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Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Cladus: Synapsida
Cladus: Eupelycosauria
Cladus: Sphenacodontia
Cladus: Sphenacodontoidea
Cladus: Theriodontia
Subordo: Cynodontia
Cladus: Mammaliaformes
Classis: Mammalia
Subclassis: Trechnotheria
Infraclassis: Zatheria
Supercohort: Theria
Cohort: Eutheria
Cohort: Placentalia
Cladus: Boreoeutheria
Superordo: Laurasiatheria
Ordo: Artiodactyla
Subordo: Whippomorpha
Infraordo: Cetacea

Familia: †Pakicetidae
Genus: †Ichthyolestes
Species: †I. pinfoldi

†Ichthyolestes Dehm & Oettingen-Spielberg, 1958: 15

Type species: †Ichthyolestes pinfoldi Dehm & Oettingen-Spielberg, 1958, by original designation and monotypy.
Primary references

Dehm, R. & Oettingen-Spielberg, T. 1958. Paläontologische und geologische Untersuchungen im Tertiär von Pakistan. 2. Die mitteleocänen Säugetiere von Ganda Kas bei Basal in Nordwest-Pakistan. Abhandlungen. Bayerische Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Klasse 91: 1–54. Reference page.

Vernacular names
日本語: イクチオレステス
中文: 魚中獸

Ichthyolestes ("fish thief") is an extinct genus of archaic cetacean that was endemic to Indo-Pakistan during the Lutetian stage.[1] To date, this monotypic genus is only represented by Ichthyolestes pinfoldi.[1]

Like other members of the Family Pakicetidae, which are considered the earliest and least specialized of the archaic cetaceans,[2] Ichthyolestes represents an early quadrupedal phase of the land-to-sea transition which occurs in the cetacean lineage.[3][4]

Discovery and Classification
Size of Ichthyolestes compared to a human.

I. pinfoldi was initially known only from teeth, which were found in the Eocene locality Ganda Kas, Pakistan.[1][5] Upon their discovery, I. pinfoldi was placed within the Family Mesonychidae,[1][3] a group of terrestrial mammals from which cetaceans were previously thought to have evolved.[6] Due to morphological distinctions from other mesonychids, I. pinfoldi was later acknowledged and described as an archaeocete,[7][8] making it the first Pakicetid to be described.[4]

Cranial and postcranial material of I. pinfoldi have also been found in the Kuldana Formation, Pakistan, from the early-middle Eocene,[8][6][9][2] and from the Subathu sediments of Jammu and Kashmir, India.[10]

Ichthyolestes is the smallest Pakicetid, approximately 29% smaller than Pakicetus, and has been considered “fox-sized.”[9] They retain many features typical of terrestrial Eocene artiodactyls, including long and gracile limb bones, a fused sacrum, small mandibular foramen, and no cranial telescoping.[9][11] The body plan of Ichthyolestes is generally similar to Pakicetus, but smaller and more gracile.[2][4] Therefore, locomotion is also thought to be reliant on quadrupedal paddling.[4]

Ichthyolestes exhibits heterodont and diphyodont dentition with cusped cheek teeth.[4] The upper second molar has pointed cusps with a high narrow paracone and a lower connate metacone behind it.[8][10] The molars are also relatively labio-lingually compressed with serrated crests along the labial cusps.[8][10]
Braincase and orbital region an Ichthyolestes pinfoldi skull (HGSP 98134)

Ichthyolestes has dorsally oriented eyes and a narrow skull, which is smaller than the skulls of both Pakicetus and Nalacetus.[3] Like other Pakicetids, Ichthyolestes lacks a supraorbital shield; however, there is some variation in supraorbital morphology between the three genera. The supraorbital region of Ichthyolestes and Pakicetus is cup-like and cradles the dorsal portion of the eye. Ichthyolestes also has a weaker incisure on the dorsal side of the supraorbital region compared to Pakicetus.[3]

