{{Short description|Geological formation in Pakistan}} {{Use Pakistani English|date=May 2024}} {{Use dmy dates|date=May 2024}} {{Infobox Rockunit | name = Kuldana Formation | type = Geological formation | age = {{fossil range|Lutetian|Lutetian|Lutetian}} | period = Lutetian | prilithology = | otherlithology = | namedfor = Kuldana Village, Hazara District<ref name=G03/> | namedby = Latif | year_ts = 1970 | location = Gilgit-Baltistan and Punjab, Pakistan | region = Kala Chitta Range | country = {{flag|Pakistan}} | coordinates = {{coord|33|43|N|72|10|E|display=inline,title}} | paleocoordinates ={{coord|14.3|N|68.7|E|display=inline}} | unitof = Subathu Group | subunits = | underlies = Kohat Formation | overlies = Ghazij, Shekhan & Chorgali Formations | thickness = | extent = | area = | map = {{Location map+ | Pakistan | map_caption = | relief = 1 | width = 250 | float = center | places = {{Location map~ | Pakistan | lat_deg = 33 | lon_deg = 72 | mark = Yellow pog.svg | marksize = 12 }} }} }}
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== {{One source section|date=May 2024}} 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, {{convert|200|km|abbr=on}} 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).<ref name=G03>{{Cite journal|last1=Gingerich|first1=P.D.|year=2003|title=Stratigraphic and micropaleontological constraints on the middle Eocene age of the mammal-bearing Kuldana Formation of Pakistan.|url=http://www-personal.umich.edu/~gingeric/PDFfiles/PDG408_ageofkuldanafm_opt.pdf|journal=Journal of Vertebrate Paleontology|volume=23|issue=3|pages=643–651|doi=10.1671/2409|bibcode=2003JVPal..23..643G|archive-date=12 September 2006|access-date=5 July 2025|archive-url=https://web.archive.org/web/20060912134206/http://www-personal.umich.edu/~gingeric/PDFfiles/PDG408_ageofkuldanafm_opt.pdf|url-status=bot: unknown}}</ref>
==Geography== {{One source section|date=May 2024}} 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.<ref name=G03/><ref name=G04>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</ref>
==Stratigraphy== {{One source section|date=May 2024}} 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.<ref name=G03/><ref name =G04/>
==Age== {{One source section|date=May 2024}} 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.<ref name=G03/>
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.<ref name=G03/>
==Paleoenvironment==
[[File: Diacodexis pakistanensis e.jpg|thumb|right|250px|upright|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.<ref name=CTH09>{{Cite journal|last1=Cooper|first1=L.N.|last2=Thewissen|first2=J. G. M.|last3=Hussain|first3=S. T.|year=2009|title=New middle eocene archaeocetes (Cetacea: Mammalia) from the Kuldana formation of northern Pakistan.|doi=10.1671/039.029.0423|journal=Journal of Vertebrate Paleontology|volume=29|issue=4|pages=1289–1299|bibcode=2009JVPal..29.1289C }}</ref>
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.<ref name=CTH09/>
===Ganda Kas===
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.<ref name=Thewissen2001/>
===Banda Daud Shah===
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 ''Gujaratia 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.<ref name=Thewissen2001/>
==Paleobiota==
===Mammals=== {{Paleobiota-key-compact}}
====Arctocyonia====
{| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |''Karakia''<ref name=Thewissen2001/> |''K. longidens'' |H-GSP Loc. 9710, Banda Daud Shah |A single, fragmentary mandible. |The first arctocyonid from Indo-Pakistan. | |- |}
