Researchgate.net. Research, DOI: 10.13140/RG.2.2.20079.74403.
Distinguishing features of manus and pes footprints and tracks of titanosaurian sauropods from South Asia (Pakistan and India) M. Sadiq Malkani1,2 1
Formerly with Geological Survey of Pakistan,C/O Post Office Retra, Tehsil Taunsa, District D.G. Khan, Punjab, Pakistan.
[email protected] February 2018 Abstract
The footprints and trackways from the Mesozoic strata of Pakistan revealed herd movements, and unique shape of manus and pes of Middle Jurassic titanosauriforms (basal titanosaurs), and Latest Cretaceous (Latest Maastrichtian) derived and most advanced titanosaurs. The footprints and tracks of titanosauriforms or basal titanosaurs from Pakistan reveal herd behaviour, confrontation scenario with a solitary large theropod, and new shape of asymmetric triangle-D-oval shape of manus (60cm*70cm) and associated giant circular pes (1.5m*1.4m) with three very broad and very long oval shaped toes (about 35cm*25cm) on digit II,III and IV, while digit 1st and 5th are reduced without toe. The footprints and tracks of Latest Cretaceous titanosaurs from Pakistan reveal herd behaviour, gliding/slipping of trackmaker on slippery muds, and also new shape of asymmetric triangle-D-oval shape of manus (50-55cm*70-75cm) and associated giant circular pes (1.27m*1.28m) with three very broad and very long oval shaped toes (its anterior and posterior ends are pointingly and broad) and also bac digit II,III and IV, while digit 1 st and 5th are reduced without toe and w shape heel. A pes footprint from Late Cretaceous of India reveal small sized subcircular pes (22.5cm*16.5cm) with three broad and oval shaped toes on digit II,III and IV, while digit 1 st and 5th are reduced without toe and rounded heel. From Pakistan and also India many fossil bones of metacarpals, metatarsals and toes. There are two types of toes / unguals are collected from IndoPakistan. The first type is narrow and long (MSM-152-3), and second type is special and unique which is very long and very broad. Previously these special and unique toes were assigned as large osteodermal ellipsoid oval plate such as MSM-84-7 (now Balochisaurus toe) and MSM-85-4 (now Pakisaurus toe). Like this many toes /unguals (long oval shaped) are also reported from India (AMNH1959 Huene and Mately 1033; Demic and Wilson 2009), Malawai (Mal 204; Gomani 2005) and Argentina (Powell 2003, p153), etc. The length, width and also depth are fit to toes shape as well preserved by many footprints of titanosaurs from Pakistan and India. This hypothesis is confirmed from one ichnite from India and two ichnite from Pakistan and also bone fossil toes from many countries like India, Pakistan, Malawai and Argentina belongs to many continents. In this way three types of titanosaurs are recognised from Indo-Pakistan on the basis of foot morphology. First type includes the pes with broad three toes and w-shape heel, Second type includes the pes with broad three toes and rounded heel, and Third type includes the narrow (may be four) toes and rounded heel. 1
Introduction Since long India was the only source of Mesozoic vertebrates while Pakistan presented first time Mesozoic vertebrates and first ever dinosaurs from Balochistan since 2000. From India only a single pes footprint (Plate 31) of titanosaurian sauropod was reported from Late Cretaceous Infratrappean limestone of Jetholi Talao area near Balasinor in Kheda district, Gujarat, India in 1986 (Mohabey 1986). While from Pakistan footprints and trackways of Malakhelisaurus mianwali Malakni 2007a titanosauriforms (Plate 1,2) sauropods were reported from middle Jurassic Samanasuk limestone since 2007 (Malkani 2007a,2008a,2015a,2017c,2018a) and latest Cretaceous Pashtosaurus zhobi Malakni 2014a titanosaurs (Plate 4-30) from latest Cretaceous Vitakri Formation since 2014 (Malkani 2014a,2015a,g,h;2017a,b,c,2018a; Malkani and Sun 2016). Further from India only two isolated footprints of theropods were reported from the Lower Jurassic strata of Thaiat ridge, Jaisalmer Basin, Rajastan, India (Pienkowski et al. 2015). While from Pakistan trackways of Samanadrinda surghari Malkani 2007a large theropod (confronted by a herd of titanosauriforms; Malkani 2007a,2008a,2015a,g,h,2017c,2018a) and a couple of Himalayadrinda potwari Malkani 2015g small theropod (Malkani 2007a,2008a,2015g,h,2017c,2018a) were reported from middle Jurassic Samanasuk limestones of Pakistan. A pterosaur (Anmolpakhi alleni) tracks were also discovered from latest Cretaceous of Pakistan (Malkani 2017a,b,2018a). Further some micro biota’s footprints were also discovered from the latest Cretaceous Vitakri formation of Sulaiman basin Pakistan (Malkani 2017a,b,2018a). Further trackway of Middle Jurassic large theropod show large theropod confronted and moved as solitary while the small theropod moved as herd or at least a couple (Malkani 2007a,2008a). Proof of footprints and tracks as biological, not man-made or not abiotic or not sedimentary structures (Malkani 2017a,b,2018a), and hypothesis of assignment of large oval ellipsoid plate (previous osteoderms) to pes unguals/toes of titanosaurs with new shape of manus associated pes are reported (Malkani 2018a). Results and Discussion A pes footprint of titanosaurian sauropods from India From India a single tritoed manus footprint (22.5cm*16.5cm; Plate 31) of sauropod dinosaur was reported by Mohabey (1986) from uppermost limestone unit of the Lameta Formation (given as Infratrappean Sequence; Maastrichtian) of the Jetholi area near Balasinor in Kheda District in Gujarat, western India (Mohabey 1986), while Mateus et al. (2011), Xing et al. (2012) and Galton and Ayyasami (2017) interpreted it as a Deltapodus like (Thyreophoran type; stegosaurian affinity) track but Malkani (2017a,b) considered this Late Cretaceous footprint of Kheda District, Gujrat, India as pes print of juvenile titanosaurian sauropod dinosaur. Further pl see description below.
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Footprints and trackways of Malakhelisaurus mianwali Malkani 2007 titanosauriforms / early titanosaurian sauropod dinosaurs from Middle Jurassic of Pakistan The Malakhelisaurus mianwali Malkani 2007a ichnotype was reported from the Middle Jurassic upper successions of Samanasuk limestone, which show the slight regression of sea and the area was exposed coastal plain. Limestone dips 52o west and strike north 50o east. This ichnite consists of exposed more than 15 footprints, and 4 short trackways (Malkani 2007a,2008a). These footprints and trackways of titanosauriforms / early titanosaurian sauropods trend in a southeast direction and are parallel to each other, which are confronted obliquely at 63o theropod trackway (Plate 1) with north-northeast trend. This site characterized the largest known pes trackmaker (pes length 150cm, width 140cm: manus length 60cm, width 70cm). The manus/pes heteropody area ratio is about 1/5. The right side track of left and right side of single individual show 0.4m internal trackway width, revealing wide gauge locomotion of Malakhelisaurus mianwali titanosaurs. Further these footprints and trackways reveal a herd of wide gauge titanosaurian sauropods composed of 3 or more individuals defending the attack of a solitary predatory theropod. The footprints are well preserved and only show overprinting some in the back area. The exposed area is small. The excavation along dipward can clear further these trackways and interaction among herbivores and carnivores. The large pes prints belong to titanosaurs are diagnosed on the basis of the associated asymmetrical triangular-D-oval shape manus impression represents that metacarpals were arranged in triangualrD-oval shape, and there is no any indication of manus claw (Plate 1,2). The Malakhel Baroch site adds significantly to our understanding of locomotors styles, potential trackmaker, echnology, behavior and evolutionary history of early titanosaurs and social interaction of saurischian and theropod dinosaurs (Malkani, 2008a). The pes toes of middle Jurassic ichnite are well preserved (Plate 1,2) due to well suited substrate while depth of footprints is less that Latest Cretaceous ichnite. But unluckily some part of these footprints and trackways are ruined by the mine owners for making road. I hope scientific and administrative authority will help to save the latest Cretaceous Sor Muzghai Zhob site. Mannion et al. (2013:149) discussed these titanosaurian sauropod trackways and concluded (according Lockley 1991 and P.L. Falkingham pers.comm in 2012 with Mannion) as three toed ornithopods. But the giant size of pes associated with asymmetric triangual-D-oval shape manus without claw which can only be assigned to early titanosaurian/ titanosauriforms sauropods.
