mesaxonians are a clade of ungulates whose weight is distributed on the third toe on all legs through the plane symmetry of their feet .

for a while it was often seen to only contain the order perissodactyla ( which includes the equines , rhinos and tapirs ) .

recent work in morphological cladistics and ancient dna suggests that several extinct lineages , like the desmostylia and some of the south american ungulates of meridiungulata ( both groups seen as afrotherian relatives ) are related to the perissodactyls .

mesaxonia

the mesaxonia , as defined here , include the horses , rhinos , elephants , and dugongs , among others . the group shares a number of foot , skull , and tooth characteristics which support its monophyly . however , no official name has been published for this group , and so we have borrowed the term mesaxonia , which is still used by some authors to refer to the perissodactyla .
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fisher ms , tassy p ( 1993 ) the interrelation between proboscidea , sirenia , hyracoidea , and mesaxonia : the morphological evidence . in : szalay fs , novacek mj , mckenna mc , editors . mammal phylogeny : placentals . new york : springer - verlag . pp . 217–234 .
fischer ms , tassy p ( 1993 ) the interrelations between proboscidea , sirenia , hyracoidea , and mesaxonia : the morphological evidence . in : szalay fs , novacek mj , mckenna mc , editors . mammal phylogeny : placentals . new york : springer - verlag . pp . 217–243 .
[ mesaxonia ] include the perissodactyla and some related fossil types . the middle toe may be the only one ( e . g . in present day horses ) , or may be the largest of three , as in some ancestral horses and in the hind - foot of the rhinoceros .
mesaxonians are a group of mammals that form the , not widely accepted , superorder mesaxonia . the superorder includes the perissodactylids , hyracoids , proboscidians , and sirenians , all part of the infraclass eutheria . however , this group is not yet published , it ` s a theory based on the number of digits , teeth , and skull characteristics . . . .
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anthracobunidae is an eocene family of large mammals from south asia that is commonly considered to be part of the radiation that gave rise to elephants ( proboscideans ) and sea cows ( sirenians ) . we describe a new collection of anthracobunid fossils from middle eocene rocks of indo - pakistan that more than doubles the number of known anthracobunid fossils and challenges their putative relationships , instead implying that they are stem perissodactyls . cranial , dental , and postcranial elements allow a revision of species and the recognition of a new anthracobunid genus . analyses of stable isotopes and long bone geometry together suggest that most anthracobunids fed on land , but spent a considerable amount of time near water . this new evidence expands our understanding of stem perissodactyl diversity and sheds new light on perissodactyl origins .
citation : cooper ln , seiffert er , clementz m , madar si , bajpai s , hussain st , et al . ( 2014 ) anthracobunids from the middle eocene of india and pakistan are stem perissodactyls . plos one 9 ( 10 ) : e109232 . urltoken
editor : andrew a . farke , raymond m . alf museum of paleontology , united states of america
copyright : © 2014 cooper et al . this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
data availability : the authors confirm that all data underlying the findings are fully available without restriction . all relevant data are within the paper and its supporting information files .
funding : the national science foundation is acknowledged for support to m . t . c . ( ear - 0847413 ) , e . r . s . ( bcs - 0819186 and bcs - 0416164 ) , and j . g . m . t . ( ear - 0207370 ) . also , s . b . acknowledges financial support received from the department of science and technology , government of india for field work in the eocene sections of gujarat . the funders had no role in the study design , data collection and analysis , decision to publish , or preparation of the manuscript .
anthracobunidae is a family of large mammals that is only known from the middle eocene of south asia . historically , anthracobunids have played a prominent role in debates surrounding the origin of tethytheria , the group that includes the living and extinct members of the placental mammalian orders sirenia ( sea cows ) and proboscidea ( elephants ) . anthracobunids are often considered to have branched off near the base of tethytheria [ 1 ] – [ 8 ] , sharing most recent common ancestry with either proboscidea [ 1 ] , [ 9 ] – [ 13 ] , sirenia [ 14 ] , [ 15 ] , or the extinct desmostylia . however , until now , no cranial parts , only some partial dentitions and two postcranial elements ( a scapula associated with a maxilla [ 11 ] and an isolated astragalus [ 13 ] ) , were known for the group .
the paucity of fossil evidence has left anthracobunidae as a phylogenetically and adaptively enigmatic group [ 2 ] , [ 8 ] , [ 16 ] – [ 20 ] , with little known about their diets , habitats , and time and place of origin . in light of this , anthracobunid genera have also been aligned with two other , non - tethytherian clades , including artiodactyla [ 21 ] – [ 23 ] and perissodactyla [ 24 ] , as well as phenacodontids [ 25 ] , [ 26 ] , one of the extinct families of basal ungulates . the convergent evolution of “ungulate” - like features in paenungulata ( containing tethytheria ) and laurasiatheria ( containing artiodactyla and perissodactyla ) revealed by molecular data [ 27 ] requires that anthracobunidae be re - evaluated within this new phylogenetic context .
here we describe new anthracobunid fossils from the kuldana and subathu formations of the early - middle eocene of pakistan and india [ 28 ] – [ 32 ] . these new findings more than double the number of anthracobunid fossils , and include cranial and postcranial material . we test the phylogenetic relationships of anthracobunids using a taxonomically rich morphological data set and investigate anthracobunid diet and habitat . our results have important implications for character transformations near the origins of perissodactyla , the habitats of stem perissodactyls , and the early biogeography of paenungulata and perissodactyla .
fossils were collected in punjab province , pakistan , under a collaborative agreement of howard university and the geological survey of pakistan ( h - gsp ) . localities where fossils were collected , with topographical coordinates , were described in thewissen et al . [ 33 ] . specimens were prepared in the u . s . a . at northeast ohio medical university ( neomed ) and are catalogued using the h - gsp acronym and accessioned into the collections of the geological survey of pakistan ( gsp ) . since the gsp is a branch of the pakistani government , all collected fossils are the property of the pakistani state . at present , fossils are for study in the u . s . a . , but they will be returned to pakistan and will be permanently housed at the natural history museum of pakistan , located in islamabad . gsp geologists are collaborators on all field work as part of their official assignment and all permitting is internal to the different pakistani government agencies involved . all necessary permits were obtained for the described study , which complied with all relevant regulations .
