goniatites bohemicus is a species of extinct cephalopods belonging to the family goniatitidae , included in the superfamily goniatitaceae .

these fast-moving nektonic carnivores lived in the silurian period , from 443.4 to 419.2 million years ago .

goniatites bohemicus

goniatites is a genus of extinct cephalopods belonging to the family goniatitidae , included in the superfamily goniatitaceae . beyrichoceras and cravenoceras are among related genera .
this lithograph of goniatites bohemicus appeared in joachim barrande’s systême silurien du centre de la bohême . 1 the artist , m . humbert from lemercier in paris , prepared the solnhofen lithographic limestone plate to barrande’s exacting requirements using fossils from his extensive collection ( as shown ) . it is one of more than a thousand similarly executed plates in the monumental systême .
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joachim barrande , the son of the shopkeeper augustin barrande and his wife charlotte , was born on 11 aug 1799 in saugues , haute loire . 2 he studied civil engineering at the école polytechnique in paris , graduated at the top of his class and in 1826 he came into the service of king charles x as the science tutor of his grandson henry , the comte du chambord . after charles x was forced to abdicate in the july revolution of 1830 barrende followed the royal court to dorset , edinburgh and finally prague . 1
charles x and his family eventually moved on to trieste but barrande stayed in prague where he began work as a road and bridge engineer . barrande had a keen interest in the natural sciences and the story goes that his interest in paleontology was piqued while observing fossils found during his surveying work on the planned radnice - plzeň - budějovice railway . whatever the reasons he started collecting fossils , however , it was the publication of murchison’s silurian system in 1839 that focused his interest . 3 he realized that his finds were the same ones that murchison was describing and soon he began a much more systematic study .
phragmoceras sp . ( left ) , trochoceras and gyroceras sp . ( right )
between 1840–50 barrande travelled on foot through the mountains between prague and beroun and created a stratigraphic map of central bohemia . he identified paleozoic rock formations and hired workers – as many as 20 or 30 at a time – to dig for fossils . he was so successful in finding fossil beds that his laborers “attributed to him a mastery of the black art of divination and a possible intimacy with the devil himself . ” by the time he died he had amassed a collection of more than 100 , 000 specimens .
he began organizing his notes and in 1852 , under the fitting motto of “c’est ce que j’ai vu” ( this is what i have observed ) , published the first folio of his remarkable systême silurien du centre de la bohême .
over the next 31 years – until his death in 1883 – he published 22 quarto folios covering trilobites , cephalopods , pteropods , brachiopods and molluscs . in all the systême included 6887 pages of text , 1078 plates and described 3560 lower palaeozoic species . it was a monumental work that simply has no parallel in the history of palentology .
monomercella and orthonychia sp . ( left ) , herecynella bohemica and philhedra humillima ( right )
barrande found it necessary to act as his own publisher and set up his own press in prague . there was no part of the project that escaped his obsessive supervision – especially the plates . he personally designed and laid out the illustrations and left extensive instructions to his artists including j . fetters ( prague ) , a . swoboda ( vienna ) and m . humbert ( paris ) . he routinely demanded revisions and even discarded perfectly usable plates at the last minute when a better specimen became available . as science noted in their 1888 obituary “ [ they ] could not have been printed with greater technical elegance by any press in paris . ” 4
of course barrande’s obsessiveness was expensive . the project required continual financial support from the academy of science in vienna and the comte du chambord . even with a prohibitive price of 1575 francs the publication never broke even .
barrande’s publication would turn out to be critically important to charles darwin . he mentioned his work no less than five times in the origin of the species . 5 barrande , on the other hand , felt that his work was “to ascertain the reality , not to create ephemeral theories” and his idea of “colonies” would become one of the chief scientific alternatives to darwin’s theory of evolution .
barrande had amassed so much material that after his death in 1883 three volumes describing an additional 1000 species were published . today , more than a century later , his work is still routinely referenced by paleontologists .
1 . barrande , joachim . systême silurien du centre de la bohême ( the silurian system of central boehmia ) ( 8 volumes ) . prague : chez l’auteur , 1852–1911 . volumes 2–8 are online . the missing volume , unfortunately , is the trilobite volume – hence no trilobite scans in the post . sorry . if a version of volume 1 ever becomes available – trust me – i’ll be all over it .
2 . for a more complete biography of barrande see radvan horný , radvan and turek vojtěch . jaochim barrande : his life , work and heritage to world palaeontology . prague : praha národní muzeum , přírodovědecké muzeum , 1999 . ( worldcat , online ) , or kříž , jiří . joachim barrande . prague : český geologický ústav , 1999 ( online )
3 . murchisen , roderick . the silurian system . london : john murray , 1839 ( online ) .
4 . joachim barrande , his life . science . 30 nov 1883 ; 2 ( 43 ) : 699–701 ( online )
5 . darwin , charles . on the origin of the species . london : john murray , 1859 ( worldcat , online ) .
the shell is generally globose with an open but narrow umbilicus , the surface commonly reticulate resulting from longitudinal lirae crossing transverse striae . the ventral lobe of the suture is rather narrow with a median saddle about or little less than half the height of entire lobe . the first lateral saddle is subangular to angular .
fossils of species within this genus have been found widespread in north america , eurasia , and north africa . in particular they have been found in the
miller , furnish , and schindewolf , 1957 . paleozoic ammonoidea , treatise on invertebrate paleontology , part l . geological society of america .
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on andrew wyeth’s 22nd birthday he ventured to cushing , maine to meet the artist merle james but instead met james’ 17 - year old daughter betsy . instantly smitten , he asked her to show him around town and she was more than happy to oblige . she thought “i’ll show him a real maine building” and as something of a test took him to the hathorne point home of her friends christina and alvaro olson . 1
throughout his life andrew had a rather contentious relationship with women ; indeed with anyone who didn’t in some way directly support his painting , but on that day in july 1939 he met what would become two of the most important women in his life . 1
like andrew , betsy had health problems as a child and grew up a loner . “i was the flawed child” she said . they married in may 1940 and she was initially dismissed by andrew’s domineering father , n . c . , as a lightweight dillitente . she would prove n . c . and the rest of wyeth’s family wrong , becoming not only andrew’s wife and model but his staunchest supporter and most severe critic , his business manager and eventually his archivist .
betsy knew that the olsen’s were difficult , reclusive people . andrew , however , was completely unfazed by their dilapidated house and as he later recalled “owing to christina‚äôs real fondness for betsy , she accepted me much more readily than i suppose she ordinarily would have . ”
by this time christina , who had an undiagnosed neuromuscular disease ( likely polio ) was reduced to crawling and urinating on stacks of discarded newspapers . andrew however felt that “she was so much bigger than all the little idiosyncrasies . ” and found her a symbol of fierce independance - an extraordinary conquest of life . the result of this friendship was christina’s world , one of the iconic paintings of the 20th century .
andrew and betsy spent their winters in chadds ford , pennsylvania and their summers in cushing where he routinely returned to the olsons and their saltwater farm as a subject . he eventually had free run of their house , even setting up his studio in an upstairs bedroom .
christina’s death in jan 1968 deeply affected andrew and marked the end of a seminal two decade long period in his painting . faced with a blank canvas – as it were – it was time for a reappraisal of his art . it was then that he met siri , the daughter of the cushing farmer george erickson . siri was exotic , untouched and had an electrifying effect on his work . “a burst of life , ” he later said , “like spring coming through the ground , a rebirth of something fresh out of death . ”
it was betsy ( much to her later chagrin ) who pushed him to do nudes . because his model was only 14 years old it became – as you could imagine – something of a scandal .
wyeth painted siri for ten years , until betsy – worried that their relationship had turned sexual – put a stop to it . she told andrew “if you do this again , don’t tell me . ” her request would have rather far - reaching consequences because andrew had just met helga .
helga was born in konigsberg , prussia in 1939 . after surviving the war her family was held as prisoners in denmark . at age 16 she entered the convent but left a few years later and met john testorf . they married , moved to philadelphia and finally settled in chadds ford . by the time she met andrew she was a lonely , unhappy , homesick housewife and posing for andrew was her answer . “i became alive , ” she said , “it shows in the pictures . i became young overnight . i’ve never done anything more worthwhile . ”
wyeth spent the next 15 years painting helga in near complete secrecy . to hide her from betsy he shuttled her from studio to studio ( “like a hamster in a cage” ) and began finding excuses to remain in chadds ford while betsy was in maine . what few pieces he showed betsy carefully concealed helga’s identity . he even changed her skin color in barracoon :
eventually he tired of the subterfuge and showed his work – some 240 pieces – to betsy . her first reaction was that “ whew , i was absolutely overcome , ” and her next was “who the fuck was this woman ? ” the wyeths sold the entire collection to the publisher leonard e . b . andrews and in a masterful piece of public relations the story spread beyond art circles , with the salacious details of wyeth’s secret model becoming the simultaneous cover story in both time and newsweek .
wyeth’s artwork remains to this day a polarizing subject – either you love it or you hate it . it would be difficult , however , to deny his love of his models . as one of them stated “its a romance to to the degree where you’re the most important thing in the world to him . ”
1 . for a complete biography see : meryman , richard . andrew wyeth : a secret life . new york : harpercollins , 1996 ( worldcat ) .
2 . all of the helga images are from wilmerding , john . andrew wyeth : the helga pictures . new york : harry abrams , 1987 ( worldcat ) .


macroscelides is a genus of small shrew-like animals , the round-eared sengis ( also called elephant shrews ) , found in western namibia and in south africa ; they are members of the clade afrotheria .

