Alloptox Dawson (1969) (1988), (1979), Ochotona, Lagomorpha Brandt, 1855 Ochotonidae Thomas, 1897 Alloptox Da wson, 1961 Alloptox gobiensis ( Y

Transcription

1 VERTEBRATA PALASIATICA pp figs. 1 4, ) 1) Alloptox gobiensis( ( ) ( ) ( Alloptox gobiensis), Al2 loptox :, 6,,,,, Alloptox Kenyalagomys, Sinolagomyinae Alloptox Kenyalagomys, Sinolagomys,,,, Q ,, Alloptox,, (1991), Al2 loptox sp., Alloptox gobiensis : ( = 28 ), Alloptox gobiensis Dawson(1961) Alloptox, 6 2 : A. xichuanensis (,1997), A. sihongensis (, 1995) ; A. gobiensis ( Young, 1932 ;, 1991), A. minor (, 1978), A. chinghaiensis (, 1981), Candir A. anatoliensis ( Unay and Sen, 1976) ; (?) Al2 loptox sp. (Boule and Teilhard, 1928), Alloptox sp. (,1991) Alloptox, 1) ( :J ) ( : ) :

2 Alloptox Dawson (1969) (1988), (1979), Ochotona, Lagomorpha Brandt, 1855 Ochotonidae Thomas, 1897 Alloptox Da wson, 1961 Alloptox gobiensis ( Young, 1932) ( 1 4 ; 1 4) 4 : (V ), ; (V ) ; (V ),, (7 11 ) : (V ), ; (V ) ; (V ), (14 15 ) : 2 (V ), ; (V ), ; (V ), ; (V ), (26 ) : 3 (V ) ; (V ), m3 ; (V ), m3, p3 ( = 28 ) : (V ), ; (V ), ; (V ), ; (V ), ; (V ),, ; (V ), ; (V ),, ; (V ), ; (V ), ; (V ),, ; (V ), ; (V ),, ; (V ), ; (V ), ; (V ),p4 ; (V ),p4 ; (V ),m2 ; (V ), m2 ; (V ), m3 ; (V ), m3 ; (V ),m3, ; ( 1 2),, P3 P4

3 2 : Alloptox gobiensis (, ) 117,,,,, Ochotona,,,,,P2,,,,, P3 P4 1 Table 1 Skull measurements of Alloptox gobiensis (mm) V V V V V V V V Incisor2palatal foramen length Palatal bridge length Length of maxillary component of palatal bridge Length of pala2 tine component of palatal bridge Upper diastema length Upper cheek tooth row length Length of bulla Width of bulla Width of maxilla Breadth of palatal Breadth of medi2 al part of nasal Breadth of pos2 terior part of nasal Bizygomatic breadth Interorbital breadth Breadth of anterior part of parietal Breath of posterior part of parietal Supraoccipital length Supraoccipital breadth Ca Ca

4 Table 2 Measurements of mandible of Alloptox gobiensis (mm) V V V V Lower diastema length Lower cheek tooth length 13 (p3 - m3) (p3 - m3) 11 (p3 - m2) Height of the ascending ramus Length of the angular process Height of the horizontal ramus 9. 9 (m3) / (p4) 8. 1 (p3) 9. 6 (m3) / (p3),,,,,,, P4, P4 M1,, M1,, Ochotona,,,,,,,,,,, V , 4,,,,,,,,,,,,,

5 2 : Alloptox gobiensis (, ) Fig. 1 Composite skull of Alloptox gobiensis (Based on : V , V , V , V , V , V ) A. dorsal view ; B. ventral view ; C. lateral view Abbreviations : b2occ, basioccipital ; bs, basisphenoid ; bu, bulla ; cho, choana ; ex caro f, external carotid foramen ; fr, frontal ; hyp for, hy2 poglossal foramen ; inc f, incisive foramen ; inf2orb f, infraorbital foramen ; ju, jugal ; jug f, jugular foramen ; medi sept, medial septum ; mx, maxilla ; na, nasal ; opt f, optic foramen ; orb, orbit ; orb fis, orbital fissure ; pa, parietal ; p f, premolar foramen ; pl bri, palatal bridge ; pl f post, palatal foramen posterior ; pmx, premaxilla ; s2occ, supraoccipital ; s orb cr, supraor2 bital crest ; sq, squamosal ; tem cr, temporal crest ; tr opn mx, triangular opening of the maxilla ; zyg mass tub, zygomatic masseter tuberosity,