we also studied fossils from the locality kalakot in jammu & kashmir state , india , the location of which is described by russell and zhai [ 32 ] . this locality was discovered by a . ranga rao , a geologist of the oil and natural gas commission , and he quarried large quantities of fossiliferous rock matrix from this locality . after his death , we were given access to these by his widow , f . obergfell . fossils were prepared at neomed in the u . s . a . , and are the property of the rangarao - obergfell trust for geosciences , located in dehra dun , india , and are catalogued using the acronym rr . they will be housed on the trust campus on rajpur road in dehra dun .
the electronic edition of this article conforms to the requirements of the amended international code of zoological nomenclature , and hence the new names contained herein are available under that code from the electronic edition of this article . this published work and the nomenclatural acts it contains have been registered in zoobank , the online registration system for the iczn . the zoobank lsids ( life science identifiers ) can be resolved and the associated information viewed through any standard web browser by appending the lsid to the prefix “ urltoken ” . the lsid for this publication is : urn : lsid : zoobank . org : pub : cd33f507 - 0d06 - 4de1 - a3c8 - 8bf9e7697114 . the electronic edition of this work was published in a journal with an issn , and has been archived and is available from the following digital repositories : pubmed central , lockss .
whenever possible , characters were scored based on analysis of original fossil material and / or casts housed in either the department of vertebrate paleontology at the american museum of natural history , the department of anatomical sciences at stony brook university , or the department of anatomy and neurobiology at northeast ohio medical university . as noted above , some taxa and / or characters were scored from the literature . some dental characters were rescored for arsinoitherium , taking into account new evidence for cusp homologies provided by the more basal embrithopod namatherium [ 52 ] . furthermore , a new character state ( “large , inflated” ) was added for the character “m1 - 2 metaconules” ( number 18532 ) to take into account the condition seen in anthracobunids and some other ingroup taxa , and the character “shape of astragalar ectal facet” was excluded , because it could not be consistently scored across the new taxa that were sampled .
midshaft cross - sections of fossil and extant large - bodied ungulate long bones and ribs [ 71 ] were visualized via paleohistological sections or high resolution micro - ct scans . bone compactness ( total amount of bone per section ) was calculated via published methods [ 72 ] .
for the analysis of stable isotopes , three or more specimens of each species were analyzed ( when available ; see tables s8 and s9 ) to provide an estimate of the population mean ± s . d . for carbon and oxygen isotope values [ 73 ] urltoken - b33 . about 5 mg of enamel powder was collected from each specimen , either by drilling directly from the tooth or by grinding enamel chips in an agate mortar and pestle . before collection , contaminants were removed by abrading the outer surface of the specimen after cleaning the outer surface to a depth of ∼0 . 5 mm using a dremel drill with a 1 mm diameter diamond - coated bit .
preparation of powders for stable isotope analysis followed published methods [ 74 ] , [ 75 ] . powders were first transferred to 1 . 5 ml microcentrifuge vials and then soaked sequentially overnight in about 0 . 20 ml of a sodium hypochlorite solution ( 1–2 g dl - 1 ) and then in about 0 . 20 ml of calcium acetate buffered acetic acid ( ph about 5 ) following published procedures [ 31 ] . on addition of each reagent , samples were agitated for 1 min on a vortex genie vortex mixer . after each soak , the supernatant was removed by aspiration and the residual powder was rinsed five times with deionized water . samples were then freeze - dried overnight and about 1 . 5 mg of powder from each was weighed into individual test tubes for analysis on a thermo - finnigan gas bench autosampler attached to a thermo - finnigan deltaplusxp continuous - flow isotope - ratio mass spectrometer at the university of wyoming stable isotope facility .
all values for stable isotopes are reported in delta ( δ ) notation , using the equation δ ( ‰ ) = 1 , 000 × ( rsample / rstandard - 1 ) , where rsample is the observed isotope ratio of the sample ( 13 c / 12 c or 18 o / 16 o ) and rstandard is the accepted ratio for an appropriate international standard ( vienna pee dee belemnite for δ 13 c ; vienna standard mean ocean water for δ 18 o ) . analytical precision is typically better than 0 . 1‰ for δ 13 c values and 0 . 2‰ for δ 18 o values ( ±1σ ) .
enamel δ 18 o values are influenced by the oxygen isotope compositions of oxygen sources ( atmospheric o 2 , food and water ingested by the animal ) , certain physiological processes that affect intake or loss of oxygen by an animal ( sweating , panting , respiration , and elimination of urine and feces ) , and body temperature [ 76 ] – [ 78 ] . for aquatic and semi - aquatic mammals , the flux of environmental water by means of direct ingestion and transcutaneous exchange [ 79 ] overwhelms all other oxygen sources , which reduces inter - individual variation [ 76 ] , [ 80 ] and causes enamel δ 18 o values of freshwater taxa ( for example hippopotamus ) to be lower than those for terrestrial mammals ( ∼2–3‰ ) ; [ 81 ] – [ 83 ] . the magnitude of offset between freshwater and terrestrial mammals can vary in response to changes in humidity , such that faunas experiencing drier conditions show a greater offset ( ∼3 . 0‰ ) than those living under wetter conditions ( < 1 . 0‰ ) [ 84 ] .
in combination with enamel δ 18 o values , we also determined enamel δ 13 c values of fossil species , which were used to infer diet and past environmental conditions . for herbivorous ungulates , enamel carbon isotopic compositions are enriched in 13 c relative to diet by ∼14‰ [ 85 ] . variation in enamel δ 13 c values among ungulate species is most strongly affected by the carbon isotopic composition of the plants they ingest . in the eocene , terrestrial ecosystems were dominated by plants performing c3 photosynthesis , a process that discriminates against the heavier carbon isotope ( 13 c ) , causing plant tissue δ 13 c values to be considerably lower than those of plants exploiting other photosynthetic pathways ( c4 plants , cam plants ) [ 86 ] , [ 87 ] . the carbon isotopic compositions of c3 plants are more variable than other plant types and are strongly affected by temperature and aridity , causing c3 plants grown under hot dry conditions to have higher δ 13 c values than those grown in cooler and wetter environments [ 88 ] , [ 89 ] . since enamel δ 13 c values reflect the carbon isotopic composition of the plants in an animal ' s diet , consumer enamel δ 13 c values can be used to infer past environmental conditions ( wet vs . dry ) as well as dietary information for extinct species .
included genera . jozaria wells and gingerich , 1983 ; and obergfellia , new genus .
diagnosis . stem perissodactyls with premolars increasing in complexity from anterior to posterior , but never molariform ; molars brachyodont , but with high cusps and low valleys between cusps ; upper molars increasing in size from m1 to m3 ; lower molars with a distinct trigonid elevated above the talonid ; large caudally - projecting angular process of the mandible .
description . unlike tethytheres and other paenungulates , anthracobunids have small and simple upper and lower incisors and relatively large canines ( figs . 1 , 2 ) . cheek teeth are bunodont and brachyodont and each cusp has a high apex , but there are deep valleys between cusps . premolars are never totally molariform , and diastemata , if present at all , are short . upper p3 and p4 are shorter than the molars , and have a paracone , protocone , and metacone , but no hypocone . the m1 and m2 have four well - developed cusps , as well as strong para - and metaconules , without lophs . the distance between proto - and paracone is longer than that between the hypo - and metacone , giving the tooth a posteriorly pinched appearance . lower molars have a trigonid with two large cusps , and large , sometimes twinned , hypoconulid on the third lobe ( figs . 1g–i , s1 ) .
