cranopsis is a genus of sea snails , marine gastropod mollusks in the family fissurellidae , the keyhole limpets .

cranopsis was previously considered a subgenus of puncturella .

cranopsis ( gastropod )

cranopsis ( gastropod ) <EOT> worms - world register of marine species - cranopsis asturiana ( p . fischer , 1882 )

puncturella ( craniopsis ) [ sic ] ( misspelling of cranopsis a . adams , 1860 )

cowan , i . m . ( 1969 ) a new species of gastropod ( fissurellidae , fissurisepta ) from the eastern north pacific . the veliger , 12 , 24–26 .

( of puncturella ( cranopsis ) asturiana ( p . fischer , 1882 ) ) pérez farfante , i . ( 1947 ) . the genera zeidora , nesta , emarginula , rimula and puncturella in the western atlantic . johnsonia . 2 : 93 - 148 . , available online at urltoken page ( s ) : 118 - 120 [ details ]

( of rimula asturiana p . fischer , 1882 ) fischer p . ( 1882 - 1883 ) . diagnoses d ' espèces nouvelles de mollusques recueillis dans le cours des expéditions scientifiques de l ' aviso " le travailleur " ( 1880 et 1881 ) . journal de conchyliologie 30 : 49 - 53 [ 1882 ] , 273 - 277 [ issued march 22 , 1883 according to fischer - piette ( 1937 ) , april 1883 according to winckworth , 1936 ] , available online at urltoken page ( s ) : 51 [ details ]

( of puncturella asturiana var . alta locard , 1898 ) locard a . ( 1897 - 1898 ) . expéditions scientifiques du travailleur et du talisman pendant les années 1880 , 1881 , 1882 et 1883 . mollusques testacés . paris , masson . vol . 1 [ 1897 ] , p . 1 - 516 pl . 1 - 22 ; vol . 2 [ 1898 ] , p . 1 - 515 , pl . 1 - 18 . , available online at urltoken page ( s ) : vol . 2 p . 78 [ details ]

gofas , s . ; le renard , j . ; bouchet , p . ( 2001 ) . mollusca . in : costello , m . j . et al . ( eds ) , european register of marine species : a check - list of the marine species in europe and a bibliography of guides to their identification . patrimoines naturels . 50 : 180 - 213 . ( look up in imis ) [ details ]

rosenberg , g . ; moretzsohn , f . ; garcía , e . f . ( 2009 ) . gastropoda ( mollusca ) of the gulf of mexico , pp . 579–699 in : felder , d . l . and d . k . camp ( eds . ) , gulf of mexico–origins , waters , and biota . texas a & m ; press , college station , texas . , available online at urltoken [ details ]

turgeon , d . ; quinn , j . f . ; bogan , a . e . ; coan , e . v . ; hochberg , f . g . ; lyons , w . g . ; mikkelsen , p . m . ; neves , r . j . ; roper , c . f . e . ; rosenberg , g . ; roth , b . ; scheltema , a . ; thompson , f . g . ; vecchione , m . ; williams , j . d . ( 1998 ) . common and scientific names of aquatic invertebrates from the united states and canada : mollusks . 2nd ed . american fisheries society special publication , 26 . american fisheries society : bethesda , md ( usa ) . isbn 1 - 888569 - 01 - 8 . ix , 526 + cd - rom pp . ( look up in imis ) page ( s ) : 57 [ details ]

( of puncturella asturiana ( p . fischer , 1882 ) ) locard a . ( 1897 - 1898 ) . expéditions scientifiques du travailleur et du talisman pendant les années 1880 , 1881 , 1882 et 1883 . mollusques testacés . paris , masson . vol . 1 [ 1897 ] , p . 1 - 516 pl . 1 - 22 ; vol . 2 [ 1898 ] , p . 1 - 515 , pl . 1 - 18 . , available online at urltoken page ( s ) : vol . 2 , 77 - 78 [ details ]

medin . ( 2011 ) . uk checklist of marine species derived from the applications marine recorder and unicorn . version 1 . 0 . [ details ]

sysoev a . v . ( 2014 ) . deep - sea fauna of european seas : an annotated species check - list of benthic invertebrates living deeper than 2000 m in the seas bordering europe . gastropoda . invertebrate zoology . vol . 11 . no . 1 : 134–155 [ in english ] . [ details ] available for editors [ request ]

to biodiversity heritage library ( 11 publications ) to biodiversity heritage library ( 16 publications ) ( from synonym rimula asturiana p . fischer , 1882 ) to clemam ( from synonym rimula asturiana p . fischer , 1882 ) to clemam to encyclopedia of life to pesi to pesi ( from synonym rimula asturiana p . fischer , 1882 ) to usnm invertebrate zoology mollusca collection ( from synonym rimula asturiana p . fischer , 1882 ) to itis

adams , a . ( 1860 ) . on some new genera and species of mollusca from japan . annals and magazine of natural history . ( 3 ) 5 : 299 - 303 [ april 1860 ] ; 405 - 413 . , available online at urltoken page ( s ) : 302 [ details ]

mclean j . h . & geiger d . l . ( 1998 ) . new genera and species having the fissurisepta shell form , with a generic - level phylogenetic analysis ( gastropoda : fissurellidae ) . contributions in science , natural history museum of los angeles county , 475 : 1 - 32 . , available online at urltoken [ details ]

mclean j . h . ( 1996 ) . the prosobranchia . in : taxonomic atlas of the benthic fauna of the santa maria basin and western santa barbara channel . the mollusca part 2 – the gastropoda . santa barbara museum of natural history . volume 9 : 1 - 160 . [ details ]

( of puncturella ( craniopsis ) [ sic ] ) nordsieck f . ( 1968 ) . die europäischen meeres - gehäuseschnecken ( prosobranchia ) . vom eismeer bis kapverden und mittelmeer . gustav fischer , stuttgart viii + 273 pp : page ( s ) : 12 [ details ]

html public " - / / w3c / / dtd html 4 . 0 transitional / / en "

html public " - / / w3c / / dtd html 4 . 01 / / en " " urltoken "

the following are opisthobranchs with shells , but are included here to aid in identification .

all material posted on this site may be used for educational or non - commercial use provided this web site is duly credited as being the source of the material . copyright of all images remains with the originator . for the use of photos , please request permission through one of the contact listings .

click on the name of the family to view the species within that category .

html public " - / / w3c / / dtd xhtml 1 . 0 transitional / / en " " urltoken "

this site uses cookies to improve performance . if your browser does not accept cookies , you cannot view this site .

there are many reasons why a cookie could not be set correctly . below are the most common reasons :

you have cookies disabled in your browser . you need to reset your browser to accept cookies or to ask you if you want to accept cookies .

your browser asks you whether you want to accept cookies and you declined . to accept cookies from this site , use the back button and accept the cookie .

your browser does not support cookies . try a different browser if you suspect this .

the date on your computer is in the past . if your computer ' s clock shows a date before 1 jan 1970 , the browser will automatically forget the cookie . to fix this , set the correct time and date on your computer .

you have installed an application that monitors or blocks cookies from being set . you must disable the application while logging in or check with your system administrator .

this site uses cookies to improve performance by remembering that you are logged in when you go from page to page . to provide access without cookies would require the site to create a new session for every page you visit , which slows the system down to an unacceptable level .

this site stores nothing other than an automatically generated session id in the cookie ; no other information is captured .

in general , only the information that you provide , or the choices you make while visiting a web site , can be stored in a cookie . for example , the site cannot determine your email name unless you choose to type it . allowing a website to create a cookie does not give that or any other site access to the rest of your computer , and only the site that created the cookie can read it .

luiz ricardo l . simone museu de zoologia da universidade de são paulo , cx . postal 42494 ; 04218 - 970 são paulo , sp , brazil .

carlo m . cunha museu de zoologia da universidade de são paulo , cx . postal 42494 ; 04218 - 970 são paulo , sp , brazil .

absalão , r . s . & pimenta , a . d . ( 2003 ) a new subgenus and three new species of brazilian deep water olivella swainson , 1831 ( mollusca , gastropoda , olivellidae ) collected by the rv marion dufresne in 1987 . zoosystema , 25 , 177–185 .

absalão , r . s . , pimenta , a . d . & caetano , c . h . s . ( 2005 ) turridae ( mollusca , neogastropoda , conoidea ) coletados no litoral sudeste do brasil , programa revizee “score” central . biociências , 13 , 19–47 .

adams , a . ( 1851 ) a catalogue of the species of emarginula , a genus of gasterepodous mollusca , belonging to the family fissurellidae ; in the collection of h . cuming , esq . proceedings of the zoological society of london , 19 , 82–92 . urltoken

adams a . ( 1860 ) on some new genera and species of mollusca from japan . annals and magazine of natural history , 3 ( 5 ) , 299–307 .

aldea , c . , zelaya , d . g . & troncoso , j . s . ( 2011 ) a new gigantic species of zeidora adams , 1860 from antarctic waters ( gastropoda : fissurellidae ) . the nautilus , 125 , 79–82 .

amaral , a . c . z . , lana , p . c . , fernandes , f . c . & coimbra , j . c . ( 2003 ) biodiversidade bêntica da região sul - sudeste da costa brasileira . revezee score sul . ministério do meio ambiente . são paulo , 156 pp .

betts , v . ( 1981 ) dicionário parintintin - português . sociedade internacional de lingüística . cuiabá , 231 pp .

clarke , a . h . ( 1961 ) abyssal mollusks from the south atlantic ocean . bulletin of the museum of comparative zoology , 125 , 343–387 , pls . 1–4 .

cunha , c . m . ( 2011 ) a new species of acteon ( opisthobranchia : acteonidae ) from northeast brazil . zoologia , 28 , 229–232 . urltoken

dall , w . h . ( 1889 ) reports on the results of dredgings , under the supervision of alexander agassiz , in the gulf of mexico ( 1877–78 ) and in the caribbean sea ( 1879–80 ) , by the u . s . coast survey steamer ' blake ' . bulletin of the museum of comparative zoology , 18 , 1–492 , pls . 10–40 .

dall , w . h . ( 1927 ) small shells from dredgings off the southeast coast of the united states by the united states fisheries steamer ' albatross ' in 1885 and 1886 . proceedings of the united states national museum , 70 , 1–134 . urltoken

dautzenberg , p . & fischer , h . ( 1896 ) dragages effectués par l ' hirondelle et par la princesse - alice , 1888 - 1895 . mémoires de la société zoologique de france , 9 , 395–498 , pls . 15–22 .

defrance , m . ( 1827 ) dictionnaire des sciences naturelles , 45 , 471–472 .

egorova , e . n . ( 1972 ) biological results of the soviet antarctic expeditions . 7 . mollusca of the davis sea . explorations of the faunas of the seas , 26 , 1–142 .

espinosa , j . , ortea , j . & fernández - garcés , r . ( 2004 ) descripción de dos nuevas especies del género zeidora a . adams , 1860 ( mollusca : gastropoda ) de las costas de cuba . avicennia , 17 , 67–70 .

