First identification of Decennatherium Crusafont, 1952 (Mammalia, Ruminantia, Pecora) in the Siwaliks of Pakistan

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Original article

First identification of Decennatherium Crusafont, 1952 (Mammalia, Ruminantia, Pecora) in the Siwaliks of Pakistan§ Marı´a Rı´os a,*, Melinda Danowitz b, Nikos Solounias c,d a Departamento de Paleobiologı´a, Museo Nacional de Ciencias Naturales-Consejo Superior de Investigaciones Cientı´ficas, 2 Gutie´rrez Abascal, 28006 Madrid, Spain b Department of Pediatrics, Alfred I. duPont Hospital for Children 19806 Wilmington, DE, USA c Department of Anatomy, New York Institute of Technology College of Osteopathic Medicine, 8000 Northern Boulevard, 11568 Old Westbury, NY, USA d Department of Paleontology, American Museum of Natural History, Central Park West at 79th Street, 10024 New York, NY, USA

A R T I C L E I N F O

A B S T R A C T

Article history: Received 7 December 2018 Accepted 2 October 2019 Available online xxx

Previously undescribed remains of a new large giraffid have been identified from the late Miocene Siwaliks Hills of the Potwar Plateau in Pakistan. This taxon is very intimately related to the late Miocene giraffid genus Decennatherium, previously only certainly identified in the Iberian Peninsula, and with some possible remains assigned to Decennatherium crusafonti from Alia¨ba¨d (Iran). The new material collected in the Siwaliks Hills shows a high morphological similarity with the Spanish remains of the genus, especially the early Vallesian Decennatherium pachecoi. Two previously undescribed ossicone fragments from the same area are also described and identified as cf. Decennatherium. These findings represent the easternmost occurrence of the genus Decennatherium and show a late Miocene migration of the genus throughout the Iberian Peninsula and Southern Asia.

C 2019 Published by Elsevier Masson SAS.

Keywords: Decennatherium Giraffidae Cetartiodactyla Vallesian Miocene Palaeobiogeography

1. Introduction Giraffids are a relict group of ruminants characterized by the possession of bilobed canines and a particular kind of cranial appendages called ossicones (Hamilton, 1978; Solounias, 2007; Davis et al., 2011). Previously undescribed remains of a new large giraffid have been identified from the late Miocene Siwaliks Hills of Pakistan. The new species is named Decennatherium asiaticum. The age range of this taxon is 13.5 to 8.2 Ma. This new species appears to be intimately related to the late Miocene giraffid genus Decennatherium, previously only surely identified in the Iberian Peninsula (Crusafont, 1952; Rı´os et al., 2016a; Rı´os et al., 2017), with some possible but very scarce remains assigned to Decennatherium crusafonti Bosscha Erdbrink, 1976, from Alia¨ba¨d (Iran) (Bosscha Erdbrink, 1976). The material studied here was collected from the Siwaliks Hills, and is currently housed at the Peabody Museum (Harvard University). It has a very good preservation and appears to show a high morphological similarity with the remains of the early Vallesian Decennatherium pachecoi Crusafont, 1952, and the late Vallesian Decennatherium rex Rı´os, Sa´nchez et Morales, 2017, both

§

Corresponding editor: Pierre-Olivier Antoine. * Corresponding author. E-mail address: [email protected] (M. Rı´os).

known from the Iberian Peninsula. It also shares the same similarities with the genus Palaeogiraffa from the late Vallesian of Greece and Turkey (Geraads, 1979; 1989; de Bonis and Bouvrain, 2003; Geraads et al., 2005; Kostopoulos and Sen, 2016). In this work we intend to:  describe the new material from the Siwaliks;  compare it to the previously described Decennatherium taxa and assess the similarities and differences between them, and if necessary their paleobiogeographical implications, and;  compare it with the genus Palaeogiraffa, and try to assess the validity of the genus.

2. Paleontological, geographical, geological and chronological settings The genus Decennatherium was named by Crusafont (1952), based on the giraffid material found in several early Vallesian sites of the Iberian Peninsula (Nombrevilla, Los Valles de ˜ a, Relea, Pedrajas de San Esteban, Saldan ˜ a). These Fuentiduen remains represented the first giraffid taxon found in the Iberian Peninsula: Decennatherium pachecoi. Six decades later, and based on the findings from Cerro de los Batallones (Madrid, Spain), a new Decennatherium species was established: Decennatherium

https://doi.org/10.1016/j.geobios.2019.10.007 C 2019 Published by Elsevier Masson SAS. 0016-6995/

Please cite this article in press as: Rı´os, M., et al., First identification of Decennatherium Crusafont, 1952 (Mammalia, Ruminantia, Pecora) in the Siwaliks of Pakistan. Geobios (2019), https://doi.org/10.1016/j.geobios.2019.10.007

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Fig. 1. Map showing the location of the sites studied in the Potwar Plateau, Pakistan.

rex (Rı´os et al., 2017). In the most comprehensive parsimony analysis available so far (Rı´os et al., 2017), this Iberian genus appears as the basal offshoot of a large sivathere-clade (also including samotheres). This group is diagnosed mainly by cranial features, and is defined as the least inclusive clade of crowngiraffids that contains Decennatherium and Sivatherium. The results of Rı´os et al. (2017) showed Decennatherium as the basalmost branch of a samotheriine-sivatheriine clade, and extended the chronostratigraphic and paleobiogeographic ranges of sivatheres/samotheres back into the early late Miocene and to the East to the Iberian Peninsula, hence rendering the samothere-sivathere clade as being one of the most successful and long-lived giraffid clades. Also, Decennatherium is the earliest known evidence of the Sivatherium ossicone-plan, a pattern of four ossicones that consists of two small, anteriorlyoriented, frontal ossicones and two much larger, caudallyoriented and curved, fronto-parietal ossicones that are covered by numerous longitudinal ridges. With Decennatherium this plan can be traced back to the late Miocene of Western Europe, disappearing with the last gigantic sivatheres by the late Pleistocene in Africa (Rios et al., 2017). The occurrence and age of the fossils is from 13.5 to 8.193 Ma, mostly late Miocene in age (dates provided by the Peabody Museum, Harvard University; Barry et al., 2002). The specimens recovered come from a series of paleontological sites located on the Siwaliks Hills of northern Pakistan (Fig. 1). The localities are: Y503, Y0311, Y0258, Y0269, Y0176, Y0215 and Y0980, and are of a late Miocene age, dating from 13.5 to 8.193 Ma (Scott, 2004).

