Early Paleogene decapod crustaceans from the Sulaimanand Kirthar Ranges, Pakistan
Crustacés décapodes du Paléogène inférieur des montagnes de Sulaimanet du Kirthar, Pakistan
Sylvain Charbonnier a,∗, Alessandro Garassino b, Giovanni Pasini c,Grégoire Métais a,∗, Didier Merle a, Annachiara Bartolini a,Imdad A. Brohi d, Sarfraz H. Solangi d, Rafiq A. Lashari d,
Jean-Loup Welcomme e, Laurent Marivaux f
a Muséum national d’histoire naturelle, département histoire de la Terre, UMR 7207 CNRS, centre de recherche sur lapaléobiodiversité et les paléoenvironnements, case postale 38, 8, rue Buffon, 75005 Paris, France
b Department of Invertebrate Palaeontology, Museo di Storia Naturale di Milano, Corso Venezia 55, 20121 Milano, Italyc Museo Civico dei Fossili di Besano, Via Prestini 5, 21050 Besano, Varese, Italy
d Centre for Pure and Applied Geology, University of Sindh, Allama I.I. Kazi Campus, Jamshoro 76080, Pakistane 57, chemin de la Mort-Aux-Ânes, 34750 Villeneuve-lès-Maguelone, France
f Institut des sciences de l’évolution de Montpellier, UMR-CNRS 5554, Cc 064, université de Montpellier-2,place Eugène-Bataillon, 34095 Montpellier cedex 5, France
Received 8 October 2012; accepted 10 December 2012Available online 21 January 2013
Fossil decapod crustaceans from Pakistan have been the subject of relatively few studies.Stoliczka (1871) and Noetling (1897) reported new species from Pakistan and India and Glaessner(1933) described several new species based upon the collections of the British Museum. Later,Gingerich et al. (1979) reported decapod crustacean remains of some Paleocene and Eoceneformations in Pakistan but without formal description. Collins and Morris (1978) contributed to themost important work on Pakistan reviewing all the previously reported fossil crabs and describingtwo new genera and eight new species. Their work was completed by Glaessner and Secrétan(1987) with fossil crabs from the Eocene of the Sulaiman Range. More recently, Schweitzer et al.(2004) reported a new occurrence of hermit crab, a new species of ghost shrimp, and descriptionof one new genus from Pakistan. Most of these previous studies concerns fossil crabs collectedin northern Pakistan.
The current work includes fossil crabs collected during the late 1970s from the middleEocene (Lutetian, Domanda Formation) of Rakhi Nala, Sulaiman Range, and northern Pakistan.It includes also new crustaceans from the Paleocene (Thanetian? Bara Formation) exposed in theLakhra Dome, southern Pakistan. These latter occurrences constitute the first report of Paleocenecrustaceans in southern Pakistan (Sind, Karachi Arc).
2. Geological setting
The Sulaiman and Kirthar Ranges of Pakistan constitute a north-south trending fold and trustbelt which connects the Himalayan continental collision zone to the northeast with the Makranaccretionary wedge system to the southwest. The Sulaiman and Kirthar Ranges are generallyseparated by the Sibi Through, and the Kirthar Range is subdivided into northern and centralsectors close to the Sibi Re-entrant, and a southern sector, also called Karachi Arc (Fig. 1).
2.1. Eocene deposits of the Sulaiman Range (Upper Indus Basin)
The Sulaiman Range is a north-south trending band of rugged hills rising to 3500 m above sealevel that extends along the borders of Balochistan and Punjab Provinces (Fig. 1). Exposures of themiddle and upper Eocene marine formations of the Kirthar Group are extensive in the SulaimanRange, and they have produced the best middle and late Eocene fossil record for the entiresubcontinent. Indo-Pakistan was located astride the equator during the early Paleogene, movingnorthward toward tectonic collision with the rest of Asia (Patriat and Achache, 1984). The east
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Fig. 1. General map of Pakistan and location of the studied localities: Rakhi Nala (Domanda Formation, middle Eocene)and Lakhira Mine (Bara Formation, upper Paleocene).Carte générale du Pakistan et localisation des sites étudiés : Rakhi Nala (Formation Domanda, Eocène moyen) et MineLakhira (Formation Bara, Paléocène supérieur).
side of the Sulaiman Range has a thick sequence of well exposed Eocene sedimentary rocks, thatwere deposited on a shallow shelf on the leading edge of the subcontinent before closure of Tethysand suturing to the rest of Asia. The abundance of carbonate and the quasi absence of sandstonesand coarser clastic elements in the different formations of the Kirthar Group strongly contrastwith the overlying Oligo-Miocene Chitarwata Formation that documents the early consequencesof the collision and the rise of Himalayan Forelands. The uppermost formation of the KirtharGroup and the lowermost Chitarwata Formation are separated by a major unconformity whichcan be correlated to an intra-Early Oligocene orogenic phase (Welcomme et al., 2001; Métaiset al., 2009).