6 120 41,, m2,, m3,p3,,, p3,, p4 m1,, 2, = 1cm, Fig. 2 Occipital view of Alloptox gobiensis (V ),, scale bar = 1cm, Abbreviations : bu, bulla ;ex occ cr,, external occipital crest ;for mag, foramen magnum ;nu cr, nuchal crest ;occ con,, occipital condyle Kenyalagomys rusingae Bellatona yanghuensis Prolagus sardus Ochotona nihewanica MacInnes (1953) Rusinga Kenyalagomys rusingae, Alloptox gobiensis : 1) K. rusingae ; A. gobi2 ensis 2) K. rusingae,, ; A. gobiensis 3) K. rusingae ; A. gobiensis, 4), K. 3 rusingae Fig. 3 Composite jaw of Alloptox gobiensis (Based on :, P4 M1 ; A. go2 V , V ) A. lingual view ; B. labial view biensis P4 Abbreviations : ang pro, angular process ;, 5 ) K. as ra, ascending ramus ; con, condyle ; rusingae, ; cor pro, coronoid process ; men for, mental A. gobiensis, Ochotona foramen ; swl, swelling

7 2 : Alloptox gobiensis (, ) 121 Kenyalagomys rusingae Alloptox gobiensis :1) 2), 3), 4) 5),, 6) 7), 8) 9),, 10), p3, m1,, p3,m3,, m2 11),, (1988) Bellatona yanghuen2 sis, Alloptox gobiensis :1) B. yanghuensis, A. gobiensis 2) B. yanghuensis ; A. gobiensis 3) B. yanghuensis ; A. gobiensis 4), B. yanghuensis ; A. gobiensis 5) B. yanghuensis,,,, ; A. gobiensis,, 6) A. gobiensis, B. yanghuensis, 7) B. yanghuensis ; A. gobiensis 8) B. yanghuensis,p3,m3, Ochotona ; A. gobiensis, Bellatona yanghuensis Alloptox gobiensis : ; ; ;,, Dawson (1969) Corsica Sardinia Prolagus sardus, Prolagus sardus Alloptox gobiensis :1) P. sardus ; A. gobiensis 2) P. sardus, ; A. gobiensis, 3) P. sardus, ; A. gobiensis 4) P. sardus, ; A. gobiensis, 5) P. sardus P4 M1 ; A. gobiensis, P4 P3 P4 6) P. sardus ; A. gobiensis 7) P. sardus,a. gobiensis 8) P. sardus,a. gobiensis 9) P. sardus, A. gobiensis 10) P. sardus ; A. gobiensis 11) P. sardus ; A. gobiensis 12) P. sardus

8 122 41, ; A. gobiensis,, 13) P. sardus ; A. gobiensis 14) A. gobiensis P. sardus Prolagus sardus Alloptox gobiensis : 1) 2) 3) 4), 5) 6) 7) 8), p3, m1, Ochotona 14 (1985) Ochotona, Ochotona nihewanica, Ochotona Alloptox gobiensis, O2 chotona :1) Ochotona cansus Ochotona thomasi, 12 Ochotona nihewanica ; A. gobiensis 2), Ochotona, A. gobiensis 3), Ochotona, A. gobiensis 4) Ochotona M2 M1 ; A. gobiensis 5) Ochotona ; A. gobi2ensis 6) Ochotona ; A. gobiensis 7) Ochotona ; A. gobiensis 8) O2 chotona,a. gobiensis 9) A. gobiensis Ochotona 10) Ochotona ; A. gobiensis 11) Ochotona, ; A. gobiensis 12) Ochotona ; A. gobiensis 13) Ochotona ; A. gobiensis 14) Ochotona, p3,m3 ; A. gobiensis, m3,p3 15) Ochotona, ; A. gobien2 sis, Alloptox gobiensis Ochotona : ; ;, ; Alloptox Alloptox Ochotona,,,, 1) Alloptox gobiensis Kenyalagomys rusingae, Prolagus sardus, Ochotona,