All three Pakicetid taxa largely retained the peripheral ear morphology of terrestrial mammals,[11] but likely used bone conducting mechanisms when hearing underwater. The tympanic bulla also differs between Pakicetid taxa.[3] The absolute size of Ichthyolestes tympanic bulla is smaller than either Pakicetus or Nalacetus; but when compared relative to their body size, the tympanic bulla of Ichthyolestes is proportionally larger than that of Pakicetus.[3]

The semicircular canal system of the vestibular apparatus in the inner ear, which is involved in neural control of locomotion, is similar in size to that of Eocene artiodactyls.[12] They do not show the size reduction as seen in other Pakicetids and Eocene cetaceans like Remingtonocetus, Indocetus, and Dorudon, which have canal sizes within the upper range of modern cetaceans.[12] This suggests that Ichthyolestes had not fully invaded aquatic ecosystems and still remained somewhat terrestrial.[12]

Although Ichthyolestes is the smallest Pakicetid, some features are larger or more robust than Nalacetus, such as the astragalus, calcaneus, and pelvis. Ichthyolestes also has proportionally longer lumbar and caudal vertebrae relative to its shorter limb segments.[2]

Ichthyolestes also has long digits and strong post-thoracic vertebrae.[2] The sacrum is composed of four elongate vertebrae with complete fusion, except for the spinous processes of S2 to S4 which are columnar with smooth ventral faces.[2] The lumbar vertebrae are narrower and shallower to Pakicetus and Nalacetus.[2] Although still relatively large compared to other related taxa, the atlas vertebrae of Ichthyolestes is smaller and more gracile than Pakicetus or Nalacetus, and the neural canal is disproportionately large.[2]
Terrestrial or Semi-aquatic Locomotion

The astragalus bone, which is present in the ankle of artiodactyls and archaic whales that retained feet, has similar dimensions in both Eocene artiodactyls and Ichtyolestes.[5] In addition to lending evidence to the theory of a direct relationship between artiodactyls and cetaceans,[11] this knowledge has led to two competing hypotheses about the locomotion of Ichtyolestes and other Pakicetids. The first states that Ichtyolestes and Pakicetus were terrestrial and cursorial, implying that aquatic locomotor adaptation occurred after the origin of Cetacea.[9][13] The second states that Ichtyolestes and Pakicetus were already semi-aquatic and cetaceans originated from an earlier unknown Eocene artiodactyl, such as Elomeryx[14][15] or Indohyus,[16][17] implying that aquatic locomotor adaptations occurred before or during the origin of Cetacea.[5]

Although the postcranial anatomy of Ichtyolestes is similar to that of Eocene artiodactyls and implies cursoriality,[2][5] the assessment of bone morphology and microstructure indicate that they, and other Pakicetids, were semi-aquatic like Protocetids.[2] Hypermineralization occurs in all regions of the skeleton; in particular, the long bones and ribs had small or absent marrow cavities due to the thick cortices which developed.[2] Hypermineralization of load-bearing skeletal elements put Ichtyolestes at an increased risk of fractures during prolonged terrestrial loading and this risk increased with velocity, implying that terrestriality was limited.[2] Additionally, the dense skeletons may have allowed bottom-walking or wading in shallow pools as it would counteract buoyancy created by inflated lungs and fur-trapped air.[2] Therefore, the retention of an artiodactyl-like astragalus does not signify full terrestriality or cursoriality in Ichtyolestes.[2][5]

Fossil findings indicate Ichthyolestes shared habitat with its relatives Pakicetus and Nalacetus.[18] Their fossils are usually found around river channel deposits, not marine deposits or fauna.[4] The regions around northern Pakistan and northwest India, where fossils have been recovered, are thought to have been an arid environment with ephemeral streams and moderately developed floodplains.[19] Ichthyolestes, therefore, had an affinity for water and were either terrestrial or semi-aquatic.[4] The fluvial facies of the lower Kuldana Formation represent shallow tropical riverine complexes and were likely habitats for the initial stages of transition into water.[5][2]
See also

