ontocetus is an extinct genus of walrus , an aquatic carnivoran of the family odobenidae , endemic to coastal regions of the southern north sea and the southeastern coastal regions of the u.s. during the miocene-pleistocene .

it lived from 13.6 mya — 300,000 years ago , existing for approximately 13.3 million years .

ontocetus

belongs to ontocetus according to n . kohno and c . e . ray 2008
comparison of type specimen ontocetus emmonsi with a tooth from type specimen albicetus oxymycterus .
ontocetus emmonsi is an extinct species of walrus which inhabited the north atlantic during the pliocene .
like the modern walrus , ontocetus emmonsi had large tusks . these tusks are the most commonly found type of fossil for this species .
j . leidy . 1859 . [ remarks on dromatherium sylvestre and ontocetus emmonsi ] . proceedings of the academy of natural sciences of philadelphia 1859 : 162
comparison of type specimen < i > ontocetus emmonsi < / i > with a tooth from type specimen < i > albicetus oxymycterus < / i > .
the diversity of tusked walruses ( odobeninae ) ; note the much smaller size of aivukus and protodobenus , and the gigantic size of ontocetus emmonsi / alachtherium cretsii .
figure 3 . uf 3274 , upper tusk of ontocetus emmonsi from de soto lakes , florida . medial view , showing characteristic osteodentine core visible near the proximal end .
full reference : j . leidy . 1859 . [ remarks on dromatherium sylvestre and ontocetus emmonsi ] . proceedings of the academy of natural sciences of philadelphia 1859 : 162
leidy , j . 1859 . remarks on dromatherium sylvestre and ontocetus emmonsi . proceeding of the academy of natural sciences of philadelphia , 1859 : 162 . ( download pdf )
ontocetus emmonsi : usnm 329064 , a tooth ( tusk ) . its type locality is murfreesboro , which is in a zanclean shallow subtidal sandstone in the yorktown formation of north carolina .
ontocetus emmonsi is an extinct species of walrus which inhabited the north atlantic during the pliocene . ontocetus emmonsi is the type species and only species in the genus ontocetus . it is placed within the family odobenidae which includes only one living species , the walrus odobenus rosmarus ( demere , 1994 ) . like odobenus , ontocetus is included in the subfamily odobeninae , or “tusked walruses” and is characterized by large , continuously growing tusks , or modified canine teeth ( kohno and ray , 2008 ) . despite strong support for relationships within odobenidae , it is currently unknown as to whether this family is more closely related to phocids ( true seals ) , or otariids ( eared seals ) ( berta and churchill , 2012 ) . initially thought to be a cetacean ( leidy , 1859 ) , the genus name ontocetus comes from the greek ontos , meaning being , and ketos , meaning whale or large sea animal ( kohno and ray , 2008 ) .
figure 6 . uf 254922 , partial upper canine of ontocetus emmonsi from the quality aggregates pit 8 . medial view , showing root of the upper incisor and a portion of the premaxilla .
figure 4 . uf 256416 , upper tusk of ontocetus emmonsi from the quality aggregates locality 8a in cross sectional view . osteodentine core , surrounded by smooth orthodentine , and a thin layer of cementum .
a pleistocene skull figured and referred to odobenus huxleyi ( = odobenus rosmarus ) , and the braincase of alachtherium cretsii / ontocetus emmonsi named as the new species trichecodon antverpiensis by rutten ( 1907 ) .
this paper aims to redescribe ontocetus oxymycterus kellogg [ 28 ] , a large fossil sperm whale described from the monterey formation of santa barbara county , california , u . s . a ( fig 1 ) . the type specimen consists of an incomplete rostrum , and isolated fragment of the upper rostrum , both mandibles with some large dental roots , and several isolated , but associated incomplete teeth ( fig 2 ) . kellogg [ 28 ] tentatively referred the species to the genus ontocetus , based primarily on its large , conspicuous teeth , which he thought resembled ontocetus emmonsi leidy [ 29 ] , a tooth taxon from the neogene of north carolina [ 30 ] ( fig 3 ) .
the paleobiogeography of ontocetus emmonsi was strongly influenced by the changing ocean configuration during the pliocene . ontocetus emmonsi is thought to have originated in the northern pacific ocean and then dispersed into the atlantic via the central american seaway between north and south america in the early pliocene ( kohno and ray , 2008 ) . later dispersal via the north atlantic gyre , a circulating ocean current moving in a clockwise fashion between north america and europe , led to their eventual appearance in europe and north africa ( kohno and ray , 2008 ; naafs et al . , 2010 ) . this dispersal pattern would account for the later appearance of ontocetus emmonsi fossils in european localities relative to those of north america .
an adorable ontocetus emmonsi juvenile maxilla ( complete with mini - tusk ! ) from the pliocene yorktown formation , described and figured by kohno and ray ( 2008 ) , photographed at the usnm . seriously , that is a cute fossil .
figure 2 . uf 125201 . upper tusk of ontocetus emmonsi from the whiddon creek site in the fort meade mine , polk county , florida . lateral view , showing fluting along the length of the tusk and characteristic curved and tapered shape .
with these traits in mind , leidy [ 30 ] later suggested that ontocetus emmonsi represented either a “cetacean like the sperm whale , ” or “perhaps to a walrus - like animal . ” brandt [ 32 ] similarly designated the type species as a toothed whale , but concurred with leidy’s alternative identification ( see spamer et al . [ 33 ] , for a detailed bibliographic history ) . although matsumoto [ 34 ] , shikama et al . [ 35 ] and okazaki [ 36 ] reported cetacean fossil material from japan belonging to this species , all other recent authors have thoroughly discounted any cetacean affinities to the type specimen ontocetus emmonsi . ray [ 37 ] , kohno and ray [ 38 ] , boessenecker and churchill [ 39 ] , and churchill et al . [ 40 ] , among others , have all confirmed the odobenid identity of ontocetus emmonsi . the genus ontocetus remains a valid stem odobenid taxon , with additional species yet to be named and described [ 40 ] .
the holotype tusk fragment of ontocetus emmonsi - and the
parent
specimen of virtually all north atlantic pliocene walruses ( = alachtherium cretsii , alachtherium antwerpiensis , trichecodon huxleyi , trichecodon / odobenus antverpiensis , prorosmarus alleni ) , photographed at the usnm .
following this realization , kohno and ray [ 38 ] provisionally regarded ontocetus oxymycterus as belonging to the genus scaldicetus de bus [ 41 ] , a cosmopolitan fossil physeteroid distinguished primarily by enamel capped and gibbous teeth [ 24 ] . however , this recommendation is unsatisfactory , as the genus scaldicetus is a form taxon representing a number of isolated teeth [ 24 ] that may or may not belong to a single physeteroid taxon [ 22 – 23 ] . moreover , the type specimen of ontocetus oxymycterus exhibits a number of diagnostic morphological characters , including proportionately large dentition ( relative to rostral size ) , retention of tooth enamel , and a closed mesorostral groove , which combined distinguish it from all other described physeteroid genera . because the type species of ontocetus is not a cetacean , we propose a new generic name , albicetus , for the species of ontocetus oxymycterus [ 28 ] , and provide a redescription of the type specimen , along with body size estimates and a phylogenetic analysis to resolve its relationship and evolutionary context among physeteroidea .
ontocetus leidy , 1859 ( miocene walrus ) the - cetus means
whale
, but the tooth it was named for belonged to a walrus . onto - probably was intended to mean
existing
( it can also mean
dung
) , adding to the error .
alachtherium africanum was named by geraads ( 1997 ) . it is not extant . its type specimen is aao - 30392 . it was recombined as odobenus africanus by bosscha erdbrink and van bree ( 1999 ) ; it was synonymized subjectively with ontocetus emmonsi by kohno and ray ( 2008 ) .
historically , many authors have attributed pliocene walruses from the atlantic to numerous different genera , largely based on isolated and fragmentary tusk fossils ( see kohno and ray , 2008 for a full discussion ) . florida specimens were typically referred to trichecodon huxleyi ( e . g . , ray , 1960 ) . recent consideration of individual and ontogenetic variation , as well as sexual dimorphism and size disparities among different age groups has led to the synonymy of all atlantic pliocene walruses into a single genus , ontocetus ( kohno and ray , 2008 ) . these authors noted that much of the variation between different taxa could be observed within one population of ontocetus emmonsi from the lee creek mine in eastern north carolina . furthermore , many of the characters used to establish these new taxa could also be found in the holotype specimen of ontocetus emmonsi ( leidy , 1859 ; kohno and ray , 2008 ) .
ontocetus emmonsi became extinct before the beginning of the pleistocene , and more importantly , before the first appearance of odobenus rosmarus in the atlantic . it is therefore unlikely that competition led to its eventual extinction . instead , environmental changes such as changing ocean temperature are thought to be the cause ( kohno and ray , 2008 ) .
in addition to fossil occurrences in florida , ontocetus emmonsi fossils have been found along the eastern coast of the united states in georgia , south carolina , north carolina and virginia , as well as europe and morocco ( kohno and ray , 2008 ) . the type locality is near the town of murfreesboro in hertford county , north carolina .
it’s admittedly a controversial decision , and some other pinniped workers have expressed the notion that alachtherium cretsii is a better name as it is founded upon a complete mandible that inherently preserves more morphological information and is thus certainly more diagnostic than a tusk . i’m on the fence ; on one hand , if the tusk morphology proves in the long run to really be that distinctive ( and so far , it seems to ) , then ontocetus emmonsi works ; on the other hand , an isolated fragmentary tusk may not be diagnostic and in cetacean paleontology most species named off of isolated teeth were shit - canned a long time ago . my mind isn’t completely made up , and i think both camps in favor of ontocetus emmonsi or alachtherium cretsii have decent arguments .
figure 5 . uf 160756 ( cast of usnm 475482 ) , partial dentary of ontocetus emmonsi from the martin marietta berkeley quarry , south carolina . lateral view ( above ) , occlusal view ( below ) . teeth present are the third incisor , lower canine , and three lower premolars . note the increasing size of the premolars and the anterior incline of the first premolar in lateral view .
illustration by joseph leidy of the type specimen of ontocetus emmonsi ( usnm 329064 ) ( left ) , published by emmons [ 40 ] , who described the material as originating from “the older miocene of white river [ which ] has furnished remarkable animals remains … [ among ] these remains ruminants are particularly worthy of note . … the cetacean , fig 187 ( 2 ) , is a remarkable form of tooth for this family—having a resemblance to the canine of the hippopotamus .
this illustration is contrasted with an isolated tooth from albicetus oxymycterus ( usnm 10923 ) ( right ) , with the layers of the dentition labeled . scale bar measures 5 cm . a 3d model of the type specimen ontocetus emmonsi is now available for viewing and download on the smithsonian x 3d website ( urltoken ) .