there are three known species : namib round-eared sengi , macroscelides flavicaudatus etendaka round-eared sengi , macroscelides micus , which is only found in gravel plains in the etendaka formation of north-west namibia karoo round-eared sengi , or short-eared elephant shrew , macroscelides proboscideus

macroscelides

macroscelides typus a . smith , 1829 ( = sorex proboscideus shaw , 1800 ) .
species account of macroscelides proboscideus on the animal diversity web ( the university of michigan museum of zoology ) .
namib round - eared sengi ( macroscelides flavicaudatus ) from wlotzkasbaken , namibia . may 2000 . photo : g . rathbun
unger , r . , h . kratochvil . 1999 . feeding preferences of short - eared elephant shrews ( macroscelides proboscideus , smith 1829 ) .
to cite this page : dohring , a . 2002 .
macroscelides proboscideus
( on - line ) , animal diversity web . accessed july 09 , 2018 at urltoken
to cite this page : dr . ted macrini , 2004 ,
macroscelides proboscideus
( on - line ) , digital morphology . accessed july 9 , 2018 at urltoken
macroscelides proboscideus , the short - eared elephant shrew , is one of 15 extant species of the clade macroscelidea . elephant shrews are small mammals ( head and body length : 95 - 315 mm ) characterized by long slender tails ( length : 80 - 265 mm ) , long slender legs , and a long , narrow , semi - flexible snout from which the name derives ( nowak , 1991 ) .
we investigated the state of dental eruption in specimens of macroscelides proboscideus and erinaceus europaeus of known age . when m . proboscideus reaches adult size and sexual maturity , few or none of its replaced permanent cheek teeth have erupted . the approximate sequence of upper tooth eruption is p1 , [ i3 , c , m1 ] , [ i1–2 ] , m2 , p4 , [ p2 , p3 ] . chronologically , e . europaeus erupts its molars and most premolars prior to m . proboscideus ; but its first two upper incisors erupt after those of m . proboscideus , and its canines erupt around the same time . the approximate sequence of upper tooth eruption in e . europaeus is [ m1 , m2 , p2 , i3 ] , c , m3 , p4 , p3 , i2 , i1 . unlike m . proboscideus , e . europaeus does not reach adult size until all permanent teeth except for the anterior incisors have erupted . while not unique among mammals , the attainment of adult body size prior to complete eruption of the permanent cheek teeth is particularly common among macroscelidids and other afrotherians .
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click on images for enlargement . click here to see the gallery of photos for the newly discovered gray faced sengi .
golden - rumped sengi rhynchocyon chrysopygus weight = 550 g . gedi ruins , kenya photo : galen rathbun
black and rufous sengi rhynchocyon petersi minja forest reserve , north pare mountains , tanzania . 28 july 2006 photo : bill stanley
checkered sengi rhynchocyon cirnei ( camera trap image ) burigi game reserve , kagera , tanzania . june 2007 photo : charles foley , tanzania mammal atlas project
checkered sengi rhynchocyon cirnei ( camera trap of likely estrus female followed by male ) burigi game reserve , kagera , tanzania . june 2007 photo : charles foley , tanzania mammal atlas project
checkered sengi rhynchocyon cirnei mareja community reserve , pemba , cabo delgado province , mozambique . 16 june 2011 . photo : galen rathbun
short - snouted sengi elephantulus brachyrhynchus weight : 50 g . mankwe wildlife reserve , northwest province , south africa photo : richard yarnell
short - snouted sengi elephantulus brachyrhynchus weight : 45 g . zambezi province ( caprivi strip ) , namibia photo : galen rathbun
cape rock sengi elephantulus edwardii weight = 50 g . traveller’s rest , clanwilliam , northern cape province , south africa photo : galen rathbun
karoo rock sengi elephantulus pilicaudus weight = 50 g . loxton , northern cape province , south africa photo : galen rathbun
bushveld sengi elephantulus intufi weight = 40g . zwartmodder farm , maltahohe district , namibia photo : jack dumbacher
bushveld sengi elephantulus intufi weight = 40g . windpoort farm , outjo , namibia photo : galen rathbun
eastern rock sengi elephantulus myurus weight = 50 g . matopos national park , zimbabwe , august 1982 photo : tim osborne
eastern rock sengi elephantulus myurus weight = 50 g . weenen game reserve , kwazulu - natal province , south africa photo : david ribble
north african sengi petrosaltator rozeti weight : 50 g midlet , morocco photo : g . rathbun
rufous sengi elephantulus rufescens mother and young resting on trail through leaf litter . kibwezi , kenya photo : galen rathbun
rufous sengi elephantulus rufescens neonate - one day old . photo : gene maliniak , national zoological park
western rock sengi elephantulus rupestris weight = 50 g . rehoboth , namibia photo : galen rathbun
western rock sengi elephantulus rupestris weight = 50 g . namibrand nature reserve , namibia photo : jack dumbacher
western rock sengi elephantulus rupestris weight = 50 g . captive eating a grasshopper . photo : chris and tilde stuart
the short - eared elephant shrew mostly inhabits namibia , southern botswana , and south africa .
the animal only lives in desert and semi - desert areas of the countries in which it is found . it hides in the sparse grass cover or bushes that are a part of these dry areas . they also burrow into the sand .
compared to members of the other elephant shrew genus , the short - eared elephant shrew has shorter and rounder ears and lacks the pale rings around the eyes that are typical of those animals . the tail is hairy , with a visible gland on the underside . on the hind feet , the first digit is small and has a claw . the fur is usually long , soft , and is an orange , brown or grayish color on top and a lighter color on the underside . adults often weigh between 40 - 50 grams with 100 - 110mm long bodies and 97 - 130mm long tails . defining skull features include an enlarged auditory bullae and the appearance of three upper incisors , as well as a short rostrum and crowded teeth . females also have six mammae .
the breeding season is in the warm , wet months of august and september . a female may have many pregnancies during one breeding season . ( shaw , 1934 )
gestation for these animals is typically about 56 days and only two young are born , sometimes one . they are born in a very precocial state ; they can run within a few hours after birth , are large in size , and are born with hair and their eyes open . babies are weaned at 16 - 25 days and reach sexual maturity after about 43 days . ( rathbun & fons )
the female does not make a nest for the young ; however , she will find a sheltered area and give birth to the young in it . the mother does not guard her young and is gone from the litter most of the time , coming back once a day to feed the young . ( smith , 1829 )
in the wild , these animals only live for 1 - 2 years . in captivity they can live as long as 3 - 4 years .
these animals are mostly diurnal except when threatened by predators . they are usually solitary animals in the wild except when they come together to mate . when mating , females fend off other females and males fight off other males .
these elephant shrews take refuge under bush or rocks but also dig burrows or use shelters previously built by other small species , typically rodents . the animals use the burrows like roads to get from place to place . they keep them clean by kicking any debris that clogs their tunnels . they also sand bathe to help keep clean .
short - eared elephant shrews typically eat insects , usually termites and ants , and other small invertebrates . they may also feed on plant parts such as roots , shoots , and berries .
the animal usually jumps from bush to bush during the day or basks in the sun , but if harassed by diurnal predators , such as hawks , it switches its schedule and looks for food at dusk , hiding in bushes during the day . also , by using their forelimbs these animals can dig tunnels very rapidly to quickly escape predators . few predators prey on the young because the young mature and leave the nest shortly after birth .
these elephant shrews help move soil around to create their burrows as well as recycle vacant burrows left from rodent species .
due to destruction of its habitat , this species is labeled “vulnerable” by the iucn .
alyce dohring ( author ) , university of michigan - ann arbor , kate teeter ( editor ) , university of michigan - ann arbor .
living in sub - saharan africa ( south of 30 degrees north ) and madagascar .
having body symmetry such that the animal can be divided in one plane into two mirror - image halves . animals with bilateral symmetry have dorsal and ventral sides , as well as anterior and posterior ends . synapomorphy of the bilateria .
in deserts low ( less than 30 cm per year ) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity . vegetation is typically sparse , though spectacular blooms may occur following rain . deserts can be cold or warm and daily temperates typically fluctuate . in dune areas vegetation is also sparse and conditions are dry . this is because sand does not hold water well so little is available to plants . in dunes near seas and oceans this is compounded by the influence of salt in the air and soil . salt limits the ability of plants to take up water through their roots .
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature . endothermy is a synapomorphy of the mammalia , although it may have arisen in a ( now extinct ) synapsid ancestor ; the fossil record does not distinguish these possibilities . convergent in birds .
the area in which the animal is naturally found , the region in which it is endemic .
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female .
lincoln park zoo ,
short - eared elephant shrew
( on - line ) . accessed october 4 , 2001 at urltoken .
regina , u .
short - eared elephant shrew
( on - line ) . accessed october 4 , 2001 at urltoken .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
corbet , g . b . and hanks , j . 1968 . a revision of the elephant - shrews , family macroscelididae . bulletin of the british museum of natural history ( zoology ) 16 : 1 - 111 .
a widespread species occurring in southern namibia , extreme southwestern botswana , and western south africa , particularly the northern , western , and eastern cape ( rathbun 2009 , dumbacher et al . 2012 ) .