====Artiodactyla==== ===== Cetaceans ===== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |''Ambulocetus''<ref name=CTH09/><ref name=Gingerich2017/> |''A. natans'' |H-GSP Loc. 9209 & 9204 |Several specimens. |An ambulocetid cetacean. |center|150px |- |rowspan=2|''Attockicetus''<ref name=CTH09/><ref name=Gingerich2017>{{Cite journal|last1=Gingerich|first1=P.D.|last2=Heissig|first2=K.|last3=Bebej|first3=R.M.|last4=von Koenigswald|first4=W.|year=2017|title=Astragali of Pakicetidae and other early-to-middle Eocene archaeocetes (Mammalia, Cetacea) of Pakistan: locomotion and habitat in the initial stages of whale evolution|journal=Paläontologische Zeitschrift|doi=10.1007/s12542-017-0362-8|volume=91|issue=4 |pages=601–627|bibcode=2017PalZ...91..601G }}</ref> |''A. praecursor'' |H-GSP Loc. 9204 H-GSP Loc. 9607, Shepherd's Lake |An incomplete cranium, worn teeth, premolars. |A remingtonocetid cetacean. |rowspan=2|center|150px |- |Cf. ''A.'' sp. |H-GSP Loc. 9607, Shepherd's Lake |Two teeth |A remingtonocetid cetacean. |- |Basilosauridae indet.<ref name=West1980/> | |Ganda Kas |Two isolated teeth. |One of the teeth resemble those of ''Ichthyolestes''. | |- |''Gandakasia''<ref name=CTH09/><ref name=West1980/> |''G. potens'' |H-GSP Loc. 58, Ganda Kas<br/>H-GSP Loc. 9607 Shepherd's Lake |A fragmentary mandible |A protocetid cetacean | |- |''Ichthyolestes''<ref name=CTH09/> |''I. pinfoldi'' |H-GSP Loc. 62, Ganda Kas |A single molar |A pakicetid. |center|150px |- |''Nalacetus''<ref name=CTH09/><ref name=Gingerich2017/> |''N. ratimitus'' |H-GSP Loc. 62, Ganda Kas |Fragments of palate, maxilla and teeth |A pakicetid cetacean | |- |rowspan=4|''Pakicetus'' |''P. attocki''<ref name=West1980/> |H-GSP Loc. 62, Ganda Kas |Complete cranial remains. |A pakicetid cetacean |center|150px |- |''P. calcis''<ref name=CTH09/> |H-GSP Loc. 9607 Shepherd's Lake<br/>H-GSP Loc. 9607, Valley E |A partial mandible, palate fragment and teeth |A pakicetid cetacean | |- |''P. chittas''<ref name=CTH09/> |Chorlakki H-GSP Loc. 9607, Shepherd's Lake |Fragments of mandibles |A pakicetid cetacean | |- |''P. inachus''<ref name=TGR87/><ref name=GR81>{{Cite journal|last1=Gingerich|first1=P.D.|last2=Russell|first2=D.E.|year=1981|title=Pakicetus inachus, a new archaeocete (Mammalia, Cetacea) from the early-middle Eocene Kuldana Formation of Kohat (Pakistan).|journal=Contributions from the Museum of Paleontology|url=https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48501/ID352.pdf|volume=25|issue=11|pages=235–246}}</ref> |Chorlakki |A partial skull, mandible and teeth |A pakicetid cetacean |center|150px |- |Protocetidae indet.<ref name=West1980/> | |Ganda Kas |Jaw fragment and two isolated teeth |One of the teeth resemble those of ''Ichthyolestes''. | |- |}
===== Dichobunids ===== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |''Chorlakkia''<ref name=TGR87/> |''C. hassani'' |Chorlakki |A left dentary and multiple teeth |A dichobunid. | |- |Dichobunidae indet.<ref name=TGR87/> | |Chorlakki |Teeth |Teeth distinct from the other known Kuldana dichobunids. One particular tooth might represent a hyopsodontid. | |- |''Dulcidon<ref name="Gingerich2017" />''<ref name=Thewissen1983/><ref name=VanValen1965>{{Cite journal|last1=van Valen|first1=L.|last2=Williams|first2=E.M.|last3=Hussain|first3=S.T.|year=1977|title=Paroxyclaenidae, an Extinct Family of Eurasian Mammals|journal=Journal of Mammalogy|doi=10.2307/1377623|volume=46|issue=3|pages=388–397|jstor=1377623 }}</ref> |''D. gandaensis'' |Ganda Kas & Chorlakki |Two right molar. |A dichobunid | |- |''Pakibune''<ref name=TGR87/> |''P. chorlakkiensis'' |Chorlakki & Lammidhan |Teeth |A moderately sized dichobunid, larger than the other dichobunid taxa from the formation. | |- |} ===== Other artiodactyls =====
{| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |Artiodactyla indet.<ref name=TGR87/> | |Chorlakki |An astragalus and two calcaneum |These fossil remains are considered too large and robust to have belonged to the Chorlakki dichobunids or ''Indohyus''. | |- |''Gobiohyus<ref name="Thewissen2001" />''<ref name=West1980/><ref name=TGR87/> |cf. ''G. orientalis'' |Ganda Kas |A single tooth. |A helohyid. | |- |''Gujaratia''<ref name=Thewissen2001/><ref name="TGR87" /><ref name=Wells1977/><ref name=Wells1983>{{Cite journal|last1=Rautela|first1=A.|last2=Bajpai|first2=S.|year=2023|title=Gujaratia indica, the oldest artiodactyl (Mammalia) from South Asia: new dental material and phylogenetic relationships|journal=Journal of Systematic Palaeontology|doi=10.1080/14772019.2023.2267553|volume=21|issue=1|bibcode=2023JSPal..2167553R }}</ref> |''G. pakistanensis'' |Chorlakki<br/>Lammidhan<br/>Barbora Banda I & II<br/>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''. | |- ||''Indohyus''<ref name=TGR87/> |''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. |200px|center |- |rowspan=2|''Khirtharia<ref name="Thewissen2001" />''<ref name=West1980/><ref name=TGR87/> |''K. dayi'' |Chorlakki, Kalakot & Panoba H-GSP Loc. 62, Ganda Kas |Fragments of maxilla and mandibles; isolated teeth |A raoellid. ''Khirtharia'' is the must abundant artiodactyl at Chorlakki and more common than ''Indohyus'', but the reverse is seen at Kalakot. | rowspan="2" |frameless | |- |''K. major'' |Chorlakki |An isolated tooth |A larger species of raoellid, that would reach about twice the size of ''K. dayi'' and ''I. indirae'', but might also represent an unrelated species of diacodexid or dichobunid. |- |}
====Chiroptera==== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |Chiropteran indet. A<ref name=TGR87/><ref name=Russell1981/> | |Chorlakki |A molar | | |- |Chiropteran indet. B<ref name=TGR87/><ref name=Russell1981/> | |Chorlakki |Two fragmentary molars |Resemble that of ''Palaeochiropteryx'', and may belong to a relatively large bat. | |- |}
====Eulipotyphla==== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |''Pakilestes''<ref name=TGR87>{{Cite journal|last1=Thewissen|first1=J.G.M.|last2=Gingerich|first2=P.D.|last3=Russell|first3=D.E.|year=1987|title=ARTIODACTYLA AND PERISSODACTYLA (MAMMALIA) FROM THE EARLY-MIDDLE EOCENE KULDANA FORMATION OF KOHAT (PAKISTAN)|journal=Contributions from the Museum of Paleontology|url=https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48532/ID385.pdf|volume=27|issue=10|pages=247–274}}</ref><ref name=Russell1981/><ref name=Ziegler2009>{{Cite journal|last1=Ziegler|first1=R.|year=2009|title=Plesiosoricids from early Oligocene fissure fillings in South Germany, with remarks on plesiosoricid phylogeny|journal=Acta Palaeontologica Polonica|doi=10.4202/app.2008.0061|volume=54|issue=3|pages=365–371|doi-access=free}}</ref> |''P. lathrius'' |Chorlakki |Two molars and a premolar |An insectivore mammal of uncertain affinities, possibly a plesiosoricid soricomorph. | |- |''Perizalambdodon''<ref name=Thewissen2005>{{Cite journal|last1=Thewissen|first1=J.G.M.|last2=Bajpai|first2=S.|last3=Hussain|first3=S.T.|year=2005|title=New insectivorous placentals from the Eocene of Pakistan|journal=Journal of the Palaeontological Society of India|volume=50|issue=2|pages=37–41}}</ref> |''P. punjabiensis'' |H-GSP Loc. 9610, Jhalar |A single molar |An indeterminate lipotyphlan. | |- |''Seia''<ref name=TGR87/><ref name=Russell1981>{{Cite journal|last1=Russell|first1=D. E.|last2=Gingerich|first2=P. D.|year=1981|title=Lipotyphla, Proteutheria(?), and Chiroptera (Mammalia) from the Early-Middle Eocene Kuldana Formation of Kohat (Pakistan)|journal=Contributions from the Museum of Paleontology|url=https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48504/ID355.pdf;sequence=2|volume=25|issue=14|pages=277–287}}</ref> |''S. shahi'' |Chorlakki |Two molars |An erinaceomorph, probably representing a new family. | |- |}
====Herpetotheriidae==== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |Herpetotheriinae indet.<ref name=Thewissen2001>{{Cite journal|last1=Thewissen|first1=J.G.M.|last2=Williams|first2=E.M.|last3=Hussain|first3=S.T.|year=1977|title=Eocene mammal faunas from Northern Indo-Pakistan |journal=Contributions from the Museum of Paleontology|doi=10.1671/0272-4634(2001)021[0347:EMFFNI]2.0.CO;2|volume=21|issue=2|pages=347–366|url=http://doc.rero.ch/record/16008/files/PAL_E3764.pdf }}</ref> | |H-GSP Loc. 62, Ganda Kas |A right molar. | | |- |}
