Reptile Remains from Tiga (Tokanod), Loyalty Islands, New Caledonia Author(s): Juan D. Daza, Aaron M. Bauer, Christophe Sand, Ian Lilley, Thomas A. Wake and Frédérique Valentin Source: Pacific Science, 69(4):531-557. Published By: University of Hawai'i Press DOI: http://dx.doi.org/10.2984/69.4.8 URL: http://www.bioone.org/doi/full/10.2984/69.4.8

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Reptile Remains from Tiga (Tokanod), Loyalty Islands, New Caledonia1

Juan D. Daza ,2,3 Aaron M. Bauer ,2,8 Christophe Sand , 4 Ian Lilley ,5 Thomas A. Wake, 6 and Fr é d é rique Valentin7

Abstract: Archaeological exca va tions on Tiga provide the fi rst vouchered her- peto log i cal re cords for this small island between Lifou and Mar é in the Loyalty Islands. Eighty- three skel etal el ements from four sites yielded mate rial assign - able to skinks (Emoia loyaltiensis, Lioscincus nigrofasciolatus ), geckos (Bavayia crassi- collis, B . sp., Gehyra georgpotthasti, Nactus pelagicus ), and a boid snake ( Candoia bibroni ) all known from elsewhere in the Loyalties, as well as unde ter mined mate rial consis tent with these and other Loyalties liz ards. Diagnostic features of geckos versus skinks for ele ments commonly re cov ered from ar chaeo log i cal sites and from owl pellets are discussed. Gehyra georgpotthasti has a lim ited dis tribu- tion in the Loyalties and its occur rence on Tiga clarifi es its range. The boid snake is the only rep tile likely to have been harvested by hu man inhab i tants of Tiga. The presence of gekkonid geckos in pre- Euro pean times is con fi rmed and con trasts with the sit u a tion of Grande Terre fos sil sites, where only diplodac- tylid geckos have been recov ered. Although it is antic ipated that all spe cies re- covered from archae o logi cal sites are still present on the island, a modern herpetofaunal survey is needed.

T he Loyalty Islands lie on the Norfolk Rise (2 km 2 ), approx ima tely 160 km southeast of the in a line roughly par allel to the long axis of the main chain. All of the is lands are rela tively low Grande Terre of New Caledonia, sepa rated lying. Their raised rims are uplifted coral from it by about 110 km across the Loyalty reefs, whereas the lower el e vation in te rior re- Basin, which has a maxi mum depth of approx- gions are uplifted la goons or reef fl ats, proba- i mately 2,000 m. The Loyalties ( Figure 1) ex- bly emer gent only since the Pleis tocene (Paris tend from Ouvé a (160 km2 ), at the northwest 1981). The highest point (138 m) in the chain is ex tent of the chain, through Lifou (1,150 km 2 ), on Mar é , where basal tic outcrops rep re sent the Tiga (10 km2 ), and a se ries of smaller islands, emergent portions of the vol ca nic base un der- Maré (650 km2 ), and on to Ile Walpole lying the Miocene reef (Paris 1981, Kroenke

1 Funding for this research was provided by the Loy- 3 Current address: Department of Biological Sciences, alty Islands Province, the Government of New Caledo- Sam Houston State University, Huntsville, Texas 77341. nia, the University of Queensland (Australia), the Centre 4 Institut d ’ arché ologie de la Nouvelle -Cal é donie et National de la Recherche Scientifi que (CNRS), the Ger- du Pacifi que, 65, Rue Teyssandier de Laubar è de, BP: ald M. Lemole Endowed Chair funds (Villanova Uni- 11423, 98802 Noumé a Cedex, Nouvelle- Cal é donie, versity), and Grant Nos. DEB 0844523 and DEB 1019443 New Caledonia. from the U.S. National Science Foundation. Manuscript 5 Aboriginal and Torres Strait Islander Studies Unit, accepted 31 March 2015. The University of Queensland, Q 4072, Brisbane, Aus- 2 Department of Biology, Villanova University, tralia. 800 Lancaster Avenue, Villanova, Pennsylvania 19085. 6 University of California— Los Angeles Cotsen In- stitute of Archaeology, Box 951510, A210 Fowler, Los Angeles, California 90095- 1510. 7 UMR 7041– ArScAn, CNRS, Maison de Pacifi c Science (2015), vol. 69, no. 4:531–557 l ’Arch é ologie et de l ’Ethnologie, 21 all é e de l ’Universit é , doi:10.2984/69.4.8 F -92023 Nanterre cedex, France. © 2015 by University of Hawai‘i Press 8 Corresponding author (e- mail: aaron.bauer@ All rights reserved villanova.edu).

531 532 PACIFIC SCIENCE · Oc to ber 2015

Figure 1. Location of Tiga Island in the Loyalty Islands archi pel a go.

1984). Rainfall is rela tively low on the Loy- Grande Terre, vis ited Mar é and northern alties, ranging from 1,700 mm per annum in Lifou, and stopped briefl y on Ouvé a. Roux’ s north ern Lifou to un der 1,300 mm on Ouvé a (1913) resulting monograph summa rized the (Sautter 1981), and the vege ta tion is highly de- entire terres trial and marine herpetofauna of pauper ate rel a tive to mainland New Caledo- New Caledonia and the Loyalty Islands. Sub- nia, with < 400 plant species (Jaffré 1993), sequently, very little herpe to log i cal research consisting largely of coastal strand veg eta tion has been conducted (mate rial evalu ated by sim ilar to that on the mainland and inland Sadlier and Bauer [1997]). Virtually all re- hu mid forests (Dä niker 1931, Schmid 1981). search has been conducted on the three main The herpetofauna of the Loyalties has been is lands of the group. Very recent ly, the larg- rel atively little - stud ied, al though its over all est gecko in the Loyalties, long con sidered spe cies compo sition is believed to be well con spe cifi c with Gehyra vorax Girard, 1858, of known (Sadlier and Bauer 1997, Bauer and Fiji, has been de scribed as new based partly on Sadlier 2000). The nineteenth - century Mar- mate rial from Î le Dudun, a small satel lite off ist mission ary Xavier Montrouzier (1820– of north ern Maré (Flecks et al. 2012), but 1897) fi rst noted the pres ence of the Loyalty other small islands, most nota bly Tiga, have Island boa, which he re ferred to as Boa austra- not been explored herpetologically and are lis (Montrouzier 1860). Bavay (1869), who represented by no mate rial in museum collec- catalogued the fauna of New Caledonia, de- tions. Interestingly, howev er, both Tiga and scribing many of the endemic species, was remote Walpole have been sites of some paleo- based on the Grande Terre, but he did ob tain herpetological work on the extinct meiola- snakes from E. Dubois, who collected in the niid turtles (Gaffney et al. 1984, Balouet 1991). Loyalty Islands. In 1911– 1912, Fritz Sarasin Although the herpetofaunal tally of the and Jean Roux traveled through much of the Grande Terre con tin ues to grow as more and Reptile Remains from Tiga, Loyalty Islands · Daza et al. 533 more microendemic geckos and skinks are larg est island in the Loyalties, but it also lies discov ered, chiefl y at higher ele vations (Bauer im medi ately north of Mar é and may be ex- and Sadlier 2000, Bauer and Jackman 2006, pected to help resolve the distri bu tional lim- Sadlier 2010), the herpetofaunal inven tory of its of the Maré area en dem ics. the Loyalty Islands has remained quite sta ble We here re port on the herpe tolog i cal fi nd- and has ac tually decreased. Sadlier and Bauer ings of an archeological study on Tiga car- (1997) and Bauer and Sadlier (2000) reported ried out by the Institut d’ arch é ologie de la 20 terres trial squamates. One of these has Nouvelle - Calé donie et du Pacifi que in collab- since been recog nized as synon y mous with o ration with the University of Queensland another on the list, leaving only 19 species. Six and the Centre National de la Recherche Sci- of these are wide spread throughout much of entifi que in Paris. The study focused on the the Pacifi c, includ ing New Caledonia, and inte gra tion between indig e nous Ka nak several of these are known or suspected to knowl edge about Tiga’ s past and clas sic sur- have reached the re gion through human vey /ex ca va tion studies around the island. The agen cy. Another six are regional endem ics, re- fi rst phase of the fi eldwork concen trated on stricted to New Caledonia s.l. (one, Caledonis- re cording oral tradi tions and lo cat ing the tra- cincus atropunctatus [Roux, 1913], also occurs di tional sites that local people were ready to in southern Vanuatu) and three spe cies— share with us. In the second phase, a classic Gehyra georgpotthasti Flecks, Schmitz, Bö hme, archae o log ical study was conducted. Over 160 Henkel, & Ineich, 2012; Emoia cyanura (Les- differ ent sites were recorded, from rock- son, 1826); and Candoia bibroni (Dum é ril & shel ters to walled enclo sures, from tradi- Bibron, 1844)— oc cur else where in the west- tional burial grounds to mythi cal sites, and ern Pacifi c but are absent from the Grande from horti cultural sites to caves where peo- Terre. The remaining four spe cies appear to ple col lected fresh wa ter. An unex pected be Loyalties endem ics. Two of these have long num ber of rock art sites have been recorded. been recog nized as such, Emoia loyaltiensis Excavations done in differ ent types of sites (Roux, 1913) and Ramphotyphlops willeyi (Bou- have shown that Tiga was fi rst settled by hu- lenger, 1900). Recent molec ular phylo genetic mans around 2,950 yr ago on the low coastal research (Bauer and Jackman 2006), howev er, plain facing the northwest ern lagoon of the has also revealed that the diplodactylid gecko island. A rapid movement in land, con cen trat- Bavayia crassicollis Roux, 1913 (pre viously be- ing on the fertile central plateau and sur- lieved to occur on other islands in the New rounding ar eas, took place in the centu ries Caledonian region) is a Loyalties endemic and follow ing fi rst set tlement. A movement to that speci mens of B. sauvagii (Boulenger, 1883), marginal zones of the island is iden tifi able from Maré ac tually repre sent an undescribed later in the se quence. This se quence is char- endemic species. acter ized by fl uc tuat ing ex changes with sur- Although the reptile species com po sition of rounding is lands, demon strated, for exam ple, the Loyalties may be con sidered well estab- by the pres ence of ce ramics and stone re- lished, the same cannot be said of intra - and mains from the Grande Terre. But the main inter is land dis tribu tion patterns. A few spe cies dy nam ics in terms of in tensi fi ca tion of pro- ap pear to be restricted to Maré (Bavayia aff. duction and cultural changes were inter nal, sauvagii ) or to Mar é and its out liers (Gehyra with oral tradi tions showing the late po liti- georgpotthasti, al though there is a single vouch- cal infl uence of in com ing groups from sur- ered record from Lifou— ZFMK 70455), but rounding is lands (Sand et al. 2010). sampling has been insuf fi cient to confi rm this We here report on the reptile re mains or to de termine whether other species with found in asso ci ation with the archeological patchy dis tri bu tions may, in fact, be ubiqui- sites inves ti gated. Because there are no vouch- tous. Any addi tional sampling in the region ered mod ern speci mens from Tiga, this re- would be desir able, but Tiga stands out as an port consti tutes the fi rst list of the reptiles of espe cially glar ing gap. Not only is it the fourth the island. 534 PACIFIC SCIENCE · Oc to ber 2015