it is believed that over much of its range its numbers are relatively low ( corbet and hanks 1968 , rathbun 2005 , smit et al . 2009 , schubert 2011 , dumbacher et al . 2012 , perrin and rathbun 2013 ) . there are no data on population dynamics of this species , but it is expected that populations will vary greatly in the arid habitats that it occupies , which is possibly correlated to climate regimes and climatic variability , and there are no reasons to believe that these variations , if they indeed occur , are abnormal .
there is no indication that this species has ever been used by people for any purposes . it rarely has been exported to various zoological gardens over the last few decades , where husbandry techniques have been developed , breeding has been achieved , and research results have been published ( olbricht 2009 ) .
there are no known major threats to the species . habitat modification to relatively small areas may occur near rivers and human population centres due to small - holder and industrial agriculture , mineral extraction , and urban development . changes in habitats due to desertification and bush encroachment and proposed wind and solar energy facilities may adversely alter habitats for sengis and displace them from such areas , but at present these changes do not appear widespread or serious , especially since the species is associated with arid habitats to begin with .
because this species is widespread , it is not in conflict with most human activities ; captive husbandry and breeding methods are well - established ; and it likely occurs in many protected areas ( i . e . , goegap nature reserve , gamkaberg nature reserve , skilpad nature reserve , tankwa karoo national park , mokala national park , ricthersveld national park , augrabies falls national park , namagua national park , and karoo national park ) , therefore there are no conservation actions recommended at present or in the foreseeable future . research topics of conservation relevance include determining the impacts of habitat shifts , including livestock grazing , on local populations , determining the proportion of the total distribution range that occurs in protected areas , and continue to accumulate information on occurrence points ( see urltoken ) .
rathbun , g . b . & smit - robinson , h . 2015 .
to make use of this information , please check the < terms of use > .
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all elephant shrews are endemic to africa . the short - eared elephant shrew is native to namibia , cape province of south africa , and southern botswana ( corbet and hanks , 1968 ; nowak , 1991 ) . the fossil record of macroscelidea also is restricted to africa and extends back to the eocene , but with only three pre - miocene taxa ( butler , 1995 ) .
recent analyses of molecular data place macroscelidea in the clade afrotheria , which also includes aardvarks , elephants , hyraxes , golden moles , tenrecs , and sirenians ( murphy et al . , 2001 ) . within afrotheria , elephant shrews form a clade with golden moles and tenrecs ( murphy et al . , 2001 ) . morphological data place macroscelidea as the sister taxon to glires ( rabbits + rodents ; novacek , 1992a , b ) .
macrosclides proboscideus inhabits sandy and gravel thornbrush plains and refuges in shallow burrows under bushes ( nowak , 1991 : 183 - 184 ) . the short - eared elephant shrew is mainly diurnal but sometimes crepuscular or nocturnal in activity , and it primarily feeds on insects along with roots , tender shoots , and berries ( nowak , 1991 : 184 ) . this species is mainly solitary in the wild ( nowak , 1991 ) . young are very precocial at birth , being covered with hair and able to move about soon after they are born ( nowak , 1991 ) . four species of elephant shrews are listed as either endangered or vulnerable by iucn , but m . proboscideus is not one of them .
this specimen ( amnh 161535 ) was collected from cape province , south africa by tom larson . it was made available to the university of texas high - resolution x - ray ct facility for scanning by ted macrini of the department of geological sciences , the university of texas at austin . funding for scanning was provided by a national science foundation ( nsf ) dissertation improvement grant to mr . macrini . funding for image processing was provided by a nsf digital libraries initiative grant to dr . timothy rowe of the department of geological sciences , the university of texas at austin .
the specimen was scanned by matthew colbert on 31 october 2003 along the coronal axis for a total of 630 slices , each slice 0 . 055 mm thick with an interslice spacing of 0 . 055 mm .
corbet , g . b . , and j . hanks . 1968 . a revision of the elephant - shrews , family macroscelididae . bulletin of the british museum ( natural history ) . zoology 16 : 47 - 111 .
murphy , w . j . , e . eizirik , s . j . o’brien , o . madsen , m . scally , c . j . douady , e . teeling , o . a . ryder , m . j . stanhope , w . w . de jong , and m . s . springer . 2001 . resolution of the early placental mammal radiation using bayesian phylogenetics . science 294 : 2348 - 2351 .
novacek , m . j . 1992a . fossils , topologies , missing data , and the higher level phylogeny of eutherian mammals . systematic biology 41 : 58 - 73 .
novacek , m . j . 1992b . mammalian phylogeny : shaking the tree . nature 356 : 121 - 125 .
nowak , r . m . 1991 . walker’s mammals of the world . volume 1 . fifth edition . the johns hopkins university press , baltimore .
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raw data on state of eruption and jaw length . “erupt / 10” indicates the number of completely erupted permanent cheek teeth with interlocking occlusion , taken as a proportion out of ten ( indicating presence of all normally erupted cheek teeth ) . in cases where the dentition was incomplete ( e . g . , dentary lost ) , the proportion was calculated out of the maximum possible number of interlocking permanent cheek teeth for that specimen and species . “symph - cond” indicates the distance in mm from the anterior margin of the mandibular symphysis to the posterior margin of the mandibular condyle ( asher and lehmann
: fig . 1 ) . “prop adult jaw” indicates a given specimen’s symphysis - condyle length divided by the median length obtained for all specimens of that species with the permanent cheek teeth completely erupted ( i . e . , with a “erupt / 10” value of ten ) , as indicated in the column “median jaw” . institutional abbreviations are as follows :
asher rj ( 2005 ) insectivoran grade placental mammals : character evolution and fossil history . in : rose kd , archibald d ( eds ) the rise of placental mammals : origin and relationships of the major clades . johns hopkins university press , baltimore , pp 50–70
asher rj , lehmann t ( 2008 ) dental eruption in afrotherian mammals . bmc biol 6 : 14 doi :
beatty b ( 2008 ) craniodental ontogeny in the desmostylia . j vertebr paleontol 283 : 49a
bronner g ( 1992 ) notes on the early postnatal development of a giant golden mole . koedoe 35 : 57–58
corbet gb , hanks j ( 1968 ) a revision of the elephant - shrews , family macroscelididae . bull brit mus nat hist zool 16 : 47–111
de magalhaes jp , costa j , toussaint o ( 2005 ) hagr : the human ageing genomic resources . nucleic acids res 33 : d537–d543 doi :
douady cj , catzeflis f , raman j , springer ms , stanhope mj ( 2003 ) the sahara as a vicariant agent , and the role of miocene climatic events , in the diversification of the mammalian order macroscelidea ( elephant shrews ) . proc natl acad sci usa 10014 : 8325 doi :
eisenberg jf , gould e ( 1970 ) the tenrecs : a study in mammalian behavior and evolution . smithson contrib zool 27 : 1–138
evans fg ( 1942 ) the osteology and relationships of the elephant shrews ( macroscelididae ) . bull am mus nat hist 80 : 85–125
holroyd p ( 2008 ) new data on dental eruption patterns in condylarths and afrotheres . j vertebr paleontol 283 : 93a
kellas l ( 1954 ) observations on the reproductive activity , measurement and growth - rate of the dik - dik . proc zool soc lond 124 : 751–784
laws r ( 1968 ) dentition and ageing in the hippopotamus . afr wildl j 6 : 19–52
leche w ( 1907 ) zur entwicklungsgeschichte des zahnsystems der säugetiere , zugleich ein beitrag zur stammengeschichte dieser tiergruppe . zoologica stuttg 49 : 1–157
morris pa ( 1970 ) a method for determining absolute age in the hedgehog . j zool ( lond ) 161 : 277–281
morris pa ( 1978 ) the use of teeth for estimating the age of wild mammals . in : butler pm , joysey ka ( eds ) development , function and evolution of teeth . academic , london , pp 483–494
olbricht g , kern c , vakhruscheva g ( 2006 ) einige aspekte der fortpflanzungsbiologie von kurzohr - ruesselspringern . zool gart 5–6 : 304–316
rathbun g ( 1979 ) the social structure and ecology of elephant - shrews . fortschr verhaltensforsch 20 : 1–76
rees j , kainer r , davis r ( 1966 ) chronology of mineralization and eruption of mandibular teeth in mule deer . j wildl manage 30 : 629–631 doi :
robinette w , jones d , rogers g , gashwiler j ( 1957 ) notes of tooth development and wear for rocky mountain mule deer . j wildl manage 21 : 134–153 doi :
sauer em ( 1973 ) zum socialverhalten der kurzohrigen elefantenspitzmaus . z saugetierkd 38 : 65–97
sauer fg , sauer em ( 1972 ) zur biologie der kurzohrigen elefantenspitzmaus . (
( afrosoricida : chrysochloridae ) from kwazulu - natal , south africa . afr zool 39 : 41–46
smith bh ( 1989 ) dental development as a measure of life history in primates . evolution 43 : 683–688 doi :
smith bh ( 2000 ) “schultz’s rule” and the evolution of tooth replacement patterns in primates and ungulates . in : teaford mf , smith mm , ferguson mwj ( eds ) development , function and evolution of teeth . cambridge university press , cambridge pp 212–227
springer ms , stanhope mj , madsen o , de jong ww ( 2004 ) molecules consolidate the placental mammal tree . trends ecol evol 19 : 430–438 doi :
tabuce r , asher rj , lehmann t ( 2008 ) afrotherian mammals : a review of current data . mammalia 72 : 2–14 doi :
van nievelt a , smith kk ( 2005 ) to replace or not to replace : the significance of reduced tooth replacement in marsupial and placental mammals . paleobiol 31 : 324–346 doi :
asher , r . j . & olbricht , g . j mammal evol ( 2009 ) 16 : 99 . urltoken


