Rodhocetus Balochistanensis
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''Rodhocetus'' (from ''Rodho'', the geological
anticline In structural geology, an anticline is a type of fold that is an arch-like shape and has its oldest beds at its core, whereas a syncline is the inverse of an anticline. A typical anticline is convex up in which the hinge or crest is the ...
at the type locality, and ''cetus'', Latin for whale) is an extinct genus of
protocetid Protocetidae, the protocetids, form a diverse and heterogeneous group of extinct cetaceans known from Asia, Europe, Africa, South America, and North America. Description There were many genera, and some of these are very well known (e.g., ''Ro ...
early whale known from the
Lutetian The Lutetian is, in the geologic timescale, a stage or 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 is sometimes referred to as the Midd ...
of Pakistan. The best-known protocetid, ''Rodhocetus'' is known from two partial skeletons that taken together give a complete image of an Eocene whale that had short limbs with long hands and feet that were probably webbed and a sacrum that was immobile with four partially fused sacral vertebrae. It is one of several extinct
whale Whales are a widely distributed and diverse group of fully aquatic placental marine mammals. As an informal and colloquial grouping, they correspond to large members of the infraorder Cetacea, i.e. all cetaceans apart from dolphins and ...
genera that possess land mammal characteristics, thus demonstrating the evolutionary transition from land to sea.


Description

left, Size of ''Rodhocetus'' relative to a human. ''Rodhocetus'' was a small whale measuring long. Throughout the 1990s, a close relationship between cetaceans and
mesonychid Mesonychia ("middle claws") is an extinct taxon of small- to large-sized carnivorous ungulates related to artiodactyls. Mesonychids first appeared in the early Paleocene, went into a sharp decline at the end of the Eocene, and died out entirely ...
s, an extinct group of cursorial, wolf-like ungulates, was generally accepted based on morphological analyses. In the late 1990s, however, cladistic analyses based on molecular data clearly placed Cetacea within the Artiodactyla near the hippopotamus. One of the diagnostic characteristics of artiodactyls is the double-pulley
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
(heel bone), and palaeontologists, unconvinced by the data from the labs, set themselves out to find archaeocete single-pulley heel bones. Hind legs from three archaeocete species were recovered within a few years, among them those of ''Rodhocetus balochistanensis'', and all three had double-pulley heel bones, thus settling the cladistic controversy. Through a
principal components analysis Principal component analysis (PCA) is a popular technique for analyzing large datasets containing a high number of dimensions/features per observation, increasing the interpretability of data while preserving the maximum amount of information, and ...
demonstrated that ''Rodhocetus'' had trunk and limb proportions similar to the
Russian desman The Russian desman (''Desmana moschata'') (russian: выхухоль ''vykhukhol'') is a small semiaquatic mammal that inhabits the Volga, Don and Ural River basins in Russia, Ukraine and Kazakhstan. It constructs burrows into the banks of ponds ...
, a foot-powered swimmer using its tail mainly as a rudder. From this Gingerich concluded that ''Rodhocetus'' was swimming mostly at the surface by alternate strokes of its hind feet, and that it was insulated by fur rather than blubber, as are ''
Dorudon ''Dorudon'' ("spear-tooth") is a genus of extinct basilosaurid ancient whales that lived alongside ''Basilosaurus'' 40.4 to 33.9 million years ago in the Eocene. It was a small whale, with ''D. atrox'' measuring long and weighing . ''Dorudon'' ...
'' and modern cetaceans, which made it buoyant and incapable of deep diving.


''R. kasrani''