the paleoecology of ontocetus emmonsi is inferred to be similar to that of odobenus rosmarus , although it inhabited far warmer waters . however , stronger sexual dimorphism in ontocetus emmonsi suggests that this species may have exhibited stronger polygynous breeding behavior , more comparable to that of otariid seals ( kohno and ray , 2008 ) . additionally , size differences between tusk fossils found in florida and further north along the atlantic coast show that florida fossils tend to be larger overall , and most likely represent mature male walruses . the tusks from the carolinas and virginia tend to be smaller and are typically referred to females and juveniles . this trend is interpreted to be the result of differential migration patterns among males and females , with the timing of seasonal migration being offset for each ( kohno and ray , 2008 ) .
ontocetus emmonsi was named by leidy ( 1859 ) [ cast ansp 11255 ] . its type specimen is usnm 329064 , a tooth ( tusk ) , and it is a 3d body fossil . its type locality is murfreesboro , which is in a zanclean shallow subtidal sandstone in the yorktown formation of north carolina . it was recombined as hoplocetus emmonsi by trouessart ( 1898 ) and trouessart ( 1904 ) .
tusk fossils are by far the most common ontocetus fossils to find , particularly in florida ( figs . 2 - 3 ; ray , 1960 ; hulbert and ray , 2001 ) . partial tusks can be identified against other common florida fossils with the same general shape and size , such as dugong ribs and large sperm whale teeth , by their characteristic osteodentine core , which has a globular texture . surrounding this core is a layer of smooth orthodentine , and the outermost part of the tusk is composed of a very thin layer of cementum ( fig . 4 ; kohno and ray , 2008 ) . tusks of ontocetus emmonsi differ from other odobenine genera in having a stronger posterior curvature and taper , exhibiting fluting ( grooves ) along their length , and being more compressed laterally ( kohno and ray , 2008 ) .
prorosmarus alleni was named by berry and gregory ( 1906 ) [ no type specimen number listed by berry and gregory ( 1906 ) . ] . its type specimen is usnm 9343 and is not a trace fossil . its type locality is yorktown , which is in a zanclean / piacenzian marine horizon in the yorktown formation of virginia . it was synonymized subjectively with ontocetus emmonsi by kohno and ray ( 2008 ) .
the type specimen of ontocetus emmonsi ( now usnm 329064 ) is represented by a single , large , laterally compressed , tusk - like tooth , which was originally deposited in 1860 at williams college , williamstown , massachusetts , and was first reported in a textbook illustration by emmons [ 31 ] . the same year , leidy [ 29 ] formally designated this specimen as the holotype of ontocetus emmonsi ( fig 3 ) . although the exact provenance of this tooth is unknown , leidy [ 30 ] mentioned “miocene [ sic ] deposits of north carolina , ” likely communicated to him by emmons . the type specimen ( usnm 329064 ) is incomplete , but it exhibits long , thick , and cambered striae of dentine , with a subrectangular outline in transverse section ( fig 3 ) . these features are utterly absent from all known physeteroid teeth , which are typically conical towards the apex , cylindrical in transverse section , and gibbous towards the root .
the greatest number of specimens found in florida derive from commercial shell pits on either side of interstate highway 75 near sarasota ; macasphalt shell pit on the west side ; richardson road ( later quality aggregates , and then smr aggregates ) shell pit on the east side ( figs . 4 , 6 ) . fossils of ontocetus emmonsi from these quarries come from beds 10 and 11 , which are pliocene in age . all walrus fossils from florida are regarded as belonging to this species .
kohno and ray ( 2008 ) further considered prorosmarus alleni to be relatively similar to the type mandible of a . cretsii , and figured a new mandible that is intermediate between the two , indicating that the perceived absence of an upturned symphysis is probably an ontogenetic feature . they also pointed out that the holotype tusk of ontocetus emmonsi is a walrus , and that it and the holotype tusk of trichecodon huxleyi both share an oval , transversely compressed cross section , longitudinal fluting , greater curvature , and are more tapered in contrast to the more elongate , straighter , smoother tusks of odobenus rosmarus , which also have more of a circular cross section . most significantly , they identify that all tusks found in sediments of pliocene age yielding remains of alachtherium / trichecodon / prorosmarus all conform to this morphology . they showed some bivariate plots of atlantic pliocene walrus tusks , and showed conclusively that all of these tusks conform to similar proportions , and cluster together to the exclusion of odobenus rosmarus ( kohno and ray 2008 : fig . 27 ) . in light of this information , they synonymized all pliocene atlantic walruses with ontocetus emmonsi .
< p > illustration by joseph leidy of the type specimen of < i > ontocetus emmonsi < / i > ( usnm 329064 ) ( left ) , published by emmons [ < a href =
urltoken
target =
_ blank
> 40 < / a > ] , who described the material as originating from “the older miocene of white river [ which ] has furnished remarkable animals remains … [ among ] these remains ruminants are particularly worthy of note . … the cetacean , fig 187 ( 2 ) , is a remarkable form of tooth for this family—having a resemblance to the canine of the hippopotamus .
this illustration is contrasted with an isolated tooth from < i > albicetus oxymycterus < / i > ( usnm 10923 ) ( right ) , with the layers of the dentition labeled . scale bar measures 5 cm . a 3d model of the type specimen < i > ontocetus emmonsi < / i > is now available for viewing and download on the smithsonian x 3d website ( < a href =
urltoken
target =
_ blank
> urltoken < / a > ) . < / p >
one of the most fascinating advances was the discovery tusks and crania assignable to odobenus from the upper pliocene of japan . aside from the aforementioned tusk described by kohno et al . ( 1995 ) , a nearly complete skull of odobenus sp . with tusk dredged from the sea of okhotsk was reported by miyazaki et al . ( 1992 ) , who found that it was associated with late pliocene microfossils . these finds indicate that while valenictus was hanging out in warm waters along the california and baja california margin , and while ontocetus / alachtherium was proliferating across virtually the entire north atlantic , the modern walrus had already evolved in the western north pacific .
alachtherium crestsii was named by du bus ( 1867 ) [ no type specimen listed by du bus ( 1867 ) . no type specimen listed by du bus ( 1867 ) . ] . its type specimen is irsnb m . 168 and is a 3d body fossil . its type locality is anvers , lillo formation , which is in a piacenzian offshore sandstone in the lillo formation of belgium . it is the type species of alachtherium . it was corrected as alachtherium cretsii by vanden broeck ( 1874 ) , palmer ( 1904 ) , kellogg ( 1922 ) , deméré ( 1994 ) and deméré et al . ( 2003 ) ; it was synonymized subjectively with trichechus rosmarus by trouessart ( 1898 ) ; it was recombined as odobenus cretsii by bosscha erdbrink and van bree ( 1999 ) ; it was synonymized subjectively with ontocetus emmonsi by kohno and ray ( 2008 ) .
an important contribution towards the evolutionary history of odobenines was published by kohno et al . ( 1995 ) , who reported several tusks from the pliocene of japan . following the definition of deméré ( 1994 ) , they identified a number of tusks with globular dentine from lower and upper pliocene localities in japan , which they identified to the tribe odobenini . most of these have an oval - shaped cross section , are more highly curved and tapering than odobenus rosmarus , and some possess longitudinal fluting – and therefore compare well with tusks of the alachtherium - ontocetus - prorosmarus - trichecodon morphotype ( see below ) . critically , they identified a single tusk of odobenus sp . from the upper pliocene ( see section on odobenus for more on this ) . the importance here is that , based on tusks , a minimum of two species of odobenini appear to have coexisted in japan during the pliocene .
trichecodon huxleyi was named by lankester ( 1865 ) [ no type specimen number listed in lankester 1865 ; nhm 46000 is the lectotype ( kohno , 2008 ) ] . its type specimen is nhm 46000 and is not a trace fossil . its type locality is felixstow , which is in a gelasian foreshore horizon in the red crag formation of the united kingdom . it was recombined as trichechus huxleyi by lankester ( 1880 ) , newton ( 1891 ) , trouessart ( 1898 ) and trouessart ( 1904 ) ; it was recombined as odobenus huxleyi by hooijer ( 1957 ) , van deinse ( 1964 ) , van der feen ( 1968 ) , deméré ( 1994 ) , bosscha erdbrink and van bree ( 1999 ) and deméré et al . ( 2003 ) ; it was recombined as odobaenus huxleyi by post ( 2004 ) ; it was synonymized subjectively with ontocetus emmonsi by kohno and ray ( 2008 ) .
trichechus antverpiensis was named by rutten ( 1907 ) [ no type specimen number is listed , but the specimen is in the musee de histoire naturelle in brussels . holotype number from kohno ( 2008 ) . no type specimen number is listed , but the specimen is in the musee de histoire naturelle in brussels . holotype number from kohno ( 2008 ) . no type specimen number is listed , but the specimen is in the musee de histoire naturelle in brussels . holotype number from kohno ( 2008 ) . no type specimen number is listed , but the specimen is in the musee de histoire naturelle in brussels . holotype number from kohno ( 2008 ) . no type specimen number is listed , but the specimen is in the musee de histoire naturelle in brussels . holotype number from kohno ( 2008 ) . no type specimen number is listed , but the specimen is in the musee de histoire naturelle in brussels . holotype number from kohno ( 2008 ) . ] . its type specimen is irsnb m . 169 , a partial skull , and it is a 3d body fossil . its type locality is heyst , which is in a calabrian marine horizon in belgium . it was recombined as alachtherium antverpiensis by van deinse ( 1964 ) and repenning and tedford ( 1977 ) ; it was recombined as odobenus antverpiensis by van der feen ( 1968 ) and bosscha erdbrink and van bree ( 1999 ) ; it was synonymized subjectively with alachtherium cretsii by deméré ( 1994 ) ; it was synonymized subjectively with ontocetus emmonsi by kohno and ray ( 2008 ) .
discover a faster , simpler path to publishing in a high - quality journal . plos one promises fair , rigorous peer review , broad scope , and wide readership – a perfect fit for your research every time .
contributed equally to this work with : alexandra t . boersma , nicholas d . pyenson
citation : boersma at , pyenson nd ( 2015 ) albicetus oxymycterus , a new generic name and redescription of a basal physeteroid ( mammalia , cetacea ) from the miocene of california , and the evolution of body size in sperm whales . plos one 10 ( 12 ) : e0135551 . urltoken
this is an open access article , free of all copyright , and may be freely reproduced , distributed , transmitted , modified , built upon , or otherwise used by anyone for any lawful purpose . the work is made available under the creative commons cc0 public domain dedication
data availability : the original third - party 3d scan datasets of usnm 10923 ( as well as usnm 329064 ) , including . obj and . stl files , have been archived in zenodo , urltoken < urltoken > ( doi : 10 . 5281 / zenodo . 23029 ) . the 3d models are also available for visualization and download at the smithsonian x 3d website ( urltoken ) .