turneria bidentata is a species of ant in the genus turneria .

described by forel in 1895 , the species is endemic to australia , mostly found in the north ends of the country .

turneria bidentata

the above specimen data are provided by antweb . please see turneria bidentata for further details
turneria bidentata occurs in the top end of the northern territory and along the east australian coast from the cape york peninsula to extreme north - eastern new south wales . it is a twig - nesting species that is reasonably common although it is infrequently encountered because of its arboreal nesting habits . it is known to nest in a wide range of tree and shrub species .
shattuck , s . o . 1990 . revision of the dolichoderine ant genus turneria ( hymenoptera : formicidae ) . syst . entomol . 15 : 101 - 117 pdf
shattuck , s . o . ( 2011 ) turneria rosschinga sp . n . ( hymenoptera : formicidae ) , a new dolichoderine ant from australia . myrmecological news 15 , 125 - 128 .
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cephalic index ( head length / head width ) < 0 . 88 , relative eye length ( eye length / head width ) > 0 . 40 , frontal lobes without erect hairs , lateral areas of head moderately imbricate and with integument opaque , area between propodeal protuberances concave . in lateral profile , the concave region of the declivitous face of the propodeum is more rounded than in other species .
the following information is derived from barry bolton ' s new general catalogue , a catalogue of the world ' s ants .
lectotype ( designated by shattuck , 1990 ) , worker , mackay , queensland , australia , m . g . turner , musee d ' histoire naturelle genève ; ( lower specimen on pin ) .
paralectotype ( designated by shattuck , 1990 ) , 1 worker , mackay , queensland , australia , m . g . turner , musee d ' histoire naturelle genève .
the clypeus possesses 8 to 14 erect hairs . color varies from uniform dark brown or black to strongly bicolored with the head , alitrunk , legs and petiole yellow and the gaster brown . in lighter colored individuals , the dorsum of the head may be slightly darker than the mesosoma .
worker measurements ( n = 25 ) : ood 0 . 15 - 0 . 21 , el 0 . 24 - 0 . 28 , ocd 0 . 10 - 0 . 13 , cl 0 . 13 - 0 . 19 , hl 0 . 65 - 0 . 75 , les 0 . 03 - 0 . 05 , ew 0 . 11 - 0 . 15 , es 0 . 24 - 0 . 28 , hw 0 . 55 - 0 . 66 , sl 0 . 41 - 0 . 46 , pnl 0 . 30 - 0 . 42 , ml 0 . 27 - 0 . 37 , ppl 0 . 23 - 0 . 30 , pnw 0 . 36 - 0 . 43 , mw 0 . 23 - 0 . 30 , ppw 0 . 25 - 0 . 29 , po - 0 . 01 - 0 . 04 , ffl 0 . 41 - 0 . 53 , ffw 0 . 16 - 0 . 21 , mh 0 . 31 - 0 . 39 , pph 0 . 22 - 0 . 29 , ci 0 . 80 - 0 . 88 , oi 0 . 43 - 0 . 57 , rel 0 . 40 - 0 . 48 , si 0 . 70 - 0 . 79 , fi 0 . 34 - 0 . 48 , pi 0 . 63 - 0 . 71 , ppi 0 . 87 - 1 . 10 , rood 0 . 28 - 0 . 33 , poi - 0 . 04 - 0 . 17 , rpo - 0 . 02 - 0 . 07 , rmw 0 . 42 - 0 . 47 , rles 0 . 05 - 0 . 09 , res 0 . 41 - 0 . 45 , rffl 0 . 66 - 0 . 94 .
body color in this species is highly variable . populations from the vicinity of cairns are uniform dark brown , mackay - area populations are strongly bicolored , while southern collections from burleigh heads are yellowish brown with the gaster slightly darker ; in extreme cases some workers are yellow - brown with a black gaster . none of these populations diverge in any of other morphological traits , although the cairns - vicinity specimens average slightly , but insignificantly , smaller . for most traits all known specimens broadly overlap . the cairns - vicinity populations diverge slightly from the more southern ones in two metric traits , mw and pph , as follows : mw 0 . 23 - 0 . 27 vs . 0 . 25 - 0 . 30 and pph 0 . 22 - 0 . 25 vs . 0 . 23 - 0 . 29 . the broad overlap in the ranges of these characters make them of little value in separating these populations .
forel , a . 1895g . nouvelles fourmis d ' australie , récoltées à the ridge , mackay , queensland , par m . gilbert turner . ann . soc . entomol . belg . 39 : 417 - 428 ( page 419 , worker described )
this page was last modified on 6 july 2018 , at 19 : 14 .
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urn : lsid : biodiversity . org . au : afd . taxon : 02aa5edd - fad6 - 4d09 - 8b7e - c2800d7ad60b
urn : lsid : biodiversity . org . au : afd . taxon : 0c7269f4 - 32a6 - 41f6 - acef - e1196012a6f4
urn : lsid : biodiversity . org . au : afd . taxon : 0cdbe1fb - 9c3e - 423a - ba46 - ef8b1724b4c3
urn : lsid : biodiversity . org . au : afd . taxon : 3f65c802 - 976d - 48a7 - 9e7d - b969c738e786
urn : lsid : biodiversity . org . au : afd . taxon : adb80b81 - 29ed - 45e1 - b5f3 - e6087c6f8d33
urn : lsid : biodiversity . org . au : afd . taxon : adbcbc0f - 6923 - 413b - baf8 - 22c0f331d576
urn : lsid : biodiversity . org . au : afd . taxon : f11267f8 - 8878 - 4548 - b313 - 7e9087b01a4c
urn : lsid : biodiversity . org . au : afd . taxon : f2b5e6b6 - 7d35 - 4845 - 91c0 - 4a359ff5bf11
urn : lsid : biodiversity . org . au : afd . taxon : 40e82464 - 30c8 - 4921 - 94d9 - 847d0bf98c82
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a tiny orange ant seen running ( light trail ) along the branches of a shrub , in a suburban yard . they were entering and leaving a hole at the end of a branch , so seems to be a tree - nesting ant . i also accidentally photographed a jumping - spider that appears to be mimicking them ( the disguise worked ) .
this sighting hasn ' t been described yet ! be the first to describe this sighting .
: 419 - worker ; type locality : mackay [ 21 / 149 ] , queensland .
forel , a . 1895b . 1895 639 nouvelles fourmis d ' australie , recoltees a the ridge , mackay , queensland par m . gilbert turner . annales de la soci _ t _ entomologique de belgique 39 : 417 - 428 .
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urltoken


margaritiferidae is a family of medium-sized freshwater mussels , aquatic bivalve molluscs in the order unionoida .

they are known as freshwater pearl mussels , because the interior of the shell of these species has a thick layer of nacre or mother of pearl , and the mussels are thus capable of producing pearls .