farfante , i . p . ( 1943a ) the genera fissurella , lucapina and lucapinella in the western atlantic . johnsonia , 1 ( 10 ) , 1–20 .

farfante , i . p . ( 1943b ) the genus diodora in the western atlantic . johnsonia , 1 ( 11 ) , 1–20 .

farfante , i . p . ( 1947 ) the genera zeidora , nesta , emarginula , rimula and puncturella in the western atlantic . johnsonia , 2 , 93–148 .

fernandes , m . r . & pimenta , a . d . ( 2013 ) taxonomic review of metaxia ( gastropoda : triphoridae ) from brazil , with description of a new species . zoologia , 28 , 819–830 . urltoken

fernandes , m . r . , pimenta , a . d . & leal , j . h . ( 2013 ) taxonomic review of triphorinae ( gastropoda : triphoridae ) from the vitória - trindade seamount chain , southeastern brazil . the nautilus , 127 , 1–18 .

garcía , e . f . ( 2011 ) a new species of eccliseogyra ( gastropoda : nystiellidae ) from southeastern brazil . the nautilus , 125 , 167–170 .

geiger , d . l . & thacker , c . e . ( 2005 ) molecular phylogeny of vetigastropoda reveals non - monophyletic scissurellidae , trochoidea , and fissurelloidea . molluscan research , 25 , 47–55 .

geiger , d . l . & thacker , c . e . ( 2006 ) molecular phylogeny of basal gastropods ( vetigastropoda ) shows stochastic colonization of chemosynthetic habitats at least from the mid triassic . cahiers de biologie marine , 46 , 343–346 .

gray , j . e . ( 1825 ) a list and description of some species of shells not taken notice of by lamarck . annals of philosophy , 25 ( n . s . 9 ) , 134–140 .

gray , j . e . ( 1847 ) a list of the genera of recent mollusca , their synonyma and types . proceedings of the zoological society of london , 12 , 129–219 .

heilprin , a . ( 1889 ) on some new species of mollusca from the bermuda islands . proceedings of the academy of natural sciences of philadelphia , 41 , 141–142 , pl . 8 .

houart , r . ( 1991 ) the southeastern brazilian muricidae collected by rv marion - dufresne in 1987 , with the description of three new species . the nautilus , 105 , 26–37 .

jeffreys , j . g . ( 1877 ) new and peculiar species of mollusca procured in the valourous expedition . annals and magazine of natural history , series 4 , 19 , 231–243 .

jeffreys , j . g . ( 1882 ) on the mollusca procured during the lightning and porcupine expeditions . iv . proceedings of the zoological society of london ( 1881 ) , 922–952 .

lamarck , j . b . p . m . ( 1798–1816 ) tableau encyclopédique et méthodique des trois règnes de la nature . vol . 3 . paris , 16 pp . , pls . 391–488 .

lamarck , j . b . p . m . ( 1801 ) systême des animaux sans vertèbres . paris , viii , 432 pp .

lamarck , j . b . p . m . ( 1822 ) histoire naturelle des animaux sans vertèbres . paris , 7 , 711 pp .

larrazábal , m . e & oliveira , v . s . ( 2003 ) thecosomata e gymnosomata ( mollusca , gastropoda ) da cadeia fernando de noronha , brasil . revista brasileira de zoologia , 20 ( 2 ) , 351–360 . urltoken

leal , j . h . ( 1991 ) marine prosobranch gastropods from oceanic islands off brazil . universal book services . dr . w . backhuys . oegstgeest , 419 pp .

leal , j . h . & bouchet , p . ( 1989 ) new deep - water volutidae from off southeastern brazil ( mollusca : gastropoda ) . the nautilus , 103 , 1–12 .

libassi , i . ( 1859 ) sopra alcune conchiglie fossile dei intorni de palermo . reale academia di scienze , lettere i belle arti di palermo , atti ( n . s . ) , 3 , 1–47 .

linné , c . ( 1771 ) mantissa plantarum . laurentii salvii . holmiae [ stockholm , sweden ] , pp . 43–588 .

lowe , r . t . ( 1827 ) on balanus punctatus , puncturella flemingii , & c . ; together with some corrections relative to turbo carneus , and some of the chitones before described . zoological journal , 3 , 76–80 .

matthews , h . r . & kempf , m . ( 1970 ) moluscos marinhos do norte e nordeste do brasil . ii . moluscos do arquipélago de fernando de noronha ( com algumas referências ao atol das rocas ) . arquivos de ciências do mar , 19 , 1–53 .

mclean , j . h . ( 1970 ) descriptions of a new genus and eight new species of eastern pacific fissurellidae , with notes on other species . the veliger , 12 , 362–367 .

mclean , j . h . ( 2011 ) reinstatement of the fissurellid subfamily hemitominae , with the description of new genera , and proposed evolutionary lineage , based on morphological characters of shell and radula ( gastropoda : vetigastropoda ) . malacologia , 54 , 407–427 . urltoken

mclean , j . h . & geiger , d . j . ( 1998 ) new genera and species having fissurisepta shell form , with a generic - level phylogenetic analysis ( gastropoda : fissurellidae ) . los angeles county museum of natural history contributions in science , 475 , 1–32 .

métivier , b . ( 1972 ) sur quelques fissurellidae ( mollusques , gastéropodes ) du nord , nordest du brésil . bulletin du muséum national d’histoire naturelle , series 3 ( zoologie ) , 32 , 405–416 .

pilsbry , h . a . ( 1943 ) floridian species of rimula . the nautilus , 57 , 37–40 .

pimenta , a . d . , santos , f . n . & absalão , r . s . ( 2011 ) taxonomic revision of the genus eulimella ( gastropoda , pyramidellidae ) from brazil , with description of three new species . zootaxa , 3063 , 22–38 .

rios , e . c . ( 1985 ) seashells of brazil . fundação universidade do rio grande . rio grande , 329 pp . , 102 pls .

rios , e . c . ( 1994 ) seashells of brazil , second edition . editora da furg . rio grande , 368 pp . , 113 pls .

rios , e . c . ( 2009 ) compendium of brazilian sea shells . universidade federal do rio grande . rio grande , 668 pp .

rios , e . c . , calvo , i . s . & barcellos , l . j . ( 1987 ) moluscos marinos de isla trinidad . comunicaciones de la sociedad malacologica del uruguay , 7 , 57–62 .

salvador , r . b . , cunha , c . m . & simone , l . r . l . ( 2013 ) taxonomic revision of the orthalicid land snails ( pulmonata : stylommatophora ) from trindade island , brazil . journal of natural history , 47 , 949–961 . urltoken

salvini - plawén , l . & haszprunar , g . ( 1987 ) vetigastropoda and the systematics of streptoneurous gastropoda ( mollusca ) . journal of zoology , a 211 , 747–770 . urltoken

seguenza , g . ( 1862 ) paleontologia malacologica dele roce terziarie del discreto di messina studiata nei suoi rapporti zooloogici e geognostici . annali dell’accademia degli aspiranti naturalisti , serie 3 , 2 , 77–95 .

simone , l . r . l . ( 1999 ) comparative morphology and systematics of brazilian terebridae ( mollusca , gastropoda , conoidea ) , with descriptions of three new species . zoosystema , 21 , 199–248 .

simone , l . r . l . ( 2000 ) [ 1998 ] a phylogenetic study of the terebrinae ( mollusca , caenogastropoda , terebridae ) based on species from the western atlantic . journal of comparative biology , 3 , 137–150 .

simone , l . r . l . ( 2008 ) a new species of fissurella from são pedro e são paulo archipelago , brazil ( vetigastropoda , fissurellidae ) . the veliger , 50 , 292–304 .

simone , l . r . l . & cunha , c . m . ( 2003 ) pseudococculina rimula , a new species ( cocculiniformia : pseudococculinidae ) from off southeastern brazil . nautilus , 117 , 69–77 .

simone , l . r . l . & cunha , c . m . ( 2012 ) taxonomic study on the mollusks collected in marion - dufresne expedition ( md55 ) to se brazil : xenophoridae , cypraeoidea , mitriforms and terebridae ( caenogastropoda ) . zoosystema , 34 , 745–781 . urltoken

sowerby , g . b . i . ( 1812–1846 ) the mineral conchology of great britain ; or coloured figures and descriptions of those reamains of testaceous animals or shells , which have been preserved at various times and depth in the earth . vols . 1 . –7 . london , 234 pp . , 251 pp . , 194 pp . , 160 pp . , 168 pp . , 250 . & 123 pp . + pls 610–648 .

sowerby , g . b . iii ( 1901 ) descriptions of five new species of shells . journal of malacology , 8 , 101–103 , pl . 9 .

swainson , w . ( 1840 ) a treatise on malacology . longman , orme , brown , green , & longmans and john taylor . london , 419 pp .

verhecken , a . ( 1991 ) description of two new species of bathyal cancellariidae ( mollusca , gastropoda ) from off brazil . bulletin du muséum national d ' histoire naturelle ( zoologie ) , 12 , 547–553 .

watson , r . b . ( 1883 ) mollusca of h . m . s . ' challenger ' expedition . part xix . zoological journal of the linnean society , 17 , 320–340 . urltoken

warning : the ncbi web site requires javascript to function . more . . .

regina l . cunha , a , 1 jorge m . assis , 1 celine madeira , 1 rui seabra , 2 fernando p . lima , 2 evandro p . lopes , 2 , 3 suzanne t . williams , 4 and rita castilho 1

this work is licensed under a creative commons attribution 4 . 0 international license . the images or other third party material in this article are included in the article’s creative commons license , unless indicated otherwise in the credit line ; if the material is not included under the creative commons license , users will need to obtain permission from the license holder to reproduce the material . to view a copy of this license , visit urltoken

remote oceanic archipelagos are the ideal setting for studying patterns and processes underlying speciation . while insular terrestrial communities can be formed either by immigration or from a few colonization events and subsequent in situ diversification 1 , phylogeographic patterns of marine species inhabiting island settings can be confounded by recurrent episodes of long - distance dispersal . the volcanic origin of oceanic islands and their circumscribed geographic boundaries enable inferring the tempo and sequence of island colonization when geological ages are known 2 .

isolated islands are expected to have reduced species richness but high levels of endemicity as a result of few colonization events , particularly in older archipelagos 3 . for instance , the reduced dispersal abilities of some indo - west pacific turbinid gastropods that lack a long - lived planktonic stage played a crucial role in developing extensive archipelagic differentiation and fine scale endemism 4 . also , the ongoing emergence and subsidence of islands over geological time is thought to have promoted speciation within this group of marine gastropods at an insular scale 5 .

allopatric speciation induced by vicariance is sometimes regarded as the main driver of differentiation 6 . nonetheless , recent developments in methodological approaches to biogeographic analysis 7 , which integrate a wider range of biogeographical processes ( i . e . dispersal , extinction , vicariance and duplication ) 8 , 9 , showed the importance of dispersal in the diversification processes 10 . for instance , divergence triggered by dispersal events and the organism - specific capacity to occupy suitable habitats and persist , has recently been identified as the main driver of the avian speciation in lowland neotropical rainforests 11 .

