tika Grit (Lower Haumurian#Lower Danian); NPB in association with theropod dinosaur and marine reptile remains and p hosphorites nodules. Description: This specimen represents a proximal pedal phalanx II-1 or III-1 of a large theropod esti- mated to b e 8#10 m in total length. Most of the dorsal surface of the p halangeal b ase is p reserved, but much of the p halangeal head is missing. Pre- served as well is the shallow depression on the lateral/medial surface of the head, u sed for the attach- ment of the collateral ligament and the f lexor digitalis brevis. Remarks: The shaft is decidedly more slender than that of similar-sized allosaurids and tyrannosaurids, indicat- ing that although the animal was very large, the pes was quite slender. Gracile, elongated hind limbs are charac- teristic of some abelisaurids, such as Carnotaurus. Thus, the Chatham Island p edal phalanx seemingly belonged to a large neotheropodan theropod, perhaps a b asal allosauroid.

4. Tectonic and palaeobiogeographic history

The position of the Chatham Rise has changed little relative to New Zealand or the rest of the Pacific Plate since Late Cretaceous time, and the region has slowly subsided to its present water depth of 200#500 m (Wood and Herzer, 1993). On the other hand, the Chatham Rise has changed position relative to Austra- lia, Antarctica, and western New Zealand, determined by r econstructions of ocean crust in the Southern Ocean and Tasman Sea (Cande et al., 1995; Sutherland, 1999; M u¨ller et al., 2000). The Pacific Plate is mainly sub- merged and, hence, has a p oorly defined apparent p olar wander p ath (APWP) for Late Cretaceous time. In contrast, A ustralian data define a reliable APWP, but require rotation into a Pacific Plate frame of reference

248 J.D. Stilwell et al. / Palaeogeography, P alaeoclimatology, P alaeoecology 230 (2006) 243#250 before they can b e compared with data from the Cha- tham Islands. Fortunately, the islands contain numerous basaltic volcanic rocks of Late Cretaceous age. Radio- metric ages from 18 sites are in the range 70.3 F 1.4 t o 81.4 F 2.0 Ma, and p aleomagnetic measurements indi- cate all sites have normal polarity (Grindley et al., 1977; Campbell et al., 1993). The data indicate a prolonged p eriod of volcanic activity during chrons 32#33 (71.7#79 Ma), during w hich time only 0.5 Ma is k nown to have had negative polarity (Cande and Kent, 1995). Paleomagnetic data from volcanic rocks at 49 sites indicate that the Chatham Rise had a similar east#west orientation in Late Cretaceous time and had a mean latitude during the p eriod 71#79 Ma of 63 F 68. Paleo- magnetic data from Detroit Seamount in the N orth Pacific y ield an 8 1 Ma Pacific Plate apparent p ole position, w hich independently confirms that the Cha- tham Rise was oriented east#west, a p aleolatitude for the islands of 59 F 128 (Cottrell and Tarduno, 2003). New Zealand separated from A ntarctica during chron 33r (83#79 Ma) and magnetic anomaly 33y (chron 33y is equivalent to 73.6 Ma) is clearly identified adjacent to the New Zealand continental margin (Cande et al., 1995; Sutherland, 1999). A linear positive magnetic anomaly immediately south of the islands may have formed as an isolated spreading r idge segment that became inactive during chron 33 (Sutherland, 1999). The close spatial relationship between scattered sea- mounts associated w ith this ocean crust and Chatham Islands volcanics is consistent with this interpretation. Reconstructions of the islands for times younger than chron 33y are b ased u pon reliably identified magnetic anomalies south of New Zealand, and the b ase of the continental slope can fit at an inferred age of 83 Ma. If so, an apparent p olar w ander p ath for the islands can b e constructed from Australian p aleomagnetic data (Veevers and Li, 1991), continental margin fits (above), magnetic anomalies in the Pacific and south- east Indian Ocean (Cande et al., 1995; Cande and Kent, 1995), and b y making some r easonable as- sumptions about internal deformation of Antarctica. The resulting reconstructions confirm that the Cha- tham Islands must have moved northwards from a latitude of about 708S at 83 Ma to a latitude of about 548S at 65 Ma, at a rate of about 0.98/m.y. Thus, the islands have b een p art of the New Zealand continent since Late Cretaceous time. The Chatham Islands p aleo- latitude of 63 F 68, determined from p aleomagnetic measurement of volcanic rocks data at 71#79 Ma, is consistent w ith k nown p aleomagnetic data. The Cha- tham Rise was probably close t o sea level for much of the Late Cretaceous w ith the emergent tracts hosting terrestrial dinosaurs.Sediments of the Takatika Grit re- flect b oth a terrigenous p rovenance and authigenic input of glaucony, deposited in a shallow marine setting. Their origin relates to divergent p late motions separating the New Zealand#Chatham Islands b lock from Antarctica (Marie Byrd Land) about 80 Ma. So, the Chatham Islands have b een isolated oceanic islands since the beginning of the Cenozoic, given the slow subsidence of the Chatham Rise since the late Maastrichtian. Con- comitant w ith this p late divergence, a large, (20#25 km wide), alkaline basalt, intraplate stratovolcano erupted on southern half of Chatham Island and Pitt Strait (Campbell et al., 1993), followed subsequently by the infilling of half-grabens that accumulated t hick volcani- clastic sequences, including the Takatika Grit. Terrige- nous sequences w ere p unctuated b y episodes of localized basaltic volcanism (Campbell et al., 1993). Towards the close of the Cretaceous, grabens were still b eing infilled, with the formation of flood p lains and deltas, whereas horsts were eroded to hills of rela- tively low topographic relief (Wood et al., 1989), b lan- keted b y a diverse, floral assemblage (Fig. 4). The Cretaceous flora of the Chatham Islands was dominated b y , remnant of austral, high-latitu- dinal forests and including southern beech, araucarians, podocarps, Proteaceae and clubmosses (Lycopodiop- sida), b ased on the microflora from the Tupuangi For- mation, Kahuitara Tuff and Takatika Grit (Wilson, 1982; Campbell et al., 1993; Wilson et al., 2005; Con- soli et al., in review). Macrofloral remains have also been described from the Chatham Islands, a cone of Araucaria from the Takatika Grit, and Mataia and P odocarpus as well as angiosperm were reported b y Campbell et al. (1993) from the Tupuangi Formation.