====Hyaenodonta==== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |rowspan=2|''Paratritemnodon''<ref name=TGR87/><ref name=Thewissen2001/> |''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====
{| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |Mesonychidae indet.<ref name=West1980>{{Cite journal|last1=West|first1=R.M.|year=1980|title=Middle Eocene Large Mammal Assemblage with Tethyan Affinities, Ganda Kas Region, Pakistan|journal=Journal of Paleontology|volume=54|issue=3|pages=508–533}}</ref> | |Ganda Kas |A single premolar |Belongs to a small mesonychid, with similarities to ''Hapalodectes''. | |- |}
====Perissodactyla==== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |rowspan=2|''Anthracobune''<ref name=Wells1983/><ref name=Wells1977>{{Cite journal|last1=Gingerich|first1=P. D.|year=1977|title=A small collection of fossil vertebrates from the Middle Eocene Kuldana and Kohat Formations of Punjab (Pakistan)|journal=Contributions from the Museum of Paleontology|url=https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48485/ID335.pdf?sequence=2|volume=24|issue=18|pages=190–203}}</ref><ref name=Wells2014>{{Cite journal|last1=Wells|first1=N.A.|last2=Gingerich|first2=P. D.|year=1983|title=Review of Eocene Anthracobunidae (Mammalia, Proboscidea) with a new genus and species, Jozaria palustris, from the Kuldana Formation of Kohat, Pakistan|journal=Contributions from the Museum of Paleontology|url=https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48515/ID367.pdf|volume=26|issue=7|pages=117–139}}</ref><ref name=Cooper2014>{{Cite journal|last1=Cooper|first1=L.N.|last2=Seiffert|first2=E.R.|last3=Clementz|first3=M.|last4=Madar|first4=S.I.|last5=Bajpal|first5=S.|last6=Hussain|first6=S.T.|last7=Thewissen|first7=J.G.M.|year=2014|title=Anthracobunids from the Middle Eocene of India and Pakistan Are Stem Perissodactyls|journal=PLOS ONE|doi=10.1371/journal.pone.0109232|volume=9|issue=10|article-number=e109232 |doi-access=free |pmid=25295875 |pmc=4189980 |bibcode=2014PLoSO...9j9232C }}</ref> |''A. pinfoldi''<ref name=West1980/> |Upper Member : Lammidhan & Ganda Kas |Relatively complete cranial remains and isolated teeth. |The largest anthracobunid in the formation. |center|150px |- |''A. wardi''<ref name=TGR87/> |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''. |center|150px |- |''"Forstercooperia"''<ref name=TGR87/> |''"F." jigniensis'' |Chorlakki |Teeth |A paraceratheriid. | |- |Isectolophidae indet.<ref name=TGR87/> | |Chorlakki |A tooth | | |- |''Jozaria''<ref name=Wells1983/><ref name=Wells2014/> |''J. palustris'' |Upper Member |Several teeth |An anthracobunid. | |- |cf. ''Kalakotia''<ref name=Thewissen2001/> |''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''<ref name=Mass2001>{{Cite journal|last1=Mass|first1=M.C.|last2=Hussain|first2=S.T.|last3=Leinders|first3=J.J.M.|year=2001|title=A New Isectolophid Tapiromorph (Perissodactyla, Mammalia) from the Early Eocene of Pakistan|journal=Journal of Paleontology|volume=75|issue=2|pages=407–417|doi=10.1666/0022-3360(2001)075<0407:ANITPM>2.0.CO;2 }}</ref> |''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. |center|150px |- |''Obergfellia''<ref name=TGR87/><ref name=Wells1983/> |''O. occidentalis'' |Ganda Kas |Several mandibles. |An anthracobunid. Includes most of the material formerly assigned to ''Pilgrimella pilgrimi''. |center|150px |- |''Palaeosyops''<ref name=TGR87/><ref name=West1980/><ref name=Thewissen2001/><ref name=Missiaen2011>{{Cite journal|last1=Missiaen|first1=P.|last2=Gunnell|first2=G.F.|last3=Gingerich|first3=P.D.