Footprints and tracks of titanosaur sauropods from the Latest Cretaceous Sor Muzghai ichnite of Pakistan Titanosauria is diverse sauropod clades that include about 65 species of worldwide distribution. However, their footprints are scarce. In Sor Muzghai paleoichnological outcrop, the author discovered five levels of footprints and tracks in the upper part of Vitakri Formation. The uppermost level consists of ichnotype of Pashtosaurus zhobi and other 3 levels show its reference manus tracks. The strike of host sandstone bed is North 65o East and its dip is 70o Northwest. The exposed area is small (about 10*6=60m2) and excavation in depth inclination may reveal best results. These ichnites 3
consist of exposed more than 15 footprints. These footprints of titanosaurian sauropods mostly trend in a north direction and are parallel to each other, however there are some signs of transverse moved tracks. These are indicative of a titanosaurian sauropods herd composed of 3 or more individuals. Here two coherent trackways of two animals and a few other tracks are reported from ichnotype of Pashtosaurus zhobi. Malkani (2014a) estimated the width of pes footprint about 0.7-0.8 meter and the length is about 0.7-0.9 meter, but the recent investigation (Malkani 2017a,b,2018a) has shown measured maximum length 1.27m and width 1.28m of pes. The pes has well developed outer rim which show suitable moisture content and cohesion of the substrate. The manus are asymmetrical triangle-D-oval shape varies in length 50-55cm and width 70-75cm (Plate 7-30, 34). The asymmetric manus has five digits (Plate 7-30). The depth of footprints vary from 10-20cm. Pes is tritoed or three toed. Its length is 1.27m and width 1.28m. The heel is w shape (Plate 5, 7-25). The central digit with toe is longest and measured as 1.12m. The lateral digit with toe is slightly varied in length like 0.9m and 0.8m. The posterior concavity is 15cm in length from posterior alignment of convexities and less than 60cm in width between the convexities. While posterior convexities are 15cm from anterior margin of concavity alignment and both convexities is 0.6m apart from each other. Both convexities may be developed by the ventral surfaces of astragalus and calcanium, while dorsal surface of both are articulated with distal tibia and distal fibula respectively. The central toe is subcircular and oval and 25cm in length and 20 cm in width. One manus is found 0.5m forward from pes while another manus is found forward 2m from same pes. The pace is more than 2m measured from anterior of pes to anterior of manus and pes stride expected is about 4.5m and manus stride expected is about 4m. The internal trackway width is 0.4m (Malkani 2017a,b). The pes may be semi plantigrade /semidigitigrade with presence of fatty foot pad (Miller et al. 2007; Riga 2011). The morphological outline of manus without claw and pes with three toes of Latest Cretaceous ichnite is well preserved than middle Jurassic ichnite. Size estimation of track maker Pashtosaurus zhobi Alexander’s (1976) observations on the gaits of extant vertebrates were extended to define three different gaits in dinosaurs with reference to stride length S and hip height H: walk (S/H < 2.0), trot (S/H between 2.0 and 2.9), and run (S/H > 2.9). Alexander (1976) was among the first to offer a simple approach to estimating the size of a sauropod trackmaker. Alexander (1976), following the skeletal proportions of the diplodocid sauropod Apatosaurus, considered that H is four times the length of the pes footprint (L). He estimated similar values (e.g., L = 0.23 − 0.38H) for many bipedal dinosaurs of a wide range of sizes, both theropods and ornithopods. He derived the formula H=4L, in which H is the height of the acetabulum and L is the length of the pes track (Alexander, 1976). For Pashtosaurus zhobi the length of pes is 1.27m. Applying the formula H=4L of Alexander (1976) to the Pashtosaurus zhobi titanosaurian sauropod track, the acetabulum height of the trackmaker is estimated to be 1.27*4=5.08m.
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Thulborn (1990) claimed that the assumption of H = 4L is likely to be incorrect for two reasons. First, the H/L ratio varies in systematic fashion among dinosaur taxa. Second, the H/L ratio certainly changes during ontogeny, on account of the allometric growth that prevails in terrestrial vertebrates. For sauropods, Thulborn (1990) has suggested that H = 5.9L. Applying the formula H=5.9L of Thulborn (1990) to the Pashtosaurus zhobi titanosaurian sauropod track, the acetabulum height of the trackmaker is estimated to be 1.27*5.9=7.493m. Riga (2011) supposed that juveniles have relatively larger feet (and therefore smaller H/L ratio) than do adults of the same taxon. Based on an analysis of the titanosaur MUCPv-1533, González Riga (2011) offered a revised formula of H= 4.586L. Applying the formula H=4.586L of Riga (2011) to the Pashtosaurus zhobi titanosaurian sauropod track, the acetabulum height of the trackmaker is estimated to be 1.27*4.586=5.82422m. For sauropods, Lockley (1986) used H = 4W where W is the foot width. Here the width of pes of Pashtosaurus zhobi is 1.28m. Applying the formula H=4W of Lockley (1986) to the Pashtosaurus zhobi titanosaurian sauropod track, the acetabulum height of the trackmaker is estimated to be 1.28*4=5.12m. In this way the acetabulum (e.g., the hip joint) of Pashtosaurus zhobi is located as average 5.879305m above the ground. Considering the stocky nature of trackmaker, the acetabulum height of the trackmaker Pashtosaurus zhobi is estimated to be in the range of 3-5m. The length and width of pes show that the trackmaker were about 3-5m upto hip joint i.e., hip height is about 3-5m. The ichnological records of trackmaker Pashtosaurus zhobi matches closely with skeletal records of giant balochisaurids Marisaurus jeffi or Balochisaurus malkani. Distinguishing features of manus and pes footprints and tracks of titanosaurian sauropods from South Asia (Pakistan and India) These footprints and tracks do not belong to any sedimentary facies and not fit into any sedimentary structures like ripple marks, flute cast, groove marks, burrows, load casts, bulges, cross bedding, planner bedding, chemical and physical weathering marks, scour marks, etc (detail discussed in Malkani 2017a,b,2018a). The continuity of some footprints under the overlying bed (Plate 7-25, 28-29) shows that these are not recent man-made structures but are original in situ marks which developed before the deposition of overlying bed (detail discussed in Malkani 2017a,b,2018a). The author discovered multiple track-bearing levels in the Latest Cretaceous Vitakri Formation of Sulaiman basin, Qila Saifullah District, Zhob Division, Balochistan Province, Pakistan showing their originality. Further detail proof of footprints and tracks are also discussed in Malkani (2017a,b,2018a),