margaritiferidae

( mollusca : pelecypoda : margaritiferidae ) . zool j linn soc 69 : 257–270
( conrad , 1838 ) ( unionacea : margaritiferidae ) . nautilus 102 : 159–163
conrad ( bivalvia : margaritiferidae ) . conserv genet . 2004 ; 5 : 271–278 .
( bivalvia , margaritiferidae ) [ in russian ] . zoologicheskii zhurnal . 1993 ; 72 : 29–36 .
( bivalvia : margaritiferidae ) from japan [ in japanese ] . venus . 2007 ; 65 : 355–363 .
( bivalvia : margaritiferidae ) in japan [ in japanese ] . venus . 2009 ; 67 : 189–197 .
the origin and phylogeny of margaritiferidae ( bivalvia : unionoida ) . a synthesis of molecular and fossil data .
( bivalvia : margaritiferidae ) in the shinano river system , japan . venus . 2005 ; 64 : 63–70 .
( spengler , 1782 ) ( mollusca : margaritiferidae ) based on museum specimens . journal of conchology 37 : 49–59 .
( bivalvia : margaritiferidae ) of japan , with description of a new species . venus . 2005 ; 64 : 135–140 .
hass 1910 im oberen und mittleren donausystem ( bivalvia : unionidae , margaritiferidae ) . nachrichtenbl ersten vorarlberger malakol ges 1 : 20–40
( bivalvia : margaritiferidae ) , with reference to the existence of two distinct species . venus . 2005 ; 64 : 55–62 .
( spengler , 1793 ) ( bivalvia : margaritiferidae ) découvertes dans le sud - ouest de la france . malaco 6 : 294–297 .
kat pw ( 1983 ) conchiolin layers among the unionidae and margaritiferidae ( bivalvia ) : microsculptural characteristics and taxonomic implications . malacologia 24 : 298–311
smith dg ( 1986 ) the stomach anatomy of some eastern north american margaritiferidae ( unionoida : unionacea ) . am malacol bull 4 : 13–19
( bivalvia : margaritiferidae ) in the chubu - nougu river , nagano prefecture [ in japanese ] . venus . 2008 ; 67 : 61–71 .
the origin and phylogeny of margaritiferidae ( bivalvia : unionoida ) . a synthesis of molecular and fossil data . | laboratory of kevin j . roe
bogatov vv , prozorova la , starobogatov yi . the family margaritiferidae ( mollusca : bivalvia ) in russia . ruthenica . 2003 ; 13 : 41–52 .
all the phylogenies inferred in this study support the reciprocal monophyly of both ( unionidae + margaritiferidae ) f - and m - type lineages ( fig .
smith dg ( 1976b ) the distribution of the margaritiferidae : a review and a new synthesis . am malacol union inc bull 1976 : 42 ( abstr )
klishko ok . sakhalin - kurile species of pearl mussels ( bivalvia : margaritiferidae ) from transbaikalye . the korean journal of malacology . 2009 ; 25 : 237–242 .
smith dg . observations on the morphology and anatomy of margaritinopsis dahurica ( middendorff , 1850 ) ( unionoida : margaritiferidae ) . j conchol . 2001 ; 37 : 119–125 .
starobogatov y ( 1995 ) the pearly freshwater mussels ( mollusca , unionoida , margaritiferidae of russia . in : valovirta i , harding pt , kirne d ( eds ) proc 9
bogatov , v . v . , prozorova , l . a . , and starobogatov , y . i . , . the family margaritiferidae ( mollusca : bivalvia ) in russia ,
, a new species of unionacea ( bivalvia : margaritiferidae ) from the mobile - alabama - coosa and escambia river systems , alabama . occasional papers on mollusks . 1983 ; 4 : 299–304 .
, a new species of unionacea ( bivalvia : margaritiferidae ) from the mobile - alabama - coosa and escambia river systems , alabama . occas pap mollusks mus comp zool harv univ 4 : 299–304
machordom a , araujo r , erpenbeck d , ramos m - a . phylogeography and conservation genetics of endangered european margaritiferidae ( bivalvia : unionoidea ) . biol j linn soc . 2003 ; 78 : 235–252 .
smith dg . systematics and distribution of the recent margaritiferidae . in : bauer g , wachtler k editors . ecology and evolution of the freshwater mussels unionoida . heidelberg : springer verlag ; 2001 . pp . 33–49 .
sp . n . ( bivalvia , margaritiferidae ) a new species of pearl mussels from transbaikalye , with remarks on the natural history of far eastern najades [ in russian ] . vestnik zoologii . 2008 ; 42 : 291–302 .
smith dg ( 1983 ) on the so - called mantle muscle scars on shells of the margaritiferidae ( mollusca , pelecypoda ) , with observations on mantle - shell attachment in the unionoida and trigonioida . zool scr 12 : 67–71
smith dg . on the so - called mantle muscle scars on shells of the margaritiferidae ( mollusca , pelecypoda ) , with observations on mantle - shell attachment in the unionoida and trigonioida . zool scr . 1983 ; 12 : 67–71 .
smith dg , wall wp ( 1984 ) the margaritiferidae reinstated : a reply to davis and fulller ( 1981 ) , genetic relationships among recent unionacea ( bivalvia ) of north america . occas pap mollusks mus comp zool harv univ 4 : 321–330
klishko ok . some data on reproductive biology of the freshwater mussels ( margaritiferidae , unionidae ) and their relationships with bitterlings ( cyprinidae ) in transbaikalye [ in russian ] . the bulletin of the russian far east malacological society . 2012 ; 15 / 16 : 31–55 .
smith d . g . ( 2001 ) systematics and distribution of the recent margaritiferidae . in : bauer g . , wächtler k . ( eds ) ecology and evolution of the freshwater mussels unionoida . ecological studies ( analysis and synthesis ) , vol 145 . springer , berlin , heidelberg
citation : bolotov in , bespalaya yv , vikhrev iv , aksenova ov , aspholm pe , gofarov my , et al . ( 2015 ) taxonomy and distribution of freshwater pearl mussels ( unionoida : margaritiferidae ) of the russian far east . plos one 10 ( 5 ) : e0122408 . urltoken
huff sw , campbell d , gustafson dl , lydeard c , altaba cr , giribet g . investigations into the phylogenetic relationships of freshwater pearl mussels ( bivalvia : margaritiferidae ) based on molecular data : implications for their taxonomy and biogeography . j mollus stud . 2004 ; 70 : 379–388 .
diagrams of the five distinct gene orders detected in unionida . in the female f - type lineage , three gene orders are depicted : unionidae f - type 1 ( uf1 ) , unionidae f - type 2 ( uf2 ) and margaritiferidae f - type 1 ( mf1 ) . in the male m - type lineage , two gene arrangements are shown : unionidae m - type 1 ( um1 ) and margaritiferidae m - type 1 ( mm1 ) . continuous lines indicate different locations of genes between mitogenomes . grey box highlights gene rearrangement region between uf1 and uf2 . yellow boxes indicate main differences in gene arrangement between female and male mitogenomes , trna ( h ) location and rearrangement of atp8 - trna ( d ) region .
stephanie w . huff , david campbell , daniel l . gustafson , charles lydeard , cristian r . altaba , gonzalo giribet ; investigations into the phylogenetic relationships of freshwater pearl mussels ( bivalvia : margaritiferidae ) based on molecular data : implications for their taxonomy and biogeography , journal of molluscan studies , volume 70 , issue 4 , 1 november 2004 , pages 379–388 , urltoken
the family margaritiferidae includes 13 extant species , which are mainly distributed in temperate latitudes of the northern hemisphere [ 1 ] , [ 2 ] , [ 3 ] . smith [ 4 ] provided a detailed diagnosis of the family . recent species are known from north america , europe , northern africa , the middle east , and throughout much of southern and eastern asia [ 2 ] , [ 4 ] . the most ancient margaritiferidae fossils are known from the upper triassic and lower jurassic fluvio - lacustrine deposits in the sichuan , southeastern china [ 5 ] , [ 6 ] . the recent margaritiferids retain the simple , unfused mantle margins from the ancestral palaeoheterodont and several other ‘plesiomorphic’ features ; therefore , these species have been regarded as the basal unionoid family [ 3 ] , [ 7 ] .
the north american , european and northern african margaritiferidae are relatively well studied [ 8 ] , [ 4 ] , [ 9 ] , [ 10 ] , [ 11 ] in contrast to the asian representatives of the family . most of the references for far eastern freshwater pearl mussel populations are from japan [ 12 ] , [ 13 ] , [ 14 ] , [ 15 ] , [ 16 ] . recently , a description of a new japanese margaritiferidae species m . togakushiensis kondo & kobayashi , 2005 was published . this species was separated from m . laevis based on the results of long - term studies , specifically including differences in the host fish preference and genetic and morphological patterns [ 17 ] , [ 18 ] , [ 19 ] , [ 20 ] , [ 21 ] .
manuel lopes - lima , miguel m . fonseca , david c . aldridge , arthur e . bogan , han ming gan , mohamed ghamizi , ronaldo sousa , amílcar teixeira , simone varandas , david zanatta , alexandra zieritz , elsa froufe ; the first margaritiferidae male ( m - type ) mitogenome : mitochondrial gene order as a potential character for determining higher - order phylogeny within unionida ( bivalvia ) , journal of molluscan studies , volume 83 , issue 2 , 1 may 2017 , pages 249–252 , urltoken
shows the topology of the bi - nuc tree ; all other phylogenetic trees figures supplied on request ) . additionally , the monophyly of unionidae ( both f - and m - type ) , margaritiferidae ( f - type ) and all represented unionidae subfamilies are well supported in all inferred mtdna trees , with the exception of the unioninae , for which monophyly was only well supported in the bi - nuc tree . the remaining phylogenetic trees ( bi - aa , ml - nuc and ml - aa ) showed conflicting results regarding the position of the clade comprising
according to kondo & kobayashi [ 18 ] , the adductor muscle scar shape is one of the key features for m . togakushiensis identification . however , according to our data , this feature has a high variability and cannot be used for species identification among studied taxa . in the review of recent margaritiferidae by smith [ 4 ] , this feature was also not mentioned as a diagnostic . the adductor muscle scar shape was not used in the morphological descriptions of other unionoid species [ 10 ] , [ 86 ] , [ 95 ] , [ 96 ] , [ 97 ] .
the dimensions of the shell are the least reliable morphological traits in unionoida , upon which to base a description [ 4 ] , [ 32 ] due to high variability and the dependence on environmental gradients [ 4 ] , [ 33 ] , [ 34 ] , [ 100 ] , [ 101 ] . however , sometimes , the shell dimensions and shape are useful for distinguishing several margaritiferidae species . for example , m . middendorffi has a small shell length ( < 100 mm ) ( fig 10 ) , and m . dahurica has the largest shell length ( up to 196 . 5 mm ) among the far eastern pearl mussels ( fig 5 ) .
the analysis of the mitochondrial coi gene showed that c . monodonta forms a separate clade , which is basal within margaritiferidae . m . dahurica , m . margaritifera , m . middendorffi , m . laevis and m . falcata cluster together in a well - supported clade ( bpp 1 . 00 ) ( fig 2b ) . among these , our bayesian inference strongly supports the sister relationships within two pairs of species , namely m . dahurica and m . margaritifera as the first pair ( bpp 1 . 00 ) , and m . middendorffi and m . laevis as the second pair ( bpp 0 . 97 ) . m . falcata shows a closer affinity to m . middendorffi and m . laevis ( bpp 0 . 93 ) . lastly , the coi phylogeny confirms that m . auricularia and m . marocana are sister taxa ( bpp 0 . 98 ) .
the length of the two newly sequenced mitogenomes of m . marocana , 16 , 001 bp for the female haplotype and 17 , 562 bp for the male haplotype , is within the typical range for each sex - specific haplotype within unionida . the sequenced haplotypes include the 13 pcgs typically found in metazoan mitochondrial genomes , the sex - specific orf described for all unionida mitogenomes with dui system ( breton et al . , 2009 , 2011a ) and 22 transfer rna ( trna ) and two ribosomal rna ( rrna ) genes . the m - type genome is the largest sequenced to date within the unionida . m - type genomes are generally larger than f - type genomes due to the larger size of the pcg cox2 and m - orf in m - type genomes compared with cox2 and f - orf in f - type genomes ( breton et al . , 2009 ) . four intergenic regions were identified in the m . marocana m - type genome between the following gene pairs : nad3–trna ( a ) 106 bp , trna ( h ) –trna ( q ) 411 bp , nd4l–trna ( d ) 255 bp and trna ( d ) –atp8 498 bp . these regions were analysed to search for the m - orf . the results of the blast search ( altschul et al . , 1997 ) retrieved a significant hit with another margaritiferidae m - orf ( margaritifera monodonta , e - value = 4e–34 ) and a fickett test score of 1 . 201 ( fickett , 1982 ; a score > 0 . 95 means the sequence is probably coding ) , suggesting that the m - orf is located in the region between the genes nd4l and trna ( d ) .