The Kirthar group comprises, in ascending order, the Habib Rahi, Domanda, Pir Koh, andDrazinda formations (Shah, 1991). The Kirthar Group is renowned as one of the richest archaeo-cete whale-bearing intervals known anywhere in the world (e.g., Gingerich et al., 2001). RakhiNala, the place where the fossils described herein were found in the late 1970’s, is the classic sec-tion of western Punjab documenting these lithological units, which were first described by Eames(1951, 1952a), and studied by successive authors (Latif, 1964; Siddiqui, 1971; Samanta, 1973;Köthe et al., 1988; Afzal, 1996; Jones, 1997). The age of the Kithar succession is constrainedby both biostratigraphic and sequence stratigraphic data (Gingerich et al., 2001: fig. 11). TheDomanda Formation, in which the presently studied decapod crustaceans were collected, consists
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of coloured (brownish, gray, green, orange, purple) shales and marls; this unit is the “Lower Choco-late Clays” of Eames (1952a). Bivalves and gastropod shell beds (Eames, 1952b,c), crab burrowsand crab fossils (Schweitzer et al., 2004), and archaeocete remains (Gingerich et al., 1997, 2001)are reasonably common in this formation. Although micropaleontological studies consistentlydescribe the Domanda Formation as barren of index microfossils, the age of the underlying HabibRahi Formation is early Lutetian based on the presence of NP14-15 nannoplankton (Köthe et al.,1988). Likewise, the Pir Koh Formation overlying the Domanda Formation has yielded plank-tonic foraminiferans corresponding to the Zone P12-13 (Samanta, 1973) that is consistent withan early Bartonian age. This indicates that the Domanda Formation is Lutetian in age. Attemptsat interpreting changes of lithology and facies in the Domanda Formation in term of sea levelchanges led Gingerich et al. (2001) to distinguish three distinct cycles of sea level flooding andshallowing that are correlative to global eustatic sequences (Haq et al., 1987), suggesting that thetime of formation of the Domanda sediments [about 300 m thick at the type section after Hemphilland Kidwai (1973: 16); same thickness at Rakhi Nala] can be estimated as Lutetian or so.
2.2. Lower Indus Coal Region (Lakhra Dome and Laki Range, also named Karachi Arc)
In the southern Kirthar Range, Paleocene and lower Eocene strata in the Lower Indus coal regionare assigned to the Ranikot Group and the Laki Formation, respectively. The name Ranikot thatrefers to a fairly enigmatic fort situated in the Laki Range was first introduced by Blanford (1879).In the sector of the Karachi Arc, between Thano Bula Khan to the South and Sehwan Sharif tothe North (Fig. 1), the area where the Ranikot Group is best exposed, it consists, in ascendingorder, of the Khadro, Bara, Lakhra, and Sohnari formations (the latter is sometimes consideredas the basalmost member of the Laki Formation). The Khadro Formation (Cardita beaumontibeds of early authors) of early Paleocene and perhaps latest Cretaceous age (Williams, 1959;Cheema et al., 1977) has also produced remains of decapod crustaceans that will be describedelsewhere. The Bara, Lakhra, and Sohnari formations, which overlie the Khadro Formation, havebeen described in several papers mainly on the basis of outcrops (Nuttall, 1932; Hunting SurveyCorporation Ltd, 1961; Cheema et al., 1977) but have become much better known as a result ofcoring in the region (Ghani et al., 1975; Outerbridge et al., 1991; Wakefield and Monteil, 2002).
A set of crustacean remains described herein comes from the Bara Formation and was bothcollected in the late 1970’s (for details see Gingerich et al., 1979) and more recently by threerecent field investigations (2009–2011). The Bara Formation (“Lower Ranikot” of early authors)includes non-marine, brackish-water, and coastal marine deposits; it consists mainly of green-grey,red-brown and grey-brown mudstones with common glauconite and occasional coals. Fossils arerare in the Bara Formation (450 m thick at the type locality Bara Nai) and the age estimation isquite uncertain. Palynological evidence indicates a shallow marginal marine, mangrove swamppalaeoenvironment (Frederiksen, 1994). At least five depositional sequences, with trangressiveand highstand sediments are interpreted from palynological evidence (Wakefield and Monteil,2002). The contact between the Bara Formation and the overlying Lakhra Formation appears tobe conformable and gradational at some places (Cheema et al., 1977; Outerbridge et al., 1991),but there is clear erosional unconformity on several sections of the Lakhra Dome1.
