geodiversitas 2022  44  7 Directeur de la publication / Publication director : Bruno David, Président du Muséum national d’Histoire naturelle Rédacteur en chef / Editor-in-chief : Didier Merle Assistant de rédaction / Assistant editor : Emmanuel Côtez (geodiv@mnhn.fr) Mise en page / Page layout : Emmanuel Côtez Comité scientifique / Scientific board : Christine Argot (Muséum national d’Histoire naturelle, Paris) Beatrix Azanza (Museo Nacional de Ciencias Naturales, Madrid) Raymond L. Bernor (Howard University, Washington DC) Alain Blieck (chercheur CNRS retraité, Haubourdin) Henning Blom (Uppsala University) Jean Broutin (Sorbonne Université, Paris, retraité) Gaël Clément (Muséum national d’Histoire naturelle, Paris) Ted Daeschler (Academy of Natural Sciences, Philadelphie) Bruno David (Muséum national d’Histoire naturelle, Paris) Gregory D. Edgecombe (The Natural History Museum, Londres) Ursula Göhlich (Natural History Museum Vienna) Jin Meng (American Museum of Natural History, New York) Brigitte Meyer-Berthaud (CIRAD, Montpellier) Zhu Min (Chinese Academy of Sciences, Pékin) Isabelle Rouget (Muséum national d’Histoire naturelle, Paris) Sevket Sen (Muséum national d’Histoire naturelle, Paris, retraité) Stanislav Štamberg (Museum of Eastern Bohemia, Hradec Králové) Paul Taylor (The Natural History Museum, Londres, retraité) Couverture / Cover : Réalisée à partir des Figures de l’article/Made from the Figures of the article. Geodiversitas est indexé dans / Geodiversitas is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Physical, Chemical, and Earth Sciences® – Scopus® Geodiversitas est distribué en version électronique par / Geodiversitas is distributed electronically by: – BioOne® (http://www.bioone.org) Les articles ainsi que les nouveautés nomenclaturales publiés dans Geodiversitas sont référencés par / Articles and nomenclatural novelties published in Geodiversitas are referenced by: – ZooBank® (http://zoobank.org) Geodiversitas est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris Geodiversitas is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie, Comptes Rendus Palevol Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 diff.pub@mnhn.fr / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2022 ISSN (imprimé / print) : 1280-9659/ ISSN (électronique / electronic) : 1638-9395 First fossil of Varanus Merrem, 1820 (Squamata: Varanidae) from the Miocene Siwaliks of Pakistan Andrea VILLA Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA/ICP, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona (Spain) andrea.villa@icp.cat (corresponding author) Massimo DELFINO Dipartimento di Scienze della Terra, Università degli Studi di Torino, via T. Valperga Caluso 35, 10125 Torino (Italy) and Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA/ICP, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona (Spain) massimo.delfino@unito.it Submitted on 23 May 2020 | accepted on 15 September 2020 | published on 14 February 2022 urn:lsid:zoobank.org:pub:B85B92E1-66B0-4BE9-8085-07CDE2140D7F Villa A. & Delfino M. 2022. — First fossil of Varanus Merrem, 1820 (Squamata: Varanidae) from the Miocene Siwaliks of Pakistan. Geodiversitas 44 (7): 229-235. https://doi.org/10.5252/geodiversitas2022v44a7. http://geodiversitas.com/44/7 ABSTRACT KEY WORDS In the ‘50s of the XX century, a German expedition lead by Richard Dehm collected a large amount Reptilia, of fossil remains from northern Pakistan. Among these was an isolated trunk vertebra of a lizard, middle Miocene, which is here referred to Varanus sp. The collecting site of this specimen is not precisely known, but late Miocene, Neogene, it most likely comes from middle to early late Miocene Siwalik sediments of the Chinji Formation. Potwar Plateau, This is the first published record of a fossil lizard from the Neogene of Pakistan and adds to the very Chinji Formation, Asia, patchy record