dispersal in most sessile marine species occurs predominantly during larval stages in which larvae remain in the water column for periods that can vary between days to months before settlement 12 . it is generally assumed that planktotrophic feeding larvae exhibit higher geographic ranges because they are able to remain longer periods in the water column , while lecithotroph yolk - fed larvae complete their metamorphosis without feeding from the plankton and thus are more prone to originate locally structured populations 13 , 14 .

cape verde represents an excellent model system for studying speciation and to infer colonization pathways given its remoteness and known geological age of most of the islands . this volcanic oceanic archipelago is approximately 500 km off mainland ( senegal - west africa ) and comprises ten major islands and several islets (

) . geochronological data place the age of the islands between 5 . 9 ± 0 . 1 million years ago ( mya ) and 25 . 6 ± 1 . 8 mya

. bi analysis retrieved two clades of cape verde fissurellidae . one clade included all specimens from cape verde that grouped with the included worldwide members of fissurellinae . the other clade grouped all cape verde samples with western atlantic and mediterranean

phylogenetic relationships of fissurellidae based on a bayesian inference analysis of mitochondrial coi and nuclear 28s rrna genes produced by beast .

bayesian posterior probabilities above 50 % are indicated over the branches ( full black circles : bpp = 100 % ; half - black circle : bpp = 86 % ) . geographical origin of the taxa is depicted in colours . photos of the shells corresponding to each of the cape verde fissurella and diodora species are shown . assignments of each cape verde sample to delineated entities from gmyc and spedestem species delimitation tests are shown by rectangles . colour of cluster boxes corresponding to each species indicates mismatch ( black ) and agreement ( white ) between both methods . figure generated in adobe illustrator cs6 ( version 16 . 0 . 0 ) ( urltoken ) and photos taken by r . l . cunha .

) . specimens from south africa and angola were recovered as new species of uncertain generic status and named as fissurellidae sp . 1 and fissurellidae sp . 2 , respectively .

) . effective sample sizes ( ess ) ≫ 200 indicated adequate sampling of the posterior . the 95 % hpd ( highest posterior density interval ) sampled by bamm after analysing 9 , 001 posterior samples comprised the eight most probable distinct shift configurations . the single best shift configurations sampled by bamm suggested an increasing of diversification rates in cape verde

) . overall extinction rates were 0 . 046 ( 0 . 006–0 . 109 ) and remained constant through time . speciation rates were 0 . 089 ( 0 . 058–0 . 132 ) and an overall increase towards the present was identified .

was estimated at 10 . 21 [ 7 . 99–12 . 72 ] mya ( node a ;

) but most of the diversification within this group occurred during the late pliocene ( 2 . 98 [ 2 . 19–3 . 89 ] mya ; node d ,

) was estimated at 6 . 74 [ 5 . 29–8 . 41 ] mya ( node g ;

beast maximum clade credibility chronogram showing main cladogenetic events within fissurellidae based on mitochondrial coi and nuclear 28s rrna genes with ancestral estimation inferred with biogeobears .

age estimates in million years and bayesian posterior probabilities ( bpp ) of cape verde clades are shown on the table . the corresponding 95 % highest posterior density intervals ( blue bars ) are depicted . internal coloured vertical bars at branches indicate main ancestral areas recovered by

under the selected bayarealike + j model immediately following a speciation event whereas at nodes indicate ancestral ranges before speciation . numbers at the coloured squares indicate present - day ( tips ) distribution of species . in the legend , islands that are not represented , individually , by a single species are not coloured .

sp . ) , which was the pacific . this model suggests boavista as the ancestral range of

comprises the northwestern islands of the archipelago ( santo antão , ilhéu raso , santa luzia , são vicente and são nicolau ) . our results suggest a dispersal event towards sal where

is sal , boavista , maio , santiago and são vicente , which implies dispersal events from the northwestern islands ( the ancestral area ) towards sal , boavista and maio .

the total amount per cell of the coastline of rocky substrate ( in grey ) and sand ( in yellow ) on each island is shown in supplementary information s3 . the percentage per cell of the coastline of rocky substrate ( in grey ) and sand ( in yellow ) on each island is shown in supplementary information s4 . boavista is the island with the highest percentage of sand whereas ilhéu raso and santiago show the highest percentage of rocky substrate ( supplementary information s4 ) .

the simulations using high - resolution ocean current fields over the 10 - year period allowed releasing 360 particles per cell ( 6 . 39e

passive particles in total ) . maximum and average distances , probabilities and drifting time produced by the particles that effectively connected different coastal cells determined for the simulations running 4 and 30 days of passive dispersal are shown in

. the mean distance that particles can reach during four days of passive dispersal is 75 . 3 ± 75 . 9 km , which is approximately the same after 30 days ( 76 . 1 ± 75 . 9 km ) . mean connectivity probabilities between pairs of islands produced with simulations running 4 and 30 days of passive dispersal are shown in

, respectively . overall probabilities of connectivity between pairs of islands are quite low ( between 2 . 3e

) . the highest probability of connectivity occurs between são vicente and santo antão and the lowest between santo antão and maio . the degree of connectivity between islands inferred in network analysis running for four or 30 days of passive dispersal are shown in

( a and b ) , respectively . only stronger links are depicted . modularity values indicated good divisions > 0 . 3

. the animation shows different source locations releasing particles every 12 hours from july to november 2010 . the particles are allowed to drift for a maximum of 30 days with effective dispersal measured when they end up on shore .

( a ) degree of connectivity between cape verde islands inferred in network analysis for the simulation running four days of passive dispersal . only stronger links are depicted ( modularity : 0 . 38 ; three putative oceanographic clusters ; significance level of clustering : 0 . 0048 ) ; ( b ) degree of connectivity between cape verde islands inferred in network analysis for the simulation running 30 days of passive dispersal . only stronger links are depicted ( modularity : 0 . 34 ; two putative oceanographic clusters ; significance level of clustering : 0 . 0291 ) ; ( c ) example of pathways resulting from all particles sent from a coastal cell ( red dot ) in the simulation running four days of passive dispersal ; ( d ) example of pathways resulting from all particles sent from a coastal cell ( red dot ) in the simulation running 30 days of passive dispersal . figures were generated with igraph package ( version 1 . 0 . 1 ; url urltoken ) implemented in r language ( r core team ( 2015 ) . r : a language and environment for statistical computing . r foundation for statistical computing , vienna , austria . url urltoken ) and adobe illustrator cs6 ( version 16 . 0 . 0 ) ( urltoken ) .

maximum and average distances , probabilities and drifting time produced by the particles that effectively connected different coastal cells determined for the simulations running 4 and 30 days of passive dispersal .

mean connectivity matrix between pairs of islands produced with the simulation running 4 days of passive dispersal .

mean connectivity matrix between pairs of islands produced with the simulation running 30 days of passive dispersal .

from the mediterranean and western atlantic . both species delimitation tests ( spedestem and gmyc ) indicated the existence of seven cape verde endemic

species of the mediterranean , western atlantic and pacific , whereas gmyc did recognise cape verde specimens as a distinct entity . however , results from both gmyc and genetic distances between cape verde

) , strongly suggest that these specimens should be considered an endemic species . these results are not in agreement with a morphology - based study that reported the existence of twelve

was unexpected , even for organisms with a theoretical pelagic larval phase of four days , considering that distances between islands are as little as 17 km ( e . g . , between ilhéu raso and s . nicolau ;

) , and according to our simulations , the mean distance that a particle can reach after four days of passive dispersal is 75 . 3 ± 75 . 9 km (

, which represents a much larger distance than between most of islands of the cape verde archipelago . nonetheless , this coastal species coexists with its sister species along the continental shore , where no strong physical barriers were identified . it was suggested that transient allopatry determined by historical episodes of climatic fluctuations and shifting currents or ecological barriers could be the main driver of speciation

. considering the extremely low estimated probabilities of connectivity between cape verde islands based on four or 30 days of passive dispersal ( e . g . , ilhéu raso

) . age estimates placed this shift at 2 . 98 [ 2 . 19–3 . 89 ] mya (

) at the plio - pleistocene boundary . key innovations or distinct habitat preferences are often invoked to explain higher rates of diversification

. all eight species occupied similar habitats and we found no evidence for niche segregation among species ( lopes , evandro p . , pers . obs . ) .

the low levels of connectivity between islands and a shift in diversification rates offer plausible explanations for the level of endemism observed in fissurella but a question still remains : why did diodora not diversify in the cape verde archipelago ? to address this question , three hypotheses were considered : ( i ) different sampling efforts for each genus ; ( ii ) differences between the pld of fissurella and diodora , and ( iii ) time of origin of each genus in the archipelago .

could have been undersampled seems unlikely considering that the sampling effort was equally distributed for both genera . field work revealed a strong bias in abundances : while more than 300 specimens of

specimens in total , all from boavista and all belonging to the same species . the type of larval development of cape verde

is unknown but under the observed oceanographic conditions , the second hypothesis of a longer pld to explain the existence of a single species owing to higher connectivity is not supported . our simulations based on patterns of ocean circulation did not significantly increase effective connectivity between islands even when pld was increased from four to 30 days of passive dispersal (

) . note that only effective dispersal events ( i . e . those that result in a larva landing on a coast ) are reported . our analyses show very low probabilities of shore - to - shore connectivity ( 0 . 003 ± 0 . 017 , on average ;

) , since most particles are pushed to open waters and only produce effective connectivity events in the very first days of ocean drifting ( between 1 . 29 ± 0 . 85 and 1 . 35 ± 1 . 02 days , on average ;

) . in fact , only 2 . 62 % of the particles travelled for more than four days , and such an increase in pld did not expand the maximum travelled distance of the particles ( 349 . 3 km ;

) . as the archipelago is separated from the nearest mainland ( the coast of senegal ) by approximately 500 km , this means that island - to - continent dispersal is very unlikely . finally , the mrca of

might have had more opportunities to disperse to other islands and speciate avoiding the severe volcanic eruptions that struck boavista between 9 . 5 and 4 . 5 mya

and likely had the greatest effect on rocky shore species such as keyhole limpets . other

species also present in boavista might have become extinct after these volcanic events , before having time to disperse to other islands .

keyhole limpets can only be found on rocky substrates , we analysed the effect of hard substrate availability on species richness . the number of

lineages showed , however , no correlation with the total area of rocky coast per island [ r ( 6 ) = −0 . 62 , p = 0 . 10 ] nor with the percentage of rocky coast per island [ r ( 6 ) = −0 . 50 , p = 0 . 22 , non - significant , see also

on the other hand , the degree of connectivity between islands provides the best predictor for present - day distribution of species . a model based on known patterns of ocean circulation and four days of passive dispersal (

) . the cluster that includes the northwestern islands ( s . nicolau , santo antão , s . vicente , santa luzia and ilhéu raso ) represents the geographic distribution of

sp . 2 ( sal and s . nicolau ) is favoured by patterns of ocean circulation that connect these two islands (

the archipelagic endemism in marine sessile organisms reported here was driven by shifts in diversification rates promoted by recurrent episodes of low sea levels during the plio - pleistocene boundary and patterns of ocean circulation favouring self - recruitment . the role of dispersal and persistence was determinant in shaping present - day geographic distribution of fissurellid keyhole limpets .

