amnicolidae

bythinella reyniesii is a species of very small freshwater snail , an aquatic gastropod mollusk in the family amnicolidae .
bythinella galerae is a species of very small freshwater snail , an aquatic gastropod mollusk in the family amnicolidae .
maackia is a genus of freshwater snails with an operculum , an aquatic gastropod mollusk in the family amnicolidae .
the amnicolidae and cochliopidae are considered as distinct families according to the taxonomy of bouchet & rocroi ( 2005 ) .
akiyoshia is a genus of freshwater snails with a gill and an operculum , aquatic gastropod mollusks in the family amnicolidae .
amnicolidae ,
4 genera , 18 species
[ north america ] ( johnson et al . , 2013 ) .
amnicolidae is a family of small freshwater snails with a gill and an operculum , aquatic gastropod mollusks in the superfamily rissooidea .
amnicolidae is a family of small freshwater snails with a gill and an operculum , aquatic gastropod mollusks in the superfamily rissooidea .
amnicola walkeri , common name canadian duskysnail , is a species of freshwater snail , an aquatic gastropod mollusk in the family amnicolidae .
amnicolidae , [ palearctic , nearctic ; 200 species worldwide , 19 nearctic species ] ( strong et al . , 2008 ) .
the family amnicolidae has been reported ( or is assumed ) to occur in fresh waters . this taxon has been reported from wisconsin .
the amnicolidae of michigan : distribution , ecology , and taxonomy . misc . publ . mus . zool . u . mich . , 57 , 1 - 68 .
gittenberger e , sherub s , stelbrink b ( 2017 ) erhaia davis & kuo ( gastropoda , rissooidea , amnicolidae ) also in bhutan . zookeys 679 : 21–28 . urltoken
thompson f . g . ( 1964 ) . a new aquatic snail of the family amnicolidae from costa rica . revista de biología tropical . 12 ( 1 ) : 97 - 105 . [ details ]
ty - jour ti - amnicolidae from oneida lake , n . y t2 - the nautilus . vl - 31 ur - urltoken pb - american malacologists , inc . , etc . cy - melbourne , fla . , etc . , py - 1917 sp - 44 ep - 46 sn - 0028 - 1344 au - pilsbry , h a er -
ty - jour ti - new and little known species of amnicolidae ( continued ) t2 - the nautilus . vl - 19 ur - urltoken pb - american malacologists , inc . , etc . cy - melbourne , fla . , etc . , py - 1906 sp - 114 ep - 117 sn - 0028 - 1344 au - walker , b er -
@ article { bhlpart95721 , title = { amnicolidae from oneida lake , n . y } , journal = { the nautilus . } , volume = { 31 } , copyright = { not _ in _ copyright } , url = urltoken publisher = { melbourne , fla . , etc . , american malacologists , inc . , etc . } , author = { pilsbry , h a } , year = { 1917 } , pages = { 44 - - 46 } , }
@ article { bhlpart95287 , title = { new and little known species of amnicolidae ( continued ) } , journal = { the nautilus . } , volume = { 19 } , copyright = { not _ in _ copyright } , url = urltoken publisher = { melbourne , fla . , etc . , american malacologists , inc . , etc . } , author = { walker , b } , year = { 1906 } , pages = { 114 - - 117 } , }
the specimens from the three populations of marstoniopsis scholtzi differ from each other in only one or two observed substitutions in 638 bp ( = 0 . 152 - 0 . 304 % ) . each one of them also differs from the specimen of m . insubrica in one or two observed substitutions ( = 0 . 152 - 0 . 304 % ) . based on the extremely low genetic divergence and the lack of morphological differentiation we consider all populations to belong to one species , marstoniopsis insubrica . maximum likelihood analysis places the genus marstoniopsis in the family amnicolidae , close to the genera erhaia and amnicola .
marstoniopsis scholtzi ( a . schmidt , 1856 ) and m . insubrica ( küster , 1853 ) , have been commonly accepted as distinct species , partly because of their allopatric distribution patterns , but there appear to be no morphological differences between them . also the phylogenetic relationships of the genus marstoniopsis van regteren altena , 1936 remain enigmatic . in the present study the authors sequenced a fragment of the mitochondrial cytochrome oxidase i gene in representatives of three populations of m . scholtzi and one population of m . insubrica to access the degree of genetic differentiation between those taxa . a phylogenetic analysis was performed with representatives of the families hydrobiidae , amnicolidae and cochliopidae in order to test the taxonomic position of marstoniopsis .
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worms - world register of marine species - rachipteron f . g . thompson , 1964
» genus gerstfeldtia clessin , 1880 accepted as godlewskia crosse & p . fischer , 1879
» genus limnorea w . dybowski , 1875 accepted as baicalia martens , 1876 ( invalid : junior homonym of limnorea goldfuss , 1826 [ porifera ] and limnorea agassiz , 1846 [ coleoptera ] . baicalia is a replacement name )
» genus pseudobaicalia b . dybowski , 1911 accepted as pseudobaikalia lindholm , 1909 ( unjustified emendation )
» genus turribaicalia b . dybowski & grochmalicki , 1917 accepted as baicalia martens , 1876
subfamily liobaicaliinae b . dybowski & grochmalicki , 1913 accepted as baicaliinae p . fischer , 1885
subfamily turribaicaliinae b . dybowski & grochmalicki , 1917 accepted as baicaliinae p . fischer , 1885
spring snails are a very diverse family of small to tiny snails inhabiting fresh or brackish water , also appearing on humid ground near the coast . very few species also actually are found in the sea , on muddy ground among seaweed . it is estimated there are about 300 species in north and south europe , about 1000 species worldwide , in australia alone 280 species are mentioned . in central europe many spring snail species inhabiting ground water , caves or springs are limited to one single geographic finding site , which means a very large ecological threat to those species .
