Kuldana Formation
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The Kuldana Formation is a
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
-bearing
geological formation A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics (lithology) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock expo ...
of
Lutetian The Lutetian is, in the geologic timescale, a stage (stratigraphy), stage or age (geology), age in the Eocene. It spans the time between . The Lutetian is preceded by the Ypresian and is followed by the Bartonian. Together with the Bartonian it ...
(Early
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
) age which crops out in northern
Pakistan Pakistan, officially the Islamic Republic of Pakistan, is a country in South Asia. It is the List of countries and dependencies by population, fifth-most populous country, with a population of over 241.5 million, having the Islam by country# ...
. The abundant fossil remains were deposited by rivers and estuaries crossing an arid to semi-arid environment, between several
marine transgression A marine transgression is a geologic event where sea level rises relative to the land and the shoreline moves toward higher ground, resulting in flooding. Transgressions can be caused by the land sinking or by the ocean basins filling with water ...
s. Its fossil fauna is best known for the early
cetacean Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
s ''
Indohyus ''Indohyus'' (Meaning "India's pig" from the Greek words ''Indos'', "from India" and ''hûs'', "pig") is an extinct genus of artiodactyl known from Eocene fossils in Asia. This small chevrotain-like animal found in the Himalayas is among the clo ...
'', ''
Pakicetus ''Pakicetus'' (meaning 'whale from Pakistan') is an extinct genus of amphibious cetacean of the family Pakicetidae, which was endemic to the Indian Subcontinent during the Ypresian (early Eocene) period, about 50 million years ago. It was a w ...
'' and ''
Ambulocetus ''Ambulocetus'' (Latin ''ambulare'' "to walk" + ''cetus'' "whale") is a genus of early Semiaquatic, amphibious cetacean from the Kuldana Formation in Pakistan, roughly 48 or 47 million years ago during the Early Eocene (Lutetian). It contains o ...
'', that helped to shed a new light on the
evolution of whales The evolution of cetaceans is thought to have begun in the Indian subcontinent from even-toed ungulates (Artiodactyla) 50 million years ago (mya) and to have proceeded over a period of at least 15 million years. Cetaceans are fully aquatic mam ...
, but it also features a large number of early
ungulate Ungulates ( ) are members of the diverse clade Euungulata ("true ungulates"), which primarily consists of large mammals with Hoof, hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined ...
s,
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s and
primate Primates is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
s.


History

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, 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 The Indus ( ) is a transboundary river of Asia and a trans-Himalayas, Himalayan river of South Asia, South and Central Asia. The river rises in mountain springs northeast of Mount Kailash in the Western Tibet region of China, flows northw ...
while providing no other indicators in how the two differed. Other studies treating the two formations as distinct include Leinders (1999) and Thewissen (2001).


Geography

Outcrops of the Kuldana Formation are found in the form of isoclinal folds across northern
Pakistan Pakistan, officially the Islamic Republic of Pakistan, is a country in South Asia. It is the List of countries and dependencies by population, fifth-most populous country, with a population of over 241.5 million, having the Islam by country# ...
'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.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


Stratigraphy

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.


Age

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 In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
(Early Eocene) or
Lutetian The Lutetian is, in the geologic timescale, a stage (stratigraphy), stage or age (geology), age in the Eocene. It spans the time between . The Lutetian is preceded by the Ypresian and is followed by the Bartonian. Together with the Bartonian it ...
(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. 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.


Paleoenvironment

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 ''Planorbis'' is a genus of air-breathing freshwater snails, aquatic animal, aquatic pulmonate gastropod molluscs in the family (biology), family Planorbidae, the sheep, ram's horn snails, or planorbids. All species in this genus have sinistral o ...
'', 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. This mix of fresh- and saltwater habitats was eventually followed by a
marine transgression A marine transgression is a geologic event where sea level rises relative to the land and the shoreline moves toward higher ground, resulting in flooding. Transgressions can be caused by the land sinking or by the ocean basins filling with water ...
that characterizes the geology of the uppermost layers of the formation.


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.


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 Artiodactyls are placental mammals belonging to the order Artiodactyla ( , ). Typically, they are ungulates which bear weight equally on two (an even number) of their five toes (the third and fourth, often in the form of a hoof). The other t ...
'' Gujaratia pakistanensis'' and the tapiroid
perissodactyl Perissodactyla (, ), or odd-toed ungulates, is an order of Ungulate, ungulates. The order includes about 17 living species divided into three Family (biology), families: Equidae (wild horse, horses, Asinus, asses, and zebras), Rhinocerotidae ( ...
'' Karagalax mamikhelensis''; the presence of adapids and arctocyonids in the area indicates that it probably had an older age than the Ganda Kas fauna.


Paleobiota


Mammals


Arctocyonia


Artiodactyla


= Cetaceans

=


= Dichobunids

=


= Other artiodactyls

=


Chiroptera


Eulipotyphla


Herpetotheriidae


Hyaenodonta


Mesonychia


Perissodactyla


Primates


Rodentia


Tillodontia


Other mammals


Fish


Actinopterygians


Chondrichthyans


Reptiles


Crocodilians


Squamates


Turtles


Mollusca


Plants


Other microfossils


References

{{Reflist Lutetian Stage Paleontology in Pakistan Eocene life Natural history of Pakistan Eocene animals of Asia