9 2 : Alloptox gobiensis (, ) 123, 2) Alloptox gobiensis, Kenyalagomys rusingae, Prolagus sardus, K. rusingae A. go2 biensis, P. sardus, Ochotona (Dawson, 1969), P. sardus A. gobiensis K. rusin2 gae,,,,, (Wood, 1940) Palaeolagus Alloptox Kenyalagomys, (Wood, 1940), Palaeolagus hypsodus Palaeolagus haydeni,,,, 3) Palaeolagus Prolagus, Dawson (1969), Alloptox gobiensis,, Dawson, Alloptox Kenyalagomys Palaeolagus, Prolagus Ochotona,, Dawson Prolagus Ochotona 4) Alloptox gobiensis, Ochotona,, Ochotona Ochotona 3, (1979),,,,, A. gobiensis 4 2,, A. gobiensis K. rusingae,,, Alloptox gobiensis Ochotona Prolagus,, Alloptox gobiensis, 5) Dawson (1969) Kenyalagomys Prolagus,, Alloptox gobiensis Dawson Alloptox gobiensis Kenyalagomys Prolagus, p3, m1, Sinolagomys (Bohlin, 1942) Palaeolagus (Wood, 1940 ; Dawson, 1958), Alloptox gobiensis (Bohlin (1942)

10 (1989) Sinolagomys, Bohlin, Bohlin, Bohlin ) (1985) Ochotona nihewanica, p3, ; m2, 14 Ochotona, O. O. cansus O. thibetana O. himalayana daurica p3,, O. nihewanica,,,,, Dawson (1969) Ochotona,, A. gobiensis ; Ochotona, A. gobiensis 6) Alloptox gobiensis, Kenyalagomys Bellatona Sinolagomys Palaeolagus,, Prolagus sardus Ochotona nihewanica Ochotona,,, Alloptox Kenyalagomys,p3,m3,, Sinolagomys (Bohlin, 1942 ;,1989) Alloptox gobiensis, Alloptox Erbajeva (1994) Ochotonidae : Sinolagomyi2 nae Ochotoninae ; Alloptox Ochotoninae, Alloptox Bellatona, Ochotona Erbajeva p3 Sinolagomyinae p3,, Ochotoninae p3, Erbajeva, Kenyalagomys Bellatona Alloptox Prolagus Ochotona, Alloptox Ochotonidae Palaeolagus ( Wood, 1940 ; Dawson, 1958), 0 30 (,Table 3),, (Un2 ordered)?, 1 21 Ochotona,? 6 (,Table 4) PAUP beta, DELTRAN, ( Exhaustive search), (Length) 44, (CI) , : 1), Kenyalagomys Alloptox ; Bellatona Prolagus Ochotona Erbajeva (1994) O2 chotonidae, Alloptox Kenyalagomys

11 2 : Alloptox gobiensis (, ) 125, Bellatona, Alloptox Kenyalagomys Sinolagomyi2 nae ; Bellatona Prolagus Ochotona Ochotoninae Erbajeva (1994) Alloptox Kenyalagomys : (6) ; (9), ; (10) ; (16) ; (17), ; (18) ; (22) Bellatona Prolagus O2 chotona : (4) ; (28) Bellatona Alloptox, Alloptox Sinolagomyinae (1995) (1989) Sinolagomys ulungurensis p3, Alloptox Sinolagomys ulungurensis Sinolagomys, Alloptox Sinolagomyinae Sinolagomys, 2) Erbajeva (1994) Ochotoninae Bellatona, Bellatona Ochotona Prolagus, Bellatona Ochotona Prolagus, Bellatona Erbajeva (1994) 3) Erbajeva (1994) Prolagus, Prolagus O2 chotona, Ochotoninae : (19) ; (20) ; (26) ; (29) ; (30) Alloptox Kenyalagomys, Alloptox Kenyalagomys, Alloptox A. xichuanensis (,1997) A. sihongensis (,1995), MN4,Alloptox, Kenyalagomys, 3, ( MN4 ) K. rusingae K. minor(macinnes, 1953), Rusinga ; ( MN7) K. mellalensis (Janvier and Muizon, 1976) Beni Mellal, Alloptox Kenyalagomys, San2tao2ho Bohlinotona pusilla ( Erbajeva and Sen, 1998) Taben2buluk Sinolagomys (Bohlin, 1942) Muizon (1977) (1989) Kenyalagomys Sinolagomys (1995) Alloptox Sinolagomys ulungurensis (,1989) Alloptox Kenyalagomys, Sinolagomys, Kenyalagomys, Agenian2Orleanian( MN32MN4),