tely 65 cm, with complex stratigraphies indi- mate ri als and methods cat ing multi ple reuses of the site. The oldest date comes from test- pit LTD -18C, with a Sampling and Excavation of the Sites result of 2,530 ± 50 yr B.P. (Beta- 227 - 996) at a The collec tion stud ied here comes from four depth of 55– 65 cm. The other dates indi cate a 1 m2 ex ca va tion pits, located at three dif fer ent con tinu ous occu pation of Dré n é , but with sites on Tiga (see Sand et al. [2010] for a sum- the main use of the rock- shel ter occur ring mary). The let ter af ter each site number des- between the end of the fi rst millen nium B.C. igna tes a pit. Excavations were made by hand, and the fi rst mil len nium A.D. mostly in arti fi cial 5 cm or 10 cm spits. All the soil was sieved in water through 2.8 mm mesh Morphological Comparisons and the remaining sed i ment sorted by hand. Site LTD- 14, called Hari, is a small rock - Because our mate rial is < 3,000 yr old, we as- shelter lo cated on the west coast of Tiga, about sume that it repre sents a fauna sim ilar to that 50 m inland and approx i mately 2 m above sea of the Loyalty Islands today. Megafaunal lev el. The exca va tion reached the be ginning rep tiles are known to have sur vived until rel- of human occu pa tion at 100 cm deep, with an a tively recent times and undescribed, ap par- initial date of use of 1,630 ± 40 yr B.P. (Beta- ently extinct species of lizards have been 227979). A to tal of four other dates range from iden ti fi ed in main land New Caledonian sites 650 ± 40 yr B.P. (Beta- 227977) at 70– 75 cm, to of com para ble age (Anderson 1925, Buffetaut 240 ± 40 yr B.P. (Beta- 227972) at 15 cm, show- 1983, Balouet and Buffetaut 1987, Balouet ing regu lar use of the site mainly dur ing the 1989, 1991, Balouet and Olson 1989, Bayless mid dle of the sec ond mil len nium A.D. and Naclerio 1993, Mead et al. 2002, Ander- Site LTD- 02A, in the cen ter of the pres- son et. al. 2010, Kennedy 2011). However, we ent- day village of Tiga, is a nearly fi lled rock- found no ev i dence of extinct line ages in our shelter on the northwest coast of the island, mate ri al. Material from Tiga was compared about 70 m from the present shore. Excava- with refer ence speci mens representing species tions reached 190 cm deep and demon strate widely distrib uted in the Pacifi c as well as that until possi bly the sec ond half of the fi rst those known to be part of the ex tant fauna of mil len nium A.D., the site opened di rectly to the Loyalty Islands. In addi tion, speci mens the sea. It was only with the building of a belong ing to some taxa documented only raised sand dune and the move ment of the from the Grande Terre of New Caledonia coast to ward the west that the shelter be came were also com pared (see Bauer and Sadlier usable to hu mans. In its fi rst stage, between 2000) (Appendix 1). The New Caledonian re- the end of the fi rst mil len nium A.D. and the gional herpetofauna is largely unique, with mid dle of the sec ond millen nium A.D., the the excep tion of a few widespread, some times shel ter was used only epi sod i cally; thin lenses human - dispersed taxa (e.g., Nactus pelagicus of ash be ing sepa rated by thick lenses of ster- [Girard, 1858]; Hemiphyllodactylus typus ile sand, in di cating the proxim ity of wave ac- Bleeker, 1860; Hemidactylus garnotii Dum é ril tion. It is only from about 530 ± 40 yr B.P. & Bibron, 1836; H. frenatus Dum é ril & Bi- (Beta- 227967), starting with layers at a depth bron, 1836; and Lepidodactylus lugubris [Du- of 65 cm, that contin u ous use of the shelter is mé ril & Bibron, 1836]). All lizards known ap parent. from the region are either skinks (Scincidae) Site LTD - 18, called Dré né , is a rock- shel ter or geckos (Gekkonidae, Diplodactylidae). We lo cated on the fi rst cliff face of the uplifted used diag nos tic charac ters previ ously em- coral plat form of Tiga, at the rear of the build- ployed to dis tin guish related groups of geckos ings of the Tiga chiefdom. This site was used and skinks in fos sil lo cali ties of New Zealand until re cently for shelter during cyclones. and Tonga (Gill 1985, Worthy 1987, 1991, Three test- pits were opened in the low- Pregill 1993, Lee et al. 2009), as well as ad di- ceilinged shel ter (Figure 2), reaching the in tional charac ters based on published os te o log- situ pre human base at a depth of approx i ma- i cal studies (Wellborn 1933, Kluge 1962, Reptile Remains from Tiga, Loyalty Islands · Daza et al. 535

Figure 2. View of a test- pit exca va tion under way at site LTD- 18 at Dré n é , Tiga, showing the low ceiling of this shel ter site.

Hä upl 1980, Bauer 1990a, Abdala 1996, Con- Terrestrial snakes in the Loyalty Islands rad 2008, Daza 2008, Daza et al. 2008, Evans are represented by two fami lies, Boidae and 2008, Gelnaw 2011, Kennedy 2011, Gauthier Typhlopidae (Sadlier and Bauer 1997). Typh- et al. 2012, Jerez 2012) and new ob ser vations lopids occur ring in the southwest Pacifi c are to identify the taxa present in our archeologi- all very small and unam big u ously iden ti fi able cal samples. It is not cur rently possi ble to use osteologically at the famil ial level. As no typh- an apomorphy- based approach to di agnose lopid ele ments were present in our sample, the two resi dent gekkotan fami lies (Diplodac- our ophid ian compar i sons were re stricted to tylidae and Gekkonidae) from one another as rel e vant boids, although laticaudine ela pids they ex hibit plesiomorphic and/ or conver gent were also taken into ac count, as the four spe- oste o log ical charac ters, at least with re spect to cies in this group oc cur ring in New Caledo- the el ements preserved in the Tiga mate rial; nian waters (Laticauda colubrina [Schneider, there fore, we have focused on char acters that 1799], L. frontalis [de Vis, 1905], L. laticaudata are vari able at the ge neric and species level for [Linnaeus, 1758], L. saintgironsi Cogger & all of the lizard mate rial from Tiga. Herein we Heatwole, 2006) regu larly come onto land and use the term “ gecko ” syn ony mously with may be found at some distance inland (Ineich “ limbed gekkotan” (i.e., nonpygopodid gekko- and Laboute 2002). tans), al though in some instances we com- Descriptive oste olog i cal terms follow Daza ment on pygopodids in a compar a tive context. et al. (2008, 2012) and Evans (2008). In this 536 PACIFIC SCIENCE · Oc to ber 2015 paper we use the term “ seam” (sensu Jones ral History Museum, London (NHMUK). et al. [2011]) to refer to the bound ary between Institutional abbre vi ations for com para tive bones in artic u lation as distinct from “ su ture, ” mate rial fol low Sabaj Pé rez (2013). refer ring to the en tire joint in clud ing the ar- To esti mate the snout- vent lengths (SVLs) ticulating surfaces of partici pating bones and of fossils, we multi plied indi vidual fossil el e- the asso ci ated connec tive tissue, if any, be- ment lengths by (SVL/ ele ment length) de- tween the seams. rived from X- rays of adult speci mens of 60 Observations were made on Leica MZ6 extant Pacifi c skinks and limbed gekkotans of and Nikon SMZ1000 dissect ing micro scopes compa ra ble size (see Appendix 1). The esti- (Leica Microsystems, Buffalo Grove, Illinois), mated size for fos sils was compared with max- the latter equipped with a dig i tal camera i mum SVLs of candi date squamate species (Nikon DS- Fi1) and the image ac quisi tion (Bauer and Sadlier 2000, Meiri 2008) for Tiga. soft ware NIS Elements D (ver. 3.1, Nikon Corporation, Tokyo, Japan). All photo graphs were cre ated us ing the Extended Depth of results Focus appli ca tion, which creates an all in- Osteological Comparisons between Geckos focus image from a se ries of Z - axis images. and Skinks Images were assem bled and measured on Adobe Illustrator CS3 software (Adobe Sys- A total of 83 bones were recov ered from four tems Incorporated, San Jose, California). sites (Figure 2, Appendix 2). The most com- High -res o lu tion X- ray computed tomogra- mon el ements in the com bined sample of liz- phies of extant species were obtained at the ards and snakes were verte brae (25.3% ), lizard Digimorph fa cil ity of the University of Texas fem ora (18.1% ), lizard dentaries (12.0 %), lizard at Austin and the Amer i can Museum of Nat- hu meri (10.8 %), liz ard fron tals (9.6% ), and liz- ural History in New York with an Xradia ard maxil lae (6.0% ). The lizard mate rial re- microXCT scanner (Xradia Inc., Pleasanton, covered includes both geckos and skinks of California; 200 kV and 100 μ A) and 2010 GE sev eral spe cies each, but snake ma te rial recov- phoe nix v| tome| x s240a microfocus system ered repre sents a sin gle spe cies. (General Electric, Fairfi eld, Connecticut; max illa : Five max illae were found, two 80 kV and 170 μ A), respec tive ly. Digital radio- from geckos, representing two species, and graphs of some speci mens were obtained at three from skinks, representing a single spe- the U.S. National Museum of Natural His- cies. This bone has a very distinct morphol ogy tory at the Museum Support Center us ing a in both groups (Figure 3), espe cially with re- Kevex PXS10- 16W X- ray source (Thermo spect to the shape of the fa cial process (Wor- Electron Corporation, Scotts Valley, Cali- thy 1987), which, unfor tu nately, is very deli cate fornia) and Varian Amorphous Silicon Digi- and eas ily lost in small fos sils (see Estes [1983]). tal X- RayDetector PaxScanH 4030 (Varian Another vari able part of this bone is the poste- Medical Systems, Inc., Palo Alto, California). rior process, which in some skinks bi furcates Fossil mate rial exam ined is housed in the into a su pe rior and infe rior pos te rior pro cesses collec tion of the Institut d’ arch é ologie de la (Gill 1985), but is always single in gekkotans. Nouvelle - Calé donie et du Pacifi que in Nou- Toot h morphol ogy can also be used to dif fer- mé a. Comparative mate rial was exam ined en tiate these groups; skinks tend to have blunt, from the collec tions of the Ameri can Museum chisel - shaped tooth crowns, whereas crowns of Natural History, New York (AMNH); Aus- are rounded and con i cal in most gekkotans, al- tralian Museum, Sydney (AMS); California though in a few groups, these can also be re- Academy of Sciences (CAS); James Ford Bell curved (Patchell and Shine 1986, Bauer and Museum, University of Minnesota, St. Paul Russell 1990, Nikitina and Ananjeva 2009). (JFBM); Museum of Comparative Zoology, Skinks may bear some stri a tions on the lin gual Harvard University, Cambridge (MCZ); Mu- side of the crown (Kosma 2004), which are not seum of Vertebrate Zoology, University of present in gekkotans. The la bial fo ramina are California, Berkeley (MVZ); and The Natu- also vari able in size (large in skinks and small Reptile Remains from Tiga, Loyalty Islands · Daza et al. 537

