euchelus bicinctus is a species of sea snail , a marine gastropod mollusk in the family chilodontidae .

many specimens of clanculus tonnerrei ( g & h nevill 1874 ) have been misdientified as belonging to this species , following a misidentification of issel ( 1869 ) ( herbert , 1996 ) .

euchelus bicinctus

worms - world register of marine species - euchelus bicinctus auct . non philippi , 1849
species euchelus proximus a . adams , 1855 accepted as euchelus circulatus ( anton , 1849 )
species euchelus bicinctus auct . non philippi , 1849 accepted as clanculus tonnerrei ( g . nevill & h . nevill , 1874 )
species euchelus angulatus pease , 1868 accepted as vaceuchelus foveolatus ( a . adams , 1853 )
species euchelus hachijoensis pilsbry , 1904 accepted as herpetopoma rubrum ( a . adams , 1853 )
species euchelus edentulus ( a . adams , 1853 ) accepted as clanculus edentulus a . adams , 1853
to biodiversity heritage library ( 2 publications ) ( from synonym clanculus gibbonsi sowerby iii , 1912 ) to biodiversity heritage library ( 2 publications ) ( from synonym euchelus bicinctus auct . non philippi , 1849 ) to biodiversity heritage library ( 4 publications ) ( from synonym euchelus erythraeensis sturany , 1903 ) to biodiversity heritage library ( 4 publications ) ( from synonym clanculus gennesi h . fischer , 1901 ) to encyclopedia of life to encyclopedia of life ( from synonym euchelus bicinctus auct . non philippi , 1849 ) to mnhn molluscs type collection ( 2000 - 31160 ) ( from synonym clanculus gennesi h . fischer , 1901 )
species euchelus clathratus ( a . adams , 1853 ) accepted as vaceuchelus clathratus ( a . adams , 1853 )
species euchelus erythraeensis sturany , 1903 accepted as clanculus tonnerrei ( g . nevill & h . nevill , 1874 )
species euchelus ruber ( a . adams , 1851 ) accepted as herpetopoma rubrum ( a . adams , 1853 )
( of euchelus bicinctus auct . non philippi , 1849 ) herbert d . g . ( 1996 ) . observations on clanculus tonnerrei ( g . nevill & h . nevill , 1874 ) ( mollusca gastropoda trochidae ) . tropical zoology 9 : 31 - 45 . [ details ]
between otaki and waikanae . at first sight i took it to be a very small specimen of the banded dotterel ( charadrius bicinctus ) , several specimens of which were lying near , but observing that the colouration of the feet , breast , and head differed very much from that of c . bicinctus , i carefully preserved the skin .
subgenus euchelus ( pareuchelus ) o . boettger , 1907 † accepted as pareuchelus o . boettger , 1907 † ( original rank )
» species euchelus ( herpetopoma ) foveolatus ( a . adams , 1851 ) accepted as vaceuchelus foveolatus ( a . adams , 1853 )
species euchelus lamberti ( souverbie in souverbie & montrouzier , 1875 ) accepted as tallorbis roseola g . nevill & h . nevill , 1869
subgenus euchelus ( nevillia ) h . adams , 1868 accepted as nevillia h . adams , 1868 accepted as alcyna a . adams , 1860
species euchelus gemmula w . h . turton , 1932 accepted as vaceuchelus gemmula ( w . h . turton , 1932 ) ( original combination )
species euchelus natalensis e . a . smith , 1906 accepted as vaceuchelus natalensis ( e . a . smith , 1906 ) ( original combination )
species euchelus scabriusculus a . adams & angas , 1867 accepted as herpetopoma scabriusculum ( a . adams & angas , 1867 ) ( original combination )
subgenus euchelus ( perrinia ) h . adams & a . adams , 1854 accepted as perrinia h . adams & a . adams , 1854 ( original rank )
» species euchelus ( herpetopoma ) pruinosus b . a . marshall , 1979 accepted as herpetopoma pruinosum ( b . a . marshall , 1979 ) ( original combination )
species euchelus seychellarum g . nevill & h . nevill , 1869 accepted as herpetopoma seychellarum ( g . nevill & h . nevill , 1869 ) ( original combination )
( of euchelus erythraeensis sturany , 1903 ) herbert d . g . ( 1996 ) . observations on clanculus tonnerrei ( g . nevill & h . nevill , 1874 ) ( mollusca gastropoda trochidae ) . tropical zoology 9 : 31 - 45 . [ details ]
( of trochus ( euchelus ) philippi , 1847 ) philippi r . a . ( 1847 ) . versuch einer systematischen eintheilung des geschlechtes trochus . zeitschrift für malakozoologie . 4 : 17 - 26 . , available online at urltoken page ( s ) : 20 [ details ]
ngu ? n : wikipedia . c c trang : 36 . ch ng : euchelus , neomphalidae , vaceuchelus , anatomidae , pendromidae , cantrainea , angaria delphinus , herpetopoma , granata , danilia , choristella , addisoniidae , perrinia , lepetella , pseudococculinidae , vetulonia , lepetellidae , leptothyra , bothropoma , neomphaloidea , scissurelloidea , vaceuchelus scrobiculatus , arene cruentata , ataphridae , agathodonta elongata , pyropeltidae , tecticrater , peltospira , crysomallon , euchelus bicinctus , cinysca , mirachelus corbis , danilia tinei , retiskenea diploura , perrinia stellata , granata sulcifera , lepetelloidea , euchelus guttarosea , danilia affinis , mirachelus clinocnemus , herpetopoma barbieri , didianema pauli , lirapex , lirapex costellata , pyropelta ryukyuensis , angaria melanacantha , sutilizonidae , chilodonta suduirauti , bichoristes , herpetopoma naokoae , hybochelus leucogranulatus , agathodonta meteorae , nodopelta , emiliotia immaculatus , vaceuchelus vangoethemi , danilia stratmanni , vaceuchelus ludiviniae , vaceuchelus pagoboorum , trochaclis , angaria nhatrangensis , peltospira smaragdina , vaceuchelus vallesi , angaria turpini , planorbidella , herpetopoma eboreum , herpetopoma bella , perrinia cecileae , vaceuchelus abdii , pyropelta sibuetae , lacunoides vitreus , nodopelta rigneae , pyropelta oluae , angaria carmencita , cantrainea globuloides , danilia discordata , perrinia docili , cataegis , lirapex humata , danilia angulosa , mirachelus urueuauau , pachydermia , haliotoidea , depressizona exorum , homalopoma imperforata , danilia galeata , trachysma , euchelus atratus , turcica , homalopoma variecostata , bathyphytophilidae , herpetopoma larochei alacerrima , homalopoma fluctuata , vetulonia paucivaricosa , symmetromphalus , choristella tenera , depressigyra globulus , choristella vitrea , cantrainea yoyottei , liotipoma wallisensis , homalopoma umbilicata , leptothyra filifer , planorbidella planispira , vetulonia parajeffreysi , choristella nofronii , lepetodriloidea , angaria rugosa , cantrainea peloritana , danilia eucheliformis , angaria n . . .
please note that the content of this book primarily consists of articles available from wikipedia or other free sources online . pages : 52 . chapters : tricolia , tricolia pullus , lepetodrilus , mikadotrochus , euchelus , neomphalidae , vaceuchelus , anatomidae , pendromidae , addisonia excentrica , cantrainea , leptogyra , herpetopoma , pleurotomaria , angaria delphinus , eulithidium , granata , danilia , addisoniidae , perrinia , perotrochus , pseudococculinidae , choristella , vetulonia , lepetella , entemnotrochus , lepetellidae , bayerotrochus , leptothyra , neomphaloidea , kaiparapelta askewi , bothropoma , scissurelloidea , leptogyra costellata , eulithidium tessellatum , bayerotrochus africanus , copulabyssia colombia , tricolia retrolineata , vaceuchelus scrobiculatus , entemnotrochus rumphii , amphiplica plutonica , pseudococculina rimula , tecticrater , ataphridae , pyropeltidae , copulabyssia riosi , crysomallon , arene cruentata , melanodrymiidae , peltospira , cinysca , agathodonta elongata , copulabyssia leptalea , mirachelus corbis , helicopelta , euchelus bicinctus , perrinia stellata , tricolia saxatilis , pilus conicus , lepetodrilus shannonae , danilia tinei , granata sulcifera , tricolia adusta , lepetelloidea , mirachelus clinocnemus , euchelus guttarosea , phasianella australis , herpetopoma barbieri , danilia affinis , entemnotrochus adansonianus , sutilizonidae , pyropelta ryukyuensis , melanodrymia , osteopelta mirabilis , addisonia brophyi , lirapex , herpetopoma bella , tricolia speciosa , gorgoleptis , lirapex costellata , didianema pauli , leptogyra inconspicua , leptogyra inflata , leptogyra eritmeta , leptogyra verrilli , herpetopoma naokoae , hybochelus leucogranulatus , vaceuchelus vangoethemi , lepetodrilus atlanticus , sutilizona , lepetodrilus gordensis , nodopelta , vaceuchelus ludiviniae , vaceuchelus pagoboorum , melanodrymia galeronae , sutilizona tunnicliffae , pyropelta sibuetae , trochaclis , chilodonta suduirauti , pyropelta oluae , peltospira smaragdina , vaceuchelus vallesi , angaria melanacantha , emiliotia immaculatu . . .