( a ) crushed skull of a . pinfoldi ( h - gsp 97106 ) in ventro - lateral view ( left maxilla detached ) and ( b ) dorsal view ; ( c ) p2 of a . wardi ( h - gsp 30229 ) in lingual view and ( d ) occlusal view ; ( e ) skull fragment of a . wardi ( rr 411 ) in occlusal view , and ( f ) lateral view ; ( g ) mandible of a . wardi ( h - gsp 96434 ) in occlusal view ; ( h ) mandible of a . wardi ( h - gsp 96258 ) in lateral view , and ( i ) occlusal view . ( j ) proximal phalanx of a . pinfoldi ( h - gsp 97106 . 105 ) in dorsal view ; ( k ) proximal phalanx of a . pinfoldi ( h - gsp 97106 . 101 ) in ventral view , and ( l ) lateral view ; ( m ) head of a metapodial of a . pinfoldi ( h - gsp 97106 . 250 ) in dorsal view ; ( n ) phalangeal fragment of a . pinfoldi ( h - gsp 97106 . 257 ) in dorsal - superior view ; ( o ) terminal phalanx of a . pinfoldi ( h - gsp 97106 . 302 ) in dorsal - superior view . scale bar is 1 cm in length .
( a ) right p2 - m2 of a . wardi ( luvp - 15006 ) in occlusal view , and ( b ) labial view ; ( c ) left m1 - m3 of a . wardi ( rr - 411 ) in occlusal view , and ( d ) labial view ; ( e ) left p3 - m3 of a . pinfoldi ( h - gsp 82 - 31p ) in occlusal view ; ( f ) left m3 of a . pinfoldi ( h - gsp 82 - 31p ) in labial view ; ( g ) right c - m3 of the cambaythere kalitherium ( iitr - sb - vlm 931 ) in occlusal view ; ( h ) right m3 of kalitherium ( iitr - sb - vlm 931 ) in labial view ; ( i ) left p4 - m2 of nakusia [ 6 ] in occlusal view ; ( j ) left m2 of nakusia [ 6 ] in labial view ; ( k ) mx of cambaytherium ( iitr - sb - vlm - 521 ) in occlusal view ; ( l ) right m3 of the phenacodontid ‘condylarth’ tetraclaenodon ( ku - 8052 ) in occlusal view ; ( m ) left p1 - m3 of a . wardi ( wif / a 1101 ) [ 15 ] in occlusal view ; ( n ) left c - m3 of a . wardi ( h - gsp 96258 ) in occlusal view , and ( o ) labial view ; ( p ) left p1 - m3 of obergfellia occidentalis ( h - gsp 1981 ) in occlusal view , and ( q ) labial view ( m1 inverted from right side ) . scale bar is 1 cm in length . illustrations by jacqueline dillard .
partial skulls are known for two anthracobunids ( h - gsp 97106 and rr 411 ) , and both show the presence of a very thick bony shelf over the orbits ( fig . 1b , f ) . the palate is concave mediolaterally , with the areas near the midline much higher than the alveolar processes . the lateral sides of the basisphenoid and basioccipital are recessed ; this is particularly true for the basioccipital , which suggests that the petrosal , which is not preserved , was also deeply recessed . the mandibular fossa is concave and is located just lateral to the ear region . the postglenoid process is oval and does not extend along the entire width of the mandibular fossa .
h - gsp 97106 includes , among other elements , metapodials and phalanges ( fig . 1j - o ) . the heads of most metapodials are wider than deep , strongly convex , and the posterior side bears a strong crest . proximal phalanges show a broad , oval articular surface for the metapodial and are squat : nearly as wide mediolaterally as they are long proximodistally . their distal articular facet is rectangular in outline and deeply incised caudally , indicating the presence of a caudal crest on the middle phalanx . most prominent on the proximal phalanges is the deeply excavated posterior side which is concave both mediolaterally and proximodistally .
the genera of the family anthracobunidae . we restrict the family anthracobunidae to the genera anthracobune ( which includes most specimens previously referred to pilgrimella and lammidhania ) , jozaria , and the new genus obergfellia . there are several other families of medium - sized bunodont “ungulates” from the eocene of india and pakistan , such as quettacyonids [ 90 ] and cambaytheres [ 4 ] , [ 91 ] . while all of these groups are brachyodont , anthracobunids are unique among them in having tall cusps on their molars , while retaining valleys between the cusps that are low and close to the cingulum ( fig . 2 ) . the new genus obergfellia differs from other anthracobunids in exhibiting the following combination of features : ( i ) lower molars broad , ( ii ) lower m3 relatively short , and ( iii ) angular process of the mandible long but shorter than that of anthracobune ( figs . 1 , 2 , s1 . ) . the last of these features is not known in jozaria .
we exclude ishatherium subathuensis from anthracobunids . its holotype consists of the lingual side of an upper molar , referred originally to sirenia [ 15 ] , [ 92 ] . wells and gingerich [ 1 ] referred ishatherium to anthracobunidae . the holotype and only specimen shares with anthracobunids the strong development of conules and the deep transverse wear along the protocone - paraconule - paracone and along the hypocone - metaconule - metacone . the specimen is decidedly unlike anthracobunids in that the distance between protocone and paracone is similar to that between hypocone and metacone , and in that its cusps are not highly raised . we also exclude nakusia shahrigensis [ 6 ] from anthracobunids ( fig . 2 i , j ) . its holotype is a maxilla with p4 - m2 and the base of p3 . the base of p3 is elongate , the cusps on the relatively unworn m2 are low , and the molars are relatively wide . anthracobunids have a short p3 , high molar cusps , and squarish upper molars . nakusia is more likely a quettacyonid or cambaythere than an anthracobunid .