in central and south europe about 40 species of spring snails are found , of which the five most important german species are defined mainly by anatomic characters , but are difficult to tell apart by shell characters . often , the single species can only be told apart by the finding location , as they are restricted to a very narrow area . this does lose importance in a growing fashion , as for example in austria there are often several species of spring snails in one place usually restricted to a narrow finding area .
the colour of spring snail shells often appears dark brown or green . this is due to diatoms ( brown ) and green algae growing on the shell surface . distribution of spring snail species often happens by insects capable of flying , on which the snails have laid eggs . spring snails live on diatoms , blue and green algae which they graze from the ground , as well as on detritus - decaying organic matter .
spring snails ( bythinella compressa ) on a fallen leaf . picture : © alexander mrkvicka , vienna ( urltoken ) .
accustomed to constantly low water temperatures , spring snails are cold - stenothermal creatures . they almost exclusively live on springs and in the uppermost areas of streams . the presence of spring snails is an indicator for very clean water ; in the springs of clean streams sometimes more than 1000 snails can be found per square metre .
spring snails are crenobionts , which means the are specialist with a very low ecological amplitude , which react to the slightest difference in environmental conditions , their density decreasing or the population disappearing altogether . on the other hand , spring snails are very well adapted to the extreme habitat of springs .
spring snails are usually threatened by fountain constructions around the springs they live in , by agricultural use as watering place for cattle . also drainage and ground water descent are a large problem to spring snails , and sadly so is overfertilisation , resulting in the eutrophication also of springs .
additionally , global warming results in the water temperatures of springs rising , which means , the cold - stenothermous populations of spring snails generally decrease .
limestone tuff spring near fischbach in the rhön mountains . habitat of the rhön spring snail ( bythinella compressa ) . picture : j . gombert ( source ) .
umweltschutz - news : klimawandel bedroht weichtier des jahres 2008 , die österreichische quellschnecke . ( in german ) urltoken : die österreichische quellschnecke ist das weichtier des jahres . ( in german ) naturschutzgroßprojekt thüringer rhönhutungen : feuchtlebensräume - naturnahe quellen und kalk - flachmoore . ( in german )
description : the austrian spring snail is smaller than bythinella bavarica ( see below ) and almost cylindrical . the whorls of its shell are flattened at the sides . also the suture is less prominent . the last whorl takes two fifths of the shell ' s overall height .
different spring snail species . right : bythinella cylindrica , left : belgrandiella wawrai ( hydro - biidae ) . picture : © alexander mrkvicka , vienna .
distribution : the species is an main indicator species for springs and low mountain streams . it is found in the eastern bavarian limestone alps and the sudetes , through moravia , austria as far as north - eastern tyrol , styria , carinthia , northern slovenia , hungary and galicia .
threat situation : in austria the austrian spring snail is classified as
near threatened
( nt ) ( see also : iucn threat categories ) .
robert a . patzner , naturschutzbund österreich : weichtier des jahres 2008 - bythinella austriaca . ( in german )
the shell is less slender , the suture noticeably deepened . the whorls are rounded at the sides , the last whorl takes two thirds of the overall shell height .
distribution : northern alps , eastern bavarian limestone alps and their foot hills . from munich east through bavaria , as far as northern tyrol and salzburg country , pottenstein near wiener neustadt ( lower austria ) and the high terrace near munich .
rhön spring snail ( bythinella compressa ) . picture : © alexander mrkvicka , vienna ( urltoken ) .
the rhön spring snail ' s shell is oval to bluntly conical , the last whorl takes four fifths of the shell height . the whorls are strongly rounded at the sides , the apex is blunted a little obliquely .
distribution : in germany : rhön and vogelsberg mountains in hesse and thuringia . also present in austria .
the rhön spring snail needs constantly cold and unpolluted water ( saprobial value of 1 . 0 ) with a temperature of around 7 - 8 °c . it feeds on bacteria growing on stones and leaves , as well as on decaying organic matter ( detritus ) , which is grazed from stones , water plants , fallen leaves ( picture ) and dead wood lying in the water .
rhön spring snails ( bythinella compressa ) on a fallen leaf . picture : © alexander mrkvicka , vienna ( urltoken ) .
while some time ago the rhön spring snail was wide spread in the open landscape shaped by man , today it appears nearly exclusively in spring exits and some hundreds of metres downstream in the streams of deciduous forest areas . in intact habitats there may be as many as 50 snails on 25 x 25 cm .
populations live isolated from each other . with their highly specialized demands to habitats ( see above ) , populations disappeared are lost forever .
description : the bavarian spring snail usually is large ( for a spring snail ) , turricular and conical with rounded whorls , the last of which is widened . the shell often is green coloured due to algae growing on it . the shell tip ( apex ) is small and blunted obliquely , the whorls are separated by a deep suture .
dimensions : h : 4 mm ; w : 2 . 3 mm ; n : 4 ½ - 5 . ( abbreviations ) .
distribution : the bavarian spring snail is a calciphile species - it needs limestone ground . its distribution area spreads over the northern alps and the foot hills ( lech and isar river areas ) . the bavarian spring snail can be found from southern württemberg as far east as the inn river and northern tyrol in austria .