The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
of ''R. kasrani'', GSP- UM 3012 found in 1992, was described by : a cranium with two dentaries, most of the vertebral column as far as the anterior tail (C2–C7; T1–13; L1–6, S1–4, Ca1–4), most ribs, parts of the sternum, both hip bones, and a left femur. Gingerich et al. 1994 referred a specimen collected in 1981, GSP-UM 1852 two dentaries with teeth, to ''R. kasrani''. The body mass of the holotype has been estimated between based on different techniques.
Derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
traits in ''R. kasrani'', relative to older archaeocetes such as ''Pakicetus'', includes high-crowned cheek teeth, larger auditory bullae, larger mandibular foramen, and mandibular canals. The higher neural spines and shorter femur (60–70%) distinguishes ''Rodhocetus'' from the more primitive ''Ambulocetus''. The convex posterior surface of the exoccipital, shorter cervical vertebrae, and unfused sacral vertebrae distinguishes ''R. kasrani'' from ''Indocetus''. In contrast to later archaeocetes such as ''Protocetus'' and later cetaceans, ''Rodhocetus'' retains external nares above upper canines, high neural spines on anterior thoracic vertebrae, and four sacral vertebrae with sacroiliac joints similar to those in land-mammals (suggesting a hip joint that could support the body weight.) Several cranial features identifies ''R. kasrani'' as an archaeocete: both the premaxillae and the dentaries are elongated, the frontal shield is wide, and the nuchal crest is high. The
auditory bulla The tympanic part of the temporal bone is a curved plate of bone lying below the squamous part of the temporal bone, in front of the mastoid process, and surrounding the external part of the ear canal. It originates as a separate bone (tympanic ...
e are large and dense but, there are no associated
pterygoid fossa The pterygoid fossa is an anatomical term for the fossa formed by the divergence of the lateral pterygoid plate and the medial pterygoid plate of the sphenoid bone. Structure The lateral and medial pterygoid plates (of the pterygoid process of ...
e or air sinuses. The mandibular foramina are large with a pan bone long and high. The specific name ''kasrani'' comes from
Qaisrani Qaisrani is a Baloch clan of Rind tribe. They live mostly in Balochistan, Pakistan Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in South Asia. It is the world's List of countries an ...
, the Baloch tribe inhabiting the type locality. The protocetid ''
Qaisracetus ''Qaisracetus'' is an extinct Protocetidae, protocetid Archaeoceti, early whale known from the Eocene (Lutetian, ) of Baluchistan, Pakistan (, paleocoordinates ). Etymology The genus is named after the Qaisrani Baloch tribe which assisted Ginge ...
'' is also named after them.


''R. balochistanensis''

The fossil remains of ''R. balochistanensis'' were found in eastern
Balochistan Balochistan ( ; bal, بلۏچستان; also romanised as Baluchistan and Baluchestan) is a historical region in Western and South Asia, located in the Iranian plateau's far southeast and bordering the Indian Plate and the Arabian Sea coastline. ...
,
Pakistan Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in South Asia. It is the world's List of countries and dependencies by population, fifth-most populous country, with a population of almost 24 ...
in 2001 by Philip Gingerich. Dating from about 47 million years ago, they are one of a series of recent discoveries, including the
pakicetid Pakicetidae ("Pakistani whales") is an extinct family of Archaeoceti (early whales) that lived during the Early Eocene in Pakistan. Description described the first pakicetid, '' Ichthyolestes'', but at the time they did not recognize it as a ce ...
s, which have thrown considerable light on the previously mysterious evolutionary origin of whales. The holotype of ''Rodhocetus balochistanensis'', GSP-UM 3485, is: * A weathered braincase found at the surface next to a partial dentary with an unfused mandibular symphysis, a characteristic of protocetids. * Large parts of the axial skeleton including cervical, thoracic and proximal caudal vertebrae, but excluding sacral vertebrae. * Forelimb material including the left distal humerus, radius and ulna, and two virtually complete hand skeletons including all carpal bones, unfused and lacking an os centrale, and phalanges. * Parts of the pelvis including an acetabular rim. * Hind limb material including right femur, patella, tibia, possible partial fibula; two virtually complete foot skeletons including tarsal and metatarsal bones and phalanges. The astragalus (heel bone) is characteristic of artiodactyls with a deep tibial trochlea restricting lateral movements and a large calcaneal tuber (posterior part of heel bone) providing leverage for powerful extension. The metatarsals and phalanges are very long and thin and can not have been weight-bearing, suggesting that ''Rodhocetus'' was predominantly aquatic and on land must have walked on the plantar surface of the tarsals. The shape of the metatarsal and phalanges reveal that these bones could be tightly compressed during flexion and widely separated during extension. The five-fingered hand of ''R. balochistanensis'' is mesaxonic (i.e. has a central digit) with three weight-bearing central digits equipped with nail-like hooves, flanked by two more slender digits lacking hooves (distal phalanges preserved on first, second, and fourth digits). The four-toed foot is paraxonic (i.e. central axis passes between the two central digits), with all four digits ending in pointed nails (distal phalanges preserved on second and third digits). With an estimated body weight of , ''R. balochistanensis'' was 13% smaller than ''R. kasrani'' (), but its femur is larger.


See also

*
Evolution of cetaceans The evolution of cetaceans is thought to have begun in the Indian subcontinent from even-toed ungulates 50 million years ago (mya) and to have proceeded over a period of at least 15 million years. Cetaceans are fully aquatic marine mammals bel ...


References


Notes


Sources

* **
Supplementary material
** * ** * * {{Taxonbar, from=Q1550234 Eocene mammals of Asia Protocetidae Fossil taxa described in 1994 Prehistoric cetacean genera