the genus indobune was included in anthracobunidae [ 4 ] , but it has low cusps on its teeth and is better classified as a cambaythere . ducrocq et al . [ 7 ] described hsanotherium parvum on the basis of two specimens with upper molars from myanmar . these specimens are bunodont , with small conules and lack a hypocone . the transverse wear pattern of anthracobunids is also absent . hsanotherium displays similarities to medium - sized bunodont artiodactyls , such as haqueina [ 22 ] , and we do not include it in anthracobunids .
referred species . anthracobune wardi ( dehm and oettingen - spielberg , 1958 ) .
diagnosis . anthracobunid with narrow lower molars and a very large angular process of the mandible ( fig . 1 , fig . s1 ) .
discussion . we refer to most of the specimens formerly included in lammidhania and pilgrimella in the past to anthracobune .
holotype . bmnh m . 15792 , left m2–3 and right m3 , from ‘ lammidhan ’ and ‘ planorbis freshwater beds . ’ pilgrim never visited the type region , and the fossils were collected by a geological surveyor ( t . g . b . davies ) . lammidhan appears on davies’ unpublished map and is located on a broad alluvial plain . however , it does not match a modern topographical landmark and its meaning is not known to local people . eocene deposits occur to the north and south of the plain , with marine limestones and muds forming ledges usually towering over the unconsolidated muds that form the bottoms of valleys . these muds are the “ planorbis freshwater beds , ” and it is our contention that the specimen may have been found on these beds , but that it rolled down from overlying marine beds . gingerich [ 23 ] improved the holotype by fitting bmnh m . 15794 ( the trigonid of the left m2 ) to it .
description . h - gsp 97106 is the most complete specimen known for any anthracobunid . upper incisors are similar in size ( based on their alveoli ) ; the crown of i3 shows that there is a single , large pointed cusp , with small basal thickening on the cingulum anterior and posterior to it . canine is long and compressed mediolaterally and p1 is two - rooted , with single cusp . p2 has a small protocone and two cuspules on the posterior crest of the paracone . p3 has a protocone , paracone , metacone and two conules , and a strong postprotocrista . upper molars with four main cusps as well as strong para - and metaconule . instead , protocone - paraconule - paracone have aligned wear surfaces , and similar wear surfaces on hypocone - metaconule - metacone . with wear , these surfaces resemble crests .
lower p1 with two roots , strong protoconid , high but narrow paraconid and low metaconid , lacking talonid . lower p2 is similar to p1 but larger . lower p3 and p4 are similar with a well - developed trigonid having three cusps , and two cusps on the talonid . lower molars with strong crests between protoconid and metaconid , and between hypoconid and entoconid ; weak hypoconulid on anterior molars . these crests are stronger than the cristid obliqua . paraconid usually absent on molars .
upper anterior teeth lined up in a parasagittal row , and rostrum ends in a narrow point . the caudal edge of the nasal opening of h - gsp 97106 is over the upper canine , and the infraorbital foramen is dorsal to p3 . the zygomatic arch projects lateral from the base of m1 and m2 , and extends caudally from there . the orbit is located over m2 and m3 . the skull roof dorsal to the orbits is a thick bony plate that is flat and stretches rostrally to the nasal aperture , and caudally to the temporal crests . left and right temporal crests form the caudal extent of this plate and join into a strong sagittal crest that delineates the large temporal fossae . this crest ends at the left and right nuchal crests , which extend ventral to the ear region .
the palate is narrow rostrally , and concave mediolaterally . a weak postpalatine torus is present , and the hard palate is indented caudally with the choana reaching rostrally medial to m3 . the pterygoid process is thick . the periotics are deeply recessed between basioccipital and mandibular fossa . basisphenoid and basioccipital slope strongly from medial to lateral . the postorbital process is strong and oval in cross - section , the mandibular fossa is cylindrical , its rostrocaudal extent is as large as its mediolateral extent . the external auditory meatus is immediately caudal to the postorbital process , there is no postglenoid foramen . the external auditory meatus is directed caudolaterally . the suture between mastoid and squamosal is on the nuchal crest , and there is a mastoid emissary foramen just below it . a large emissary foramen also occurs on the parietal in the temporal fossa . the foramen magnum faces more caudal than ventral and the supraoccipital immediately dorsal to it is flat . near the sagittal crest , the supraoccipital plane is concave .
discussion . specimens referred to a . pinfoldi are known from the upper part of the kuldana formation . these are coastal beds , as indicated by the sedimentology and invertebrate paleontology of their localities , although isotopic evidence implies that the individuals fed on land . the upper part of the kuldana formation is well known for its fossil whales : ambulocetus and attockicetus .
holotype . bmnh m . 15799 , isolated talonid , most likely of m1 based on its size . locality given as ‘ lammidhan … planorbis freshwater beds of chharat stage’ ( see discussion under holotype of anthracobune pinfoldi ) . unlike other specimens from pilgrim ' s ‘ lammidhan , ’ this specimen is probably indeed from the planorbis beds , an inference based on its preservation .
diagnosis . lower molars with high length / width ratio , the m3 short , paraconid and metaconid of lower premolars small , anterior premolars long .
description . new material referred to this species includes two complete lower jaws that include the entire post - incisor dentition ( h - gsp 96258 and 96434 ) , a skull fragment that includes orbit and supraorbital region ( rr 411 ) , and part of the deciduous dentition .
lower incisors are similar in size to each other , based on their alveoli . canines with short crown , larger than the incisors , small diastemata on either side of p1 , but no other diastemata present . lower p1 and p2 with one main cusp and two roots , and small cusp on pre - and postprotocristid . lower p3 and p4 with well - developed trigonid and talonid . protoconid and metaconid on these teeth similar in height and paraconid lower , much lower on p4 . talonid of posterior premolars with high hypoconid , and entoconid high or barely distinguishable . lower molars with distinct trigonid and talonid . protoconid and metaconid similar in height , paraconid absent , but paracristid well developed . hypoconid and entoconid similar in height , cristid obliqua strong , hypoconulid weak on m1 and m2 , but strong and placed on third lobe in m3 , and sometimes twinned . coronoid process of the mandible small and with steep anterior slope . length of mandible anterior to anterior edge of coronoid process ( to rostral end of the fused symphysis ) is approximately two times as long as that posterior from this edge ( to the tip of the angular process ) . this indicates that the angular process is enormous . lower p3 with three cusps on trigonid , and two on talonid ; trigonid long . lower p4 with shorted trigonid , resembling lower m1 ( h - gsp 96052 ) . mandibular angle of juveniles small ( h - gsp 30349 , fig . s1 ) .
