Kuldana Formation

Geological formation in Pakistan

Kuldana Formation

Stratigraphic range: Lutetian

PreꞒ

Type| Geological formation
Unit of| Subathu Group
Underlies| Kohat Formation
Overlies| Ghazij, Shekhan & Chorgali Formations
Location
Location| Gilgit-Baltistan and Punjab, Pakistan
Coordinates| 33°43′N 72°10′E﻿ / ﻿33.717°N 72.167°E﻿ / 33.717; 72.167
Approximate paleocoordinates| 14°18′N 68°42′E﻿ / ﻿14.3°N 68.7°E﻿ / 14.3; 68.7
Region| Kala Chitta Range
Country|  Pakistan
Type section
Named for| Kuldana Village, Hazara District[1]
Named by| Latif
Year defined| 1970

Kuldana Formation (Pakistan)

The Kuldana Formation is a fossil-bearing geological formation of Lutetian (Early Eocene) age which crops out in northern Pakistan. The abundant fossil remains were deposited by rivers and estuaries crossing an arid to semi-arid environment, between several marine transgressions. Its fossil fauna is best known for the early cetaceans Indohyus, Pakicetus and Ambulocetus, that helped to shed a new light on the evolution of whales, but it also features a large number of early ungulates, rodents and primates.

## History

[edit]

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Fossils from the Kuldana Formation have been studied since long before the name was established, with some of the earliest papers being those of Pilgrim, published in 1940. At that point in time, the sediments were known as the Chharat beds. The name Kuldana Formation as such was first coined by Mir Abdul Latif in 1970, prior to which the sediments had already been known as the Kuldana Beds and Kuldana Series, deriving its name from a nearby village. Around the same time, Meissner and colleagues studied similar outcrops near the village of Mami Khel, 200 km (120 mi) southwest of Kuldana, naming it the Mami Khel Clay. Although study began prior to the publication of Latif's work, this paper was not published until later. Several subsequent papers comparing the two generally agreed that they were synonyms and represented a single geological formation, that being the Kuldana Formation. A 1996 publication by Pivnik and Wells meanwhile used the name Mami Khel Formation and Maas et al. (2001) treated the two as distinct formations primarily based on their geographic separation through the Indus River while providing no other indicators in how the two differed. Other studies treating the two formations as distinct include Leinders (1999) and Thewissen (2001).[1]

## Geography

[edit]

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Outcrops of the Kuldana Formation are found in the form of isoclinal folds across northern Pakistan's Kohat plateau and Potwar plateau. The Formation stretches from Barbora and Mami Khel at the western edge of the Kohat plateau to the name-giving locality of Kuldana at the eastern end of the Potwar plateau. Between these points are a variety of other localities as well, such as Chorlakki, located within the eastern-most Kohat plateau, as well as Lammidhan and Ganda Kas (Kala Chitta) at the western end of the Potwar plateau.[1][2]

## Stratigraphy

[edit]

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While the stratigraphy within the Kuldana Formation is poorly understood, its relation to under- and overlying formations has been much better studied thanks to the succession being consistently visible across various localities. At Banda Daud Shah, the Kuldana Formation overlies the Ghazij Formation, the Shekhan Formation and the Jatta Gypsum. The Ghazij and Shekhan Formations also underlie the Kuldana Formation at Chrolakki while at Ganda Kas and Gali Jhagir the formation preceding the Kuldana was the Chorgali Formation. The Kohat Formation consistently overlies the Kuldana Formation.[1][2]

## Age

[edit]

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The age of the Kuldana Formation has been subject to repeated revision and multiple back and forths among researchers. Two early estimates, proposed by Cotter and Pilgrim respectively, suggested that the Kuldana Formation was either Ypresian (Early Eocene) or Lutetian (early Middle Eocene) in age, with subsequent authors generally following one of these hypothesis. After officially establishing the name Kuldana Formation, Latif suggested that the formation dates to the lower to middle Eocene on the basis of shallow benthic foramins, somewhat straddling the line between both the results of Cotter and Pilgrim. This would come to be the general result of subsequent papers as well, which often recovered an age within that general range.[1]