, d . : unsere land - und süßwassermollusken , stuttgart , 1927 , s . 165 ff .
wide - lipped spring snail ( emmericia patula ) . picture : © alexander mrkvicka , vienna ( urltoken ) .
description : the wide - lipped spring snail has a bluntly conical shell , the last whorl of which has a distinctive palatal callus . the apertural lip , as the name states , is folded back to form a wide whitish lip . the whorls grow fast .
dimensions : h : 6 mm ; w : 4 mm . n : 4 ½ - 5 . ( abbreviations ) .
habitat and distribution : emmericia patula lives in springs and rivers rich in limestone .
the species ' distribution area spreads along the italian and yugoslavian adriatic coast from venetia as far south as central dalmatia . in southern france and bavaria ( central franconia ) , emmericia patula has been introduced in 1960 ; how and where from , is unknown . in the meantime , it has also been found in dachau and munich .
wide lipped spring snail ( emmericia patula ) from the tinavo springs near trieste , italy .
francisco welter - schultes : emmericia patula species homepage . dr . stefan nehring : neozoa ( makrozoobenthos ) in den deutschen gewässern - eine einführung . ( in german ) . molluscs with immigration background ( neobiota ) .
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kento furui added the japanese common name
イヌエンジュ属
to
maackia
.
jennifer hammock split the classifications by nmnh invertebrate zoology resource from amnicola to their own page .
jennifer hammock split the classifications by urltoken import from bythinella to their own page .
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disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
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this family is in the clade littorinimorpha ( according to the taxonomy of the gastropoda by bouchet & rocroi , 2005 ) . this family was previously considered to be a subfamily of hydrobiidae .
the atlantic drainages of georgia , comprising approximately 40 % of the waters of the state , are presently covered . . . ››
the four ecoregions of north carolina drain toward the atlantic through six major river systems . . . ›› more
although perhaps not as environmentally heterogeneous as neighboring states , south carolina does include . . . ›› more
the survey reported here is focused on the eastern 75 % of the state , draining from the blue ridge to the atlantic . . . ›› more
all waters draining into the tennessee river above the alabama line , including sw virginia , w north carolina , and n georgia . . . ›› more
a study area of approximately 150 , 000 km2 , including delaware , maryland , new jersey , eastern pennsylvania and the west virginia panhandle . . . ›› more
is widespread throughout most of north america , ranging north to the hudson bay , and as far west as utah ( berry 1943 , clarke 1981 ) . populations are not typically found in lotic environments , such as prevail in east tennessee . but in atlantic drainages of the southern united states ,
populations are common in the slow - moving rivers of the lower piedmont and coastal plain , often associated with detritus or woody debris . such waters may be darkened by high concentrations of organic compounds , but are not necessarily acidic . further north ,
are not found in waters that become hot , stagnant , or anaerobic . fwgna incidence rank
populations appear to be efficient grazers of diatoms and other periphyton ( kesler 1981 , cattaneo & kalff 1986 ) . they in turn may be eaten by crayfish ( lewis 2001 ) and sunfish ( osenberg 1989 , bronmark et al 1992 )
of dillon 2000 : 156 – 162 ) . but dynamics appear to be sensitive to aspects of water chemistry , especially hardness and ph ( shaw & mackie 1989 , 1990 ) . jokinen’s ( 1987 ) analysis of the distribution of
in connecticut and new york led her to classify it as a “c - d tramp , ” potentially present in nearly every community . dillon’s ( 2000 : 360 - 363 ) reanalysis of these data suggested that
( s . s . ) is distinguished by its ( entirely ) paucispiral operculum . an
co1 mtdna sequence appeared among the outgroups in the molecular phylogenetic study of liu and colleagues ( 2001 ) .
earlier versions of this website , online until august of 2016 , adopted the large , broadly - inclusive concept of the hydrobiidae ( sl ) following kabat & hershler ( 1993 ) . more recently the fwgna project has shifted to the wilke et al . ( 2013 ) classification system , distinguishing a much smaller hydrobiidae ( ss ) and elevating many hydrobioid taxa previously ranked as subfamilies to the full family level . for more details , see the classification of the hydrobioids .
) indirect effects of predation in a freshwater , benthic food chain . ecology 73 : 1662 – 1674 .
the freshwater molluscs of canada . national museums of canada , ottawa . 446 pp .
the ecology of freshwater molluscs . cambridge university press , cambridge , united kingdom . 509 pp .
( say , 1817 ) ( gastropoda : hydrobiidae ) with comments on status of the subfamily amnicolinae . malacol . rev . 21 : 81 - 92 .
structure of freshwater snail communities : species - area relationships and incidence categories . amer . malac . bull . 5 : 9 - 19 .
the prosobranch snail family hydrobiidae ( gastropoda : rissooidea ) : review of classification and supraspecific taxa . smithsonian contributions to zoology 547 : 1 - 94 .
( hydrobiidae ) eggs and individuals in a rhode island pond . nautilus 94 : 25 - 26 .