) . all specimens were preserved in 96 % ethanol . dr . rolán , e . , a recognized expert in cape verde invertebrate fauna and author of

using the - - auto option that automatically selects the appropriate strategy according to data size . alignments of both coi ( 540 bp ) and 28s rrna ( 826 bp ) were unambiguous , and amino acid translations in coi were checked using

, respectively . we used the combined data set ( coi : 145 taxa , 540 bp ; 28s rrna : 145 taxa , 826 bp ) for both analyses . this data set included all 120 unique coi and 28s haplotypes from cape verde samples and the remaining fissurellidae retrieved from genbank ( accession numbers in

selected the trn + i + g as the evolutionary model that best fits both data sets . the bayesian analysis was performed using two partitions , coi and 28s rrna . we used a yule tree prior

that assumes a constant rate of speciation among lineages and is more appropriate for species - level phylogenies . we used an uncorrelated relaxed , lognormal clock . mcmc analyses were run twice ( each run with 100 , 000 , 000 generations and a sample frequency of 10 , 000 ) following a discarded burn - in of 10 , 000 , 000 steps . length of burn - in was determined by visual examination of traces in

using the “maximum clade creditability” option and mean node height . the convergence to the stationary distribution was confirmed by inspection of the mcmc samples and of effective sample sizes ( ess ) . ess values above 200 indicate convergence

) and on the r2c2 research group cluster facility , both at the university of algarve .

, code written by t . ezard , t . fujisawa and t . barraclough in r v . 3 . 0 . 2

to compare the number of esus obtained from the single gene ( coi : 540 bp ; 145 taxa ) and the two - gene ( coi : 540 bp ; 28s : 826 bp ; 145 taxa ) data sets . the ultrametric trees based on the coi and the combined data sets were obtained with

using a strict clock model without fossil calibrations and a yule tree prior . both data sets included all 120 unique haplotypes from cape verde samples and remaining fissurellidae retrieved from genbank ( accession numbers in table 1 suplementary material ) . the analyses ran for 10 , 000 , 000 generations with sample frequency of 1000 , after a burn - in of 1000 , 000 .

to delimit the number of fissurellid species in cape verde by comparing the probability of models where putative evolutionary lineages are separate entities to the probability of models where putative lineages are collapsed into a single lineage using a maximum likelihood approach . spedestem takes as an input ultrametric trees and a user - supplied estimate of theta returning a table of models ranked according to their probability . ultrametric trees based on the coi and 28s data sets were produced by

using the coi ( 540 bp ; 145 taxa ) and the 28s ( 826 bp ; 145 taxa ) data sets . the analyses ran for 10 , 000 , 000 generations with sample frequency of 1000 , after a burn - in of 1000 , 000 . we used

that allows incorporation of fossil uncertainties . the data set used in this analysis ( 35 taxa ; coi : 540 bp ; 28s rrna : 826 bp ) included a single representative from each cape verde species inferred by abgd and spedestem and the remaining fissurellidae used in previous analyses . the calibration points used in this analysis are described in the

. this program uses mcmc simulations and reversible - jump sampling to estimate time - varying rates of speciation and extinction , and to find the optimal set of rate - shift configurations .

ultrametric tree was used for this analysis . we set four reversible jumping - mcmc running for 10 million generations sampled every 1000 generations and a burn - in of 10 % . the function setbammpriors in r was used to choose more appropriate prior values . we used ess ( effective sample size ) to assess the convergence of the runs and considered values above 200 as indicating convergence .

to estimate the ancestral ranges of fissurellidae . full description of the method is available on

. we defined 13 geographical areas : ( 1 ) sal ; ( 2 ) boavista ; ( 3 ) maio ; ( 4 ) santo antão ; ( 5 ) santiago ; ( 6 ) ilhéu raso ; ( 7 ) santa luzia ; ( 8 ) são vicente ; ( 9 ) são nicolau ; ( 10 ) mediterranean ; ( 11 ) western atlantic ; ( 12 ) pacific , and ( 13 ) africa . the maximum number of areas that any species may occur was set to five because it is the maximum number of areas where the species may occur .

to characterize shore substrate composition along the studied area , we prepared a 0 . 01° arc - degree ( approx . 1 . 6 km at 16° n ) mesh using r ( r development core team , 2014 ) . the mesh grid was then imported to google earth , and all tiles covering both ocean and land were assigned a substrate type by means of visual inspection . substrate types used were “rock” , “sand” or “both” . lastly , the substrate type layers produced in google earth were rasterised using the r package raster 54 , and the amount of each substrate was quantified .

how to cite this article : cunha , r . l . et al . drivers of cape verde archipelagic endemism in keyhole limpets . sci . rep . 7 , 41817 ; doi : 10 . 1038 / srep41817 ( 2017 ) .

publisher ' s note : springer nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations .

we thank the direcção geral do ambiente , ministério do ambiente , habitação e ordenamento do território from cape verde for providing sampling permits . we are very grateful to dr . emílio rolán for the help provided in species identification . we also thank katy nicastro and gerardo zardi for providing samples from south africa . rlc and ja were supported by post - doctoral fellowships from fct - portuguese science foundation ( sfrh / bpd / 109685 / 2015 and sfrh / bpd / 111003 / 2015 , respectively ) . rs was supported by marinfo - norte - 01 - 0145 - feder - 000031 , funded by norte portugal regional operational program ( norte2020 ) , under the portugal 2020 partnership agreement , through the european regional development fund ( erdf ) .

author contributions r . l . c . and r . c . designed this study . e . p . l . collected the samples . c . m . sequenced all the samples . r . l . c . , j . a . , r . s . , and f . l . analyzed data . r . c . prepared figures and tables . r . l . c . wrote the manuscript and s . t . w . , r . c . , and f . l . revised it . all authors contributed with their ideas and reviewed the final version of the manuscript .

reynoso - granados t . , monsalvo - spencer p . & serviere - zaragoza e .

& guzman del proo , s . a . larval and early juvenile development of the volcano keyhole limpet , fissurella volcano

maddison w . p . & maddison d . r . mesquite : a modular system for evolutionary analysis . urltoken ( 2015 ) .

a mathematical theory of evolution , based on the conclusions of dr . j . c . willis , f . r . s

r : a language and environment for statistical computing . r foundation for statistical computing ( vienna , austria , 2016 ) .

matzke n . j . biogeobears : biogeography with bayesian ( and likelihood ) evolutionary analysis in r scripts . r package , version 0 . 2 . 1 , published july 27 , 2013 at : urltoken ( 2013 ) .

raster : geographic data analysis and modeling . r package version 2 . 1 - 25 . urltoken ( 2013 ) .


striatolamia is an extinct genus of sharks belonging to the family odontaspididae .

these extinct sharks lived in the paleocene and miocene periods ( from 61.7 to 10.3 ma ) .

striatolamia

striatolamia <EOT> carcharias cf . macrota , carcharias macrota , carcharias macrotus , eugomphodus macrota , lamna cf . compressa , lamna compressa , lamna depressa , odontaspis macrota , odontaspis macrota striata , odontaspis ( otodus ) macrota , odontaspis ( synodontaspis ) macrota , striatolamia cf . macrota , striatolamia compressa , striatolamia depressa , striatolamia elegans elegans

lateral striatolamia macrota . when view on profile the lateral teeth are very compressed .

excellent examples of striatolamia macrota - anterio - lateral teeth . a large species !

compared with the striated - crown carcharias species , the striatolamia crowns tend be more erect ( less sigmoid ) with stronger striations and more greatly reduced cusplets . in contrast to striatolamia , the striations on

about 65 million years ago , just as the dinosaurs were ending , this shark , striatolamia , appears .

fig . 1 - striatolamia striata - lateral 16 . 5 x 14 . 0 mm aquia formation ( paleocene ) , maryland

eugomphodus striatus , lamna striata , odontaspis striata , odontaspis striatus , odontaspis ( synodontaspis ) striata , otodus striatus , striatolamia cf . striata

the genus striatolamia left a fossil record with teeth that can be easily confused with those from other genera . i find it particularly disconcerting to need to know the stratigraphic position before tendering an opinion as to whether an anterior tooth might be scapanorhynchus or striatolamia , or at other times , carcharias or striatolamia . the experts may find the subtleties " obvious " , but i ' ve never achieved that comfort level .

appearing again , and the back curving shape is beginning to re - appear . this tooth is actually from a later striatolamia , a giant version

compared with the striated - crown carcharias - like species , the striatolamia crowns tend be more erect ( less sigmoid ) with stronger striations and more greatly reduced cusplets . in contrast to striatolamia , the striations on scapanorhynchus tend to be stronger and extend beyond the basal margin of the crown ' s enameloid .

purdy , r . ( 2005 ) . " is striatolamia a junior synonym of mitsukurina ? " . journal of vertebrate paleontology 25 ( 3 ) : 102a .

it is more likley that they intended to refer to siverson ( 1995 ) where the author described striatolamia cederstroemi from the danian of sweden and ascribed the genus to .

striatolamia is an extinct genus of sharks from the paleocene and eocene . its teeth are notably big and rather common in sediments of these epochs . = = systematics = = this genus had been assigned to families mitsukurinidae , odontaspididae and striatolamiidae by different authors . most widespread species of striatolamia are s . striata and s . macrota . . . .

fig . 2 - striatolamia macrota - anterior left 31 . 5 x 15 . 0 mm and right : 27 . 5 x 15 . 0 mm nanjemoy formation ( eocene ) , virginia

i have one shark tooth from washington . it ' s an incomplete sand tiger that appears to be a striatolamia tooth . the locality data says " late eocene , tukwilla formation , duwamish river , seattle , washington . " striatolamia is not known elsewhere from the late eocene . however , i ' ve read that part of the tukwila is considered late middle eocene and that would be around the time of the most recent occurrence of the genus .

this is a large anterior fossil tooth of the extinct sand tiger shark , striatolamia macrota . these teeth come from the eocene aged phosphate deposits near khouribga , morocco . the distinctive side cusps are well preserved on this beautiful tooth .

this catalog contains teeth from a large extinct sand tiger shark , striatolamia macrota . all teeth are complete . exceptional specimens . note - 1 ) all teeth sizes are slant height unless otherwise noted . click on photos to enlarge .

the muddy creek material employed by cunningham ( 2000 ) and this author , includes teeth that appear to be stunted anteriors ( la1 ) and intermediate teeth of a sand tiger - design . based on the similarities of the striatolamia macrota and carcharias taurus dental morphologies , as expressed in cunningham ( 2000 ) , it is reasonable to expect that many of these teeth belong to the most common sand tiger in the fauna . the presence of strong striations helps confirm those that are likely striatolamia in origin .

striatolamia is represented by two named species in north america , s . striata ( winkler 1874 ) from the paleocene & early eocene and s . macrota agassiz 1843 from the eocene . these two species have also been reported from europe and north africa .