genus margaritinopsis accepted as margaritanopsis f . haas , 1910 accepted as margaritifera schumacher , 1815 ( incorrect subsequent spelling )
henderson j . ( 1929 ) . non - marine mollusca of oregon and washington . the university of colorado studies . 17 ( 2 ) : 47 - 190 . page ( s ) : 53 [ details ]
invertebase . ( 2015 ) . authority files of u . s . and canadian land and freshwater mollusks developed for the invertebase project ( invertebase . org ) . [ details ]
( of margaritanidae ortmann , 1910 ) graf , d . ; cummings , k . ( 2007 ) . review of the systematics and global diversity of freshwater mussel species ( bivalvia : unionoida ) . journal of molluscan studies . 73 ( 4 ) : 291 - 314 . [ details ]
( of margaritanidae ortmann , 1910 ) invertebase . ( 2015 ) . authority files of u . s . and canadian land and freshwater mollusks developed for the invertebase project ( invertebase . org ) . [ details ]
( of cumberlandiidae heard & guckert , 1970 ) graf , d . ; cummings , k . ( 2007 ) . review of the systematics and global diversity of freshwater mussel species ( bivalvia : unionoida ) . journal of molluscan studies . 73 ( 4 ) : 291 - 314 . [ details ]
( of cumberlandiidae heard & guckert , 1970 ) invertebase . ( 2015 ) . authority files of u . s . and canadian land and freshwater mollusks developed for the invertebase project ( invertebase . org ) . [ details ]
discover a faster , simpler path to publishing in a high - quality journal . plos one promises fair , rigorous peer review , broad scope , and wide readership – a perfect fit for your research every time .
copyright : © 2015 bolotov et al . this is an open access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited
data availability : data are available from : rmbh - russian museum of the biodiversity hotspots of institute of environmental problems of the north of the ural branch of russian academy of science , arkhangelsk , 163000 , russia . inrec - institute of natural resources , ecology and cryology of the siberian branch of russian academy of sciences , chita , 672000 , russia . zisp - zoological institute of the russian academy of sciences , st . petersburg , 199034 , russia . smf - forschungsinstitut und natur - museum senckenberg , senckenberg - anlage 25 , 6000 frankfurt - am - main 1 , germany .
funding : this study was supported by the federal agency for scientific organisations ( no . 0410 - 2014 - 0028 ) , the ural branch of russian academy of sciences ( nos . 15 - 12 - 5 - 3 and 15 - 2 - 5 - 7 ) , grants from the president of russia ( no . md - 6465 . 2014 . 5 and мк - 4735 . 2015 . 4 ) and the russian foundation for basic research ( nos . 14 - 04 - 98801 and 15 - 04 - 05638 ) and the ministry of science and education of russia ( no . p362 ) . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
thus , the available information on the asian freshwater pearl mussels is limited . their taxonomy is not completely clear and reliable data on the species’ ranges are sparse . however , freshwater pearl mussels are extremely demanding regarding habitat quality and can exist only in a narrow range of environmental conditions [ 11 ] . the majority of these species are listed by iucn as endangered or vulnerable taxa [ 8 ] , [ 9 ] , [ 10 ] , [ 11 ] , [ 40 ] . freshwater mussels have experienced one of the highest rates of extinction of any group of organisms in the past 100 years [ 41 ] . therefore , reliable information on the taxonomy and distribution ranges is still needed for the conservation of the margaritifera species .
we conducted field studies within several regions of eastern russia in the period 2004–2012 ( see fig 1 for the location of the study areas ) . sampling on the russian federation territory was permitted within the framework of the special grants of the russian foundation for basic research ( rfbr , no . 12 - 04 - 00594 ) , the ministry of science and education of russia ( no . № p362 ) , and the scientific program of the ural branch of russian academy of sciences ( no . 12 - p - 5 - 1014 ) . sampling on the protected sites of the kunashir island was permitted by the directorate of the state nature protected area “kurilsky” ( scientific agreement no . 5 / 11 of 17 . 08 / 2010 between the institute of ecological problems of the north of the ural branch of russian academy of sciences and the kurilsky nature reserve ) .
1—kamchatka peninsula ( 2012 , i . bolotov , y . bespalaya , i . vikhrev , m . gofarov ) , 2—central sakhalin ( 2012 , same team ) , 3—southern sakhalin ( 2011–2012 , same team & y . kolosova , o . aksenova ) , 4—kunashir island ( 2011 , same team & y . kolosova , o . aksenova ) , 5—primorye ( 2012 , same team ) , 6—transbaikalia ( 2004–2011 , o . k . klishko ) . data on the studied river sites are presented in s1 – s4 tables .
for molecular analyses , the tissue samples were collected from 52 live bivalves in 15 localities ( see s1 table ) . samples were immediately preserved in 96 % ethanol . dead shells were collected from each field site for morphological investigation ; additional shells from some museum collections were studied ( see s2 – s4 tables ) . a total of 554 shells from 44 localities were studied ( 426 from our collection and 128 museum specimens ) .
the total length of the live specimens and dead shells were measured to the nearest 0 . 1 mm with dial calipers [ 8 ] . the comparative morphologies of the shells were analyzed using shape and structure of the pseudo - cardinal and lateral teeth in the valves , shell shape , umbo position and the patterns of distribution of mantle attachment scars on the inner shell surface . we calculated the plane projection squares of the inner surface of the right valve for the estimation of the density of mantle attachment scars . ten specimens of each species were taken for analysis . the squares were calculated using shape v . 1 . 3 software [ 42 ] . the scars were counted by the gnu image manipulation program ( gimp ) v . 2 . 8 . a normality test of the obtained density values and a one - way anova welsh’s test ( s5 table ) were calculated using statistica v . 10 software .
the soft tissue morphologies were analyzed and compared based on the shape and structure of the inhalant siphon , gills and labial palps . shell and soft tissue photos were taken using a dslr camera ( canon eos 7d , japan ) with a 24–70 mm lens ( sigma af 24–70 mm f / 2 . 8 if ex dg aspherical hsm , japan ) , and photos of the shell structure details and inhalant siphons were taken using a stereomicroscope ( leica m2c , germany ) and a dslr camera with a 100 mm macro lens ( canon macro lens ef 100 mm 1 : 2 , 8 l is usm , japan ) . descriptions of the shell morphologies were based on an analysis of the collected specimens and museum samples using the original species descriptions [ 18 ] , [ 38 ] , [ 43 ] , [ 44 ] , [ 45 ] , [ 46 ] and a few other works [ 23 ] , [ 24 ] , [ 47 ] , [ 48 ] .
we studied the type specimens for the following taxa : unio ( alasmodonta ) dahuricus middendorff , 1850 ; margaritana middendorffi rosén , 1926 ( including specimens of unio ( alasmodonta ) complanatus middendorff , 1851 ) ; and m . sachalinensis zhadin , 1938 ( zisp ) . the type series of several comparatory “taxa” were also investigated , including dahurinaia sujfunensis moskvicheva , 1973 ; d . kurilensis zatravkin & starobogatov , 1984 ; d . tiunovae bogatov & zatravkin , 1988 ; d . komarovi bogatov et al . , 2003 ; d . ussuriensis bogatov et al . , 2003 ; d . prozorovae bogatov et al . , 2003 ; d . transbaicalica klishko , 2008 ; kurilinaia kamchatica bogatov et al . , 2003 ; and k . zatravkini bogatov et al . , 2003 ( zisp ) . in general , the zisp systematic catalog has a very complicated numeration because v . v . bogatov has transferred many specimens from different samples , including true type series , to his newly described comparatory “taxa” .
the mapping was based on the data of our field studies ( see s2 – s4 tables ) . we determined the precise coordinates of investigated river sites using a geographical positioning system ( gps ) . data on other localities were obtained from museum collections and reliable published studies ( see s2 – s4 tables ) . in the cases where a particular map scale did not allow separated precision pointing of closely situated localities , we marked those localities one by one with points and joined them under the one number in the respective table . the arrangement of the localities was digitized and mapped using esri arcgis 10 . the presumed error of determination of the locality coordinates is around ±1–2 km , because published records and collection labels are usually ascribed to an approximate location . the layers of digital maps were added from standard esri data & maps 10 dataset .
) . genomic dna was extracted from the foot or mantle tissues using the diatom dna prep 200 reagents kit ( “laboratoriya isogen” llc , russia ) following the manufacturer’s protocol . the standard forward primer lco 1490 [
] was used to amplify the mitochondrial coi gene from every species . as reverse coi primers , hco 2198 in a modified version ( without the first 6 bases at the 5´ end ) was applied to
) were used to amplify the nuclear 18s rrna gene from every species . the pcr mix contained approximately 200 ng of total cellular dna , 10 pmol of each primer , 200 μmol of each dntp , 2 . 5 μl of pcr buffer ( with 10×2 mmol mgcl
) , 0 . 8 units of taq dna polymerase ( sibenzyme ltd . , novosibirsk , russia ) , and h
o , which was added up to a final volume of 25 μl . thermocycling included one cycle at 95°c ( 4 min ) , followed by 30–35 cycles of 95°c ( 50 sec ) , 52°c ( 50 sec ) , and 72°c ( 50 sec ) and a final extension at 72°c ( 5 min ) . forward and reverse sequencing was performed on an automatic sequencer ( abi prism 3730 , applied biosystems ) using the abi prism bigdye terminator v . 3 . 1 reagent kit . the resulting sequences were checked using a sequence alignment editor ( bioedit version 7 . 2 . 5 , [
the electronic edition of this article conforms to the requirements of the amended international code of zoological nomenclature , and hence the new names contained herein are available under that code from the electronic edition of this article . this published work and the nomenclatural acts it contains have been registered in zoobank , the online registration system for the iczn . the zoobank lsids ( life science identifiers ) can be resolved and the associated information viewed through any standard web browser by appending the lsid to the prefix “ urltoken ” . the lsid for this publication is : urn : lsid : zoobank . org : pub : 46c93315 - 18e0 - 45dd - b121 - 1f225d6e200f . the electronic edition of this work was published in a journal with an issn , and has been archived and is available from the following digital repositories : pubmed central and lockss .
rmbh—russian museum of the biodiversity hotspots of institute of ecological problems of the north of the ural branch of russian academy of science , arkhangelsk , 163000 , russia .
inrec—institute of natural resources , ecology and cryology of the siberian branch of russian academy of sciences , chita , 672000 , russia .
zisp—zoological institute of the russian academy of sciences , st . petersburg , 199034 , russia .
smf—forschungsinstitut und natur - museum senckenberg , senckenberg - anlage 25 , 6000 frankfurt - am - main 1 , germany .