Figure 3. Maxillae: ( A, B ) Lioscincus nigrofasciolatus (MCZ A- 27345), ( C, D ) Bavayia robus ta (MCZ A - 27347). ( A, C ) Lateral and ( B, D ) medial views of right maxil la. Scale bar equals 5 mm.

Figure 4. Frontals: (A, B ) Lioscincus nigrofasciolatus (MCZ A- 27345), (C, D ) Bavayia ro bus ta (MCZ A - 27347). ( A, C ) Dorsal and (B, D ) ven tral views. Scale bar equals 5 mm. in geck os) (Estes 1983, Greer 1989, Evans tans and in lygosomatine skinks (Greer 2008). Number of tooth loci is also in forma- 1970, 1989, Abdala 1996, Evans 2008, Skin- tive, but these vary with age and size (Arnold ner et al. 2011), this bone can be eas ily differ- 1980, Bauer and Russell 1990), and this vari a- enti ated on the ba sis of the crista cranii. In tion must be eval uat ed. skinks and many other lizards, the crista cra- frontal: One skink frontal and seven nii are two ventral downgrowths that do not gecko frontals, the latter assign able to three meet ventrally (Conrad 2008, Evans 2008, spe cies, were identi fi ed. Although the fron- Gelnaw 2011, Gauthier et al. 2012) (Figures tals are unpaired in the major ity of gekko- 4 A, B ); whereas in gekkotans they fuse 538 PACIFIC SCIENCE · Oc to ber 2015

ventrally forming a subolfactory pro cess that of the most conspic u ous differ ences are the com pletely en closes the olfac tory nerve presence of a fora men prootico for cranial (Wellborn 1933, Kluge 1962, Bauer 1990 a, nerve V instead of an incisura or notch (al- Daza et al. 2013) (Figures 4C, D ). though there is a notch in pygopodids and a pari e tal: One skink pari etal was found. few limbed geck os, e.g., Gekko gecko [Lin- Parietals of gekkotans and skinks are sim i lar naeus, 1758], Coleonyx variegatus [Baird, but they can be distin guished eas ily be cause 1858]), lon ger basipterygoid pro cess of the skinks have a pa ri etal fora men, fused pari etals, parabasisphenoid forming an ob tuse angle and a well- de vel oped pari etal fossa, whereas in (acute in skinks and elon gated in pygopo- gekkotans, the pari etal fora men is missing dids), lack of basal spines or verti cal lami nae (Gill 1985, Estes et al. 1988, Daza et al. 2013), that fl are out an teri orly from the crista alaris and the pa ri etals are typ i cally unfused (there and that overlie the basipterygoid stalks are excep tions, e.g., the pygopodid Lialis and (some limbed geckos have a clinoid pro cess in some spe cies of the gekkonid gen era Gekko and this po sition, but this struc ture is wider than Luperosaurus, in which partial or complete fu- a narrow spine), presence of a well- devel oped sion occurs in large spec imens, but not among crista prootica (lacking in pygopodids and re- diplodactylids or insu lar Pacifi c gekkonids). duced in some gekkonids, e.g., Matoatoa ), and quad rate: One skink quad rate was a bi cip i tal occip i tal condyle (single in Lialis ). found. The quad rates of limbed gekkotans dentary : Ten dentaries were found, seven and skinks have a simi lar deep concave conch, are from geckos, representing three taxa, a rela tively narrow central pillar, little medial and three from skinks of a single species. expan sion, and a small pter ygoid lappet These two groups typi cally have a closed (Evans 2008). Despite these simi lar i ties, the Meckelian canal (Kluge 1967, Greer 1989, quadrate of skinks differs from that of gek- Shea 1999, Evans 2008) (Figure 5), although kotans in its type of sus pension; whereas in some skinks retain an open Meckelian groove, skinks the squa mosal and supratemporal but they can be easily differ enti ated by the contrib ute to the sus pension by contacting mandib u lar symphy sis. Geckos gener ally have this bone dorsal ly, in gekkotans the sus pen- a re duced intermandibular ar ticu lation, taper- sion is almost exclu sively through ar ticu la- ing lat erally away from the mid line and giv- tion with the otooccipital (or paroccipital ing the joint a ˄ shape (Holliday et al. 2010). abut ting) (Rieppel 1984), via a synchondrosis The ven tral border of the dentary in skinks is (Sensu Payne et al. [2011]). Although sus- strongly curved (Figures 5A, B ), whereas in pension cannot be directly assessed in disar- geckos this edge is gen er ally straight (Figures ticulated and frag men tary mate ri al, the 5 C, D ); the dentary is also strongly curved differ ences in the de gree of os sifi ca tion of the mediolaterally in skinks (Gill 1985). Geckos postero lat eral part of the cephalic condyle of also have a shelf for the anterolateral process the quadrate can help to dis tinguish these of coronoid on the labial side, whereas in groups. In many limbed gekkotans, this area skinks the dentary has an upwardly directed ap pears irreg u larly notched in the dried skull coronoid process that prevents the exten sion (Rieppel 1984), while cleared -and - stained of the coronoid onto the la bial surface of the prepa ra tions reveal the notch to be fi lled by dentary (Figure 5). Additional charac ters carti lage, often with a small imbedded os sifi - from tooth morphol ogy can be used to dif fer- cation (Daza et al. 2012, Daza pers. obs.). enti ate dentaries among these groups. Among skinks, the postero lat eral part may compound bone: Three compound have only a slight notch or an apical fora men bones were found, two are from geckos and at the confl uence of the tym panic crest, me- one from a skink, representing one taxon dial pillar, and the cephalic condyle (Evans each. Geckos and skinks both expe ri ence fu- 2008, Gelnaw 2011). sion among some postdentary bones. In gek- braincase: One gecko basicranium was kotans, the fusion of the artic ular and found, and it can easily be differ enti ated from prearticular and the partial fusion of the those of skinks by sev eral charac ters. Some surangular are common. The an gular has Reptile Remains from Tiga, Loyalty Islands · Daza et al. 539

Figure 5. Dentaries: (A, B ) Lioscincus nigrofasciolatus (MCZ A- 27345), ( C, D ) Bavayia ro bus ta (MCZ A- 27347). (A, C ) Lateral and (B, D ) medial views. Scale bar equals 5 mm. only been reported in a few species of eu- cally shaped, rather than a cordate ischiopu- blepharid geckos and in the genus Teratoscin- bic fe nes tra of some geckos (Kluge 1962, cus (Kluge 1962; Arnold 1977; Grismer 1988; Bauer 1990b, Daza et al. 2012). The ace tab u- Evans 2008). Among skinks, fusion of the lar area is also proportionally larger in artic ular (ar tic ular complex, i.e., artic ular geckos than in skinks (Daza et al. pers. obs.). and prearticular) and angu lar is also common ap pen dicu lar el ements: Nine humeri (Evans 2008), but the an gular is re duced, be- were recov ered, four from skinks and fi ve ing either persis tent or absent in some lygo- from geckos. When limbs are not re duced, somines (Greer 1970, Gelnaw 2011). Although the humerus is proportionally lon ger in skinks the mor phol ogy of the retroarticular pro cess than in gekkotans. Another dif fer ence lies is somewhat conver gent between skinks and in the shaft, which is more cylin dri cal and gekkotans, these two groups can eas ily be straight (Figures 6A, B ) in skinks, in com- differ enti ated because gekkotans have a pari son to geckos in which the preax ial tu- unique lateral notch that forms a waist proxi- beros ity and ectepicondyle em inence are mally (Estes et al. 1988) and a spoon -shaped consid er ably more well devel oped (Figures retroarticular process (rod- like in pygopods), 6 C, D ). Another differ ence is that the ra dial whereas skinks have a narrower retroarticu- condyle is twice as large as the ul nar condyle lar process with only a narrow, shallow cup is in geckos, whereas in skinks the radial con- (Kluge 1976, Hutchinson 1997, Evans 2008). dyle is only slightly larger than the ulnar verte brae : Twent y -two procoelous ver- condyle (Daza pers. obs.). te brae were found, 5 are from skinks, repre- Seventeen femora were re cov ered, 10 from senting two taxa and 17 from a single snake skinks and 7 from geckos. Femora of skinks spe cies. Gekkotan ver te brae, with the excep- and geckos can be identi fi ed because skinks tion of the major ity of New World sphaero- ex hibit a well- de fi ned U -shaped notch be- dactyls, are charac terized by amphicoelous tween the fem o ral head and the inter nal tro- verte brae (Hoffstetter and Gasc 1969, Kluge chanter (Figure 6; see also Worthy [1991: fi g. 1967, 1995). 8]). Morphological vari ation at the ge nus or pel vic gir dle : Five in nom inate bones spe cies level is subtler, and here we limit were re covered, three from skinks and two our selves to size compar i sons, al though in from geckos. These two groups can be dif- many cases the bones recov ered did not ex- fer enti ated because the pubis in skinks is hibit epiphy seal ter mi nal fusion, therefore more elongat ed, thus de fi n ing a more el lipti- skele tal matu rity cannot be assumed. 540 PACIFIC SCIENCE · Oc to ber 2015

Figure 6. Humeri (A – D ) and femora (E – I ). ( A, B, E– G ) Marmorosphax tricol or (MCZ A- 27320); (C, D, H, I ) Cor- relophus ciliatus (JFBM 15825). ( A, C ) Anterodorsal view of right humeri; ( B, D ) posteroventral view of right hu- meri; ( E, H ) ante rior view of right femo ra; (F, I ) poste rior view of right femo ra; (G ) posteroventral view of right humer us. Scale bar equals 5 mm.