remains from 10 vertebrate species were recovered by screen - washing ( george and westgate , 1998 ) . five shark species include striatolamia macrota ( sand tiger shark ) , tmm 42986 - 1 ; abdounia enniskilleni ( requiem shark ) , tmm 42986 - 2 ; . . .

well preserved teeth from the shark species - striatolamia macrota . they come from the phosphatic mines of morocco and date to the eocene epoch of around 50 million years ago . you will receive one boxed tooth as shown or similar and they measure around 25 - 30mm long .

most widespread species of striatolamia are s . striata and s . macrota . s . macrota anterior teeth have smaller roots than s . striata , and they are often recurved . another difference between these two spieces is the length of their teeth . teeth of striata are generally smaller ( 13 - 51mm ) than macrota ( 19 - 38mm )

db = nuccore | term = striatolamia % 20rossica % 20rossica | query = 1 | qty = 23 | blobid = ncid _ 1 _ 267939926 _ 130 . 14 . 22 . 215 _ 9001 _ 1531162914 _ 1251276653 _ 0meta0 _ s _ megastore _ f _ 1 | ismultiple = false | min _ list = 5 | max _ list = 20 | def _ tree = 20 | def _ list = | def _ view = | url = / taxonomy / backend / subset . cgi ? | trace _ url = / stat ?

striatolamia is represented by two named species in north america , s . striata ( winkler 1874a ) from the palaeocene & early eocene and s . macrota agassiz 1843 from the eocene . purdy ( 1998 ) attempted to synonymized s . striata with s . macrota and reported the latter from the williamsburg formation ( thanetian - late palaeocene ) of south carolina . these two species have also been reported from europe and north africa . in western europe , they are found in early ( middle ypresian ) and middle eocene sediments . ( in kazakhstan very large specimens are found in the bartonian . )

striatolamia macrota is one of the few species from nj to posse moderately strong striations on the lingual side of the tooth . these teeth range in size from 1 / 2 inch to 2 inches plus , with the average being a little over an inch . the anterior teeth have a moderately narrow elongated crown with one small ( sometimes even completely lacking ) , cusplet on each shoulder . the moderately strong striations tend to weaken the larger the tooth gets . there is a pronounced lingual protuberance and obvious nutrient grove on the root . the lateral teeth are broader and compressed , the striations are weaker or lacking entirely and the cusplets reduced , often appearing as no more than a scalloped nub . although not that uncommon , these teeth are very prone to root damage .

html public " - / / w3c / / dtd html 4 . 01 transitional / / en " " urltoken "

the anterior teeth of this genus have an elongated crown , sigmoid in shape when viewed laterally , which bears striations on the lingual face . lateral cusplets are greatly reduced or absent . kent ( 1993 ) reports these teeth ( from s . macrota ) reaching 5 . 6 cm in height . the lateral teeth are lower and broader , the striations are weaker or lacking entirely and the cusplets reduced , often appearing as no more than a scalloped heel . teeth from the eocene species , s . macrota , are larger than their paleocene counterpart and bear shorter striations . it is arguable whether there are actually two species or just an evolutionary trend toward larger teeth within a single species ( the eocene teeth termed s . striata actually being juvenile s . macrota ) .

tend to be stronger and extend beyond the basal margin of the crown ' s enameloid .

html public " - / / w3c / / dtd html 4 . 01 transitional / / en "

javascript is required to use this web site . please turn it on before proceeding .

teeth , which can in most cases , help differentiate these teeth from other sand tigers , yet showed the dentition itself to be more similar to the odontaspids rather than mitsukurinids .

another view on its phylogenetic position has been raised by siverson ( 1996 : 835 - 36 , pers . com . july 2008 ) - - it may have been derived from

is neither an odontaspid nor a mitsukurinid but a lamniform of uncertain familial affinity having evolved an odontaspid - like dentition convergently . "

. text ( 5 vols ; i . , xlix + 188 pp . , ii xii + 310 + 366 pp . , iii viii + 390 pp . , iv xvi + 296 pp . , v xii + 122 + 160 pp . ) and atlas ( 5 vols ; i 10 pl . , ii . , 149 pl . , iii 83 pl . , iv , 61 pl . , v , 91 pl . ) . neuchâtel .

chondrichthyan fishes from the paleocene of south carolina . in : paleobiology of the williamsburg formation ( black mingo group ; paleocene ) of south carolina , u . s . a . , albert sanders ed .

lamniform sharks of the mid - cretaceous alinga formation and beedagong claystone , western australia . palaeontology , vol 39 : 4 , pp 813 - 49 . 1997 . sharks from the mid - cretaceous gearle siltstone , southern carnarvon basin , western australia .

, 1878 ; vol . iv , ( fasc . 1 , 1876 ) , pp 1 - 15 ; extraits [ 1874 ] . les héritiers loosjes , haarlem , belgium .

lingual view of anterior tooth . the striations are a good characteristic for identification . 1 1 / 4 inch monmouth county , nj .

the striations on the lateral teeth are reduced , but present on most teeth . note the scalloped cusplets .

average tooth size is a about one inch but larger teeth are not uncommon . the tooth on the right is just shy of two inches

blades are the order of the day in the new jersey area . expect to find a significant amount of blades or damaged teeth , laterals seem to hold up a little better .

html public " - / / w3c / / dtd html 4 . 0 transitional / / en "

be the first to find out when we add items to this site ! join the lowcountry geologic mailing list .

the modern sand tiger shark reaches about 10 ft in length and about 600 lb in weight . the bar under each shark image indicates 4 ft in length .

it still retains the cusplets , but there is a growing difference between the front teeth ( right ) and the side teeth ( left ) .

notice how the cusps are still big on the side teeth , but are reduced to a simple sharp point on the front teeth . and here are the striations

about 35 million years ago . these teeth date to 40 million years ago , from the piney point formation in virginia .

to at least 35 million years ago . notice the front tooth ( on the right ) is now showing small barb - like cusplets , and the side tooth ( left )

is looking very similar . these teeth date to about 55 million years ago , and are from morocco .

which brings us to the current end - result : the modern sand tiger shark . they appear about 29 million years ago ,

but the cut - off dates become blurred as the teeth now look so similar to the prior shark . this is carcharias taurus , and as you can see the

front and side tooth look quite identical , and both show small sharp cusplets and have the recurve shape .

this is a side view of the same teeth to show the curve in the teeth . the inside of the mouth is to the image left .

this tooth style is ideal for grabbing and holding a small fish , and then just swallowing it whole . they seem a refinement of the

original old shark ` s teeth design . these are not teeth for biting out chunks of a victim . . .

html public " - / / w3c / / dtd xhtml 1 . 0 transitional / / en " " urltoken "

warning : the ncbi web site requires javascript to function . more . . .

format summary genbank genbank ( full ) fasta asn . 1 xml insdseq xml tinyseq xml feature table accession list gi list gff3

umbilicaria rossica isolate agred52 18s small subunit ribosomal rna gene , partial sequence ; internal transcribed spacer 1 , 5 . 8s ribosomal rna gene , and internal transcribed spacer 2 , complete sequence ; and 28s ribosomal rna gene , partial sequence

umbilicaria rossica isolate agred33 18s small subunit ribosomal rna gene , partial sequence ; internal transcribed spacer 1 , 5 . 8s ribosomal rna gene , and internal transcribed spacer 2 , complete sequence ; and 28s ribosomal rna gene , partial sequence

umbilicaria rossica isolate agred014 18s small subunit ribosomal rna gene , partial sequence ; internal transcribed spacer 1 , 5 . 8s ribosomal rna gene , and internal transcribed spacer 2 , complete sequence ; and 28s ribosomal rna gene , partial sequence

umbilicaria rossica isolate agred008 18s small subunit ribosomal rna gene , partial sequence ; internal transcribed spacer 1 , 5 . 8s ribosomal rna gene , and internal transcribed spacer 2 , complete sequence ; and 28s ribosomal rna gene , partial sequence

umbilicaria rossica isolate agred006 18s small subunit ribosomal rna gene , partial sequence ; internal transcribed spacer 1 , 5 . 8s ribosomal rna gene , and internal transcribed spacer 2 , complete sequence ; and 28s ribosomal rna gene , partial sequence

lasallia rossica internal transcribed spacer 1 , 5 . 8s ribosomal rna gene , and internal transcribed spacer 2 , complete sequence ; and 28s ribosomal rna gene , partial sequence

finds sub - sequence or patterns in the sequence and highlights the matching region . the tool works with standard single letter nucleotide or protein codes including ambiguities and can match prosite patterns in protein sequences . more . . .

we use cookies to enhance your experience on our website . by continuing to use our website , you are agreeing to our use of cookies . you can change your cookie settings at any time .

first report of costacoplumacollins and morris , 1975 ( decapoda : brachyura : retroplumidae ) from the eocene of alabama , u . s . a . | journal of crustacean biology | oxford academic

first report of costacopluma ( decapoda : brachyura : retroplumidae ) from the eocene of alabama , u . s . a .

( rmf , correspondence ) department of geology , kent state university , kent , ohio 44242 , u . s . a .

( rwp ) florida museum of natural history , p . o . box 117800 , university of florida , gainesville , florida 32611 - 7800 , u . s . a .

rodney m . feldmann , roger w . portell ; first report of costacopluma ( decapoda : brachyura : retroplumidae ) from the eocene of alabama , u . s . a . , journal of crustacean biology , volume 27 , issue 1 , 1 january 2007 , pages 90–96 , urltoken

nine specimens of retroplumid crabs collected from the late early or early middle eocene tallahatta formation in southern alabama form the basis for description of a new species , costacopluma grayi . the discovery confirms the extension of the range of the genus into the eocene and represents the first occurrence of costacopluma in the united states . as a result of the geologic range extension , the genus is now known to be contemporary with two other retroplumid genera , retrocypoda , and retropluma .

the retroplumidae gill , 1894 , is a relatively small family of decapod crustacea which was originally named for a single extant genus of deep - water crabs from the indo - pacific region . subsequently , another extant genus has been recognized and as many as five genera , known only from the fossil record , have been assigned to the family . the stratigraphic range has been extended into the late cretaceous as a result . the family has been re - evaluated by de saint laurent ( 1989 ) , vega and feldmann ( 1992 ) , and schweitzer and feldmann ( 2001 ) , and the position of the extinct genera has been debated .