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These footprints from India and also Pakistan belongs to titanosaurs and not Deltapodus or any ornithischian. The known stratigraphic range of Deltapodus is at present restricted to the Middle to Late Jurassic (Aalenian to Tithonian) although the skeletal record of stegosaurs extends into the Lower Cretaceous (Maidment et al. 2008). Considerable doubt (Chatterjee and Rudra 1996; see also Maidment et al. 2008) has been cast on records of stegosaurs in the Late Cretaceous of India (Yadagiri and Ayasami 1979; Galton and Upchurch 2004). Deltapodus brodricki Whyte & Romano 1995 based on a supposed sauropod track from the Middle Jurassic of England, identified three toed stegosaur like Stegosaurus by Whyte and Romano (2001). The Indian pes print of Deltapodus sp. (Plate 42) agrees with those of Deltapodus brodricki in several characters, viz., it is triangular with a well-developed heel, digitigrade, three very wide blunt digits, toes radiating and not separated by well developed hypicies (angles), and weakly mesaxonic (digit III only slightly longer than II and IV) (Whyte & Romano 2001; Li et al. 2012). Deltapodus also occurs in the Lower Cretaceous of Spain (Cobos et al. 2010; Pascual et al. 2012), western USA (Milân et al. 2015) and China (Xing et al. 2013). The present author (Malkani) considered this Late Cretaceous tritoed footprint of Kheda District, Gujrat, India as pes print of juvenile titanosaurian sauropod dinosaur. This interpretation is based and confirmed by the circular giant pes with broad oval tritoed pes tracks associated with asymmetric triangle-D-oval shape manus of latest Cretaceous titanosaurs and middle Jurassic titanosauriformes found from Pakistan. The large tritoed pes has broad toes on 2nd, 3rd and 4th digits, while 1st and 5th digit are reduced and without toes. The asymmetric triangle-D-oval shape manus without claw is the autopomorphy of titanosaurs only. So when giant tritoed pes associated with typical (characterized for titanosaurs only) manus show titanosaur assignment only. This association is confirmed on many ichnites from Pakistan like middle Jurassic Malakhel ichnites and also latest Cretaceous Sor Muzghai sites. Further the giant pes with 1.27m length and 1.28m width and associated manus with 50-55cm length and 7075width can only be assigned to titanosaurian sauropod because most of the Deltapodus footprints are less than or close to half meter in length and also half meter in width (Mateus et al. 2011) and have no asymmetric D shape manus without claw (Mateus et al. 2011). Further the Deltapodus / stegosaurs are found upto Late Jurassic and not found in Late Cretaceous. The Latest Cretaceous was the kingdom age of titanosaurs in Indo-Pak and Gondwana but also in the world. As Cobos et al. (2012) the characterization of Deltapodus are kidney shaped manus and manus prints immediately in front of the pes print, so in the case of Indo-Pak the asymmetric triangle-D shape manus and away separation of associated manus again show titanosaurian affinity. The length of pes of Deltapodus is half of the manus of Pakistani titanosaurs, and same is true for the pes. Further the fossils found so far from Indo-Pak belongs to titanosaurs. No any confirmed fossils of stegosaurs or any ornithopods are not found from Indo-Pak subcontinent. Some osteoderms assigned to ornithischian may belong to pes toe of titanosaurs. Sauropod pes tracks typically are sub-circular or elliptical in shape, with well-preserved examples showing evidence of up to five anterolaterally directed digits and a large, rounded heel 6
(Farlow et al., 1989; Thulborn, 1990; Farlow, 1992). In Sor Muzghai case the pes is well preserved showing central 3 digits with broad toes and 1 st and 5th digits are relatively short but without toes. The heal is generally rounded but w shape. In Sor Muzghai case, here three pes (Plate 10-28) are preserved (due to small exposed area) in the first three tracks. In 1st track pes, the two toes marks of right pes are well preserved. In the 2 nd track a complete left pes is preserved with three toes and w shape heel. In this way they form a coherent trackway. These phenomenon show original footprints on regular pattern. In this way the w shape heel of 2nd track full pes is matches with the heel of pes of 3rd track (Plate 10-28 ). These phenomenons again show original footprints on regular pattern. The asymmetric triangle-D-oval shape of manus is also consistent with the titanosaurs footprints in above mentioned tracks. Sauropod manus tracks are smaller than associated pes tracks, and typically appear as posteriorly concave, crescent shapes with no distinct digit impressions except for rare pollex claw marks (Thulborn, 1990; Farlow, 1992). In our Sor Muzghai footprints and tracks case, the manus is asymmetric triangle-D-oval shape but without any claw. The pes has 3 robust digits with broad toes remaining two digits are reduced and have no toes/claws. The heel of pes is w shape. The manus without claw of titanosaurs were confirmed by many footprints and revealed by distal metacarpals having no articular distal surface reported from Pakistan. Other important identifying features that are discernable where whole trackways are preserved include placement of the manus relative to the pes (Farlow, 1992), trackway gauge (the lateral position of the tracks relative to the trackway midline; Farlow et al., 1989; Farlow, 1992; Lockley et al., 1994a), and heteropody (manus-pes area ratio; Lockley et al., 1994a). Regarding the placement of pes and associated manus, here two different patterns are observed and reported. In Latest Cretaceous Sor Muzghai case, the manus and pes are alternately placed generally with equal distances, while the Middle Jurassic Malakhel site show relatively close placement of pes and manus. These trackways reveal less wide gauge movements and about same distance away from midline both in middle Jurassic Malakhel and latest Cretaceous Sor Muzghai tracks. Regarding heteropody (manus-pes ratio) is about 1:5-7 same for both middle Jurassic and Latest Cretaceous sites. The phalangeal and footpads also have been described from sauropod pes prints (Gallup, 1989; Romano et al., 1999), but these do not show the distinct nodes present in some tracks attributed to other saurischian dinosaurs (e.g., Hitchcock, 1858; Rainforth, 2003). In our case the relevant is titanosaurian sauropods saurischian dinosaur’s footprints and tracks which reveal three toes in pes. The possible phalangeal pad formula 1-2-2-2-0 including phalangeal toe formula 0-1-11-0. The central 3 digits are strong and broad while left and right digits are reduced and also without toe/ungual.