In 1983 Gingerich used the fact that the Kuldana Formation is preceded and followed by marine strata to compare it with then available maps of sea levels during the Eocene, arguing for a late Early Eocene age, once again falling into the convention established by prior work. Gingerich did however come to revise his 1983 age estimate in a paper published in 2003 that dealt with the stratigraphy observed across different localities in order to obtain a more reliable result. Another aim of this work was to provide counter arguments to some claims made around the turn of the century, when several papers claimed that the sediments at Mami Khel represented a distinct formation that dated to the early Early Eocene. To do so, foramins of underlying formations were used to establish a maximum age for the Kuldana strata. At Banda Daud Shah, Gingerich recovered a late Early Eocene to early Middle Eocene (P9 or P10) age based on the planktonic foraminifera of the underlying Shekhan Formation, with similar results being recovered based on shallow benthic foramins at Chorlakki, Ganda Kas and Gali Jhagir (all dated to SB12 to SB13). Gingerich further utilizes updated knowledge on global sea levels to find similar results. Unlike in 1983, Gingerich now recognized two distinct periods of low sea levels, one during the late Early Eocene and a second, longer-lasting period during the early Middle Eocene. Gingerich argues that, since only one terrestrial phase is observed within the stratigraphy of Eocene Pakistan, it is likely that the Kuldana Formation represents the longer period of low sea levels, placing it within the early Late Eocene.[1]

## Paleoenvironment

[edit]

Life reconstructions of Pakicetus inachus and Gujaratia pakistanensis.

The Kuldana Formation featured several different environments across its time of deposition, with the oldest layers showing fluvial conditions. The strata overlying these sediments suggest that the environment shifted to a mix of freshwater and shallow marine habitats, which is supported by multiple lines of evidence. Evidence for freshwater biomes is present through isotopic analysis as well as the local fauna, for example the freshwater snail Planorbis, whereas the presence of sharks and pycnodontoid fish shows the presence of marine habitats. It is thought that the archaeocetes that inhabited the formation at the time stuck to freshwater biomes, even if some localities were evidently closer to the coast.[3]

This mix of fresh- and saltwater habitats was eventually followed by a marine transgression that characterizes the geology of the uppermost layers of the formation.[3]

### Ganda Kas

[edit]

Fossiliferous sediments in and around the Ganda Kas localities were deposited in semi-arid Eocene floodplains and freshwater channels. The H-GSP Locality 62, from which an abundance of material is known, was deposited in a stillwater environment, while other localities in the area represent the remains of prehistoric rivers. Localities around Ganda Kas deposited in a marine environment appears to be of a later age.[4]

### Banda Daud Shah

[edit]

Like in Ganda Kas, the depositional environment in the areas presenting a Banda Daud Shah-type fauna, such as the eponymous Banda Daud Shah and Barbora Banda localities, represents an arid environment crossed by various rivers. The most common fossil mammals are the early artiodactyl Diacodexis pakistanensis and the tapiroid perissodactyl Karagalax mamikhelensis ; the presence of adapids and arctocyonids in the area indicates that it probably had an older age than the Ganda Kas fauna.[4]

## Paleobiota

[edit]

### Mammals

[edit]

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Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

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| Notes
Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.

#### Arctocyonia

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Karakia[4] | K. longidens | H-GSP Loc. 9710, Banda Daud Shah  | A single, fragmentary mandible.  | The first arctocyonid from Indo-Pakistan.  |

#### Artiodactyla

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Ambulocetus[3][5] | A. natans | H-GSP Loc. 9209 & 9204  | Several specimens.  | An ambulocetid cetacean.  |
Artiodactyla indet.[6] |  | Chorlakki  | An astragalus and two calcaneum | These fossil remains are thought to be too large and robust to have belonged to the Chorlakki dichobunids or Indohyus.  |
Attockicetus[3][5] | A. praecursor | H-GSP Loc. 9204