funding : this work was supported by vassar college ( ab ) , and the nmnh remington kellogg fund and basis foundation ( ndp ) ( urltoken ) . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
competing interests : ndp is an academic editor for plos one . this does not alter the authors’ adherence to plos one policies on sharing data and materials .
abbreviations : usnm , department of paleobiology , national museum of natural history , smithsonian institution , washington , district of columbia , u . s . a . ; lacm , natural history museum of los angeles county , los angeles , california , u . s . a . ; be - bvm , bob ernst collection , buena vista museum of natural history , bakersfield , california , u . s . a .
a , a map of the state of california , showing santa barbara county and a box indicating the 2015 united states geological survey 7 . 5 minute topographic map for santa barbara quadrangle , used for b and c . b , a general outline of the vicinity , showing the city of santa barbara , with major highways , santa barbara point , and the type locality . dashed box indicates the area in c , a map of the coast around the type locality , with the location of the original santa barbara lighthouse , using a modified basemap from the usgs topographic map ( available at urltoken ) . isobars are 50 meters , and cartographic north for all panels points to the top .
a , the rostrum and mandibles together in ventral view , b in dorsal view , c in right lateral view , d in anterior view , and e in posterior view .
anatomical terminology was taken from mead and fordyce [ 42 ] , with modifications from fitzgerald [ 24 ] and velez - juarbe et al . [ 9 ] .
no permits were required for the described study because the material was collected from an undocumented locality between 1879 and 1909 .
a , rostrum and mandibles in oblique anterior - left lateral view , b , isolated upper rostrum fragment in posterior view , and c , isolated tooth fragment with enamel cap . scale bars measure 5 cm . the 3d model is accessible through the smithsonian x 3d website ( urltoken ) .
we used two methods to estimate the condylobasal length and total length of albicetus . since the type specimen of albicetus lacked other skull measurements needed for partial least squares ( pls ) mulivariate regression equations , both approaches used a bivariate ordinary least squares ( ols ) linear regression with antorbital notch width as a single body length proxy [ 43 ] . the antorbital notch was not preserved on the left side of the rostrum , and thus we collected this measurement by measuring from the right side of the specimen , and then doubled it for a minimum estimate of the antorbital notch width ( also often referred to as width at rostrum base ) ( see s2 fig for a visual guide to measurements ) . the antorbital notch is not preserved in its entirety in the type specimen , and the measurement therefore likely corresponds to a level just slightly anterior to the antorbital notch , though we refer to it as antorbital notch width in this paper .
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 : 828c6915 - 3aa2 - 45ae - 99c0 - e97e7f6e1a0a . 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 , and lockss .
physeteroidea , gray [ 49 ] , sensu velez - juarbe et al . [ 9 ]
albicetus oxymycterus , new combination , urn : lsid : zoobank . org : act : 0a979799 - 2e74 - 4c2e - 9e23 - 8940a036ae18 ,
etymology : combining the latin words albus ( white ) and cetus ( whale ) . the name pays tribute to h . melville [ 50 ] ’s classic american novel moby - dick ; or , the whale . in the novel , melville refers to moby - dick as “the white whale” , a creature of “unwonted magnitude” with a “remarkable hue” and “deformed lower jaw” [ 50 ] . these traits are coincidentally similar to the type specimen of albicetus , a white fossil sperm whale whose jaws have been displaced due to diagenetic processes , providing apt inspiration for the connection to the famous literary whale .
illustrated with a low opacity mask and interpretive line art . tooth fragments retained in the alveoli are emphasized with a nearly opaque white layer . to view the 3d model of the specimen , visit the smithsonian x 3d website at ( urltoken ) .
illustrated with a low opacity mask and interpretive line art . from top : a , left lateral , b , ventral , and c , right lateral views abbreviations : d indicates dorsal direction , l indicates left lateral , a indicates anterior , p indicates posterior , r indicates right lateral . ai 1 indicates first incisor alveoli , ac 1 indicates first canine alveoli , apc 1 indicates first post - canine alveoli , dr indicates alveolus with dental root . the mandibles ( ma ) , maxillae ( mx ) and premaxillae ( pmx ) are all labeled accordingly . tooth fragments in alveoli are emphasized with a nearly opaque white layer . to view the 3d model of the specimen , visit the smithsonian x 3d website at ( urltoken ) .
illustrated with a low opacity mask and interpretive line art . tooth fragments retained in alveoli are emphasized with a nearly opaque white layer . abbreviations : d indicates dorsal direction , l indicates left lateral , a indicates anterior , ai , , indicates incisor alveolus , ac 1 indicates first canine alveoli , apc 1 indicates first post - canine alveoli , dr indicates alveoli with dental roots . to view the 3d model of the specimen , visit the smithsonian x 3d website at ( urltoken ) .
illustrated with a low opacity mask and interpretive line art . tooth fragments retained in alveoli are emphasized with a nearly opaque white layer . to view the 3d model of the specimen , visit the smithsonian x 3d website at ( urltoken ) .
illustrated with a low opacity mask and interpretive line art . a indicates anterior direction , r indicates right lateral direction . the line of dashed line emphasizes the asymmetry in the premaxillae . the dotted line delineates the premaxillary ( pmx ) sac fossae , which are also asymmetrical . to view the 3d model of the specimen , visit the smithsonian x 3d website at ( urltoken ) .
illustrated with a low opacity mask and interpretive line art . to view the 3d model of the specimen , visit the smithsonian x 3d website at ( urltoken ) .
measurements for main rostrum segment , right mandible , and upper rostrum fragment of albicetus oxymycterus ( usnm 10923 ) , in centimeters .
measurements for alveoli in right mandible of albicetus oxymycterus ( usnm 10923 ) , in millimeters .
holotype : usnm 10923 , the incomplete extremity of the rostrum and mandibles , with 10 or 11 roots or portions of teeth in situ in each mandible , several incomplete teeth fragments found loose in the matrix , and a separate piece of the upper rostrum . archival typewritten notes at usnm state that the specimen was first observed by charles o . roe ( 1867–1923 ) when he was a boy , and was collected by him some thirty years later in 1909 , implying an initial discovery as early as 1879 . these notes are consistent with kellogg [ 28 ] ’s report about the type specimen’s discovery around 1884 , and it being subsequently moved to roe’s home in 1909 . the specimen was received by usnm on 16 february 1924 from roe’s wife , after his death in 1923 [ 51 ] .
etymology : according to kellogg [ 28 ] , oxymycterus derives from the greek words oxy ( sharp ) and mycter ( nose ) .
the original santa barbara lighthouse , however , was destroyed in an earthquake on 29 june 1929 [ 53 ] , and kellogg [ 28 ] ’s published coordinates correspond to a location about 5 km directly south of the original lighthouse , offshore in santa barbara bay . the published locality account by kellogg [ 28 ] corresponds today to sea cliffs located near the property of the current lighthouse [ 54 ] , which is 100 m northeast of the original lighthouse , at the following coordinates ( n 34° 23 ' 44
, w 119° 43 ' 23
) . we argue that this general vicinity , within less than a 100 m radius , likely represents the type locality of albicetus oxymycterus ( fig 1 ) .
formation : kellogg [ 28 ] described the stratigraphic provenance of the type specimen of albicetus as a unit of bituminous dolomite in the sea cliffs of santa barbara county , likely belonging the monterey formation . this description is consistent with minor et al . ’s [ 55 ] geologic mapping of the santa barbara coastal plain , which at the reported locality shows the underlying lowest three subunits of the marine siliceous and calcareous mudstone and shale belonging to the monterey formation . these marine rocks are mapped as the lower calcareous unit of the monterey formation [ 55 ] . overlying marine terrace deposits of pleistocene age in this area do not match the lithology of the matrix with usnm 10923 .
age : minor et al . [ 55 ] summarized biostratigraphic findings for lower calcareous unit outcrops of the monterey formation in the santa barbara coastal plain . benthic foraminiferal assemblages point to relizian to saucesian stages , and calcareous nannofossil zones cn1 - cn5 , with ages generally tending to be younger in the northwestern localities , and older heading to the southeast . for santa barbara point , a locality less than 1 km due east of original santa barbara lighthouse , minor et al . [ 55 ] reported benthic foraminiferal assemblages consistent with relizian and luisian stages of kleinpell [ 56 – 57 ] and calcareous nannofossils of lower middle miocene zone cn4 . these data constrain the age of the type specimen of albicetus to the early middle miocene ( ~ 16–14 ma ) , or langhian .
diagnosis : albicetus is a large odontocete ( about 6 m in total length ) that belongs in physeteroidea based on the following features : large , single - rooted upper and lower teeth , with a ratio of condylobasal length to greatest tooth diameter greater than 0 . 03 ; anterior elongation of the premaxillae onto the rostrum ; sinistral asymmetry of the posterior processes of the premaxillae ; and a posterior section of the rostrum that is wide and relatively flat . albicetus differs from all other known physeteroid genera in the following combination of character states : retention of enameled lower tooth apices with longitudinal striations ; large lower and upper teeth with a ratio of condylobasal length to greatest tooth diameter of approximately 0 . 05 ; a mesorostral groove completely roofed over at the level of the antorbital notches ; and the premaxillae angled downward into the midline , creating a trough down the middle of the rostrum .
the type specimen ( usnm 10923 ) consists of an incomplete skull that includes a partial rostrum and incomplete mandibles , with a number of isolated tooth fragments also found in the surrounding matrix ( figs 2 and 5 – 10 ) . much of the area between the ventral surface of the rostrum and the mandibles is filled with very dense , heavy , and nearly aphanitic grey sediments . these sediments severely hinder any ability to further prepare matrix from bone , which is poorly differentiated from the surrounding matrix . the teeth are also especially soft and chalky , which hinders any further mechanical preparation . the specimen itself is extremely heavy , and the bones of the type specimen are heavily mineralized . the cortical surfaces are also extremely weathered and eroded , likely mostly from exposure as the specimen protruded from the cliff face , which lasted at least a couple decades . these wear patterns are too extreme to differentiate primary weathering from abrasion via transport , or other diagenetic effects [ 58 ] . almost all the bone has been permineralized or replaced by phosphatization in deep - sea sediments , and , in some cases , the original shape of the bone is difficult to distinguish from the surrounding matrix , especially along in posterior view of the rostrum , where it contacts the mandibles .