Y503 is dated 13.5 Ma, which corresponds to the end of the early Serravallian in the MN European zones (MN6). Y0311 and Y0258 are dated 10.063 Ma and 10.159 Ma, respectively, close to the end of the equivalent early Vallesian (MN9), while Y0269 falls within the MN10 with an age of 9.415 Ma. Finally, Y0176, Y0215 and Y0980 are dated 8.193 Ma, 8.481 Ma and 8.891 Ma, respectively, which corresponds to MN11 (Johnson et al., 1982; Gradstein et al., 2012). The depositional environments of the Siwaliks layers are primarily derived from small and larger riverine environments. These Siwalik sediments are found along the southern margin of the Himalayas (Patnaik, 2013), and are exposed best in northern Pakistan at the Powar Plateau where most of the collections under study originate. Significant exposures of fluvial and fluviodeltaic deposits are also known in southwestern Punjab, Baluchistan, and Sind (Barry et al., 2002; Patnaik, 2013). According to Scott (2004) there is evidence of diverse but primarily forested habitats at Y0311 (Nagri Fm.).

3. Material and methods 3.1. Material The studied material includes:  Ossicone fragments YGSP 42429 and YGSP 27682 (locality Y503, 13.5 Ma);

Please cite this article in press as: Rı´os, M., et al., First identification of Decennatherium Crusafont, 1952 (Mammalia, Ruminantia, Pecora) in the Siwaliks of Pakistan. Geobios (2019), https://doi.org/10.1016/j.geobios.2019.10.007

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 m2 YGSP 6399 (10.063 Ma);  mandible fragment YGSP 12210 (locality Y0311, 10.063 Ma);  mandible fragment with dentition YGSP 13201 (locality Y0311, 10.063 Ma);  metacarpal YGSP 11455 (locality Y0311, 10.063 Ma);  metacarpal YGSP 14580 (locality Y0269, 9.415 Ma);  distal tibia YGSP 4124 (locality Y0176, 8.193 Ma);  distal tibia YGSP 52743 (locality Y0980, 8.891 Ma);  astragalus YGSP 9472 (locality Y0258, 10.159 Ma);  metatarsal YGSP 15184 (locality Y0311, 10.063 Ma);  proximal phalanx YGSP 5807 (locality Y0215, 8.481 Ma). The material comes from the Harvard-Geological Survey of Pakistan Collection, an historical collection curated at the Peabody Museum (Harvard University). 3.2. Comparative material The comparative material comprises the entire Decennatherium ˜ a and the pachecoi collection from Los Valles de Fuentiduen complete sample of Decennatherium rex from Batallones-10, both stored at the MNCN-CSIC (Madrid, Spain), listed in Appendix A. It also includes the published data on the Vallesian Palaeogiraffa major from Ravin de la Pluie (Northern Greece; Geraads, 1979; de Bonis and Bouvrain, 2003), as well as Palaeogiraffa macedoniae from Pentalophos (Northern Greece; Geraads, 1989; de Bonis and Bouvrain, 2003) and Yulafli (Turkey; Geraads et al., 2005), and Palaeogiraffa pamiri from Xirochori (Northern Greece;de Bonis and Bouvrain, 2003), Yassio¨ren (Ozansoy, 1965; Geraads et al., 2005), and Ku¨c¸u¨kc¸ekmece West (Turkey; Kostopoulos and Sen, 2016). 3.3. Methods We mostly follow the set of measurements proposed by Ba¨rmann and Ro¨ssner (2011) for the dentition, Quiralte (2011) for the tibia, Rı´os et al. (2016b) for the metapodials, and Solounias and Danowitz (2016) for the astragalus. All were taken with digital calipers. Abbreviations: YGSP, Geological Survey of Pakistan (Peabody Museum, Harvard University); MNCN, Museo Nacional de Ciencias Naturales-CSIC; BAT, Cerro de los Batallones; AM, American Museum of Natural History.

4. Systematic paleontology Class Mammalia Linnaeus, 1758 Order Cetartiodactyla Montgelard, Catzeflis et Douzery, 1997 Suborder Ruminantia Scopoli, 1777 Family Giraffidae Gray, 1821 Genus Decennatherium Crusafont, 1952 Decennatherium asiaticum nov. sp. Fig. 2 Derivation of the name: Refer to Asia, where this new species has been found. Holotype: Metatarsal YGSP 15184 (locality Y0311, 10.063 Ma). Paratype: Metacarpal YGSP 11455 (locality Y0311, 10.063 Ma). Material: All material listed above (Section 3.1) with the exception of the ossicone fragments. Type locality and horizon: Y0311 (equivalent to the end of the early Vallesian (MN9), dating 10.063 Ma). Occurrence: From 10.1 to 8.1 Ma. Measurements: See Tables 1 and 2. Differential diagnosis: Large giraffid. Lower molars longer than wider where the lingual wall shows a developed metastylid. The metacarpal is longer and with a slightly slenderer diaphysis