H-GSP Loc. 9607, Shepherd's Lake

| An incomplete cranium, worn teeth, premolars.  | A remingtonocetid cetacean.  |
Cf. A. sp.  | H-GSP Loc. 9607, Shepherd's Lake  | Two teeth  | A remingtonocetid cetacean.
Basilosauridae indet.[7] |  | Ganda Kas  | Two isolated teeth.  | One of the teeth resemble those of Ichthyolestes.  |
Chorlakkia[6] | C. hassani | Chorlakki  | A left dentary and multiple teeth  | A dichobunid.  |
Dichobunidae indet.[6] |  | Chorlakki  | Teeth  | Teeth distinct from the other known Kuldana dichobunids. One particular tooth might represent a hyopsodontid.  |
Dulcidon[8][9][5] | D. gandaensis | Ganda Kas & Chorlakki  | Two right molar.  | A dichobunid |
Gandakasia[3][7] | G. potens | H-GSP Loc. 58, Ganda Kas
H-GSP Loc. 9607 Shepherd's Lake  | A fragmentary mandible  | A protocetid cetacean |
Gobiohyus[7][6][4] | cf. G. orientalis | Ganda Kas  | A single tooth.  | An helohyid.  |
Gujaratia[6][4][10][11] | G. pakistanensis | Chorlakki
Lammidhan
Barbora Banda I & II
H-GSP Loc. 300, 9710 & 9712, Banda Daud Shah  | Teeth  | A diacodexeid. Gujaratia is rare at Chorlakki and may be the only artiodactyl at Barbora Banda. Formerly classified within the European and American genus Diacodexis.  |
Ichthyolestes[3] | I. pinfoldi | H-GSP Loc. 62, Ganda Kas  | A single molar  | A pakicetid.  |
Indohyus[6] | I. indirae | Ganda Kas, Chorlakki & Kalakot  | Numerous specimens  | A raoellid. The most abundant artiodactyl at Kalakot, but less common at Chorlakki where Khirtharia is more prominent.  |
I. major | Chorlakki  | An isolated tooth  | A larger species of raoellid, that would reach about twice the size of I. indirae, but might also represent an unrelated species of diacodexeid or dichobunid.
Khirtharia[7][6][4] | K. dayi | Chorlakki, Kalakot & Panoba

H-GSP Loc. 62, Ganda Kas

| Fragments of maxilla and mandibles ; isolated teeth  | A large helohyid. Khirtharia is the must abundant artiodactyl at Chorlakki and more common than Indohyus, but the reverse is seen at Kalakot.  |
Nalacetus[3][5] | Nalacetus ratimitus | H-GSP Loc. 62, Ganda Kas  | Fragments of palate, maxilla and teeth  | A pakicetid cetacean |
Pakibune[6] | P. chorlakkiensis | Chorlakki & Lammidhan  | Teeth  | A moderately sized dichobunid, larger than the other dichobunid taxa from the formation.  |
Pakicetus | P. attocki[7] | H-GSP Loc. 62, Ganda Kas  | Complete cranial remains.  | A pakicetid cetacean |
P. calcis[3] | H-GSP Loc. 9607 Shepherd's Lake
H-GSP Loc. 9607, Valley E  | A partial mandible, palate fragment and teeth  | A pakicetid cetacean |
P. chittas[3] | Chorlakki

H-GSP Loc. 9607, Shepherd's Lake

| Fragments of mandibles  | A pakicetid cetacean |
P. inachus[6][12] | Chorlakki  | A partial skull, mandible and teeth  | A pakicetid cetacean |
Protocetidae indet.[7] |  | Ganda Kas  | Jaw fragment and two isolated teeth  | One of the teeth resemble those of Ichthyolestes.  |

#### Chiroptera

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Chiropteran indet. A[6][13] |  | Chorlakki  | A molar  |  |
Chiropteran indet. B[6][13] |  | Chorlakki  | Two fragmentary molars  | Resemble that of Palaeochiropteryx, and may belong to a relatively large bat.  |

#### Eulipotyphla

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Pakilestes[6][13][14] | P. lathrius | Chorlakki  | Two molars and a premolar  | An insectivore mammal of uncertain affinities, possibly a plesiosoricid soricomorph.  |
Perizalambdodon[15] | P. punjabiensis | H-GSP Loc. 9610, Jhalar  | A single molar  | An indeterminate lipotyphlan.  |
Seia[6][13] | S. shahi | Chorlakki  | Two molars  | An erinaceomorph, probably representing a new family.  |

#### Herpetotheriidae

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Herpetotheriinae indet.[4] |  | H-GSP Loc. 62, Ganda Kas  | A right molar.  |  |

#### Hyaenodonta

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Paratritemnodon[6][4] | P. indicus | Ganda Kas & Chorlakki  | A mandible (now lost), several isolated teeth  | Represent a smaller form of hyaenodont.  |
P. jandewalensis | H-GSP Loc. 9205, Ganda Kas  | Fragmentary maxilla and isolated tooth.  | The teeth are twice as large than those of P. indicus, and it was probably much larger.  |

#### Mesonychia

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Mesonychidae indet.[7] |  | Ganda Kas  | A single premolar  | Belongs to a small mesonychid, with similarities to Hapalodectes.  |