: an enclosure - exclosure experiment . j . freshwat . ecol . 1 : 51 - 59 . kesler , d . h . & tulou , c . a . g . ( 1980 ) cellulase activity in the freshwater gastropod
trade - offs between growth and survival : responses of freshwater snails to predacious crayfish . ecology 82 : 758 - 765 .
intraspecific gastropod shell strength variation among north temperate lakes . can . j . fish . aquat . sci . 56 : 1687 - 1695 .
liu , h - p , hershler , r . & thompson , f . g . ( 2001
) phylogenetic relationships of the cochliopinae ( rissooidea : hydrobiidae ) : an enigmatic grop of aquatic gastropods . molec . phylog . evol . 21 : 17 - 25 .
relationships between buffering capacity of water and the size and calcium content of freshwater mollusks . freshwat . invertebr . biol . 2 : 48 - 55 .
osenberg , c . ( 1989 ) resource limitation , competition and the influence of life history in a freshwater snail community . oecologia 79 : 512 – 519 .
( say ) ( mollusca , prosobranchia ) from lake st - louis , near montreal ) . can . j . zool . 51 : 311 - 313 .
( gastropoda ) in low alkalinity lakes in south - central ontario . can . j . fish . aquat . sci . 46 : 863 - 869 . molluscan family planorbidae . urbana : university of illinois press .
wilke t . , haase m . , hershler r . , liu h - p . , misof b . , ponder w . ( 2013 )
pushing short dna fragments to the limit : phylogenetic relationships of “hydrobioid” gastropods ( caenogastropoda : rissooidea ) .
please cite as : dillon , r . t . , jr . , b . t . watson , t . w . stewart & w . k . reeves 2006 . the freshwater gastropods of north america . internet address : urltoken
department of biology , college of charleston , charleston , sc 29424 p : 843 . 953 . 8087 f : 843 . 953 . 5453
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the occurrence of at least one species of erhaia in bhutan , viz . erhaia wangchuki sp . n . , is confirmed by dna sequencing . a second unnamed species from bhutan , that might be congeneric , is known from only a single shell . according to the molecular analysis , e . wangchuki is most closely related to a still undescribed erhaia species from china . these two species together with e . jianouensis and akiyoshia kobayashii , both also from china , form a well supported clade . awaiting additional molecular data , the apparent inconsistency regarding erhaia versus akiyoshia is not dealt with here . the extant true sister species of e . wangchuki could be among the four se himalayan species from bhutan and nepal that are classified with erhaia on the basis of conchological data only .
the extremely speciose superfamily rissooidea , with over 400 recent genera ‘one of the largest gastropod families’ ( wilke et al . 2001 : 1 ) , encompasses very many species that cannot be identified on the basis of only shell characters , distribution and ecology . anatomical characters may additionally be used , when the equipment for dissection is available , but even that technically demanding approach does not always bring conclusive results . therefore , molecular analyses are advisable in cases like this . this has resulted in a more reliable classification allowing these snails to be used as ecological indicator species on the one hand and model organisms in historical biogeography on the other hand .
after its description by davis and kuo in davis et al . ( 1985 ) , the genus erhaia turned out to be widespread in asia . from nepal , two species of erhaia are reported by nesemann et al . ( 2007 ) on the basis of shell shape and ecology . here , we report the occurrence of a species of erhaia in bhutan , confirmed by a molecular analysis . the shell of a species that might be congeneric is additionally described but without naming it .
the first author coincidentally collected a minute snail in a marshy source area in bhutan , in the district thimphu , w of geneykha at 2825 m altitude . the locality could not yet be visited again . the specimen got lost after being photographed with a ricoh wg - 4 digital camera , using the extreme macro facility . some equally small snails were found by jigme wangchuk of the ugyen wangchuck institute for conservation and environment , bumthang , in a source in the district wangdue phodrang at 2883 m altitude , and preserved in alcohol 70 % . one specimen ( fig .
) of the latter sample , a paratype , was photographed with a digital microscope system ( keyence vhx - 2000 ; keyence corp . , itasca , il , usa ) and then used for a molecular analysis . another shell , the holotype ( fig .
) , was photographed with a canon eos 7d , using a canon extension tube ef25 and a sigma dc 18 - 50 mm lens .
erhaia wangchuki sp . n . , sequenced paratype ; scale bar 0 . 5 mm ( photographs by b . s . ) ; bhutan , district wangdue phodrang , gangchhu , 2883 m alt . ; 27°26 ' n , 90°11 ' e ; jigme wangchuk leg . 21 . iii . 2015 .
erhaia wangchuki sp . n . , holotype nbcb1013 , measurements 2 . 15×1 . 77 mm ( photographs by e . g . ) ; bhutan , district wangdue phodrang , gangchhu , 2883 m alt . ; 27°26 ' n 90°11 ' e ; jigme wangchuk leg . 21 . iii . 2015 .
maximum likelihood tree based on the 16s rrna dataset of liu et al . ( 2014 ) . numbers on branches denote bootstrap values > 50 .
: 22 ) . the sister taxon of these two species is unclear , but a clade formed by four species , viz .
erhaia daliensis davis & kuo , in davis , kuo , hoagland , chen , yang and chen , 1985 .