12 Fig. 4 Phylegenetic relationship of Alloptox ( Gomphotherium Landbridge),, (Bernor et al., 1988 ; R ogl, 1998 ; Harrison and Gu, 1999), Kenyalagomys,, ; ; ;, ;,,! THE CRANIAL MORPHOLO GY AND PHYLO GENETIC RELATIONSHIP OF ALLOPTOX GOBI ENSIS ( LAGOMORPHA, OCHOTONIDAE) WU Shao2Yuan ( Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences Beijing ) ( The Graduate School of Chinese Academy of Sciences Beijing ) Key words Tongxin of Ningxia, Middle Miocene, Ochotonidae, Alloptox, skull Summary Lagomorpha Brandt, 1885 Ochotonidae Thomas, 1897 Alloptox Da wson, 1961 Alloptox gobiensis ( Young, 1932)

13 2 : Alloptox gobiensis (, ) 127 Locality and horizon Tongxin County, Ningxia Hui Autonomous Region, China ; Hongliugou Formation, Middle Miocene. Description Skull : The incisive foramen, narrowing anteriorly, is confluent with the palatal foramen. Inside the joint2foramen a bony lamina, which is formed by nasal septum and vomer, com2 pletely separates this foramen in the middle line. The nasal bone is slightly narrower posteriorly than anteriorly. The nasal2frontal suture terminates at a point between the anteroventral margin and an2 tero2dorsal margin of the orbit. A large acute triangular opening occurs on the lateral surface of the maxilla. The smallest angle of this triangle points anteriorly. Below the opening is a lacework of bone. The infraorbital foramen is situated above P2 and below the lacework of bone, in a more forward position to the anterior zygo2 matic root than in Ochotona. The strongly hypsodont cheek teeth extend into the maxillary tuberosity that contacts the zygoma along the entire length of the tuberosity. The palatine component of the pa2 latal bridge is reduced, as opposed to Ochotona in which the maxillary component of the palatal bridge is reduced. A pair of premolar foramina is situated in the maxilla, the position of which va2 ries in different specimens at somewhere between P3 talon and P4 trigon. The palatine2maxillary su2 ture appears like shallow W. The choana is narrow, and ends anteriorly at the level of M1. The two walls of choana are nearly parallel. The anterior lacerum foramen is a long fissure. The basisphe2 noidale and the presphenoidale extend a process into this foramen respectively. The process from the basisphenoidale is much longer and thinner than that from presphenoidale, and a very small foramen appears at the posterior root of the basisphenoidale process. The zygoma is a vertical tabulate bone, with a deep anterolateral masseter concavity and a well2 developed ventral process. The jugal has a long tapering process posterior to the posterior zygomatic root. The jugal process of squamosal borders the orbit ventroposteriorly. Posterior to the zygomatic root the squamosal extends back along the parietal and terminates above the meatus. The roof of the skull formed by frontal and parietal is relatively flat. The frontal is wide. The two sides of frontal, which form the dorsal border of the orbit, are nearly parallel. The most con2 spicuous feature of the frontal of Alloptox is that the supraorbital ridge starts near the premaxilla2 frontal suture and extends backwards along the upper margin of the orbit. The temporal crest is well developed, which starts after the posterior zygomatic root, extends backwards along the whole length of the parietal2squamosal suture. There is a distinct depression in the posterior area of parietal. This is bounded posteriorly by a ridge along the parietal2supraoccipital suture, and a distinct crest is present along the posterior part of the sagittal parietal suture. No specimens of Alloptox show a sepa2 rate interparietal. The supraoccipital extends back in approximately the same plane as the parietals for a long distance. The nuchal crest along the posterior margin of supraoccipital is well developed. The occipital is almost vertical, and considerably higher than that of Ochotona. There is a conspicu2 ous external occipital crest from the nuchal crest to the foramen magnum along the sagittal line. La2 teral to the external occipital crest the occipital surface is pitted, which seems to serve for the mus2 cle attachment. The basioccipital surface is slightly depressed. The medial ridge of basioccipital is well developed, which is higher posteriorly than anteriorly, and terminates half length of the basioc2 cipital. In the orbitosphenoid the optic foramen is confluent with that of the other side, and forms a large inter2orbital fenestra as in other lagomorphs. Behind the orbit fissure five foramina pierce the alisphenoid in a row. Between the first two foramina there is also a small foramen on a relatively high position. The third and fourth foramina are separated externally by a narrow bony splint, but connect with each other internally. The bulla is rounded and high, and proportionately larger than that in modern Ochotona. The meatus is short, as in Ochotona. Lateral to the basioccipital the walls of bulla are pierced by the external carotid foramen. The internal structure of bulla is not preserved very well, but it seems that the wall of bulla is not a mass of spongy bone, as in Ochotona. The bulla is thin walled. The jugu2