#### Perissodactyla

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Anthracobune[11][10][16][17] | A. pinfoldi[7] | Upper Member : Lammidhan & Ganda Kas  | Relatively complete cranial remains and isolated teeth.  | The largest anthracobunid in the formation.  |
A. wardi[6] | Chorlakki & Ganda Kas  | Several mandibles, fragments of a skull and isolated teeth.  | An anthracobunid. Formerly the distinct genus Lammidhania, also present in the Subathu Formation. Includes the holotype of Pilgrimella pilgrimi.  |
"Forstercooperia"[6] | "F." jigniensis | Chorlakki  | Teeth  | A paraceratheriid.  |
Isectolophidae indet.[6] |  | Chorlakki  | A tooth  |  |
Jozaria[11][16] | J. palustris | Upper Member  | Several teeth  | An anthracobunid.  |
cf. Kalakotia[4] | K. sp.  | H-GSP Loc. 62, Ganda Kas

H-GSP Loc. 9613, Thatta

| A fragmentary maxilla and isolated teeth  | A lophialetid tapiroid, probably representing a new species.  |
Karagalax[18] | K. mamikhelensis | H-GSP Loc. 300, Barbora Banda  | Several relatively well-preserved skulls ; isolated postcranial elements tentatively referred to the genus.  | An isectolophid tapiromorph, more cursorial than its contemporary American relatives.  |
Obergfellia[6][11] | O. occidentalis | Ganda Kas  | Several mandibles.  | An anthracobunid. Includes most of the material formerly assigned to Pilgrimella pilgrimi.  |
Palaeosyops[6][7][4][19] | P. dayi | Chorlakki

H-GSP Loc. 64, 9613 and 227, Ganda Kas & Thatta

| Fragmentary maxilla and mandible ; isolated teeth  | Formerly belonging to the genus Eotitanops, it seems to be intermediate between the two genera. A small and primitve brontothere. Also present in the Baska Formation.  |
Pakotitanops[7][4] | P. latidentatus | H-GSP Loc. 9205 & 9206, Ganda Kas  | A fragmentary maxilla and isolated teeth  | A brontothere, distinctly larger and more derived than Eotitanops.  |

#### Primates

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Cf. Agerinia[6][20] | Cf. A. sp. | Chorlakki  | A tooth.  | A notharctid, with similarities to A. roselli.  |
Jattadectes[4] | J. mamikheli | H-GSP Loc. 9712, Banda Daud Shah  | A premolar and an incisor  | The first plesiadapid from Indo-Pakistan.  |
Kohatius[21][22] | K. coppensi | Chorlakki  | Teeth.  | An omomyid. Might also be present in the Ghazij Formation.  |
cf. K. sp.  | Barbora  | Mandible fragment.  | An omomyid ; the smallest primate in Indo-Pakistan.  |
K. sp. A  | H-GSP Locality 223, Jhalar  | A premolar.  | An omomyid with similarities with Altanius.  |
Panobius[6][4][21][20] | P. afridi | Chorlakki  | Two isolated teeth.  | An adapid.  |
P. amplior | Either from Chorlakki,[21] or from H-GSP Loc. 9712, Banda Daud Shah.[4] | A fragmentary mandible with two associated molars and isolated teeth.  | An adapid, much larger than P. russelli and P. afridi, to which it was firstly attributed.  |
Parvocristes[4] | P. oligocollis | H-GSP Loc. 225, Jhalar  | A premolar and an incisor  | A carpolestid.  |

#### Rodentia

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Birbalomys[6][4][23] | B. (Basalomys) ijlsti | H-GSP Loc. 62, Ganda Kas  | Teeth  | A chappatimyid, the most common rodent in its locality.  |
B. (Birbalomys) sondaari | Chorlakki

H-GSP Loc. 57, 62 and 144, Ganda Kas

| Teeth  | A chappatimyid abundant in Chorlakki and Ganda Kas.  |
B. (Birbalomys) woodi | Chorlakki  | Three teeth.  | A chappatimyid. Relatively rare in Chorlakki, but abundant in other similarly-aged formations.  |
B. (Basalomys) vandermeuleni[24] | Shekhan, Chorlakki

H-GSP Loc. 57 & 144, Ganda Kas

|  | A chappatimyid. Formerly Saykanomys.  |
Chapattimys[6][23] | C. debruijni | Chorlakki