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index than in other Decennatherium species with a value of 9.18. The lateral and medial epicondyles of the metacarpal are slightly asymmetrical with a more triangular medial epicondyle. The metacarpal palmar shaft shows rounded medial and lateral ridges. Distally, the tibia has a large, flat and circular ventral fibular facet and an elongated and thin medial malleolus. The distal APD/ distal TD tibia index is lower than in the other Decennatherium taxa with a value of 0.74. The astragalus intratrochlear notch is broad and shows a distinct medial scala and distal intracephalic fossa. The metatarsal shows no distinct pygmaios and has a slightly slenderer diaphysis index than other Decennatherium species with a value of 8.44. The proximal phalanx shows a distinct notch in lateral view that separates the lateral palmar eminence from the articular surface of the base. Description: Lower molar (YGSP 6399, YGSP 13201, and YGSP 12210): The lower molars (Fig. 2(A–D): Table 1) have bucco-lingual compression, which is diagnostic of this species. They have a lingual wall with a moderate metastylid. The metaconid and entoconid are aligned and similar in size. The buccal wall has a well-developed, V-shaped protoconid and hypoconid. The protoconids are smaller than the hypoconids. The medial and anterior notches are thin and ‘‘Crescent-moon’’-shaped. There is a buccal basal pillar between the anterior and posterior lobes with a variable degree of development. The lingual surfaces of these teeth are exceptionally flat. Metacarpal (YGSP 14580, YGSP 11455): The lateral and medial epicondyles are slightly asymmetrical (Fig. 2(E, F); Table 2). The lateral epicondyle is triangular. The proximal articular surface slightly extends onto the palmar surface of the lateral epicondyle. The medial epicondyle is larger than the lateral epicondyle, and is triangular. There are two small, rounded protrusions on the palmar surface of the medial epicondyle. The lateral and medial epicondyles are separated by a notably narrow groove that connects distally with the central trough. Both the medial and lateral ridges are rounded. There is an elongated oval fossa on the outer edge of the lateral ridge. The central trough is intermediate in depth, and flattens towards the distal shaft. There is slight outward flaring of the distal shaft that is more pronounced medially. The keels of the distal condyles slightly protrude onto the palmar shaft. Tibia (YGSP 4124, YGSP 52743): Distally, the ventral fibular facet is large, circular, and flat (Fig. 2(G); Table 2). The dorsal fibular facet is massive and flattened. The ventral and dorsal fibular facets are separated by a deep and narrow fibular incisure. The lateral tibial cochlea is large, and is separated from the medial tibial cochlea by a narrow wall. There is a small tibial tubercle. The medial malleolus is elongated and thin. Astragalus (YGSP 9472): Dorsally, the lateral and medial trochleae are separated centrally by a wide and flattened groove (Fig. 2(H, I); Table 2). The lateral edge of the trochlea is flat without any arching. There is an intermediately-sized and flat central fossa. The trochlea and the head of the astragalus are on the same plane in dorsal view. The collum tali is tall. At the medial collum tali, the astragalus bulges outward with a faint ridge on the surface that extends towards the central fossa. At the proximal edge of the articular surface, there is a sigmoid medial edge, flat central surface, and vertical lateral edge. The lateral aspect of the head is notably larger than the medial aspect. There is a notch at the lateral astragalar surface between the trochlea and the head. Ventrally, there is a small notch at the lateral aspect of the proximal trochlea. The intratrochlear notch is broad and the proximal triangular fossa is pronounced and deep. The interarticular groove is exceptionally wide. The medial edge of the ventral articular surfaced is directed obliquely towards the intratrochlear notch. There is a distinct medial scala and distal intracephalic fossa. The astragalus is tall and narrow.

Please cite this article in press as: Rı´os, M., et al., First identification of Decennatherium Crusafont, 1952 (Mammalia, Ruminantia, Pecora) in the Siwaliks of Pakistan. Geobios (2019), https://doi.org/10.1016/j.geobios.2019.10.007

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Fig. 2. Skeletal remains of the giraffid Decennatherium asiaticum nov. sp., from the Miocene of Pakistan. A, B. Mandible fragment with dentition YGSP 13201/Left in occlusal (A) and lingual (B) views. C, D. Lower m2 YGSP 6399/Right in occlusal (C) and lingual (D) views. E, F. Metacarpal YGSP 14580/Left in proximal (E) and plantar (F) views. G. Tibia YGSP 4124/Right in distal view. H, I. Astragalus YGSP 9472/Left in dorsal (H) and plantar (I) views. J–L. Metatarsal YGSP 15184/Left in proximal (J), dorsal (K) and plantar (L) views. M–O. Proximal phalanx YGSP 5807/Left digit in dorsal (M), medial (N) and proximal (O) views. Scale bars: 2 cm (A–D), 4 cm (E, J), 5 cm (F–I, K–O).

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GEOBIO-891; No. of Pages 14 M. Rı´os et al. / Geobios xxx (2019) xxx–xxx Table 1 Measurements (in mm) of the teeth of the Miocene giraffid Decennatherium asiaticum nov. sp. from the Siwaliks of Pakistan. Specimen No.