Species Identifi cation Maré (Sadlier and Bauer 1997, Bauer and Sadlier 2000). Scincidae ma teri al : Maxillae (n = 3) ( Figures 7 A, C ) and compound bone (n = 1) (Figures 7D, E ). Emoia loyaltiensis (Roux, 1913), Loyalty Is- Using the length of two complete maxil lae lands Emoia . and a compound bone, we esti mate SVL val- ues be tween 66.5 mm and 68.5 mm. The dis tribu tion in new caledonian re- maxi mum SVL for this species is 83.2 mm gion: Recorded only from Ouvé a, Lifou, and (Zug and Ineich 1995). Reptile Remains from Tiga, Loyalty Islands · Daza et al. 541

Figure 7. Material identi fi ed as belong ing to Emoia loyaltiensis: (A – C ) maxilla (LTD- 14A 65– 70 cm, LTD- 18C 55– 65 cm); (D, E ) com pound bone (LTD -18C 55 – 65 cm). (A, C, D ) Lateral view; (B, E ) medial view. Scale bar equals 5 mm.

re marks : One left (Figure 7A, B ) and terolat eral tri angu lar pro cess, and the max illa one right com plete max illa and a frag ment of contacts the ventral and labial surfaces of the posteromedial portion of a left maxilla the jugal. The pal atal shelf is irreg u lar and (Figure 7C ). The most complete ele ment mea- some what folded ventrally (forming a thick sures 8.6 mm and bears 20 tooth loci. Teeth la bial edge for the tooth row, being deeply have slightly ex panded tooth ba ses and chisel - troughed). shaped crowns. The facial process bends The compound bone is from the right side dorsomedially. The facial process slopes grad- and is incom plete (Figures 7D, E ), lacking the u ally from the poste rior edge of the exter nal ante rior process of the surangular portion and naris and ends in a pointed pro cess. There is the total ity of the retroarticular process, but a well- de fi ned tri an gu lar de pression me dial the out line has no indi ca tion of a proxi mal to the ex ternal border of the naris, this de- waist as in geck os. This bone is compressed pressed area is defi ned by two ridges, the mediolaterally, the labial sur face with an equal crista lateralis, which is perpen dic u lar to the an terior taper ing dentary- surangular seam, la bial margin of the maxil la, and the crista the ante rior surangular fora men oval and transversalis, which extends from the dor sal more than four times the diam e ter of the pos- area of the maxil lary lap pet to the bor der of te rior surangular fora men, a longi tudi nal the nares. Below the junc tion of these two ridge ventral to the poste rior surangular fo- ridges there is a trian gu lar transverse wall ramen, and an oval but elon gated man dib ular that is pierced by a fora men that opens ante- fos sa. riorly and corre sponds to the an te rior al ve o- compar i sons : The fossil maxilla has four lar fora men (Smith and Gauthier 2013). The vascu lar foram i na, but some of these are very pos terior margin is mainly con cave but also en larged and seem to be confl uent as result of has a short prefron tal pos terior process. The compact bone erosion. Five and six vascu lar pos terior process above the tooth row is bi- foram ina were observed in compar a tive mate- fur cated, the supe rior projec tion forms a pos- rial Lioscincus nigrofasciolatus and E. loyaltiensis, 542 PACIFIC SCIENCE · Oc to ber 2015 re spec tively, although the number of foram- ti mate SVL val ues be tween 98.1 mm and ina may be intraspecifi cally vari able. 108.9 mm. The maxi mum adult SVL for this spe cies is 112 mm (Bauer and Sadlier 2000). Lioscincus nigrofasciolatus (Pe ters, 1869), re marks : The frontal bone, mea suring Green- bellied tree skink. 7.7 mm in length, is almost complete (Figures 8A, B ). Similarities with the frontal of the dis tribu tion in new caledonian re- compar a tive ma terial from this species in- gion: Widespread on the Grande Terre; Ile cludes dor sal sculptur ing and a broad nasal des Pins and its satel lite islands; and on Ou- shelf in dicat ing that na sal bones cov ered most v é a, Lifou, and Maré (Bauer and Sadlier of the ante rior portion. 2000, Geneva et al. 2013). The pa rie tal (Figures 8 C, D ) shows signif- ma teri al : Frontal (n = 1) (Figures 8 A, B ), i cant inter spe cifi c vari ation among some skink pari e tal (n = 1) (Figures 8 C, D ), quadrate (n = 1) spe cies (Worthy 1987, 1991). This bone was (Figures 8 E, F ), dentary ( n = 3) (Figures 8 G, found asso ci ated with the frontal and an in- H ), and innom inate bone (n = 2) (Figure 8I ), nomi nate bone, and they might cor respond to and verte brae (n = 3) (Figure 8J ). Using the the same indi vid u al. The main body of this length of complete frontal and pari etal, we es- bone (exclud ing the pos terolat eral pro cess) is

Figure 8. Material identi fi ed as belong ing to Lioscincus nigrofasciolatum: (A, B ) frontal (LTD- 14A 70– 75 cm); (C, D ) pa rie tal (LTD - 14A 70– 75 cm); (E, F ) quad rate (LTD- 18C 55 –65 cm); (G, H ) dentary (LTD 18C 45– 55 cm); (I ) in- nomi nate bone (LTD - 14A 70– 75 cm); (J ) verte brae, pygal series (LTD- 02A 60– 70 cm). (A, C ) Dorsal view; ( B, D ) ventral view; (E ) poste rior view; (F, H ) medial view; (G, I ) lateral view; (J ) dor sal (above) and ventral (below) views. Scale bar equals 5 mm. Reptile Remains from Tiga, Loyalty Islands · Daza et al. 543 almost rectan gu lar (7.5 mm long × 5.5. wide) cor respond ing to the third or the fourth py- and the pa rie tal fo ramen is located in the an- gal. The third ver tebrae has both transverse terior half as in other skinks (Greer 1970, Gel- pro cesses broken, there fore it is not possi ble naw 2011). The lateral mar gins are almost to infer its posi tion. The neu ral arch in all of straight with a small lateral pro jec tion an te- these verte brae is mod er ately high and has rior to the postero lat eral pro cess es. In ventral marked dor sal ridges on the neu ral arch lam- view, the pos terior margin has a ∩ -shaped ina, although in compar i son with the mate- notch with no posteromedial process, defi n ing rial of L. nigrofasciolatus, these ridges tend to the posterodorsal margin of the pari etal fossa. be somewhat abrad ed. The left quadrate recov ered (Figures 8 E, compar i sons : The frontal re sembles F ) measures 5.0 mm long and has an ex panded both Lioscincus nigrofasciolatus and Emoia loy- pos terior concav ity with stout me dial pillar. altiensis in the degree of nasal overlap and On the dor sal surface, this quad rate of this sur face sculptur ing. However the frontona- spe cies has a large quad rate fora men or this sal seam in the compar a tive mate rial of may perfo rate the margin of the bone to form L. nigrofasciolatus (Figure 4A ) is charac ter- a small notch. ized by a less well - de fi ned super fi cial median One of the dentaries is nearly in tact (Fig- pro jec tion of the bone. The bi furca tion of ures 8G, H ). The Meckelian canal is fused the anteromedial process of the frontal is but re mains open up to the 11th last tooth. well defi ned, as in the compar a tive ma te rial The most complete one is a left bone with 35 of Lioscincus and Emoia. The two prongs are tooth lo ci, 5 men tal foram i na, measur ing asym met ri cal in L. nigrofasciolatus and sub- 9.5 mm in length. equal in E. loyaltiensis. The fossil frontal The innom inate bones are from the left seems to have subequal prongs, but it is pos si- and right sides (Figure 8I ). These bones are ble that one of the sides is in com plete; there- iden tifi ed as L. nigrofasciolatus based on size fore, this dif fer ence can not be used and morpho logi cal char ac ters, includ ing a conclu sively. Additionally, although charac- short and wide pectineal process, an antero- ters from the ante rior margin of the frontal medially ori ented metischial pro cess, and an may be used to compare differ ent species, expanded boot - like is chial foot. these charac ters in skinks should be used The verte brae are procoelous and are all with some re serve because it has been demon- from the caudal region. Each measures about strated that in Oligosoma (as Cyclodina ) this 3 mm long (Figure 8 J ). They have single structure varies with de gree of os sifi ca tion transverse pro cesses with expanded bases; this (Worthy 1987). marks the divi sion between a wide ante rior part and a narrow pos terior part. They are Scincidae, incertae sedis nonautotomic and have no hemal arches; therefore, we identify these as part of the ma teri al : One dentary, one innom inate pygal series using two previ ous defi ni tions bone, fi ve presacral and three cau dal verte- of this verte bral segment (Holder 1960, Rus- brae, four humeri, and seven femo ra. sell 1967). Although the most proxi mal cau- re marks : The innom inate bone is from dal verte brae are simi lar in all lizards the right side and is asso ci ated with an 8 mm (Etheridge 1967), these verte brae have vari- femur; it is tenta tively identi fi ed as belong ing able trans verse processes that may be used to to Emoia cyanura . The innom i nate bone is iden tify their posi tion. We iden tify as the somewhat sim ilar to that of Marmorosphax fi rst pygal ver tebra (Figure 8J, left) an ele- tricol or, but is smaller in size. The pectineal ment that retains a posterolaterally directed pro cess is broken and the metischial pro cess left transverse pro cess that is wide and long is dig it- like in shape and more posteriorly di- (1.5 times centrum length) and does not taper rected than in M. tricol or, where this process later ally. Another verte bra (Figure 8J, mid- is more lami nar and slightly bent ante rior ly. dle) exhib its a left transverse pro cess that is The is chial foot is rectan gu lar, but the ter mi- narrower and is almost later ally direct ed, nus of this struc ture seems to be bro ken. 544 PACIFIC SCIENCE · Oc to ber 2015