( of euchelus erythraeensis sturany , 1903 ) albano p . g . , bakker p . a . j . , janssen r . & eschner a . ( 2017 ) . an illustrated catalogue of rudolf sturany’s type specimens in the naturhistorisches museum wien , austria ( nhmw ) : red sea gastropods . zoosystematics and evolution . 93 ( 1 ) : 45 - 94 . , available online at urltoken [ details ]
fairly common . the distinctness of this form is not beyond doubt , vittata , littorinoides , and strebeli all seem to intergrade somewhat on the mainland , and investigation of specific values in this genus must be left for another occasion . i have , however , used strebeli for a number of chatham shells , evidently not bicinctus , of a uniformly dull colour , and very solid habit ; i have identical shells from dunedin harbour ( the type locality of strebeli ) and other forsterian localities . it may be noted , however , that reeve ' s figure of the type of his littorinoides ( conch . icon . , vol . 3 , pl . 12 , f . 94 ) looks as much like strebeli as it does the form commonly accepted as littorinoides .
this seems to be needed for the various minolioid genera such as talopia gray , 1842 , minolia a . ad , 3860 , talopena iredale , 1918 , spectamen iredale , 1924 , antisolarium , conominolia , zeminailia , and zetela , all of finlay , 1926 , but not ethminolia iredale , 1924 ; of these talopia is the oldest name , and may be taken as the foundation of the family . solariella wood , 1842 , and machaeroplax friele , 1877 , perhaps belong here , but , being northern groups , may be more closely related to other associations . thiele has placed these in his subfamily margaritinae , but iredale has noted ( rec . austr . mus . , vol . 14 , no . 4 , p . 258 , 1925 ) that , as far as austral species are concerned , this should be termed stomatellinae , and to the euchelus - stotnatella series the austral minolioids show little resemblance .
manual of conchology , structural and systematic : with illustrations of the species : tryon , george w . ( george washington ) , 1838 - 1888 : free download , borrow , and streaming : internet archive
tryon , george w . ( george washington ) , 1838 - 1888 ; pilsbry , henry augustus , b . 1862 ; sharp , b
vols . 11 - 17 ( published 1889 - 1898 / 8 ) by george w . tryon , continued by henry a . pilsbry vols . 11 - 17 have imprint : published by the conchological section , academy of natural sciences originally issued quarterly , in 68 pts consists of
first series , marine univalves .
cf . p . [ 4 ] of covers of individual pts includes bibliographical references and index
there are no reviews yet . be the first one to write a review .
herbert d . g . ( 1996 ) . observations on clanculus tonnerrei ( g . nevill & h . nevill , 1874 ) ( mollusca gastropoda trochidae ) . tropical zoology 9 : 31 - 45 . [ details ]
herbert d . g . ( 2012 ) a revision of the chilodontidae ( gastropoda : vetigastropoda : seguenzioidea ) of southern africa and the south - western indian ocean . african invertebrates , 53 ( 2 ) : 381–502 . [ 6 november 2012 ] , available online at urltoken [ details ]
worms - world register of marine species - clanculus tonnerrei ( g . nevill & h . nevill , 1874 )
clanculus ( clanculopsis ) tonnerrei ( g . nevill & h . nevill , 1874 ) · accepted , alternate representation
( of clanculus gennesi h . fischer , 1901 ) fischer h . ( 1901 ) . liste des coquilles recueillies par m . de gennes à djibouti et ali - sabieh , avec description de plusieurs formes nouvelles . journal de conchyliologie . 49 : 96 - 130 , pl . 4 . , available online at urltoken page ( s ) : 123 - 125 [ details ]
( of clanculus gennesi h . fischer , 1901 ) herbert d . g . ( 1996 ) . observations on clanculus tonnerrei ( g . nevill & h . nevill , 1874 ) ( mollusca gastropoda trochidae ) . tropical zoology 9 : 31 - 45 . [ details ]
( of clanculus gennesi h . fischer , 1901 ) bosch d . t . , dance s . p . , moolenbeek r . g . & oliver p . g . ( 1995 ) seashells of eastern arabia . dubai : motivate publishing . 296 pp . [ details ]
( of clanculus gibbonsi sowerby iii , 1912 ) herbert d . g . ( 1996 ) . observations on clanculus tonnerrei ( g . nevill & h . nevill , 1874 ) ( mollusca gastropoda trochidae ) . tropical zoology 9 : 31 - 45 . [ details ]
( of clanculus gibbonsi sowerby iii , 1912 ) petit , r . e . ( 2009 ) . george brettingham sowerby , i , ii & iii : their conchological publications and molluscan taxa . zootaxa . 2189 : 1–218 . , available online at urltoken [ details ] available for editors [ request ]
( of clanculus ( clanculopsis ) tonnerrei ( g . nevill & h . nevill , 1874 ) ) herbert d . g . ( 1996 ) . observations on clanculus tonnerrei ( g . nevill & h . nevill , 1874 ) ( mollusca gastropoda trochidae ) . tropical zoology 9 : 31 - 45 . [ details ]
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transactions and proceedings of the new zealand institute , 1879 . [ electronic resource ]
, ph . d . , f . r . s . , director of the canterbury museum .
[ read before the philosophical institute of canterbury , 26 th november , 1879 . ]
, i offered a description of the skeleton of this interesting southern ziphioid whale . i then stated on the authority of the late mr . f . fuller , taxidermist of the canterbury museum , who went to secure the skeleton of that specimen , stranded in lyttelton harbour , some details about the characteristic form and colour of the skin of the animal in question . when my informant arrived where the fishermen were at work , he found that the blubber had nearly all been taken off , so that he could only partially obtain the required measurements . from the observations i am about to offer to the society , on two more specimens stranded since then on our sea - beach , it will be seen that some of the statements were far from being correct .
in fact , the animal was so much cut about that its lower part was taken for the upper , and vice versâ and consequently no dorsal fin could be found where it was looked for . the first of the specimens under review was stranded on sunday , the 17th of november , 1878 , near new brighton . there were numerous visitors at the time who observed another whale ( according to other lookers - on , two whales ) in the offing , by which the animal was driven into the surf , where soon it became helpless . gradually it was drifted upon the low sandy beach , where it died only after a long struggle . having received prompt information , i arrived early next morning on the scene , and found the animal quite intact , so that i could not only take the necessary measurements , but also have a careful sketch prepared , which , as the sequel will show , is of importance , in offering us some curious information as to the habits of this species of ziphioids .
colour : head , neck , and anterior portion of the back , as far as the dorsal fin , white ; the rest of the body black ; a white narrow line running along the edge of the dorsal fin , which is otherwise black . the line of division between the two colours is everywhere well marked , except upon
the cheeks , where blackish blotches advance some distance towards the nose . the cylindrical form of the animal for its length is rather slender , its height at the occiput being only 2 feet 3 inches , and at about nine feet from the tip of the lower jaw 3 feet 3 inches , after which it tapers gradually to the tail . the animal proved to be a young female .
the two teeth at the termination of the lower jaw stood half an inch above the gums , having a diameter of one inch where they rose above the latter . they are conical , and have a sharp apex , and are not covered anywhere with enamel , not even on the tip . the dentine shows a number of horizontal lines one above the other , running round the tooth . they are therefore quite different from the teeth of the two specimens described in vol . ix . ( transactions of the new zealand institute ) , which were found to be covered with a rough cement . they are also different from those of another specimen , of which i shall give some details further on .
a single fold begins below the throat , 1 foot 1 inch from the top of the lower jaw . after rising rapidly for four inches , it continues for seven inches more at a smaller angle , ceasing where the black colour of the throat begins . this fold is separated into two portions by a ridge of the breadth of half an inch below the centre of the throat .
lips flesh - coloured ; roof of mouth slaty - black ; no signs of teeth along the jaws ; there is , however , a hardened ridge along both sides of the palate . the extremity of the lower jaw projects about two inches beyond the upper . the head rises steeply above the upper lip to the forehead .
the blow - hole is situated on the vertex of the head just above the eye . both the form and the size of the dorsal fin and of the tail - lobes , show that this species must be a remarkably swift swimmer .
before giving a description of the external appearance of the specimen under review , i wish to allude to another female , 21 feet 6 inches long , of the same species , stranded on may 15th , 1879 , on the sea - beach near kaiapoi , and of which the skeleton was also secured .
this was , doubtless , a full - grown , aged animal , the terminal epiphyses being so well anchylosed to the body of the vertebræ that even the line of junction could be scarcely distinguished , while in the new brighton specimen these discs were still unanchylosed , and detached themselves readily during maceration .
in form of the body , and colouration , this animal resembled in every respect the new brighton specimen . however , the two teeth existing at the tip of the lower jaw could not be felt when passing the fingers over the gums , and were only disclosed when making incisions .
the teeth are the smallest of all those known to me , being 1·98 and 2 inches long , and only ·46 of an inch broad . the left tooth weighs 66 and the right 62 grains . the flattened root is square , and somewhat constricted a quarter of an inch above the base , after which the tooth expands , being broadest about the middle . it then contracts rapidly , running out to a sharp point . this is thus confirmatory evidence that the teeth , with age , are absorbed , and disappear gradually below the gums ; although it is possible that even below the gums they may still be of some use to the animal . it is a peculiar character of the small teeth of the kaiapoi specimen that they should be so very thin , and terminate in a sharp point ; and that the latter should be covered with real enamel , different from any observed upon the dentine in any other teeth of the same species .
is no doubt in my mind that with them the teeth in front of the lower jaw are both permanent and of larger size than those of the females , just in the same manner as they exist in other ziphioid genera . fortunately , however , there is some evidence at hand strengthening such an hypothesis .
dr . hector , in his account of the skull of epiodon chathamiensis , * obtained in the chatham islands , describes the teeth of this species , as follows : —“the lower jaw * * * terminates in two , short , stout , slightly compressed teeth , 2 inches long , and 4 inches in circumference , implanted in shallow sockets . the teeth have slight , irregular striæ , and are worn down into two lateral facets , divided by an acute ridge . the position of the teeth , when the jaws are closed , is 2 inches beyond the upper mandible , and unless they are applied against callosities on the upper lip , it is difficult to conceive how they are worn down to this acute form . weight of teeth 817 and 836 grains . ”
finally , i should like to make a few observations on the nomenclature , and the changes proposed .
in plate viii . , b is the vent , c the pudendum , and d the fold .