Element

Length

Width

W/L

YGSP 13201 YGSP 6399

m2 m2

35.02 39.21

23.89 26.14

1.46 1.50

Metatarsal (YGSP 15184): The proximal articular surface shows an asymmetry between the lateral and the medial profile. The kidney-shaped articulation facet for the ectomesocuneiform is similar in size to the navicular-cuboid facet, which is semicircular. The entocuneiform facet is relatively big and oval and it stands out over the plantar profile. The medial epicondyle is separated into a distinct dorsal and plantar head by a wide, obliquely oriented groove (Fig. 2(J–L); Table 2). The lateral epicondyle is separated into dorsal and plantar heads by a narrow, deep groove. The dorsal heads flare outward and the plantar heads are longitudinally oriented and rounded. The inner surface of the medial epicondyle contains a small, obliquely oriented ridge, which laterally displaces the central foramen. There is no distinct pygmaios. There is a rounded medial ridge and sharp lateral ridge that create an intermediate-to-shallow central trough. The central trough abruptly flattens at the distal shaft. The pyramidal rise is absent. There is pronounced outward flaring at the lateral distal shaft. The keels of the distal condyles are pronounced and sharp. Proximal phalanx (YGSP 5807): There is a massive lateral palmar eminence (Fig. 2(M–O); Table 2). There is a distinct notch in lateral view that separates the lateral palmar eminence from the articular surface of the base. The head is notably rounded and is dorso-ventrally wider than the distal shaft. The bone is robust and elongated. ?cf. Decennatherium Fig. 3(A) Material: Ossicone fragments YGSP 42429 and YGSP 27682. Locality and horizon: Locality Y503 (end of the early Astaracian (MN6), dating 13.5 Ma). Description: The ossicone fragments YGSP 42429 and YGSP 27682 can be connected to form an incomplete ossicone where the proximal part is missing (Fig. 3(A)). The ossicone is curved and is ornamented by a high number of deep ridges on its surface, which run irregularly and longitudinally to the apex of the ossicone. The apex is blunt and the section is oval. Though it is smaller in size than the large posterior ossicones of D. rex males (Fig. 3(B)), they share the same morphology regarding both the curvature, cross section, and surface details.

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5. Discussion 5.1. Anatomical comparisons The ossicone is morphologically similar to those of D. rex, despite having smaller dimensions. Both species show deep longitudinal irregular ridges and possess a blunted apex and a curvature on the distal part of the ossicone. The cross section is also very similar and different from other giraffids known from the area as Bramatherium or Sivatherium. Regarding the possible similarity between YGSP 42429 and YGSP 27682 and Giraffokeryx (which is present in the Siwaliks), although they have a similar size, the morphology differs as the ridges on the ossicones of Giraffokeryx are fainter and there is barely no curvature at the apex (Fig. 3(C)). Also the shaft near the apex is rounder in the Siwaliks specimen while in Giraffokeryx it is more compressed. However, the ossicone found in the Siwaliks is closer in age to those of Giraffokeryx than to Decennatherium, making the assignement of the material found to this genus still uncertain. As for the material ascribed to Decennatherium asiaticum nov. sp., comparisons with other Decennatherium species discussed below are summarized in Appendix B. The lower molars show many similarities with the Iberian Decennatherium taxa, as they share the same size and proportions as well as numerous morphological features. They all have welldeveloped and V-shaped protoconid and hypoconid, thin and ‘‘crescent-moon’’-shaped medial and anterior notches, a buccal basal pillar between the anterior and posterior lobes, a buccal cingulum, and rugose enamel. These particular features are also seen in Palaeogiraffa pamiri from Ku¨c¸u¨kc¸ekmece West, Turkey (Ozansoy, 1965; Kostopoulos and Sen, 2016) and Palaeogiraffa major from Ravin de la Pluie, Greece (Geraads, 1979; deBonis and Bouvrain, 2003). Decennatherium asiaticum nov. sp., however, has a lingual wall with a well-developed metastylid, strongest in the specimen from the Siwaliks, intermediate in D. rex, and least developed in D. pachecoi and P. pamiri (Kostopoulos and Sen, 2016; Fig. 4). The Siwalik giraffid also differs from other forms in having exceptionally flat lingual surfaces. Regarding the lower m2 Length/ width index, the specimens from the Siwaliks fall within the range of D. rex and P. pamiri; they have smaller values than D. pachecoi and P. major, which have the highest index values as well as the largest overall size (Table 3; Fig. 5(A)). The medial epicondyle of the metacarpal is triangular in the Siwaliks form, whereas in Decennatherium pachecoi and D. rex it is semicircular (Fig. 6). The Siwalik species also differs in showing a medial epicondyle that barely protrudes contrary to the developed and protruding medial condyles of D. pachecoi and P. pamiri (Kostopoulos and Sen, 2016) and the large and very protruding medial condyles of D. rex. Also, the palmar medial and lateral ridges are rounded in the Siwaliks specimen, whereas in D. pachecoi and

Table 2 Measurements (in mm) of the postcranial of the Miocene giraffid Decennatherium asiaticum nov. sp. from the Siwaliks of Pakistan. Specimen No. YGSP YGSP YGSP YGSP YGSP

14580 4124 15184 11455 5807

Element

Total Length

Proximal TD

Mc III–IV Tibia Mt III–IV Mt III–IV Proximal phalanx

469.64

80.27

456.72

65.45

102

49

Specimen No.

Element

YGSP 9472

Astragalus

Medial Length

Lateral Length 94.44

Proximal APD

Diaph. TD

Diaph. APD

43.00 64.44

78.28 102.3 68.77 64.51

40.01

Medial APD

Distal TD

Lateral APD

Distal DT

Distal APD 76.16

Proximal DT

61.15

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Fig. 3. Ossicones of selected fossil giraffids. A. ?cf. Decennatherium, YGSP 42429 and YGSP 27682. B. Decennatherium rex, posterior ossicone BAT-10’07-G4-127. C. Giraffokeryx punjabiensis, posterior ossicone AM 19475. Scale bar: 5 cm.