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The Pakistani pes print of titanosaur characterized general rounded circular shape with w shape heel, semi digitigrade/semi plantigrade, spreading digits, broad toes on II,III,IV digits, while Digit I and V are reduced and without toe, central toe and digit is long and forward inserted. Instead stegosaur include triangular with a well-developed heel, digitigrade, three very wide blunt digits, toes radiating and not separated by well developed hypicies (angles), and weakly mesaxonic (digit III only slightly longer than II and IV). Deltapodus occurs in Spain, western USA and China only. In our case two coherent trackways and remaining tracks are reported. Most of the tracks are well preserved and appear as the shape of manus without claw and pes with their three toes with good external foot morphology of the trackmaker. The presence of dinosaur toe mark like three toes on pes footprints, and manus without claw show its typical recognition of titanosaurian sauropod dinosaurs. The Pashtosaurus zhobi Malkani 2014 titanosaurian sauropod dinosaurs based on ichnotype of Sor Muzghai footprints and tracks characterized by asymmetric triangle-D-oval shape manus without any claw, pes in general circular shape but also robust, pes digits spreading, pes with robust and spreading tritoed print (prominent and forward inserted central toe, straight forward insertion created posterior central concavity while lateral digits are backward inserted (with reference to central main digit) creating posterior convexities in their relevant axis (convexities may be due to orientation of astragalus and calcanium), diverging anteriorly in this way forming w shape in posterior view, 1st and 5th digits are lost or reduced without toes), latest Cretaceous age and finding of bones in the same age, same formation and same basin. All these evidences indicate that the trackmaker were derived titanosaurs. The distal end of metacarpals of Balochisaurids and also Pakisaurids have no articular surfaces showing absence of phalanges and claws. The manus tracks and footprints also show no claw. In this way skeletal and ichnological data of Indo-Pak supported the titanosaurs. The oval shaped ellipsoid which are previously assigned to Balochisaurus and Pakisaurus. Now these oval shaped ellipsoid are completely (in size, shape and morphology) fit into the large and oval toe of pes prints of middle Jurassic titanosauriforms / early titanosaurs and also advanced derived titanosaurs. Further from India such plate assigned to ornithischian (Huene 1923; Huene and Matley 1933) and that is also fit in to toe. Previously most of titanosaur footprints have been indentation but the present track have well preserved shape of entire asymmetric triangle-D-oval shape manus without claw and circular giant pes with three robust and wide toe on digit II,III,IV and the remaining two digit I and V are reduced and without toe. In general tacks with digit imprints are rare, so footprints from Pakistan are significant. Tracks provide clues about the scrolling of titanosaurian sauropod dinosaurs along sandy muddy path near the shores of sea. In Sulaiman basin, the fossils and tracks of titanosaurs are found among the world latest or latest or youngest dinosaurs. There are expulsion rims around the footprints suggesting the substrate were suitable for footprints development. From here we can measure the footprint length and width, footprint rotation, pace angulations, pace length, stride 8
length, inner and outer trackway width, etc. New sauropod footprints show less wide gauge locomotion and anatomical features that relates the footprints with adult titanosaurian sauropods (Balochisaurus) as the best candidates to be the trackmaker. The less wide gauge conditions, manus pes distance, asymmetrical triangle-D shape to oval morphology of manus, symmetrical tritoed (with median toe large inserted forward) almost w shape heel of pes, large length and more broader width suggest the trackmaker are stocky titanosaurian sauropods. In this way Pashtosaurus zhobi ichnotype matches with balochisaurs (Balochisaurus). Pashtosaurus zhobi is an excellent case study of less wide gauge style of locomotion produced by Latest Maastrichtian derived titanosaurs that have no impressions of manual phalanges and claws. These feature and the fossil records from the Vitakri Formation, suggest that the trackmaker were probably small to medium sized and stocky bodied balochisaurs like Balochisaurus malkani. Sor Muzghai ichnite is youngest titanosaurian sauropod track recorded in the world and just below the Cretaceous-Tertiary (K-T) extinction event. The acetabular height is approximate 3-5m estimated from length and width of pes footprint. The pes is well preserved revealing new and useful information. Pes is spreading. Three main digits of pes are not equidimensional, and the central digit with toe is prominent, largest, longest and inserted forward in movement direction, forming backward central concavity, while its adjoining right and left lateral digits are inserted backward/posteriorly forming convex shape in their axis (Malkani 2017a,b,2018a). The central ungual or claw is prominent, large, subcircular. The shape of pes in posterior view is ‘W’ shape. In the left and right lateral most side of pes print showing straight left and right side may be reduced 1st and 5th digits but seems without toe (Plate 7-25). This discovery is significant due to revealing of new shape of manus and pes of titanosaurs, less wide gauge locomotion style, herd movements and gliding/slipping of legs, etc, and occurrence of titanosaurs body fossils in coeval strata and also in the same middle Indus/Sulaiman basin. From tracks we learn how the large and massive animal walked on earth and glided or slipped on slippery muds. A few manus footprints show glide (slipped) mark due to slippery muddy surface of thick sandstone bed (Plate 7-25). Four or five manus footprints show gliding (slipped) mark of trackmaker Pashtosaurus zhobi due to slippery muddy surface of thick sandstone bed. The gliding print of left and right manus is represented by two parallel pair of print. One pair (left and right manus prints) show gliding in the forward direction, when ended this gliding prints, other pair (left and right manus prints) print started in transverse directions or about perpendicular to first pair gliding movements. One another manus slipped and meet one of the parallel slipped lines. The manus/pes heteropody area ratio is about 1/5 - 1/7 of titanosaurian sauropods found from Pakistan and as compared with ratios ½ of middle Jurassic sauropods (Galinha, Portugal), 1/3 of titanosaur from Argentina (Riga 2011), 1/3 of Cretaceous Brontopodus birdi (Texas), and ¼ to 1/5 of Late Jurassic sauropods (Colorado) tracks (Santos et al. 1994). The apparent absence of pes prints from many known sauropods trackways may be due to overlapping (Day, et al., 2004) but here the pes along with toes are best preserved (Plate 7-25).
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The Mesozoic vertebrates are described and reported by Malkani (2003a,b,c; 2004; 2006a,b,c; 2007a,b,c; 2008a,b,c,d; 2009; 2010a,b,c; 2011; 2012a,b,c,d; 2013a,b,c; 2014a,b,c,d,e; 2015a,b,c,d,e,f,g,h,i; 2016a,b,c; 2017a,b,c,d,e,f,g,h,i), Malkani et al. 2001, Wilson et al. (2001,2005), Malkani and Sun (2016). The Pakititanosaurs are classified into 3 clans like Pakisaurids (relatively large-sized and slender bodied titanosaurs) which include Pakisaurus, Sulaimanisaurus and Khetranisaurus; Marisaurs (medium-sized and stocky bodied titanosaurs) which include Marisaurus, Maojandino and Gspsaurus; and Balochisaurs (small-sized and stocky bodied titanosaurs) which include Balochisaurus, Nicksaurus and Saraikimasoom. The footprints of middle Jurassic Malakhel (Broach) site titanosauriforms/ titanosaurs Malakhelisaurus closely resemble or match with Pakisaurus (or any member of Pakisaurids or its ancestors while the latest Cretaceous Sor Muzghai site titanosaurian Pashtosaurus zhobi closely resemble with Balochisaurs especially with Balochisaurus. Both these sites show herd behavior for defending against predatory animals. The predatory animals from latest Cretaceous of Sulaiman basin include theropods (Vitakridrinda and Vitakrisaurus), mesoeucrocodiles (Pabwehshi, Sulaimanisuchus and Induszalim) and pterosaur (Saraikisaurus). The herbivorous Pakititanosaurs used herd behaviors and osteodermal armor spines and plates for defending these predatory animals. The calculated acetabulum (e.g., the hip joint) height of Pashtosaurus zhobi is average 5.879305m (Malkani 2018a). Considering the stocky nature of trackmaker, the acetabulum height of the trackmaker Pashtosaurus zhobi is estimated to be in the range of 3-5m. The ichnological records of trackmaker Pashtosaurus zhobi matches closely with skeletal records of giant balochisaurids Marisaurus jeffi or Balochisaurus malkani.