the left mandible shows the remains of 10 dental roots from teeth that have been fragmented through diagenesis , while the right has 11 ( fig 6a and 6c ) . the type specimen also includes an isolated fragment of the upper rostrum , which was not described by kellogg [ 28 ] ( figs 9 – 11 ) . this triangular fragment is composed of premaxillae , maxillae , and vomer , although it does not have direct , patent contact with the large rostrum .
a , dorsal view of main rostrum section and upper rostral fragment . dotted line indicated hypothesized non - direct connection between the main section and fragment . b , anterior view of upper rostral fragment and c , posterior view of main rostrum section . both have a dotted line indicating the similarity between the transverse profiles of the vomer and mesorostral groove .
though sutures are indistinct , they appear to be closed . this , in combination with the size of the teeth and retention of dental roots in the mandibles , suggest that the specimen belonged to a mature whale .
in dorsal view , the premaxillae dominate the majority of the rostrum , alone forming the anterior ~ 30 % of the rostrum ( fig 5 ) . the premaxilla - maxilla suture is about 27 . 4 cm posterior from the anterior tip of the rostrum , where the premaxillae measure 20 . 7 cm wide . anterior to the premaxillae - maxillae suture , the premaxillae narrow in width to the tip of the rostrum , and maintain approximately the same width from the suture to the posterior end of the specimen . although sutures are indistinct , the right and left premaxillae meet at the midline for the entirety of the preserved rostrum , roofing over the mesorostral groove ( visible in posterior , transverse section ( fig 8 ) . no premaxillary foramen is visible on either premaxilla . in lateral view , the premaxillae gradually thin dorsoventrally from the anterior extremity to the posterior end of the rostrum ( fig 6a and 6c ) . in posterior view , the premaxillae shows a slightly arched profile , angling downwards towards the midline to create a trough down the middle of the rostrum ( fig 8 ) . on this portion of the rostrum , there is no patent asymmetry in the dimensions of the right and left premaxillae ( see description of isolated rostral fragment for further discussion of asymmetry ) .
in dorsal view , the maxillae are most narrow where they meet the premaxillae , 27 . 4 cm from the anterior tip ( fig 5 ) . from this level , the maxillae gradually widen until the posterior of the rostrum , where they rapidly expand laterally to a width of at least 14 cm from its medial margin ( as measured from the right maxilla ) along the maxillary flange ( sensu mead and fordyce [ 42 : 62 ] ) . based on comparison to other fossil physeteroids , this is just anterior to the antorbital notch . in lateral view , the maxillae increase in dorsoventral depth from the premaxilla - maxilla suture towards the base of the rostrum ( fig 6a and 6c ) . in posterior view , the maxillae underlie the premaxillae , meeting the latter dorsally , though this relationship may be due to increased erosion further from the midline ( fig 8 ) . from the lateral margin in posterior view , the maxillae curve dorsally towards the midline to meet the premaxillae .
the mesorostral groove is not visible in dorsal view , being completely sealed over by the premaxillae ( fig 5 ) . in posterior view , the vomer forms the floor of this basin , along with the lateral walls of the mesorostral groove ( fig 8 ) . though it is unclear whether the mesorostral groove was ossified ; in posterior view , it is now filled with sediment , and was dorsoventrally low in life , as in living physeter . see description of the isolated rostral fragment for more on the vomer and mesorostral groove further posterior to the antorbital notch .
the mandibles are both incomplete , but they preserve the entire mandibular symphysis , along with mandibular rami 15 cm posterior of the symphysis . although the extremities of both mandibles end abruptly , the anterior orientation of the anterior - most alveoli ( likely incisors ) suggests that the type specimen preserves most of the anterior portion of the mandibles in life ( fig 7 ) . the orientations of the right and left mandibles are not preserved in life position , especially relative to the anatomical planes of the overlying rostrum ( fig 5 ) . from 57 . 8 cm posterior of their anterior tips , the right and left mandibles are disarticulated , exposing the symphyseal plate along the medial surfaces of the right mandible , whereas the counterpart surface on the left mandible remains covered in sediment ( fig 7 ) . both mandibles are also displaced anteriorly relative to the overlying rostrum , so that they extend past the tip of the rostrum ( fig 5 ) . in dorsal view , the mandibles have also been rotated outwards , so that the alveoli face dorsolaterally and the medial surfaces of the mandibles face the ventral surface of the rostrum . the dorsal curvature of the preserved anterior margins of the mandibles , along with the anterior orientation of terminal alveoli , suggest that , in life , the mandibles had a lateral profile that gradually curved dorsally along the symphysis to the anterior tip , however there is the possibility that this morphology is diagenetic from sediment compaction and deformation ( fig 6a and 6c ) . there is no evidence of the presence of mental foramina on either mandible .
the right mandible retains evidence of 11 alveoli , while the left retains evidence of 10 ( fig 6 ) . both mandibles are heavily eroded on the ventral side . the rami of the mandibles , posterior of the symphysis , remain laterally flattened at the level of the base of the rostrum . although the bone is very difficult to separate from matrix at visual inspection , we suspect that the dorsal rise of the mandibles at this level ( fig 2e , fig 6b ) possibly represents the beginning of the acoustic pan bone region . given all of the aforementioned observations , we propose that the majority of the mandibles are indeed preserved with the rostrum .
in ventral view , numerous black , branching , and likely phosphatized infilled canals about 2–3 cm wide , extend along the ventral surface of the mandibles ( fig 6b ) . ostensibly , these features are exposed from erosion of the overlying cortical bone , and likely represent branches of the mandibular canal . in life , these branches would have once housed nerves and the inferior alveolar arteries connecting to alveoli of the lower dentition , although to date there are only published descriptions of cranial and postcranial sperm whale arterial systems to evaluate this notion , with no particular detail on physeteroid mandibles [ 59 ] . towards the posterior end of the mandibles , there is a patent posteroventral boss on the right mandible , implying the termination of the symphysis in life .
included with the type specimen is a large cranial fragment that was collected in the surrounding matrix , representing a posterior portion of the rostrum separate from the skull and mandible block ( figs 9 and 10 ) . this fragment measures 39 . 5 cm down its anteroposterior length , narrowing from 25 cm wide at its anterior end to 16 . 5 cm at its posterior termination ( fig 9 ) . although it does not have a patent contact with the larger portion of the rostrum , the plane of the anterior breakage is subtransverse , and we suspect it would have articulated within 5–25 cm of a gap with the larger portion ( fig 11 ) . the fragment consists of the premaxillae along the dorsal surface , showing putative premaxillary sac fossae , overlying sediment infilling the mesorostral groove , and finally , along the ventral side , a thin layer of vomer and maxillae along the lateral aspect . the narrowing of the premaxillae and maxillae posteriorly suggests that both are incomplete laterally .
in dorsal view , there are no premaxillary foramina visible ( fig 9 ) . the right and left premaxillae of the fragment are asymmetrical , with the left premaxillary sac fossa showing a more elongate outline in dorsal view ( around 17 cm lengthwise ) and being displaced anteriorly by about 10 cm relative to the right one ( around 14 cm lengthwise ) ( fig 9 ) . this asymmetry is consistent with the sinistral displacement of skulls bones near the narial opening in physeteroids [ 15 , 16 , 22 ] . the dorsal surfaces of the premaxillae are slightly concave and eroded , but towards the posterior end more cortical bone is visible grading through to the underlying maxillae . ventrally , the vomer bridges the right and left premaxillae and maxillae via the mesorostral groove , which is mostly filled with a thick layer of matrix ( fig 10 ) . at this level posterior , the mesorostral groove is open , separating the premaxillae by 9 cm at the anterior end and 6 cm and the posterior end . we can therefore say that the mesorostral groove was roofed over by the premaxillae from the antorbital notches anterior to the tip of the rostrum as seen in the main rostrum fragment ( fig 5 ) , but that the mesorostral groove opens within 5–25 cm posterior of the antorbital notches ( fig 11 ) .
right lateral close up of septal bone material retained in the third premaxillary incisor alveoli ( a ) of albicetus oxymycterus ( usnm 10923 ) , and a illustration of the same area with a low opacity mask and interpretive line art ( b ) . scale bar measures 10 cm in both a and b .
kellogg [ 28 , 60 ] also indicated that 8 alveoli were present in the “distal end” of each maxilla , and he argued that as many as 18 upper teeth would have been present in life . the evidence for upper alveoli in the maxillae is difficult to pinpoint in lateral view , as kellogg [ 28 ] noted , given the diagenetic wear to the specimen . however , a regular scalloped pattern suggesting alveoli can be observed along the lateral margin of the maxillae , with a more prominent pattern on the right side than the left ( fig 12 ) . despite the poor condition of the bony surface of the rostrum , the visible morphology of the lateral margin of the maxillae is largely consistent with much better preserved fossil physeteroids , such as acrophyseter deinodon lambert et al . [ 61 ] , which shows a similar pattern with in situ upper dentition [ 21 ] . at close inspection , clear septa separate the anterior - most alveoli of the right maxilla , especially for c1 ( first canine ) and pc1 and pc2 ( post - canines 1 and 2 ) ( fig 12 ) . at most , we count 6 alveoli for the upper right maxilla , and no more than 5 alveoli for the upper left maxilla , all oriented ventrally and with diastema of at least 1 cm between each ( fig 6a and 6c ) .
the lower alveoli in the mandibles of albicetus are very large , occupying more than half the lateral width of the mandible . the transverse sections of the alveoli along the mandibular border are more ovate towards the anterior end of the mandibles , and more circular posteriorly , likely indicating that the anterior - most teeth were more obliquely oriented than the posterior teeth ( fig 6a and 6c ) . most of the alveoli are filled with matrix , but several retain tooth roots . these alveoli have matrix filled in around the roots around 1–2 cm in thickness , suggesting that , in life , the gum tissue would have actively kept the teeth rooted in the alveoli . each alveolus is distinct from one another , with spacing ranging from virtually none to 3 . 7 cm .