|year=1977|title=New Brontotheriidae (Mammalia, Perissodactyla) from the Early and Middle Eocene of Pakistan with implications for mammalian paleobiogeography |journal=Journal of Paleontology|volume=85|issue=4|pages=665–677|doi=10.1666/10-087.1 }}</ref> |''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 primitive brontothere. Also present in the Baska Formation. | |- |''Pakotitanops''<ref name=West1980/><ref name=Thewissen2001/> |''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==== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |Cf. ''Agerinia''<ref name=TGR87/><ref name=Russell1987a/> |Cf. ''A. sp.'' |Chorlakki |A tooth. |A notharctid, with similarities to ''A. roselli''. | |- |''Jattadectes''<ref name=Thewissen2001/> |''J. mamikheli'' |H-GSP Loc. 9712, Banda Daud Shah |A premolar and an incisor |The first plesiadapid from Indo-Pakistan. | |- |rowspan=3|''Kohatius''<ref name=Gunnell2008/><ref name=Thewissen1997>{{Cite journal|last1=Thewissen|first1=J.G.M.|last2=Hussain|first2=S.T.|last3=Arif|first3=M.|year=1997|title=New Kohatius (Omomyidae) from the Eocene of Pakistan|journal=Journal of Human Evolution|volume=32|issue=5 |pages=473–477|doi=10.1006/jhev.1996.0115 |pmid=9169995 |bibcode=1997JHumE..32..473T }}</ref> |''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''. | |- |rowspan=2|''Panobius''<ref name=TGR87/><ref name=Thewissen2001/><ref name=Gunnell2008>{{Cite journal|last1=Gunnell|first1=G.F.|last2=Gingerich|first2=P.D.|last3=Ul-Haq|first3=M.|last4=Bloch|first4=J.I.|last5=Khan|first5=I.H.|last6=Clyde|first6=W.C.|year=2008|title=New Primates (Mammalia) From The Early and Middle Eocene Of Pakistan And Their Paleobiogeographical Implications|journal=Contributions from the Museum of Paleontology|volume=32|issue=1|pages=1–14}}</ref><ref name=Russell1987a>{{Cite journal|last1=Russell|first1=D.E.|last2=Gingerich|first2=P.D.|year=1987|title=Nouveaux primates de l'Éocène du Pakistan|journal=Comptes Rendus de l'Académie des Sciences, Série 2|url=https://gallica.bnf.fr/ark:/12148/bpt6k5664623d/f215.item.r=Panobius.zoom|volume=5|pages=209–214}}</ref> |''P. afridi'' |Chorlakki |Two isolated teeth. |An adapid. | |- |''P. amplior'' |Either from Chorlakki,<ref name=Gunnell2008/> or from H-GSP Loc. 9712, Banda Daud Shah.<ref name=Thewissen2001/> |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''<ref name=Thewissen2001/> |''P. oligocollis'' |H-GSP Loc. 225, Jhalar |A premolar and an incisor |A carpolestid. | |- |}
====Rodentia==== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |rowspan=4|''Birbalomys''<ref name=TGR87/><ref name=Thewissen2001/><ref name=Hartenberger1982>{{Cite journal|last1=Hartenberger|first1=J.L.|year=1982|title=A review of the Eocene rodents of Pakistan|journal=Contributions from the Museum of Paleontology|url=https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48510/ID362.pdf?sequence=2|volume=26|issue=2|pages=19–35}}</ref> |''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''<ref name=Gingerich1979/> |Shekhan, Chorlakki H-GSP Loc. 57 & 144, Ganda Kas | |A chappatimyid. Formerly ''Saykanomys''. | |- |rowspan=2|''Chapattimys''<ref name=TGR87/><ref name=Hartenberger1982/> |''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''<ref name=TGR87/><ref name=Hartenberger1982/> |''G. asifi'' |H-GSP Loc. 62, Ganda Kas; Chorlakki |Several teeth |A rare chappatimyid. | |- |Paramyidae indet.<ref name=Thewissen1983/><ref name=Leinders1999/> | |Barbora Banda I |Teeth | | |- |rowspan=2|Cf. ''Petrokozlovia''<ref name=TGR87/><ref name=Hartenberger1982/> |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''<ref name=Gingerich1979>{{Cite journal|last1=Gingerich|first1=P.D.|last2=Russell|first2=D.E.|last3=Sigogneau-Russell|first3=D.|last4=Hartenberger|first4=J.-L.|last5=Ibrahim Shah|first5=W.|last6=Hassan|first6=M.|last7=Rose|first7=K.D.|last8=Holt Ardrey|first8=R.|year=1979|title=Reconnaissance survey and vertebrate paleontology of some Paleocene and Eocene formations in Pakistan|journal=Contributions from the Museum of Paleontology|volume=25|issue=5|pages=105–116}}</ref> |Cf. ''T.'' sp. |Chorlakki | |A Tamquammyidae. | |- |rowspan=2|Tamquammyidae indet.<ref name=Thewissen1983/><ref name=Thewissen2001/><ref name=Leinders1999>{{Cite journal|last1=Leinders|first1=J.J.M.|last2=Arif|first2=M.|last3=de Bruijn|first3=H.|last4=Hussain|first4=S.T.|last5=Wessels|first5=W.|year=1999|title= Tertiary continental deposits of northwestern Pakistan and remarks on the collision between the Indian and Asian plates|journal=Deinsea|volume=7|pages=199–213}}</ref> |spe. indet. 1 |Barbora Banda I |Teeth | | |- |spe. indet. 2 |Barbora Banda I |Teeth | | |- |}