D. rex the medial ridge is sharper and the lateral ridge is rounder. They also differ in the diaphysis index with the Siwaliks specimen showing a slightly lower value than D. pachecoi (Diaphysis index of 9.15 vs. 9.25). D. rex shows slightly more robust metacarpals than the other two forms (Diaphysis index = 10.58–12.76, with an

average of 11.52, n = 9), similar to P. pamiri (11.25) (Fig. 5(B)). The Siwaliks specimen also differs in showing longer metacarpals than the other two taxa, with a length of 469 mm (n = 1) when D. pachecoi only complete specimen does not exceed 391 mm and D. rex ranges from 387 to 425 mm, with an average of 412.55

Fig. 4. Lower m2 of selected fossil giraffids in occlusal (top) and lingual (bottom) views. A. Decennatherium asiaticum nov. sp., YGSP 13201/Left. B. Decennatherium pachecoi, MNCN-43481/Left. C. Decennatherium rex, BAT-10’10-F2-18/Left. Scale bar: 5 cm.

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GEOBIO-891; No. of Pages 14 M. Rı´os et al. / Geobios xxx (2019) xxx–xxx Table 3 Descriptive parameters of the giraffids Decennatherium pachecoi, D. rex and D. asiaticum nov. sp. Abbreviations: Max., maximum; Min., minimum; ntotal, total sample size. Measurements taken from Crusafont (1952), Bosscha Erdbrink (1976), Geraads (1979, 1989), de Bonis and Bouvrain (2003), Geraads et al. (2005), Kostopoulos and Sen (2016), and Rı´os et al. (2017). m2 L/W

D. pachecoi D. rex D. asiaticum P. major P. pamiri

Max.

Min.

Average

ntotal

1.477 1.794 1.470 1.588 1.558

1.424 1.386 1.470 1.479 1.517

1.450 1.564 1.470 1.534 1.533

2 23 1 2 3

Mc III–IV total length

D. pachecoi D. rex D. asiaticum P. macedoniae P. pamiri

Max.

Min.

Average

ntotal

391 425 469 465 ? 400

391 387 469 465 ? 400

391 412.6 469 465 ? 400

1 9 1 1 1

Mc III–IV Diaph TD/Total length  100

D. pachecoi D. rex D. asiaticum P. macedoniae P. pamiri

Max.

Min.

Average

ntotal

9.251 12.767 9.186 17.634 21.650

9.251 10.580 9.186 17.634 21.650

9.251 11.525 9.186 17.634 21.650

1 9 1 1 1

Tibia distal index

D. D. D. D.

pachecoi rex asiaticum crusafonti

Max.

Min.

Average

ntotal

0.816 0.891 0.744 0.800

0.768 0.743 0.744 0.800

0.790 0.792 0.744 0.800

4 5 1 1

Astragalus Distal TD/Lateral length  100

D. pachecoi D. rex D. asiaticum D. crusafonti P. pamiri

Max.

Min.

Average

ntotal

0.729 0.715 0.648 0.648 0.669

0.573 0.652 0.648 0.648 0.605

0.631 0.681 0.648 0.648 0.637

23 8 1 1 3

Mt III–IV total length

D. pachecoi D. rex D. asiaticum P. macedoniae P. pamiri

Max.

Min.

Average

ntotal

427.5 465 450 480 ? 435

406.1 428 450 480 ? 435

416.8 446.1 450 480 ? 435

2 8 1 1 1

Mt III-IV Diaph TD/Total length  100

D. pachecoi D. rex D. asiaticum P. pamiri

Max.

Min.

Average

ntotal

10.455 10.336 8.444 9.011

8.269 8.700 8.444 9.011

9.362 9.411 8.444 9.011

2 8 1 1

Proximal phalanx total length

D. pachecoi D. rex D. asiaticum P. pamiri

Max.

Min.

Average

ntotal

99.1 110.6 102 98.7

80.5 86.5 102 88

89.7 96.9 102 93.25

23 40 1 4

7 Proximal phalanx Prox. TD/Total length  100

D. pachecoi D. rex D. asiaticum P. pamiri

Max.

Min.