macasinia minifurcata is a species of moth of the tortricidae family .

it is found in carchi province , ecuador .

the wingspan is 16-17.5 mm .

the ground colour of the forewings is brownish creamy , but darker in the basal half of the wing .

the hindwings are creamy brownish , but darker in the distal half .

macasinia minifurcata

have a fact about macasinia minifurcata ? write it here to share it with the entire community .
have a definition for macasinia minifurcata ? write it here to share it with the entire community .
this is the place for minifurcata definition . you find here minifurcata meaning , synonyms of minifurcata and images for minifurcata copyright 2017 © urltoken
here you will find one or more explanations in english for the word minifurcata . also in the bottom left of the page several parts of wikipedia pages related to the word minifurcata and , of course , minifurcata synonyms and on the right images related to the word minifurcata .
encyclo . co . uk , online since 2007 , is a search engine for english meanings and definitions . the website aims to publish all wordlists , big and small , on the internet , making it much easier to find the word you need . follow us on facebook
sign in to disable all ads . thank you for helping build the largest language community on the internet .
have a better pronunciation ? upload it here to share it with the entire community .
simply select a language and press on the speaker button to listen to the pronunciation of the word . leave a vote for your preferred pronunciation .


homo rhodesiensis refers to an extinct hominin species of the genus homo , first described in 1921 by arthur smith woodward in reference to the kabwe skull fossil recovered at broken hill , or kabwe , in zambia ( once known as northern rhodesia ) .

a number of morphologically-comparable fossil remains came to light in east africa ( bodo , ndutu , eyasi , ileret ) and north africa ( salé , rabat , dar-es-soltane , djbel irhoud , sidi aberrahaman , tighenif ) during the 20th century and were classified as homo rhodesiensis .

smith woodward decided the fossils represented an extinct hominid species as the " ... thick skull , sloped forehead and giant brow ridges made the species distinct from living people " .

the finds were dated between 300,000 and 125,000 years bc .

