Rodhocetus Balochistanensis
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''Rodhocetus'' (from ''Rodho'', the geological
anticline In structural geology, an anticline is a type of Fold (geology), fold that is an arch-like shape and has its oldest Bed (geology), beds at its core, whereas a syncline is the inverse of an anticline. A typical anticline is convex curve, c ...
at the type locality, and ''cetus'', Latin for whale) is an extinct genus of
protocetid Protocetidae, the protocetids, form a paraphyletic, 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 kno ...
early whale known from the
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 ...
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 The sacrum (: sacra or sacrums), in human anatomy, is a triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part of the pelvic cavity, ...
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 animal, aquatic placental mammal, placental marine mammals. As an informal and Colloquialism, colloquial grouping, they correspond to large members of the infraorder Cetacea ...
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 mesonychians, 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 may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
(ankle 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 Linear map, linear dimensionality reduction technique with applications in exploratory data analysis, visualization and Data Preprocessing, data preprocessing. The data is linear map, linearly transformed ...
demonstrated that ''Rodhocetus'' had trunk and limb proportions similar to the
Russian desman The Russian desman (''Desmana moschata''; ''vykhukhol'') is a small semiaquatic mammal that inhabits the Volga river, Volga, Don River, Russia, Don and Ural River basins in Russia. Some authorities, citing old Soviet sources, claim the animal ca ...
, 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'' 41.03 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 (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
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 traits in ''R. kasrani'', relative to older archaeocetes such as ''Pakicetus'', include high-crowned cheek teeth, larger auditory bullae, larger mandibular foramen, and mandibular canals. The higher neural spines and shorter femur (60–70%) distinguish ''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 b ...
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 ...
e or air sinuses. The
mandibular foramina The mandibular foramen is an opening on the internal surface of the ramus of the mandible. It allows for divisions of the mandibular nerve and blood vessels to pass through. Structure The mandibular foramen is an opening on the internal surfac ...
are large with a pan bone long and high. The specific name ''kasrani'' comes from
Qaisrani Qaisrani is a Baloch clan of the Rind tribe. They live mostly in Balochistan, Pakistan Balochistan (; ; , ) is a province of Pakistan. Located in the southwestern region of the country, Balochistan is the largest province of Pakistan by ...
, the Baloch tribe inhabiting the type locality. The protocetid ''
Qaisracetus ''Qaisracetus'' is an extinct protocetid early whale known from the Eocene (Lutetian, ) of Baluchistan, Pakistan (, paleocoordinates ). Etymology The genus is named after the Qaisrani Baloch tribe which assisted Gingerich and his team during ...
'' is also named after them.


''R. balochistanensis''

The fossil remains of ''R. balochistanensis'' were found in eastern
Balochistan Balochistan ( ; , ), also spelled as Baluchistan or Baluchestan, is a historical region in West and South Asia, located in the Iranian plateau's far southeast and bordering the Indian Plate and the Arabian Sea coastline. This arid region o ...
,
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# ...
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 early whales that lived during the Early Eocene in Pakistan. Unlike modern cetaceans, they had well-developed limbs and were capable of walking. The species included were fox to wolf-sized. ...
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 includes the right femur, patella, tibia, and possible partial fibula; two virtually complete foot skeletons include 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 (Artiodactyla) 50 million years ago (mya) and to have proceeded over a period of at least 15 million years. Cetaceans are fully aquatic mam ...


References


Notes


Sources

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