The presacral ver te brae are very frag men- ma teri al : Dentary (n = 1) (Figures 9 A, tary and measure about 4 mm each. These el- B ), maxilla (n = 1) (Figures 9C, D ), frontal e ments cannot be identi fi ed be yond famil ial (n = 1) (Figures 9E, F ), and compound bone level based on previ ous descrip tions (Hoff- (n = 2) (Figures 9G, H ), and in nom inate stetter and Gasc 1969). bone (incom plete, n = 1) (Figure 9I ). The es- The lack of diag nos tic features at spe cies ti mated SVL us ing the most com plete bones level limits the precise al loca tion of the appen- are frontal (72.7 mm), dentary (66. 7 mm), dic ular el ements. Using known lengths of maxilla (68.1 mm), and com pound bone bones from disarticulated speci mens and the (74.3 mm). The maxi mum SVL for this spe- maxi mum SVL size for can di date spe cies, we cies is 86 mm (Bauer and Sadlier 2000). were able to pro vide tenta tive iden ti fi ca tions. re marks : The dentary (Figures 9A, B ) is The four hu meri recov ered were mostly nearly complete. It is robust and has rel a tively com plete. The esti mated SVL in ter val for the stout teeth in 28 –30 tooth loci. The number liz ards asso ciated with these bones is 55.8 mm of tooth loci in adult speci mens of B. crassicol- to 76.9 mm. This falls within the size range of lis over 70 mm SVL is usually approx i mately adult Emoia loyaltiensis or subadult to small 35, so the low num ber in the Tiga speci men adult Lioscincus nigrofasciolatus . The much is consis tent with the esti mated size of the larger Phoboscincus garnieri (Bavay, 1869) (to spec i men since tooth loci increase with size 200 mm SVL) has been recorded from Lifou intraspecifi cally (Bauer and Russell 1990). and Ouvé a (Sadlier 1987, Bauer and Sadlier The sin gle left max illa lacks most of the fa- 2000), and it is also possi ble that the larger hu- cial process. The piece of bone mea sures meri and fem ora could repre sent ju ve nile ma- 8.5 mm and has 34 tooth loci and six vascu lar te rial of this species. fo ramina (Figures 9C, D ). Features we tenta- Seven femora were re cov ered, and the es- tively con sider diag nos tic of B. crassicollis in- timated SVL in terval for the lizards asso ci- clude simi lar tooth locus counts and a simi lar ated with these bones is 44.9 mm to 76.8 mm. transverse shelf of the pre maxilla (straight, be- Within this range some of the likely species tween a short anterolabial process and the are Caledoniscincus atropunctatus, C. haplorhinus maxil lary lappet). (G ü nther, 1872), or C. austrocaledonicus (Bavay, The fron tal is incom plete, lacking the an- 1869), all of which attain maxi mum sizes of terodorsal part. The pre served part measures 57 mm or less, or Emoia cyanura (maxi mum 7 mm in length (Figures 9 E, F ). This bone 56 mm SVL), and Emoia loyaltiensis (maxi mum has a dor sal sur face that is slightly furrowed, 83 mm SVL). The larger fem ora are also con- large dis tally ex panded anterolateral processes sis tent with ju venile or subadult Lioscincus of the crista cranii, short anteromedial process nigrofasciolatus. of the crista cranii, poste rior margin of the crista cranii trans verse in stead of V notched, a Diplodactylidae broad recess lateral to the tu bu lar portion of the bone, and a constricted pos terolat eral Bavayia crassicollis Roux 1913, Strand Bavayia . pro cess of the fron tal to fi t the postorbito- frontal. dis tribu tion in new caledonian re- The compound bone (Figures 9 I, J ) lacks gion: Although pre vi ously thought to be most of the retroarticular process and the me- present on the Ile des Pins as well as at least dial margin of the man dibu lar fos sa. It is one near- coastal is let off the northeast 10– 11 mm in length, has a dentary- surangu- Grande Terre (Bauer and Sadlier 2000), on- lar seam with rounded out line, and lacks an go ing molec ular phylo ge netic studies re veal ante rior surangular fora men. that B. crassicollis is restricted to Lifou and The innom inate bone lacks the dis tal por- Mar é (its absence from Ouv é a may be an ar ti- tions of pubis and ischium and exhib its the il- fact of poor sam pling), except for proba ble iac blade shape and ace tab u lar contour typi cal acci dental translo cations to the Ile des Pins of this spe cies (Figure 9K ). The more incom- re gion (Geneva et al. 2013). plete el ements can only be assigned with cer- Reptile Remains from Tiga, Loyalty Islands · Daza et al. 545

Figure 9. Material identi fi ed as be longing to Bavayia crassicollis (A – I ) and B. sp. (J – M ): ( A, B ) dentary (LTD 18C 55– 65 cm); (C, D ) maxilla (LTD- 18C 55 –65 cm); (E, F ) frontal (LTD - 14A 60– 65 cm); (G, H ) compound bone (LTD- 18C 55– 65 cm); (I ) in nomi nate bone (LTD- 14A 70– 75 cm); (J, K ) dentary (LTD 18A 50– 55 cm); (L, M ) braincase (LTD- 18C 55– 65 cm). (A, C, G, J, I ) lateral view; ( B, D, H, K ) medial view; (E, L ) dor sal; (F, M ) ventral. Scale bar equals 5 mm. tainty to the B. cyclura complex, but their tially larger than the largest known B. sauva- spe cifi c allo cation is highly prob a ble. gii ( ∼ 62 mm SVL). The more robust dentary compar i son : The fron tal of B. crassicollis and larger, less numer ous teeth also dis tin- and other members of the B. cyclura group guish B. crassicollis from this species. has more pronounced anterolateral pro- The poste rior surangular fora men lo cation, cesses of the crista cranii and broader lateral medial offset of the ar ticu lar, quadrate groove, recesses than B. sauvagii . The Tiga frontal and anterodorsal incli nation of the ar tic ular implies a SVL of ∼ 73 mm, which is sub stan- surface are also simi lar to other large - bodied 546 PACIFIC SCIENCE · Oc to ber 2015 mem bers of the B. cyclura group, in clud ing tively lit tle intrageneric vari a tion in Bavayia, B. ro busta Wright, Bauer, & Sadlier, 2000. may be attrib ut able to ei ther this spe cies or B. crassicollis . At least the larg est of the den- Bavayia sp. taries implies a size greater than that reached by B . aff. sauvagii, but is more sim ilar to this ma teri al : Dentary (n = 3) (Figures 9J, spe cies than to B. crassicollis in morphol o gy. K ) bones, braincase (n = 1) (Figures 9L, M ). Using the length of the most complete den- Gekkonidae tary and the braincase, we esti mate SVL val- ues of 69.3 mm and 50.3 mm, respec tive ly. Gehyra georgpotthasti Flecks, Schmitz, B ö hme, re marks : The dentary is represented by Henkel & Ineich, 2012, Vanuatu giant gecko. two incom plete left rami and one nearly com- plete right ramus (Figures 9J, K ). These bones dis tribu tion in new caledonian re- have numer ous slender teeth (to tal tooth loci gion: This species is known from Maré , ranges of 38– 44, which are con sis tent with Dudun, and a sin gle in di vidual from Lifou, counts from living speci mens). The brain- as well as from Malakula, Pen tecost, Epi, case (Figures 9L, M ) has a marked su ture Espiritu Santo, Ambae, Erromango, and Efate be tween the parabasisphenoid and the basi- in Vanuatu (Hamilton et al. 2009, Ineich occipital ante rior to the sphenooccipital tu- 2011, Flecks et al. 2012). Two speci mens from bercles and the sphenooccipital tuber cle Fakarava, Tuamotu Archipelago, are cer tainly partially cover ing the lateral aper ture of the the re sult of human intro duc tion (Flecks et al. recessus scalae tympani in ventral view. 2012), and we consider a record from Norfolk compar i sons : The dentary differs Island (Boulenger 1885) to be in er ror. slightly from the com par a tive ma terial of Ba- ma teri al : Maxilla ( n = 1) (Figures 10A, vayia sauvagii in hav ing a lower height and a B ), dentary (n = 3) (Figures 10C, D ), in nom i- very well- defi ned lat eral shelf for the coro- nate bone (n = 1) (Figure 10E ). The esti mated noid poste rior to the tooth row. The slen der SVL val ues us ing the most com plete bones are: den tition somewhat resem bles the larg er, ex- dentary (137.5 mm), and maxilla (123.8 mm). tralim ital species Oedura marmorata Gray, The maxi mum known SVL for this species is 1842. The braincase strongly resem bles com- 142 mm (Flecks et al. 2012). para tive ma te rial of Bavayia . It is too large to re marks : The maxilla is one left incom- be refer able to Lepidodactylus lugubris (maxi- plete bone with out most of the fa cial process mum SVL 47 mm), to which it also bears and preserv ing the en tire basal part includ- some simi lar i ty. The paroccipital process is ing the pal a tal shelf (Figures 10A, B ). The long and slen der as in Bavayia, Gehyra, and tooth row measures 16.5 mm, bears 33 tooth Nactus, but differs from the latter two in the loci, and has nine vascu lar foram i na. extent of the sella turcica, be ing small in Ba- Two dentaries from the left and one from vayia, in ter me diate in size in Gehyra, and the right were recov ered; the ones from the broad and elongated in Nactus . left are the most com plete ones (Figures 10C, Bavayia sauvagii was previ ously thought to D ), whereas the one from the right is largely be widespread on the Grande Terre as well as incom plete. The most complete dentary mea- on the Ile des Pins and on Mar é (Bauer and sures 21 mm and bears 34 tooth loci. The other Sadlier 2000); howev er, ongo ing molec u lar more complete el ement measures 16 mm and phy lo genetic studies re veal that B. sauvagii is bears 31 tooth loci, and the most in com plete a com plex of microendemic cryp tic species el ement only preserves the ante ri or - most part (Bauer and Jackman 2006). The form occur- of the tooth row and a splint of bone corre- ring on Maré ap pears to be restricted to this spond ing to the ven tral margin of the dentary. island and perhaps other islands in the south- A frag ment of a large right innom i nate bone ern Loyalties. The max i mum size of B . aff. (Figure 10E ) preserv es the ilium blade, ac etab- sauvagii is 62 mm SVL (Bauer and Sadlier ulum, and minor parts of is chium and pubis. 2000). Thus the braincase, which shows rela- On the pu bic part, there is no indi cation of Reptile Remains from Tiga, Loyalty Islands · Daza et al. 547