; jigme wangchuk leg . 21 . iii . 2015 . national biodiversity centre , serbithang , thimphu [ holotype nbcb1013 , paratypes nbcb1014 / 2 ] .
the gangzetem brooklet ( 5 ) , with the watertank at the source ( 6 ) , and the site where the brooklet crosses the road ( 7 ) . photographs by damber bdr chhetri .
conical , broader than high , with a flat apex because the initial ¾ - 1 whorl is planispiral ; 3¼ whorls in total . body whorl large , the height of the aperture exceeds that of the spire . aperture with a broadly rounded outer lip and a nearly straight parietal side , so that a columellar border is hardly recognizable . growthlines moderately strong , with a more prominent periostracal ridge at more or less regular distances . teleoconch whorls broadly shouldered and separated by a deeply incised suture . aperture oblique ovoid , smooth inside ; apertural edge not touching the penultimate whorl . umbilicus widely open . the holotype is the largest shell and measures 2 . 15×1 . 77 mm .
the shell differs from the shells of the three ‘ erhaia ’ species reported from nepal by nesemann et al . ( 2007 ) by the large body whorl , the relative height of the aperture , and by being broader than high . the other species that are referred to as erhaia in the literature , from areas that are further apart than bhutan and nepal , also have different combinations of character states .
) . the stream bed substrate , viz . pebbles , small rocks and parts of plants , is covered with dark - green algae , housing an abundant diversity of aquatic invertebrates . alongside the brooklet are rhododendrons (
a farm road to the villages of gangphel and zizi crosses over the stream . the source is very close ( ~ 50m ) to that road . the stream also spins a chhukhor , i . e . a water powered prayer wheel . at the very outlet of the stream is a water tank , which supplies drinking water to damchu lhakhang . the brooklet meanders into the phobji main stream , and measures about 1100 meters . during the pre - monsoon ( 21 . 03 . 2015 ) and post - monsoon ( 29 . 11 . 2015 ) , physiochemical properties of the stream were measured . the water is almost neutral ( ph 7 . 06 , 7 . 58 ) and has a nearly stable temperature ( 6 . 76 , 6 . 20 º c ) .
erhaia spec . , measurements c . 2 . 0×1 . 35 mm ( photograph by e . g . ) . bhutan , district thimphu , 4 . 5 km e of chuzom , w of genekha , 2750 m alt . ; 27°19 ' n 89°36 ' e ; e . gittenberger leg . 21 . vi . 2012 .
district thimphu , 4 . 5 km e of chuzom , w of genekha , 2750 m alt . ; 27°19 ' n 89°36 ' e ; e . gittenberger leg . 21 . vi . 2012 .
elongated ovoid , higher than broad , with a last whorl measuring more than ¾ of the total shell height ; aperture attached to the penultimate whorl for less than ⅓ of the parietal - columellar side . umbilicus very narrow . shell height ca . 2 mm .
the shell is most similar in size and shape to ‘ erhaia ’ chandeshwariensis nesemann & sharma , 2007 , and ‘ erhaia ’ banepaensis nesemann & sharma , 2007 , as figured by nesemann et al . ( 2007 : 78 , figs 4–5 ) . erhaia wangchuki sp . n . differs clearly by the broader shell with a lower spire .
erhaia is known by several species in china , from the yangtze river drainage and , by one species , from the mekong river drainage ( davis and kang 1995 , davis and rao 1997 ) . the probability that erhaia is represented in nepal indeed ( nesemann et al . 2007 ) is considerably enhanced by the anatomically confirmed occurrence of e . nainitalensis davis & rao , 1997 , in nainital in northern india west of nepal , and e . wangchuki sp . n . in bhutan east of nepal . it may be hypothesized that there is a radiation in erhaia along the southern border of the himalaya , far west and northwest of its large range in china . with hardly any spring area in bhutan searched for micro - snails , additional species may wait for discovery there at least .
we cordially thank prof . dr . t . wilke ( gießen ) for valuable information regarding erhaia and jigme wangchuk ( bumthang ) for placing interesting specimens at our disposal for research . damber bdr chhetri , forester in phobjikha , is gratefully acknowledged for providing photographs and detailed data about the source and its environment . we are grateful to dr . b . páll - gergely for constructive comments on the manuscript .
davis gm , kuo y - h , hoagland ke , chen p - l , yang h - m , chen d - j . ( 1985 )
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erhaia ( gastropoda : rissooidea ) : phylogenetic relationships and the question of paragonimus coevolution in asia .
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this family is in the clade littorinimorpha ( according to the taxonomy of the gastropoda by bouchet & rocroi , 2005 ) . this family was previously considered to be a subfamily of hydrobiidae .
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genus amnicola - - wisconsin ! genus lyogyrus - - wisconsin ! genus antrobia - - north america . genus antroselates - - north america . genus colligyrus - - north america . genus dasyscias - - north america . genus marstoniopsis genus rachipteron genus baicalia
johnson , p . d . , a . e . bogan , k . m . brown , n . m . burkhead , j . r . cordeiro , j . t . garner , p . d . hartfield , d . a . w . lepitzki , g . l . mackie , e . pip , t . a . tarpley , j . s . tiemann , n . v . whelan & e . e . strong . 2013 . conservation status of freshwater gastropods of canada and the united states . fisheries 38 ( 6 ) : 247 - 282 .
brown , k . m . & c . lydeard . 2010 . mollusca : gastropoda . [ in ] j . h . thorp & a . p . covich ( eds . ) . ecology and classification of north american freshwater invertebrates , 3rd edition : 277 - 307 .
strong , e . e . , o . gargominy , w . f . ponder & p . bouchet . 2008 . global diversity of gastropods ( gastropoda ; mollusca ) in freshwater . hydrobiologia 595 : 149 - 166 . [ available online ]
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picodynastornithes is a clade that contains the orders coraciiformes ( rollers and kingfishers ) and piciformes ( woodpeckers and toucans ) .

this grouping also has current and historical support from the molecular and morphological studies .