" however , this [ african ] group of fossils has [ also ] been known by many other now-obscure names " , none ever having been accepted universally , including the taxon homo rhodesiensis .

the saldanha cranium , found in 1954 in south africa was subject to at least three taxonomic revisions from 1955 to 1996 .

consensus is confined to informal taxonomic categories , such as " pre-modern " .

most scientists now regard homo rhodesiensis as to be the local ( african ) representative of homo heidelbergensis and as such the direct ancestor of anatomically modern humans .

homo rhodesiensis

the majority of the scientists now consider homo rhodesiensis as the local / african representative of the group homo heidelbergensis . they were also considered direct ancestor of anatomically modern humans ( homo sapiens ) . archaic homo sapiens and homo sapiens rhodesiensis were the other designations proposed for homo rhodesiensis .
homo rhodesiensis was robust , with very large brow ridges and a broad face .
there is , however , an alternative case that can be made concerning the evidence presented . homo habilis , homo rudolfensis and homo georgicus , are all apes , completely unrelated to homo sapiens . homo ergaster and homo erectus , homo cepranensis and homo antecessor , homo heidelbergensis , homo rhodesiensis , homo neanderthalensis , homo sapiens and now extinct homo floresiensis were all fully human without any evolutionary connection to the other allegedly earlier fossil specimens .
arthur smith woodward assigned kabwe 1 or the broken hill cranium as the type specimen for homo rhodesiensis . most researchers do not consider taxon “rhodesiensis” and rather place it in group of homo heidelbergensis .
archaeology in zambia : skull of ' homo rhodesiensis ' discovered in zambia 1921 and placed . . .
homo heidelbergensis fossils tend to have features that are intermediate between those of homo ergaster and either homo neanderthalensis or homo sapiens .
h . rhodesiensis , estimated to be 300 , 000 - 125 , 000 years old , most current experts believe rhodesian man to be within the group of homo heidelbergensis though other designations such as archaic homo sapiens and homo sapiens rhodesiensis have also been proposed .
archaeology in zambia : skull of ' homo rhodesiensis ' discovered in zambia 1921 and placed . . . | the national archives
homo rhodesiensis was robust , with very large brow ridges and broad face . it is also referred to as ' african neanderthal ' , although it demonstrates intermediate features between homo sapiens and
its cranial capacity , however , has been estimated at 1300cc , close to that of a modern human and considerably greater than that of homo ergaster . it has variously been classed as archaic homo sapiens , homo sapiens c . f . rhodesiensis , but is now generally classed as homo heidelbergensis .
homo ergaster and homo erectus – 1 . 8 ma to 70 thousand years ago ( ka )
homo heidelbergensis began to develop regional differences that eventually gave rise to two species of humans . european populations of homo heidelbergensis evolved into homo neanderthalensis ( the neanderthals ) while a separate population of homo heidelbergensis in africa evolved into our own species , homo sapiens .
the anthropologist rupert murrill analyzed the relation between archanthropus skull of petralona ( chalcidice , greece ) and rhodesian man . many contemporary experts assume that rhodesian belonged to homo heidelbergensis group . some other proposed that they belong to other designations such as homo sapiens arcaicus and homo sapiens rhodesiensis .
both anatomically and chronologically , the herto skulls seem intermediate between earlier and more primitive skulls such as bodo ( homo heidelbergensis , dated around 600 , 000 years ago ) and homo rhodesiensis , and the first completely modern human skulls .
this was in fact a sample of what was commonly referred to as the group homo sapiens ( archaic ) . it was a complete cranium that was very robust , with large brow ridges and a receding forehead , similar to homo neanderthalis , but with a cranium intermediate between advanced homo sapiens and neanderthalis . most current experts believe rhodesiensis to be within the group of homo heidelbergensis though other designations such as archaic homo sapiens and homo sapiens rhodesiensis have also been proposed . its estimated age is between 200 , 000 and 125 , 000 years . the brain size was about 1280cc .
homo rhodesiensis are the archaic species of genus ‘homo’ who belonged to the tribe ‘hominini’ . they date back to 300 , 000 to 125 , 000 bc . in 1921 , this species was described with reference to kabwe cranium fossil that was retrieved from the broken hill or kabwe in northern rhodesia ( now known as zambia ) . an english paleontologist arthur smith woodward is credited for describing homo rhodesiensis species for the first time .
the teeth and lower body skeleton suggest homo naledi may have had a lifestyle and diet similar to other hunter - gatherers , such as late homo heidelbergensis / rhodesiensis and early homo sapiens , that were present elsewhere in africa at the same time . how then did h . naledi survive against the bigger - brained competition ?
one of the african archaic species ( now extinct ) was homo rhodesiensis . read here to know what it is , how it was discovered , how it is classified and where does its specimen currently reside .
homo heidelbergensis is an extinct species of the genus homo which may be the direct ancestor of both homo neanderthalensis in europe and homo sapiens . the fossils have been dated to between 600 , 000 and 400 , 000 years ago . the tools used were very similar to those of the acheulean tools used by homo erectus .
some european fossils have features that indicate they were intermediate between earlier homo heidelbergensis and the later neanderthal people . their classification is therefore debated – are they homo heidelbergensis or are they early homo neanderthalensis ?
see
homo naledi , a new species of the genus homo from the dinaledi chamber , south africa
in volume 4 , e09560 .
excavations at kabwe in 1921 revealed the almost complete skull of homo sapiens rhodesiensis ( “broken hill man” ) , which may be well over 100 , 000 years old . however , by 20 , 000 bce the only surviving type of human throughout the old world was the ancestor of modern humans , homo sapiens… …
homo sapiens , ( latin : “wise man” ) the species to which all modern human beings belong . homo sapiens is one of several species grouped into the genus homo , but it is the only one that is not extinct . see also human evolution . the name homo sapiens was…
homo heidelbergensis means ‘heidelberg man’ . homo , is the latin word for ‘human’ or ‘man’ and heidelbergensis is the latinised word for ‘heidelberg’ , the city in germany where the first homo heidelbergensis fossil was discovered in 1907 .
comparison of neanderthal and modern human dna suggests that the two lineages diverged from a common ancestor , most likely homo heidelbergensis , sometime between 350 , 000 and 400 , 000 years ago – with the european branch leading to h . neanderthalensis and the african branch ( sometimes called homo rhodesiensis ) to h . sapiens .
most fossils now known as homo heidelbergensis were previously known as either homo erectus , homo neanderthalensis or ‘archaic’ homo sapiens . with the discovery of many more fossils over the last few decades , many researchers now accept homo heidelbergensis as a separate species , although the designation of some fossils is still debated since they possess features that are transitional between earlier and later species .
before the naming of this species , scientists referred to early human fossils showing traits similar to both homo erectus and modern humans as ‘archaic’ homo sapiens .
homo antecessor ( known only from spain ) has recently been touted as ancestral to homo heidelbergensis , suggesting that either the latter was a european species that later migrated back into africa , or that homo antecessor evolved in africa .
it was considered to be the first species of the genus homo until may 2010 , when a new species , homo gautengensis was discovered in south africa , that most likely arose earlier than homo habilis ( viegas 2010 ) .
earliest homo : temporal bone from chemeron fm , kenya ; 2 . 4 ma
homo erectus : paratype ; first h . erectus found from africa but not oldest
homo heidelbergensis : paratype ; more rounded cranium like h . sapiens ; from europe
olduvai gorge volume 4 : the skulls , endocasts and teeth of homo habilis .
olduvai gorge volume 4 : the skulls , endocasts and teeth of homo habilis .
a skull was discovered by a labourer in 1921 at broken hill in northern rhodesia ( now kabwe in zambia ) . it was initially named homo rhodesiensis ( ' rhodesian man ' ) . it is now commonly referred to as the broken hill skull or the kabwe cranium .
until about ten years ago , the rather unsatisfactory term “archaic homo sapiens” was used to describe any mid - pleistocene hominin that wasn’t homo erectus , homo sapiens , or a neanderthal . the latter was usually classified as a subspecies of homo sapiens , i . e . h . s . neanderthalensis , but they are now generally regarded as a separate species . consequently “archaic homo sapiens” is itself regarded as a separate species , with the 1907 name homo heidelbergensis having seniority under the rules of taxonomy .
this species name is highly debated with many considering the remains to be homo heidelbergensis . whatever species they come from , these fossils are the oldest homo found in europe .
smithsonian national museum of natural history ( smithsonian nmnh ) . 2007 . homo habilis .
see commentary
the many mysteries of homo naledi
in volume 4 , e10627 .
h . heidelbergensis ( heidelberg man ) lived from about 800 , 000 to about 300 , 000 years ago . also proposed as homo sapiens heidelbergensis or homo sapiens paleohungaricus .
a great deal of speculation on this topic has led nowhere . no one really knows what the true significance of this skull might be , or even how old it is . all that anyone can say for sure is that homo rhodesiensis is a clear example of a caveman of uncertain age and status .
homo rhodesiensis is an extinct hominin species living between 400 , 000 and 125 , 000 years ago . it is known as ' rhodesian man ' . the fossil kabwe skull is the type specimen of this species , found at broken hill in northern rhodesia , now zambia , by tom zwiglaar in 1921 .
homo erectus : paratype ; cranium ; shows consistent h . erectus cranial anatomy ; asian variant
homo sapiens : oldest h . sapiens in s . africa ? ; not quite anatomically modern
homo erectus and h . ergaster were the first of the hominina known to leave africa .
during the 20th century , several morphologically comparable fossil remnants were discovered in the regions of north africa ( djebel irhoud , rabat , sidi abderrahman , dar - es - soltane , tighennif , salé ) and east africa ( ileret , ndutu , bodo , eyasi ) . and they were identified as homo rhodesiensis .
more recently , fossils remains found at gran dolina in spain have cast doubt on this interpretation . although many experts consider these remains to be part of an early and variable homo heidelbergensis population , the discoverers believe the fossils are different enough to be given a new species name homo antecessor . they also claim they are the last common ancestor of neanderthals and homo sapiens . this scenario sees european homo heidelbergensis moved to a side branch from modern humans as they are the descendants of homo antecessor and the ancestor of neanderthals . african homo heidelbergensis would require a name change .
the american paleoanthropologist and professor tim d . white believes that the rhodesian man was supposedly the ancestor of homo sapiens idaltu ( herto man ) which itself was the ancestor of homo sapiens sapiens .
importance for reconstructing the early homo skull knm - er 1470 .
1 : j clin pediatr
see
homo naledi and pleistocene hominin evolution in subequatorial africa
in volume 6 , e24234 .
take a tour through seven million years of human evolution and explore the origin of homo sapiens .
l ' homo erectus et la place de l ' homme de tautavel parmi les hominides fossiles .
this taxon is disputed , and most workers consider it to be a junior synonym of homo heidelbergensis .
earliest homo : mandible from ledi geraru ; 2 . 8 ma ( older than a . garhi )
homo erectus : paratype ; the turkana boy ; partial skeleton ; potentially h . ergaster ; african variant
location in algeria where the remains of a homo erectus were found . relatively large teeth and no chin
arthur smith woodward said that the fossils were representative of an extinct hominid species because the
. . . thick skull , sloped forehead and giant brow ridges made the species distinct from living people
. this african fossil group is known by several other ( now obscure ) names but none of them have been accepted widely including the taxon homo rhodesiensis .
earliest homo : maxilla from hadar fm , ethiopia ; 2 . 3 ma ; potential oldest h . habilis
homo erectus : paratype ; skull ; reference skull of
classic
h . erectus features ; asian variant
homo heidelbergensis : paratype ; cranium ; asian h . erectus and h . sapiens like features ; from africa
mounier , aurélien . , marchal , françois and condemi , silvana . ( 2009 ) . “is homo heidelbergensis
the issue of whether the denisovans are a unique species or a subspecies of homo sapiens ( homo sapiens ssp . ' denisova ) has not been settled , just as it has not been settled for the neanderthals .
it was originally though that this transitional species was to disappear within 10 , 000 years as full - blown homo sapiens emerged in africa , but it is known known that homo sapiens appeared much earlier , by 195 , 000 , so homo sapiens idaltu must have been one of several contemporary variations which existed for a time .
while the skull is much similar to that of kabwe , the nomenclature by smith woodward was dropped . the explorers attributed to h . heidelbergensis . it represented the intermediate features between homo ergaster / erectus and homo sapiens .
from africa , based on the close morphology . however , because homo heidelbergensis had a larger average brain capacity than
four species of homo you ' ve never heard of , part ii
. ( smithsonianmag . com )
homo erectus : paratype ; child ' s cranium ; found in java ; oldest h . erectus in southeast asia
homo heidelbergensis : share cranial features with h . erectus , the neandertals , and h . sapiens ; from europe
arsuaga , juan luis ( 2006 ) . “el ' homo antecessor ' de atapuerca era diestro . ” el mundo
homo erectus : a man for all seasons . baker books . grand rapids , michigan . p . 115 .
kimbel wh , johanson dc , rak y . systematic assessment of a maxilla of homo from hadar , ethiopia .
de ruiter dj , churchill se , hawks j , berger lr . late australopiths and the emergence of homo .
kreger , c . d . 2005 . homo sapiens : introduction . urltoken . retrieved february 9 , 2014 .
the next in line is homo heidelbergensis . h . heidelbergensis , e . g . “heidelberg man” , named after the university of heidelberg , is an extinct species of the genus homo which may be the direct ancestor of both homo neanderthalensis in europe and homo sapiens ( mounier et al . 2009 ) . homo heidelbergensis is thought to have likely descended from the morphologically very similar homo ergaster from africa . but because h . heidelbergensis had a larger brain - case — with a typical cranial volume of 1100 - 1400 cm³ overlapping the 1350 cm³ average of modern humans — and had more advanced tools and behavior , it has been given a separate species classification ( mounier 2009 ) .
etymology : the latin word homo means
man
or
human being .