The Lakhra Formation (“Upper Ranikot” of early authors) appears to represent mainly shallowmarine to brackish-water deposition. This formation is widely exposed in the Lakhra Dome and
1 Métais et al., in prep.
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near Jhirak where rich assemblage of fossil molluscs was collected by E.W. Vredenburg andreported in Cossmann and Pissarro (1909). The Lakhra Formation contains some carbonaceousdetrital strata like the Bara, but the distinctive rocks of the Lakhra are foraminiferal limestone,marl, and shell hash beds. The Lakhra is generally much more calcareous, glauconitic, and shellythan the Bara. The Lakhra was deposited in a deeper-marine palaeoenvironment than the Bara,perhaps in front of some mangrove swamps.
The Bara and Lakhra formations have long been considered to be Paleocene in age, mainly onthe basis of larger foraminifers (Williams, 1959; Hunting Survey Corporation Ltd, 1961; Cheemaet al., 1977). According to Cheema et al. (1977), the Bara is not younger than early Selandian, andit is underlain by the Khadro Formation, which is early Paleocene (Danian) at least in its upperpart. Based on palynological data, Frederiksen (1994) concluded that the lower part of the BaraFormation is middle Paleocene. The Lakhra Formation, which unconformably overlies the Bara,has been assigned a late Paleocene (Thanetian) age based on planktonic foraminifera (Mohan,1982; Usmani, 1983). Wakefield and Monteil (2002) identified two Maximum Flooding Surfaces(MFS) in the Lakhra Formation, and possibly five MFSs in the Bara Formation on the basis ofsubsurface data (including lithology, biostratigraphy of pollens, nanno- and microfossils), seismicand gamma ray) on the Duljan-1 well-core, situated between the central and lower portions ofthe Indus Basin and about 100 km NE of the fossil localities studied here. It is worth noting thatthey did not recognize the Khadro or the Sohnari Formations in the Duljan succession. The firstMFS identified in the Lakhra Formation is near its base and correlates with the top of planktonicforaminiferal Zone P4 which is probably equivalent of MFSPg10 of Sharland et al. (2001). Thesecond MFS lies in the upper Lakhra and occurs in planktonic foraminiferal Zone P5/6. Thisbiostratigraphic calibration involves that the Lakhra Formation is straddling the Paleocene-Eoceneboundary, which is inconsistent with the chronostratigraphic studies published so far. The probableearly Eocene age of the upper Lakhra has to be tested from outcrops data in the area where thefossil described herein were collected. Fossil crustaceans described herein come from a ligniticlevel near the top of the Bara Formation. Consequently, it is reasonable to think that these fossilsare early Thanetian.
3. Material and methods
The palaeontological arrangement of this study follows the systematics proposed by De Graveet al. (2009).
3.1. Specimens from the Domanda Formation
The whole sample of 52 brachyurans includes complete and fragmentary carapaces rep-resenting different onthogenetic stages, and loose disarticulated chelipeds, all preserved asthree-dimensionally molds. The specimens were collected in the late 1970s from the middleEocene (Lutetian, Middle Domanda Formation) in different outcrops of Rakhi Nala, SulaimanRange, Northern Pakistan.
3.2. Specimens from the Bara Formation
The sample includes seven specimens of decapod crustaceans (axiideans and brachyurans)collected by different missions in the late 1970s and 2009–2011’s field seasons from the latePaleocene (Thanetian? Bara Formation) in two different localities situated on the northeastern
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rim of the Lakhra Dome, southern Kirthar Range, southern Pakistan. The specimens collectedby the 2011’s field season come from the Lakhira Mine (N25◦ 43′ 00.57′′, E68◦ 10′ 33.41′′) thatexploits the coal beds in subsurface at the top of the Bara Formation. The specimens collected bythe late 1970s’ expeditions come from another very close coal mine (N25◦ 40′, E68◦ 12′). All thespecimens are preserved as three-dimensional internal molds including complete carapaces andfragmentary chelipeds.