Hypothesis regarding the assignment of large ellipsoidal oval plate (previous osteoderms) to pes toes of titanosaurs, shape and anatomy of manus and pes of titanosauriforms or early titanosaurs and derived titanosaurian sauropods Malkani (2017a,b,2018a) presented asymmetric triangle-D shape to asymmetric oval shape of manus and associated pes with very broad and long three toes on II,III, IV while digit I and V is reduced and without toes and w shape heel. Malkani (2018a) presented the hypothesis that the broad and long osteodermal oval plates belong to toes of titanosaurs. The toes prints of Middle Jurassic titanosauriformes / basal titanosaurs ichnite have length 35cm and width 25cm. In this way the bone fossil oval plates also reported from different continents variesgenerally in length from 19-32, width from 13-23 and depth from 10-13. So the length, width and also morphological shape of toes (Plate 32,33) with skin are exactly fit and matches into toes prints (Plate 1.2,7-25). From Pakistan and also India many fossil bones of metacarpals, metatarsals and toes. There are two types of toes / unguals are collected from Indo-Pakistan. The first type is narrow and long (MSM-152-3), and second type is special and unique which is very long and very broad. Previously 10
these special and unique toes were assigned as large osteodermal ellipsoid oval plate such as MSM84-7 (now Balochisaurus toe) and MSM-85-4 (now Pakisaurus toe). Like this many toes /unguals are also reported from India (AMNH1959 Huene and Mately 1033; Demic and Wilson 2009), Malawai (Mal 204; Gomani 2005) and Argentina (Powell 2003, p153), etc. The length, width and also depth of these fossils are fit to toes shape as well preserved by many footprints of titanosaurs from Pakistan and India. This hypothesis is confirmed from one ichnite from India and two ichnite from Pakistan and also bone fossil toes from many countries like India, Pakistan, Malawai and Argentina belongs to many continents. The footprints and tracks of Latest Cretaceous titanosaurs from Pakistan reveal new shape of asymmetric triangle-D-oval shape of manus (50-55m*70-75m) and associated giant circular pes (1.27m*1.28m) with three broad toes on digit II,III and IV, while digit 1 st and 5th are reduced without toe and w shape heel. This hypothesis is also confirmed from Middle Jurassic titanosauriforms/ early titanosaur trackways from Pakistan and Late Cretaceous sauropod track from India. This hypothesis was presented by Malkani (2007a, 2008a) and this hypothesis is shifted into confirmed theory by Malkani (2014a,2015a,g,h, 2017a,b,c) with new discoveries of latest Cretaceous footprints and tracks of titanosaurs found from same basin and same formation of Sulaiman basin which also yielded rich bones of titanosaurs. The toe or ungual are large oval shape. Now again problem arises to assign large oval plate because the ornithopods have half large and broad ungual while in our case is the full oval shape ungual in all three digits (digit II, digit III and digit IV). The other lateral digits are reduced or lost and have no toe or ungual. A relatively thin ungual is reported for Pakisaurus from Pakistan (Malkani 2010a) and also from Malawisaurus from Malawi (Gomani 2005), further both these genera include the oval osteoderms. Malkani (2010c,2015a,2017c) reported an oval osteoderms assigned to Balochisaurs. Powell (2003) also reported an oval plate osteoderms of titanosaur from Argentina. Cerda et al. (2015) reported many type of osteoderms of titanosaurs from Patagonia. Vidal et al. (2014) reported many types of osteoderms of titanosaurs from Spain. However the large oval ellipsoid plate (Plate 32,33) or some large bulb with root type osteoderms may belong to pes toes of titanosaurs. Like this many oval plates reported from Pakistan, India, Argentina, and also from Europe. So the present author (Malkani) propose the new hypothesis that these oval osteoderms (Plate 43, Row 1) belong to ungual of pes of titanosaurs based on the fit shape, size and morphology, and evidence from trackmaker of these type of pes tracks found from middle Jurassic and latest Cretaceous of Pakistan. The already reported relatively thin unguals are shifting to osteoderms and large oval osteodermal plates are being shifted from osteoderms to pes unguals (Plate 32, Row 1; Plate 33). This hypothesis arises by middle Jurassic tracks and verified by latest Cretaceous tracks, now it needs its confirmation from articulated pes skeleton. From India Huene and Matley (1933) reported a proximal left metacarpal II (K27/506) and a proximal very large right metacarpal V (K27/507) with diameters of are 13 and 8cm and proximal metatarsal I (K27/509) with diameter of proximal facet is 15.5 and 9 cm (Huene and Matley 1933). 11
An oval plate of ornithischian is also reported by Huene (1923) and Huene and Matley (1933), and present author proposed this plate as pes ungual. From Pakistan metacarpals and metatarsal (Plate 43) were assigned to Marisaurus (Malkani 2008a), Balochisaurus (Malkani 2009a) and Pakisaurus (Malkani 2010a). Metacarpals are very large. Metacarpals are very large, elongated, triangular and rugose articular proximal surfaces. Distal condyle shape is undivided and has no articular rugosities. A proximal and distal end is expanded with triangle shape (Malkani 2008a). Metatarsals represent irregular concave proximal and distal expanded surfaces (Malkani 2008a). The metatarsals may be arranged in semicircular fashion diverging outward and downward in contrast to the vertically oriented metacarpals as reveals by footprints of titanosaurs (Malkani, 2007a, 2008a, 2014a,2015a,2017a,b,c). According to McIntosh (1990) the metatarsal V is shorter with characteristically much larger proximal than distal ends. The first three digits bore claws of decreasing size but in Pakistani titanosaurs footprint these are nearly equal. It is possible there is no claw on digit I and V, may be deduced from the footprint of titanosaurs from Pakistan (Malkani, 2007a; 2008a). Only one possible relatively thin ungual was assigned to Pakisaurus (Malkani 2010) but now it is shifted to osteoderms. Malkani (2003) reported osteodermal plates including the two large oval shape osteoderms (Plate 43). The first oval plate (Plate 43, row 1) assigned to Balochisaurus is found from Nala Basti (Zubra) locality PDL 7 is sub circular plate which has slight concavity and rugosities on smooth ventral surface, and rugosities on lateral and dorsal surfaces. Dorsal surface has an asymmetrical low ridge/cone. The ossicles on the ridge area are tightly packed and directing toward the tip of ridge making radial pattern, having centre at tip; and remaining dorsal and lateral area has irregularly shaped ossicles and internal ducts. The ridge seems to be directed dorsoposterior. The length and width of ventral sub circular surface are 19 and 16 cm respectively; and dorsoventral depth of dorsal ridge from ventral surface to preserved apex is 12 cm. Apex/tip of cone/ridge is not preserved. The second oval plate (Plate 32, row 1; Plate 33) included in the type series of Pakisaurus is found from Kinwa locality. Its ventral surface is semi plain slightly convexing downward, with irregularly rugose ossicles and internal ducts pattern. The dorsal surface has well developed asymmetrical, low and sharp cone/ridge having possibly dorsoposterior directed tip. The ossicles on the ridge area are tightly packed and directing toward the tip of ridge making radial pattern, having centre at tip. In the centre of dorsal surface of plate has a transverse median concavity belt just in front of ridge. The remaining dorsal and lateral surfaces have irregular rugosities of ossicles and internal ducts. The length and width of ventral oval surface are about 21 and 13 cm respectively, and dorsoventral depth of dorsal ridge/cone is 13 cm i.e., from ventral surface to apex or tip of ridge/cone. The first and second oval plates have resemblance in shape but differ in nature of smooth, concave, convex, circular and oval ventral surfaces, and transverse median concavity belt on dorsal surface. The ventral surface of first type is slightly concave and smooth having corroded rugosities on circular ventral surface. The second type shows slight convexity and well developed, irregular rugosities on oval ventral surface, and it bears a transverse median concave belt on dorsal surface. Now the present author (Malkani) proposed that these both oval plates being assigned to pes 12
toes of titanosaurian sauropods because its shape, length, width, robustness and general outlines closely matches with the toes found in pes footprints found from Pakistan. Further Ostrom and McIntosh (1966:pl.78) mentioned osteoderms are generally similar in robustness, size, and shape to sauropod carpal and tarsal elements. The present author is encouraged by the above remarks of Ostrom and McIntosh (1966:pl.78) and assigned the oval plate (previously called oval osteodermal plate or ellipsoid) to toes or unguals of pes of titanosaurs. The general line sketch of pes is shown in Plate 8.