the latin suffix - ensis was added to rhodesi - ( from the former country of rhodesia in which the type specimen was found ) to produce rhodesiensis . so the name of this hominid means
man , or human being , from rhodesia .
rhodesia itself was named for the diamond magnate cecil rhodes .
h . naledi differs from middle pleistocene ( mp ) and late pleistocene ( lp ) homo ( here we include specimens sometimes attributed to the putative early pleistocene taxon homo antecessor , and mp homo heidelbergensis , homo rhodesiensis , as well as archaic homo sapiens and neandertals ) in exhibiting a smaller cranial capacity . h . naledi has its maximum cranial width in the supramastoid region , rather than in the parietal region . it has a clearly defined canine fossa ( similar to h . antecessor ) , a shallow anterior palate , and a flat and a squared nasoalveolar clivus . h . naledi lacks the bilaterally arched and vertically thickened supraorbital tori found in mp and lp homo . h . naledi also differs in exhibiting a root of the zygomatic process of the temporal that is angled downwards approximately 30° relative to fh , a projecting entoglenoid process , a weak vaginal process , a weak crista petrosa , a prominent eustachian process , a laterally inflated mastoid process , and a small eam . the h . naledi mandible tends to be more gracile than specimens of mp homo . the mandibular canine retains a distinct accessory distal cuspulid not seen in mp and lp homo . molar cuspal proportions for h . naledi do not show the derived reduction of the entoconid and hypoconid that characterizes mp and lp homo . the mandibular m 3 is not reduced in dh1 , thus revealing an increasing molar size gradient that contrasts with reduction of the m 3 in mp and lp homo .
homo erectus : paratype ; cranium ; compare to knm - er - 3883 ; potentially h . ergaster ; african variant
homo erectus : paratype ; cranium ; compare to knm - er - 3733 ; potentially h . ergaster ; african variant
wood , b . & collard , m . ( 1999 ) the changing face of genus homo . evolutionary anthropology .
the early human species homo naledi lived much more recently than many scientists had suspected , according to research released today .
homo erectus , ( latin : “upright man” ) extinct species of the human genus ( homo ) , perhaps an ancestor of modern humans ( homo sapiens ) . h . erectus most likely originated in africa , though eurasia cannot be ruled out . regardless of where it first evolved , the species seems to have dispersed quickly , starting about 1 . 9 million…
it seems likely now that this was either an example of late heidelbergensis or an intermediate stage between heidelbergensis and homo sapiens .
earliest homo : mandible from chiwondo beds , malawi ; 2 . 4 ma ; potential h . habilis or h . rudolfenesis
homo habilis . ( 2008 , august 21 ) . new world encyclopedia . retrieved 18 : 35 , october 20 , 2009
another possibility is that african and european homo heidelbergensis are different species , with the bodo cranium an early example of the former . on this view , the african species would take the name originally assigned to the bodo cranium , homo rhodesiensis . however the sudden increase in brain size to 90 % of the modern average seen in the fossil record at around 600 , 000 years ago , after remaining more or less static at 65 % during the previous 1 . 2 million years , does suggest a punctuated event which in turn suggests a single species .
homo rhodesiensis = = = = = = = image - copyright - info = = = = = = = image is in public domain author - info : j . arthur thomson . image source : urltoken = = = = = = = image - copyright - info = = = = = = = = ☆video is targeted to blind users attribution : article text available under cc - by - sa image source in video
argue , d . , d . donlon , c . groves , and r . wright . 2006 . homo floresiensis : microcephalic , pygmoid , australopithecus , or homo ? j . hum . evol . 51 ( 4 ) : 360–74 . pmid 16919706 .
because of increasing evidence of sophisticated material culture , new fossil material , and refined dating techniques , paleoanthropologists have renewed investigations of modern human origins in africa , in particular the question of when early homo sapiens †† became fully modern ( 1 – 6 ) . whereas cranial and dental features demonstrate that african middle pleistocene homo underwent an anatomical transition from a more primitive form ( i . e . , homo heidelbergensis / homo rhodesiensis ) to a relatively modern form ( h . sapiens ) ( 1 ) , little is known about changes in the timing of somatic development , reproductive scheduling , or lifespan ( 7 ) . analyses of dental development , molar eruption , and age at death based on incremental features of the dentition are the most accurate means of identifying developmental change in the human fossil record ( 7 – 10 ) .
australopithecus afarensis and australopithecus africanus are among the most famous of the extinct hominids . the australopithecines are now thought to be immediate ancestors of the genus homo , the group to which modern humans belong . both australopithecines and homo sapiens are part of the tribe hominini .
this is a controversial species designation . most researchers consider these to be part of an early and variable homo heidelbergensis population . however , its discoverers suggest that it shares more traits with modern humans than does european h . heidelbergensis so consider h . antecessor to be the last common ancestor of neanderthals and homo sapiens . some dental and cranial features suggest h . antecessor is descended from homo ergaster .
the species homo heidelbergensis is central to many discussions about recent human evolution . for some workers , it was the last common ancestor for the subsequent species homo sapiens and homo neanderthalensis ; others regard it as only a european form , giving rise to the neanderthals . following the impact of recent genomic studies indicating hybridization between modern humans and both neanderthals and
denisovans
, the status of these as separate taxa is now under discussion . accordingly , clarifying the status of homo heidelbergensis is fundamental to the debate about modern human origins .
schoetensack , o . , 1908 . der unterkiefer des homo heidelbergensis aus den sanden von mauer bei heidelberg . leipzig : wilhelm engelmann .
homo habilis : paratype ; large brain , big teeth , robust face ; controversial species variation ( potentially lectotype of h . rudolfensis )
homo naledi sp . nov . urn : lsid : zoobank . org : pub : 00d1e81a - 6e08 - 4a01 - bd98 - 79a2ceae2411
see
the age of homo naledi and associated sediments in the rising star cave , south africa
in volume 6 , e24231 .
berger lr , hawks j , dirks p , elliott m , roberts em . homo naledi and pleistocene hominin evolution in subequatorial africa .
voisin j - l . évolution de la morphologie claviculaire au sein du genre homo . conséquences architecturales et fonctionnelles sur la ceinture scapulaire .
smithsonian institution , human origins program . 2007a . homo sapiens . smithsonian national museum of natural history . retrieved february 9 , 2014 .
smithsonian institution , human origins program . 2007b . homo erectus . smithsonian national museum of natural history . retrieved february 9 , 2014 .
homo heidelbergensis is associated with the same acheulian ( mode ii ) technology as that originated by homo ergaster 1 . 65 million years ago ; however later acheulian hand - axes are thinner , more symmetric and more extensively trimmed . some authorities describe this technology as “late acheulian” . it is possible that its appearance is connected with the emergence of homo heidelbergensis and is a product of this species greater cognitive abilities .
and used advanced tools , it has achieved its own species classification . homo heidelbergensis was taller - averaging 7 ft - and more robust than
whether or not homo heidelbergensis is a genuine species or simply a grade of “version 3 . 0 human
containing several species remains controversial .
kabwe ( broken hill ) , zambia : skull and several postcranial bones including a femur and a tibia ( broken hill 1 ) . it was found in an iron and zinc mine in broken hill , northern rhodesia ( now kabwe , zambia ) in 1921 by a swiss miner named tom zwiglaar . dating is uncertain , but probably between 700 , 000 and 400 , 000 years old . it has a cranial capacity of around 1100cc and was originally described as homo rhodesiensis .
this species lived between 300 , 000 and 600 , 000 years ago . the african fossils tend to be older than those from europe . fossils from gran dolina in spain date to 800 , 000 years old , and may be homo heidelbergensis or a different species , homo antecessor .
homo heidelbergensis , or “archaic homo sapiens” , is the name given to the large - brained hominins that appeared in africa 600 , 000 years ago and migrated into europe and possibly asia . it is conventionally regarded as having given rise to modern humans in africa and the neanderthals in europe .
see
geological and taphonomic context for the new hominin species homo naledi from the dinaledi chamber , south africa
in volume 4 , e09561 .
in modern taxonomy , homo sapiens is the only extant ( living ) species of its genus , homo . however , the ongoing study of the origins of homo sapiens often demonstrates that there were other homo species , all of which are now extinct . while some of these other species might have been ancestors of h . sapiens , many were likely
cousins ,
having speciated away from our ancestral line . there is not yet a consensus as to which of these groups should count as separate species and which as subspecies of another species . in some cases , this is due to the paucity of fossils ; in other cases , it is due to the slight differences used to classify species in the homo genus .
the use of primitive tools and the greater cranial capacity , e . g . 680 - 800 cm³ greater than the australopithecines but approximately ½ the size of homo sapiens , homo habilis was placed firmly into the genus homo . the fact that ancient and modern apes , chimpanzees , etc . , have been seen using primitive tools does not seem to bother those who defend the placement of this species in the same general category as man .
this brings us to the relatively new kids on the evolutionary block , homo cepranensis and homo antecessor . both fossil specimens were discovered in 1994 . the h . cepranensis fossil consisted of one skullcap and the best representative fossil of h . antecessor was the mandible of an alleged 10 year old boy .
saptomo , e . , brunsden , b . & prior , f . ( 2005 ) .
the brain of lb1 , homo floresiensis
.
feuerriegel em , green dj , walker cs , schmid p , hawks j , berger lr , churchill se . the upper limb of homo naledi .
this brings us to one of the most famous and well documented of the fossil hominids , the neanderthal ( also spelled neandertal ) is an allegedly extinct member of the homo genus that is known from pleistocene specimens found in europe and parts of western and central asia . i say allegedly because the timeline neanderthals are either classified as a subspecies of humans , homo sapiens neanderthalensis , or as a separate species , homo neanderthalensis ( tattersal and schwartz 1999 ) .
homo rhodesiensis (
rhodesian man
or
broken hill man
) was based on a fairly complete cranium , known as the kabwe skull or broken hill 1 , of a large - brained individual found in a mine at broken hill ( now kabwe ) , in northern rhodesia ( now zambia ) by swiss miner tom zwiglaar in 1921 . the specimen was first described by arthur smith woodward ( woodward 1921 ) , the palaeontologist who is one of the chief suspects in the perpetration of the piltdown man hoax .
because until 2006 the liang bua human fossil remains were not available to the entire paleoanthropological community , the taxonomic position of homo floresiensis was only a matter of opinion in publications . from the beginning , two schools of thought prevailed , and this situation persists today . one purports that the liang bua human series belongs to a local modern human ( homo sapiens sapiens ) with anatomical particularities or pathologies that may be due to insular isolation / endogamy . the second argues in favour of the existence of a new species that , depending on the authors , is either a descendant of local homo erectus , or belongs to a much more basal taxon , closer to archaic homo or to australopithecines . because there are no postcranial remains confidently attributed to homo erectus in the fossil record , and because the homo erectus type specimen is a single and partial calvaria , a cladistic analysis was undertaken using both nonmetric morphological features and metrics of the calvariae of human fossil specimens including lb1 to test if it belongs to this taxon . our results indicate that lb1 is included in the homo erectus clade .
so what has science been able to ascertain in the years since the leakey’s discovery concerning homo habilis ? some scientists have proposed moving this species out of homo and into australopithecus due to the morphology of its skeleton being more adapted to living on trees rather than to moving on two legs like homo sapiens ( wood & collard 1999 ) . “a picture is worth a thousand words” is a wise saying attributed to the likes of confucius and napoleon bonaparte , neither of whom can be proven to have ever uttered those words . here is a likeness of homo habilis . see if you see any resemblance to modern man .
homo neanderthalensis : paratype ; largest brain capacity ( 1740 cc ) ; more similar to shanidar and h . sapiens from israel than european neandertals ; from israel
even if woodward ' s now tainted reputation is left out of consideration , the validity of h . rhodesiensis as a distinct type of hominid is not well accepted . it has been variously suggested that the skull on which it is based should be assigned to one or the other of h . erectus , h . neanderthalensis , h . sapiens , or h . heidelbergensis .
rightmire , g . p . , 1998 . human evolution in the middle pleistocene : the role of homo heidelbergensis . evolutionary anthropology 6 , 218 - 227 .
the cranial capacity is typically between 1100 - 1300cc , around 90 % of that of modern humans , a considerable increase on that of homo erectus / ergaster .
fossil bones of homo naledi discovered in the first cave and announced in 2015 . credit berger et al ( 2015 ) elife under cc by 4 . 0 .
homo erectus is considered to have arrived around 1 . 8 million years ago , with fossils supporting its existence to 143 , 000 years ago . often the early phase , from 1 . 8 to 1 . 25 mya , is considered either to be a separate species , h . ergaster , or it is seen as a subspecies of erectus , homo erectus ergaster . that is , h . ergaster is viewed as possibly ancestral to h . erectus , or sharing a common ancestor with h . erectus , or is simply the african variety of h . erectus ; regardless , it is widely considered to be the direct ancestor of later hominids such as homo heidelbergensis , homo sapiens , neanderthals , denisovans , and the asian homo erectus . homo erectus and h . ergaster were the first of the hominina known to leave africa . for example , h . erectus is known to have spread as far as georgia , india , sri lanka , china and java .
this brings us to homo ergaster and homo erectus . h . ergaster is now considered a sub - species of h . erectus or a separate species altogether . for this reason we will examine h . erectus as our next candidate in the human evolutionary ladder . homo erectus is latin for “upright man” and researchers have identified fossils that come from east africa , ethiopia , indonesia , china , the caucasus , and elsewhere in eurasia as remains of homo erectus . depending on the definition of species , a term that from darwin to today has eluded a definitive explanation , h erectus is considered to be either a direct ancestor of modern humans , or a separate species which co - existed with the distinct homo neanderthalensis and a third branch which is thought to be a direct ancestor of modern humans ( heng 2009 ) .