3.3. Abbreviations
MNHN.F: Collection de Paléontologie, Muséum national d’histoire naturelle, Paris, France.hpa: height of propodus.lpa: length of propodus (excluding fixed finger).lcxp: length of carapace.wcxp: width of carapace.
4. Systematic palaeontology
Class: MALACOSTRACA Latreille, 1802Order: DECAPODA Latreille, 1802Infraorder: AXIIDEA de Saint Laurent, 1979Family: CALLIANASSIDAE Dana, 1852Subfamily: EUCALLIACINAE Manning and Felder, 1991Genus Calliax de Saint Laurent, 1973
Type species: Callianassa lobata de Gaillande and Lagardère, 1966, by original designation.Included fossil species: see discussion by Hyzny and Hudácková (2012).
Calliax sp.Fig. 2A, B
Stratigraphic range:Late Paleocene, Thanetian? (Bara Formation).Occurrence: Lakhira Mine (N25◦ 43′ 00.57′′, E68◦ 10′ 33.41′′), northeastern rim of the Lakhra
Dome, southern Kirthar Range, southern Pakistan.Material and measurements: One incomplete minor right propodus (MNHN.F.A47685:
preserved lpa = 16 mm; hpa = 16 mm).Description: An incomplete minor right propodus, eroded proximally and at the lower distal
edge (the overall shape is retained); dactylus not preserved; index preserved only at the base;broken proximal tooth. Propodus ovoid in transversal section; smooth outer and inner surfaces;dorsal margin nearly straight with very smooth small alternate rounded spinules or teeth, wherethe distal last one more marked. Outer surface slightly convex, inner surface more compressed;both side bearing six to seven small sparse tubercles along the lower middle part of the propodus.Large triangular proximal tooth (broken distally) at the middle of the posterior cutting edge,separate by one rounded notch from the undentate index that is slightly directed downward. Infrontal view, the proximal tooth and the index are slightly divergent laterally each to other fromthe vertical median axis of the propodus.
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Fig. 2. Callianassid from the coal beds of the Bara Formation (upper Paleocene, Thanetian?) exploited in the LakhiraMine situated on the northeastern rim of the Lakhra Dome, southern Kirthar Range, southern Pakistan. A, B. Calliax sp.,incomplete right minor propodus (MNHN.F.A47685) in outer and inner views. Scale bar: 1 cm.Callianassidé des niveaux de charbon de la Formation Bara (Paléocène supérieur, Thanétien?) exploités dans la MineLakhira située sur la bordure nord-est du Dôme de Lakhra, sud de la montagne du Kirthar, sud du Pakistan. A, B.Calliax sp., propode mineur droit incomplet (MNHN.F.A47685) en vues externe et interne. Barre d’échelle : 1 cm.
Discussion: The specimen has oblique proximal margin, the index is very short and triangular inshape and there is a distinct tooth above the base of the index. All these characters fits to minor chelaof Calliax de Saint Laurent, 1973 sensu Ngoc-Ho (2003), in which the index is shorter than, andseparated from, the dactylus by a wide gap, bearing a large triangular proximal tooth (Ngoc-Ho,2003: 490, fig. 17e; Sakai, 1999: figs. 28, 29c; see also Hyzny, 2012: fig. 2h). No other callianassidgenus has similarly shaped major or minor cheliped. So, due to the poor preservation, and in lackingof others specimens for comparison, we ascribe tentatively the specimen to Calliax sp.
According to Ngoc-Ho (2003) and our own observations, the generic re-assignment toCalliax de Saint Laurent, 1973 proposed by Schweitzer et al. (2003) for Callianassa whiteavesiWoodward, 1886 (Campanian, Canada) is not justified and the taxon can not be considered as amember of Calliax. Indeed, the absence of the diagnostic proximal tooth on the distal marginbetween the fingers (dactylus and pollex) in C. whiteavesi questioned the affinity to Calliax.Moreover, according to Matús Hyzny (pers. comm. 2012), there are few reports on unidentifiedcallianassid material which might represent Calliax but none of them is as old as the studiedspecimen. Consequently, the specimen presently reported in the Bara Formation, Pakistan is theoldest record for the genus, enlarging the fossil range to the Paleocene.
Infraorder: BRACHYURA Latreille, 1802Section: EUBRACHYURA de Saint Laurent, 1980Subsection: HETEROTREMATA Guinot, 1977Superfamily: CARPILIOIDEA Ortmann, 1893Family: CARPILIIDAE Ortmann, 1893Genus Proxicarpilius Collins and Morris, 1978
Type species: Proxicarpilius planifrons Collins and Morris, 1978, by monotypy.Included fossil species: Proxicarpilius planifrons.