[ read before the wellington philosophical society , 11 th october , 1879 . ]
having since made a minute examination , i have not the slightest hesitation in pronouncing it to be a specimen of gould ' s hiaticula ruficapilla ; it appears to be a very common australian species .
mr . gould , in his “handbook to the birds of australia , ” * states : —“the red - capped dotterel is universally dispersed over every part of the sea - shores of australia that i have visited , and everywhere evinces a greater preference for the shingly beach of the ocean , and especially for deep salt - water bays , than for the sides of rivers and inland waters ; it is very numerous in tasmania , on flinders ' island , on the sand - banks at the mouth of the hunter in new south wales , and at port adelaide in south australia ; and gilbert states that it is equally abundant in western australia , where it is likewise so strictly a bird of the coast that he never saw it inland . it is usually met with in pairs , but may be occasionally observed associating in small companies : —
“like the tringœ , this bird resorts to every possible device in order to lure the intruder from its nest ; throwing itself down upon its breast and flapping its wings , as if in the agonies of death , it will so continue until he has approached almost near enough to place his hand upon it , when it moves along for several yards , dragging one of its legs behind , and , if still followed , attempts to fly , and so well imitates the motion of a bird wounded in the wing , that the intruder is easily misled , and the eggs remain undiscovered . ”
“the male has the forehead crossed by a broad band of white , which gradually diminishes to a point at the posterior angle of the eye ; above , a band of black , which also diminishes to a point at the same place ; from the angle of the mouth to the eye , a line of black , which is continued from the posterior angle of the eye down the sides of the neck ; crown of head , nape , and back of neck , rich rusty red ; all the upper surface and wings pale brown , each feather margined with a still lighter tint : primaries , blackish - brown ; the shafts and extreme edge of the inner webs white ; four central tail - feathers dark brown , the remainder white ; all the under surface white ; irides very dark brown ; bill dark reddish brown ; naked part of legs above the tarsi dark greenish grey , tarsi light grey ; feet blackish brown . ”
the example before us is probably an accidental straggler to our shores from australia , it is , however , a very interesting addition to our list of new zealand birds .
it will be remembered that , in 1876 , dr . buller read before this society descriptions of several varieties of the common , wood - pigeon ( carpophaga novœ - zealandiœ ) . i have now the pleasure of bringing under your notice two additional examples of albinism in this species .
no . 1 is a beautiful albino , the whole plumage being pure white , with the exception of the lesser wing - coverts , which are a delicate yellowish - brown colour , but much more decided than in the specimen mentioned by dr . buller . the claws are yellow instead of black , which is the normal colour . this specimen was shot at springhill station , upper whareama , by mr . a . cameron , and by him presented to the museum ; he says it has frequently been seen about the station during the last four years .
no . 2 is a partial albino . the head , neck , back , and fore - part of the breast are light brown , stained in places with coppery - purple ; lesser wing - coverts , coppery - purple ; quills and their coverts , light brown ; quills tipped and margined with white . tail - feathers brown , tipped with white ; under - surface steel grey , changing to brown towards the extremities ; under - parts from breast downwards , white , slightly tinged with brown ; eyes and feet the usual carmine pink ; claws yellowish - pink , tipped with black . this specimen was procured at pahautanui , and presented to the museum by mr . wise , a very old resident in the district .
the next specimen i have to draw your attention to is a curious and interesting variety of the kotuku , or white heron ( ardea syrmatophora ) .
on the right wing , near the “bend , ” is a patch of dark feathers ; thence a band of black and brown passes right over the back and joins a much larger patch of the same colour on the left wing , and then extends obliquely across the breast , becoming fainter as it again approaches the left side . inner webs of primaries , lining of wings and flank - feathers , more or less marked with brown , passing in places into black . a black patch about an inch in length will also be noticed on the outer web of one of the “secondary plume feathers . ”
i have never before heard of a specimen of this species possessing a single coloured feather , and indeed i am informed that “white as a kotuku ” has passed into a proverb amongst the natives . i was therefore surprised , when , on proceeding to examine the six specimens contained in the “type collection , ” in the colonial museum , i found that no less than three of them had the wings , especially the under - surfaces , more or less spotted or dashed with brown and black .
museum collection . it is evidently the result of an accident , and from its appearance i should say that a shot had just passed below the nostril , splitting the bill in the manner shown . the left side of the upper mandible has also been broken off , but this was evidently a subsequent misfortune , as the broken edge is still somewhat sharp ; while the top of the bill and “spike” are smooth and polished . this unfortunate bird was presented to the museum several years ago , by mr . j . d . enys , who shot it at akitea .
[ read before the wellington philosophical society , 27 th september , 1879 . ]
( orange - wattled crow ) have been as yet entirely unknown , and the author , having chanced to find , towards the end of february last , two nests of this species near the ko - i - te - rangi hill , on the hokitika river , forwards the following description of them .
the nests , which were 15 inches externally , were somewhat loosely constructed of twigs and roots , and had well - formed cup - shaped interiors , lined with pine roots and twigs ; they were built in the branches of the coprosma , or “black” scrub , which grows upon the low river - flats of westland , near the mountain ranges . the average height of the scrub in this instance was about 15 feet , while the nests were about 9 feet above the ground , and 200 feet distant from each other ; one contained an egg , the other , two nearly fledged birds . the egg has been presented to the colonial museum . the two young birds were kept for some weeks in a cage for the
purpose of studying their habits ; their wattles were of a light - rose tint , changing into a violet colour towards the base , but after death , when their skins were dried , the wattles assumed a dull orange tint . the parent birds had wattles of the usual rich crimson - lake colour , the base being tinted with violet as in the young birds .
the egg has almost similarly rounded extremities . length 1·7 inch , breadth 1·1 inch ; the under tint of the egg is brownish , mottled with grey and dark brown blotches which are larger and darker at the larger ends .
. translated from the zoology of the voyage of the “novara , ” by professor hutton .
passage imperfectly developed ; transverse processes of the sacral vertebræ triangular , flat ; no paratoids . ( fingers and toes not dilated at the tips ; maxillary teeth ) .
tympanum , sacs , and auditory tubes wanting ; teeth in the upper jaw , and in two faint oblique rows on the palate , behind and between the interior nares . tongue roundish , more or less margined . fingers free . toes half webbed . projection of the navicular bone small .
fitz . verhandl . d . zool - bot . gesellschaft zu wien , jahrg 1861 ; xi ; pag . 218 , taf . vi
body moderately compressed . eyes rather large ; muzzle rather longer than the eye ; exterior nostrils rather nearer to the eye than to the end of the muzzle .
a glandular fold between the posterior corner of the eye and the shoulder ; a second from the eyes , along the sides of the body to the thighs . several warts about the corner of the mouth , and a few smaller ones on the back and on the sides of the rump . two smooth , yellow callosities on the palm of the hand , one on the metacarpus of the thumb , and one near that of the fourth finger . fingers and toes depressed . toes connected by the web for about half their length , the free part bordered by the membrane . projection of the navicular bone faintly elevated . a dark - grey triangular band between the eyes , in front of which is a broadish and lighter oblique stripe ; hind limbs with broad cross bands on a yellowish - brown ground . belly and sides of rump dirty grey - violet , marbled with yellowish - brown ; a
light oblique streak running down from the anterior and posterior corners of the eye , and diverging to the rim of the upper jaw . male without sound - bag .