14 lar foramen, which occurs between the bulla and occipital condyle, is small and rounded. This con2 dition is in contrast to other lagomorphs, in which the jugular foramen is a long fissure. Above the jugular foramen, there is another small foramen penetrating the bulla. Other lagomorphs have no such a foramen in the same area. The hypoglossal foramen, which is located just behind the jugular foramen, is smaller than the jugular foramen. Mandible : The mandible is more similar to that of leporids than that of Ochotona. The hori2 zontal ramus is higher posteriorly than anteriorly on the labial surface, its greatest depth is at the level of m3. There is a distinct groove along the anterior surface of the ascending ramus, and the coronoid process is a laminal bone derived from the outer flange of the groove, and curves slightly inwards. On the lingual surface of the horizontal ramus the incisor2socket forms a massive curved swelling. This swelling becomes apparent posterior to the symphysis and ends below m2. The di2 astema is relatively long, and bends upwards. Two distinct mental foramina are present, one below p3 and the other is lower and below m1. Between the two mental foramina there are numerous small2 er subsidiary foramina. The condyle protrudes laterally at the anterior part and tapers posteriorly. The angular process is relatively long. Phylogenetic analysis According to Erbajeva (1994) the family Ochotonidae consists of two subfamilies: Sinolagomyinae and Ochotoninae. Alloptox was assigned to Ochotoninae, which is probably derived from a primitive species of Bellatona. In order to test her hypothesis, I conducted a preliminary phylogenetic analysis based on cranial characters (Table 3 4). Table 3 List of the transformation series 1) Skull roof flat (0) arched (1) 2) Nasal widens posteriorly (0) equal wide anteroposteriorly (1) narrow posteriorly (2) 3) Lateral margin of the nasal extends further caudad than any of the rest of the bone and ends in a sharp point (0) posterior end of the nasal gently rounded and its most posterior point is medially positioned (1) 4) Frontal wide anteroirly (0) equally wide anteroposteriorly (1) wide posteriorly (2) contract in the middle of the interorbit re2 gion(3) 5) Supraorbital process present (0) absent (1) 6) Supraorbital ridge absent (0) present (1) 7) One infraorbital foramen (0) two infraorbital foramina, of which the lower one is slightly in advance of the upper one (1) two infraobital foramina, of which the small one is just in front of the large one (2) 8) On lateral surface of maxilla a large triangular opening is absent (0) present (1) 9) The triangular opening on the lateral surface of the maxilla is an acute triangle, the smallest angle points forwards (1) acute tri2 angle, the smallest angle points backwards (2) right triangle, the smallest angle points forwards (3) right triangle, the smallest angle points backwards (4) 10) A lacework of fenestrated bone below the large triangular opening absent (0) present (1) 11) The maxillary tuberosity is isolated from zygoma (0) part of the tuberosity free of the zygoma (1) the tuberosity contacts the zygoma completely (2) 12) The ventral process of the zygoma scarcely developed (0) well developed (1) 13) On the anterior part of the lateral surface of the zygoma a circular pit present (0) absent (1) 14) The posterior extension of zagoma short (0) long (1) 15) The optic foramina confluent with each other (0) remain separate (1) 16) An apparent crest along the parietal2supraoccipital suture absent (0) present (1) 17) A deep depression in the posterior area of parietal and a sagittal ridge absent (0) present (1) 18) Interparietal bone present (0) absent (1) 19) The supraoccipital well developed (0) scarcely developed (1) 20) Nuchal crest and external occipital crest well developed (0) scarcely developed (1) 21) The incisive foramen confluent with the palatal foramen (0) separated from palatal foramen (1) 22) The maxillary component is reduced in palatal bridge (0) the palatine component is reduced in palatal bridge (1) 23) In palatal bridge the maxilla2palatine suture appears like a shallow W shape (0) straight (1) 24) Premolar foramen absent (0) present (1) 25) External carotid foramen present (0) absent (1) 26) Jugular foramen is a small rounded one (1) a long fissure (2) 27) The auditory meatus long (0) short (1) 28) The labial surface of the mandibular horizontal ramus is higher posteriorly than anteriorly (0) highest at the level of p3 (1) 29) A distinct groove on the anterior border of the ascending ramus present (0) absent (1)