: 50 - 51 ) . these shallow - marine sediments are part of a transgression - related condensed section in the lower tallahatta formation of eocene age .

location map showing the costacopluma collecting area along the southeastern bank of the conecuh river behind the point a dam in covington county , alabama .

the name “tallahatta” was first applied by w . h . dall ( 1898 : 344 ) as a replacement for the colloquial name “buhrstone . ” dall , at the suggestion of e . a . smith , renamed the formation after the tallahatta hills in choctaw county , alabama , which exhibited the lithologic character and fauna of the unit .

in alabama , the tallahatta formation is the basal member of the lower to middle eocene claiborne group which also consists ( in ascending order ) of the lisbon formation and gosport sand . typically , the tallahatta is disconformably underlain by either the lower eocene hatchetigbee or bashi formations ( upper wilcox group ) . elsewhere in the eastern gulf coast plain , the tallahatta formation has been mapped in mississippi and georgia . in alabama , the unit typically consists of massive siliceous claystone ( buhrstone ) interbedded with layers of glauconitic sand and sandy clay . according to

: 50 - 51 ) consists of very coarse to medium grained quartz sand with glauconite , clay clasts , and fine clay particles . here , evidence of bioturbation is abundant throughout the entire stratigraphic section . the bluish - green sediments of bed 5 , contain a diverse assemblage of fossils , consisting mainly of remains of sharks and other fishes . teeth of shark and ray species collected in situ with

archaeopus rathbun , 1908 ; bathypluma de saint laurent , 1989 ; costacopluma collins and morris , 1975 ; cristipluma bishop 1983 ; loerenthopluma beschin , busulini , de angeli , and tessier , 1996 ; retrocypoda via boada , 1959 ; retropluma gill , 1894 .

archaeopus originated in the cretaceous and is the oldest of the genera assigned to the family . today , the family is represented in the indo - pacific region by retropluma and bathypluma .

costacopluma concava collins and morris , 1975 , p . 823 , pl . 97 , figs . 1–9 , by original designation .

costacopluma australis feldmann , casadío , chirino - gálvez , and aguirre - urreta , 1995 ; c . bifida collins , higgs , and cortitula , 1994 ; c . binodosa collins and rasmussen , 1992 ; c . bishopi vega and feldmann , 1992 ; c . concava collins and morris , 1975 ; c . mexicana vega and perrilliat , 1989 ; c . nordestina feldmann and martins - neto , 1995 ; c . salamanca feldmann , rodriguez , martinez , and aguirre - urreta , 1997 ; c . senegalensis ( rémy in gorodiski and rémy , 1959 ) , as archaeopus .

small , rectangular to ovoid carapace ; wider than long ; with distinctly flattened surface crossed by three elevated ridges , the anteriormost complete and the medial and posterior ridges converging mesially to define depressed , triangular mesobranchial region . carapace flanks distinct , nearly perpendicular to dorsal surface . sternal plates well defined ; sternites 4 - 7 each with prominent transverse , beaded ridge . transverse ridges also present on abdominal somites .

new species . a , dorsal view of holotype , uf 113749 . b , oblique frontal view of paratype , uf 113748 , showing the downturned , rimmed , pustulose rostrum ( arrow ) , rectangular orbit , and steep lateral flanks . c and d , dorsal and ventral views of paratype , uf 115672 , illustrated at the same scale and showing well developed carapace regions and , on the ventral surface , pterygostomial regions and left mxp3 . e , dorsal view of worn and exfoliated paratype , uf 113750 , on which the carapace ornamentation is not in evidence . f , dorsal view of

rémy , 1959 , holotype , ro 3785 , deposited in the muséum national d’histoire naturelle , paris . scale

. b , enlargement of part of mesogastric and epibranchial regions showing the different sculpture exhibited on surface of exocuticle and endocuticle . scale

. c , ventral view of paratype , uf 115794 , showing nearly complete sternum and male abdomen . scale

. d , ventral view of paratype , uf 115796 , showing posterior part of sternum and proximal segments of male abdomen . scale

typical costacopluma with strongly downturned , triangular rostrum with beaded surface ; sharp , distinctly beaded lateral margin ; nearly circular posterior element of mesogastric region .

carapace small to moderate size for genus ; subovate , wider than long , length about 80 percent maximum width measured at midpoint of mesobranchial region ; flattened transversely and longitudinally ; lateral flanks distinct , downturned at right angles to carapace surface ; three transverse ridges of which anteriormost is complete ; posterior and medial ridges converge in advance of cardiac region .

front narrow , about 17 percent maximum carapace width ; rostrum strongly downturned , triangular , axially sulcate ; surface pustulose . orbits deep , concave , forward - directed , rectangular when viewed from front ; bounded dorsally by distinct , beaded rim . fronto - orbital margin about 65 percent maximum width . anterolateral margin straight to weakly convex with distinct beaded rim extending from outer - orbital corner to merge with more convexly rounded , rimmed posterolateral margin . posterolateral corner and posterior margins poorly preserved , appearing to be gently convex .

epibranchial regions transversely elongate , swollen , separated by axial sulcus extending onto rostrum . protogastric and hepatic regions not distinguishable , depressed . mesogastric region circular , swollen , beaded , 27 percent maximum width , bounded laterally by deep , broad arcuate depressions . metagastric and urogastric region indistinct , narrow , with subtle longitudinal axial elevation . cardiac region transversely ovoid , 40 percent maximum width . urogastric region elongate - oval , inflated , separated from posterior margin by shallow depression . epibranchial and mesobranchial regions depressed .

transverse ridges distinct , elevated well above remainder of carapace . anterior - most ridge a sinusoidal curve , concave axially and convex laterally , surface beaded ; medial ridge straight , beaded , extending posteromesially to anterior end of cardiac region where it merges with concave forward , beaded , anteromesially - directed posterior ridge ; medial and posterior ridges define a triangular mesobranchial region .

surface of exocuticle very finely beaded , nearly smooth . surface of endocuticle with coarser , evenly spaced pustules .

buccal frame broad , widening anteriorly , bounded by inflated and beaded pterygostomial regions . ischium of third maxilliped generally rectangular , tapering slightly anteriorly ; surface smooth .

sternites 1 - 3 forming isosceles triangle , directed dorsally ; sutures 1 - 2 , 2 - 3 , and 3 - 4 fused but elevated and distinct in position . sternite 4 much wider than sternite 3 with prominent anterolaterally directed projection . lateral margin of sternite 4 elevated into rim . sutures between somites 4 - 5 , 5 - 6 , and 6 - 7 appear to be unfused laterally and obscurred by abdomen axially . somite 5 directed laterally ; somites 6 and 7 directed posterolaterally ; each with prominent nodose keel . somite 8 not exposed . axis of somites 2 - 7 deeply depressed , weakly rimmed on somite 4 . male abdomen apparently unfused throughout , each abdominal somite transversely keeled , telson triangular , extends onto depression in sternal somite 3 .

measurements , in millimeters , taken on specimens of costacopluma grayi are given in table 1 .

carapace measurements ( mm ) . key : maximum length ( l ) , maximum width ( w ) , fronto - orbital width ( w1 ) .

the trivial name honors mark m . gray , for bringing the first specimens to the attention of the authors and for generously donating his specimens to the florida museum of natural history ( flmnh ) .

the holotype , uf 113749 , and paratypes , uf 113748 , 113750 , 114747 , 115672 115793 , 115794 , 115795 , and 115796 , are deposited in the invertebrate paleontology division at the flmnh , university of florida , gainesville , florida .

the type series was collected from the upper lower to lower middle eocene tallahatta formation behind point a dam , sw1 / 4 , ne1 / 4 , sec . 35 , t5n , r15e , river falls quadrangle ,

costacopluma grayi conforms to the diagnostic features of the carapace of the genus in all regards . the ventral surface of the carapace is not preserved on six of the nine specimens and only the pterygostomial region and one of the maxillipeds is preserved on one specimen . two specimens exhibit parts of the sternum and abdomen of male specimens so it is possible to describe the morphology . the morphology of the sternum and abdomen , coupled with that of the dorsal carapace , makes placement within costacopluma and the retroplumidae certain .

a single specimen , uf 115793 , retains the exocuticule over much of the dorsal carapace ( fig . 3a , b ) . as is the case with many decapods , the sculpture exhibited by the exocuticle is different from that seen on the surface of the endocuticle , which is visible when the exocuticle is exfoliated . therefore it is important to note which surface that observations of sculpture are made , if possible . the contrast in ornamentation is even more striking when comparing the sculpture of the cuticular surfaces with that of the mold of the interior as seen on uf 113750 ( fig . 2e ) , which lacks ornamentation at a fine scale . these observations provide a note of caution when employing fine details of surface ornamentation in systematic work .

species within the genus are distinguished from one another on the basis of shape of the rostrum , development of granulation on the carapace elevations , carapace outline , and various dimensional ratios including length / width , posterior width / maximum width , and frontal width / maximum width . because the margins of the alabama specimens are not completely preserved , it is difficult to employ these ratios with certainty ; however , morphological features of the surface of the carapace are well exhibited so that detailed comparisons can be made . examination of types , or illustrations of types , of all other species within the genus confirms that the form of the carapace ridges , shape of the rostrum , and outline of the posterior element of the mesogastric region provide ample evidence that c . grayi is unique . all species within the genus , except c . concava , c . senegalensis , and c . grayi , possess ridges that are broadly rounded . the three exceptions have narrower , more sharply defined , granular ridges . among these three species , only c . senegalensis and the new species have downturned , triangular rostra . costacopluma concava , as with all other species of the genus for which the rostrum is known , bears a rostrum that is broadened or clearly bifid distally . all species of costacopluma , with the exception of c . senegalensis and c . grayi , have rostra that are substantially narrower proximally than it is distally . costacopluma grayi can be readily distinguished from c . senegalensis because the former exhibits a beaded rostral surface , a distinctly beaded lateral margin , and a nearly circular posterior element of the mesogastric region . the posterior part of the mesogastric region on c . senegalensis is transversely ovoid and lacks beaded ornamentation .

retroplumoidea ( crustacea , brachyura ) nel terziario del vicentino ( italia settentrionale ) .

fossil decapod crustacea from the late cretaceous coon creek formation , union county , mississippi .

the eocene tallahatta formation of alabama and georgia : its lithostratigraphy , biostratigraphy , and bearing on the age of the claibornian stage .

a new crab , costacopluma bifida ( crustacea , decapoda ) from the palaeocene of venezuela .

a table of the north american tertiary horizons , correlated with one another and with those of western europe , with annotations .

costacopluma nordestina n . sp . ( decapoda : retroplumidae ) from the maria farinha formation ( paleocene ) of brazil .

fossil decapod crustaceans from the jagüel and roca formations ( maastrichtian - danian ) of the neuquén basin , argentina .

costacopluma salamanca new species ( decapoda , retroplumidae ) from the salamanca formation ( danian ) of patagonia , argentina .

histoire naturelle , générale et particulière , des crustacés et des insectes . vol . 3 .

catalogue of the fossil reptilia and amphibia in the british museum ( natural history ) .

natural history notes from the r . i . m . s . “investigator” , series iii , no . 6 . an account of the new and some of the rarer decapod crustacea obtained during the surveying season , 1901 - 1904 .

la nouvelle superfamille des retroplumoidea gill , 1894 ( decapoda , brachyura ) : systématique , affinités , et évolution . pp . 103 - 179 .

occurrence of costacopluma ( decapoda : brachyura : retroplumidae ) in the maastrichtian of southern mexico and its paleobiogeographic implications .

una especie nueva de cangrejo del género costacopluma ( crustacea : decapoda : retroplumidae ) del maastrichtiano del estado de nuevo león .