[ read before the hawke ' s bay philosophical institute , 12 th may , 1879 . ]
in the winter of 1878 , i received a glass jar from hampden , in this provincial district , containing three full - grown living green lizards . they were pretty nearly alike in size ; two of them were spotted with large irregular - shaped light - green spots , or markings , and one was wholly green . they had been found together , a short time before , in a hole , with a fourth , which was accidentally killed ; and , on their capture , were put carefully into a jar , and packed loosely in moss . on my receiving them i found them apparently very well , but unwilling to move or to face the light , seeking to bury themselves more and more in their mossy bed , so i left them alone , believing they were hybernating . meanwhile , i made many enquiries , by letter , as to their “hole , ” its linings , etc . , but gained little reliable information , save that “in it , and with them , was a lot of stuff like blasting powder ; ” this , i have reason to believe , was the fæcal debris . i greatly regretted the loss of the fourth , as i think that would have proved to be a green male .
during the winter i looked at them three or four times , but they always acted in the same manner , as if averse to having their quiet sleep disturbed . on again looking at them early in october , i found them wholly altered ; they were now desirous of coming to the light , restless , and pawing against the glass , and had increased in number , having four little ones ! two being spotted with white , and two entirely green ; their lovely little bodies looking as if cased in silk velvet instead of scales ; this appearance continued for some weeks . i now lost no time in removing them to more suitable
early in november i was sorry to observe that the young ones , although all four had grown rapidly in length , were daily becoming more weak , especially the two entirely green ones ; this , of course , was owing to their not eating . on the 3rd of november one of the young green ones died . at this time , too , the head of one of the adult lizards ( as i believe , the female one ) swelled much , changed to a livid colour , and grew to an unshapely size , with a bloody discharge distilling from its ears . i thought , that something being the matter with its head , the other lizards in their scrambling about over each other ( which they commonly do ) had fixed their sharp claws in its ears , being now tender , and so caused them to bleed , & c . the sick lizard , however , was very patient under it ; and as its disorder increased , the skin of its head became more and more stretched with the swelling , and great and irregular throbbings or undulations were very apparent . ( here i should mention , that the regular pulsation in their throats is always prominently seen ) . and so , as this diseased lizard became
offensive , yet still living ( though not eating ) , dirtying the others with its discharges , anal now as well as aural , i threw it out into the field .
one of the spotted young ones ( which i shall term no . 1 . ) also cast its skin on the 6th december ; like that of the large male it commenced at the snout , but it came away in fragments—perhaps owing to its being both young and tender .
on the 8th december the second young green lizard died , just as the former young one died , from starvation . this one had , in common with the two young spotted ones , plenty of small flies ( now more easily obtained as the summer advanced ) , but it wanted the power to catch any .
about the 12th december the two remaining adult lizards seemed to be getting into a diseased state ; the handsome male , which had so lately shed its outer skin , had something the matter with its ear , from which a bloody discharge was oozing ( resembling in a smaller degree the early diseased state of the adult one that died ) , while the adult female was restless , swelled in the lower abdomen , and discharging a bloody mixture from its anus ; finding this one getting rapidly worse , with its anus greatly swelled and blotchy—starred all round the margin as it were in a curious regular manner—i lost no time in putting it into a bottle of spirits , and , on my going to look at it some ten minutes after , i found , to my astonishment , no less than 26 large living larvæ of that red - brown flesh - fly had been discharged from its anus ! these were each 5 lines long , and it was their posterior ends compacted together and jutting out from the lizard ' s anus which had given it in that part its peculiar appearance . now it flashed across my
mind , —their evident dislike and dread on their first seeing that flesh - fly in their cage ; and that this was also the cause of the death of my first lizard , into which the living larvæ had been deposited through its ears ! causing its head to possess and show those ugly , unnatural throbbings or semi - undulations . i now hastened to the adult male lizard , and caught it , and on gently squeezing its head i saw the posterior end of a larva presenting itself within its ear ; i took a needle and extracted it ; it was much larger than those in the spirits , and gorged with blood . after this the male lizard soon recovered and became lively , though that aural orifice completely closed up , and so remained until the next shedding of its skin , when i was glad to find that it resumed its former appearance . from the time of this discovery i was careful not to give them any more viviparous female flesh - flies , consequently i have had no more similar diseases to notice .
the big male lizard again shed its skin on the 24th january ; this time , however , in fragments , yet done quickly , all being over within two hours . and again this lizard shed its skin on the 15th of march , this time in large pieces ; finding that while it had extricated its hind - legs it could not draw out its tail , i caught it and helped it to do so . it was pleasing to see how quietly it remained in my hand , when it found out what i was doing , and how naturally it moved its long tail in an easy wriggling manner , and with
strong muscular power pulling against me , so that the whole outer skin of the tail came off , as at first , in one unbroken piece . the cast skin is damp , soft , and slightly clammy , on its being shed , but it quickly dries and hardens .
the young lizard ( no . 1 ) next cast off its skin on the 31st december , having assumed the milky appearance already mentioned the day before ; and to my great surprise this same lizard again put on the cloudy milky appearance on the 13th january , and again shed its skin on the following day when its scurf was just a fortnight old ! as before , it began to break away at its snout , but on this occasion , somehow , possibly owing to its fineness , it got rolled up together and backwards behind its eyes , giving the animal with its white wig the drollest appearance imaginable , so that i often laughed outright ! this time it was very slow in casting off its rags , as parts of its skin were still hanging on its sides on the 24th january—just ten days—when i caught it and helped it . this lizard again shed its skin on the 1st march , when it was two days in getting it wholly off : often biting it and tearing at it with its claws . the next time it did so was on the 19th april , having assumed the usual milky appearance two days before ; on this occasion its old scurf first broke through over its back .
the other young lizard ( no . 2 ) again cast off its outer skin on the 5th february , having the day before put on the peculiar milky appearance .
big adult male , 1878 , november 16 ; 1879 , january 24 ; march 15 . * young one , no . 1 , 1878 , december 6 , december 31 ; 1879 , january 14 , march 1 , april 19 . young one , no . 2 , 1878 , december 16 ; 1879 , february 5 .
their manner of taking their prey ( flies ) is peculiar : when the lizard clearly sees the fly , and makes sure it is living , it steals towards it in the most stealthy manner . as the lizard nears the fly , and when within two inches of it , then is the time closely to notice its actions . first it arches its neck to a tolerably sharp angle , and its eyes swell and bulge out , or rather upwards , over their orbits , and the expression of its countenance alters greatly , taking on a fierce look ; next it lifts its little hand - like paws and moves them , only a toe or a finger at a time and often in the air , very slowly and cautiously ( much like a little child does its hands when stealing along on tip - toe ) , and then it nears its head towards its prey , but so very slowly that i have better detected its movement by watching its shadow cast on marked paper by strong sunlight , —reminding me of the almost imperceptible movement of the hour - hand of a clock . at last it has got to
which they also did . the large lizard often puts its tongue out ( “licking its lips” ) when it goes after a fly , especially if a big one ; i am inclined to think that it is hungry then . it is pretty to see the two young lizards going together after the same fly , especially if the fly is crawling above them , within , on the glass roof ; to see them walking slowly , side by side , with measured gait , and step by step , like a pair of hounds in a leash or a couple of miniature fairy - like little creatures , with their heads up , and their little black eyes glistening ; at such times , too , when they at last near the fly , they often trample on each other in their eagerness , but whenever they do so , they always take it very quietly , the one underneath neither struggling nor retaliating .
it has often seemed to me as if it were a natural law , or rule , of these lizards ( a thing understood by them ) , that whenever they trample on , or walk slowly over , each other , or stand , or lie , or even sleep on each other , the under one , or ones , always take it patiently , and rarely ever move at all—not even when the sharp claws of the upper lizard are pressing on the eyes of the one under him : i have often been surprised at this . i have never once seen them fight or fall out , or attempt to bite each other , although confined in so small a compass . they often spend hours lying on each other ' s backs , which is a favourite posture with them , and sometimes sleep , or spend , the whole night thus . i have seen the whole seven thus together in one lump , with , sometimes , the little ones underneath .
they don ' t seem very timid nor easily startled to any great degree with noises , or sights , or sounds . i keep them on the table in my sitting - room , at which i take my meals , etc . , and i have often thrown down a newspaper by their side , or struck the table with a book pretty strongly , yet they never start ; it is the same when the candles are lit . they appear , too , as if they liked to snugly ensconce themselves in their cage under the koromiko branch , or ( the two young ones ) stretched out at full length on the upper side of the twigs .
i believe them to be inoffensive , peaceful , and sociable ; and if , as i have already surmised , the fourth one ( which was killed ) was also a male , then there would have been two couples , at least , hybernating together in one “hole ; ” or that “hole” may have been their usual dwelling - place , seeing there were found in it “ lots of black stuff”—no doubt their dry and hardened fæces , which could not , i think , have been so largely deposited during the short period then passed of their hybernation . an intelligent friend in the country , who is also an observer of nature , has informed me that he has found them , in clearing , “six or seven together , cuddled up under the roots of a flax - bush” ( phormium ) .
and then passing their broad , thin , and large tongue right over their eyes , as if washing them , and always so finishing their drinking . i have also seen them lick the wet koromiko leaves when fresh ; and the young ones , more than once , lick the adult male . their tongue and palate are of a deep purple colour , much like that of some plums , and the tongue , when fully extended ( as in licking ) , has an emarginate appearance , which may , however , be owing to the action of the hyoid muscle .
they seem to like the water , as they often go singly into their water - trough , and remain extended in the water for some time . they can swim very fast , too , but clumsily , as if they were in a great hurry about it ; i have occasionally tried them at swimming in a large vessel of water .