picodynastornithes

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yuri , t . ; kimball , r . t . ; harshman , j . ; bowie , r . c . k . ; braun , m . j . ; chojnowski , j . l . ; han , k . - l . ; hackett , s . j . ; huddleston , c . j . ; moore , w . s . ; reddy , s . ; sheldon , f . h . ; steadman , d . w . ; witt , c . c . ; braun , e . l . parsimony and model - based analyses of indels in avian nuclear genes reveal congruent and incongruent phylogenetic signals . biology 2013 , 2 , 419 - 444 .
yuri t , kimball rt , harshman j , bowie rck , braun mj , chojnowski jl , han k - l , hackett sj , huddleston cj , moore ws , reddy s , sheldon fh , steadman dw , witt cc , braun el . parsimony and model - based analyses of indels in avian nuclear genes reveal congruent and incongruent phylogenetic signals . biology . 2013 ; 2 ( 1 ) : 419 - 444 .
yuri , tamaki ; kimball , rebecca t . ; harshman , john ; bowie , rauri c . k . ; braun , michael j . ; chojnowski , jena l . ; han , kin - lan ; hackett , shannon j . ; huddleston , christopher j . ; moore , william s . ; reddy , sushma ; sheldon , frederick h . ; steadman , david w . ; witt , christopher c . ; braun , edward l . 2013 .
parsimony and model - based analyses of indels in avian nuclear genes reveal congruent and incongruent phylogenetic signals .
biology 2 , no . 1 : 419 - 444 .
1 department of neurobiology , howard hughes medical institute ( hhmi ) , and duke university medical center , durham , nc 27710 , usa .
2 department of computer science , the university of texas at austin , austin , tx 78712 , usa .
3 scientific computing group , heidelberg institute for theoretical studies , heidelberg , germany .
5 college of medicine and forensics , xi’an jiaotong university xi’an 710061 , china .
6 centre for geogenetics , natural history museum of denmark , university of copenhagen , øster voldgade 5 - 7 , 1350 copenhagen , denmark .
7 department of biology , new mexico state university , las cruces , nm 88003 , usa .
8 school of biological sciences , university of sydney , sydney , new south wales 2006 , australia .
9 department of ecology and evolutionary biology , university of california , los angeles , ca 90095 , usa .
10 department of biological sciences , louisiana state university , baton rouge , la 70803 , usa .
11 cnrs umr 5554 , institut des sciences de l’evolution de montpellier , université montpellier ii montpellier , france .
12 department of evolutionary biology , evolutionary biology centre , uppsala university , se - 752 36 uppsala sweden .
13 biodiversity and biocomplexity unit , okinawa institute of science and technology onna - son , okinawa 904 - 0495 , japan .
14 department of statistics and institute of bioinformatics , university of georgia , athens , ga 30602 , usa .
15 laboratoire de biométrie et biologie evolutive , centre national de la recherche scientifique , université de lyon , f - 69622 villeurbanne , france .
16 environmental futures research institute , griffith university , nathan , queensland 4111 , australia .
17 bioinformatics and genomics programme , centre for genomic regulation , dr . aiguader 88 , 08003 barcelona , spain .
20 joint institute for computational sciences , the university of tennessee , oak ridge national laboratory , oak ridge , tn 37831 , usa .
21 bioinformatics research centre , aarhus university , dk - 8000 aarhus c , denmark .
22 the genome institute , washington university school of medicine , st louis , mi 63108 , usa .
23 department of biochemistry , molecular biology , entomology and plant pathology , mississippi state university , mississippi state , ms 39762 , usa .
24 institute for genomics , biocomputing and biotechnology , mississippi state university , mississippi state , ms 39762 , usa .
25 department of biological sciences , texas tech university , lubbock , tx 79409 , usa .
26 department of ecology and evolutionary biology , university of california santa cruz ( ucsc ) , santa cruz , ca 95064 , usa .
27 organisms and environment division , cardiff school of biosciences , cardiff university cardiff cf10 3ax , wales , uk .
28 key laboratory of animal ecology and conservation biology , institute of zoology , chinese academy of sciences , beijing 100101 , china .
30 college of medicine and forensics , xi’an jiaotong university xi’an , 710061 , china .
31 state key laboratory for agrobiotechnology , china agricultural university , beijing 100094 , china .
32 department of ecology and evolutionary biology , tulane university , new orleans , la 70118 , usa .
33 museum of natural science and department of biological sciences , louisiana state university , baton rouge , la 70803 , usa .
34 center for zoo and wild animal health , copenhagen zoo roskildevej 38 , dk - 2000 frederiksberg , denmark .
35 department of behavioral neuroscience , oregon health and science university , portland , or 97239 , usa .
36 brazilian avian genome consortium ( cnpq / fapespa - sisbio aves ) , federal university of para , belem , para , brazil .
37 institute of biological sciences , federal university of para , belem , para , brazil .
38 institute of medical biochemistry leopoldo de meis , federal university of rio de janeiro , rio de janeiro rj 21941 - 902 , brazil .
39 centre for biological sequence analysis , department of systems biology , technical university of denmark kemitorvet 208 , 2800 kgs lyngby , denmark .
43 trace and environmental dna laboratory department of environment and agriculture , curtin university , perth , western australia 6102 , australia .
44 department of integrative biology , university of california , berkeley , ca 94720 , usa .
45 laboratory of genomic diversity , national cancer institute frederick , md 21702 , usa .