Average

ntotal

48.618 49.237 48.039 46.477

36.649 35.859 48.039 44.640

42.826 43.208 48.039 45.550

23 40 1 4

(n = 9) (Table 3). As for the length of Palaeogiraffa, P. pamiri metacarpals from Ku¨c¸u¨kc¸ekmece West are similar in size to those of D. pachecoi and D. rex metacarpals, while a specimen of P. macedoniae from Pentalophos reaches 465 mm in length, very similar to the Siwaliks specimen. However, P. macedoniae has a much higher proximal index, closer to that of D. pachecoi, while the Siwaliks specimen has a proximal index that falls within the range of D. rex and P. pamiri. The tibia of the Siwalik specimen is the youngest tibia with a Decennatherium morphology found in the fossil record so far. It shows distally a large, circular, and flat ventral fibular facet as found in D. pachecoi whereas D. rex shows a ventral fibular facet which is also large and circular, but much more prominent than in other taxa (Fig. 7). They also differ in the development of the medial malleolus, which is elongated and thin in the Siwalik specimen, strong and narrowing distally in D. rex and in D. pachecoi, slightly weaker than in the Siwaliks form and much weaker than in D. rex. Finally, the distal tibia index is 0.74 (n = 1), slightly lower than D. pachecoi (0.76–0.81; average 0.79, n = 5), D. rex (0.74–0.89, average 0.78, n = 5), and D. crusafonti (0.8, n = 1), though it is larger in absolute dimensions (Table 3; Fig. 5(C)). The astragalus from the Siwaliks exhibits an intratrochlear notch broader than other Decennatherium species (Fig. 8). It also differs in the distinct medial scala and distal intracephalic fossa, which is faint in other taxa. The lateral length of the astragalus is shortest in D. pachecoi, and longest in D. rex. Decennatherium asiaticum nov. sp. has a medial length closest to the longest D. rex specimens, very similar to that of D. crusafonti (Bosscha-Erbrink, 1976), while Palaeogiraffa pamiri shows slightly lower values (Kostopoulos and Sen, 2016). All the taxa mentioned above show a similar Distal TD/Lateral length index close to 0.6 (Rı´os, 2018; Table 3; Fig. 5(D)). The metatarsal shows no distinct pygmaios in the Siwaliks specimen, whereas the pygmaios in Decennatherium pachecoi is present as a distinct, oval protrusion with a rounded surface, which is oriented medially and protrudes proximally. In D. rex it is barely developed (Fig. 9). They also differ in the diaphysis index with the Siwaliks specimen (8.75, n = 1) showing a slightly lower value than the average of D. pachecoi. (8.26–10. 45, average 9.36, n = 2) and P. pamiri (9.01, n = 1) (Kostopoulos and Sen, 2016). Decennatherium rex shows slightly more robust metatarsals than the other forms (Diaphysis Index = 8.69–10. 36, with an average of 9.41, n = 8; Table 3; Fig. 5(E)). The Siwalik specimen also differs in possessing metatarsals, which are similar in length to those of D. rex and P. pamiri (Kostopoulos and Sen, 2016), but are longer than those of D. pachecoi and shorter than those of P. macedoniae (Geraads, 1989). They all share an intermediate-to-shallow central trough except D. crusafonti, which has a slightly more marked plantar metatarsal trough than the rest (Bosscha Erdbrink, 1976). The proximal phalanx of the Siwalik specimen shows a distinct notch in lateral view that separates the lateral palmar eminence from the articular surface of the base. This notch is more marked than in other Decennatherium species (Fig. 10). The proximal phalanx of D. pachecoi differs from D. rex in showing less developed rugose proximopalmar/plantar areas for ligament insertion, which

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Fig. 5. Bivariate plots showing different biometric relationships for Decennatherium and Palaeogiraffa species. A. Lower m2. B. Metacarpal III–IV. C. Tibia. D. Astragalus. E. Metatarsal III–IV. F. Proximal phalanx. Abbreviations: TD, transversal diameter; APD, anteroposterior diameter; diaph., diaphysis. Measurements taken from Crusafont (1952), Bosscha Erdbrink (1976), Geraads (1979), de Bonis and Bouvrain (2003), Geraads et al. (2005), Kostopoulos and Sen (2016), and Rı´os et al. (2017).

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Fig. 6. Metacarpals of selected fossil giraffids in plantar view. A. Decennatherium asiaticum nov. sp., YGSP 14580/Left. B. Decennatherium pachecoi, MNCN-42769/Left. C. Decennatherium rex, BAT-10’07-G2-28/Left. Scale bar: 5 cm.

are remarkably prominent in D. rex and in the Siwalik specimen. Regarding the length, D. rex is longer on average, followed but the Siwaliks specimen, P. pamiri and D. pachecoi (Table 3). As for the proximal robustness index, D. rex shows the highest value (average = 49.23, n = 40), followed by D. pachecoi and the Siwaliks specimen (Table 3; Fig. 5(F)). P. pamiri shows much lower values (average = 45.55, n = 4). Decennatherium, the Siwaliks specimens and Palaeogiraffa represent medium-large sized giraffids from the Vallesian sharing numerous morphological and biometrical features (Fig. 5), which also are pretty distinct from those of other giraffids from the same period and areas. But there are also some differences between them, as we have listed in the comparisons above. In addition, these comparisons do not include other skeletal remains that have been mentioned in the past as key to separate both genera.

Kostopoulos and Sen (2016) separated P. pamiri from Ku¨c¸u¨kc¸ekmece West from Decennatherium on the basis of dental dimensions (Spanish taxa teeth are ca. 20% larger than the Ku¨c¸u¨kc¸ekmece ones), as well as the p4 morphology (presence of a continuous eocristid on the p4), and limb proportions (metacarpals appear to be wider that those of D. pachecoi). However, their study could not include D. rex and our analysis reveals that both the dental dimensions and the limb proportions of P. pamiri are actually quite similar to those of D. rex (Fig. 5). As for the p4 morphology, Rı´os et al. (2016a) showed the high variability of the p4 posterior lobe in D. pachecoi so we have to take the identifications made regarding the differences of this tooth cautiously. Rı´os et al. (2016a) included Palaeogiraffa in their phylogenetic analysis and they analyzed other anatomical elements, linking

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(B. schaubi). However, in both P. macedoniae and P. major the p3 presents a strong mesolingual conid (Montoya and Morales, 1991: fig. 5a-c; Rı´os et al., 2016a: fig. 5L-N) that contrasts with D. pachecoi, which has an isolated mesolingual conid (only in two of its morphotypes). Also, the decidual canines of P. major have a larger second lobe than the ones from D. pachecoi (de Bonis and Bouvrain, 2003). Facing the lack of Palaeogiraffa ossicones and the scantiness of this material, the differences listed above prevent us from synonymizing Palaeogiraffa with Decennatherium for the moment. Our results show that there is a strong argument for placing the Pakistan specimens within the Decennatherium genus and the points indicated in this discussion suffice, in our opinion, to assign the Siwalik material to a separate, new species. As for the comparisons with the other most abundant large giraffids of the Siwalik region, namely Bramatherium and Sivatherium, these have much larger values regarding the length of all the elements analyzed, as well as much higher robusticity indexes indicating the specimens analyzed in this paper are slenderer and not as massive. They also differ in many morphological features as the characteristic cranial, metapodial and tarsal features. 5.2. Phylogenetics