they can run very swiftly , as i have often proved . when they merely walk , their tails are always straight ; but when they make haste their tails are undulated laterally throughout their whole length . here , no doubt , their under - squamæ help them ; this , indeed , i have in a measure ascertained , in my taking the large lizard into my hands and holding it vertically , when , to aid its ascent in crawling , all the squamæ below are used strongly , and one feels them curiously applied against the hand . * this , also , i think , will account for their being able to climb up on the outside of their glass dome , which they can do—in which feat they are no doubt also materially aided by the large transverse scales on their toes , which are a beautiful object , and admirably adapted for climbing purposes . their claws , too , are exceedingly sharp , having a translucent or semi - crystalline appearance , and are set on at almost a right angle to their toes . one can hardly bear to hold them in one ' s hand when they struggle and use their sharp claws .
their tails have also a strong prehensile power , as i have found in their clasping my fingers with them very closely , and so holding on . on one occasion i had to clear the tail of one which was fast , having taken a half - turn over itself in the sharp angle of a twiggy branch of half - withered and flaccid koromiko , which , i suppose , it had pressed down by lying upon it .
they sometimes spring a short distance very nimbly when they wish to get away from any little obstructions ; and they also jump down fearlessly and without hesitation . i have taken them up and allowed them to run over a book , etc . , held horizontally , 2–3 feet above the table , when they would run straight over the edge of the book and drop on the table on all - fours , like a weasel or a cat , and so continue to run as before .
they assume all manner of curious and grotesque positions , some of them being most extraordinary , and some apparently painful , but in reality i suppose are not so . whatever posture they assume they both can and
[ footnote ] * the sensation being just as if every single scale was being forcibly moved forwards in rapid succession by the muscles of the animal .
8 p . m . it had not moved , so also it was at 11 p . m . , when i went to bed , and when i came down the next morning it was still in exactly the same strange position ; i now thought it could not easily get out , so i lifted the glass to help it , but the moment i did so it scuttled away very fast .
they always take a most peculiar attitude to void their fæces , which , however , they do not perform frequently . i always know when they are about to do so ( if on the look - out ) , for with young and old their preparation is pretty much alike . they first lift up their tails in a semi - curvature towards their backs , then they lift their hind - feet from the floor , and so slowly void their one pellet ; which done they gently lower their hind - feet , and then their tail , and move away . on one occasion i saw the adult male lizard , which was quietly at ease among the koromiko twigs , leave its lair and climb up into the water - trough ; at first i thought it was going to drink , or to bathe in the water , but i was agreeably surprised in noting its actions ; having got into the salt - cellar , it placed its feet on both sides , cocked - up its big tail , and voided its pellet into the water ! that over , it leisurely descended to its former resting - place . in their voiding the fæcal pellet the anus of the animal is produced much more than would be supposed . their dung is of a long oval shape 4–5 lines long , and not unlike that of a sheep ; it is black in colour , but always with a white adjunct ( uric acid ) , somewhat resembling that of a fowl , which portion always appears first ; they void rather slowly . sometimes , especially after eating “bluebottle flies , ” the portions of the fly in rather coarse fragments are very plain in the deposit .


pseudosophronia exustellus is a moth of the gelechiidae family .

it was described by zeller in 1847 .

it is found in portugal , spain , france , italy , and on sicily .

pseudosophronia exustellus

pseudosophronia corley , 2001 ; entomologists gaz . 52 : 214 ; ts : ypsolophus exustellus zeller
sophronia cosmella constant , 1885 ; ann . soc . ent . fr . ( 6 ) 4 : 258 , pl . 10 , f . 19 ; tl : corsica
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oligobuninae is an extinct subfamily of the family mustelidae .

the subfamily was described by j. a. baskin in 1998 ; of the genera that he assigned to this clade , eight are recognized today - brachypsalis , megalictis , oligobunis , potamotherium , promartes , zodiolestes , floridictis and parabrachypsalis - representing fifteen separate species .

oligobuninae

oligobuninae is an extinct subfamily of the family mustelidae known from miocene deposits in north america .
citation : valenciano a , baskin ja , abella j , pérez - ramos a , álvarez - sierra má , morales j , et al . ( 2016 ) megalictis , the bone - crushing giant mustelid ( carnivora , mustelidae , oligobuninae ) from the early miocene of north america . plos one 11 ( 4 ) : e0152430 . urltoken
the new specimens of megalictis ferox described here ( f : am 54079 , f : am 25430 and amnh 54076 ) give us a broader understanding of the morphology of m . ferox and lead us to conclude that the holotypes of both m . ferox ( amnh 12880 ) and aelurocyon brevifacies ( cm 1590 ) are conspecific and thus the latter should be subsumed into m . ferox . we argue that there are 3 species ascribed to megalictis : m . ferox , m . frazieri and m . simplicidens . however , the fourth potential congener , “m” . petersoni , might be best ascribed to a different genus . our cladistic analysis suggests that m . ferox is the sister taxon of the clade composed by m . simplicidens — m . frazieri . our phylogenetic hypothesis supports the subfamily oligobuninae as being a stem mustelid .
megalictis ferox matthew , 1907 [ 1 ] is a giant mustelid of the subfamily oligobuninae and belongs to the paraphyletic group of “paleomustelids” [ 2 ] . it lived in the early miocene during the late arikareean ar4 north american land mammal age 22 . 7–18 . 5 mya [ 3 , 4 ] of the central great plains of united states in the states of nebraska , south dakota , and wyoming [ 1 , 5 – 7 ] . the ar4 lithostratigraphic units containing giant oligobunines have been revised . hunt [ 8 ] named the anderson ranch formation for the terminal formation of the arikaree group in nebraska and wyoming formerly referred to as the upper harrison beds of peterson [ 5 , 9 ] and the lower marsland formation of schultz [ 10 ] . the black bear formation replaces the upper rosebud formation of south dakota [ 11 ] .
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full reference : j . a . baskin . 1998 . mustelidae . evolution of tertiary mammals of north america 152 - 173
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copyright : © 2016 valenciano et al . this is an open access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
data availability : all relevant data are within the paper and its supporting information files .
funding : a . v . has received funding from the european union’s seventh framework programme ( fp7 / 2007 - 2013 ) under grant agreement n° 226506 ( synthesys ; se - taf - 3637 ) , the usc school of medicine ( columbia , south carolina , usa ) , the amnh ( collection study grant program 2014 ) and the international travel grant 2015 from the vertebrate paleontology department of flmnh from uf . a . v . is researcher in formation in the csic program jae - pre _ cp2011 ( csic program
junta para la ampliación de estudios
) , co - funded by the european social fund . a . p . r . is a pre - doctoral fpi fellowship ( bes - 2013 - 065469 ) of the project cgl2012 - 37866 . this study was also supported by the spanish ministerio de economía y competitividad ( research project cgl2015 - 68333 , cgl2011 - 28877 and cgl2011 - 28681 ) , the research group bsch - ucm 910607 and university of south carolina school of medicine ( columbia , south carolina , usa ) .
abbreviations : acm , amherst college beneski museum of natural history , massachusetts , usa ; ahr , comparative anatomy research collection , university of south carolina school of medicine , columbia , usa ; amnh , american museum of natural history , division of paleontology and division of mammalogy , new york , usa ; cm , carnegie museum of natural history , pittsburgh , usa ; f : am , collection housed in the frick collection of the division of paleontology , amnh , new york , usa ; lgput , laboratory of geology and palaeontology , university of thessaloniki , greece ; mncn , museo nacional de ciencias naturales madrid , spain ; mncncomp , comparative anatomy research collection of paleobiology department of museo nacional de ciencias naturales madrid , spain ; nrm , naturhistoriska rikmuseet , stockholm , sweden ; pin , palaeontological institute , russian academy of sciences , moscow , russia ; sam - pql , iziko south african museum , cape town south african museum , south africa ; tmm , texas memorial museum at the vertebrate paleontology laboratory , the jackson school of geosciences , university of texas , austin , usa ; uf , vertebrate paleontology collection of the florida museum of natural history ( flmnh ) , university of florida , gainesville , usa ; unsm , university of nebraska state museum , lincoln , nebraska , usa ; usnm , united states national museum of natural history , smithsonian institution , washington , d . c . , usa
megalictis ferox [ 1 ] was described from the black bear formation , stanley county , south dakota , usa . a second giant oligobunine , aelurocyon brevifacies peterson , 1907 [ 5 ] , was described from the niobrara canyon local fauna , anderson ranch formation in sioux county , nebraska , usa . hunt and skolnick [ 7 ] established that the actual publication date for a . brevifacies was one week after matthew described m . ferox in 1907 , not in 1906 as indicated in the journal . after these initial descriptions , riggs [ 6 ] described new cranial and postcranial material of both taxa . hunt and skolnick [ 7 ] synonymized megalictis ferox , aelurocyon brevifacies , and the large oligobunine mustelid paroligobunis simplicidens ( peterson , 1907 ) [ 5 ] .
here , we describe an important unpublished sample of craniomandibular remains of megalictis ferox ( f : am 25430 , f : am 54079 , and amnh 54076 ) , housed at the american museum of natural history ( new york , usa ) . although f : am 25430 and f : am 54079 were found in the late 1930s and have been used to obtain metric , morpho - functional and phylogenetic data ( e . g . , [ 2 , 7 , 12 – 16 ] ) , they have never been fully described . therefore , the main objective of the present paper is to describe these unpublished skulls and mandibles , and provide new data on the taxonomy and systematics of the genus in order to shed new light on the paleobiology of megalictis .