cuticular structure in costacopluma mexicana vega and perrilliat , from the difunta group ( maastrichtian ) of northeastern mexico , and its paleoenvironmental implications .

fossil crabs ( crustacea : decapoda ) from the late cretaceous cárdenas formation , east - central mexico .

oxford university press is a department of the university of oxford . it furthers the university ' s objective of excellence in research , scholarship , and education by publishing worldwide

for full access to this pdf , sign in to an existing account , or purchase an annual subscription .

educalingo cookies are used to personalize ads and get web traffic statistics . we also share information about the use of the site with our social media , advertising and analytics partners .

is the study of the origin of words and their changes in structure and significance .

is a type of word the meaning of which determines reality . nouns provide the names for all things : people , objects , sensations , feelings , etc .

any shark of the family carcharhinidae , occurring mostly in tropical seas and characterized by a nictitating membrane and a heterocercal tail . the family includes the tiger shark and the soupfin .

from english to other languages presented in this section have been obtained through automatic statistical translation ; where the essential translation unit is the word «requiem shark» in english .

the map shown above gives the frequency of use of the term «requiem shark» in the different countries .

of the word «requiem shark» during the past 500 years . its implementation is based on analysing how often the term «requiem shark» appears in digitalised printed sources in english between the year 1500 and the present day .

and brief extracts from same to provide context of its use in english literature .

set in the 18th century ' s golden age of piracy , the requiem shark is the tale of a young recruit , william williams and his forced apprenticeship to bartholomew roberts , slaver turned pirate captain .

please note that the content of this book primarily consists of articles available from wikipedia or other free sources online .

four taxa , however , make up over 99 % of the identified specimens : requiem shark ( carcharhinidae ) , bat ray ( myliobatis californica ) , salmon ( oncorhynchus sp . ) , and sturgeon ( acipenser sp . ) . each of these species can reach substantial sizes , . . .

last & stevens ( 1994 ) reviews the australian requiem shark fauna , and compagno & niem ( 1998 ) provide an overview of indo - west pacific species . compagno ( 1999 ) is an updated list of living carcharhinid ( and chondrichthyan ) species . . .

. . . shark sand shark tope hammerhead shark mackerel shark porbeagle requiem shark school shark soupfin shark bull shark lemon shark bronze whaler sixgill shark frill shark nurse . . .

dutertref described a requiem shark which , according to the description and drawing of it given by him , must have been a carcharias . dutertre says that its young were fixed by means of a cord to a large membrane . cuvier | also says briefly in . . .

high quality content by wikipedia articles ! the silvertip shark ( carcharhinus albimarginatus ) is a large species of requiem shark , family carcharhinidae , with a fragmented distribution throughout the tropical indian and pacific oceans .

a young , great white shark had a near - death experience when the retreating tide . . . sharks that live close to shore , like the

it is estimated that 160 species of sharks are known to occur in india ' s commercial fishing zone .

, with a rounded nose , dark grey coloration , large teeth , and a curved dorsal fin .

that inhabits deep waters in the world ' s temperate and tropical oceans . preferring cooler waters ,

visitors watch the fish and sharks inside the shark tank at the loveland . . . the reef sharks and sandbar sharks are from the family of

as we brought it in closer we could see there were actually two sharks on the line . . . . renowend experts think it was made by a carcharhinus or

« educalingo . requiem shark [ online ] . available < urltoken > . jul 2018 » .

this email address is being protected from spambots . you need javascript enabled to view it .

final citation : cicimurri , david j . and ebersole , jun a . 2015 . two new species of pseudaetobatus cappetta , 1986 ( batoidei : myliobatidae ) from the southeastern united states . palaeontologia electronica 18 . 1 . 15a : 1 - 17 . urltoken urltoken

the hatchetigbee formation specimens discussed herein were all recovered from an upper unnamed member of the formation from three distinct localities in alabama and mississippi : site abu - 3 in butler county , al ; site awa - 1 in washington county , al ; and the whynot locality located in lauderdale county , ms ( figure 1 ; more detailed geographic information on these localities is provided in the systematic paleontology section ) .

in both states , the hatchetigbee comprises the upper unit of the upper paleocene - to - lower eocene wilcox group . in alabama , the thickness of the hatchetigbee ranges from 76 m in the western part of the state to 11 m in the east , and the formation is subdivided into two members , the lower bashi marl and an unnamed upper member ( mancini and tew , 1995 ; raymond et al . , 1988 ; toulmin , 1977 ) . in mississippi , the bashi sequence is more complete than in alabama and is considered a distinct formation from the hatchetigbee ( dockery et al . , 1994 ) .

both the hatchetigbee in mississippi and the corresponding unnamed upper member in alabama are composed of a series of non - marine and marine beds that consist of olive green fine - grained , micaceous and carbonaceous fossiliferous sand , with silty clay , silt , and sandy clay ( raymond et al . , 1988 ; toulmin , 1977 ) . bounded by unconformities both above and below , the hatchetigbee is interpreted to represent a tagc - 2 . 4 type 1 depositional sequence , and strata represent inner neritic deposits ( gibson , 1982 ; mancini and tew , 1990 ; 1995 ) .

the tallahatta formation specimens presented in this study , which includes the type material of a new pseudaetobatus species , were collected from site adl - 1 in dale county , southeast alabama ( figure 1 ; more detailed geographic information is provided in the systematic paleontology section ) . in alabama , the tallahatta formation represents the basal unit of the lower - to - middle eocene claiborne group . in ascending order , this group is composed of the tallahatta , lisbon , and gosport sand formations ( raymond et al . , 1988 ) . the contact between the claiborne group and the underlying wilcox group represents a type 1 unconformity , which lies at the contact between the tallahatta and underlying hatchetigbee formation ( bybell and gibson , 1985 ; mancini and tew , 1991 ) .

the aerial extent of the tallahatta formation forms an arcuate belt that extends through northern and central mississippi , east to west across southern alabama , and into western georgia . in mississippi , the tallahatta formation is divided into three members : the meridian sand , basic city shale , and neshoba sand . although outcrops of the basal meridian sand are exposed in southwestern alabama ( bybell and gibson , 1985 ) , the tallahatta thins in the southeastern part of the state and is generally not differentiated into members ( raymond , 1988 ; savrda et al . , 2010 ) . bybell and gibson ( 1985 ) , however , described a unit at the base of the tallahatta in southeastern alabama that they referred to as “meridian sand equivalent . ”

in aiken county , the dry branch formation disconformably overlies kaolin deposits ( which are up to 12 . 5 m thick ) of the lower - to - middle eocene huber formation ( figure 3 ) . the contact between these two formations has been reported as a sequence boundary , and the base of the dry branch formation consists of a lag deposit ( harris et al . , 2002 ; schroeder et al . , 2002 ) that formed during the initial transgression of the jackson sea into the region ( huddleston , 1993 ; huddleston and hetrick , 1986 ) .

the dry branch formation is at least 28 m thick in aiken county , and twiggs clay , griffins landing sand ; and irwinton sand lithologies have been reported in the area ( mittwede , 1982 ; nystrom and willoughby , 1982 ; zullo and kite , 1985 ) . to the south of aiken in the jackson quadrangle , mittwede ( 1982 ) described mustard - yellow to orange - yellow loose , fine - to - medium grained quartz sand containing thin , discontinuous clay beds that he attributed to the irwinton sand , and he noted that the unit thickened markedly towards the aiken area . the dry branch formation deposit yielding pseudaetobatus teeth is consistent with the irwinton sand , and the stratum occurs approximately 2 . 0 m below the contact with the overlying tobacco road formation .

msc , mcwane science center , birmingham , alabama ; sc , south carolina state museum , columbia .

type species . pseudaetobatus casieri cappetta , 1986 , lower eocene ( ypresian ) phosphate deposits , sidi daoui , morocco .

etymology . species named for gorden l . bell , jr . in honor of his contributions to alabama vertebrate paleontology . gorden also collected the matrix sample from which this new species was recovered .

hypodigm . msc 35048 ( holotype ) , lower median tooth ( figure 4 . 1 ) ; msc 35054 ( paratype ) , lower right distal - most lateral tooth ( figure 4 . 2 ) ; msc 35058 ( paratype ) , right half of upper ? median tooth ( figure 4 . 5 ) , msc 35059 ( paratype ) , upper right distal - most lateral tooth ( figure 4 . 3 ) ; msc 35062 ( paratype ) , intermediate lateral tooth ( figure 4 . 4 ) . all tallahatta formation , site adl - 1 .

type locality . site adl - 1 , 31 . 317309 n , - 85 . 71598 w ( n 31° 19’ 02” lat . , w 85° 42’ 57” long . ) , dale county , alabama .

type horizon . lower tallahatta formation , just above the “meridian sand member equivalent” beds .

diagnosis . teeth are larger than those of the type species , pseudaetobatus casieri . as in p . casieri , upper median teeth fairly straight , whereas lower median teeth are arcuate , both have crowns with rectangular cross sections , angular distal ends and labial / lingual ornamentation of fine vertical ridges ; tooth base thick , with sharply oblique labial face and lingual lobes that extend distally past the crown foot . distal - most lateral teeth differ from p . casieri in being less wide , with shorter and less pronounced distal projection , but more sharply basally curved margin . intermediate lateral tooth morphology is six - sided and nearly symmetrical .

the holotype of pseudaetobatus belli sp . nov . , a lower median tooth , is 20 % larger than equivalent teeth of p . casieri illustrated by cappetta ( 1986 , plate 3 , figure 9 ) . in addition , the distal - most lateral teeth of p . belli sp . nov . are not as wide as those of p . casieri , the distal corner of the crown are not as elongated , but the distal edge is more tightly curled . the intermediate lateral tooth morphology represented by msc 35602 may not have been available to cappetta ( 1986 ) for his description of p . casieri , but it was likely present because it has been identified in another new species of pseudaetobatus from south carolina .

stratigraphic and geographic range . hatchetigbee formation ( np 11 ) of eastern mississippi and southern alabama ; tallahatta formation ( np 12 - 13 ) of southern alabama .

hypodigm . sc 2013 . 38 . 91 ( holotype ) , upper right distal - most lateral tooth ( figure 6 . 1 ) ; sc 2013 . 38 . 84 ( paratype ) , lower median tooth ( figure 6 . 6 ) ; sc 2013 . 38 . 86 ( paratype ) , upper median tooth ( figure 6 . 8 ) ; sc 2001 . 1 . 5 . 1 ( paratype ) , lower right distal - most lateral tooth ( figure 6 . 5 ) ; sc 2001 . 1 . 5 . 2 ( paratype ) , intermediate lateral tooth ( figure 6 . 4 ) .

type locality . 33 . 504444 n , - 84 . 742778 w ( n 33° 30 ' 16 " lat . , w 81° 44 ' 34 " long . ) , aiken , aiken county , south carolina .

type horizon . dry branch formation , approximately 2 . 0 m below contact with overlying tobacco road formation .