Proxicarpilius planifrons Collins and Morris, 1978
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Figs. 3–5, 6A–IProxicarpilius planifrons Collins and Morris, 1978: 972–976, pl. 117, figs. 5, 7, pl. 118, figs.
1–4, 7.Proxicarpilius minor Collins and Morris, 1978: 976, pl. 117, figs. 6, 8, 9, pl. 118, figs. 5, 6, 11.Palaeocarpilius minor – Glaessner and Secrétan, 1987: 5, 6, pl. 1, fig. 1.Proxicarpilius planifrons – Schweitzer et al., 2004: 101, 102, fig. 6. – Schweitzer et al., 2010:
(Middle Domanda Formation).Occurrences, material and measurements:Middle Domanda Formation (Middle Eocene, Lutetian). – All the specimens come from differ-
ent outcrops at Rakhi Nala, Sulaiman Range, northern Pakistan. The measurements are reportedonly for the complete smaller male (MNHN.F.A47736) and the bigger female (MNHN.F.A47740)specimens. – Locality 1: one complete carapace (MNHN.F.A47728); one complete male carapaceshowing abdomen (MNHN.F.A47729); one left palm (MNHN.F.A47730); three disarticulatedchelipeds and one incomplete carapace (MNHN.F.A47731). – Locality 2: one complete femalecarapace with abdomen (MNHN.F.A47732). – Locality 3: one complete female carapace withabdomen (MNHN.F.A47733); three carapaces with abdomen, seven nearly complete cara-paces, 12 fragmentary carapaces, and two loose carpi (MNHN.F.A47734); one complete malecarapace with abdomen (MNHN.F.A47735); three complete male carapaces with abdomen(MNHN.F.A47736: lcxp = 14.8 mm; wcxp = 20.0 mm, A47737, A47738); two right chelae(MNHN.F.A47739, A47868); one complete female carapace with abdomen (MNHN.F.A47740:lcxp = 23.3 mm; wcxp = 33.0 mm); two carapaces (MNHN.F.A47726, A47727). – Locality 4:one carapace (MNHN.F.A47741); one complete carapace with abdomen (MNHN.F.A47742);one nearly complete carapace (MNHN.F.A47743). – Localities unreported: two incompleteright palms (MNHN.F.A47724, A47869); one right palm, one left palm, and one right carpus(MNHN.F.A47725).
Bara Formation (Late Paleocene, Thanetian?). – All the specimens come from an unnamedcoal mine (N25◦ 40′, E68◦ 12′) situated on the northeastern rim of the Lakhra Dome, southernKirthar Range, southern Pakistan: one complete carapace (MNHN.F.A47691: lcxp = 28.5 mm;wcxp = 43.0 mm); one nearly complete carapace preserving part of the abdomen (MNHN.F.A47692: lcxp = 23.0 mm; wcxp = 30.5 mm); one carapace preserving the abdomen and incompleteright propodus (MNHN.F.A47693); one incomplete carapace (MNHN.F.A47694).
Discussion: Proxicarpilius has been erected by Collins and Morris (1978), including twospecies, P. planifrons Collins and Morris, 1978 and P. minor Collins and Morris, 1978. Later,Schweitzer et al. (2004: 101, 102), based on new observations of additional specimens fromPakistan, suggested that P. minor is junior synonym of P. planifrons.
According to Schweitzer et al. (2004), all the studied specimens from Rakhi Nala (DomandaFormation) are assigned to P. planifrons for the following characters: ovate carapace wider thanlong, slightly vaulted transversely, strongly vaulted longitudinally; produced front, directed down-ward; inflated protogastric region; depressed hepatic region; anterolateral margin with three bluntteeth and triangular postorbital tooth; epibranchial tooth longer, directed laterally; sinuous meso-gastric and metabranchial ridges across the axis more or less marked. The characters of male andfemale abdomens, well-preserved in some specimens, have been also previously described byCollins and Morris (1978: 974). Moreover the loose palm and carpus share close affinities withthe specimens figured by Collins and Morris (1978: figs. 5, 6).