dental nomenclature follows ginsburg [ 17 ] and smith and dodson [ 18 ] . anatomical descriptions are based primarily on scapino [ 19 ] , turnbull [ 20 ] , barone [ 21 , 22 ] , waibl et al . [ 23 ] , evans and de lahunta [ 24 , 25 ] , and hartstone - rose et al . [ 26 ] . the terminology conforms to the standard of the nomina anatomica veterinaria [ 23 ] with the exception of the masseter and temporalis muscle complexes for which we follow hartstone - rose et al . [ 26 ] . the megalictis material ( figs 1 – 4 ) has been compared to all the other material of megalictis and paroligobunis on the basis of published descriptions , figures , measurements and photographs . we have re - measured the dentition of amnh 12880 and 22632 ( cast of cm 1590 ) measured initially by matthew [ 1 ] and peterson [ 5 ] and completed the measures of paroligobunis petersoni loomis , 1932 [ 27 ] using a cast tmm 40966–1 . measurements were made using mitutoyo absolute digital calipers to the nearest 0 . 1 mm ( tables 1 and 2 ) .
a lateral view ; b rostral view ; c dorsal view ; d caudal view ; e ventral view . scale bar equals 5 cm .
a right mandible lateral view ; b occlusal view ; c left mandible lingual view of lower dentition . scale bar equals 5 cm .
a1–4 cranium f : am 54079 , lateral view ( a1 ) , dorsal view ( a2 ) , ventral view ( a3 ) , and caudal view ( a4 ) ; b1–3 right hemimandible f : am 54079 , lateral view ( b1 ) , medial view ( b2 ) , and occlusal view ( b3 ) ; c1–3 left hemimandible f : am 54079 lateral view ( c1 ) , medial view ( c2 ) , and oclussal view ( c3 ) ; d1–3 right hemimandible of amnh 54076 , lateral view ( d1 ) , medial view ( d2 ) , and occlusal view ( d3 ) . scale bar equals 5 cm .
a , and b megalictis ferox holotype amnh 12880 , lateral view ( a ) , ventral view ( b ) ; c , and d megalictis ferox cm 1590 ( genotype of aelurocyon brevifacies ) , lateral view ( c ) , ventral view ( d ) ; megalictis ferox f : am 25430 lateral view ( e ) , ventral view ( f ) ; g , and h megalictis ferox f : am 54079 lateral view ( g ) , ventral view ( h ) . scale bar equals 5 cm . c and d courtesy of the carnegie museum of natural history .
f : am 25430 ( figs 1 and 2 , s1 video ) : relatively complete skull with i1 - 3 , c , p1 - 4 and m1 - 2 , missing only its left zygomatic arch , a broken frontal area plus a portion of the right parietal region missing and filled with plaster , a hole in its right parietal bone , and a complete mandible with i1 - 3 , c , p1 - 4 and m1 - 2 ; f : am 54079 ( fig 3 , s2 video ) : right side of a partial skull without the premaxilla , with worn c , p2 - m1 and partial mandibles with a nearly complete right one with c - m2 and a broken mandibular symphysis and a left one just with the mandibular corpus preserved and a broken p2 , and a complete both p3 and m1 ; amnh 54076 ( fig 3 ) : partial mandibular corpus with m1 - 2 .
in order to better understand the phylogenetic relationships of the oligobunines megalictis ferox ( amnh 12880 , cm 1590 , f : am 25430 and f : am 54079 ) , m . simplicidens ( = paroligobunis simplicidens ) ( cm 1553 and cm 2389 ) , m . frazieri ( = paroligobunis frazieri ) ( uf 23928 ) , “ m . ” petersoni ( = paroligobunis petersoni ) ( acm 2011 ) , and oligobunis crassivultus ( amnh 6903 ) , we have performed a cladistic analysis ( fig 5 ) including 18 taxa ( m . ferox is represented in the analysis as 4 separate operational taxonomic units ( otu ) ) and 73 equally weighted and unordered craniomandibular characters ( s1 – s3 appendices ) . cladistic analysis was performed using in paup * 4 . 0b10 [ 38 ] . the analysis was rooted using c . lupus as the outgroup .
searches were performed using the branch and bound and a bootstrap analysis through 1000 replicates to test the clade support in the analysis . the outgroup was c . lupus . strict consensus tree of 6 trees ( length 194 steps , consistency index ( ci ) = 0 . 41 , retention index ( ri ) = 0 . 65 ) for knowing the relationships between the different specimens of megalictis ferox , megalictis simplicidens , megalictis frazieri , “ megalictis ” petersoni , oligobunis crassivultus , and a sample of extant musteloids and a canid . numbers below nodes are bremer indices , and numbers above nodes are bootstrap support percentages ( only shown when ≥ 50 ) . character / taxa matrix is detailed in the s1 – s3 appendices . silhouette of megalictis ferox based on hunt and skolnick [ 7 ] , silhouette of megalictis simplicidens , megalictis frazieri , “ megalictis ” petersoni and oligobunis crassivultus based on megalictis ferox but rescaled according the size of the dentition .
virtual models of the mandibles and skulls of megalictis ferox ( f : am 25430 , f : am 54079 , amnh 12880 and amnh 22632 ) as well as megalictis frazieri uf 23928 and megalictis simplicidens ( cast of cm 1553 ) were derived by means of a 3d nextengine hd laser surface scanner ( s1 – s6 videos ) .
diagnosis : large to giant size mustelid ; robust mandible with a high , wide and distally curved ascending ramus ; deep masseteric fossa with a stout crest that extends from the dorsal border of the coronoid process to below the m2 ; robust dentition ; p1 present ; p2–4 with distal cingula high - crowned ; p4 relatively enlarged with mesial and distal accessory cuspids ; m1 trigonid widened , with a strong lingual concavity between the paraconid and protoconid ; m1 metaconid reduced to absent , present in the older and smaller forms and absent in the giant forms , m1 talonid low and narrow with a short , trenchant and labially located hypoconid ; and m2 with reduced metaconid .
type species : megalictis ferox matthew , 1907 p1 . ii , fig . 1 [ 1 ]
included species : megalictis simplicidens ( = paroligobunis simplicidens ) ( peterson , 1907 ) [ 5 ] and megalictis frazieri ( = paroligobunis frazieri ) ( frailey , 1978 ) [ 28 ] .
synonyms : senior subjective synonym [ 7 ] of aelurocyon brevifacies peterson , 1907 , p . 68 [ 5 ] , “upper harrison formation” , sioux county , nebraska and paroligobunis peterson , 1910 [ 29 ] . hunt and skolnick [ 7 ] synonymized megalictis , aelurocyon , and paroligobunis simplicidens into a single , sexually - dimorphic chronospecies m . ferox . this hypothesis has been generally accepted ( e . g . , [ 3 , 13 , 42 ] ) .
megalictis ferox , hunt and skolnick , 1996 ( pars ) . [ 7 ]
holotype : amnh 12880 , a partial reconstructed skull ( fig 4 , s3 video ) with right p4 , m1 - 2 , a fragmented right mandible with almost complete coronoid process , m1 trigonid and m2 , and very few postcranial remains figured by matthew , 1907 , p . 196 , fig . 10–13 , 15 [ 1 ] .
type locality : rosebud 22 , porcupine butte , black bear formation , stanley county , south dakota .
other localities : rosebud 5 , porcupine butte , stanley county , south dakota , usa ( amnh 12881 ) ; niobrara canyon local fauna , sioux county , nebraska , usa ( cm 1590 ) , “high daemonelix beds” , niobrara canyon local fauna , sioux county , nebraska , usa ( f : am 25430 ) ; j - m district , south of lusk , goshen county , wyoming , usa [ 6 ] ; “high brown sand” , 16 mile district , goshen county , wyoming , usa ( f : am 54079 ) ; 8 north of lusk , goshen county , wyoming , usa ( f : am 54076 ) .
age : upper part of the anderson ranch formation and its equivalents , south dakota , nebraska , and wyoming , late late arikareean ( ar4 ) , 22 . 7–18 . 5 mya [ 4 ] early miocene .
diagnosis : baskin [ 2 ] diagnosed of aelurocyon brevifacies ( which he considered the senior subjective synonym of megalictis ferox because of the presumed earlier publication date at the time he submitted the chapter ) . new or revised characters follow . megalictis ferox is the largest of the oligobunines ; coronoid process high and caudally curved ; enlarged masseteric fossa with a robust crest extending from the dorsal border of the coronoid process to below the m2 ; laterocaudal area of the ventral edge of the mandibular corpus laterally projected ; p2 with distal accessory cusp ; robust p3 ; robust p4 with strong parastyle and protocone ; p4 carnassial notch present ; m1 with enlarged stylar area ; m2 with paracone and protocone ; p2–4 with high - crowned distal cingula ; p3 with mesial and distal accessory cuspid ; p4 relatively enlarged with presence of mesial accessory cuspid and stout distal accessory cuspid ; m1 trigonid widened ; m1 with strong lingual concavity between paraconid and protoconid ; m1 protoconid higher than paraconid ; m1 hypoconid short , trenchant and buccally located ; m1 with a lingual cingulum in the entoconid position ; m2 reduced with metaconid .