Figure 10. Material identi fi ed as be long ing to Gehyra georgpotthasti ( A – E ) and Nactus pelagicus (F, G ). ( A, B ) Max- illa (LTD- 14A 45– 50 cm); (C, D ) dentary (LTD- 14A 65– 70 cm); (E ) innom i nate bone (LTD- 18C 55– 65 cm); (F, G ) fron tal (LTD - 14A 85– 90 cm). (A, C, E ) lateral view; (B, D ) medial view; (F ) dor sal view; ( G ) ventral view. Scale bar equals 5 mm. an obtu rator fora men right next to the ac etab- den tition is moder ately recurved as in u lum. The il iac blade tapers distally grad ual ly. G. georgpotthasti and R. trachyrhynchus and compar i son : Although this mate rial re- not strongly recurved as in R. auriculatus . sem bles both large- sized gekkonids and di- The dentaries also re semble Rhacodactylus plodactylids, we favored an iden ti fi ca tion of auriculatus and R. trachyrhynchus although this ma te rial with G. georgpotthasti based on they differ from Rhacodactylus in having more com pari sons with museum mate rial and dis- tooth loci. The splenial extends as far an te ri- tribu tion re cords of this spe cies (Bauer and orly as the 10th tooth lo cus from the back. Sadlier 2000, Flecks et al. 2012), previ ously The me sial teeth are re curved and the pos te- only known from Vanuatu and Lifou, Maré , rior straight, but the ante rior - most are not as and Dudun among the Loyalties (Ineich procum bent as in R. auriculatus . There are 2011, Flecks et al. 2012). seven mental foram i na, the last one slit -like In terms of size and shape, the maxilla is and hav ing a mark edly shallow furrow di- also compa ra ble to that of Rhacodactylus auric- rected to ward the ante rior point of the labial ulatus (Bavay, 1869) and R. trachyrhynchus Bo- coronoid shelf as in Gehyra georgpotthasti and cage, 1873, includ ing two rows of vas cular Rhacodactylus spp. Ventral border of dentary is foram i na, a pos terolat eral margin slightly fl attened, and bone surface is slightly rugose. rectan gu lar and not ta pering mark ed ly, mid- dle teeth 3 – 4 times larger than the pos te rior Nactus pelagicus (Girard, 1858), Pelagic gecko. teeth, pala tal shelf with a shal lowly troughed ven trally, pal a tal shelf ante rior to the pal a tine dis tribu tion in new caledonian re- su ture straight with maxil lary lappets forming gion: This unisex ual species is widespread an inter nal an gle of about 140º. However, the in the New Caledonian region, in clud ing 548 PACIFIC SCIENCE · Oc to ber 2015

Ouvé a, Lifou, and Maré (Bauer and Sadlier 7.0 – 9.0 mm) recov ered imply SVLs of 48.2– 2000) and is wide spread in southern Vanuatu, 58.4 mm, and 47.1– 60.7 mm, respec tively. Fiji, and parts of Micronesia and Polynesia These could belong to any of sev eral candi- (Bauer and Henle 1994, Heinicke et al. 2010). date gekkonid or diplodactylid species (Ba- ma teri al: Frontals (n = 5) (Figures 10F, vayia crassicollis, B. aff. sauvagii, Nactus G ). The es ti mated SVL us ing the most com- pelagicus, Hemidactylus garnotii, H. frenatus, plete fron tal (7.8 mm) is 75.1 mm. The maxi- Lepidodactylus lugubris ) except Gehyra georgpot- mum SVL for this species is 86 mm (Zug and thasti or Hemiphyllodactylus typus, which can Fisher 2012). be excluded on the basis of size. re marks : The fron tal (Figures 10F, G ) has a smooth, fl at dor sal surface and wide an- Boidae te rior end (Pregill 1993). The nasofrontal seam is W -shaped, and the ante rior and dor- Candoia bibroni (Dumé ril & Bibron, 1844), sal part of the frontal remains unfused, de- Pacifi c boa. fi ned anterolateral shelves for the prefron tal, an te rior portion of the bone lami nar with lit- dis tribu tion in new caledonian re- tle or no inden ta tion (i.e., no discrete antero- gion: This snake has been recorded from medial and anterolateral process), slightly Ouvé a, Lifou, and Maré (Bauer and Sadlier concave frontoparietal suture, pos terolat eral 2000) and is the only rep tile previ ously re- pro cesses termi nate in broad, slightly poste- ported in the liter a ture from Tiga (Oliva and riorly angled fl anges, anterolateral pro cess of Talon 1996). It is oth erwise widely distrib- the crista cranii trian gu lar and poorly de vel- uted from the east ern Solo mon Islands to oped, with a longer process in between (i.e., Ameri can Samoa (Smith et al. 2001). anteromedial pro cess). ma teri al : Procoelous trunk verte brae compar i sons : Diplodactylids and Hemi- ( n = 17) ( Figures 11A, D ). No ex act es ti ma- dactylus frenatus possess a rela tively broader tions of the size are possi ble with the iso- inter or bital re gion of the fron tal. The pos- lated verte brae, but the sizes of these tero lat eral processes of the frontal (which ver te brae are compa ra ble to the skel etons of reaches a maxi mum SVL of only 47 mm) in the compar a tive mate rial. Lepidodactylus lugubris taper to a fi ne point. re marks : All the verte brae have the diag- nos tic charac ters of boine snakes such as the Gekkota, incertae sedis pres ence of paracotylar fossae and foram ina and the vaulted, biangled pos te rior margin of ma teri al : Six hu meri and seven femo ra. the neural arch (Head et al. 2006, Head re marks : Using the size of the el ements et al. 2009). Candoia bibroni is the only boine recov ered to infer the SVL, the six humeri known from the Loyalty Islands (Bauer and (range 6.5– 7.0 mm) and seven femora (range Sadlier 2000, Noonan and Sites 2010).

Figure 11. Material identi fi ed as be long ing to Candoia bibroni : ( A– D ) trunk verte bra of Candoia bibroni (18C 45 – 55 cm). (A ) Dorsal; ( B ) ven tral; (C ) an teri or; (D ) poste rior views. Scale bar equals 5 mm. Reptile Remains from Tiga, Loyalty Islands · Daza et al. 549

discus sion recov ered on Tiga con fi rm at least the his tor- i cal presence of the snake on the island. The boid Candoia bibroni is the only extant Both Gehyra and Candoia were found in rep tile to have previ ously been reported from deeper as well as more su perfi cial layers of ex- Tiga, but without any vouchered spec i mens cava tion (Appendix 2). Although the skel etal (Oliva and Talon 1996). The anal ysis of the ele ments themselves have not been dated, it present mate rial, obtained from ar chaeo log i- may be as sumed that the most re cent mate rial cal exca va tions, thus adds substan tially to the cor responds to the histor i cal pe riod, whereas known reptile fauna of Tiga. Of a total of 19 the older mate rial extends well into the pre - spe cies pre viously recorded from the Loyalty Euro pean period of Tiga’ s occu pation. Al- Islands (Sadlier and Bauer 1997, Bauer and though most herpe tol o gists have assumed Sadlier 2000), we are able to in fer the presence that Candoia bibroni is native to the Loyalties, of six spe cies with some surety, includ ing two one published source has proposed that the skinks and three geckos, and the boa. Another snakes were intro duced as food items by Poly- two species, Bavayia aff. sauvagii and Emoia cy- ne sians in the six teenth cen tury (Logan and anura are proba bly represented. On the basis Cole 1997) and other theo ries propose its hu- of size, post cra nial mate rial is poten tially con- man- me di ated in troduc tion as a rat preda tor sis tent with up to six addi tional species ( Fig- (Oliva and Talon 1996). Although Loyalties ure 12 ). All of the con fi rmed spe cies are ma terial was not included in a molec u lar study present elsewhere in the Loyalty Islands and of vari ation in Pacifi c boas (Austin 2000), it most are common where they occur. The does appear that there are deep diver gences most in ter esting re cords are Gehyra georgpot- among popu lations of C. bibroni in Fiji and Sa- thasti and Candoia bibroni . The fi rst of these moa, and there is no reason to suspect that spe cies, very recently described based in part these snakes are not native to the region. on Loyalty mate rial previ ously assigned to Ge- Previously Bavayia sauvagii was regarded as hyra vorax, is the only gekkonid gecko en- a widespread diplodactylid that oc curred over demic to the greater New Caledonian region much of the Grande Terre and its satel lite is- (it is also present in Vanuatu). It has previ ously lands as well as on Maré . Its absence in the been recorded from Maré and Dudun, to the other Loyalties was ten tatively ascribed to a south east of Tiga (Flecks et al. 2012), with lack of suit able rocky sub strates for di urnal re- only one record, with out precise local i ty, from fu gia (Sadlier and Bauer 1997). However, re- Lifou to the north. The con fi rma tion of the cent inves ti ga tions (Bauer and Jackman 2006) presence of this species on Tiga helps to clar- suggest that the sauvagii - like spe cies on Mar é ify its limited range in the Loyalties. Although are proba bly endemic to the Loyalties. Its oc- the simi larly sized G. vorax was tradi tion ally cur rence on Tiga, suggested but not con fi rmed con sumed by humans in Fiji (Gibbons and by ma te rial from archae olog i cal sites, is of Clunie 1984), there is no evi dence of this prac- both biogeo graphic and con ser vation inter est. tice in the Loyalties, so if the spe cies is no Interestingly, at least one spe cies of gekko- lon ger present on Tiga today, it is likely that nid gecko, Nactus pelagicus, is present in both habi tat alter ation would be the under lying older and younger screenings (Appendix 2). cause. Candoia bibroni, on the other hand, is Grant -Mackie et al. (2003) hypoth esized, known to have been regu larly eaten by peo- based on a small sam ple of reptile remains ple in the Loyalties, and this practice has con- from owl pellets at M é Aur é on the west coast tin ued on a limited scale to the pres ent, at of the New Caledonian mainland, that gekko- least on Lifou (Delauw 1990). The species is nid geckos may have arrived on the Grande now un common, at least in portions of its Terre some time during the period of human range in the Loyalties (R. Shine, pers. comm., col o ni za tion, perhaps even after the ar rival of Jan. 2014), although it is not known if this is Eu ro peans. This hypoth e sis has received fur- as so ci ated with human hunting pressure, hab- ther sup port from much more exten sive owl i tat modi fi ca tion, or in troduced mammals, pel let sam pling from Pinda ï , where > 5,000 liz- al though all could play a role. The verte brae ard bones spanning the period before and