15 2 : Alloptox gobiensis (, ) ) The coronoid process is a lamina bone derived from the outer flange of the anterior surface of the ascending ramus, and curves slightly inwards (0) the coronoid process is a low, thickened tuber, situated on the midway of the anterior surface of the ascending ra2 mus (1) I code these characters for Alloptox, Kenyalagomys, Bellatona, Prolagus and Ochotona. Palaeolagus is defined as outgroup on the basis of descriptions given by Wood (1940) and Dawson (1958). The character state polarity is determined by the method of outgroup comparison. Code? indicates the characters that were not preserved. The two character states of character 1 and charac2 ter 21 both exist in Ochotona. I also code those two characters as? for Ochotona. The data2ma2 trix has been conducted by implementation of Exhaustive search ( all multistate characters un2 ordered) using PAUP beta. One most parsimonious tree is produced, with a tree length of 44 steps and a consistency index of Table 4 Data Matrix Palaeolagus ?0000 Kenyalagomys 011? ??0000 Bellatona ?11?0????? 0011???100 Alloptox Prolagus Ochotona? ? The result of the phylogenetic analysis demonstrates that Alloptox and Kenyalagomys form a monophyletic group. Bellatona, Prolagus and Ochotona form another monophyletic group. The cur2 rent analysis supports Erbajeva s (1994) view that the family Ochotonidae consists of two subfami2 lies. But Alloptox is closer to Kenyalagomys than to Bellatona, and should be assigned to subfamily Sinolagomyinae; Bellatona and Prolagus should belong to subfamily Ochotoninae, which differs from Erbajeva s ( 1994) classification. This revised analysis excludes that Alloptox derived from Bellatona. Alloptox is probably derived from a primitive species of the subfamily Sinolagomyinae. As mentioned above, the Asiatic form Alloptox is closely related with the North African form Kenyalagomys. This indicates a fauna exchange between both continents. The genus Alloptox consists of six species and shows a wide Asian distribution from Turkey to Eastern China. The genus Kenyalagomys consists of three species and distributes in North and East2 ern Africa. The Early Miocene (MN4 or earlier) forms, A. xichuanensis (Liu and Zheng, 1997) and A. sihongensis (Wu, 1995), are the earliest known fossil records of Alloptox. K. rusingae and K. minor (MacInnes, 1953) from Early Miocene (MN32MN4) of Eastern Africa are the oldest known species of Kenyalagomys. The Asiatic form Alloptox and the African form Kenyalagomys ap2 peared almost at the same geological age. Muizon ( 1977 ) and Tong ( 1989 ) conclude that Kenyalagomys was derived from Sinolagomys and had an Asian origin in consideration of the simi2 larity in dental morphology. Wu (1995) considered that Alloptox was probably derived from Sino2 lagomys ulunguernsis ( Tong, 1989) or a species close to it based on dental similarity. Therefore, Alloptox and Kenyalagomys are probably derived from a common ancestor, Sinolagomys or a form closely related to Sinolagomys. It seems that the ancestor of Kenyalagomys originated in Asia and migrated to Africa in the early Neogene. The collision of the Afro2Arabian plate with Eurasia during Agenian2Orleanian established an intermittent land connection between Arabia and Southwest Asia. That allowed successive migrations of African mammals into Asia and Europe during the early Miocene. This introduced the African proboscideans and primates to Eurasia (Bernor et al., 1988 ; R ogl, 1998 ; Harrison and Gu, 1999). The ancestor of Kenyalagomys probably immigrated to Africa in this migration event. References Bernor R L, Flynn L J, Harrison T et al., Dionysopithecus from southern Pakistan and the biochronology and biogeography of

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