Fig. 7. Tibia of selected fossil giraffids in distal view. A. Decennatherium asiaticum nov. sp., YGSP 4124/Right. B. Decennatherium pachecoi, MNCN-30785/Right. C. Decennatherium rex, BAT-10’14-E2-26/Right. Scale bar: 5 cm.

most of the alleged Decennatherium findings and Palaeogiraffa to the samothere clade with a very low support (Rı´os et al., 2016a: fig. 14, Node M). Their most parsimonious tree linked P. macedoniae to D. pachecoi as sister taxa, while it placed P. major next to Birgerbohlinia schaubi, due to the striking similarities between the p3 from Ravin de la Pluie (P. major) and the ones from Crevillente

The anatomical and biometrical evidence, together with the biochronology of the sites suggests that D. pachecoi from the early Vallesian of the Iberian Peninsula (MN9) is ancestral to D. rex from the late Vallesian (MN10) of the Iberian Peninsula, while D. asiaticum nov. sp. from the Middle Siwaliks (MN9-11) may be more derived or not. ?cf. Decennatherium may represent the first Decennatherium appearance in the fossil record and represent a common ancestor of the Decennatherium clade. Further findings in the Pakistan area are necessary to shed light on the relationships within the Decennatherium taxa. Regarding the dentition, the development of the lingual wall and the metastylid of the lower molars appear to point that way, as the development in D. rex is intermediate between D. pachecoi and the specimen from the Siwaliks, suggesting D. pachecoi exhibits the plesiomorphic state of the character. The lingual wall would have become stronger, developing the metastylid to increase the resistance of the tooth. The metapodial length also suggests a similar scenario, with D. pachecoi having the shortest metapodials and the specimen from Siwaliks the longest ones, indicating a trend towards longer metapodials. The same occurs with the metatarsal, with the development of the pygmaios, as it is distinct and prominent in D. pachecoi, less developed in D. rex and not present in the Siwalik form, indicating a trend towards a lesser development of the plantar interosseous muscles. However, there are some anatomical and biometrical features that may suggest a different scenario where both D. rex and D. asiaticum nov. sp. are derived from a common ancestor close to D. pachecoi in different ways. For example, the robustness index of the metapodials is very similar in D. pachecoi and in the specimen from the Siwaliks, while the metapodials of D. rex are more robust. Regarding the development of the metacarpal medial epicondyle, it is more developed in D. rex than in D. pachecoi, and barely protruding in the Siwaliks taxon, indicating less developed carpal flexor radial and extensor oblique muscles than in the other two forms. The same occurs with other features as the tibia distal index and the metatarsal robustness index, which indicate a less robust distal tibia and metatarsals for the Siwalik specimen than for the Iberian ones. So far, available evidence appears to indicate that both D. rex and D. asiaticum nov. sp. are more derived than D. pachecoi, with D. rex remaining plesiomorphic in the dentition and several

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Fig. 8. Astragalus of selected fossil giraffids in dorsal (left) and plantar (right) views. A. Decennatherium asiaticum nov. sp., YGSP 9472/Left. B. Decennatherium pachecoi, MNCN42554/Left. C. Decennatherium rex, BAT-10’09 G2-59/Left. Scale bar: 5 cm.

postcranial features but evolving towards a more robust shape, and D. asiaticum nov. sp. retaining the ancestral postcranial slenderness of D. pachecoi but showing other derived features as longer limb bones and stronger molars. Regarding the phylogenetic affinities between Decennatherium species and the rest of the Giraffidae, in previous phylogenetic analysis (Rı´os et al., 2016a, 2017) Decennatherium has been nested within the crown-giraffids, in a sister clade to all the large late

Miocene giraffids called the Samotheriinae-Sivatheriinae group. The ossicones of D. rex linked this genus to the base of this clade, having the same type of ossicone bauplan than the larger and more modern sivatheres. In addition, apart from the rare remains of D. crusafonti in Iran, Decennatherium-like giraffids have been found in the Eastern Mediterranean, as there are several findings of Palaeogiraffa in Greece and Turkey (Ozansoy, 1965; Geraads, 1979, 1989; de Bonis and Bouvrain, 2003; Geraads et al., 2005;

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Fig. 9. Metatarsal of selected fossil giraffids in dorsal (left picture for each specimen) and plantar (right picture for each specimen) views. A. Decennatherium asiaticum nov. sp., YGSP 15184/Left. B. Decennatherium pachecoi, MNCN-42764/Left. C. Decennatherium rex, BAT-10’07-G2-22/Left. Scale bar: 5 cm.

Fig. 10. Proximal phalanx of selected fossil giraffids in medial (left) and proximal (right) views. A. Decennatherium asiaticum nov. sp., YGSP 5807/Left digit. B. Decennatherium pachecoi, MNCN-42667/Left digit. C. Decennatherium rex, BAT-10’08-G2-39/Left digit. Scale bar: 5 cm.

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˜ a; BAT, Fig. 11. Map showing the geographical distribution of Decennatherium and Palaeogiraffa findings. Abbreviations: NOM, Nombrevilla; LVF, Los Valles de Fuentiduen ¨ C¸, Ku¨c¸u¨kc¸ekmece West). Data from Batallones-10; ALI, Alia¨ba¨d; SIW, Siwaliks; RDLP, Ravin de la Pluie; PNT, Pentalophos; YUL, Yulafli; XIR, Xirochori; YAS, Yassio¨ren; KU Crusafont (1952), Ozansoy (1965), Bosscha Erdbrink (1976), Geraads (1979, 1989), de Bonis and Bouvrain (2003), Geraads et al. (2005), Kostopoulos and Sen (2016), and Rı´os et al. (2017). Map adapted from Popov et al. (2006).