No distal surfaces of metacarpals marks are clear in many manus prints, however one manus footprint shao the marks of metacarpal I-V impressions (Plate 31-32, 45). In this way estimated sketch of orientation of five metacarpals are prepared (Plate 45). The pes has spreading digits (Plate 8, 10-28) like diplodocus but the toes are subcircular. The middle Jurassic and latest Cretaceous large and giant pes footprints associated with asymmetric D shape manus revealed the new theory. The middle Jurassic and latest Cretaceous large and giant pes footprints associated with asymmetric triangle-D-oval shape manus without claw show titanosaur affinity. Further these large and giant pes footprints have well preserved three large subcircular oval toes. These subcircular toe matches closely with plates (Plate 44) which were considered previously osteodermal plate revealed the new theory. Further the narrow toe (previously assigned to Pakisaurus; Plate 43, row 3) is may be assigned to another genus of titanosaurs. The possible phalangeal pad formula 1-2-2-2-0 including phalangeal toe formula 0-1-1-1-0. The central 3 digits are strong and broad while left and right digits are reduced and also without toe/ungual (Plate 8, 10-28). As conclusions there are three types of titanosaurian sauropods in Indo-Pakistan subcontinent based on ichnites and foot bone fossils. First type includes the pes with broad three toes and w-shape heel. Second type includes the pes with broad three toes and rounded heel Third type includes the narrow (may be four) toes and rounded heel.
Acknowledgements The author (MSM) is very thankful to the sponsor Factum Arte and lconem to preserve this geoasset. The author is thankful to Director General, directors and other officer and official of Pakistan Museum of Natural History (PMNH) for collaborating the project with Mr. Nicholas Allen and his foreign team members. The author is very thankful to the field team of PMNH namely Mr. Aamir Yaseen, Mr. Khalil and Mr. Mirani for their kind cooperation during field work. I am also thankful 13
to Administration of Zhob and Qila Saifullah districts for providing security and accommodation during field work. I am thankful to Mr. Jan Mohammad Khan and other local peoples for their kind cooperation during filed work. Further I am also thankful for the kind suggestions of Prof Dr. Jeffrey A. Wilson, Prof. Dr. Lockley, Prof. Dr. Peter Falkingham and Prof. Dr. Lida Xing. On the basis of these suggestions, the author has provided the proofs and evidence for these footprints and tracks as these are not non-biological structures, not sedimentary structures and not man-made structures. The author further provided the proof for the demonstration of these footprints and tracks and their assignment to titanosaurs.
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Malkani M.S. 2012b. A review of Coal and Water resources of Pakistan. Science, Technology and Development, 31(3), 202-218.
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Malkani M.S. 2012d. New Look of titanosaurs: Tail Special of Pakisauridae and Balochisauridae, Titanosauria from Pakistan. In: Huh M., Kim H.J., and Park J.Y., Eds. Abstract volume of 11 th Symposium on Mesozoic Terrestrial Ecosystems, August 15-18, Gwanju, Korea, 113-119.
Malkani M.S. 2011. Stratigraphy, Mineral Potential, Geological History and Paleobiogeography of Balochistan Province, Pakistan. Sindh University Research Journal (Science Series) 43 (2):269-290.
Malkani M.S. 2010a. New Pakisaurus (Pakisauridae, Titanosauria, Sauropoda) remains and Cretaceous Tertiary (K-T) boundary from Pakistan. Sindh University Research Journal (Science Series) 42 (1):39-64.
Malkani M.S. 2010b. Stratigraphy and Mineral potential of Sulaiman (Mid. Indus) basin, Pakistan. Sindh University Research Journal (Science Series) 42 (2):39-66.
Malkani M.S. 2010c. Osteoderms of Pakisauridae and Balochisauridae (Titanosauria, Sauropoda, Dinosauria) in Pakistan. Journal of Earth Science, Vol. 21, Special Issue 3, 198-203. doi: 1007/s12583-010-0212-z.
Malkani M.S. 2009. New Balochisaurus (Balochisauridae, Titanosauria, Sauropoda) and Vitakridrinda (Theropoda) remains from Pakistan. Sindh University Research Journal (Science Series) 41 (2):65-92.
Malkani M.S. 2008a. Marisaurus (Balochisauridae, Titanosauria) remains from the latest Cretaceous of Pakistan. Sindh Univ. Res. Journ. (Sci. Series) 40 (2):55-78.
Malkani M.S. 2008b. First articulated Atlas-axis complex of Titanosauria (Sauropoda, Dinosauria) uncovered from the latest Cretaceous Vitakri member (Dinosaur beds) of upper Pab Formation, Kinwa locality of Sulaiman Basin, Pakistan. Sindh Univ. Research Journal (Science Series) 40 (1):55-70.
Malkani M.S. 2008c. Mesozoic Continental Vertebrate Community from Pakistan-An overview. Journal of Vertebrate Paleontology Volume 28, Supplement to Number 3, 111A.
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Malkani M.S. 2008d. Mesozoic terrestrial ecosystem from Pakistan. In, Sundquist, B. (Science Programme coordinator), Abstracts of the 33rd International Geological Congress, Oslo, August 6-14, 2008, 1p. (Abstract: CD-ROM).
Malkani M.S. 2007a. Trackways evidence of sauropod dinosaurs confronted by a theropod found from Mid Jur. Samana Suk Limestone of Pakistan. Sindh University Research Journal (Science Series) 39 (1):1-14.
Malkani M.S. 2007b. First diagnostic fossils of Late Cretaceous Crocodyliform (Mesoeucrocodylia, Reptilia) from Vitakri area, Barkhan District, Balochistan, Pakistan. In; Ashraf, M., Hussain, S. S., and Akbar, H. D. eds. Contribution to Geology of Pakistan (2007), Proceedings of 5th Pakistan Geological Congress (2004), A Publication of the National Geological Society of Pakistan, Pakistan Museum of Natural History, Islamabad, Pakistan, 241-259
Malkani M.S., 2007c. Lateral and vertical rapid variable Cretaceous depositional environments and Terrestrial dinosaurs from Pakistan. In; Huang, Y., Wang, P., Gu, J., & Jing, S. eds. Abstracts volume, Joint Workshop on Rapid Environmental/Climate Change in Cretaceous Greenhouse World: Ocean-Land Interaction and Deep Terrestrial Scientific Drilling Project of the Cretaceous Songliao Basin, August 28-30, 2007, Daqing, China, 44-47
Malkani M.S. 2006a. Biodiversity of saurischian dinosaurs from the latest Cretaceous Park of Pakistan. Journal of Applied and Emerging Sciences, 1(3):108-140.