H-GSP Loc. 9205, Ganda Kas

| Teeth  | One of the largest chappatimyid in Indo-Pakistan, reaching twice the size of C. wilsoni. Quite rare in all the deposits where it is found.  |
C. wilsoni | Chorlakki

H-GSP Loc. 62 & 144, Ganda Kas

| Teeth  | A chappatimyid.  |
Gumbatomys[6][23] | G. asifi | H-GSP Loc. 62, Ganda Kas ; Chorlakki  | Several teeth  | A rare chappatimyid.  |
Paramyidae indet.[8][25] |  | Barbora Banda I  | Teeth  |  |
Cf. Petrokozlovia[6][23] | Cf. P. sp. indet. 1  | Chorlakki

H-GSP Loc. 57, Ganda Kas

| One tooth.  | Less derived than P. notos, but similar to a more primitive specimen from Kazakhstan.  |
Cf. P. sp. indet. 2  | Chorlakki  | One tooth.  | Seemingly closer to P. notos, from Mongolia, than the other species.  |
Cf. Tamquammys[24] | Cf. T. sp.  | Chorlakki  |  | A Tamquammyidae.  |
Tamquammyidae indet.[8][4][25] | spe. indet. 1  | Barbora Banda I  | Teeth  |  |
spe. indet. 2  | Barbora Banda I  | Teeth  |  |

#### Tillodontia

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Basalina[6][26] | B. basalensis | Ganda Kas & Chorlakki  | A fragmentary mandible and associated teeth.  | A small estonychid tillodont, firstly identified as a taeniodont.  |
cf. B. basalensis | Ganda Kas  | A jaw fragment.  | The heavy wear on the specimen renders the identification as B. basalensis only tentative.  |

#### Other mammals

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Mammalia indet.[8] |  | Barbora Banda II  | A single, large incisor.  | May belong to a small artiodactyl.  |

### Fish

[edit]

#### Actinopterygians

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Acanthopterygii indet.[27] |  | Chorlakki & Shekhan Nala  | Several isolated teeth.  | Some of the teeth might belong to an Osteoglossiforme indeterminate.  |
Amiidae indet.[28] |  | H-GSP Loc. 56, Ganda Kas redbeds  | Partial right premaxilla with associated teeth.  | From predominantly marine deposits.  |
Anchichanna[28] | S. kuldanensis | H-GSP Loc. 62, Ganda Kas  | Several relatively complete cranial remains.  | A snakehead.  |
Ariidae indet.[27] |  | Chorlakki  | Several abdominal vertebra.  | Despite being primarily marine, ariid catfish are known to frequently enter freshwater environments.  |
Bagridae indet.[27] |  | Chorlakki  | An angular bone, a cleithrum and pectoral spines.  |  |
Cf. Bagridae indet.[28] |  | H-GSP Loc. 62, Ganda Kas  | Fragments of the skull and spines.  | Might represent several species of catfish.  |
Clariidae indet.[27] |  | Chorlakki & Shekhan Nala  | An articular bone and pectoral spines.  | The low angular resemble that of Heterobranchus.  |
Cyprinidae indet.[27] |  | Chorlakki  | A pharyngeal tooth.  |  |
Cyprinodontidae indet.[27] |  | Chorlakki  | Several teeth and a quadrate bone.  | The teeth are similar to those of Aphanius.  |
Eotrigonodontidae indet.[28] |  | H-GSP Loc. 229 & 9607, Shepherd's Lake  | Two teeth.  | Tentatively referred to the Mesozoic genera Hadrodus or Stephanodus.  |
Lepisosteus[27] | L. sp.  | Chorlakki  | A tooth and a scale.  | Appears to be closely related or identical to the modern genus Lepisosteus osseus.  |
Macquaria[27] | M. antiquus | Chorlakki  | Various bone fragments and isolated spines  | A temperate perch.  |
Osteoglossidae indet.[27] |  | Scales : Chorlakki & Shekhan Nala

Maxilla : H-GSP Loc. 9611

| A maxilla, several scales of various size and shape.  |  |
Perciformes indet.[28] |  | H-GSP Loc. 62, Ganda Kas  | Isolated fin spine.  | Distinct from Macquaria antiquus.  |
Pycnodontoidea indet.[3][28] |  | H-GSP Loc. 9206 & 9608, Ganda Kas
H-GSP Loc. 9607, Shepherd's Lake  | Several isolated teeth and a jaw fragment.  | From predominantly marine deposits.  |
Siluriforme indet.[27] |  | Chorlakki  | A basioccipital.  |  |
Teleostei indet.[28] |  | H-GSP Loc. 62, Ganda Kas  | Isolated teeth.  | From predominantly freshwater deposits, possibly representing several species.  |
Cf. Varohstichthys[27] | Cf. V. sp.  | Chorlakki  | A pharyngeal tooth.  | A Cyprinidae.  |