====Tillodontia==== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |rowspan=2|''Basalina''<ref name=TGR87/><ref name=Lucas>{{Cite journal|last1=Lucas|first1=S.G.|last2=Schoch|first2=R.M.|year=1981|title=Basalina, a Tillodont from the Eocene of Pakistan|url=https://www.zobodat.at/pdf/Mitt-Bayer-Staatsslg-Pal-hist-Geol_21_0089-0095.pdf|journal= Mitt. Bayer. Staatsslg. Paläont. Hist. Geol. |volume=21|issue= |pages=89–95}}</ref> |''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 ==== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |Mammalia indet.<ref name=Thewissen1983/> | |Barbora Banda II |A single, large incisor. |May belong to a small artiodactyl. | |- |}
=== Fish === ==== Actinopterygians ==== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |Acanthopterygii indet.<ref name=Gayet,1987/> | |Chorlakki & Shekhan Nala |Several isolated teeth. |Some of the teeth might belong to an Osteoglossiforme indeterminate. | |- |Amiidae indet.<ref name=Murray2008/> | |H-GSP Loc. 56, Ganda Kas redbeds |Partial right premaxilla with associated teeth. |From predominantly marine deposits. | |- |''Anchichanna''<ref name=Murray2008/> |''S. kuldanensis'' |H-GSP Loc. 62, Ganda Kas |Several relatively complete cranial remains. |A snakehead. | |- |Ariidae indet.<ref name=Gayet,1987/> | |Chorlakki |Several abdominal vertebra. |Despite being primarily marine, ariid catfish are known to frequently enter freshwater environments. | |- |Bagridae indet.<ref name=Gayet,1987/> | |Chorlakki |An angular bone, a cleithrum and pectoral spines. | | |- |Cf. Bagridae indet.<ref name=Murray2008/> | |H-GSP Loc. 62, Ganda Kas |Fragments of the skull and spines. |Might represent several species of catfish. | |- |Clariidae indet.<ref name=Gayet,1987/> | |Chorlakki & Shekhan Nala |An articular bone and pectoral spines. |The low angular resemble that of ''Heterobranchus''. | |- |Cyprinidae indet.<ref name=Gayet,1987/> | |Chorlakki |A pharyngeal tooth. | | |- |Cyprinodontidae indet.<ref name=Gayet,1987/> | |Chorlakki |Several teeth and a quadrate bone. |The teeth are similar to those of ''Aphanius''. | |- |Eotrigonodontidae indet.<ref name=Murray2008>{{Cite journal|last1=Murray|first1=A.M.|last2=Thewissen|first2=J.G.M.|year=2008|title=Eocene Actinopterygian Fishes from Pakistan, with the Description of a New Genus and Species of Channid (Channiformes)|journal=Journal of Vertebrate Paleontology|volume=28|issue=1|pages=41–52|doi=10.1671/0272-4634(2008)28[41:EAFFPW]2.0.CO;2 }}</ref> | |H-GSP Loc. 229 & 9607, Shepherd's Lake |Two teeth. |Tentatively referred to the Mesozoic genera ''Hadrodus'' or ''Stephanodus''. | |- |''Lepisosteus''<ref name=Gayet,1987/> |''L.'' sp. |Chorlakki |A tooth and a scale. |Appears to be closely related or identical to the modern genus ''Lepisosteus osseus''. | |- |''Percalates''<ref name=Gayet,1987>{{Cite journal|last1=Gayet|first1=M.|last2=De Broin|first2=F.|last3=Rage|first3=J.C.|year=1987|title=Lower Vertebrates from the Early-Middle Eocene Kuldana Formation of Kohat (Pakistan): Holostei and Teleostei, Chelonia, and Squamata|journal=Contributions from the Museum of Paleontology|url=https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48528/ID381.pdf?sequence=2|volume=27|issue=7|pages=151–193}}</ref><ref>{{Cite journal |last=Rix |first=Alan |date=2022 |title=Coal, bees and fossils: The history and significance of the Redbank plains formation fossil sites, south East Queensland |url=https://search.informit.org/doi/abs/10.3316/informit.237428598989426 |journal=The Proceedings of the Royal Society of Queensland |volume=131 |pages=131–144 }}</ref> |''P. antiquus'' (=''Macquaria antiquus'')<ref name="Gayet,1987" /> |Chorlakki |Various bone fragments and isolated spines |A percalatid. | |- |Osteoglossidae indet.