Figure 12. ( A – M ) Silhouettes of the can di date squamate species recov ered from Tiga archae o log ical sites. All sil- houettes are drawn at the same scale based on the maxi mum values for each species. Species shown in white are those confi rmed to be pres ent in Tiga ar chaeo log ical sites; oth ers may be pres ent based on ma terial identi fi able only to higher order groups. ( A ) Phoboscincus garnieri, (B ) Lioscincus nigrofasciolatus, (C ) Emoia loyaltiensis, (D ) Emoia cyanura, ( E ) Caledoniscincus atropunctatus, (F ) Caledoniscincus haplorhinus, ( G ) Caledoniscincus austrocaledonicus, ( H ) Bavayia crassicollis, ( I ) B. aff. sauvagii, ( J ) Nactus pelagicus, ( K ) Lepidodactylus lugubris, (L ) Gehyra georgpotthasti, ( M ) Candoia bibroni . Scale bar equals 100 mm. Reptile Remains from Tiga, Loyalty Islands · Daza et al. 551 af ter the ar rival of the fi rst Pacifi c Islanders and the island for permis sion to under take this re- predating Euro pean settle ment have revealed search program. Permission for exca va tions in no gekkonid re mains, only diplodactylids and Tiga was granted by the President of the Loy- skinks (Kennedy 2011). The data from Tiga alty Islands Province. We thank Ross Sadlier dem on strate that at least in the Loyalties, Nac- and Cecilie Beatson (Aus tra lian Museum); José tus does pre date the Eu ropean peri od. This Rosado (Museum of Comparative Zoology, parthe no genetic species has a broad dis tribu- Harvard University); Darrel Frost and Edward tion in the Pacifi c, much of which is certainly Stanley (Amer i can Museum of Natural His- not human - medi at ed, so it is prob a ble that tory); Jens Vindum (California Academy of this gecko is native to the Loyalties. Differ- Sciences); Tony Gamble (James Ford Bell Mu- ences in the means of bone accu mu la tion, seum, University of Minnesota); Da vid Wake habi tat, and lizard commu nity structure could (Museum of Vertebrate Zoology, University of all con trib ute to differ ences seen on the California, Berkeley); Colin McCarthy (The Grande Terre and in the Loyalties. However, Natural History Museum, London); Kenneth the Loyalties, unlike the mainland, shares a Tighe, Jeremy Jacobs, and Kevin de Queiroz num ber of line ages with other island groups (Smithsonian Institution, National Museum of of the west ern Pacifi c (Bauer 1989, Sadlier and Natural History); and Elizabeth Glynne (Sam Bauer 1997, Bauer and Sadlier 2000) and Houston State University) for ac cess to com- might be expected to have histor ically har- par a tive mate rial used in this study. We are bored species absent on the main land. grateful to Alicia Kennedy who shared data Tiga presents an inter est ing case in that from her master ’s thesis. fresh samples of reptiles are unknown and ar- chaeo log i cally derived mate rial provi des the Literature Cited only ev i dence for the recent oc cur rence of liz- ards and snakes on the island. On the Grande Abdala, V. 1996. Osteolog í a craneal y relacio- Ter re, reli able mod ern re cords provide a base- nes de los geconinos sudamericanos (Rep- line against which subfos sil remains can be tilia: Gekkonidae). Rev. Esp. Herpetol. com pared and re veal that local and even island - 10:41– 54. wide ex tinc tions have taken place since hu man Anderson, A., C. Sand, F. Petchey, and T. H. oc cupa tion (Kennedy 2011). On Tiga, we start Worthy. 2010. Faunal extinc tion and hu- with the os te o log ical mate rial, which provi des a man habi tation in New Caledonia: Initial hy poth e sis about what may be present today. results and im pli cations of new re search at Some taxa (e.g., Lioscincus nigrofasciolatus, Cale- the Pindai Caves. J. Pac. Archaeol. 1:89– doniscincus spp.) are wide spread throughout 109. New Caledonia or on the Loyalties alone Anderson C. 1925. Notes on the extinct Che- ( Emoia cyanura, E. loyaltiensis ) and have gen er al- lonian Meiolania, with re cord of a new oc- ized hab itat re quirements allowing them to cur rence. Rec. Aust. Mus. 14:223 –342, pls. thrive in even dis turbed habi tats (Bauer and xxx – xl. Sadlier 2000) and may certainly be expected to Arnold, E. N. 1977. Little known geckos be present and locally abundant today. How- (Reptilia: Gekkonidae) from Arabia with ever, the re quirements of Gehyra georgpotthasti de scriptions of two new species from the for a more special ized forest hab itat (Flecks Sultanate of Oman: The scien tifi c re sults of et al. 2012) and the human harvesting of Can- the Oman fl ora and fauna survey 1975. J. doia bibroni (Delauw 1990) leave the current sta- Oman Stud. Special Rep. 1:81– 100. tus of these taxa on Tiga uncer tain and call for — — — . 1980. Recently ex tinct reptile popu- a thorough herpetofaunal sur vey of the island. la tions from Mauritius and Reunion, In- dian Ocean. J. Zool. 191:33 – 47. Austin, C. C. 2000. Molecular phylog e ny, acknowl edg ments popu la tion structure, and histor i cal bioge- We are grateful to the chief dom of ogra phy of Pacifi c island boas (Candoia ). Tiga/ Tokanod and to the dif fer ent clans of Copeia 2000:341– 352. 552 PACIFIC SCIENCE · Oc to ber 2015