#### Chondrichthyans

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Selachimorpha indet.[3] |  | H-GSP Loc. 9607, Shepherd's Lake  | Teeth.  |  |

### Reptiles

[edit]

#### Crocodilians

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Crocodilia indet.[8][29] |  | Barbora Banda II & H-GSP Loc. 62, Ganda Kas  |  |  |

#### Squamates

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Tinosaurus[27] | T. sp.  | Chorlakki  | Two isolated teeth.  | Similarities can be observed with Eocene species of Tinosaurus from North America and China.  |
Sauria indet.[27] |  | Chorlakki  | A fragmentary vertebra.  |  |
Boidae indet.[27] |  | Chorlakki  | Five fragmentary vertebrae.  |  |
Erycinae indet.[27] |  | Chorlakki  | Two fragmentary vertebrae.  | The material probably represent a new genus and species of sand boa.  |
Boinae indet.[27] |  | Chorlakki  | Two fragmentary vertebrae.  | The material probably represent a new genus and species of boa resembling the Eocene genera Paleryx and Palaeopython.  |

#### Turtles

[edit]

Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Trionychinae indet.[27] |  | Chorlakki  | Four fragments of pleural plates.  | The fragments belonged to a shell reaching 30 cm in length and are similar to plates found in the Kala Chitta Hills and Lammidhan localities.  |
"Chorlakkichelys"[27] | "C. shahi"  | Chorlakki  | Fragmentary shell belonging to a single individual.  | A Carretochelyinae. Material from Chharat, Jhalar and Lammidhan can be tentatively assigned to the genus. A 2014 study established that the genus Chorlakkichelys is a nomem dubium due to its lack of diagnostic apomorphy.[30] |

### Mollusca

[edit]

Name  | Species  | Locality/Member/Microfacies  | Material  | Notes  | Image

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Bivalvia indet.[31] |  | Molluscan microfacies
Ostracode wackestone  | "Oyster" shell fragments  | Composes more than half of the molluscan microfacies.  |
Gastropoda indet.[31] |  | Molluscan microfacies
Ostracode wackestone  |  |  |
Planorbis[3][24][29] | P. sp.  | Chorlakki
Shekhan Nala
H-GSP Loc. 62, Ganda Kas
H-GSP Loc. 9607, Shepherd's Lake  |  | A freshwater snail.  |

### Plants

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Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

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| Notes
Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.
Name  | Species  | Locality/Member  | Material  | Notes  | Image

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Cf. Ajunginucilla[8] | Cf. C. sp.  | Barbora Banda I  | Seeds.  | A sage.  |
Celtis[8] | C. sp.[8] | Barbora Banda I  | Seeds.  | A hackberry.  |
Chara[8] | C. sp.  | Barbora Banda I  | Seeds.  | A charophyte.  |

### Other microfossils

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Name  | Species  | Microfacies  | Material  | Notes  | Image

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Assilina[31] | A. spp.  | Nummulitic wacke-packstone