<ref name=Gayet,1987/> | |Scales : Chorlakki & Shekhan Nala Maxilla : H-GSP Loc. 9611 |A maxilla, several scales of various size and shape. | | |- |Perciformes indet.<ref name=Murray2008/> | |H-GSP Loc. 62, Ganda Kas |Isolated fin spine. |Distinct from ''Macquaria antiquus''. | |- |Pycnodontoidea indet.<ref name=CTH09/><ref name=Murray2008/> | |H-GSP Loc. 9206 & 9608, Ganda Kas<br/>H-GSP Loc. 9607, Shepherd's Lake |Several isolated teeth and a jaw fragment. |From predominantly marine deposits. | |- |Siluriforme indet.<ref name=Gayet,1987/> | |Chorlakki |A basioccipital. | | |- |Teleostei indet.<ref name=Murray2008/> | |H-GSP Loc. 62, Ganda Kas |Isolated teeth. |From predominantly freshwater deposits, possibly representing several species. | |- |Cf. ''Varohstichthys''<ref name=Gayet,1987/> |Cf. ''V.'' sp. |Chorlakki |A pharyngeal tooth. |A Cyprinidae. | |- |}
==== Chondrichthyans ==== {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |Selachimorpha indet.<ref name=CTH09/> | |H-GSP Loc. 9607, Shepherd's Lake |Teeth. | | |- |}
===Reptiles=== ==== Crocodilians ====
{| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |Crocodilia indet.<ref name=Thewissen1983/><ref name=Aslan1996>{{Cite journal|last1=Aslan|first1=A.|last2=Thewissen|first2=J.G.M.|date=1996|title=Preliminary evaluation of paleosols and implications for interpreting vertebrate fossil assemblages, Kuldana Formation, Northern Pakistan|journal=Palaeovertebrata|language=en|volume=25|issue=2–4|pages=261–277}}</ref> | |Barbora Banda II & H-GSP Loc. 62, Ganda Kas | | | |- |}
====Squamates====
{| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |''Tinosaurus''<ref name=Gayet,1987/> |''T.'' sp. |Chorlakki |Two isolated teeth. |Similarities can be observed with Eocene species of ''Tinosaurus'' from North America and China. | |- |Sauria indet.<ref name=Gayet,1987/> | |Chorlakki |A fragmentary vertebra. | | |- |Boidae indet.<ref name=Gayet,1987/> | |Chorlakki |Five fragmentary vertebrae. | | |- |Erycinae indet.<ref name=Gayet,1987/> | |Chorlakki |Two fragmentary vertebrae. |The material probably represent a new genus and species of sand boa. | |- |Boinae indet.<ref name=Gayet,1987/> | |Chorlakki |Two fragmentary vertebrae. |The material probably represent a new genus and species of boa resembling the Eocene genera ''Paleryx'' and ''Palaeopython''. | |- |}
====Turtles====
{| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- |Trionychinae indet.<ref name=Gayet,1987/> | |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''"<ref name=Gayet,1987/> |"''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.<ref name="Joyce2014">{{Cite journal|last=Joyce|first=Walter G.|date=April 2014|title=A Review of the Fossil Record of Turtles of the Clade Pan-Carettochelys|url=http://www.bioone.org/doi/abs/10.3374/014.055.0102|journal=Bulletin of the Peabody Museum of Natural History|language=en|volume=55|issue=1|pages=3–33|doi=10.3374/014.055.0102|s2cid=59382889 |issn=0079-032X|url-access=subscription}}</ref> | |- |}
=== Mollusca === {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member/Microfacies ! Material ! Notes ! Image |- |Bivalvia indet.<ref name=Shahzad2023/> | |Molluscan microfacies<br/>Ostracode wackestone |"Oyster" shell fragments |Composes more than half of the molluscan microfacies. | |- |Gastropoda indet.<ref name=Shahzad2023/> | |Molluscan microfacies<br/>Ostracode wackestone | | | |- |''Planorbis''<ref name=CTH09/><ref name=Gingerich1979/><ref name=Aslan1996/> |''P.'' sp. |Chorlakki<br/>Shekhan Nala<br/>H-GSP Loc. 62, Ganda Kas<br/>H-GSP Loc. 9607, Shepherd's Lake | |A freshwater snail. | |- |}