of Varanus Merrem, 1820 in Asia. It further supports previous reconstructions of a new record. warm climate for the middle to early late Miocene of the Siwaliks. RÉSUMÉ Premier fossile de Varanus Merrem, 1820 (Squamata: Varanidae) du Miocène des Siwaliks du Pakistan. Dans les années 50 du XXème siècle, une expédition allemande dirigée par Richard Dehm a recueilli MOTS CLÉS une grande quantité de restes fossiles dans le nord du Pakistan. Parmi ceux-ci se trouvait une vertèbre Reptilia, isolée du tronc d’un lézard, qui est ici désigné comme Varanus sp. Le site de collecte de ce spécimen Miocène moyen, n’est pas connu précisément, mais il provient très probablement de sédiments des Siwaliks du Miocène Miocène supérieur, Néogène, moyen à supérieur (Formation de Chinji). Il s’agit de la première occurrence publiée d’un lézard fossile Plateau de Potwar, du Néogène du Pakistan. Il s’ajoute à l’enregistrement très disparate du genre Varanus Merrem, 1820 Chinji Formation, Asie, en Asie. Cette occurrence soutient également les reconstructions précédentes d’un climat chaud, du signalement nouveau. Miocène moyen au début du Miocène supérieur, dans les Siwaliks. GEODIVERSITAS • 2022 • 44 (7) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 229 Villa A. & Delfino M. A B Fig. 1. — Labels related to SNSB-BSPG 1956 II 2010: original label (A) and inventory label (B). INTRODUCTION between Marianwala and Kas. Both these villages are located south of Chinji (Fig. 2), in an area where the Chinji Forma- In 1955-1956, a palaeontological joint expedition of the tion crops out. Numerous collecting sites are marked in the Institut für Paläontologie und historische Geologie of the surroundings of these two villages in the original maps from Ludwig-Maximilians-Universität München (today Depart- the expedition (AV, pers. obs.), but neither on the maps nor ment für Geo- und Umweltwissenschaften, Paläontologie on the original diaries any lizard vertebra is noted despite the und Geobiologie) and the Bayerische Staatssammlung für original label clearly states that the fossil was recognised as Paläontologie und Historische Geologie (today Staatliche such (Fig. 1A). The Chinji Formation is currently considered Naturwissenschaftliche Sammlungen Bayerns-Bayerische middle to early late Miocene in age (11.2-14.2 Ma; Barry Staatssammlung für Paläontologie und Geologie, SNSB- et al. 2002; Patnaik 2016). BSPG) lead by Prof. Richard Dehm explored northern Paki- stan in search for fossils (Dehm et al. 1958). They returned to Munich with an extensive collection of remains, coming MATERIAL AND METHODS from Neogene sediments exposed near the villages of Chinji, Dhok Pathan, and Nagri (Siwalik Group) and now stored The subject of this study is a single vertebra, numbered SNSB- in the SNSB-BSPG. Studies on the fossil mammal remains BSPG 1956 II 2010. The vertebra was photographed with a recovered in this and previous expeditions in the area by the Leica M205 microscope equipped with the Leica Application Munich team reported the presence of deinotheres (Dehm et al. Suite V 4.10 at the Università degli Studi di Torino. Photo- 1963), equids (Hussain 1971), primates (Dehm 1983, 1984; graphs of the labels were taken with a Samsung WB252F Pickford 2010), rhinocerotids (Heissig 1972), and tragulids Digital Camera. The terminology used for the vertebral (Guzmán-Sandoval & Rössner 2021). Pickford (2010) further laminae follows that used by Tschopp (2016). mentioned still unpublished occurrences of other groups, including anthracotheres, carnivorans, equids, gomphotheres, ruminants, and suids. The Pakistani collection in the SNSB- SYSTEMATICS BSPG lacks any trace of amphibian remains (AV, pers. obs.), whereas reptiles are represented by crocodylians, turtles, and Order SQUAMATA Oppel, 1811 squamates. Of these, only snakes were studied by Hoffstetter Family Varanidae Gray, 1827 (1964), who