differential diagnosis : megalictis ferox differs from m . simplicidens , m . frazieri , “m . ” petersoni and oligobunis crassivultus in its larger size , m1 without metaconid and m1 talonid with a closed lingual morphology with a lingual cingulum between the metacristid and entocristid . additionally , it differs from m . simplicidens and m . frazieri in having a higher and more robust mandibular symphysis , a reduced p2 and a more robust p4 and m1 . it further differs from “m . ” petersoni in much larger size and p3–4 with mesial accessory cuspids . it further differs from oligobunis crassivultus in having a more rectangular p2 , smaller m1 than p4 , enlarged m1 stylar area , higher paracone than metacone on the m1 , reduced p2 , p2–3 high - crowned distal cingula , more developed p3 distal accessory cuspid , relatively enlarged p4 , and higher protoconid than paraconid on the m1 .
comments : specimens that can be referred to m . ferox s . s . are from the latest arikareean ( ar4 ) upper part of the anderson ranch formation and its equivalents .
a nearly complete skull with i1 - 3 , c , p1 - 4 and m1 - 2 ( fig 1 , s1 video ) and a complete mandible with i1 - 3 , c , p1 - 4 and m1 - 2 ( fig 2 , s1 video ) . the left zygomatic arch is missing . part of the frontal and a region of the right parietal bones are missing and filled with plaster . there is a subrectangular and anthropogenic hole in its right parietal bone located above the most caudal area of the zygomatic arch .
locality : “high daemonelix beds” , anderson ranch formation , niobrara canyon local fauna , sioux county , nebraska , usa .
skull and upper dentition : the very well preserved skull f : am 25430 ( fig 1 ) has a basicranial length of 189 . 5 mm . it is slightly domed dorsally at the frontal bone , the bullae are broken and the left zygomatic arch is missing . in general terms the skull is high , domed with a short rostrum and high snout ( fig 1a ) . the nasal aperture is large ( fig 1b ) , and the crushed nasal bones are robust . they are crushed in the mid - sagittal plane and anteriorly the left nasal bone is partially above the right one . the reconstructed frontal region is quite domed . the interorbital region is broad . the postorbital processes are absent . the moderately developed infraorbital foramen is rounded and located above the distal accessory cusp of the p3 . the rostral margin of the orbit ends at the level of the distal margin of the p4 paracone . the orbits are large and rounded . the lacrimal foramen is rounded and relatively large . the sagittal crest is moderately developed and extends caudally where it divides into the nuchal crests , forming a y - pattern ( fig 1a , 1c and 1d ) . in lateral view , the outline of the skull is convex in the temporal region and concave between the temporal bone and nuchal crests .
the zygomatic arches are robust , especially caudally near the glenoid cavity . both m . masseter pars superficialis and m . masseter pars profunda have their origin on the ventrolateral side of the zygomatic arch . the frontal processes of the zygomatic arches are triangular and dorsoventrally high .
mandible and lower dentition : the mandible of f : am 25430 is very robust ( fig 2a and 2b ) . it has a total length of 149 . 0 mm . the tooth row is slightly convex and is aligned with the articular process . the mandibular corpus is high and robust . the ventral margin is convex at the level of the m1 . there is single rounded mental foramen under p2 . the ascending ramus is tall and rostrocaudally broad ( fig 2a ) . its tip is distally oriented . the coronoid process is laterally rotated with an angle of ~ 75 degrees , compared to the articular process . there is a robust crest from the dorsal border of the coronoid to beneath the m2 where the tendon of the m . temporalis is attached . this area is especially enlarged and laterally projected around the area of the m2 ( fig 2a and 2b ) . the masseteric fossa is large and deep . its rostral margin lies at the level of talonid of m1 , and ventrolaterally is limited by a strong area where the m . masseter pars superficialis and m . masseter pars profunda insert . the articular process is large and robust . the angular process is robust and shows a medial crest for the muscular attachment of the m . pterygoideus medialis .
the lower dentition ( 3 / 1 / 4 / 2 ) is also preserved in its entirety ( fig 2 ) . the lower incisors are heavily worn . the canine is large , stout and markedly curved distally ( fig 2a and 2b ) . it has a swollen base and is oval in cross - section . the p1 is oval , single - cusped and distally wide ( fig 2b ) . the p2–4 are stout , subrectangular and wider distally . these premolars have strong cingula at their bases , and the distal cingula are high - crowned . the p2 has a single messially - located cuspid . the p3 has a low mesial accessory cuspid and a more developed distal one . the p4 is the largest lower premolar and has more strongly developed mesial and distal accessory cuspids . the m1 is a relatively short and stout tooth ( fig 2 ) . the very robust trigonid occupies almost three fourths of the total length of the tooth , with the greatest width at the base of the protoconid . the paraconid is lower than the protoconid and there is no metaconid . the m1 shows a markedly lingual concavity in the base of the crown between the trigonid cuspids ( fig 2b ) . the stout talonid lacks an entoconid . the hypoconid is low , trenchant and buccally located . there is a smooth cristid from the top of the protoconid to the hypoconid that encloses a deep lingual depression ( fig 2c ) . the m2 is rounded and low ( fig 2b ) . the paraconid is low , and located in the mesial corner . the protoconid is the highest cuspid , located buccally in the middle of the tooth . the metaconid is situated over the lingual corner . it is less developed than the protoconid . the hypoconid is low and located in the distal corner . there is a cingulum around the whole tooth .
partial skull with worn c , p2–m1 and partial mandible with worn p1–4 and m1–2 ( fig 3a1–4 , 3b1–3 and 3c1–3 , s2 video ) .
locality : “high brown sand” , 16 mile district , anderson ranch formation , goshen county , wyoming , usa .
skull and upper dentition : the skull f : am 54079 ( fig 3a1–4 ) only preserves its right side . it has a maximum length of 180 . 2 mm . the premaxilla is missing , so the basicranial length is unknown . in general terms , f : am 54079 resembles f : am 25430 ( fig 1 ) . the frontal bone and dorsal area of the parietal bone are absent ( fig 3a2 ) . the zygomatic arches are more robust than those of f : am 25430 , especially in the rostral and the central part of the arches , and the origin of m . masseter pars superficialis and m . masseter pars profunda are also more developed . however , the frontal processes of the zygomatic arches are lower than those of f : am 25430 . the glenoid fossa is stout with a very well developed postglenoid process ( fig 3a3 ) . the complete right auditory bulla is large and swollen . the external auditory meati are rounded . the postglenoid foramen , the rostral foramen lacerum and the foramen ovale are similar to those of f : am 25430 . the mastoid process is also robust and expanded . the right occipital condyle is preserved but the caudodorsal area of the skull is not . the paroccipital process is triangular , stout and caudally oriented ( fig 3a1 and 3a3–4 ) .
c , p2–4 and are preserved . the p1 is missing . they are more worn than are those of f : am 25430 . the c has a large lingual wear facet . the morphology of p2–4 ( fig 3a3 ) is almost identical to that of f : am 25430 . the p3 is more quadrangular than that of f : am 25430 , but the mesiolingual corner of the p3 is missing . the p4 paracone , protocone and metastyle are greatly - worn ( fig 3a3 ) . the m1 ( fig 3a3 ) has the same development of the cusps as that found in f : am 25430 , and shows a very similar morphology as that of amnh 12880 . the m2 and its alveoli are not preserved .
mandible and lower dentition : the right hemimandible ( fig 3b1–3 ) has a fragmented corpus that is missing its symphyseal end but includes a complete ascending ramus with p1–4 and m1–2 . its morphology is almost identical to that of f : am 25430 . the left hemimandible ( fig 3c1–3 ) is missing its ascending ramus but includes a complete mandibular corpus , a complete p1 , a fragmented p2 , a highly worn p3 , a complete m1 and a fragmented m2 . the p1–4 and m1 are almost identical to those of f : am 25430 though there is more substantial occlusal wear in p2–4 and m1 than in f : am 25430 . the m2 is oval and has a more developed metaconid than the m2 of f : am 25430 .
locality : 8 north of lusk , goshen county , anderson ranch formation , wyoming , usa .
mandible and lower dentition : amnh 54076 is a fragmented mandibular corpus missing its symphysis ( fig 3d1–3 ) . it has roots for p2–3 , and complete m1–2 . the mandibular corpus is high and robust . the m1 is identical to those of f : am 54079 and f : am 25430 . it has a stout trigonid , and a low talonid composed of a trenchant hypoconid , lingually located and a lingual depression . the m2 is rounded and low . it has a distinguishable protoconid and metaconid , and a continuous basal cingulum .
hunt and skolnick [ 7 ] did not consider the other two species referred to paroligobunis : the small p . petersoni loomis , 1932 [ 27 ] and p . frazieri frailey , 1978 [ 28 ] . as discussed below , we consider the material referred to both p . simplicidens and p . frazieri to be valid species : megalictis frazieri and m . simplicidens .