Assilina wacke-packstone

| A foraminifera | Present in minor quantity in the nummulitic wacke-packstone, and as the dominant fossil in the Assilina wacke-packstone. Its presence in large numbers indicates deeper environments than the Nummulites deposits.  |
Biantholithus[31] | B. sparsus | Ostracode wackestone (Q-2)
Nummulitic wacke-packstone (Q-15)  |  |  |
Brachiopoda indet.[31] |  | Molluscan microfacies  |  |  |
Coccolithus[31] | C. foraminis | Nummulitic wacke-packstone (Q-15)  | Typical of warm waters  |  |
C. formusus | Nummulitic wacke-packstone (Q-15)  |  |  |
C. pauxillus | Ostracode wackestone (Q-2)  |  |  |
C. pelagicus | Lime mudstone (Q-1)
Ostracode wackestone (Q-2)
Nummulitic wacke-packstone (Q-15)  |  | Typical of warm , low-latitude waters  |
Cyclicargolithus[31] | C. luminis | Nummulitic wacke-packstone (Q-15)  |  |  |
Echinodermata indet.[31] |  | Molluscan microfacies  | Fragments  |  |
Fasciculithus[31] | F. lillianae | Molluscan microfacies (Q-13)  |  |  |
F. tympaniformis | Molluscan microfacies (Q-13)  |  |  |
F. clinatus | Molluscan microfacies (Q-13)
Assilina wacke-packstone (Q-17)  |  |  |
Lockhartia[31] | L. spp.  | Assilina wacke-packstone  | A rotaliid | Present in minor quantity in the Assilina wacke-packstone.  |
Neococcolithes[31] | N. protenus | Nummulitic wacke-packstone (Q-15)  |  |  |
Nummulites[31] | N. spp.  | Nummulitic wacke-packstone  |  | Present as the dominant fossil in the nummulitic wacke-packstone, and in lower numbers in the Assilina wacke-packstone. Typical of a low energy marine environment.  |
Ostracoda indet.[31] |  | Ostracode wackestone  |  |  |
Rhomboaster[31] | R. brameletti | Ostracode wackestone (Q-2)
Molluscan microfacies (Q-13)
Lime mudstone (Q-14)  |  |  |
Sphenolithus[31] | S. anarrhopus | Lime mudstone (Q-1, Q-14)
Ostracode wackestone (Q-2)  |  |  |
S. moriformis | Molluscan microfacies (Q-13)  |  |  |
S. primus | Ostracode wackestone (Q-2)
Molluscan microfacies (Q-13)  |  |  |
S. sp.  | Lime mudstone (Q-14)  | Typical of warm waters  |  |
Tribrachiatus[31] | T. absidatus | Assilina wacke-packstone (Q-22)
Nummulitic wacke-packstone (Q-27)  |  |  |
T. lunatus | Assilina wacke-packstone (Q-22)  |  |  |
T. brameletti-contortus intergrade | Nummulitic wacke-packstone (Q-27)  |  |  |
T. contortus | Lime mudstone (Q-1)
Molluscan microfacies (Q-13)  |  |  |
Zeughrabdotus[31] | Z. sigmoides | Nummulitic wacke-packstone (Q-15)  |  | More typical of colder waters.  |