Malkani M.S. 2006b. Cervicodorsal, Dorsal and Sacral vertebrae of Titanosauria (Sauropod Dinosaurs) discovered from the Latest Cretaceous Dinosaur beds/Vitakri Member of Pab Formation, Sulaiman Foldbelt, Pakistan. Journal of Applied and Emerging Sciences 1(3):188-196.
Malkani M.S. 2006c. Lithofacies and Lateral extension of Latest Cretaceous Dinosaur beds from Sulaiman foldbelt, Pakistan. Sindh University Research Journal (Science Series) 38 (1):1-32.
Malkani M.S. 2004. Saurischian dinosaurs from Late Cretaceous of Pakistan. In Hussain SS, and Akbar HD, eds. Fifth Pakistan Geological Congress, National Geological Society of Pakistan, Pakistan Museum of Natural History (Pakistan Science Foundation), April 14-15, Islamabad, 71-73.
Malkani M.S. 2003a. First Jurassic dinosaur fossils found from Kirthar range, Khuzdar District, Balochistan, Pakistan. Geological Bulletin University of Peshawar 36:73-83.
Malkani M.S. 2003b. Pakistani Titanosauria; are armoured dinosaurs?. Geological Bulletin Univ. Peshawar 36:85-91.
Malkani M.S. 2003c. Discovery of Partial Skull and Dentary of Titanosauria (Sauropod dinosaur) from the Late Cretaceous Pab Formation of Vitakri area, Barkhan district, Balochistan, Pakistan. Geological Bulletin Univ.Peshawar 36:65-71.
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Malkani M.S., Sun G. 2016. Fossil biotas from Pakistan with focus on dinosaur distributions and discussion on paleobiogeographic evolution of Indo-Pak Peninsula. Proceeding volume of 12th Symposium on “Mesozoic Terrestrial Ecosystems (MTE-12) and 3rd Symposium of International Geoscience Program (IGCP 608) “Cretaceous Ecosystem of Asia and Pacific” August 15-20, 2015, Paleontological Museum of Liaoning/Shenyang Normal University, Shenyang, China, Global Geology 19 (4), 230-240. Article ID: 1673-9736 (2016) 04-0230-11: Digital Obj Identifier/DOI: 10. 3969 / j. issn. 1673-9736. 2016. 04. 04.
Malkani M.S., Wilson, J.A., Gingerich, P.D. 2001. First Dinosaurs from Pakistan. Journal of Vertebrate Paleontology (USA), Volume 21(3): 77A.
Mannion P.D., Upchurch P., Barenes R.N., Mateus O. 2013. Osteology of the Late Jurassic Portugues sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms. Zoological Journal of Linnean Society 168:98-206.
Meteus O., Milan J., Romano M., Whyte M.A. 2011. New finds of stegosaur tracks from the Upper Jurassic Lourinhã Formation, Portugal. Acta Palaeontologica Polonica, 56 (3): 651-658. Milan J., Chiappe L.M., Loope D.B., Kirkland J.L., Lockley M.G. 2015. First report on dinosaur tracks from the Burro Canyon Formation, San Juan County, Utah, USA – evidence of a diverse, hitherto unknown Lower Cretaceous dinosaur fauna. – Annales Societatis Geologorum Poloniae, 85: 515-525. Miller C.E., Basu C., Fritsch G., Hildebrandt T., Hutchinson J.R. 2007. Ontogenetic scaling of foot musculoskeletal anatomy in elephants. JR Soc Interface 5: 465–475. Mohabey D.M. 1986. Note on dinosaur footprint from Kheda district, Gujarat. Journal Geological Society of India, 27:456-459.
Pascual C., Canudo J.I., Hernandez N., Barco J.L., Castanera D. 2012. First record of stegosaur dinosaur tracks in the Lower Cretaceous (Berriasian) of Europe (Oncala Group, Soria, Spain). – Geodiversitas, 34 (2): 297-312. Pienkowski G, Branski P., Pandey K.D. Schlogl J., Alberti M., Fursich F.T. 2015. Dinosaur footprints from the Thaiat ridge and their palaeoenvironmental background, Jaisalmer Basin, Rajastan, India. Volumina Jurassica, 2015, XIII (1): 17–26. DOI : 10.5604/17313708 .1148553
Powell J. E. 2003. Revision of South American Titanosaurids dinosaurs: paleobiological, paleobiogeographical and phylogenetic aspects. Records of the Queen Victoria Museum, Launceston. 111:173p. Riga B.J.G. 2011. Speeds and stance of titanosaur sauropods: analysis of Titanopodus tracks from the Late Cretaceous of Mendoza, Argentina. Anais da Academia Brasileira de Ciências (Annals of the Brazilian Academy of Sciences) 83(1): 112. Printed version ISSN 0001-3765 / Online version ISSN 1678-2690 www.scielo.br/aabc. Romano M., Whyte M., Jackson S.J. 2007. Trackway Ratio: A New Look at Trackway Gauge in the Analysis of Quadrupedal Dinosaur Trackways and its Implications for Ichnotaxonomy. Ichnos 14: 257–270. Russel D. A. 1981. Estimated speed of a giant bipedal dinosaur. Nature, v. 292, p. 274.
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Sahni A. 2001. Dinosaurs of India. National book Trust, Delhi, 110pp.
Santos V.F.D., Lockley M.G., Meyer C.A., Carvalho J. de, Carvalho A.M.G., Moratalla J.J. 1994. A new sauropod tracksite from the Middle Jurassic of Portugal. Gaia 10, 5-13. Thulborn R.A. 1990. Dinosaur tracks. Chapman and Hall, London, 410 p. Vidal D., Ortega, F. and Sanz, J.L. 2014. Titanosaur Osteoderms from the Upper Cretaceous of Lo Hueco (Spain) and Their Implications on the Armor of Laurasian Titanosaurs. PLoS ONE 9(8): e102488. doi:10.1371/journal. pone.0102488. Wilson J.A., Carrano M.T. 1999. Titanosaur locomotion and the origion of “wide gauge” trackways: a biomechanical and systematic perspective on sauropod locomotion. Paleobiology, (25): 252-263.
Wilson J.A. Malkani M.S., Gingerich P.D. 2001. New Crocodyliform (Reptilia, Mesoeucrocodylia) form the Upper Cretaceous Pab Formation of Vitakri, Balochistan (Pakistan), Contributions form Museum of Paleontology, University of Michigan, 30 (12):321-336. Wilson J.A., Malkani M.S., Gingerich P.D. 2005. A sauropod braincase from the Pab Formation (Upper Cretaceous, Maastrichtian) of Balochistan, Pakistan. Gondwana Geological Magazine, Special Volume 8:101-109. Xing L., Lockley M.G., McCrea R.T., Gierlinski G.D., Buckley L.G., Qi L. Jia C. 2012. Thyrophoran tracks of presumed stegosaurian affinity from the Early Cretaceous, Xingjiang, China. In: Min Huh, Hyun Joo Kim and Jin-Young Park, Eds. The 11th Symposium on “Mesozoic Terrestrial Ecosystems (MTE 2012), 423-426, Gwanju, Korea, August 15-18, 2012. Xing L., Lockley M.G., McCrea R.T., Gierlinski G.D., Buckley L.G., Zhang J., Qi L., Jia C. 2013. First record of Deltapodus tracks from the Early Cretaceous of China. Cretaceous Research, 42: 55-65. Yadagiri P., Ayyasami K. 1979. A new stegosaurianWhyte M.A., Romano M. 1993. Footprints of a sauropod dinosaur from the Middle Jurassic of Yorkshire. Proceedings of the Geologists’ Association, 104: 195- 199. Whyte M.A., Romano M. 1995. Probable sauropod footprints from the Middle Jurassic of Yorkshire, England. Gaia, 10: 15-26. Whyte M.A., Romano M. 2001. Probable stegosaurian dinosaur tracks from the Saltwick Formation (Middle Jurassic) of Yorkshire, England. – Proceedings of the Geologists’ Association, 112: 45-54.