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Fig. 3. Specimens of Proxicarpilius planifrons Collins and Morris, 1978 from the Domanda Formation (middle Eocene,Lutetian) collected at different localities at Rakhi Nala, Sulaiman Range, northern Pakistan. A–C. Complete male cara-pace with abdomen (MNHN.F.A47735), dorsal, ventral and frontal views. D. Complete female carapace with abdomen(MNHN.F.A47733), ventral view. E–F. Complete male carapace with abdomen (MNHN.F.A47737), dorsal and ventralviews. G–I. Complete male carapace with abdomen (MNHN.F.A47738), dorsal, ventral and frontal views. J. Completecarapace (MNHN.F.A47727), dorsal view. K–L. Complete carapace (MNHN.F.A47726), dorsal and frontal views. Scalebars: 1 cm.Spécimens de Proxicarpilius planifrons Collins and Morris, 1978 de la Formation Domanda (Eocène moyen, Lutétien)récoltés dans différents affleurements à Rakhi Nala, montagne de Sulaiman, nord du Pakistan. A–C. Carapace mâlecomplète avec l’abdomen (MNHN.F.A47735), vues dorsale, ventrale et frontale. D. Carapace femelle complète avecl’abdomen (MNHN.F.A47733), vue ventrale. E–F. Carapace mâle complète avec l’abdomen (MNHN.F.A47737), vuesdorsale et ventrale. G–I. Carapace mâle complète avec l’abdomen (MNHN.F.A47738), vues dorsale, ventrale et frontale.J. Carapace complète (MNHN.F.A47727), vue dorsale. K–L. Carapace complète (MNHN.F.A47726), vues dorsale etfrontale. Barres d’échelle : 1 cm.
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Fig. 4. Specimens of Proxicarpilius planifrons Collins and Morris, 1978 from the Domanda Formation (middle Eocene,Lutetian) collected in different localities at Rakhi Nala, Sulaiman Range, northern Pakistan. A–C. Complete female cara-pace with abdomen (MNHN.F.A47740), dorsal, ventral and frontal views. D–F. Complete male carapace with abdomen(MNHN.F.A47736), dorsal, ventral and frontal views. G–I. Complete carapace with abdomen (MNHN.F.A47742), dorsal,ventral and frontal views. J–K. Near-complete carapace (MNHN.F.A47743), dorsal and frontal views. Scale bars: 1 cm.Spécimens de Proxicarpilius planifrons Collins and Morris, 1978 de la Formation Domanda (Eocène moyen, Luté-tien) récoltés dans différents affleurements à Rakhi Nala, montagne de Sulaiman, nord du Pakistan. A–C. Carapacefemelle complète avec l’abdomen (MNHN.F.A47740), vues dorsale, ventrale et frontale. D–F. Carapace mâle complèteavec l’abdomen (MNHN.F.A47736), vues dorsale, ventrale et frontale. G–I. Carapace complète avec l’abdomen(MNHN.F.A47742), vues dorsale, ventrale and frontale. J–K. Carapace subcomplète (MNHN.F.A47743), vues dorsale etfrontale. Barres d’échelle: 1 cm.
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Fig. 5. Chelipeds of Proxicarpilius planifrons Collins and Morris, 1978 from the Domanda Formation (middleEocene, Lutetian) collected in different localities at Rakhi Nala, Sulaiman Range, northern Pakistan. A–C. Right chela(MNHN.F.A47739), outer, inner and ventral views. D–F. Right chela (MNHN.F. A47868), outer, inner and ventral views.G–H. Right carpus (MNHN.F.A47725), outer and inner views. Scale bars: 1 cm except G–H: 5 mm.Chélipèdes de Proxicarpilius planifrons Collins and Morris, 1978 de la Formation Domanda (Eocène moyen, Luté-tien) récoltés dans différents affleurements à Rakhi Nala, montagne de Sulaiman, nord du Pakistan. A–C. Pince droite(MNHN.F.A47739), vues externe, interne et ventrale. D–F. Pince droite (MNHN.F. A47868), vues externe, interne etventrale. G–H. Carpus droit (MNHN.F.A47725), vues externe et interne. Barres d’échelle : 1 cm excepté G–H : 5 mm.
The specimens from the northeastern rim of the Lakhra Dome (coal mine, Bara Formation)show the main characters of the monospecific genus Proxicarpilius Collins and Morris, 1978. Theyare confidently ascribed to P. planifrons Collins and Morris, 1978, as emended by Schweitzer et al.(2004).
P. planifrons was previously reported from the Eocene of several northern Pakistan localitiesin the Derha Ismail Khan district (Collins and Morris, 1978; Schweitzer et al., 2004) and from theSanghach Nala section of Sulaiman Range (Glaessner and Secrétan, 1987). The new occurrencesat Rakhi Nala attest that P. planifrons is the most widespread and common carpiliid crab in theEocene faunal assemblage from northern Pakistan. The occurrence of P. planifrons in the coalbeds of the Bara Formation on the northeastern rim of the Lakhra Dome, southern Kirthar Range,southern Pakistan, is the first report from the Paleocene of Pakistan, enlarging the stratigraphicand the palaeobiogeographical ranges for this species.