upper dentitions of a . wardi are mostly known from india ( luvp 15006 , rr 411 , wif / a 616 ) . the p2 is longer than wide , unlike the p3 and p4 . p2 has connate paracone and metacone , and a lingual bulge with a small protocone ( luvp 15006 , h - gsp 30229 ) . p3 and p4 are similar in shape , with connate para - and metacone , and large protocone . paraconule and metaconule are strong in p3 ( luvp 15006 ) , whereas in p4 transverse crests are well developed and conules less so . upper molars have four strongly developed cusps , with smaller but distinct paraconule and metaconule . transverse crests are absent , but cusps are lined up in such a way that they form an anteriorly convex arch .
tooth wear in anthracobunids tends to be distinct on the posterior side of the protoconid - metaconid , and the anterior side of the protocone - protoconule - paracone , and is accentuated by the deep valleys behind these rows . this transverse wear pattern is consistent with the large and caudally - projecting angular process of the mandible , the area of insertion for the medial pterygoid and masseter muscles .
specimen rr 411 reveals some details about the orbit and zygomatic arch . the zygomatic arch has a weak postorbital process and the dorsal part of the zygomatic arch is made up of the maxilla in the orbital rim , but not posterior to the orbit . the ventral edge of the zygomatic arch is made up of the jugal . the morphology of the root of the zygomatic arch , the postorbital process of the frontal , and the mandibular fossa in this specimen resembles that of h - gsp 97106 .
the anterior edge of the ascending ramus of the mandible of h - gsp 96259 is vertical , whereas it slopes slightly caudal in h - gsp 96434 : the superior part overhanging the inferior part . h - gsp 30349 is a juvenile in which the anterior edge of the ramus slopes slightly rostral . in h - gsp 96149 this ramus is vertical . the mandibular symphysis of all jaws , except h - gsp 30349 , is strongly fused , and slopes caudal ending ventral to the premolars .
discussion . most specimens of anthracobune wardi are known from the lower redbeds of the kuldana formation of pakistan , where pakicetid cetaceans are common . these are freshwater deposits and the specimens of anthracobune found here pertain to a . wardi . additional specimens of a . wardi are from the freshwater deposits of the subathu formation of india .
obergfellia , new genus . urn : lsid : zoobank . org : act : 952704f2 - f029 - 4643 - b791 - b30e7c71aa0a .
anthracobune ( in part ) , west , 1980 , 1981 , 1983 , 1984 .
pilgrimella ( in part ) , wells and gingerich , 1983 ; kumar , 1991 .
etymology . in honor of the late vertebrate paleontologists dr . friedlinde obergfell and her husband a . ranga rao . both were instrumental in the initial discovery and description of the kalakot fauna . this fauna became the best - known eocene land mammal fauna from india , mostly due to the efforts of a . sahni and k . kumar .
obergfellia occidentalis , new species , urn : lsid : zoobank . org : act : b5a9a6da - db8f - 4498 - 8bf2 - 3812b98458d9 .
anthracobune pinfoldi ( in part ) , west , 1980 , p . 518 , pl . 2 . 5 .
anthracobune pilgrimi , west , 1981 ; west , 1983 ( in part ) , p . 367 , fig . 5 , fig . 6 ; west , 1984 ( in part ) , p . 187 , fig . 4 , fig . 5 .
adams consensus of all trees recovered from parsimony analyses that included some ordered multistate characters , with continental geography ( see states in upper left - hand corner ) optimized onto the tree using parsimony ( relationships among extant taxa were constrained by a “molecular scaffold” ) .
bootstrap support for clades derived from each analysis contributing to the consensus is depicted by colored circles . a1 = atlantogenata constraint , transitions between polymorphic and “fixed” states in ordered morphoclines weighted as 0 . 5 steps ; a2 = atlantogenata constraint , transitions between polymorphic and “fixed” states in ordered morphoclines weighted as one step ; e1 = exafroplacentalia constraint , transitions between polymorphic and “fixed” states in ordered morphoclines weighted as 0 . 5 steps ; e2 = exafroplacentalia constraint , transitions between polymorphic and “fixed” states in ordered morphoclines weighted as one step . across all trees , anthracobunids and desmostylians were placed as perissodactyls , along with two enigmatic asian taxa , the late paleocene “condylarth” radinskya and early eocene cambaytherium .
( d ) bar diagram of bone compactness , a quantification of the amount of bone per midshaft cross - section , compared here between fossil and extant ungulates .
c values for enamel samples of early and middle eocene mammals from india and pakistan .
results shown as mean ±s . d . for the sample populations . data from ( a ) early eocene and ( b , c ) middle eocene taxa from india and pakistan . circles , perissodactyls ; red circles , anthracobunids ; squares , artiodactyls ; triangles , cetaceans ; inverse triangles , creodonts ; diamonds , condylarths . see suppl . info . for details .
pilgrimella pilgrimi , wells and gingerich , 1983 ( in part ) , p . 122 , fig . 2c–d ; kumar , 1991 ( in part ) , p . 230 .
etymology . specific epithet is from occidentalis , latin for west , in honor of robert m . west , who discovered and described the holotype .
holotype . h - gsp 1981 , mandibles with left p1 - m3 and right p4 - m3 ( fig . 2p–q ) .
type locality . h - gsp locality 62 , ganda kas area , punjab province , pakistan .
formation and age . kuldana formation , kala chitta hills of northeastern pakistan , early lutetian in age ( ≈48 ma ) .
diagnosis . differs from other anthracobunids in exhibiting the following combination of features : ( i ) lower molars broad , ( ii ) lower m3 short , and ( iii ) angular process of the mandible long but shorter than that of anthracobune .
description . the most complete specimen for this species is h - gsp 1981 , a left and right mandible not larger than a . wardi ( h - gsp 96258 and 96434 ) , but with lower molars much broader than a . wardi . h - gsp 1981 lacks the angular process , due to post - mortem damage but it is preserved in h - gsp 96214 , a specimen with an erupting m3 . in this specimen , the angular process is 80 % as long , rostro - caudally , as in a . wardi . the angular process is not preserved in h - gsp 96149 , but its root is preserved and the slope thereof indicates that the process is small h - gsp 96149 has extremely worn teeth , but its molars are similar in size to h - gsp 1981 ( fig . s1 ) , premolars and canine are larger and the depth and robusticity of the jaw is much greater than in h - gsp 1981 . the wear stage is most consistent with advanced age , and makes it unlikely that the smaller angular process is the result of young age . eventually this specimen may be shown to pertain to a new species of obergfellia .