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Following Plate 1,2 show footprints and trackways of Malakhelisaurus mianwali Malkani 2007a titanosaurian sauropod confronted by Samanadrinda surghari Malkani 2007a large theropod dinosaur found on Middle Jurassic Samanasuk Limestone, Upper Indus Basin/Kohat-Potwar basin, Malakhel ichnite, Mianwali District, Punjab Province, Pakistan
Plate 1
Malakhelisaurus mianwali titanosaurian sauropod trackways confronted by Samanadrinda surghari theropod track
Plate 2
Left manus Left Pes with three large toes. Large oval ellipsoid osteoderms are being assigned as toe of pes Right manus
Asymmetric D shape right manus of Malakhelisaurus mianwali Malkani 2007a.
Right Pes with toes
Right Pes with toes
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Following Plate 3 show footprints and trackway of a couple of Himalayadrinda potwari Malkani 2015g found on found on Middle Jurassic Samanasuk Limestone, Upper Indus Basin/Kohat-Potwar basin, Malakhel site, Mianwali District, Punjab Province, Pakistan
Trackways of a couple (left and right) of small/ noasaurian theropod Himalayadrinda potwari Malkani 2015 found from Middle Jurassic Limestone
Plate 3
(central arrows show movement direction)
Trackway of left small/ noasaurian theropod Himalayadrinda potwari Malkani 2015
Subparallel Trackway of right small/ noasaurian theropod Himalayadrinda potwari Malkani 2015
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Age
Formation
Lithologic log
Descriptions
LATEST VITAKRI
Level 5 (Pashtosaurus zhobi ichnotype bed)
FORMATION
Level 4 (micro fauna prints on burrowed & ripple mark surface) Level 3 (Pashtosaurus zhobi referred footprints bed) Level 2 (Pashtosaurus zhobi referred footprints bed and ichnotype of Anmolpakhi alleni new genus and species of pterosaur-flying reptile
CRETACEOUS
Level 1 (Pashtosaurus zhobi referred footprint bed)
Legend Sandstone
Mudstone/shale
Plate 4: Lithologic section through the 5 levels of footprints and trackways found in the Latest Cretaceous Vitakri Formation, Qila Saif ullah District, Zhob Division, Balochistan Province, Pakistan
25
Plate 5 Upper, Tentative Map of Latest Cretaceous Sor Muzghai ichnite site of Pashtosaurus zhobi titanosaurs
Lower , Line drawings of left pes with digits of Pashtosaurus zhobi titanosaurs
26
Following Plate 6 shows filled matrix plate in footprint of Pashtosaurus zhobi (level 5)
Plate 9
27
Following Plate 7-25 show ichnotype (footprints and trackway) of Pashtosaurus zhobi (on level 5) Pes?
manus
pes with toe marks
Plate 7
Complete Pes Pes with posterior convexities,median concavity, w shape heel
Manus gliding line
Manus Right Manus gliding line, parallel to left manus glide line
Left Manus gliding line, parallel to right manus glide line
28
Central toe of right pes
Plate 8
Left toe of right pes
W shape heel of two pes
1st digit 2nd digit and toe 3rd digit and toe 4th digit and toe left pes
W shape heel of pes
5th digit of
Asymmetric D shape manus without claw
Toe of pes
29
Plate 9
Plate 10 Footprint continuing under the overlying bed showing these are not man-made.
Pes footprint continuing under the overlying bed showing these are not man-made.
30
Plate 11
Footprint continuing under the overlying bed showing these are not man-made.
31
Plate 12
Pes footprint of Pashtosaurus zhobi Malkani 2014 vide Malkani 2015
32
Plate 13
33
Plate 14
34
Plate 15
35
Plate 16
36
Plate 17
37
Plate 18
38
Plate 19
39
Plate 20
40
Plate 21
41
Plate 22
42
Plate 23
43
Plate 24
44
Plate 25
45
Following Plate 26-27 show referred (footprints and trackway) of Pashtosaurus zhobi Malkani 2014 found on bed level 3
Plate 26
D-Oval manus footprints of titanosaurs
46
Plate 27
D-Oval manus footprints of titanosaurs
47
Following Plate 28-29 show footprints and trackway of Pashtosaurus zhobi Malkani 2014 and pterosaur (trackmaker may be Saraikisaurus minhui-a flying reptile) found on bed level 2
Plate 28 Pterosaur footprints and trackway
Digit 3rd, 4th,
5th of right manus footprint of titanosaur
Asymmetric D shape right Manus footprint of titanosaur
Digit 2nd Digit 1st of right manus footprint of titanosaur
Right Manus track /footprint of titanosaur showing continuity under the overlying bed
48
Plate 29 Pterosaur footprints and trackway
Asymmetric D shape right Manus footprint of titanosaur (length 55cm, width +60cm exposed while about 15cm is covered under the overlying bed) showing continuity just under the overlying bed. It represents it is not man-made structure.
Pterosaur footprint
49
Following Plate 30 show footprint of Pashtosaurus zhobi Malkani 2014 found on bed level 1
Plate 30 Manus footprint of titanosaur
50
Plate 31 of a plaster cast of a footprint of sauropod from India (after Mohabey 1986)
51
,
Plate 32, Row 1, large oval ellipsoid MSM-84-7 ( 3 views) of Balochisaurus and osteodermal ellipsoid MSM-85-4 (one view) of Pakisaurus now assigned as their pes toes (for different views pl.see plate 44). Row 2, Pakisaurus proximal metacarpal MSM-280-4 in two views, Pakisaurus partial metacarpals MSM-350-4 and MSM-970-4, Pakisaurus proximal metacarpal MSM-296-7. Row 3, Pakisaurus ungual, now may belong to another genera or its osteoderms MSM-152-3 in 4 views. Row 4, Photo 1,2, Balochisaurus metacarpal MSM-297-15 in twoviews. Photo 3, Marisaurus metacarpals, upper row, proximal metacarpals MSM-295-2, MSM-279-2, MSM-685-2, MSM-566-2, MSM-278-2, MSM-686-2, MSM-1029-2, MSM-688-2; lower row, distal metacarpals MSM-277-2, MSM-1028-2, MSM-285-2, MSM-370-2, MSM-684-2, MSM-687-2, MSM-361-2, MSM-683-2. Photo 4, Marisaurus metatarsals, upper metatarsal MSM-643-2, and lower metatarsals MSM-1031-2 and MSM-1030-2 Photo 5, Marisaurus distal metacarpal MSM-1036-16. Scale, every black or white digit is 1cm.
52
Plate 33. Very broad and very long toe of Balochisaurus MSM-84-7 in 3 views (left), and Pakisaurus MSM-85-4 in 3 views (right) titanosaurian sauropods previously assigned as osteoderms. The size of fossil toes is fit to footprint toes. Scale every black/white unit is 1cm.
53
Plate 34, line drawing of manus with digits I-V
54