## References

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1. ^ a b c d e f Gingerich, P.D. (2003). "Stratigraphic and micropaleontological constraints on the middle Eocene age of the mammal-bearing Kuldana Formation of Pakistan" (PDF). Journal of Vertebrate Paleontology. 23 (3): 643–651. Bibcode:2003JVPal..23..643G. doi:10.1671/2409.
2. ^ a b Bilal, Ahmer, Muhammad Saleem Mughal, Hammad Tariq Janjuhah, Johar Ali, Abrar Niaz, George Kontakiotis, Assimina Antonarakou, Muhammad Usman, Syed Asim Hussain, and Renchao Yang. 2022. "Petrography and Provenance of the Sub-Himalayan Kuldana Formation: Implications for Tectonic Setting and Palaeoclimatic Conditions" Minerals 12, no. 7: 794. https://doi.org/10.3390/min12070794
3. ^ a b c d e f g h i j k l Cooper, L.N.; Thewissen, J. G. M.; Hussain, S. T. (2009). "New middle eocene archaeocetes (Cetacea: Mammalia) from the Kuldana formation of northern Pakistan". Journal of Vertebrate Paleontology. 29 (4): 1289–1299. Bibcode:2009JVPal..29.1289C. doi:10.1671/039.029.0423.
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5. ^ a b c d Gingerich, P.D.; Heissig, K.; Bebej, R.M.; von Koenigswald, W. (2017). "Astragali of Pakicetidae and other early-to-middle Eocene archaeocetes (Mammalia, Cetacea) of Pakistan: locomotion and habitat in the initial stages of whale evolution". Paläontologische Zeitschrift. 91 (4): 601–627. Bibcode:2017PalZ...91..601G. doi:10.1007/s12542-017-0362-8.
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10. ^ a b Gingerich, P. D. (1977). "A small collection of fossil vertebrates from the Middle Eocene Kuldana and Kohat Formations of Punjab (Pakistan)" (PDF). Contributions from the Museum of Paleontology. 24 (18): 190–203.
11. ^ a b c d Rautela, A.; Bajpai, S. (2023). "Gujaratia indica, the oldest artiodactyl (Mammalia) from South Asia: new dental material and phylogenetic relationships". Journal of Systematic Palaeontology. 21 (1). Bibcode:2023JSPal..2167553R. doi:10.1080/14772019.2023.2267553.
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13. ^ a b c d Russell, D. E.; Gingerich, P. D. (1981). "Lipotyphla, Proteutheria(?), and Chiroptera (Mammalia) from the Early-Middle Eocene Kuldana Formation of Kohat (Pakistan)". Contributions from the Museum of Paleontology. 25 (14): 277–287.
14. ^ Ziegler, R. (2009). "Plesiosoricids from early Oligocene fissure fillings in South Germany, with remarks on plesiosoricid phylogeny". Acta Palaeontologica Polonica. 54 (3): 365–371. doi:10.4202/app.2008.0061.
15. ^ Thewissen, J.G.M.; Bajpai, S.; Hussain, S.T. (2005). "New insectivorous placentals from the Eocene of Pakistan". Journal of the Palaeontological Society of India. 50 (2): 37–41.
16. ^ a b Wells, N.A.; Gingerich, P. D. (1983). "Review of Eocene Anthracobunidae (Mammalia, Proboscidea) with a new genus and species, Jozaria palustris, from the Kuldana Formation of Kohat, Pakistan" (PDF). Contributions from the Museum of Paleontology. 26 (7): 117–139.
17. ^ Cooper, L.N.; Seiffert, E.R.; Clementz, M.; Madar, S.I.; Bajpal, S.; Hussain, S.T.; Thewissen, J.G.M. (2014). "Anthracobunids from the Middle Eocene of India and Pakistan Are Stem Perissodactyls". PLOS ONE. 9 (10): e109232. Bibcode:2014PLoSO...9j9232C. doi:10.1371/journal.pone.0109232. PMC 4189980. PMID 25295875.
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20. ^ a b Russell, D.E.; Gingerich, P.D. (1987). "Nouveaux primates de l'Éocène du Pakistan". Comptes Rendus de l'Académie des Sciences, Série 2. 5: 209–214.
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22. ^ Thewissen, J.G.M.; Hussain, S.T.; Arif, M. (1997). "New Kohatius (Omomyidae) from the Eocene of Pakistan". Journal of Human Evolution. 32 (5): 473–477. Bibcode:1997JHumE..32..473T. doi:10.1006/jhev.1996.0115. PMID 9169995.
23. ^ a b c d Hartenberger, J.L. (1982). "A review of the Eocene rodents of Pakistan" (PDF). Contributions from the Museum of Paleontology. 26 (2): 19–35.
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27. ^ a b c d e f g h i j k l m n o p q r Gayet, M.; De Broin, F.; Rage, J.C. (1987). "Lower Vertebrates from the Early-Middle Eocene Kuldana Formation of Kohat (Pakistan): Holostei and Teleostei, Chelonia, and Squamata" (PDF). Contributions from the Museum of Paleontology. 27 (7): 151–193.
28. ^ a b c d e f g Murray, A.M.; Thewissen, J.G.M. (2008). "Eocene Actinopterygian Fishes from Pakistan, with the Description of a New Genus and Species of Channid (Channiformes)". Journal of Vertebrate Paleontology. 28 (1): 41–52. doi:10.1671/0272-4634(2008)28[41:EAFFPW]2.0.CO;2.
29. ^ a b Aslan, A.; Thewissen, J.G.M. (1996). "Preliminary evaluation of paleosols and implications for interpreting vertebrate fossil assemblages, Kuldana Formation, Northern Pakistan". Palaeovertebrata. 25 (2–4): 261–277.
30. ^ Joyce, Walter G. (April 2014). "A Review of the Fossil Record of Turtles of the Clade Pan-Carettochelys". Bulletin of the Peabody Museum of Natural History. 55 (1): 3–33. doi:10.3374/014.055.0102. ISSN 0079-032X. S2CID 59382889.
31. ^ a b c d e f g h i j k l m n o p q Shahzad, A.; Khan, J.; Hanif, M.; Baumgartner-Mora, C.; Sarfraz, Y.; Ahmed, K.S.; Riaz, M.T.; Baumgartner, P.O.; ul Hassan Munir, M.; Wazir, A. (2024). "Eocene nannofossils and paleoenvironmental reconstruction of the Kuldana Formation in Yadgar area, Muzaffarabad, northern Pakistan". Palaeoworld. 33 (1): 205–215. Bibcode:2024Palae..33..205S. doi:10.1016/j.palwor.2023.01.003.