the bayesian analysis of the nuclear 18s rrna gene revealed that the nw pacific species m . middendorffi and m . laevis belong to a well - supported monophyletic clade ( bpp 1 . 00 ) together with the north american species m . marrianae and m . falcata ( fig 2a ) . among these species , m . middendorffi is a sister to m . marrianae ( bpp 1 . 00 ) . m . dahurica , m . auricularia and c . monodonta cluster together in an unresolved clade ( bpp 0 . 79 ) , but there is a sister relationship between the two latter species ( bpp 0 . 99 ) . m . dahurica and m . auricularia both have an intra - specific variation of the nuclear 18s rrna gene with two close haplotypes in each species .
the scale bar indicates the branch length . asterisks : posterior probabilities ≥0 . 95 ; other significant support node values are mentioned in the figure . for detailed locality and specimen data for analyzed haplotypes , see the supplementary materials ( s1 and s2 tables ) . a—the 18s rdna gene dataset . b—the coi gene dataset .
the structure of hinge teeth and the shape and relative dimensions of the shell have specific characteristics that are described in detail below in the taxonomic account of each species . the density of mantle scars has no significant differences ( welch’s test : f = 2 . 45 , df = 16 , 38 , p = 0 . 12 ; ( s5 table ) ) whereas muscle scar has a high variability among studied taxa .
a— margaritifera dahurica ( middendorff , 1850 ) . b— m . laevis ( haas , 1910 ) . c— m . middendorffi ( rosén , 1926 ) . scale bar—1 cm .
a— margaritifera dahurica ( middendorff , 1850 ) . b— m . middendorffi ( rosén , 1926 ) . c— m . laevis ( haas , 1910 ) . scale bar—2 mm .
unio ( alasmodonta ) dahuricus middendorff , 1850 : [ 43 ] : 109 , [ 44 ] : 275–276 , pl . 26 , figs 3–5 .
unio ( margaritana ) dahuricus middendorff , 1850 : [ 58 ] : 699–700 .
unio margaritiferus simpson 1895 , partim ( ident . err . , non linnaeus , 1758 ) : [ 59 ] : 328 .
margaritana dahurica ( middendorff , 1850 ) : [ 23 ] : 109–112 , fig 35 , [ 24 ] : 289 , fig 250 .
margaritifera margaritifera dahurica ( middendorff , 1850 ) : [ 60 ] : 120 , [ 61 ] : 11 .
dahurinaia sujfunensis moskvicheva , 1973 ( comparatory “taxa” ) : [ 62 ] : 1468 , fig 3 , e - h .
dahurinaia sujfunica moskvicheva , 1973 ( inadv . err . in the species description ) . [ 62 ] : 1468 .
dahurinaia tiunovae bogatov & zatravkin , 1988 ( comparatory “taxa” ) : [ 63 ] : 156–158 , fig 1 , a - b .
dahurinaia komarovi bogatov et al . , 2003 ( comparatory “taxa” ) : [ 26 ] : 45–47 , figs 3a & 3d .
dahurinaia ussuriensis bogatov et al . , 2003 ( comparatory “taxa” ) : [ 26 ] : 47 , figs 3b & 3g .
dahurinaia prozorovae bogatov et al . , 2003 ( comparatory “taxa” ) : [ 26 ] : 48 , figs 3c & 3i .
dahurinaia transbaicalica klishko , 2008 ( comparatory “taxa” ) : [ 28 ] : 292–296 , figs 1–4 , 5a .
dahurinaia ( kurilinaia ) laevis klishko , 2009 ( ident . err . , non haas , 1910 ) : [ 64 ] : 237–238 , figs 1–2 .
dahurinaia ( kurilinaia ) zatravkini klishko , 2009 ( ident . err . , non bogatov et al . , 2003 ) : [ 64 ] : 238–239 , fig 3 .
margaritifera dahurica inoue et al . , 2013 ( inadv . err . ) : [ 65 ] : fig 3 ( a ) .
a—lectotype ( zisp : no . 7a ) . b—specimen from komarovka river , razdolnaya river basin , primorye . photos by i . v . vikhrev . scale bar—2 cm .
a— margaritifera dahurica ( middendorff , 1850 ) . b— m . middendorffi ( rosén , 1926 ) . c— m . laevis ( haas , 1910 ) . scale bar—1 cm .
microphotographs of the mantle attachment scars on shells of four eastern asian margaritiferid species .
a— margaritifera dahurica ( middendorff , 1850 ) . b— m . middendorffi ( rosén , 1926 ) . c— m . laevis ( haas , 1910 ) .
green circles are representing recent viable populations ( observed since 2000 ) , white circles—old records ( until 2000 ) . question mark is indicated an uncertain record from the langry river [ 69 ] , [ 26 ] . grey areas are indicated an approximate modern species range ( it is shown only for the large river systems ) . species locality numbers on the map correspond to numbers in s2 table .
a— margaritifera dahurica ( middendorff , 1850 ) : the ilystaya river , primorsky kray . b— m . middendorffi ( rosén , 1926 ) : the nachilova river , kamchatka . c— m . middendorffi ( rosén , 1926 ) & m . laevis ( haas , 1910 ) : the golovnina river , kunashir island . photos by y . v . bespalaya , y . s . kolosova & i . v . vikhrev .
“transbaikalien” [ 43 ] : 109 ; “am zusammenflusse des argunj mit der schilka” [ 44 ] : 276 ( confluence of the rivers argun and shilka , the upper amur basin : see fig 8 & s2 table , locality no . 36 ) .
the middendorff’s type series includes two specimens that are deposited in the zisp collection . bogatov et al . [ 26 ] : 45 designate specimen no . 7a as a lectotype , because it was pictured by middendorff [ 44 ] : pl . 26 , figs 3 – 5 ; and assigned the second specimen no . 7b ( single valve ) as a paralectotype . the lectotype shell has 105 mm length , 32 mm height and 25 mm width . it is noteworthy , that bogatov et al . [ 26 ] were not using the protologue of middendorff [ 43 ] for their revision , and cited only middendorff’s later work [ 44 ] .
the amur basin ( within the territories of russia , mongolia and china ) , the razdolnaya basin ( upper tributaries ) , the peschernaya ( kulumbe ) river , the iska river and the arey lake ( fig 8 & s2 table ) .
different types of watercourses , such as small streams , small and medium sized rivers , as well as large rivers ( rivers ussuri , arkhara , shilka , etc . ) ( fig 9a ) . the sole lake population was found in arey lake , transbaikalia , siberia , russia [ 28 ] ; but divers recorded only a few dead shells here in the year 2013 ( o . k . klishko , pers . comm . ) . populations prefer sand - gravel and gravel - pebble grounds at the riffles and runs . sometimes , individual specimens were observed on the clay and silt - sand bottom of the pool river sites .
not known . actually , a list of seven salmonid species , which can serve as hosts of the m . dahurica [ 36 ] , has not been verified experimentally .
all of the referred comparatory “taxa” of the genus dahurinaia were synonymized with m . dahurica based on shell morphology [ 4 ] , [ 8 ] , [ 31 ] , [ 39 ] . our molecular data confirmed this decision . for example , bogatov [ 29 ] noted that the single known dahurinaia sujfunensis population inhabits razdolnaya basin , which is separate from the amur drainage . bogatov reports that “…between the amur basin and basins of rivers of the south of primorsky kray , no common species of large bivalves has been found yet , which is explained by the historic development of these basins” [ 29 ] : 674 . however , according to new molecular and morphological data , the specimens from the razdolnaya drainage are identical to typical m . dahurica specimens from different parts of the river amur basin .
unio ( alasmodonta ) complanatus middendorff , 1851 ( ident . err . , non dillwyn , 1817 ) : [ 44 ] : 273–274 , pl . 27 , figs 1–6 .
margaritana middendorffi rosén , 1926 : [ 46 ] : 269–270 , [ 23 ] : 112–114 , fig 36 , [ 24 ] : 289 , fig 251 .
dauhrinaia middendorffii buyanovsky , 1993 ( inadv . err . ) : [ 67 ] : 29 .
kurilinaia kamchatica bogatov et al . , 2003 ( comparatory “taxa” ) : [ 26 ] : 48 , figs 4a & 4c .
kurilinaia zatravkini bogatov et al . , 2003 , partim ( comparatory “taxa” ) : [ 26 ] : 49–50 , figs 4b & 4f .
margaritifera laevis huff et al . , 2004 ( ident . err . , non haas , 1910 ) : [ 38 ] : 381 , genbank acc . no . ay579124 .
a—holotype of m . togakushiensis [ 18 ] : 137 , figs 5 – 8 . b—lectotype of m . middendorffi ( zisp : no . 6 ) . photo by i . v . vikhrev . scale bar—2 cm .
circles— m . middendorffi locations , squares— m . togakushiensis locations . green circles and squares are representing recent viable populations ( observed since 2000 ) , white circles—old records ( until 2001 ) , yellow squares—records without exact dates . grey areas indicate approximate modern species ranges ( showing only the large river systems ) . species locality numbers on the map correspond to numbers in s3 table .
“reka golyshka” [ the river golygina , kamchatka peninsula : see fig 11 & s3 table , locality no . 1 ] [ 46 ] : 269 ; “kamtschatka ; im see mjäkéshino des südendes ( lopatka ) dieser halbinsel” ( kamchatka ; the lake mekeshino on the southernmost extremity of the peninsula : see fig 11 & s3 table , locality no . 4 ) [ 44 ] : 274 .
rivers of the kamchatka peninsula , kurile archipelago , sakhalin island ( fig 11 & s3 table ) , and likely of japan .
small streams , small - and medium - sized rivers ( fig 9b ) . usually , populations were recorded on sand - gravel grounds of the runs but , in the extreme north of the range , the species inhabited only pool river sites . in some rivers of the sakhalin and kunashir islands , far east of russia , the species coexists with m . laevis ( see fig 9b & s3 table ) .
not known . kondo & kobayashi [ 18 ] stated salvelinus leucomaenis as a host fish for m . togakushiensis . as far as we suppose m . togakushiensis and m . middendorffi conspecific , they may have the same host fish . [ 18 ] .
then , simpson [ 59 ] : 328 noted that middendorf’s specimens are “…without lateral teeth , and appear to be a stunted form of unio margaritiferus ” . finally , rosén [ 46 ] described m . middendorffi as a separate species and not as an intra - specific form of m . margaritifera . however , haas [ 61 ] , [ 60 ] incorrectly cited rosén’s name as margaritana margaritifera middendorffi . kurilinaia kamchatica bogatov et al . , 2003 is a comparatory “species” that was recently synonymized with m . middendorffi [ 31 ] . kurilinaia zatravkini bogatov et al . , 2003 is also a comparatory “species” ; its holotype belongs to m . laevis , but among the paratypes , we found four specimens of m . middendorffi ( zisp : nos . 5 , 5a , 7 and 8 ) . records of margaritifera specimens on kunashir island [ 68 ] : 134 can pertain to m . middendorffi as well as to m . laevis . therefore , some published references [ 8 ] , [ 67 ] , [ 69 ] , [ 70 ] with records from the sakhalin and southern kurile islands cannot be used without revision of the specimen samples .
margaritana dahurica kobelt , 1879 ( ident . err . , non middendorff , 1850 ) : [ 71 ] : 427–428 , pl . 13 , figs 1–2 .
unio margaritifer schrenck , 1867 , partim ( ident . err . , non linnaeus , 1758 ) : [ 58 ] : 700–704 .
unio margaritiferus simpson , 1895 , partim ( ident . err . , non linnaeus , 1758 ) : [ 59 ] : 303 .
margaritana sachalinensis zhadin , 1938 : [ 23 ] : 114–115 , fig 37 , [ 24 ] : 289–291 , fig 252 .
margaritifera margaritifera laevis ( haas , 1910 ) : [ 60 ] : 120 , [ 61 ] : 12 .
dahurinaia kurilensis zatravkin & starobogatov , 1984 ( comparatory “taxa” ) : [ 72 ] : 1789–1790 , figs 11–14 , [ 66 ] : 21 .
dahurinaia shigini zatravkin & bogatov , 1987 : ( comparatory “taxa” ) : [ 66 ] : 23–24 , fig 4b .
margaritifera ( dahurinaia ) kunahiriensis habe , 1991 ( inadv . err . ) : [ 73 ] : 3 .
dauhrinaia kurilensis buyanovsky , 1993 ( inadv . err . ) : [ 67 ] : 29 .
margaritana sacchariensis bába , 2000 ( inadv . err . ) : [ 74 ] : 133 .
kurilinaia kurilensis ( zatravkin & starobogatov ) : [ 26 ] : 42 , [ 27 ] : 25 .
kurilinaia zatravkini bogatov et al . , 2003 : partim ( comparatory “taxa” ) : [ 26 ] : 49–50 , figs 4b & 4f .
a—sennaya river , kunashir island . b—tym’ river , sakhalin island . photos by i . v . vikhrev . scale bar—2 cm .
green circles are representing recent viable populations ( observed since 2000 ) , white circles—old records ( until 2001 ) , yellow circles—records without exact dates . grey areas indicate the approximate modern species range ( showing only the large river systems ) . species locality numbers on the map correspond to numbers in s4 table .
type specimen ( holotype ) of haas [ 45 ] is deposited in the senckenberg museum , frankfurt ( no . smf3626 ) [ 39 ] .
sakhalin island , south kurile islands ( kunashir , iturup and shikotan ) , honshu and hokkaido islands ( fig 13 ) . correspondence on the species occurrence in the upper amur basin , transbaikalia [ 64 ] is based on an erroneous identification ; however , all of these specimens belong to m . dahurica based on morphological and molecular data ( see s1 table ) .