Genus and species indeterminateFig. 7Stratigraphic range: Middle Eocene, Lutetian (Middle Domanda Formation).
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Fig. 6. Crabs from the coal beds of the Bara Formation (upper Paleocene, Thanetian?) exploited in an unnamed mine (A–I,K–L) and in the Lakhira Mine (J) both situated on the northeastern rim of the Lakhra Dome, southern Kirthar Range,southern Pakistan. A–C. Proxicarpilius planifrons Collins and Morris, 1978, complete carapace (MNHN.F.A47691), dor-sal, ventral and frontal views. D–F. P. planifrons Collins and Morris, 1978, nearly complete carapace (MNHN.F.A47692),dorsal, ventral and frontal views. G–I. P. planifrons Collins and Morris, 1978, carapace (MNHN.F.A47693) preser-ving the abdomen and incomplete right propodus, dorsal, ventral and frontal views. J. Pakicarcinus orientalis (Collinsand Morris, 1978), carapace (MNHN.F.A47687), dorsal view. K–L. Xanthoidea indeterminate, incomplete left palm(MNHN.F.A47690), outer and inner views. Scale bars: 1 cm.Crabes des niveaux de charbon de la Formation Bara (Paléocène supérieur, Thanétien?) exploités dans une mine sansnom (A–I, K–L) et dans la Mine Lakhira (J) situées toutes les deux sur la bordure nord-est du Dôme de Lakhra, sud dela montagne du Kirthar, sud du Pakistan. A–C. Proxicarpilius planifrons Collins and Morris, 1978, carapace complète(MNHN.F.A47691), vues dorsale, ventrale et frontale. D–F. P. planifrons Collins and Morris, 1978, carapace presque
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Fig. 7. Carpiliid crab from the Domanda Formation (middle Eocene, Lutetian) collected at Rakhi Nala, Sulaiman Range,northern Pakistan. A–C. Genus and species indeterminate, carapace MNHN.F.A47744 in dorsal, ventral and frontal views.Scale bars: 5 mm.Crabe carpiliidé de la Formation Domanda (Eocène moyen, Lutétien) récolté à Rakhi Nala, montagne de Sulaiman, norddu Pakistan. A–C. Genre et espèce indéterminés, carapace MNHN.F.A47744 en vue dorsale, ventrale et frontale. Barresd’échelle : 5 mm.
Occurrence, material and measurements: Rakhi Nala, Sulaiman Range, northern Pakistan.– Locality 3: one complete carapace preserving the abdomen (MNHN.F.A47744: lcxp = 12.0 mm;wcxp = 19.5 mm).
Description: small carapace, poorly preserved dorsally but with well-preserved abdomen;smooth, convex carapace, wider than long, oval transversally; smooth, indistinct regions; wide,straight?bilobate front turned downward; moderately elongate orbits; entire anterolateral mar-gin, ending in a lateral smooth rounded tooth; rounded posterolateral margin, slightly straightposteriorly.
Discussion: the specimen, although poorly preserved dorsally, differs from P. planifrons forwider and shorter smooth carapace lacking transverse ridges; less produced front and only twoanterolateral smooth teeth (including epibranchial tooth). The poor preservation of the specimen,however, does not allow closer assignment within the Carpiliidae.
Type species: Lobonotus orientalis Collins and Morris, 1978, by monotypy.Included fossil species: Pakicarcinus orientalis (Collins and Morris, 1978); P. whittenensis
(Glaessner, 1980).
Pakicarcinus orientalis (Collins and Morris, 1978)Fig. 6JLobonotus orientalis Collins and Morris, 1978: 970–972, pl. 116, figs. 10, 11, pl. 117, fig. 1.
complète (MNHN.F.A47692), vues dorsale, ventrale et frontale. G–I. P. planifrons Collins and Morris, 1978, carapace(MNHN.F.A47693) préservée avec l’abdomen et le propode droit incomplet, vues dorsale, ventrale et frontale. J. Paki-carcinus orientalis (Collins and Morris, 1978), carapace (MNHN.F.A47687), vue dorsale. K–L. Xanthoidea indéterminé,paume gauche incomplète (MNHN.F.A47690), vues externe et interne. Barres d’échelle : 1 cm.