46 institute of molecular and cellular biology , sb ras and novosibirsk state university , novosibirsk , russia .
47 smithsonian institution national museum of natural history , washington , dc 20013 , usa .
49 department of biological sciences , national university of singapore , republic of singapore .
50 department of vertebrate zoology , national museum of natural history , smithsonian suitland , md 20746 , usa .
51 center for macroecology , evolution and climate , natural history museum of denmark , university of copenhagen , universitetsparken 15 , dk - 2100 copenhagen ø , denmark .
52 bell museum of natural history , university of minnesota , saint paul , mn 55108 , usa .
53 department of life sciences , natural history museum , cromwell road , london sw7 5bd , uk .
54 department of life sciences , imperial college london , silwood park campus , ascot sl5 7py , uk .
55 smithsonian conservation biology institute , national zoological park , front royal , va 22630 , usa .
56 smithsonian conservation biology institute , national zoological park , washington , dc 20008 , usa .
57 theodosius dobzhansky center for genome bioinformatics , st . petersburg state university , st . petersburg , russia 199004 .
58 oceanographic center , nova southeastern university , ft lauderdale , fl 33004 , usa .
59 center for biomolecular science and engineering , ucsc , santa cruz , ca 95064 , usa .
60 san diego zoo institute for conservation research , escondido , ca 92027 , usa .
61 department of vertebrate zoology , mrc - 116 , national museum of natural history , smithsonian institution , washington , dc 20013 , usa .
62 department of environmental health science , university of georgia , athens , ga 30602 , usa .
63 moore laboratory of zoology and department of biology , occidental college , los angeles , ca 90041 , usa .
64 department of genomics and genetics , the roslin institute and royal ( dick ) school of veterinary studies , university of edinburgh , easter bush campus , midlothian eh25 9rg , uk .
65 swedish species information centre , swedish university of agricultural sciences box 7007 , se - 750 07 uppsala , sweden .
66 key laboratory of zoological systematics and evolution , institute of zoology , chinese academy of sciences , beijing 100101 , china .
67 department of organismic and evolutionary biology and museum of comparative zoology , harvard university , cambridge , ma 02138 , usa .
68 institute of theoretical informatics , department of informatics , karlsruhe institute of technology , d - 76131 karlsruhe , germany .
69 department of biochemistry and biophysics , university of california , san francisco , ca 94158 , usa .
70 department of ornithology , american museum of natural history , new york , ny 10024 , usa .
71 department of biology and genetics institute , university of florida , gainesville , fl 32611 , usa .
72 departments of bioengineering and computer science , university of illinois at urbana - champaign , urbana , il 61801 , usa .
73 department of biology , university of copenhagen , ole maaløes vej 5 , 2200 copenhagen , denmark .
74 princess al jawhara center of excellence in the research of hereditary disorders , king abdulaziz university , jeddah 21589 , saudi arabia .
75 macau university of science and technology , avenida wai long , taipa , macau 999078 , china .
77 centre for social evolution , department of biology , universitetsparken 15 , university of copenhagen , dk - 2100 copenhagen , denmark .
science 12 dec 2014 : vol . 346 , issue 6215 , pp . 1320 - 1331 doi : 10 . 1126 / science . 1253451
department of neurobiology , howard hughes medical institute ( hhmi ) , and duke university medical center , durham , nc 27710 , usa .
for correspondence : jarvis @ neuro . duke . edutandywarnow @ gmail . commtpgilbert @ gmail . comwangj @ urltoken zhanggj @ urltoken
department of computer science , the university of texas at austin , austin , tx 78712 , usa .
centre for geogenetics , natural history museum of denmark , university of copenhagen , øster voldgade 5 - 7 , 1350 copenhagen , denmark .
department of biology , new mexico state university , las cruces , nm 88003 , usa .
school of biological sciences , university of sydney , sydney , new south wales 2006 , australia .
department of ecology and evolutionary biology , university of california , los angeles , ca 90095 , usa .
department of biological sciences , louisiana state university , baton rouge , la 70803 , usa .
cnrs umr 5554 , institut des sciences de l’evolution de montpellier , université montpellier ii montpellier , france .
department of evolutionary biology , evolutionary biology centre , uppsala university , se - 752 36 uppsala sweden .
biodiversity and biocomplexity unit , okinawa institute of science and technology onna - son , okinawa 904 - 0495 , japan .
department of statistics and institute of bioinformatics , university of georgia , athens , ga 30602 , usa .
laboratoire de biométrie et biologie evolutive , centre national de la recherche scientifique , université de lyon , f - 69622 villeurbanne , france .
environmental futures research institute , griffith university , nathan , queensland 4111 , australia .
bioinformatics and genomics programme , centre for genomic regulation , dr . aiguader 88 , 08003 barcelona , spain .
joint institute for computational sciences , the university of tennessee , oak ridge national laboratory , oak ridge , tn 37831 , usa .
bioinformatics research centre , aarhus university , dk - 8000 aarhus c , denmark .
the genome institute , washington university school of medicine , st louis , mi 63108 , usa .
department of biochemistry , molecular biology , entomology and plant pathology , mississippi state university , mississippi state , ms 39762 , usa .
institute for genomics , biocomputing and biotechnology , mississippi state university , mississippi state , ms 39762 , usa .
department of biological sciences , texas tech university , lubbock , tx 79409 , usa .
department of ecology and evolutionary biology , university of california santa cruz ( ucsc ) , santa cruz , ca 95064 , usa .