small streams , small - and medium - sized rivers . usually , populations were recorded on sand - gravel grounds at the run and pool sites . in several rivers of sakhalin , kunashir , honshu and hokkaido islands , islands , the species coexists with m . middendorffi ( see fig 9c & s4 table ) .
the masu salmon ( oncorhynchus masu ( brevoort , 1856 ) ) [ 18 ] .
in the main , species synonymy was provided in previous reviews [ 4 ] , [ 18 ] , [ 31 ] , [ 39 ] . m . perdahurica ( yokoyama , 1932 ) , m . otatumei ( suzuki , 1942 ) and m . owadaensis ( noda , 1970 ) , three fossil cenozoic margaritiferid species from japan , were also assumed as m . laevis representatives [ 39 ] . however , the condition of specimens of these species is very poor [ 75 ] , [ 76 ] , [ 77 ] , [ 78 ] . it is impossible to reliably compare these fossils with shells of recent margaritiferids . moreover , most of the known fossil specimens , including m . perdahurica and m . otatumei , are ancient and belong to the eocene or oligocene by stratigraphic classification [ 78 ] , [ 79 ] . it is unlikely that these fossils belong to recent species , but thorough revision of all specimens and collections of fossil margaritiferids would be necessary , in order to clarify the relationships of these bivalves [ 80 ] .
three margaritifera species inhabit the rivers of the russian far east , as discerned from morphological and molecular data : m . dahurica ( middendorff , 1850 ) , m . middendorffi ( rosén , 1926 ) and m . laevis ( haas , 1910 ) . however , the rivers of northeastern russia , where large mainland regions remain unexplored , are still not completely studied ; therefore the same species that were previously noted by smith [ 4 ] or additional species , may inhabit the north of the khabarovsky kray , the magadan oblast , and the koryakia and chukotka districts . these areas are difficult to access and require future studies .
m . dahurica has the largest range among the eastern asian species in the genus ( see fig 8 ) . this species is found in almost all of the major tributaries of amur basin , an area of approximately two million km 2 , as well as in razdolnaya basin and in two separate small rivers . a similar distribution pattern was found in some freshwater fish species , among which approximately 16 were endemic to this area [ 81 ] . we found only a single old record of m . dahurica from sungari river , the largest chinese tributary of the amur , and a few occurrences from mongolia . however , this species is likely widely distributed in part of the amur river system within the territory of those countries . dashi - dorgi [ 82 ] reported m . dahurica as one of the most common mussel species in the rivers of eastern mongolia . this species is not found in the rivers of the kurile islands and the most of sakhalin island . bogatov [ 69 ] and bogatov et al . [ 26 ] reported a few specimens from the langry river , which is situated on the extreme northwest of the sakhalin island . it is possible that because this river empties into the amur estuary , this river might have belonged to the ancient system of the paleo - amur [ 81 ] . the range of some amurean fish species also includes rivers in northwestern sakhalin island [ 83 ] . nevertheless , the species identification is in need of revision .
from our data , the range of m . laevis is significantly narrower than the distribution of the previous species ( see fig 13 ) . this species has a more southerly distribution , and has not been found north of central sakhalin . however , the host fish oncorhynchus masu is much more widespread northward up to kamchatka peninsula . most of the known localities of m . laevis are situated on the honshu island , and their number is linearly reduced to the north . a few fish species have a distribution resembling this , including the sakhalin taimen ( hucho perryi ( brevoort , 1856 ) ) [ 83 ] .
in this study , we analyzed the applicability of specific shell features that were indicated in previous studies for reliable species identification . three pearl mussel species were identified using a complex of morphology patterns and molecular data . the development and expression of hinge teeth , the shape and the relative dimensions of the shell , the mantle attachment scars , and the muscle scar shape are the most used conchological key features [ 4 ] , [ 10 ] , [ 18 ] , [ 23 ] , [ 24 ] , [ 34 ] , [ 38 ] , [ 47 ] , [ 43 ] , [ 44 ] , [ 45 ] , [ 46 ] , [ 95 ] , [ 96 ] , [ 97 ] , [ 98 ] , [ 99 ] .
s1 table . list of sequenced margaritifera specimens including species , localities and voucher details as well as ncbi genbank accession numbers .
s2 table . list of known localities of margaritifera dahurica ( middendorff , 1850 ) .
s3 table . list of known localities of margaritifera middendorffi ( rosén , 1926 ) and margaritifera togakushiensis ( kondo and kobayashi , 2005 ) .
s4 table . list of known localities of margaritifera laevis ( haas , 1910 ) .
s5 table . variance and significance tests results for mantle attachment scars density within three margaritiferid species .
s6 table . additional species sequences that were used in the analyses with ncbi genbank accession numbers .
we are thankful to to dr . p . v . kijashko and l . l . jarochnovich ( zoological institute of russian academy of sciences , st . petersburg , russia ) for assistance in studies of margaritifera specimens in zisp collection . the authors express their gratitude to mr . e . p . dekin ( russia ) , mr . m . a . antipin ( state nature protected area “kurilsky” , russia ) . special thanks go to mr . o . n . bespaliy ( russia ) for his excellent fieldwork assistance .
conceived and designed the experiments : inb yvb ivv . performed the experiments : inb yvb ivv ova myg okk ysk avk isp esk st nib isv . analyzed the data : inb yvb ivv avk . contributed reagents / materials / analysis tools : yvb ivv ova pea myg okk ysk avk aal isp esk st nib isv . wrote the paper : inb yvb ivv avk .
graf dl , cummings ks . review of the systematics and global diversity of freshwater mussel species ( bivalvia : unionoida ) . j mollus stud . 2007 ; 73 : 291–314 .
bogan ae , roe kj . freshwater bivalve ( unioniformes ) diversity , systematics , and evolution : status and future directions . j n am benthol soc . 2008 ; 27 : 349–369 .
graf dl . patterns of freshwater bivalve global diversity and the state of phylogenetic studies on the unionoida , sphaeriidae , and cyrenidae . am malacol bull . 2013 ; 31 : 135–153 .
liu x - z . on some newly discovered non - marine pelecypods from the late triassic wuzhongshan formation in sichuan basin [ in chinese ] . bulletin of the chengdu institute of geology and mineral resources , chinese academy of geological sciences . 1981 ; 2 : 121–136 .
fang z - j , chen j , chen c , sha j , lan x , wen s . supraspecific taxa of the bivalvia first named , described , and published in china ( 1927–2007 ) . the university of kansas paleontological contributions , new series . 2009 ; 17 : 1–157 .
graf dl , cummings ks . palaeoheterodont diversity ( mollusca : trigonioida + unionoida ) : what we know and what we wish we knew about freshwater mussel evolution . zool j linn soc lond . 2006 ; 148 : 343–394 .
ziuganov v , zotin a , nezlin l , tretiakov v . the freshwater pearl mussels and their relationships with salmonid fish . moscow : vniro , russian federal institute of fisheries and oceanography ; 1994 . 135 p .
araujo r , toledo c , van damme d , ghamizi m , machordom a .
( pallary , 1918 ) : a valid species inhabiting moroccan rivers . j mollus stud . 2007 ; 75 : 95–101 .
l . ) : a synthesis of conservation genetics and ecology . hydrobiologia . 2010 ; 644 : 69–88 .
akiyama y , iwakuma t . survival of glochidial larvae of the freshwater pearl mussel ,
( bivalvia : unionoida ) , at different temperatures : a comparison between two populations with and without recruitment . zool sci . 2007 ; 24 : 890–893 . pmid : 17960993
kondo t . monograph of unionoida in japan ( mollusca : bivalvia ) [ in japanese ] . special publication of the malacological society of japan . 2008 ; 3 : 1–69 .
. the biodiversity center of japan , the nature conservation bureau , the ministry of the environment , japan ; 2010 . p . 43
kurihara y , sakai h , kitano s , kobayashi o , goto a . genetic and morphological divergence in the freshwater pearl mussel ,
kobayashi o , kondo t . difference in host preference between two populations of the freshwater pearl mussel
kobayashi o , kondo t . comparative morphology of glochidia and juveniles between two species of freshwater pearl mussel
buldovsky at . about harvested freshwater mussels of the soviet far east [ in russian ] . proceedings of the far eastern branch of the ussr academy of sciences . 1935 ; 12 : 39–65 .
zhadin vi . fam . unionidae [ in russian ] . faune de l’urss , new series . 1938 ; 4 : 1–170 . pmid : 25389754
zhadin vi . mollusks of fresh and brackish waters of the ussr [ in russian ] . identification guides on the ussr fauna , issued by zoological institute of the ussr academy of sciences . 1952 ; 46 : 1–376 .
kurenkov ii . on distribution of the kamchatka freshwater pearl mussel [ in russian ] . questions of kamchatka geography . 1966 ; 4 : 110–112 .
starobogatov yi , prozorova la , bogatov vv , saenko em . mollusks [ in russian ] . in : freshwater invertebrates of russia and adjacent territories : an identification guide . vol . 6 : mollusks , polychaetes , and nemertines . st . petersburg : nauka publ . ; 2004 . pp . 9–422 .
) from the amur river basin . biol bull . 2012 ; 39 : 672–675 .
schum . , 1915 [ sic ] ( mollusca , bivalvia ) . biol bull + . 2013 ; 40 : 488–481 .
graf dl . palearctic freshwater mussel ( mollusca : bivalvia : unionoida ) diversity and the comparatory method as a species concept . p acad nat sci phila . 2007 ; 156 : 71–88 .
preston sj , harrison a , lundy m , roberts d , beddoe n , rogowski d . square pegs in round holes—the implications of shell shape variation on the translocation of adult
bolotov in , makhrov aa , bespalaya yuv , vikhrev iv , aksenova ov , aspholm pe , et al . the results of comparatory method testing : frontal section curvature of shell valve could not be a systematic indicator for freshwater pearl mussels of
akiyama b , kimura r , nomoto k , usui t , machida y . new record of the freshwater pearl mussel
from northern sakhalin , the russian far east . venus . 2013 ; 71 : 191–198 . pmid : 23542829
graf dl , cummings ks . the freshwater mussels ( unionoida ) of the world ( and other less consequential bivalves ) , updated 8 august 2013 . mussel project web site . available :
bogan ae . freshwater bivalve extinctions ( mollusca : unionoida ) : a search for causes . am zool . 1993 ; 33 : 599–609 .
haag wr . past and future patterns of freshwater mussel extinctions in north america during the holocene . in : turvey s . , editor . holocene extinctions . new york : oxford university press ; 2009 . pp . 107–128 .
iwata h , ukai y shape : a computer program package for quantitative evaluation of biological shapes based on elliptic fourier descriptors . j hered . 2002 ; 93 : 384–385 . pmid : 12547931
middendorff atv . beschreibung einiger neuer mollusken - arten , nebst einem blicke auf den geographischen charakter der land - und süsswasser - mollusken nord - asiens . bulletin de la classe physico - mathématique de l ' académie impériale des sciences de st . - pétersbourg . 1850 ; 9 : 108–112 .
middendorff atv . wirbellose thiere : annulaten . echinodermen . insecten . krebse . mollusken . parasiten . reise in den äussersten norden und osten sibiriens , zoologie . 1851 ; 2 : 163–508 . pmid : 24870494
haas f . new unionidae from east asia . the annals and magazine of natural history , 8th series . 1910 ; 6 : 496–499 .
rosén ov . terrestrial and freshwater mollusks , collected by kamchatka expedition of f . p . riabushinsky in 1908–1909 [ in russian ] . annuaire du musée zoologique de l’academie des sciences de l’urss . 1926 ; 27 : 261–274 .