114 S. Charbonnier et al. / Annales de Paléontologie 99 (2013) 101–117
Pakicarcinus orientalis – Schweitzer et al., 2004: 110, 111, fig. 8a,b. – Schweitzer et al., 2010:123.
Type horizon: Ghazij Formation (Lower Eocene).Stratigraphic range: Late Paleocene, Thanetian? (Bara Formation) – Early Eocene, (Ghazij
Formation).Occurrence, material and measurements: the material comes from the Late Paleocene,
(Thanetian? Bara Formation) of the Lakhira Mine (N25◦ 43′ 00.57′′, E68◦ 10′ 33.41′′), north-eastern rim of the Lakhra Dome, southern Kirthar Range, southern Pakistan. – One completesmall-sized carapace preserved in matrix (MNHN.F.A47687: lcxp = 10.0 mm; wcxp = 12.5 mm).
Discussion: Schweitzer et al. (2004) described the new genus Pakicarcinus to accommodateLobocarcinus orientalis Collins and Morris, 1978 from the lower Eocene Ghazij Formation, southof Nila Kund, Dera Ghazi Khan, Punjab Province, Pakistan.
The new specimen MNHN.F.A47687 is assigned to P. orientalis for the hexagonal carapace,with well-defined regions, wide subtruncate front, U-shaped median notch, spiniform externalorbital tooth, anterolateral margins with four teeth, and posterolateral margins nearly straight,converging to the posterior margin. It is the first report of P. orientalis in the Paleocene ofsouthern Pakistan, enlarging the stratigraphic and the palaeobiogeographical ranges of this species.
Xanthoidea indeterminateFig. 6K, LStratigraphic range: Late Paleocene, Thanetian? (Bara Formation).Occurrence, material and measurements: unnamed coal mine (N25◦ 40′, E68◦ 12′) situated
on the northeastern rim of the Lakhra Dome, southern Kirthar Range, southern Pakistan. – oneincomplete left palm (MNHN.F.A47690: lpa = 27.0 mm; hpa = 28.4 mm).
Discussion: fragmentary preserved stout subquadrate palm, slightly higher than long, lackingboth the robust fingers. Smooth edges; inflated dorsally and with curved smooth dorsal margin.The specimen is poorly preserved as an internal mould and partially washed. The general formof the palm and the absence of spines on the upper margin do not exhibit affinities to any of thealready known genera of xanthoids from Pakistan. Due to the poor preservation, the specimen istentatively ascribed to an indeterminate xanthoid sensu lato.
5. Conclusions
This study enlarges notably our knowledge of the stratigraphical and palaeobiogeogaphicalranges of carpiliid and xanthoid crabs and axiideans from the Paleogene of Pakistan. The taxadescribed in the Eocene of northern Pakistan are now reported in the Paleocene of southernPakistan. The future field seasons in Sind (Karachi Arc) will probably provide new discoverieson the Cenozoic decapod crustaceans from Pakistan.
Acknowledgements
We thank all the participants to the Ranikot field Project since 2009, including Pierre-OlivierAntoine, Stéphane Guillot, Asghar Hakro, Mashooq Warar, Kashif Memoon, Kashif Samoon,and Muhammad Hassan Agheem. This project benefited from the enthusiasm and early sup-port of Dr. Parveen Usmani (then head of the Geology Department of the University of Sindh).The French Consuls Pierre Seillan and Christian Ramage, and Kamal Majidullah (Karachi) are
S. Charbonnier et al. / Annales de Paléontologie 99 (2013) 101–117 115
thanked for their interest and support. The French Embassy and CampusFrance, and the MNHN-Paris financially supported study stay of Rafiq A. Lashari and Sarfraz H. Solangi, respectively.We thank Philippe Loubry (UMR-CNRS 7207, MNHN) for assistance in photographic work.This collaborative research programme was supported by the Depatment of pure and appliedGeology (University of Sindh), the Action Transversale du Muséum “Biodiversité actuelle etfossile – crises, stress, restaurations et panchronismes : le message systématique”, and the ANR-Palasiafrica program (ANR-08-JCJC-0011-01-ANR-ERC). This paper is a contribution fromthe UMR-CNRS 7207 Centre de Recherche sur la Paléobiodiversité et les Paléoenvironnements(CR2P) as well as the Département Histoire de la Terre (MNHN, Paris). We thank the editor andthe two reviewers Francisco J. Vega and Matús Hyzny for their constructive remarks on the initialversion of the manuscript.
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