described vertebrae of an indeterminate Python Genus Varanus Merrem, 1820 and of a new large acrochordid, Acrochordus dehmi. Fossils of crocodylians and turtles will be the subject of Varanus sp. future dedicated works, but we here add to the squamate fauna by describing the only other fossil of this group of reptiles Description that we were able to find in the SNSB-BSPG collections: an SNSB-BSPG 1956 II 2010 (Fig. 3) is a large and elongated isolated lizard vertebra. No precise locality was available for trunk vertebra, with a procoelous centrum. It is moderately this fossil, but some information can be retrieved from the well preserved, but the anterior part and most of its left lat- labels (Fig. 1). It was collected by members of Dehm’s expe- eral surface are covered by a hard sandstone matrix that was dition on the first day of February 1956 (Fig. 1A), somewhere not removed in order not to damage the fossil. In ventral 230 GEODIVERSITAS • 2022 • 44 (7) Fossil Varanus from the Miocene of Pakistan A B 100 km 60 mi Dhok Pathan Chinji Lahore 200 km PUNJAB 100 mi Fig. 2. — Map of Pakistan (A) and of northern Punjab (B), showing the area where the villages of Marianwala and Kas are located (small square in B) south of Chinji. view (Fig. 3A), the centrum is subtriangular, with posteriorly the prespinal lamina sensu Tschopp 2016), but it thickens converging and straight lateral margins. The ventral surface is slightly by the posterior end of the neural arch where it is flat, without keel. In spite of the matrix covering, the ventral broken off (Fig. 3D). The dorsal surface of the neural arch margin of the anterior cotyle is clearly located posteriorly com- displays a distinct striation (Fig. 3D), composed of fine and pared to the dorsal margin (Fig. 3A); thus, the anterior cotyle dense striae that run posterodorsally from the sides of the arch was facing anteroventrally. In posterior view (Fig. 3B), the towards the neural spine. On each side of the same surface, centrum is dorsoventrally compressed. The posterior condyle a robust and dorsally-rounded lamina connects the posterior is strongly eroded, but a distinct precondylar constriction is corner of each prezygapophysis with the base of the neural spine still recognisable in ventral view (Fig. 3A). The width of the around midlength of the vertebra (Fig. 3C, D, F). We named centrum at the constriction is 5 mm, whereas the anterior this structure mediospinal lamina. It marks the anterior mar- cotyle can be estimated as about 8 mm wide. This would gin of the raised posterior half of the neural arch. Posteriorly, indicate a constriction/cotyle width ratio of 0.63. On the the spinopostzygapophyseal laminae are robust but not sharp right lateral surface (Fig. 3C), the base of the synapophysis (Fig. 3B). Due to the covering sediment, it is not possible to is visible, even though the rest of the latter is not preserved describe the centropostzygapophyseal laminae. Further, the anymore. Nevertheless, it was dorsoventrally elongated. The broken neural spine hinders the description of a postspinal vertebra has no postzygoprezygapophyseal lamina, but a lamina, in case it was present (as is usually the case at least in very poorly distinct posterior centrosynapophyseal lamina Varanus exanthematicus; Tschopp 2016). The preserved mini- is present (Fig. 3C). In dorsal view (Fig. 3D), a distinct mum length of the vertebral centrum (measured at midline interzygapophyseal constriction is visible. Being covered by starting from the ventral margin of the cotyle) is 11 mm. sediment, the prezygapophyses are not clearly visible. The postzygapophyses are better exposed. Zygapophyses are sub- Identification circular to suboval and dorsally inclined of about 45°. Each The following combination of features clearly indicates that prezygapophysis is separated by the pars tectiformis of the the Pakistani trunk vertebra