the results of the cladistic analysis indicate that the specimens we assign to m . ferox form a monophyletic group ( fig 5 ) . we agree with hunt and skolnick [ 7 ] in that m . ferox and a . brevifacies are the same taxon , and that m . ferox has priority . morphologically , the specimens f : am 54079 , f : am 25430 and amnh 54076 , as well as cm 1590 and amnh 12880 , are practically identical to each other ( figs 1 – 4 ) . f : am 54079 differs from f : am 25430 and cm1590 in having a more robust p3 and a relatively longer m2 . cm 1590 has a reduced lingual expansion of p3 and a stronger parastyle of p4 than f : am 54079 , f : am 25430 and amnh 12880 . the morphology of f : am 25430 is clearly different from the skull of amnh 12880 , and shows that the reconstructed parts of the latter were incorrect , in which the temporal , frontal and a part of the zygomatic arch bones are misinterpreted ( fig 4a and 4b ) . f : am 25430 allows us to complete the knowledge about the morphology of the skull of m . ferox and showing that the holotype of m . ferox ( amnh 12880 ) and the holotype of a . brevifacies ( cm 1590 ) belong to the same species . consequently , f : am 54079 , f : am 25430 and amnh 54076 should be assigned to m . ferox . we agree with hunt and skolnick [ 7 ] that the difference observed in the specimens of m . ferox can be explained by intraspecific variability ( sexual dimorphism and intrapopulational differences ) or small temporal differences .
megalictis ferox ( figs 1 – 4 ) is characterized by several traits : long external auditory meatus ; high and caudally curved coronoid process ; enlarged masseteric fossa with a robust crest from the dorsal border of the coronoid process to just beneath the m2 ; latero - caudal area of the ventral edge of the mandibular corpus is laterally projected , with the ventral edge of the angular process also laterally projected ; i3 is enlarged ; p2 with a distal accessory cusp ; robust p3 ; robust p4 with carnassial notch ; enlarged stylar area of m1 , and a m2 with paracone and protocone differentiated ; p2–4 distal cingula high - crowned ; distal accessory cuspid on p3 ; relatively enlarged p4 with a stout mesial accessory cuspid ; relatively stout m1 with a widened trigonid , a strong lingual concavity between the paraconid and protoconid , no metaconid , protoconid higher than paraconid , with a short , trenchant and buccally located hypoconid and a lingual rim in the entoconid position ; reduced m2 with a metaconid .
all the three species that have been referred to paroligobunis ( fig 6 ) are known from limited material . the genotype of paroligobunis , megalictis simplicidens ( cm 1590 , peterson , 1907 , 1910 ) [ 5 , 29 ] comes from the “agate stock farm” , sioux county nebraska . the exact locality is unknown and it is either from the harrison formation ( ar3 ) or the basal part of the anderson ranch formation [ 7 ] . additional material first referred to p . simplicidens [ 29 ] and later to megalictis ferox [ 7 ] is from quarry 3 , beardog hill , agate fossil beds national monument , from the basal part of the anderson ranch formation . the small “m” . petersoni ( loomis , 1932 ) [ 27 ] is from a locality near van tassel , wyoming , “upper harrison beds” ( = anderson ranch formation ) and p . frazieri frailey , 1978 [ 28 ] is from the sb - 1a local fauna , florida , latest oligocene , early late arikareean ( ar3 ) . hunt ( in tedford et . al , 2004 : p . 205 [ 3 ] ) recognized that “‘ paroligobunis’ frazieri is an earlier form preceding the late arikareean species of megalictis ” .
( a ) megalictis simplicidens , type specimen , cm1553 ( peterson , 1907 ) [ 5 ] , lateral view of the mandible , ( b ) megalictis simplicidens cm 1553 ( peterson , 1907 ) [ 5 ] , medial view , ( c ) megalictis simplicidens cm 2389 ( peterson , 1910 ) [ 29 ] , lateral view of the mandible , ( d ) megalictis simplicidens cm 2389 ( peterson , 1910 ) [ 29 ] , medial view , ( e ) megalictis frazieri ( frailey , 1978 ) [ 28 ] , holotype uf 23928 , lateral view of the mandible , ( f ) megalictis frazieri ( frailey , 1978 ) [ 28 ] , uf 23928 , medial view , ( g ) “ megalictis” petersoni ( loomis , 1932 ) [ 27 ] , holotype acm 2011 , lateral view of the mandible , ( h ) “ megalictis” petersoni ( loomis , 1932 ) [ 27 ] acm 2011 , medial view , ( i ) megalictis simplicidens cm1553 ( peterson , 1907 ) [ 5 ] , occlusal view , ( j ) megalictis simplicidens cm 2389 ( peterson , 1910 ) [ 29 ] , occlusal view , ( k ) megalictis frazieri ( frailey , 1978 ) [ 28 ] , uf 23928 , occlusal view , ( l ) “ megalictis” petersoni ( loomis , 1932 ) [ 27 ] acm 2011 , occlusal view . scale bar equals 5 cm . a - d , i and j courtesy of the carnegie museum of natural history . e - f and k courtesy of the florida museum of natural history . g - h and l , beneski museum of natural history at amherst college , courtesy of the trustees of amherst college .
there are no derived characters uniting the three named species of paroligobunis that are not shared with megalictis ( s2 appendix ) . our phylogenetic analysis ( fig 5 ) shows that these three species are paraphyletic with m . ferox . the larger p . frazieri and p . simplicidens are both referred to megalictis . the differences in morphology and size between the three species of megalictis with respect to “ m . ” petersoni ( fig 6 ) suggest that “ m . ” petersoni could be excluded from the genus megalictis .
megalictis simplicidens and m . frazieri ( fig 6 ) resemble m . ferox in several characters , such as a high , wide and distally curved ascending ramus , and a deep masseteric fossa with a robust crest that extends from the dorsal border of the coronoid process to below the m2 . both taxa have a p1 , the distal cingula of p2–4 are high - crowned , and the p4 is relatively enlarged with mesial and distal accessory cuspids . the m1 trigonid is widened , with a strong lingual concavity between the paraconid and protoconid , a low , and narrow talonid with a short , trenchant and labially located hypoconid , and a reduced m2 with presence of a metaconid . however they differ from m . ferox in having a non - reduced p2 , the presence of a stout m1 metaconid , relatively more slender p4 and m1 , m1 talonid with an open lingual morphology between the metacristid and entocristid , and a lower and more slender mandibular symphysis .
hunt and skolnick [ 7 ] partially described and measured some of the unsm and cm specimens of megalictis from the basal part of the anderson ranch formation at beardog hill that we refer to m . simplicidens . aside from their more primitive morphology ( e . g . , presence of a metaconid on m1 ) , they are smaller than m . ferox from the upper anderson ranch formation . the upper and lower dental measurements indicate a size similar to g . gulo .
megalictis frazieri ( fig 6e , 6f and 6k ) differs from m . simplicidens ( fig 6a–6d , 6i and 6j ) in having a less massive mandible and a more distinctive distal cingulum with a higher crown in p2–4 than m . simplicidens . the c and p2 of m . frazieri are also more robust . the m1 hypoconid is higher and the talonid is relatively larger , slightly basined with a very low internal rim .
“megalictis” petersoni ( fig 6g , 6h and 6l ) differs from m . simplicidens and m . frazieri in the absence of mesial accessory cuspids on p3–4 , a relatively stouter p4 with a shorter mesial part and a relatively more robust m1 with a taller and stouter metaconid .
metrically the new megalictis ferox sample described above ( f : am 54079 , f : am 25430 and amnh 54076 ) together with amnh 12880 and cm 1590 form a single picture of m . ferox with dental biometric variability similar to the largest extant terrestrial mustelids gulo and mellivora ( figs 7 and 8 ) . however , if m . simplicidens is considered as a synonym of m . ferox , this variability exceeds the extant one . such variability is much more pronounced when all the specimens of m . simplicidens , m . frazieri and the small “ m . ” petersoni ( figs 7 and 8 ) are included .
a life appearance ; b , reconstructed skull and mandible ; c , skull and mandible f : am 25430 . artwork by adam hartstone - rose .
the preservation of the of m . ferox specimen f : am 25430 represents by far the most complete and best preserved craniomandibular specimen of any giant mustelids . based on the size of the skull , m . ferox emerges as the largest terrestrial mustelid ever known–even larger than the extinct late miocene giant mustelid ekorus , eomellivora , and plesiogulo [ 13 , 32 , 33 , 35 , 37 , 70 ] . this new material sheds light on a new paleobiological interpretation of megalictis as a hyena - like , bone - crushing mustelid , instead of the cat - like ecomorphotype previously ascribed to the genus .
s1 table . list of the extant specimens of carnivorans used in this paper .
s1 video . video of the cranium and mandible of megalictis ferox f : am 25430 .
s2 video . video of the cranium and mandible of megalictis ferox f : am 54079 .
s3 video . video of the cranium and mandible of the holotype of megalictis ferox amnh - 12880 .
s4 video . video of the cranium and mandible megalictis ferox amnh - 22632 ( cast of cm 1590 ) .
s5 video . video of the mandible of megalictis simplicidens ( cm 1553 ) and megalictis frazieri uf 23928 .
s6 video . video of the reconstructed head of megalictis ferox f : am 25430 .
conceived and designed the experiments : av jm ahr . performed the experiments : av jab jm . analyzed the data : av jab ja maas jm ahr . contributed reagents / materials / analysis tools : av ja apr . wrote the paper : av jab ja maas jm ahr .
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the subfamily was described by j . a . baskin in 1998 ; of the genera that he assigned to this clade , six are recognized today -
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the subfamily was described by j . a . baskin in 1998 , who assigned seven genera to it -
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