additional material . abandoned clay pit north of the aiken city limit , 33 . 624867 n , - 81 . 681713 w ; aiken county , sc ; dry branch formation - sc 96 . 97 . 52 , 22 median tooth fragments ; sc 96 . 97 . 53 , lateral tooth . type locality - sc 2001 . 1 . 3 , 110 partial median teeth ; sc 2001 . 1 . 4 . 1 , upper left distal - most lateral tooth ( figure 6 . 2 ) ; sc 2001 . 1 . 4 . 2 , lower right distal - most lateral tooth ; sc 2001 . 1 . 4 . 3 , lateral tooth ; sc 2001 . 1 . 5 . 3 , 8 lateral teeth ; sc 2001 . 1 . 6 , lateral tooth ; sc 2001 . 1 . 7 , lower left distal - most lateral tooth ; sc 2013 . 38 . 85 , upper median tooth ( figure 6 . 10 ) ; sc 2013 . 38 . 87 , upper median tooth ; sc 2013 . 38 . 88 , incomplete upper median tooth ; sc 2013 . 38 . 89 , incomplete upper median tooth ; sc 2013 . 38 . 90 , lower median tooth ( figure 6 . 7 ) ; sc 2013 . 38 . 92 , lower right distal - most lateral tooth ; sc 2013 . 38 . 93 . 1 , upper right distal - most lateral tooth ; sc 2013 . 38 . 93 . 2 , upper left distal - most lateral tooth ; sc 2013 . 38 . 93 . 3 , lower left distal - most lateral tooth ( figure 6 . 3 ) ; sc 2013 . 38 . 93 . 4 , two upper distal - most lateral teeth ; sc 2013 . 38 . 94 , 12 lower distal - most lateral teeth ; sc 2013 . 38 . 95 , intermediate lateral tooth ; sc 2013 . 38 . 96 . 1 , intermediate lateral tooth ; sc 2013 . 38 . 96 . 2 , intermediate lateral tooth ; sc 2013 . 38 . 96 . 3 , three intermediate lateral teeth ; sc 2013 . 38 . 97 , 88 partial median teeth .

diagnosis . upper and lower tooth morphologies as in pseudaetobatus casieri and p . belli sp . nov . , but smaller in overall size than the latter species . the distal - most lateral teeth are the most diagnostic , and these are distinguished from equivalent teeth of all other pseudaetobatus species by their sinuous outline in labial / lingual view .

remarks . the median teeth of pseudaetobatus undulatus sp . nov . differ from those of p . belli sp . nov . in that they are smaller in overall size ( sc 2013 . 38 . 84 is only half as large as the p . belli holotype ) , the lingual crown face is more vertical , and the groove above the lingual longitudinal ridge is more deeply impressed . the distal - most lateral teeth of p . undulatus sp . nov . , uppers and lowers , are easily distinguished from p . casieri and p . belli sp . nov . in having undulating crowns ( those of the latter two species are rather flat ) . in addition , the distal - most lateral teeth of p . casieri are wider and have more elongated disto - lingual projections than either p . belli sp . nov . or p . undulatus sp . nov . ( cappetta , 1986 ) .

stratigraphic and geographic range . the species is thus far only known from the upper eocene ( priabonian ) dry branch formation of aiken county , south carolina .

the distal ends of pseudaetobatus median teeth are angular and have straight edges ( i . e . , figure 4 . 1 and figure 5 . 7 - 8 ) , and these areas articulate with lateral teeth . in contrast , aetobatus has no lateral teeth , the distal ends of median teeth curl basally ( figure 7 . 8 ) , and the straight margin is perpendicular to the tooth width ( see also hovestadt and hovestadt - euler , 2013 ; purdy et al . , 2001 ) . in pseudaetobatus , these features are observed on the distal - most lateral teeth ( i . e . , figure 4 . 2 - 3 and figure 5 . 3 , 5 . 10 ) . it may be difficult to distinguish these two genera if the distal ends of a median tooth are missing ( figure 5 . 10 ; also compare figure 7 . 7 to figure 5 . 14 - 15 ) , and the presence of pseudaetobatus in other deposits could go unnoticed without the aid of more complete material and / or the recognition of lateral teeth .

albin , e . f . 1999 . regional stratigraphic correlation of north american tektites . lunar and planetary science , 30 : 1 - 2 .

albin , e . f . and wampler , j . m . 1996 . new potassium - argon ages for georgiaites and the upper eocene dry branch formation ( twiggs clay member ) : inferences about tektite stratigraphic occurrence . lunar and planetary science , 27 : 5 - 6 .

arambourg , c . 1952 . les vertébrés fossils des gisements de phosphates ( maroc - algérie - tunisie ) . notes et mémoires du service géologique du maroc , 92 : 1 - 372 .

baum , g . r . and vail , p . r . 1988 . sequence stratigraphic concepts applied to paleogene outcrops , gulf and atlantic basins , p . 309 - 327 . in wilgus , c . k . , hastings , b . s . , posamentier , h . , van wagoner , j . , ross , c . a . , and kendall , c . g . s . c . ( eds . ) , sea level change - an integrated approach . sepm ( society for sedimentary geology ) special publications 42 .

bonaparte , c . l . 1838 . selachorum tabula analytica . nuovi annali delle scienze naturali , 1 ( 2 ) : 195 - 214 .

bybell , l . m . and gibson , t . g . 1985 . the eocene tallahatta formation of alabama and georgia : its lithostratigraphy , biostratigraphy , and bearing on the age of the claibornian stage . united states geological survey bulletin , 1615 : 1 - 20 .

cappetta , h . 1986 . myliobatidae nouveaux ( neoselachii , batomorphii ) de l’yprésien des ouled abdoun , maroc . geologica et palaeontologica , 20 : 185 - 207 .

cappetta , h . 2012 . handbook of paleoichthyology , vol . 3e : chondrichthyes . mesozoic and cenozoic elasmobranchii : teeth . verlag dr . friedrich pfeil .

cappetta , h . and nolf , d . 1981 . les sélaciens de l’auversien de ronquerolles ( eocène supérieur du bassin de paris ) . mededelingen van den werkgroep voor tertiaire en kwartaire geologie , 18 ( 3 ) : 87 - 107 .

cappetta , h . and nolf , d . 2005 . révision de quelques odontaspididae ( neoselachii : lamniformes ) du paléocène et de l’eocène du bassin de la mer du nord . bulletin de l’institut des sciences naturelles de belgique , science de la terre , 75 : 237 - 266 .

carter , b . d . 1987 . paleogene echinoid distributions in the atlantic and gulf coastal plains . palaios , 2 : 390 - 404 .

case , g . r . and cappetta , h . 1990 . the eocene selachian fauna from the fayum depression in egypt . palaeontographica abteilung a , 212 : 1 - 30 .

case , g . r . , udovichenko , n . i . , nessov , l . a . , averianov , a . o . , and borodin , p . d . 1996 . a middle eocene selachian fauna from the white mountain formation of the kizylkum desert , uzbekistan , c . i . s . palaeontographica , abteilung a , 242 ( 4 - 6 ) : 99 - 126 .

casier , e . 1946 . la faune ichthyologique de l’ypresien de la belgique . memoires de musée royal d’histoire naturelle de belgique 104 .

casier , e . 1966 . faune ichthyologique du london clay . british museum ( natural history ) , london , england .

clayton a . a . , ciampagalio , c . n . , and cicimurri , d . j . 2013 . an inquiry into the stratigraphic occurrence of a claibornian ( eocene ) vertebrate fauna from covington county , alabama . bulletin of the alabama museum of natural history , 31 ( 2 ) : 60 - 73 .

compagno , l . j . v . 1973 . interrelationships of living elasmobranchs . zoological journal of the linnean society , 53 ( 1 ) : 15 - 61 .

dames , w . 1883 . über eine tertiäre wirbelthierfauna von der westlichen insel des birket - el - qurūn im fajum ( aegypten ) . sitzungsberichte der königlich preussischen akademie der wissenschaften zu berlin , 6 : 129 - 153 .

darteville , e . and casier , e . 1959 . les poissons fossils du bas - congo et des regions voisines ( part iii ) . annales du musée congo belge , minéralogique géologique , paléontologique , 3 ( 2 ) : 257 - 568 .

dockery , d . t . , iii , copeland , c . w . , jr . , and huddlestun , p . f . 1994 . reply to a revision of the hatchetigbee and bashi formations . mississippi geology , 4 ( 3 ) : 11 - 15 .

duthiel , d . b . 1991 . a checklist of neoselachii ( pisces , chondrichthyes ) from the palaeogene of the paris basin , france . tertiary research , 13 ( 1 ) : 27 - 36 .

edwards , l . e . , bybell , l . m . , gohn , g . s . , and frederiksen , n . o . 1997 . paleontology and physical stratigraphy of the usgs - pregnall no . 1 core ( dor - 208 ) , dorchester county , south carolina . united states geological survey open - file report 97 - 145 .

edwards , l . e . , gohn , g . s . , bybell , l . m . , chirico , p . g . , christopher , r . a . , frederiksen , n . o . , prowell , d . c . , self - trail , j . m . , and weems , r . w . 2000 . supplement to the preliminary stratigraphic database for subsurface sediments of dorchester county , south carolina . united states geological survey open - file report 00 - 049 - b .

eversull , l . g . 2005 . the twiggs clay : mineralogy , origin , and industrial properties of an upper eocene opaline claystone in the coastal plain province of georgia , u . s . unpublished phd dissertation , louisiana state university , baton rouge , louisiana .

falls , w . f . and prowell , d . c . 2001 . stratigraphy and depositional environments of sediments from five cores from screven and burke counties , georgia . usgs professional paper 1603 - a .

gibson , t . g . 1982 . revision of the hatchetigbee and bashi formations ( lower eocene ) in the eastern gulf coastal plain . united states geological survey bulletin , 1529 - h : h33 - h41 .

gradstein , f . , ogg , j . , and smith , a . 2004 . a geologic time scale . cambridge university press , massachusetts .

harris , r . s . , duncan , m . s . , holland , s . m . , roden , m . f . , and schroeder , p . a . 2002 . probable shocked quartz as evidence for an upper eocene impact horizon in coastal plain strata , warren county , georgia , u . s . a . geological society of america , abstracts with programs , 2002 annual meeting , paper 178 - 9 .

herman , j . , hovestadt - euler , m . , hovestadt , d . c . , and stehman , m . 2000 . contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites in living supraspecific taxa of chondrichthyan fishes . part b : batomorphii 4c : order rajiformes - suborder myliobatoidei - superfamily dasyatoidea - family dasyatidae - subfamily dasyatinae - genus : urobatis , subfamily potamotrygoninae - genus : paratrygon , superfamiliy plesiobatoidea - family plesiobatidae - genus : plesiobatis , superfamily myliobatoidea - family myliobatidae - subfamily myliobatinae - genera : aetobatus , aetomylaeus , myliobatis and pteromylaeus , subfamily rhinopterinae - genus : rhinoptera and subfamily mobulinae - genera : manta and mobula . addendum 1 to 4a : erratum to genus pteroplatytrygon . bulletin de l’institut royal des sciences naturelles de belgique , 70 : 5 - 67 .