homo heidelbergensis , extinct species of archaic human ( genus homo ) known from fossils dating from 600 , 000 to 300 , 000 years ago in africa , europe , and possibly asia . the name first appeared in print in 1908 to accommodate an ancient human jaw discovered in 1907 near the town of mauer , 16 km ( 10…
when brain size is used as an indicator of intelligence , we can easily find ourselves underestimating or overestimating the intelligence of the hominids . there is a now extinct species of homo sapiens known as homo floresiensis , latin for flores man . h . floresiensis was nicknamed the hobbit due to the short stature of these men and women .
wood , b ( 1999 ) . “‘homo rudolfensis ' alexeev , 1986 - fact or phantom ? ” . j . hum . evol . 36 ( 1 ) :
' despite the young date for the homo naledi fossils , and the more modern anatomical features , ' says prof stringer , ' their anatomy indicates that in evolutionary terms they could lie close to the origin of the genus homo , suggesting that this is a relic species , retaining many primitive traits from a much earlier time . '
the conclusion of the discoverers was that the herto skulls ' sample a population that is on the verge of anatomical modernity but [ is ] not yet fully modern ' . they therefore assigned it to a new subspecies called idaltu , ' because the herto hominids are morphologically just beyond the range of variation seen in amhs [ anatomically modern homo sapiens ] , and because they differ from all other known fossil hominids , we recognise them here as homo sapiens idaltu , a new palaeosubspecies of homo sapiens ' .
modern humans , or homo sapiens , are now the only living species in their genus . but as recently as 100 , 000 years ago , there were several other species that belonged to the genus homo . together with modern humans , these extinct human species , our immediate ancestors and their close relatives , are collectively referred to as ‘hominins’ .
over the course of human history , archaic human admixture has occurred with modern humans ( homo sapiens ) , namely through interbreeding of modern humans with neanderthals and denisovans respectively .
scientific american ( sa ) . 2005 . fossil reanalysis pushes back origin of homo sapiens .
scientific american
february 17 , 2005 . retrieved december 19 , 2013 .
wood , b . , and m . collard . 1999 . the changing face of genus homo . evol . anth . 8 ( 6 ) : 195 - 207 .
most recently numerous homo erectus individuals have been found in europe at the site of dmanisi , georgia ( 1 . 75 mya ) and ceprano , italy ( 800 , 000 years ) . the dmanisi material , if properly dated makes them some of the oldest homo erectus remains known . would link up to mandible found at mauer looks like typical homo erectus earliest homo erectus from africa considered to be 1 . 8 - - how do you get from middle parts of africa to europe and asia ? hominin remains come from pipes and gullies , which creates problems from dating , layers missing due to erosion etc . paleomagnetic sequence points to it being more like a million years old ( missing layers )
the earliest delineated member of the genus homo has generally been considered to be h . habilis , which lived from 2 . 33 to 1 . 44 million years ago , although some authorities do not consider it should be included in homo , considering it more worthy , for example , to be retained in australopithecus ( wood and richmond 2000 ) . there also are proposals for a new discovery , homo gautengensis , to be considered the world ' s earliest recognized species of human , predating h . habilis ( viegas 2010 ) .
the word homo is latin for
person ,
chosen originally by carolus linnaeus in his classification system . it is often translated as
man ,
although this can lead to confusion , given that the english word
man
can be generic like homo , but can also specifically refer to males . latin for
man
in the gender - specific sense is vir , cognate with
vir ile
and
wer ewolf .
the word
human
is from humanus , the adjectival form of homo .
h . erectus lived from about 1 . 8 mya to 70 , 000 years ago . homo erectus was apparently very successful , with fossils found in africa , asia ( indonesia , and china ) , georgia ( caucasus region of europe ) , and eastern and southern africa ( mayr 2001 ) . a famous example of homo erectus is peking man .
kabwe 1 ( 1921 ) : at the broken hill ( now kabwe ) iron and zinc mine in zambia , swiss miner tom zwiglaar came across several fossils , including a skull , jaw and leg bones . the specimens looked human , but the skull also had features that didn’t resemble any modern people , such as heart - shaped browridges and a sloping forehead . the bones were sent to british paleontologist arthur smith woodward . he decided the fossils represented an extinct hominid species he called homo rhodesiensis ( zambia was once part of the british colony northern rhodesia ) . today , the kabwe 1 skull , dating to 300 , 000 to 125 , 000 years ago , is classified in the species homo heidelbergensis , which some paleoanthropologists think was the common ancestor of neanderthals and modern humans .
this is a difficult fossil to classify , given its mixture of traits . the skull is classified by some scientists as late homo erectus and by others as homo neanderthalensis . the brain size is 1220 cc . - - large for h . erectus , but small for h . sapiens - - and the face is large , with a particularly wide upper mandible .
the discovers offer this scenario for the evolutionary and temporal relationships of h . antecessor to other species . h . ergaster gave rise to h . antecessor in africa . about one million years ago , h . antecessor spread via the middle east to europe , including gran dolina . in europe , h . antecessor evolved into h . heidelbergensis , who were the ancestors of the neandertals . in africa , h . antecessor evolved into homo sapiens via an unknown species , although possibly represented by such fossils as the bodo and kabwe skulls . in this scenario h . heidelbergensis is off the line leading to modern humans as it is the descendant of h . antecessor in europe . african h . heidelbergensis would require a name change , probably to homo rhodesiensis , the name originally given to the kabwe skull .
homo naledi is a previously - unknown species of extinct hominin discovered within the dinaledi chamber of the rising star cave system , cradle of humankind , south africa . this species is characterized by body mass and stature similar to small - bodied human populations but a small endocranial volume similar to australopiths . cranial morphology of h . naledi is unique , but most similar to early homo species including homo erectus , homo habilis or homo rudolfensis . while primitive , the dentition is generally small and simple in occlusal morphology . h . naledi has humanlike manipulatory adaptations of the hand and wrist . it also exhibits a humanlike foot and lower limb . these humanlike aspects are contrasted in the postcrania with a more primitive or australopith - like trunk , shoulder , pelvis and proximal femur . representing at least 15 individuals with most skeletal elements repeated multiple times , this is the largest assemblage of a single species of hominins yet discovered in africa .
although we do have another example of a small - brained species surviving until relatively recently - the diminutive homo floresiensis in indonesia - that probably survived as it was isolated on an island .
now berger et al . report the recent discovery of an extinct species from the genus homo that was unearthed from deep underground in what has been named the dinaledi chamber , in the rising star cave system in south africa . the species was named homo naledi ; ‘naledi’ means ‘star’ in sotho ( also called sesotho ) , which is one of the languages spoken in south africa .
h . naledi exhibits anatomical features shared with australopithecus , other features shared with homo , with several features not otherwise known in any hominin species . this anatomical mosaic is reflected in different regions of the skeleton . the morphology of the cranium , mandible , and dentition is mostly consistent with the genus homo , but the brain size of h . naledi is within the range of australopithecus . the lower limb is largely homo - like , and the foot and ankle are particularly human in their configuration , but the pelvis appears to be flared markedly like that of au . afarensis . the wrists , fingertips , and proportions of the fingers are shared mainly with homo , but the proximal and intermediate manual phalanges are markedly curved , even to a greater degree than in any australopithecus . the shoulders are configured largely like those of australopiths . the vertebrae are most similar to pleistocene members of the genus homo , whereas the ribcage is wide distally like au . afarensis .
leakey mg , spoor f , dean mc , feibel cs , antón sc , kiarie c , leakey ln . new fossils from koobi fora in northern kenya confirm taxonomic diversity in early homo .
fossil evidence regarding body size and shape is currently limited but leg bones indicate they were tall , reaching about 180 centimetres in height and had relatively long legs like their earlier ancestor , homo ergaster .
falk d , hildebolt c , smith k , morwood mj , sutikna t , brown p , jatmiko , saptomo ew , brunsden b , prior f . the brain of lb1 , homo floresiensis .
early humans arose in africa between two and three million years ago , but tracing the history and evolution of the various homo species that have existed since then is tricky as the fossil record is patchy . although homo sapiens are the only surviving members of the group , there was no simple progression towards modern humans , and many different human types and body plans co - existed at different times .
as manzi ( 2004 ) notes however this is no more than a revision of the old paradigm , with the substitution of a grade “archaic homo sapiens” with a clade , homo heidelbergensis , accompanied by the recognition of three distinct species , i . e . h . heidelbergensis , h . neanderthalensis and h . sapiens , with corresponding speciation events between the middle and late pleistocene in africa and eurasia .
the kabwe cranium , found in 1921 at broken hill , northern rhodesia ( now kabwe , zambia ) , and originally called rhodesian man . the skull is now considered to be representative of homo heidelbergensis .
modern human beings , neanderthals , and denisovans are believed to have shared a common ancestor about 400 , 000 to 500 , 000 years ago ( marshall 2013 ; green et al . 2010 ) . one theory is that these three groups all descended from homo heidelbergenesis , which lived between 600 , 000 to 250 , 000 years ago ( marshall 2013 ) ( other species suggested as ancestral are h . rhodesiensis and h . antecessor ) . one branch of h . heidelbergenesis are theorized to have left africa about 400 , 000 years ago and split shortly thereafter to become neanderthals , which settled in west asia and europe , and denisovans , which settled farther to the east ( ng 2013 ) .
the remains of at least 6 individuals found at the site of gran dolina , atapuerca , in spain . they lived about 800 , 000 to 1 million years ago in europe and are the oldest human remains found in that continent . although many experts consider these remains to be part of an early and variable homo heidelbergensis population , the discoverers believe the fossils are different enough to be given a new species name homo antecessor .
this skull shows a mixture of features of h . heidelbergensis and homo erectus , to which it is sometimes assigned . it consists of a fairly complete face , with five molars and part of the brain case .
homo heidelbergensis almost certainly used fire . clear evidence of the controlled use of fire , however , is very difficult to establish at archaeological sites , so the origin of this practice among hominids may never be known .
jusqu’en 2006 , plusieurs opinions ont été exprimées à propos du statut taxonomique de homo floresiensis , avant que les paléoanthropologues n’aient directement accès au matériel original de liang bua . de nos jours , les points de vue débattus sont davantage fondés sur des études anatomiques quand , par le passé , seules des opinions s’exprimaient sur les travaux des deux équipes ayant eu accès aux fossiles . il demeure que deux écoles s’affrontent . l’une est partisane de l’existence d’une population d’hommes modernes ( homo sapiens sapiens ) , dont les particularités ou les pathologies sont liées à un isolement et / ou à une endogamie insulaire ; l’autre école défend l’existence d’une nouvelle espèce , qu’elle soit issue d’un homo erectus régional ou qu’il s’agisse de l’ultime représentant d’une lignée archaïque du genre homo , ou même d’un genre différent qui serait proche des australopithèques . dans la mesure où aucun reste postcrânien n’est attribuable de manière certaine à homo erectus et , comme le spécimen - type de cette espèce est une calvaria incomplète , une analyse cladistique a été réalisée à partir des données morphologiques métriques et non métriques de la calvaria de plusieurs spéminens fossiles , dont lb1 , pour tester son appartenance à ce taxon . nos conclusions suggèrent une alternative aux hypothèses précédemment discutées et indiquent que ce spécimen n’est , ni un homme moderne ayant une pathologie , ni une nouvelle espèce , mais est inclus dans le clade homo erectus .
regarding social behaviour , homo heidelbergensis may have been the first species to bury their dead , based on 28 skeletons found at atapuerca , spain . evidence suggests the development and use of a proto - language . the discovery of red ochre at terra amata indicates personal adornment . dental analysis suggests they may have been right - handed . homo heidelbergensis was a sophisticated hunter , suggested by wooden projectile spears found at schoningen in germany attributed to this species .
the skull is huge and has largest of the brow - ridges indicating that it belongs to some really tough individual . similar to homo neanderthalensis , it also has a broad face , large nose and thick protruding brow ridges . as a result of this , the researchers came up with interpretations such as
african neanderthal
. but the latest research with regards to skull’s hardness shows that it had many intermediate features between modern homo sapiens and neanderthal .
double - sided , teardrop - shaped tools , like this lanceolate hand ax , had sharp edges and were sharp enough to slice through tough animal hides . whether or not these tools were made by homo heidelbergensis is debated .
casts of the jaw and hand of homo naledi . the hand has a very human shape , especially if you look at the wrist and palm . but the curved finger bones and strong thumb set it apart from ours .
homo habilis , e . g . “handy man” , is the earliest of the ancestors to modern man that has been classified in the genus homo . remember that evolutionary science places this genus in the family tree of the great apes including the most recent candidate for the missing link between man and monkey , the elder stateswoman of the australopithecines , ardipithecus ramidus , or as we have come to know her as ardi . this particular discovery is traced back to mary and louis leakey and their son john . from 1961 to 1964 the leakeys unearthed fossils of homo habilis , the oldest known primate with allegedly human characteristics and discovered yet another species in 1967 called kenyapithecus africanus . the leakey’s claimed that homo habilis had walked upright . “until then the idea that two hominids could occupy the same area at the same time had been unacceptable to most scientists , ” mary leakey wrote in disclosing the past ( leakey 1984 ) .