found in the SNSB-BSPG col- neural arch by a deep groove (Fig. 3D). The pars tectiformis lections pertains to Varanus (Hoffstetter 1943; Estes 1983; does not bear a zygosphene, and no zygantrum is visible on Smith et al. 2008; Holmes et al. 2010): anterior cotyle facing the posterior edge of the neural arch. The posterior half of ventrally due to posteriorly-located ventral margin; presence of the latter is raised compared to the pars tectiformis (Fig. 3E, precondylar constriction; complete absence of a more-or-less 3F). The neural spine is well developed, even though broken developed zygosphene/zygantrum complex; and presence of along its entire length. It starts at the anterior margin of the dorsal striation. The newly-named structure, the mediospinal neural arch and is laminar for most of its length (originating lamina, also supports the identification of the vertebra as a GEODIVERSITAS • 2022 • 44 (7) 231 Villa A. & Delfino M. varanid. This structure has been also recognised, but not named, Miocene of the Chinji Formation. To our knowledge, this is by Smith et al. (2008), who reported its sharp development the first published record of a non-snake squamate from the in Varanus and Saniwa, but not in Heloderma. In contrast, it Neogene of Pakistan. Another occurrence of Pakistani Vara- appears to be absent in other lizard groups, such as lacertids nus is listed in the fosFARbase database (Böhme & Ilg 2003) (Tschopp 2016), scincids (Camaiti et al. 2019), and anguids from Winnewala. The only formation reported to crop out at (Čerňanský et al. 2019). A more extensive survey of the distri- Winnewala is the Dhok Pathan Formation (Hoffstetter 1964; bution and development of this structure in lizards could help Head 2005), which is dated between 10.1 and c. 3.5 Ma, late in the future to better understand its taxonomic significance. Miocene to early Pliocene, by Barry et al. (2002). Given that, Isolated vertebrae of Varanus are not diagnostic at the species fossils coming from this locality should be younger than the level (see literature in Villa et al. 2018). Thus, SNSB-BSPG herein studied vertebra supposedly coming from the Chinji 1956 II 2010 cannot be further identified. The development Formation. Unfortunately, we were unable to find Varanus of the precondylar constriction compared to the posterior remains from Winnewala in the SNSB-BSPG collection, condyle is considered useful to discriminate between dif- where they should be stored, and consequently to confirm this ferent Varanus subclades (Conrad et al. 2011; Delfino et al. occurrence and potentially make comparisons with SNSB- 2013), but this character is not evaluable in SNSB-BSPG BSPG 1956 II 2010. 1956 II 2010 because of its complete erosion. Nevertheless, In the Asian continent, Varanus fossils are known from the if this vertebra represents an adult (or subadult) individual, early Miocene of China and Kazakhstan (Böhme & Ilg 2003; the minimum length of the vertebral centrum suggests that Malakhov 2005; Vasilyan et al. 2017), the middle Miocene it probably belonged to a small-bodied monitor species (see of Kazakhstan (Böhme & Ilg 2003), the middle to early late measurements in e.g., Conrad et al. 2012, and Vasilyan & Miocene of Pakistan (this work), the late Miocene of Armenia, Bukhsianidze 2020), with a snout-vent length possibly higher Georgia, India, Kazakhstan, and Russia (Chkhikvadze 1985; than 300 mm and not exceeding 500 mm. A confident esti- Rage et al. 2001; Böhme & Ilg 2003; Vasilyan et al. 2017; mate of the size is not advisable due to uncertainties regarding Vasilyan & Bukhsianidze 2020; Čerňanský et al. 2020), the the real position of the vertebra in the column, as well as the Pliocene of India and Turkey (Falconer 1868; Lydekker 1886, unknown intracolumnar variation. 