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Appendix 1 dachner, 1870): Sk, AMS R2084, XR, USNM 322722, USNM 322723. Emoia atrocostata (Lesson, Comparative Specimens Examined 1830): Sk, AMS R40833, MCZ R15080, XR USNM Abbreviations for the type of prepa ra tion: EtOH = eth- 195779. Emoia caeruleocauda (De Vis, 1892): Sk, anol - pre served spec imens, C&S = cleared and stained AMS R25720, XR, USNM 122797, USNM 122857. prepa ration, HRXCT = high - reso lu tion X- ray computed Emoia concolor (Dumé ril, 1851): Sk, MCZR 16931, tomog ra phy, Sk = dry skele ton, XR = X - rays. XR, USNM 333338, USNM 333340. Emoia cyano- gaster (Lesson, 1830): Sk, MCZR15135, XR, USNM Diplodactylidae : Bavayia crassicollis: C&S, AMS 533324, USNM 533332. Emoia cyanura: Sk, MCZ R - R78285. Bavayia cyclura (Gü nther): C&S, AMS 14582, R- 72287, XR, USNM 28203, USNM 28204. R78092, MCZ A -27309, XR, USNM 146331, USNM Emoia kordoana (Meyer, 1874): Sk, AMS R64319, 515879. Bavayia montana Roux, 1913: C&S, AMS XR, USNM 499639. Emoia longicauda (Macleay): Sk, R77666, Sk, AMS R77665. Bavayia ro busta: Sk, AMS AMS R23905, USNM 336680, USNM 336681. R93710, MCZ A- 27346 – 27348, 27356 – 27357. Ba- Emoia loyaltiensis (Roux, 1913): HRXCT, AMNH R- vayia sauvagii: C&S, AMS R- 78350 –78351, CAS 60463. Emoia maculata Brown, 1954: Sk, AMS 16590 – 16696, Sk, AMS R77661, CAS 80823. Correlo- R80134, XR, USNM 121807. Emoia nigra (Jacquinot phus ciliatus Guichenot, 1866: Sk, NHMUK & Guichenot, 1853): Sk, AMS R138857, MCZR 85.11.16.7, JFBM 15825. Eurydactylodes vieillardi (Ba- 72514, XR, USNM 215235. Emoia pallidiceps (De Vis, vay, 1869): HRXCT, CAS 231986. Nebulifera robus ta 1890): Sk, AMS R14834, AMS R127604, XR, (Boulenger, 1885): Sk, CAS 75675. Oedura marmo- USNM 166268, USNM 166276. Emoia physicae (Du- rata: Sk, CAS 75405. Rhacodactylus auriculatus: C&S, mé ril & Bibron, 1839): Sk, AMS R60458, XR, CAS 165895– 165902; HRXCT, CAS 205486; Sk, USNM 567019. Graciliscincus shonae Sadlier, 1987: NHMUK 86.3.11.10, CAS 165891, CAS 165892. Sk, AMS R146534. Kanakysaurus viviparus Sadlier, Rhacodactylus trachyrhynchus: Sk, NHMUK 86.3.11.4. Bauer, Smith & Whitaker, 2004: Sk, AMS R162921. Gekkonidae: Gehyra australis (Gray, 1845): XR, USNM Lacertoides pardalis Sadlier, Shea & Bauer, 1997: Sk, 128538, USNM 203930. Gehyra baliola (Dumé ril, AMS R152644. Lioscincus maruia Sadlier, Whitaker & 1851): XR, USNM 167701. Gehyra borroloola King, Bauer, 1998: Sk, AMS R149958. Lioscincus nigrofas- 1983: XR, USNM 128537. Gehyra brevipalmata (Pe- ciolatus: Sk, AMS R78348, AMS R93712, MCZ A- ters, 1874): XR, USNM 546190, USNM 546197, 27345, MCZ A- 27310. Lioscincus novocaledoniae USNM 546203. Gehyra georgpotthasti : EtOH, AMS (Parker, 1926): Sk, AMS R78349. Lioscincus stein- R3558, AMS R13256, AMS B3834, AMS R1158, dachneri Bocage, 1873: Sk, AMS R153495. Lioscincus AMS R6668, AMS R7244, AMS R7257. Gehyra tillieri Ineich & Sadlier, 1991: Sk, AMS R148033. membranacruralis King & Horner, 1989: XR, Lioscincus vivae Sadlier, Bauer, Whitaker & Smith, USNM 518565. Gehyra mutilata (Wiegmann, 1835): 2004: Sk, AMS R167230. Marmorosphax tricol or: Sk, Sk, JFBM 15819, XR, USNM 584526, USNM AMS R93713, MCZ A - 27319 –27321. Nannoscincus 584527. Gehyra oceanica (Lesson, 1830): Sk, AMNH garrulus Sadlier, Bauer & Smith, 2006: Sk, AMS R - 27048, CAS 50045 [434], XR, USNM 322415, R151502, AMS R151521. Nannoscincus gracilis (Ba- USNM 322419. Gehyra papuana Meyer, 1874: XR, vay, 1869): Sk, AMS R77745, AMS R135195, XR, USNM 159828. Gehyra variegata (Dum é ril & Bi- USNM 268764. Nannoscincus greeri Sadlier, 1987: bron, 1836): XR, USNM 570629. Gehyra vorax: Sk, Sk, AMS R149495, AMS R165974. Nannoscincus AMS R179016. Hemidactylus frenatus: Sk, AMNH manautei Sadlier, Bauer, Whitaker & Smith, 2004: R - 71551, AMNH R - 71589, AMS R123990, JFBM Sk, AMS R167225. Nannoscincus mariei (Bavay, 15817, CAS 21784. Hemiphyllodactylus typus: C&S, 1869): Sk, AMS R146485, XR, USNM 515886, CAS 127969, Sk, AMS R125810. Lepidodactylus chris- USNM 515887. Nannoscincus rankini Sadlier, 1987: tiani Taylor, 1917: XR, USNM 305956, USNM Sk, AMS R149387. Nannoscincus slevini (Loveridge, 305957, USNM 305959. Lepidodactylus lugubris: Sk, 1941): Sk, AMS R135149. Sigaloseps deplanchei (Bavay, AMS R26611, AMS R179020, MCZ A- 27306 – 1869): Sk, AMS R125837, XR, USNM 515889, 27308, XR, USNM 494236, USNM 494238, USNM USNM 525888. Simiscincus aurantiacus Sadlier & 287039, USNM 287040, USNM 287045. Nactus ebo- Bauer, 1997: Sk, AMS R164368. Tropidoscincus varia- racensis (Macleay, 1877): Sk, MVZ 77610, MVZ bilis (Bavay, 1869): Sk, AMS R164374, uncatalogued 77615, XR, USNM 561877, USNM 561882. Nactus spec imen, XR, USNM 149680, USNM 268762. pelagicus: Sk, AMS R179023, AMS (no number), XR, Boidae: Candoia bibroni: Sk, MCZ R14516, MCZ R15018. USNM 343831, USNM 343833, USNM 343836. Scincidae: Caledoniscincus austrocaledonicus: Sk, uncata- Appendix 2 logued, MCZ A - 27301 –27305, XR, USNM 267845, USNM 515855. Caledoniscincus festivus Sadlier, 1987: Material by Excavation Site and Depth of Recovery Sk, AMS R116036 . Celatiscincus euryotis (Werner, 1910): Sk, AMS R153990. Cryptoblepharus egeriae SVL* indi cates the es ti mated size based on com pa ra ble (Boulenger, 1888): Sk, AMS R83160. Cryptoblepharus ele ments of artic ulated speci mens. plagiocephalus (Cocteau, 1836): Sk, AMS R142565, LTD - 14A: Rock- shel ter lo cated on the west coast of XR USNM 54063. Cryptoblepharus virgatus (Gar- Tiga, about 50 m in land. Human occu pa tion starts man, 1901): Sk, AMS R85791. Emoia adspersa (Stein- at 100 cm below surface, dated to the middle of the Reptile Remains from Tiga, Loyalty Islands · Daza et al. 557

fi rst millen nium A.D. Most of the fi ll is dated to below surface, dated to the second half of the fi rst the second mil len nium A.D. 45– 50 cm: Lioscincus ni- mil len nium A.D. The long- term occu pa tion fi ll, grofasciolatum, dentary (1L, > 9 mm, SVL* = 52.0 mm). concen trated in the upper 70 cm of the stratig ra phy, Gehyra georgpotthasti, maxilla (1L, 16.5 mm, is dated to the last 600 yr. 50 – 60 cm: Scincidae, SVL* = 123.8 mm); 50– 55 cm: Nactus pelagicus, frontal incertae sedis, femur (1L, 13 mm, SVL* = 76.6 mm); (1, > 7 mm, SVL* = 67.3 mm). Gehyra georgpotthasti, 60– 70 cm: Candoia bibroni, verte bra (3, ∼ 5 mm). Scin- dentary (1L, 16.5 mm, SVL* = 113.4 mm); 55– 60 cm: cidae, incertae sedis, fe mur (1R, 8 mm, SVL* = Nactus pelagicus, frontal (1, 7.8 mm, SVL* = 75.1 mm); 50.8 mm; 1R, 12.5 mm, SVL* = 76.1 mm) and in- 60 –65 cm: Bavayia crassicollis, frontal (1, > 7 mm, nomi nate bone (1), and cau dal verte brae (3); 100 – SVL* = 60.8 mm), innom i nate bone (1L). Gekkota, 110 cm: Nactus pelagicus, frontal (1, > 8 mm; SVL* = fe mur (2L, > 8.5 mm. SVL* = 57.2 mm); 65– 70 cm: 80.3 mm). Emoia loyaltiensis, max illa (1L, 1R, 8.6 mm, SVL* = LTD - 18C: Rock- shel ter located on the northwest coast 66.5 mm). Gekkota, hu merus (1L, 1R, 7 mm, of Tiga. Human occu pa tion in pit C starts at 65 cm SVL* = 54.4), femur (1R, 7 mm, SVL* = 47.1 mm). below surface, dated to 2530 yr BP. Most of the fi ll is Gehyra georgpotthasti, dentary (1L, 20 mm, SVL* = dated to the fi rst millen nium A.D. 0– 15 cm: Candoia 137.5 mm); 70– 75 cm: Lioscincus nigrofasciolatum bibroni, trunk ver tebra (2, ∼ 5mm). Scincidae, incertae fron tal (1, 7.7 mm, SVL* = 108.86 mm), pari etal sedis, fe mur (1L, 11 mm, SVL* = 68.0 mm). Gekkota, (1, 7.5 mm length × 5.4 wide, SVL* = 98.13), innom i- fe mur (1L, 8 mm, SVL* = 50.5 mm); 15– 25 cm: Gek- nate bones (1L, 1R). Scincidae, incertae sedis, humerus kota, humerus (1R, 7 mm, SVL* = 48.2), femur (1R, (1R, 10.5 mm, SVL* = 76.8 mm), femur (1R, 13 mm, 9 mm, SVL* = 60.7 mm); 35– 45 cm: Candoia bibroni, SVL* = 76.6 mm). Bavayia sp., dentary (1L, > 9 mm, trunk ver tebra (2, ∼ 5 mm). Gekkota, femur (1R, SVL* = 67.4 mm). Gehyra georgpotthasti, dentary (1R, 8.5 mm, SVL* = 56.7). Two unde ter mined appen dic- ante rior portion only); 75– 80 cm: Scincidae, incertae ular bones (incom plete). Scincidae, incertae sedis, sedis, hu merus (1L, 1R, 6.5 mm, SVL* = 55.8 mm; presacral ver tebra (3); 45– 55 cm: Candoia bibroni, 1L, 11.1 mm, SVL* = 78.6 mm); 85– 90 cm: Scincidae, trunk ver tebrae (8), ribs (3). Undetermined femur incertae sedis, humerus (1L, 7.1 mm, SVL* = 57.2 mm), (1, > 8.5 mm). Emoia loyaltiensis, dentary (1L, > 9 mm, fe mur (1R, 6 mm, SVL* = 44.9 mm; 1L, 7.6 mm, SVL* = 63.7), Gekkota, humerus (1R, 6.5 mm, SVL* = 49.2 mm). Nactus pelagicus, frontals (2, 7 mm, SVL* = 58.4), femur (1L, 9 mm; SVL* = 60.7 mm); SVL* = 67.37 mm). Scincidae, incertae sedis, presacral 55– 65 cm: Candoia bibroni, trunk ver tebrae (2, verte brae (2, ∼ 4 mm). ∼ 5mm). Emoia loyaltiensis, max illa (1L, > 5 mm), com- LTD - 18A: Rock- shel ter lo cated on the northwest Coast pound bone (1L, 8 mm, SVL* = 68.5 mm). Lioscincus of Tiga. Human occu pa tion in pit A starts at 65 cm nigrofasciolatum, quadrate (1L). Bavayia crassicollis, below surface, dated to 2,100 yr B.P. Most of the fi ll max illa (1L, < 8.5 mm, SVL* = 68.1 mm), dentary is dated to the fi rst millen nium A.D. 50– 55 cm: Ba- (1L, 8.9 mm; SVL* = 66.7 mm), compound bone (1L, vayia sp. dentary (1R, 9.2 mm, SVL* = 69.33 mm); 7 mm; 1R, 9 mm, SVL* = 74.3 mm). Bavayia sp., 65 –70 cm: Bavayia sp., dentary (1L, > 8.5 mm, braincase (1, 4.6 mm, SVL* = 50.3 mm). Gehyra SVL* = 64.06 mm). georgpotthasti, innom i nate bone (1). Two inde ter mi- LTD -02A: Rock - shel ter lo cated on the northwest coast nate appen dic u lar bones; 65 –75 cm: Scincidae, incer- of Tiga. Human occu pa tion starts at over 190 cm tae sedis, dentary (1L, > 9 mm, SVL* = 38.0 mm).