Evolution of cetaceans


Dehm, R.; zu Oettingen-Spielberg, T. (1958). Paläontologische und geologische Untersuchungen im Tertiär von Pakistan. 2. Die mitteleocänen Säugetiere von Ganda Kas bei Basal in Nordwest-Pakistan (PDF). Bayerische Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Klasse, Abhandlungen, Neue Folge. 91. Munich: C. Beck’sche Verlagsbuchhandlung.
Madar, S. (2007). "The postcranial skeleton of early Eocene Pakicetid cetaceans". Journal of Paleontology. 81 (1): 176–200.
Nummela, S.; Hussain, S. T.; Thewissen, J. G. M. (2006). "Cranial anatomy of Pakicetidae (Cetacea, Mammalia)". Journal of Vertebrate Paleontology. 26 (3): 746–759.
Thewissen, J. G. M.; Williams, E. M. (2002). "The early radiation of Cetacea (Mammalia): Evolutionary pattern and development correlations". Annual Review of Ecology and Systematics. 33: 73–90.
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". PalZ. 91: 601–627.
Gingerich, P. D.; Wells, N. A.; Russell, D. E.; Ibrahim Shah, S. M. (1983). "Origin of whales in epicontinental seas: New evidence from the early Eocene of Pakistan". Science. 220 (4595): 403–406.
Gingerich, P. D.; Russell, D. E.; Sigogneau-Russell, D.; Hartenberger, J. -L.; Ibrahim Shah, S. M.; Hassan, M.; Rose, K. D.; Ardrey, R. H. (1979). "Reconnaissance survey and vertebrate paleontology of some Paleocene and Eocene formations in Pakistan". Contributions from the Museum of Paleontology University of Michigan. 25 (5): 105–116.
West, R. M. (1980). "Middle Eocene large mammal assemblage with Tethyan affinities, Ganda Kas Region, Pakistan". Journal of Paleontology. 54: 508–533.
Thewissen, J. G. M.; Williams, E. M.; Roe, L. J.; Hussain, S. T. (2001). "Skeletons of terrestrial cetaceans and the relationship of whales to artiodactyls". Nature. 413: 277–281.
Kumar, K.; Sahni, A. (1985). "Eocene mammals from the Upper Subathu group, Kashmir Himalaya, India". Journal of Vertebrate Paleontology. 5 (2): 153–168.
Marx, F. G.; Lambert, O.; Uhen, M. D. (2016). Cetacean Paleobiology. Chichester, UK; Hoboken, NJ: John Wiley & Sons Ltd.
Spoor, F.; Bajpai, S.; Hussain, S. T.; Kumar, K.; Thewissen, J. G. M. (2002). "Vestibular evidence for the evolution of aquatic behaviour in early cetaceans". Nature. 417: 163–166.
Fish, F. E. (2016). "Secondary evolution of aquatic propulsion in higher vertebrates: Validation and prospect". Integrative and Comparative Biology. 56: 1285–1297.
Gingerich, P. D.; Haq, M.; Zalmout, I. S.; Khan, I. H.; Malkani, M. S. (2001). "Origin of whales from early artiodactyls: Hands and feet of Eocene Protocetidae from Pakistan". Science. 293: 2239–2242.
Gingerich, P. D. (2003). "Land-to-sea transition of early whales: Evolution of Eocene Archaeoceti (Cetacea) in relation to skeletal proportions and locomotion of living semiaquatic mammals". Paleobiology. 29: 429–454.
Thewissen, J. G. M.; Cooper, L. N.; Clementz, M. T.; Bajpai, S.; Tiwari, B. N. (2007). "Whales originated from aquatic artiodactyls in the Eocene epoch of India". Nature. 450: 1190–1195.
Cooper, L. N.; Thewissen, J. G. M.; Bajpai, S.; Tiwari, B. N. (2011). "Postcranial morphology and locomotion of the Eocene raoellid Indohyus (Artiodactyla: Mammalia)". Historical Biology. 24: 279–310.
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): p. 746-759. DOI:10.1671/0272-4634(2006)26[746:CAOPCM]2.0.CO;2
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.

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