organisms and environment division , cardiff school of biosciences , cardiff university cardiff cf10 3ax , wales , uk .
key laboratory of animal ecology and conservation biology , institute of zoology , chinese academy of sciences , beijing 100101 , china .
college of medicine and forensics , xi’an jiaotong university xi’an , 710061 , china .
state key laboratory for agrobiotechnology , china agricultural university , beijing 100094 , china .
department of ecology and evolutionary biology , tulane university , new orleans , la 70118 , usa .
museum of natural science and department of biological sciences , louisiana state university , baton rouge , la 70803 , usa .
center for zoo and wild animal health , copenhagen zoo roskildevej 38 , dk - 2000 frederiksberg , denmark .
department of behavioral neuroscience , oregon health and science university , portland , or 97239 , usa .
brazilian avian genome consortium ( cnpq / fapespa - sisbio aves ) , federal university of para , belem , para , brazil .
institute of biological sciences , federal university of para , belem , para , brazil .
institute of medical biochemistry leopoldo de meis , federal university of rio de janeiro , rio de janeiro rj 21941 - 902 , brazil .
centre for biological sequence analysis , department of systems biology , technical university of denmark kemitorvet 208 , 2800 kgs lyngby , denmark .
trace and environmental dna laboratory department of environment and agriculture , curtin university , perth , western australia 6102 , australia .
department of integrative biology , university of california , berkeley , ca 94720 , usa .
laboratory of genomic diversity , national cancer institute frederick , md 21702 , usa .
institute of molecular and cellular biology , sb ras and novosibirsk state university , novosibirsk , russia .
smithsonian institution national museum of natural history , washington , dc 20013 , usa .
department of vertebrate zoology , national museum of natural history , smithsonian suitland , md 20746 , usa .
center for macroecology , evolution and climate , natural history museum of denmark , university of copenhagen , universitetsparken 15 , dk - 2100 copenhagen ø , denmark .
bell museum of natural history , university of minnesota , saint paul , mn 55108 , usa .
department of life sciences , natural history museum , cromwell road , london sw7 5bd , uk .
department of life sciences , imperial college london , silwood park campus , ascot sl5 7py , uk .
smithsonian conservation biology institute , national zoological park , front royal , va 22630 , usa .
smithsonian conservation biology institute , national zoological park , washington , dc 20008 , usa .
theodosius dobzhansky center for genome bioinformatics , st . petersburg state university , st . petersburg , russia 199004 .
oceanographic center , nova southeastern university , ft lauderdale , fl 33004 , usa .
center for biomolecular science and engineering , ucsc , santa cruz , ca 95064 , usa .
san diego zoo institute for conservation research , escondido , ca 92027 , usa .
department of vertebrate zoology , mrc - 116 , national museum of natural history , smithsonian institution , washington , dc 20013 , usa .
department of environmental health science , university of georgia , athens , ga 30602 , usa .
moore laboratory of zoology and department of biology , occidental college , los angeles , ca 90041 , usa .
department of genomics and genetics , the roslin institute and royal ( dick ) school of veterinary studies , university of edinburgh , easter bush campus , midlothian eh25 9rg , uk .
swedish species information centre , swedish university of agricultural sciences box 7007 , se - 750 07 uppsala , sweden .
key laboratory of zoological systematics and evolution , institute of zoology , chinese academy of sciences , beijing 100101 , china .
department of organismic and evolutionary biology and museum of comparative zoology , harvard university , cambridge , ma 02138 , usa .
institute of theoretical informatics , department of informatics , karlsruhe institute of technology , d - 76131 karlsruhe , germany .
department of biochemistry and biophysics , university of california , san francisco , ca 94158 , usa .
department of ornithology , american museum of natural history , new york , ny 10024 , usa .
department of biology and genetics institute , university of florida , gainesville , fl 32611 , usa .
departments of bioengineering and computer science , university of illinois at urbana - champaign , urbana , il 61801 , usa .
department of biology , university of copenhagen , ole maaløes vej 5 , 2200 copenhagen , denmark .
princess al jawhara center of excellence in the research of hereditary disorders , king abdulaziz university , jeddah 21589 , saudi arabia .
macau university of science and technology , avenida wai long , taipa , macau 999078 , china .
centre for social evolution , department of biology , universitetsparken 15 , university of copenhagen , dk - 2100 copenhagen , denmark .
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yuri , t ( 2013 ) .
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hackett , s . j . et al . ( 2008 ) .
a phylogenomic study of birds reveals their evolutionary history
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naish , d . ( 2012 ) .
birds .
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kimball , r . t . et al . ( 2013 ) identifying localized biases in large datasets : a case study using the avian tree of life . mol phylogenet evol . doi : 10 . 1016 / j . ympev . 2013 . 05 . 029
jarvis , e . d . ; mirarab , s . ; aberer , a . j . et al . ( 2014 ) .
whole - genome analyses resolve early branches in the tree of life of modern birds
. science 346 ( 6215 ) : 1320–1331 . doi : 10 . 1126 / science . 1253451 .
livezey , bradley c . ; zusi , richard l . ( 2007 ) .
higher - order phylogeny of modern birds ( theropoda , aves : neornithes ) based on comparative anatomy . ii . analysis and discussion
. zoological journal of the linnean society 149 ( 1 ) : 1–95 . pmc 2517308 . pmid 18784798 . doi : 10 . 1111 / j . 1096 - 3642 . 2006 . 00293 . x .
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