1888; Hocknull et al. 2009; Sen et al. 2017), as well as the Quaternary of India, Indonesia, and Thailand (Falconer 1868; Lydekker 1886, 1888; Hooijer 1972; Patnaik et al. 2008; DISCUSSION Hocknull et al. 2009; Suraprasit et al. 2016). Most of the Quaternary remains are referred to extant species, whereas all The squamate fossil record in Pakistan is so far dominated by the Neogene ones are not identified at the species level. The snakes. Most occurrences come from Neogene (i.e., Siwalik) only exception to this frame is Varanus sivalensis, an extinct sediments, with only few others coming from the Paleogene. species described from the Pliocene to Early Pleistocene of The oldest squamate fossils from Pakistan are two vertebrae India (Falconer 1868; Lydekker 1886, 1888; Hocknull et al. of Gigantophis from the Danian (Rage et al. 2014). Younger 2009). However, Hocknull et al. (2009) casted doubts on the Paleogene snakes are then found in Lutetian (indeterminate real taxonomic identity of the three specimens assigned to erycines, boines and unidentified snakes from Chorlakki; this taxon, a humerus and two vertebrae: the authors stated Rage 1987) and Chattian (indeterminate snakes from Paali that the humerus, which has Pliocene age, might indeed Nala; Böhme & Ilg 2003) sediments. Snakes found in the show sufficient differences to sustain its own taxonomic Neogene Siwaliks recorded the presence of various different status, whereas the dorsal vertebrae, found in Early Pleisto- taxa (Hoffstetter 1964; Böhme & Ilg 2003; Head 2005): the cene deposits, are comparable with Varanus salvator in their acrochordid A. dehmi (Burdigalian to Tortonian); the boids morphology and size. Considering all of this, our knowledge Python sp. (Burdigalian to Messinian), cf. Erycinae indet. of the evolutionary history of Varanus in Asia is still rather (Serravallian), and ?Boidae indet. (Serravallian); indeterminate patchy and clearly needs more in-depth studies based on the booids s.l. (Burdigalian to Messinian); the colubroids Gansophis recovery of further material. potwarensis (Messinian), Colubroidea morph. A (Serravallian), This bears significance not only for the history of this Colubroidea morph. B (Tortonian to Messinian), as well as clade in this continent, but also for its European and Afri- indeterminate colubroid snakes (Burdigalian to Messinian); can representatives. The biogeographical origin of the genus the “colubrines” Chotaophis padhriensis (Tortonian), Sivaophis Varanus is still a debated issue (Villa et al. 2018), with both downsi (Serravallian to Messinian), and indeterminate “colu- African (Fuller et al. 1998; Ast 2001; Holmes et al. 2010) brine” snakes (Langhian to Tortonian); indeterminate natricines and Eurasian (Amer & Kumazawa 2008; Conrad et al. 2012; (Tortonian); and the elapids Bungarus sp. (Tortonian) and Ivanov et al. 2018) origins proposed as possible alternatives. ?Elapidae indet. (Tortonian to Messinian). Compared to the Nevertheless, the origin is to be found in the Paleogene, as snake record, lizards are extremely underrepresented among indicated by the oldest Varanus remains coming from the Pakistani fossils, with only the agamid Tinosaurus sp. reported Eocene of Egypt (Holmes et al. 2010). The absence of confi- from the Lutetian of Chorlakki (Rage 1987) and indeterminate dently diagnosable Varanus species prior to the early Miocene scincids from the Chattian of Paali Nala (Böhme & Ilg 2003). and, in Asia, even to pre-Quaternary times, however, hinders a To these, we now add Varanus sp. from the middle to early late better understanding of this problem. Detailed identification 232 GEODIVERSITAS • 2022 • 44 (7) Fossil Varanus from the Miocene of Pakistan A pl B spl C pcl s D ml E F pt ml Fig. 3. — Varanus sp. from the Miocene of Pakistan: trunk vertebra (SNSB-BSPG 1956 II 2010) in ventral (A), posterior (B), right lateral (C), dorsal (D), anterior (E), and left lateral (F) views. Abbreviations: ml, mediospinal lamina; pcl, posterior