few upper molars have been discovered for o . occidentalis . h - gsp 538 is a large tooth , and an interstitial facet on its posterior side indicates that it is not an m3 . m1 and m2 of p . pilgrimi do not reach this size , and we therefore refer this tooth to p . obergfelli .
discussion . west [ 10 ] , [ 93 ] recognized that his collection of specimens from the ganda kas region of pakistan contained two species of anthracobunids from the freshwater part of the formation and used the names anthracobune pilgrimi and lammidhania wardi for them . unfortunately , the type specimen for l . wardi is an isolated trigonid , and pertains to the same species as several large complete lower jaws collected by us ( h - gsp 96258 and 96434 ) , and these match upper dentition material referred to pilgrimella pilgrimi ( the type of which is an upper molar ) . hence a new genus and species name is needed . a specimen described by west , h - gsp 1981 , is the most appropriate holotype for this species .
discussion . wells and gingerich [ 1 ] described jozaria palustris on the basis of a single specimen which shows the diagnostic features of anthracobunids . the lower premolars of this specimen are large compared to the molars , confirming it as distinct from other anthracobunid species and genera .
parsimony analyses were first run to determine the most parsimonious placement of xenarthra ( represented in this analysis by dasypus ) given different placements in placental phylogeny – i . e . , as either the sister group either of afrotheria ( the atlantogenata hypothesis ) or of laurasiatheria + euarchontoglires ( the exafroplacentalia hypothesis ) , as this is one of the nodes that was not convincingly resolved by meredith et al . ' s [ 27 ] analysis , and continues to be debated in the literature on placental phylogenetics [ 94 ] . a placement with afrotheria was more parsimonious [ ( 3704 . 5 versus 3716 steps ( i . e . , constraint a1 versus e1 , with 0 . 5 weighting of ordered multistate characters with intermediate polymorphic states ; fig . s2a versus fig . s2c ) and 5758 versus 5773 steps ( a2 versus e2 , no weighting of ordered multistate characters with intermediate polymorphic states , fig . s2b versus fig . s2d ) ] , and all subsequent alternative constraints accordingly employed an atlantogenata , rather than an exafroplacentalia , constraint .
the placement of cambaytherium within perissodactyla is consistent with bajpai et al . ' s [ 38 ] , [ 91 ] assessment of the genus . confusion surrounding the dental similarities that cambaytherium shares with perissodactyls and anthracobunids [ 102 ] is resolved by the placement of cambaytheriids and anthracobunids as closely related sister taxa of crown perissodactyla , thereby indicating that similarities to tethytheres are entirely due to convergence . bajpai et al . [ 38 ] argued that the middle eocene european genus hallensia , which was originally identified as a “condylarth” [ 50 ] and later placed within perissodactyla as either a possible equoid [ 51 ] , [ 103 ] , might be a cambaytheriid . in our analyses hallensia was usually placed as a stem perissodactyl ( a1 , e1 , e2 ) or in an unresolved position relative to these taxa ( a2 ) . the monophyly of a clade containing anthracobunidae , cambaytherium , hallensia , radinskya , and “advanced” perissodactyls is supported by 11 unambiguous synapomorphies ( table s2 ) . the clade containing anthracobunids , cambaytheriids , radinskya , crown perissodactyla , and hallensia was reconstructed as having been equivocally of either asian or european origin ( table s3 ) , while the anthracobunid - desmostylian - hippomorph - tapiromorph clade was optimized as having been of asian origin .
we quantified the midshaft cross - sectional geometries of limb and rib bones of anthracobunids and other ungulates . long bone and rib cross - sectional phenotypes are a reliable indicator of vertebrate habitat , with most taxa occupying a shallow - water habitat displaying the greatest amount of bone per cross - section via thickened cortices and / or bone - filled medullary cavities [ 71 ] , [ 104 ] , [ 105 ] . in contrast , bones of terrestrial taxa typically display thin cortices and vacant medullary cavities . bone compactness values , a combined measure of cortical and medullary bone area [ 72 ] , indicate that anthracobune limb bones display values between 0 . 85–0 . 96 . this range is greater than values observed in a sample of fossil and extant artiodactyls and perissodactyls ( 0 . 42–0 . 83 ) , with the exception of modern hippopotamus ( 0 . 94 , 0 . 96 ) and rhinoceros ( 0 . 64–0 . 85 ) , and semi - aquatic pakicetid and remingtonocetid eocene cetaceans ( 0 . 93–0 . 96 ) ( fig . 4 ) . similar results were documented in rib elements ( table s7 ) .
we also studied the oxygen and carbon isotope composition of structural carbonate in tooth enamel [ 75 ] , [ 106 ] for anthracobunids and coeval taxa from the early and middle eocene of india and pakistan ( fig . 5 , table s8 , table s9 ) . mammal fossils sampled from early eocene localities ( fig . 5c ) , including cambaytherium , spanned a narrow range of δ 18 o values , and are consistent with occupation of a wet and forested habitat . enamel δ 13 c values showed more than 3‰ range in individual values ( −12 . 4‰ to −9 . 1‰ ) , and , when corrected to present - day atmospheric δ 13 c values [ 107 ] , suggest foraging in a relatively wet environment . conversely , taxa sampled from middle eocene deposits of northern pakistan displayed a much wider range of δ 18 o values ( 21 to 27‰ ) and much higher enamel δ 13 c values ( fig . 5b , c ) .
two specimens of a . wardi collected from middle eocene deposits of northern india had low enamel δ 18 o values ( 21 . 3‰ and 21 . 8‰ ) that approached values for aquatic taxa , but also overlapped with values for terrestrial rhinocerotoids ( 20 . 6‰ to 24 . 7‰ ) from the same formation . these low enamel δ 18 o values are more consistent with a freshwater habitat for these individuals , but , again , low sample size complicates differentiating δ 18 o values for these specimens from δ 18 o values for taxa assumed to be fully terrestrial .
anthracobunids have not previously been incorporated into taxonomically broad phylogenetic analyses of morphological data that incorporate the molecular evidence for placental supraordinal relationships . we tested for the possibility that anthracobunidae and various laurasian “condylarths” ( phenacodontids , louisinids ) are not members of afrotheria by adding a number of living and extinct laurasiatherians to a morphological character matrix that has previously been employed to reconstruct afrotherian phylogeny [ 19 ] , [ 35 ] , [ 108 ] . we constrained relationships of extant taxa , based largely on the results of meredith et al . [ 27 ] , using the “molecular scaffold” technique proposed by springer et al . [ 109 ] . we also employed several alternative constraints to obtain tree lengths for competing topologies .