Kostopoulos and Sen, 2016). These new Decennatherium remains in the Siwaliks during the late Miocene, whereas larger sivatherines appear for the first time in the fossil record later towards the end of the late Miocene, opens a new line of research on the origin of this clade. The fragmented ossicone YGSP 42429 and YGSP 27682, which is dated as old as 13.5 Ma (MN6), shows mosaic features between Giraffokeryx and Decennatherium, but is much older than the first occurrence of D. pachecoi in the Iberian Peninsula, so more material from the same site would be necessary to make an informed statement on the nature of this material. 5.3. Paleobiogeographical distribution of Decennatherium The earliest known occurrence of Decennatherium in the fossil record is D. pachecoi, in the Iberian Peninsula, in the early ˜a Vallesian sites of Nombrevilla and Los Valles de Fuentiduen

(MN9, 11.1–9.7 Ma; Crusafont, 1952). However, the new occurrence of Decennatherium in the Siwaliks (MN9-11, 10.1– 8.1 Ma) is very close in age (Vallesian to early Turolian, and the ossicone is Astaracian). Decennatherium rex is known so far only from the Cerro de los Batallones, also in the Iberian Peninsula, dated late Vallesian (MN10, ca. 9 Ma; Morales et al., 2008). As for D. crusafonti from the Iranian site of Alia¨ba¨d (Bosscha Erdbrink, 1976), the site was dated tentatively to the early Vallesian (MN9, 11.1–9.7 Ma) but later Ataabadi et al. (2016) palaeomagnetically dated the Iranian area as mammal unit MN11 or possibly early MN12 (7.9–6.8 Ma), resulting to be younger than the Iberian findings and possibly implying a high mobility of this fossil giraffid genus, as we see in modern giraffes (Dagg, 2014) as well as late Miocene putative dispersals to the Middle East and Southern Asia. The finding of Decennatherium sp. in the Siwaliks (MN9-11, 10.1–8.1 Ma) has deep paleobiogeographical implications as it

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represents the easternmost occurrence of the genus (Fig. 11). This would suggest that this genus originally described from Iberia was widespread throughout all Eurasia during the Miocene, reaching the areas where we find their larger relatives Samotherium, Bramatherium and the later Sivatherium. 6. Conclusions We have described and compared anatomically and biometrically the previously unknown remains of a new large giraffid from the Siwaliks Hills of Pakistan. This taxon is attributed to the late Miocene giraffid genus Decennatherium, previously only surely identified in the early and late Vallesian of the Iberian Peninsula, with only some possible remains assigned to Decennatherium crusafonti from Alia¨ba¨d (Iran). Decennatherium asiaticum nov. sp. shows a high morphological similarity with the remains of the genus from Spain, and represents a more derived form than the ancestral D. pachecoi, with longer limbones and more developed metastylids on the lower molars. Decennatherium asiaticum nov. sp. also shares some similarities with Palaeogiraffa, but the differences support a separation of the two genera until further analysis. The findings analyzed in this study represent the easternmost occurrence of Decennatherium. They allow for naming a new species, D. asiaticum nov. sp., and show that this genus had a much wider distribution than expected, ranging the westernmost part of Eurasia through the Middle East, and reaching the Punjab area in Pakistan. Acknowledgements M.R. acknowledges an FPI predoctoral grant (Spanish Government MINECO) as well as the EEBB-FPI grant program 2013, 2014 and 2015. We thank P. Perez and S. Fraile (MNCN-CSIC, Madrid, Spain), J. Galkin, J. Meng and E. Westwig (AMNH; New York, USA), P. Brewer and S. Pappa (NHM, London, UK), C. Argot and the late S. Peigne´ (MNHN, Paris, France), J.C. Barry, D. Pilbeam, L. J. Flynn and M. E. Morgan (Harvard; Massachusetts; USA) and their respective home institutions for giving us access to their giraffid fossil collections. We thank the Anatomy Department of the NYIT-COM. We thank E. Cantero, B. Go´mez and P. Gutierrez for the preparation of the BAT10 fossils. We also thank A. Gentry, A. Xafis and an anonymous reviewer as well as the editors for their thoughtful comments and efforts towards improving our manuscript.

Appendices A, B. Supplementary information Supplementary information (including a list of comparative material [A] and a comparative table of morphological characters [B]) associated with this article can be found, in the online version, at https://doi.org/10.1016/j.geobios.2019.10.007. References Ataabadi, M.M., Kaakinen, A., Kunimatsu, Y., Nakaya, H., Orak, Z., Paknia, M., Sakai, T., Salminen, J., Sawada, Y., Sen, S., 2016. The late Miocene hominoid-bearing site in the Maragheh Formation Northwest Iran. Palaeobiodiversity and Palaeoenvironments 96, 349–371. Ba¨rmann, E.V., Ro¨ssner, G.E., 2011. Dental nomenclature in Ruminantia: Towards a standard terminological framework. Mammalian Biology 76, 762–768. Barry, J.C., Morgan, M.E., Flynn, L.J., Pilbeam, D., Behrensmeyer, A.K., Raza, S.M., Imran, A., Khan, I.C., Badgley, C., Hicks, J., Kelley, J., 2002. Faunal and environmental change in the late Miocene Siwaliks of northern Pakistan. Paleobiology 28, 1–71.

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Please cite this article in press as: Rı´os, M., et al., First identification of Decennatherium Crusafont, 1952 (Mammalia, Ruminantia, Pecora) in the Siwaliks of Pakistan. Geobios (2019), https://doi.org/10.1016/j.geobios.2019.10.007