centrosynapophyseal lamina; pl, prespinal lamina; pt, pars tectiformis; s, syn- apophysis; spl, spinopostzygapophyseal lamina. Scale bar: 5 mm. of the Asian fossil monitor lizards could have a direct role in Indian subcontinent and both Africa and likely Europe in the unveiling the origin of the populations that inhabited Europe considered time interval. Further data are therefore needed at least from the early Miocene (Delfino et al. 2013) to the to shed light on the biogeographic relationships of Neogene Middle Pleistocene (Georgalis et al. 2017). A recent revision lizards from Pakistan and the Indian subcontinent as a whole. of European monitors by Villa et al. (2018) suggested the In palaeoenvironmental terms, Varanus indicates a warm presence of at least two lineages in Europe, the second one palaeoclimate (mean annual temperature ranging from 14.8°C (represented by Varanus marathonensis from the latest mid- to 28.1°C according to Böhme 2003) for the Chinji Forma- dle to the late Miocene) originating from an Asiatic ancestor. tion. This partially agrees with the landscape reconstructions It would be, thus, interesting to comprehend if and which based on palaeobotanical studies (Srivastava et al. 2014; Patnaik of the Asian Varanus, including the one from Pakistan here 2016) suggesting dominance of evergreen to deciduous forests described, could be related to this dispersal wave. reflecting warm and humid conditions with high rainfall during This cannot be stated considering only morphological the middle Miocene of the Siwaliks. The Siwalik sediments in characters, given that known Asian remains of Varanus are Pakistan deposited in a fluvial environment (Barry et al. 2002; represented in most cases by poorly-significant vertebral Head 2005). As far as the Chinji Formation is concerned, the material. Nevertheless, Siwalik snakes could be potentially of riverine system originated fine-grained sandstone deposits that are help in drawing some hypotheses about relationships between still present as a covering on at least some of the fossil remains, European and Pakistani squamate faunas. Most Siwalik snake such as the herein studied Varanus vertebra but also other fossils genera and species are not shared with Europe (Head 2005), stored in the SNSB-BSPG (AV, pers. obs.). The most abundant thus suggesting ecological barriers and different biogeographic squamate found in the whole Pakistani Siwalik sequence is A. units. Based on stratigraphic distribution of snake fossils, dehmi (Hoffstetter 1964; Head 2005), which is considered a Head (2005: 30) stated that “partition of European and South fully-aquatic animal. This is particularly evident in the Chinji Asian biogeographic theaters occurred no later than middle and Nagri Formations, but subsequently, there is a rise in the Miocene”. However, it should be also noted that the pattern occurrence of terrestrial and semi-aquatic taxa from around 10 reported for snakes does not reflect the mammalian palaeo- Ma onwards (Dhok Pathan Formation; Head 2005). Based on biogeographic dynamics (Patnaik 2016): as a matter of fact, rough correlations between monsoonal precipitation patterns, mammal faunal exchanges were indeed present between the the transition towards a C4-dominated vegetation, and the GEODIVERSITAS • 2022 • 44 (7) 233 Villa A. & Delfino M. increasing diversity in Siwalik snakes, Head (2005) suggested Böhme M. & Ilg A. 2003. — fosFARbase, http://www.wahre- a possible link between the latter and increasing seasonality staerke.com/ (accessed April 2020). in the area and diversification of the available habitats, which Camaiti M., Villa A., Wencker L. C. M., Bauer A. M., Stan- ley E. L. & Delfino M. 2019. — Descriptive osteology and would have favoured the onset of a more diverse reptilian fauna. patterns of limb loss of the European limbless skink Ophiomorus Varanus currently displays a wide