when continental geography is mapped onto the adams consensus in fig . 3 , crown and advanced stem perissodactyls ( including cambaytherium and radinskya ) are optimized as having originated in asia ( fig . 3 ) , while the perissodactyl stem lineage is of ambiguous ( but certainly laurasian ) continental origin . later colonization of north america by crown perissodactyls was presumably facilitated by global warming at the paleocene - eocene boundary [ 111 ] , [ 112 ] . afrotheria is unambiguously of afro - arabian origin on the adams topology , and the recently described paleocene african “condylarth” ocepeia [ 58 ] was placed as either a stem paenungulate or as a stem atlantogenatan , depending on whether atlantogenata or exafroplacentalia was constrained ( respectively ) . the consistent recovery of the european louisinids paschatherium and teilhardimys as stem perissodactyls challenges the view that they are afrotherian macroscelideans [ 62 ] .
some early tethytheres are thought to have had a semi - aquatic lifestyle ( proboscideans [ 81 ] , [ 116 ] , desmostylians [ 117 ] , sirenians [ 104 ] , [ 118 ] – [ 120 ] ) , but no evidence suggests that stem perissodactyls occupied an aquatic habitat . to reconstruct the ancient habitat preferences of anthracobunids , we analyzed bone geometry of the postcranial skeleton as well as stable isotope values within tooth enamel .
we quantified the midshaft cross - sectional geometries of limb and rib bones of anthracobunids and other ungulates . anthracobune limb bones displayed bones that were more hyperostotic and therefore more compacted compared to most extant ungulates . exceptions included extant taxa that are known to frequently swim or wallow in shallow water habitats ( i . e . , hippopotamus and rhinoceros ) [ 121 ] , and fossil eocene cetaceans that are thought to be semi - aquatic and also occupy shallow water habitats ( i . e . , pakicetid and remingtonocetids ) ( fig . 4 ) . thus , anthracobunids display a variety of skeletal modifications consistent with taxa that exploit shallow - water habitats , including thickened postcranial elements and widened elements of the autopodium , in addition to large body size .
we also studied the oxygen and carbon isotope composition of structural carbonate in tooth enamel [ 75 ] , [ 106 ] for anthracobunids and coeval taxa ( fig . 5 ) . the anthracobunids recovered from the kuldana formation ( anthracobune and obergfellia ) displayed enamel δ 18 o values suggesting some evidence of aquatic habits , and enamel δ 13 c values showed they fed on land . anthracobunids recovered from northern india displayed enamel δ 18 o values consistent with occupation of freshwater habitats . although all sampled anthracobunids may have had some aquatic preferences , stable isotopic evidence alone is inconclusive as there is overlap in range of isotopic values between terrestrial and aquatic taxa .
taken together , skeletal and isotopic evidence are most consistent with an interpretation of anthracobunids ( especially anthracobune ) as having had ecological preferences similar to those of modern rhinos and tapirs , but not exhibiting the same degree of commitment to aquatic habits as hippopotamus [ 84 ] , [ 122 ] . most rhinos and tapirs have a restricted range near permanent bodies of water in which they frequently wallow and wade [ 121 ] , and obtain most of their drinking water [ 84 ] , [ 122 ] . modern rhinos may be a good model for the anthracobunid lifestyle as both display thickened limbs and intermediate oxygen isotope values relative to hippopotamus and terrestrial ungulates .
stable isotopic evidence also showed mammal fossils sampled from early eocene localities ( fig . 5c ) occupied and foraged within a wet and forested habitat . conversely , taxa sampled from middle eocene deposits of northern pakistan ( fig . 5b , c ) indicated shifts in the climate toward increased aridity . this climate change was most likely caused by the northward movement of the indian plate , which pushed these areas out of an equatorial zone of high precipitation in the middle eocene [ 123 ] , [ 124 ] . vegetation would have opened up , and shifted to a drier , tropical forest or savanna habitat .
based on new dental , cranial , and postcranial material , our phylogenetic analysis is the first to suggest that anthracobunids are stem members of the laurasiatherian order perissodactyla , rather than members of the afrotherian clades tethytheria , proboscidea , or sirenia . the strictly asian distribution of anthracobunids is consistent with the isolation of afrotherian and laurasiatherian clades through the early paleogene [ 19 ] , [ 113 ] , while the phylogenetic result is consistent with the recent recovery of very primitive basal stem proboscideans such as phosphatherium [ 125 ] and eritherium [ 18 ] , from the early eocene and paleocene of africa . analyses of postcranial bones and stable isotopes also indicate that anthracobunids were large and lumbering , and suggest at least some taxa spent time in the water similar to modern rhinos , and that all known anthracobunids fed on land . our results therefore identify an old world radiation of large , non - cursorial , partly aquatic perissodactyls that convergently came to occupy a basal tethythere - like niche on the northern coast of the tethys sea .


thomas 's water mouse ( rheomys thomasi ) is a species of rodent in the family cricetidae found in el salvador , guatemala , and mexico at altitudes of 400 to 2700 m .

it lives near forest streams and is semiaquatic ; its carnivorous diet includes both invertebrates and small vertebrates .

the conservation status of the species is rated as " near threatened " because of the small size of its range and the threat of degradation of its habitat , including the water quality of the streams it lives along .

thomas ' s water mouse

the thomas ' s water mouse is listed as least concern ( lr / lc ) , lowest risk . does not qualify for a more at risk category . widespread and abundant taxa are included in this category , on the iucn red list of threatened species
stirton , r . a . 1944 . tropical mammal trapping i : the water mouse rheomys . journal of mammalogy 25 : 337 - 343 .
a young / baby of a thomas is called a ' pinkie , kitten or pup ' . the females are called ' doe ' and males ' buck ' . a thomas group is called a ' nest , colony , harvest , horde or mischief ' .
stream degradation and water quality degradation are major threats due to its reliance on stream habitats . other potential threats include landslides .
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note : data presented in infonatura at urltoken were updated to be current with natureserve ' s central databases as of april 2007 . note : this report was printed on .
this rodent is found along streams through most forest types and second growth ( reid 1997 ) . it is sometimes found on very small streams and steeps in forests , and may be more generalized in diet and habits than other water mice . in el salvador , foods eaten include insects , birds , salamanders , mammals ( possibly another rheomys ) , and starchy pulp ( stirton 1944 ) . one individual from guatemala had eaten stonefly and mayfly larvae , beetles , and part of a catfish ( hooper 1968 ) . a captive mouse readily consumed small fish and aquatic insect larvae ( reid 1997 ) .
information indicates evolutionary relationships at both the ordinal and the family level . this indispensable reference work belongs in public and academic libraries throughout the world and on the shelf of every biologist who works with mammals .