=== Plants === {{Paleobiota-key-compact}} {| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Locality/Member ! Material ! Notes ! Image |- style="background:#D1FFCF;" |Cf. ''Ajunginucilla''<ref name=Thewissen1983/> |Cf. ''C.'' sp. |Barbora Banda I |Seeds. |A sage. | |- style="background:#D1FFCF;" |''Celtis''<ref name=Thewissen1983>{{Cite journal|last1=Thewissen|first1=J.G.M.|last2=Russell|first2=D.E.|last3=Gingerich|first3=P.D.|last4=Hussain|first4=S.T.|year=1983|title=A new dichobunid artiodactyl (Mammalia) from the Eocene of North-West Pakistan|journal=Proceedings of the Koniklijke Nederlandse Akademie van Wetenschappen|volume=86|issue=2|pages=153–180}}</ref> |''C.'' sp.<ref name=Thewissen1983/> |Barbora Banda I |Seeds. |A hackberry. | |- style="background:#D1FFCF;" |''Chara''<ref name=Thewissen1983/> |''C.'' sp. |Barbora Banda I |Seeds. |A charophyte. | |- |}
=== Other microfossils ===
{| class="wikitable" align="center" width="75%" |- ! Name ! Species ! Microfacies ! Material ! Notes ! Image |- |''Assilina''<ref name=Shahzad2023/> |''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''<ref name=Shahzad2023/> |''B. sparsus'' |Ostracode wackestone (Q-2)<br/>Nummulitic wacke-packstone (Q-15) | | | |- |Brachiopoda indet.<ref name=Shahzad2023/> | |Molluscan microfacies | | | |- |rowspan=4|''Coccolithus''<ref name=Shahzad2023>{{Cite journal|last1=Shahzad|first1=A.|last2=Khan|first2=J.|last3=Hanif|first3=M.|last4=Baumgartner-Mora|first4=C.|last5=Sarfraz|first5=Y.|last6=Ahmed|first6=K.S.|last7=Riaz|first7=M.T.|last8=Baumgartner|first8=P.O.|last9=ul Hassan Munir|first9=M.|last10=Wazir|first10=A.|year=2024|title=Eocene nannofossils and paleoenvironmental reconstruction of the Kuldana Formation in Yadgar area, Muzaffarabad, northern Pakistan|journal=Palaeoworld|volume=33|issue=1 |pages=205–215|doi=10.1016/j.palwor.2023.01.003 |bibcode=2024Palae..33..205S }}</ref> |''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)<br/>Ostracode wackestone (Q-2)<br/>Nummulitic wacke-packstone (Q-15) | |Typical of warm, low-latitude waters | |- |''Cyclicargolithus''<ref name=Shahzad2023/> |''C. luminis'' |Nummulitic wacke-packstone (Q-15) | | | |- |Echinodermata indet.<ref name=Shahzad2023/> | |Molluscan microfacies |Fragments | | |- |rowspan=3|''Fasciculithus''<ref name=Shahzad2023/> |''F. lillianae'' |Molluscan microfacies (Q-13) | | | |- |''F. tympaniformis'' |Molluscan microfacies (Q-13) | | | |- |''F. clinatus'' |Molluscan microfacies (Q-13)<br/>''Assilina'' wacke-packstone (Q-17) | | | |- |''Lockhartia''<ref name=Shahzad2023/> |''L.'' spp. |''Assilina'' wacke-packstone |A rotaliid |Present in minor quantity in the ''Assilina'' wacke-packstone. | |- |''Neococcolithes''<ref name=Shahzad2023/> |''N. protenus'' |Nummulitic wacke-packstone (Q-15) | | | |- |''Nummulites''<ref name=Shahzad2023/> |''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.<ref name=Shahzad2023/> | |Ostracode wackestone | | | |- |''Rhomboaster''<ref name=Shahzad2023/> |''R. brameletti'' |Ostracode wackestone (Q-2)<br/>Molluscan microfacies (Q-13)<br/>Lime mudstone (Q-14) | | | |- |rowspan=4|''Sphenolithus''<ref name=Shahzad2023/> |''S. anarrhopus'' |Lime mudstone (Q-1, Q-14)<br/>Ostracode wackestone (Q-2) | | | |- |''S. moriformis'' |Molluscan microfacies (Q-13) | | | |- |''S. primus'' |Ostracode wackestone (Q-2)<br/>Molluscan microfacies (Q-13) | | | |- |''S.'' sp. |Lime mudstone (Q-14) |Typical of warm waters | | |- |rowspan=4|''Tribrachiatus''<ref name=Shahzad2023/> |''T. absidatus'' |''Assilina'' wacke-packstone (Q-22)<br/>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)<br/>Molluscan microfacies (Q-13) | | | |- |''Zeughrabdotus''<ref name=Shahzad2023/> |''Z. sigmoides'' |Nummulitic wacke-packstone (Q-15) | |More typical of colder waters. | |- |}
==References==
{{Reflist}}
Category:Lutetian Stage Category:Paleontology in Pakistan Category:Eocene life Category:Natural history of Pakistan Category:Eocene animals of Asia