variety of ecological adaptation punctatissimus (Squamata, Scincidae). Journal of Morphology 235: and includes species strongly linked with water environments 313-345. https://doi.org/10.1111/joa.13017 (e.g., V. salvator, which nowadays is widely spread in Southeast ČerŇanský A., Yaryhin O., Ciceková J., Werneburg I., Hain M. & Asia; Pianka et al. 2004; Eidenmullen 2007). Acquiring infor- Klembara J. 2019. — Vertebral comparative anatomy and mor- phological differences in anguine lizards with a special reference mation on the ecological preference of the species represented to Pseudopus apodus. The Anatomical Record 302: 232-257. https:// by SNSB-BSPG 1956 II 2010 is impossible, even though the doi.org/10.1002/ar.23944 finding of only one remain in the Pakistani Siwaliks may suggest ČerŇanský A., Syromyatnikova E. V. & Jablonski D. 2020. — The a low relation with water for the represented monitor when first record of amphisbaenian and anguimorph lizards (Reptilia, considering the pattern showed by snakes. Further studies of Squamata) from the upper Miocene Solnechnodolsk locality in Russia. Historical Biology 32 (7):869-879. https://doi.org/10.10 squamate fossils from the area are strongly needed to enhance 80/08912963.2018.1539973 our knowledge of their distribution, evolution, and ecology in Chkhikvadze V. M. 1985. —Preliminary results of studies on the Neogene of Pakistan. tertiary amphibians and squamate reptiles of the Zaisan Basin, in Darevsky I. (ed.), Problems in Herpetology, Proceedings of the 6th All-Union Herpetological Conference, Tashkent, 18-20 September 1985. Leningrad, Nauka: 234-235. Acknowledgements Conrad J. L., Ast J. C. A., Montanari S. & Norell M. A. The study of SNSB-BSPG 1956 II 2010 benefited significantly 2011. — A combined evidence phylogenetic analysis of Angui- from the help of Gertrud Rössner (Munich), who assisted AV morpha (Reptilia: Squamata). Cladistics 27: 230-277. https:// in defining the possible provenance and age of the specimen doi.org/10.1111/j.1096-0031.2010.00330.x and shared information about Dehm’s expedition to Pakistan. Conrad J. L., Balcarcel A. M. & Mehling C. M. 2012. — Earliest example of a giant monitor lizard (Varanus, Varanidae, We also thank Madelaine Böhme (Tübingen) for discussions Squamata). PLoS ONE 7: e41767. https://doi.org/10.1371/ about the Winnewala Varanus occurrence, the two reviewers, journal.pone.0041767 Davit Vasilyan (Porrentruy) and Martin Ivanov (Brno), for Dehm R. 1983. — Miocene hominoid primate dental remains from their valuable comments that improved an earlier version of the the Siwaliks of Pakistan, in Ciochon R. L. & Corruccini R. S. manuscript, as well as the editor Emmanuel Côtez (Paris). This (eds), New Interpretations of Ape and Human Ancestry. Springer, Boston: 527-537. https://doi.org/10.1007/978-1-4684-8854-8_20 work originated during a visit to the SNSB-BSPG collection in Dehm R. 1984. — Molar size enlargement in Siwalik hominoid 2017 supported by an EAVP Research Grant provided to AV primates and its significance. Courier Forschungsinstitut Senck- by the European Association of Vertebrate Palaeontologists. enberg 69: 49-52. Oliver Rauhut (Munich) is thanked for help while visiting Dehm R., Oettingen-Spielberg T. zu & Vidal H. 1958. — the collection back then. During part of the development of Paläontologische und geologische Untersuchungen im Tertiär von Pakistan. I. Die Münchner Forschungsreise nach Pakistan this work, AV was funded by the Alexander von Humboldt 1955-1956. Abhandlungen der Mathematisch-Naturwissenschaftliche Foundation through a Humboldt Research Fellowship. 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Palevol 15: 889-902. https://doi.org/10.1016/j.crpv.2015.11.004 https://doi.org/10.1371/journal.pone.0207719 Submitted on 23 May 2020; accepted on 15 September 2020; published on 14 February 2022. GEODIVERSITAS • 2022 • 44 (7) 235