5. Discussion

Although our k nowledge of the geology of the Cha- tham Islands has improved markedly over the last decade, relatively little is known of the terrestrial (Campbell et al., 1993) and marine (Stilwell, 1997, 1998) b iota t hat inhabited the region after it separated from the A ntarctic margin (see reconstructions b y Huber and Watkins, 1992; Lawver et al., 1992). The Takatika Grit is p articularly significant in allowing the reconstruction of the palaeoenvironment and b iota for the latest Cretaceous, j ust p rior to the K#T b oundary, and into the early Cenozoic, representing the dawn of a bbrave new worldQ dominated b y b irds and mammals. This was a time of intense, global change. Furthermore, J.D. Stilwell et al. / Palaeogeography, P alaeoclimatology, P alaeoecology 230 (2006) 243#250 249

topography dominated the landmass along the Chatham Rise w hich comprised low-lying hills and broad flood-plains, surrounded by a shallow sea to north and south. Remnant volcanoes p ersisted forming seamounts, hills and islands. Revised from Wood et al. (1989). the Takatika Grit consists of three horizons containing bones, and these increase in abundance u p-section. Whether or not these mass b one accumulations of reptiles represent individual m ass mortality events remains to b e determined. The b one horizons of terres- trial and marine r eptiles, sharks, teleost fish, ammonites and n autiloids (Consoli and Stilwell, 2005), b ivalves, gastropods, , and p lant matter, including exqui- sitely p reserved cones are laterally p ersistent, u nique in the Southern Hemisphere for this time. Until 1979 w hen the first vertebra of a Late Creta- ceous dinosaur was recovered from New Zealand, sev- eral hypotheses were advanced to explain their absence in the widespread Cretaceous terrestrial and marine sequences on t his small m icrocontinent. To date, sparse terrestrial and flying reptile b ones have been r ecovered from the slightly older to coeval Mangahouanga Stream site, including a t ail vertebra (Molnar, 198 1), p edal phalanx from a terrestrial b ird or theropod (Scarlett and Molnar, 1984) and manual phalange from a thero- pod (Molnar et al., 1998), a rib fragment of a small sauropod (Molnar and Wiffen, 1994), and a rib and tail vertebrae of an ankylosaur (Molnar and Wiffen, 1994). Discovery of dinosaurs on the Chatham Islands demon- strates that these successful reptiles inhabited the most isolated p arts of the Southern Hemisphere, and dino- saurs w ere definitely present at high-latitudes in the temperate to warm, low-lying coastal habitats n early 1000 km east of New Zealand, an isolated and unique, K#T boundary environment.

Acknowledgments

We w ish t o give hearty thanks to Terry and Donna Tuanui of Chatham Island for access to the Takatika Grit fossil localities, their u nwavering enthusiasm through- out this study, and for their logistical support, also to A. Grebneff, A. Morrell and W. Zinsmeister for assistance in the field. E. Nicholls (deceased), L. Chiappe and Y. Kobayashi provided comments on dinosaur identifica- tions. S. Morton, D. Gelt, D. Pickering and L. Kool provided valuable expertise throughout this investiga- tion. W. Zinsmeister, J. Long, J. Snelleman, and F. Koning greatly improved the manuscript and w e appre- ciate their input. This research was supported b y an Australian Research Council Fellowship to JDS, Com- mittee for Research and Exploration of the National Geographic Society Grant, James Cook U niversity (Townsville, Queensland) Merit Research Grant, and Monash University, (Melbourne, Victoria), Support Grant (all t o JDS).

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