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wilson and reeder ' s mammal species of the world is the classic reference book on the taxonomic classification and distribution of the more than 5400 species of mammals that exist today . the third edition includes detailed information on nomenclature and , for the first time , common names . each concise entry covers type locality , distribution , synonyms , and major reference sources . the systematic arrangement of information indicates evolutionary relationships at both the ordinal and the family level . this indispensable reference work belongs in public and academic libraries throughout the world and on the shelf of every biologist who works with mammals .
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disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
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wilson and reeder ' s mammal species of the world is the classic reference book on the taxonomic classification and distribution of the more than 5400 species of mammals that exist today . the third edition includes detailed information on nomenclature and , for the first time , common names . each concise entry covers type locality , distribution , synonyms , and major reference sources . the systematic arrangement of
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i thought you might be interested in this item at urltoken title : mammal species of the world : a taxonomic and geographic reference author : don e wilson ; deeann m reeder publisher : baltimore : johns hopkins university press , ©2005 . isbn / issn : 0801882214 9780801882210 0801882389 9780801882388 0801882397 9780801882395 oclc : 57557352
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mcknight , m . ( global mammal assessment team ) & amori , g . ( small nonvolant mammal red list authority )
justification : listed as near threatened because its extent of occurrence is probably not much greater than 20 , 000 km 2 , and the extent and quality of its habitat are probably declining , thus making the species close to qualifying for vulnerable under criterion b1b ( iii ) .
this species occurs in the highlands of south méxico ( chiapas ) , guatemala , el salvador and honduras ( musser and carleton 2005 ) . it is found from 400 m to 2 , 700 m ( reid 1997 ) .
occurs in el triunfo national park in mexico and in el imposible national park in el salvador . expected to occur in sierra de las minas national park in guatemala .
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hooper , e . t . 1968 . habitats and food of amphibious mice of the genus rheomys . journal of mammalogy 49 : 550 - 553 .
musser , g . g . and carleton , m . d . 2005 . superfamily muroidea . in : d . e . wilson and d . a . reeder ( eds ) , mammal species of the world : a geographic and taxonomic reference , pp . 894 - 1531 . the john hopkins university press , baltimore , usa .
reid , f . 2009 . a field guide to the mammals of central america and southeast mexico . oxford university press , new york , usa .
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musser , guy g . , and michael d . carleton / wilson , don e . , and deeann m . reeder , eds .
mammal species of the world : a taxonomic and geographic reference , 3rd ed . , vol . 2
mammal species of the world : a taxonomic and geographic reference , 2nd ed . , 3rd printing
with contributions by bernadette n . graham , adam p . potter , and mariana m . upmeyer
infonatura species index : 351 - 400 of 479 records in family muridae of order rodentia .
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citation for mammal range maps : patterson , b . d . , g . ceballos , w . sechrest , m . f . tognelli , t . brooks , l . luna , p . ortega , i . salazar , and b . e . young . 2005 . digital distribution maps of the mammals of the western hemisphere , version 2 . 0 . natureserve , arlington , virginia , usa .
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the finescale razorbelly minnow ( salmophasia phulo ) is a species of ray-finned fish in the genus salmophasia .

it is a species of freshwater fish native to bangladesh and india .

it lives in the lower reaches of various bodies of water including rivers , canals , ponds , and ditches .

with a maximum length of only 12 centimetres ( 4.7 in ) , the fish is of little commercial or dietary value to humans .

finescale razorbelly minnow

latin , salmo , plinius = salmon + greek , stoma = mouth ( ref . 45335 )
maturity : l m ? range ? - ? cm max length : 12 . 0 cm tl male / unsexed ; ( ref . 41236 )
occurs in the lower reaches of rivers , ponds , beels , ditches and canals ( ref . 41236 ) .
menon , a . g . k . , 1999 . check list - fresh water fishes of india . rec . zool . surv . india , misc . publ . , occas . pap . no . 175 , 366 p . ( ref . 41236 )
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 5001 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 00724 ( 0 . 00250 - 0 . 02102 ) , b = 3 . 06 ( 2 . 82 - 3 . 31 ) , in cm total length , based on lwr estimates for this ( sub ) family - body shape ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 3 . 2 ±0 . 4 se ; based on size and trophs of closest relatives
resilience ( ref . 69278 ) : high , minimum population doubling time less than 15 months ( preliminary k or fecundity . ) .
vulnerability ( ref . 59153 ) : low vulnerability ( 23 of 100 ) .
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occurs in the lower reaches of rivers , ponds , beels , ditches and canals ( ref . 41236 ) .
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cyprinus phulo hamilton , 1822 , fishes of the ganges , p . 262 .
oxygaster phulo rahman , 1974 , banglaesh j zool . 2 ( 2 ) . p . 193 .
samostoma phulo phulo banarescu , 1968 , rev . roum . biol . zool . 13 ( 1 ) . p . 5 .
this species is a surface feeder and available distributed in the study area . various types of dip nets are used for its fishing .
bhuiyan , a . l . 1964 . fishes of dacca . asiatic society of pakistan , dacca . 48 pp .
day , f . 1878 . fishes of india , being a natural history of fishes known to inhabit the seas and freshwaters of india , burma and ceylon . william dawson & sons ltd . , london , vol . i : 778 pp .
hamilton , f . , 1822 . an account of the fishes found in the river ganges and its branches , edinburgh & london , fishes ganges , 262 p .
rahman , 1974 , banglaesh j zool . 2 ( 2 ) . p . 193 .
rahman , a . k . a . 1989 . freshwater fishes of bangladesh . the zoological society of bangladesh , department of zoology , university of dhaka , dhaka - 1000 . 364 pp .
rahman , a . k . a . 2005 . freshwater fishes of bangladesh ( second edition ) . the zoological society of bangladesh , department of zoology , university of dhaka , dhaka - 1000 . 394 pp .
talwar , p . k . and jhingran , a . g . , 1991 . inland fishes of india and adjacent countries . volume 1 and 2 . oxford & ibh publishing co . pvt . ltd . new delhi , calcutta . 1158pp .
ex - student , department of fisheries , university of rajshahi , rajshahi - 6205 , bangladesh . more . . .
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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .