this family is in the superfamily truncatelloidea and in the clade littorinimorpha ( according to the taxonomy of the gastropoda by bouchet & rocroi , 2005 ) .

lithoglyphidae

lithoglyphidae is a family of small freshwater snails with gills and an operculum , aquatic gastropod mollusks .
extensive diversification of pebblesnails ( lithoglyphidae : fluminicola ) in the upper sacramento river basin , northwestern united states .
and several other north american genera in the family lithoglyphidae . its anatomy has been described by thompson ( 1984 ) .
two new species of fluminicola ( caenogastropoda , lithoglyphidae ) from southwest oregon , usa , and a range extension for f . multifarius
two new species of fluminicola ( caenogastropoda , lithoglyphidae ) from southwest oregon , usa , and a range extension for f . multifarius
molecular phylogeny of the western north american pebblesnails , genus fluminicola ( rissooidea : lithoglyphidae ) , with description of a new species .
ebscohost | 82975756 | molecular phylogeny of the western north american pebblesnails , genus fluminicola ( rissooidea : lithoglyphidae ) , with description of a new species .
hershler r , liu h - p , hubbart n ( 2017 ) two new species of fluminicola ( caenogastropoda , lithoglyphidae ) from southwest oregon , usa , and a range extension for f . multifarius . zookeys 679 : 1–20 . urltoken
fluminicola was previously hypothesized to be paraphyletic based on morphological data ( hershler & frest , 1996 ) . our findings , although based on a single gene fragment , provide independent , robust evidence that this genus is a composite of two separate lineages , one containing f . virens and the new species described below , and one containing 21 other congeners . the two lineages are genetically divergent ( differing by 17 . 1±1 . 3 % coi sequence divergence ) and do not appear to be closely related ( fig . 1 ) , although additional studies will be needed to resolve further their relationships within the lithoglyphidae . the average coi sequence divergence within clade a is 2 . 4±0 . 3 % ( congeners ranging from 6 . 2 to 6 . 6 % ) and 8 . 0±0 . 6 % within clade b ( congeners ranging from 1 . 5 to 14 . 3 % ) , suggesting that these clades are either fairly old or have accelerated evolutionary rates .
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museum für naturkunde leibniz - institut für evolutions - und biodiversitätsforschung invalidenstr . 43 10115 berlin germany e - mail : fauna - eu ( at ) mfn - berlin . de website : urltoken
» genus pristinicola hershler , frest , johannes , bowler & f . g . thompson , 1994
» genus taylorconcha hershler , frest , johannes , bowler & f . g . thompson , 1994
bouchet p . & rocroi j . - p . ( 2005 ) . classification and nomenclator of gastropod families . malacologia . 47 ( 1 - 2 ) : 1 - 397 isbn 3 - 925919 - 72 - 4 . [ details ]
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in this paper , we further investigate the phylogenetic relationships of fluminicola and test for monophyly of the genus using mitochondrial dna sequence data . this study was prompted in part by the discovery of a new species belonging to the f . virens lineage , which is described herein .
fresh material of the new species was collected by hand and by washing rocks . subsamples were anaesthetized overnight with menthol crystals , fixed in dilute formalin and preserved in 70 % ethanol for morphological study , and in 90 % ethanol for molecular study . shells , radula , opercula and soft anatomy were studied using methods employed in recent taxonomic studies of fluminicola ( hershler & frest , 1996 ; hershler , 1999 ; hershler et al . , 2007 ) .
samples of fluminicola and rissooidean outgroups used for molecular analysis , with codes ( used in fig . 1 , table 2 ) , locality details and genbank accession numbers for coi .
ash creek , south culvert at ash valley road crossing , lassen co . , ca
willow creek at lower end of lower mcbride springs , lassen co . , ca
spring at west end of upper rush creek campground , modoc co . , ca
springs 1 . 8 km east of lower kalama hatchery , cowlitz co . , wa
choctawatchee river , 3 . 2 km north of geneva , geneva co . , al
the posterior distribution of trees was used statistically to test fluminicola monophyly using the methods outlined by weisrock et al . ( 2006 ) and mcmahon , geheber & piller ( 2010 ) . a constraint ( fluminicola monophyly ) created in paup * v . 4 . 0 ( swofford , 2002 ) was used to count the number of bayesian trees that resolved fluminicola as a monophyletic group . monophyly was rejected under a bayesian criterion if < 5 % of the total number of generated trees resolved a clade containing all of the fluminicola sequences .
fifteen specimens of the new species described herein were newly sequenced for this study . five haplotypes were resolved and used in the phylogenetic analysis . the new sequences ( one example of each haplotype per population ) were deposited in genbank under accession numbers jq731609 - jq731615 ( table 1 ) . the alignment of coi sequences yielded 658 bp , of which 263 sites were variable ( 39 . 97 % ) and 242 were parsimony informative ( 36 . 78 % ) . average base frequencies for coi were 27 . 3 % a , 34 . 5 % t , 19 . 2 % c and 19 . 0 % g . base frequencies were homogeneous across all taxa ( χ 2 = 109 . 16 , df = 141 , p = 0 . 98 ) .
the fluminicola sequences were resolved as two evolutionarily distinct , well supported ( ≥95 % posterior probability ) clades in the bayesian tree ( fig . 1 ) . clade a contained f . virens and the new species described herein and was positioned as sister to the rest of the lithoglyphids ; clade b was composed of the rest of the fluminicola sequences and was positioned distally as sister to somatogyrus sp . although the basal branches of the tree were generally poorly supported ( < 0 . 95 posterior probability ) , only 1 , 248 out of 500 , 001 trees ( 0 . 25 % ) delineated fluminicola as a single clade , thus the hypothesized monophyly of this genus can be rejected .
phylogenetic analysis of fuminicola species . bayesian tree based on the coi dataset . posterior probabilities are provided when > 95 % . terminals and specimen codes are labelled as in table 1 .
our results suggest that fluminicola needs to be revised . however , nomenclatural changes cannot be made until the phylogenetic relationships of the type species , f . nuttallianus , are resolved . this species is known only from specimens that were collected in the lower willamette river during the 1800s ( hershler & frest , 1996 ) . although this watershed has been severely degraded by urbanization , pebblesnails of uncertain taxonomic status have been recently collected in many of its streams ( waite et al . , 2008 ) . we are currently reviewing this material to determine whether f . nuttallianus is extant .
the discovery of these two divergent lineages underscores our limited knowledge of the diversity of the western north american component of the lithoglyphid radiation . more than half of the currently recognized species of fluminicola ( 15 / 24 ) have been described since the genus was last reviewed in 1996 ( hershler , 1999 ; hershler et al . , 2007 ) . the overwhelming majority of the pebblesnail populations in western north america are currently unassigned and the recognition of > 40 putatively new fluminicola species in recent contract reports ( e . g . frest & johannes , 1993 , 1995 , 2000a , 2005 ) suggests a considerable amount of undescribed diversity that may possibly include additional divergent lineages .
lithoglyphus nuttallianus — taylor , 1966 : fig . 14 ( clearwater river record ) .
fluminicola sp . — hershler & frest , 1996 : 12 ( salmon river records ) .
types : holotype ( fig . 2 a ) , usnm 905409 , salmon river at pine bar rapids , idaho county , idaho ( n 5082021 , e 551897 ) , coll . 23 april 2001 , coll . dg . paratypes ( from same lot ) usnm 1175400 .
scanning electron micrographs of shells and opercula of fluminicola gustafsoni new species . a . holotype , salmon river at pine bar rapids , idaho county , idaho ( usnm 905409 ) . b . clearwater river at spaulding mission pull - off , nez perce county , idaho ( usnm 883458 ) . c . clearwater river , jim ford creek , clearwater county , idaho ( usnm 1082623 ) . d . salmon river at pine bar rapids , idaho county , idaho ( paratype , usnm 1175400 ) ( shell cleaned with bleach ) . e , f . shell apex , clearwater river , jim ford creek , clearwater county , idaho ( usnm 1082623 ) . g , h . opercula ( outer , inner sides ) , clearwater river , orofino , clearwater county , idaho ( usnm 883460 ) . abbreviation : rm , rim along outer margin . scale bars a–d = 1 . 0 mm ; e , f = 200 µm , g , h = 1 . 0 mm .
etymology : named after daniel l . gustafson ( montana state university ) , who collected much of the material used in this project , including the type material .
diagnosis : a medium - sized fluminicola having a trochoidal to ovate - conic shell and large , gently tapered penis . differs from closely similar f . virens in its smaller size , medium - convex and prominently shouldered teleoconch whorls , weakly angled shell aperture , well developed rectangular wing on the outer marginal radular teeth , vertically oriented coiled oviduct , and in its mtdna sequences ( see remarks below ) .
description : shell trochoidal to ovate - conic , spire usually eroded ( fig . 2 a–d ) ; height 6 . 5–7 . 2 mm ; whorls about 4 . 0 . apex flat , slightly tilted . protoconch of 1 . 5 whorls , diameter 0 . 95 mm ; spiral striae numerous , well developed ( fig . 2 e ) , apical region sometimes also having weak transverse elements ( fig . 2 f ) . teleoconch whorls medium convex , shouldered , sculptured with well developed collabral growth lines . aperture ovate , weakly angled adapically . parietal lip complete , adnate , thin and curved across body whorl , or thickened and nearly straight . columellar lip usually somewhat thickened , straight , columellar shelf usually broad . outer lip usually thin , prosocline . umbilicus absent , umbilical region sometimes excavated . shell clear - white ( fig . 2 d ) , periostracum tan or light brown ( fig . 2 a–c ) . shell parameters are given in table 2 .
operculum thin , light amber , ovate , multispiral , nucleus eccentric , last 0 . 25–0 . 50 whorl weakly frilled ( fig . 2 g ) , outer margin sometimes having weak rim ( fig . 2 h : rm ) . attachment scar margin smooth ( fig . 2 h ) .
radula having about 65 well - formed rows of teeth ( fig . 3 a ) . central tooth ( fig . 2 b , c ) 75 µm wide , cutting edge concave , lateral cusps 3–7 ; median cusp rounded or weakly pointed , usually parallel - sided proximally , broader and longer than laterals ; basal cusps 2–3 , innermost cusp largest ; basal tongue rounded or v - shaped , approximately even with lateral margins . lateral tooth ( fig . 3 d ) face rectangular , outer wing flexed , about 125 % length of cutting edge , central cusp u - shaped , lateral cusps 2–6 ( inner ) , 3–7 ( outer ) . inner marginal teeth ( fig . 3 e ) with 11–17 cusps . outer marginal teeth ( fig . 3 f , g ) with 10–16 cusps ; inner edge of tooth having rectangular wing . radular data are from usnm 883460 and usnm 1175400 .
scanning electron micrographs of radula of fluminicola gustafsoni new species . a . portion of radular ribbon , salmon river at pine bar rapids , idaho county , idaho ( usnm 1175400 ) . b , c . central teeth , salmon river at pine bar rapids , idaho county , idaho ( usnm 1175400 ) ; clearwater river , orofino , clearwater county , idaho ( usnm 883460 ) . d . lateral and inner marginal teeth ( usnm 1175400 ) . e . inner marginal tooth ( usnm 1175400 ) . f , g . outer marginal tooth ( usnm 1175400 , usnm 883460 ) . scale bars a = 100 µm , b = 10 µm , c–g = 20 µm .
snout brown dorsally , ventral surface , small areas around eyes and distal tips pale . foot generally pale , anterior portion brown , sole pale . pallial roof and visceral coil black , pigment somewhat lighter on genital ducts . ctenidium abutting pericardium posteriorly ; ctenidial filaments 30–31 ( n = 5 ) , apices centrally positioned , lateral surfaces weakly ridged . osphradium narrow , elongate , anteriorly hooked , positioned along posterior half of ctenidium . renal organ with large pallial component . hypobranchial gland thin , large , overlapping most of rectum and pallial roof . pedal commissure short . cephalopedal ganglia pale or very lightly pigmented .
renal oviduct having connective tissues pigmented with black granules . ovary composed of 3–4 compound lobes occupying about 0 . 5 whorl , slightly overlapping stomach anteriorly . glandular oviduct and associated structures shown in figure 4 a , b . coiled oviduct ( cov ) narrow , nearly vertical , proximal arm kinked , posterior arm sometimes containing sperm . bursa copulatrix ( bu ) small , globose , partly overlapped by albumen gland ( ag ) . bursal duct ( dbu ) as long or slightly longer than bursa copulatrix , narrow , originating near anterior edge . seminal receptacle ( sr ) small , globose , duct ( dsr ) short , positioned between coiled oviduct and bursa copulatrix , completely overlapped by albumen gland . albumen gland ( ag ) having short pallial component . capsule gland ( cg ) about as long as albumen gland , composed of two glandular zones , ovate in section . rectum forming well - developed furrow on capsule gland ( not shown in fig . 4 a ) . ventral channel ( vc ) narrow . genital aperture ( ga ) a small , subterminal pore .
reproductive anatomy of fluminicola gustafsoni new species , salmon river at pine bar rapids , idaho county , idaho ( usnm 1175400 ) . a . female glandular oviduct and associated structures ( viewed from left side ) . b . bursa copulatrix and seminal receptacle . c . penis , dorsal surface ( pigment darkly stippled ) . abbreviations : ag , albumen gland ; bu , bursa copulatrix ; cg , capsule gland ; cov , coiled oviduct ; dbu , bursa copulatrix duct ; dsr , seminal receptacle duct ; ga , genital aperture ; pa , penial papilla ; pd , penial duct ; pw , posterior wall of pallial cavity ; sr , seminal receptacle ; vc , ventral channel of capsule gland . scale bars ( a–c ) = 500 µm .
testis composed of compound lobes , occupying 1 . 25 whorls , overlapping stomach chambers . seminal vesicle composed of thick coils occupying about 0 . 5 whorl , abutting posterior edge of stomach , opening little behind anterior edge of testis . prostate gland large , pear - shaped , with about 25 % of length in pallial roof . visceral vas deferens opening to ventral edge of prostate gland just behind pallial wall . pallial vas deferens opening from ventral edge of prostate gland in front of pallial wall , duct narrow , undulating . penis ( fig . 4 c ) large , nearly straight , gently tapering . base and median sections deeply folded along inner edge , distal tip weakly pointed . penial duct ( pd ) near centrally positioned , narrow , undulating , opening through small terminal papilla ( pa ) . distal section of penis having dense core of internal black pigment .
distribution and habitat : fluminicola gustafsoni is distributed in the clearwater river ( and its south fork ) , lower salmon river ( downflow from riggins ) , and the reach of the snake river between the mouths of these two streams ( fig . 5 ) . this species was collected in shallow water on rocks and cobbles . it occurs sympatrically with f . fuscus in both the snake river and salmon river .
map showing the geographic distributions of fluminicola gustafsoni new species and f . virens . data for the latter are from hershler & frest ( 1996 ) .
remarks : five haplotypes were detected for f . gustafsoni which differed from each other by 1–4 base pairs ( table 3 ) . one of the haplotypes ( ii ) was observed in three samples , the others were either private alleles ( iii , v ) or singletons ( i , iv ) ( table 3 ) . the five samples of this species that were analysed differed from each other by 0 . 1–0 . 6 % sequence divergence .
fluminicola gustafsoni is distributed well upriver in the columbia river basin from its sister species , f . virens ( fig . 5 ) . these two species differed by 6 . 5±0 . 1 % coi sequence divergence . a close relationship between f . virens and clearwater river pebblesnails was previously postulated by ( former usnm curator ) joseph p . e . morrison , who was asked to identify specimens of the former by parasitologist stewart schell ( university of idaho ) in 1969 . morrison responded that ‘the species you sent from the clearwater river is close to , but not identical to lithoglyphus virens ( lea ) 1838 … it may be a new species ; at the moment i ' m not sure … list it as lithoglyphus cf . virens ( lea ) , and you will be correct’ ( morrison letter to schell , april 9 , 1970 ) .
karolyn darrow ( smithsonian institution ) inked drawings ; yolanda villacampa ( smithsonian institution ) prepared scanning electron micrographs , counted radula cusps and measured shells . peter hovingh assisted the first author in the field . this project was supported , in part , by an award from the idaho department of fish and game ( agreement no . t - 3 - 7 0814 ) . the manuscript was improved by comments provided by david reid and an anonymous reviewer .
final report prepared for u . s . department of agriculture , forest service , pacific northwest region , forest ecosystem management working group , portland , oregon
a baseline mollusk survey of southwestern oregon , with emphasis on the rogue and umpqua river drainages . year 2000 report
springsnails of the cascade - siskiyou national monument and vicinity , oregon . 2004 report
a systematic review of the hydrobiid snails ( gastropoda : rissooidea ) of the great basin , western united states . part ii . genera colligyrus , fluminicola , pristinicola , and tryonia
the life history of neopaleorchis catostomi gen . et sp . n . ( trematoda : monorchiidae ) , an intestinal parasite of the coarsescale sucker , catostomus macrocheilus girard
paup * : phylogenetic analysis using parsimony ( and other methods ) . version 4 . 0b10
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this dataset is made available under the open database license urltoken the open database license ( odbl ) is a license agreement intended to allow users to freely share , modify , and use this dataset while maintaining this same freedom for others , provided that the original source and author ( s ) are credited .
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gravel snail ( lithoglyphus naticoides ) from upper austria ( ri - ver danube between plesching and steyregg ) . picture : jan steger ( source ) .
description : the gravel snail has a bulbous shell with a strong wall coloured a brownish grey green . the apertural whorl dominates the whole spire . the animal itself has a characteristically broad foot . males are often smaller than females . the animal is coloured dark grey in its upper part and lighter grey on the bottom . the body is covered in golden yellow dots and spots .
dimensions : h : 7 - 10 mm ; w : 7 - 10 mm ; n : 4 ½ - 5 . ( abbreviations ) .
habitat and distribution : the gravel snail lives in slow flowing rivers . they can be found on firm muddy ground , on stones and on wood . lithoglyphus naticoides needs a high content of oxygen and calcium carbonate in the water .
the gravel snail originally came from tributaries of the black sea . at least since the pleistocene it has been found in the danube river , where it spread as far west as regensburg . in the neolithic era , gravel snail shells were used for ornamental purposes . only since 1883 gravel snails have also been introduced by ship into the north of central europe , in the rivers rhine and elbe , as well as in the baltic rivers .
threat situation : since 1960 the species is declining fast in central european waters because of water pollution . in saxony - anhalt and baden - württemberg , it is endangered ( en ) , in the rest of germany it is classified as vulnerable ( vu ) . in some places , though , the populations are recovering ( see also : iucn threat categories ) .
1828 ) - oder wie aus einem inländer ein zuwanderer wurde
. nachr . bl . d . e . vorarlb . mal . ges . , 19 , p . 27 ff .
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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 .
johnson , p . d . , bogan , a . e . , brown , k . m . , burkhead , n . m . , cordeiro , j . r . , garner , j . t . , hartfield , p . d . , lepitzki , d . a . w . , mackie , g . l . , pip , e . , tarpley , t . a . , tiemann , j . s . , whelan , n . v . and strong , e . e . 2013 . conservation status of freshwater gastropods of canada and the united states . fisheries 38 ( 6 ) : 247 - 282 .
( taylor , 1966 ) , but is considered a valid species by hershler and frest ( 1996 ) .
dyer , e . , soulsby , a . - m . , whitton , f . , mcguinness , s . , de silva , r . , milligan , h . t . , kasthala , g . , herdson , r . , thorley , j . , mcmillan , k . & collins , a .
justification : fluminicola dalli has been assessed as data deficient due to insufficient information on the species population trends . it is therefore not possible to accurately assess the effects of increasing salinity and water abstraction on the species . further research is required to clarify whether this constitutes a serious threat before the species can be accurately listed .
this species is only known from the type locality and a small surrounding area in the northern area of pyramid lake , washoe county , nevada ( hershler and frest 1996 ) . the extent of occurrence of this species is estimated as no more than 3 , 050 km² .
this species has been collected from small streams ( hershler and frest 1996 ) .
the salinity of pyramid lake has risen from 3 . 75 to 5 . 5 g l - 1 between 1933 and 1980 ( williams 1999 ) , likely as a result of ground water abstraction for agriculture , upstream water diversion programs , and decreased rain fall . the species is found in an area of decreasing water availability with increased salinities as development progresses , however , the species has been collected repeatedly over the years , is still present and often referred to as common ( j . cordeiro pers . comm . 2011 ) . therefore there is no real evidence that the salinity change has affected the population , although precise data on the population effects of increased salinity is lacking .
this species has been given a global heritage status rank of g1 - critically imperilled ( natureserve 2009 ) . there is insufficient information regarding population numbers to assess the effect of increased salinities on this species , so that further research is recommended .
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1 department of invertebrate zoology , smithsonian institution , washington , d . c . 20013 - 7012 , usa ,
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) from southwest oregon based on morphologic and mitochondrial dna ( coi , cytb ) evidence .
prior to the recent restriction of the latter to the lower columbia river drainage , lives in lotic habitats in the umpqua river basin . this species is readily distinguished from closely related
ranges among lotic habitats in the north umpqua river basin , and in the upper rogue river drainage north of little butte creek . this species differs from other congeners by > 9 . 1 % for both genes and is distinguished from closely similar and geographically proximal
from the upper rogue river basin south of little butte creek , which extend the geographic range of this species about 80 km northward from the sacramento river headwater region . this continues a recent series of taxonomic papers on the poorly known and little studied pebblesnail fauna of the vast pacific watershed from northern california to southern british columbia .
the broad geographic range of fluminicola includes most of the pacific coastal watershed from northern california to southern british columbia . pebblesnail populations are scattered throughout much of this huge area , yet are currently undescribed aside from six species in the columbia river basin ( hershler and frest 1996 , hershler and liu 2012 , liu et al . 2013 ) and 14 species in the sacramento river basin ( hershler et al . 2007 ) . this taxonomic knowledge gap is hampering efforts by the conservation community to obtain legal protection for pebblesnails ( e . g . , usfws 2012 ) , which are groundwater - dependent and threatened by various anthropogenic activities .
the fluminicola fauna of southwestern oregon includes a relatively large pebblesnail in the umpqua river basin that was traditionally identified as f . virens prior to the restriction of the latter to the lower columbia river drainage ( hershler and frest 1996 ) , and numerous unstudied populations of smaller pebblesnails in the upper reaches of both the rogue and umpqua river basins that were recently reported in grey literature ( frest and johannes 1999 , 2000 , 2004 , 2005 ) . herein we utilize dna sequences from two mitochondrial genes in delineating the fluminicola species in the rogue - umpqua basins . this continues our recent series of integrative taxonomic studies of the pebblesnails of the pacific coastal drainages ( hershler et al . 2007 , hershler and liu 2012 , liu et al . 2013 ) .
we sequenced specimens from 35 sites in the rogue - umpqua basins , including eight localities containing the large pebblesnail resembling f . virens and 27 localities containing smaller pebblesnails . thirty - two of these localities were sampled by rh and hpl during september 2015 specifically for this project . specimens were collected by hand or with a small sieve and preserved in 90 % ( non - denatured ) ethanol in the field . vouchers were deposited in the smithsonian institution’s national museum of natural history ( usnm ) collection . other relevant material from the usnm and the bell museum of natural history ( bellmnh ) was also examined during the course of this study .
a total of 155 and 146 sequences of cytochrome c oxidase subunit i ( coi ) and cytochrome b ( cytb ) , respectively were obtained from 161 analyzed rogue - umpqua pebblesnails . genomic dna was extracted from entire snails using a ctab protocol (
) were used to amplify a 708 base pair ( bp ) fragment of coi ; cytb427f ( 5’tga ggk gcn act gtt att act aa3’ ) and cytb1049r ( 5’gtg aaa act tgs ccr att tgc tc3’ ) were used to amplify a 644 bp fragment of the cytb gene . the cytb427f and cytb1049r primers were designed based on conserved regions of cytb in an alignment using previously published sequences from
. sequences were determined for both strands and then edited and aligned using sequencher™ version 5 . 4 . 1 ( gene codes corporation , ann arbor , mi ) . our analysis of the coi dataset also included 35 previously published sequences from 23 congeners , two taxonomically indeterminate
) . in order to generate easily readable topologies , one example of each detected haplotype was used in the phylogenetic analyses , which were performed separately for the coi and cytb datasets . sample codes , locality and voucher details , and genbank accession numbers for the sequences used in the molecular phylogenetic analyses are in suppl . material
large adult females were used for shell measurements . the total number of shell whorls ( wh ) was counted for each specimen ; and the height and width of the entire shell ( sh , sw ) , body whorl ( hbw , wbw ) , and aperture ( ah , aw ) were measured from camera lucida outline drawings ( hershler 1989 ) . descriptive statistics were generated using systat for windows 11 . 01 ( ssi 2004 ) . other methods of morphological study were routine ( hershler et al . 2007 ) .
forty - one coi and 55 cytb haplotypes were detected in the analyzed specimens from the rogue - umpqua basins ( suppl . material
, respectively ) . the molecular phylogenetic analyses of both the coi and cytb datasets consistently resolved the rogue - umpqua haplotypes into three distinct clades . the trees generated by the three methods of phylogenetic analysis were closely similar ; the bayesian topology based on the coi dataset is shown in figure
; rogue - umpqua haplotype codes ( in parentheses ) are from suppl . material
) contained the smaller rogue - umpqua pebblesnails . clade a , which was weakly supported generally , but well supported in the cytb bayesian topology ( 100 % posterior probability ) , was composed of the haplotypes from populations in the upper rogue river basin to the south of little butte creek ( fig .
( 2 . 6 % for coi and 3 . 6 % for cytb ,
) was slightly greater than the variation within these two groups ( 2 . 2 and 1 . 4 % for coi ; and 3 . 5 and 2 . 4 % for cytb , respectively ) and falls into the range of pairwise differences among currently recognized
species : 1 . 4–18 . 7 % for coi and 1 . 6–25 . 7 % for cytb (
) . however , we could not differentiate these geographically disjunct yet phylogenetically intermixed groups of populations based on morphologic criteria and thus are treating them as conspecific .
clade b , which was well supported in all but the cytb bayesian analysis , is composed of the small pebblesnails in the north umpqua basin , and the upper rogue river basin north of little butte creek ( fig .
by several anatomical characters . based on the sum of this evidence we recognize clade b as a somewhat variable new species which is described below .
the third clade ( c ) , which was moderately supported in most of the analyses and well supported in the coi ml tree ( 100 % ) , contained the haplotypes detected in the large pebblesnails from the umpqua river basin ( fig .
) . the sequence divergence within this clade was slight – 0 . 3 % for coi and 0 . 6 % for cytb , respectively . clade c , in turn , formed a well - supported monophyletic group with
, and is most similar genetically to the latter , from which it differs by 3 . 6 % for coi and 3 . 9 % for cytb . the large pebblesnail in the umpqua river basin is also readily differentiated morphologically from both
, and thus we recognize it as a new species which is described below .
shells , opercula and radula , f . multifarius . a shell , usnm 1144567 b shell , usnm 1144997 c shell , usnm 1145051 d , e opercula ( outer , inner sides ) , usnm 1144567 f portion of radular ribbon , usnm 1144567 g central teeth , usnm 1144567 h lateral teeth , usnm 1144567 . scale bars : a–b 1 . 0 mm ; d–e 250 µm ; f , 100 µm ; g–h 10 µm .
reproductive anatomy . a–d f . multifarius a , b , d usnm 1144567 , c usnm 1070753 e–g f . fresti , sp . n . , usnm 1422223 h–j f . umpquaensis , sp . n . , usnm 1144736 . a , e , g female glandular oviduct and associated structures ( viewed from left side ) b , c , f , i bursa copulatrix and seminal receptacle , d , g , j penis , dorsal surface . the bursa copulatrix and seminal receptacle of type material of f . multifarius ( c ) , which was not illustrated by hershler et al . ( 2007 ) , is shown for comparative purposes . scale bars , 500 µm . ag albumen gland bu bursa copulatrix cg capsule gland cov coiled oviduct dsr seminal receptacle duct ga genital aperture pd penial duct pw posterior wall of pallial cavity sr seminal receptacle vc ventral channel of capsule gland .
fluminicola multifarius . – hershler et al . 2007 : 415 , 417 , 419 , figs . 7m , 24 , 25 ( big springs ( source ) at big springs city park northwest of the city of mount shasta , south of spring hill , siskiyou county , california ( [ utm zone 10 ] 556400 e , 4575265 n , 1092 m ) .
sacramento river headwater region ( as far downflow as conant ) , and a few sites in the upper reaches of the mccloud river drainage ( hershler et al . 2007 ) .
california . siskiyou county . usnm 1070753 , big springs ( source ) at big springs city park northwest of the city of mount shasta , south of spring hill , usnm 1020771 , bundoora spring , west of access road off fs40n44 .
in all respects aside from a slightly larger maximum shell height ( 5 . 02 vs . 4 . 64 mm ) . representative rogue basin specimens are illustrated in figures
( shells ) , 3d–e ( opercula ) , 3f–h ( radula ) and 4a , b , d ( reproductive anatomy ) . ten coi and cytb haplotypes were detected in the rogue basin populations ( suppl . material
is also distributed in the intervening klamath river basin ; the pebblesnail fauna of this large watershed is currently undescribed . populations of
shells , opercula and radula , f . fresti , sp . n . a holotype , usnm 1144376 b , c sympatric ovate - conic and trochiform shell morphs , respectively , usnm 1297149 d shell , usnm 1297155 e , f opercula ( outer , inner sides ) , usnm 1422223 g portion of radular ribbon , usnm 1422223 h central teeth , usnm 1422223 i lateral teeth , usnm 1422223 . scale bars : a–d 1 . 0 mm ; e–f 250 µm ; g 100 µm ; h–i 20 µm .
holotype , usnm 1144376 , diversion from big butte springs through butte falls hatchery , just south of butte falls - fish lake road ( jackson county 321 ) and 0 . 16 km west of butte falls - prospect road ( jackson county 922 ) , jackson county , oregon , 42 . 5389°n , 122 . 5551°w , 10 / 22 / 1994 , terrence j . frest and edward j . johannes . paratypes , usnm 1422223 ( a large series of dry shells and alcohol - preserved specimens ) , from same lot .
a small to medium - sized fluminicola ( 2 . 3–5 . 5 mm shell height ) having a trochoidal to ovate - conic shell and small , gently tapered penis . differs from closely similar and geographically proximal f . multifarius in the hooked shape of the anterior end of the osphradium , larger number of ctenidial ( gill ) filaments , smaller seminal receptacle , and in its mtdna sequences .
) trochoidal to narrow - conic , whorls 3 . 5–4 . 0 . teleoconch whorls medium convex , sometimes weakly shouldered . aperture ovate , slightly angled above ; inner lip complete , variably thickened and reflected , sometimes forming a rather wide parietal - columellar shelf that sometimes covers the umbilical region . outer lip thin , prosocline . umbilicus very small or absent , umbilical region sometimes excavated . shell white , periostracum brown , sometimes covered with thick black deposits . shell measurements and whorl count data are summarized in table
) as for genus ; muscle attachment margin thickened on inner side . radula ( fig .
) as for genus ; dorsal edge of central teeth concave , lateral cusps two–five , basal cusp one–two . lateral teeth having two–three cusps on inner side and three–four cusps on outer side ; length of outer wing 175–185 % length of cutting edge . inner marginal teeth with 23–31 cusps , outer marginal teeth with 27–40 cusps . radula data are from
snout , cephalic tentacles , pallial roof , visceral coil usually medium pigmented ( brown ) ; foot varying from near pale to medium pigmented along anterior edges . distal section of penis having dense core of internal black pigment . ctenidial filaments 21–24 (
= 5 ) , lateral surfaces smooth . anterior end of osphradium distinctly hooked ( not illustrated ) . glandular oviduct and associated structures shown in figure
. coiled oviduct circular , proximal arm kinked , posterior arm sometimes having small pouch containing sperm . bursa copulatrix large , reniform , partly overlapped by albumen gland . bursal duct slightly shorter than bursa copulatrix , narrow . seminal receptacle small , sac - like , almost completely overlapped by albumen gland . albumen gland having short pallial component . capsule gland longer than albumen gland , composed of two glandular zones . genital aperture a small , sub - terminal pore . penis ( fig .
) small , slightly curved , gently tapering , distal section abruptly narrowing to small pointed tip . medial section having a few weak folds along inner edge . penial duct near centrally positioned , straight , narrow .
this species name is a patronym ( in the genitive singular ) honoring recently deceased malacologist terrence frest for his many contributions to the documentation of molluscan biodiversity in the northwestern united states .
fluminicola fresti is distributed in spring - fed habitats in the north umpqua river drainage and in the rogue river basin north of little butte creek .
vary in overall shape and in the width of the inner apertural lip . although this variation is generally continuous in the material that we examined , two rather distinct forms – ovate - conic with a narrow inner apertural lip ( fig .
) – can be identified in one of the springs in joseph stewart state park . we sequenced specimens from this locality and found that the two shell forms ( samples ru18 and ru17 , respectively ) differed by 0 . 7 % for coi and 1 . 3 % for cytb , which was less than the variation within these two groups ( 0 . 2 and 1 . 2 % for coi ; and 0 . 2 and 1 . 5 % for cytb , respectively ) . the scant genetic differentiation of the two shell forms when in sympatry provides additional evidence that they are conspecific . additional studies incorporating rapidly evolving nuclear markers ( e . g . , microsatellites ) may help sift through the possible explanations for the interesting variation in the shells of this species ( e . g . , ecophenotypic plasticity , incipient speciation ) . twenty - three coi haplotypes and 34 cytb haplotypes were detected in the sequenced specimens of
populations of f . fresti were referred to as the beaverdam pebblesnail , camp creek pebblesnail , clark pebblesnail , evergreen pebblesnail , rogue pebblesnail , stewart pebblesnail , and umpqua pebblesnail by frest and johannes ( 1999 , 2000 , 2004 , 2005 ) . in order to avoid confusion , we suggest that “frest’s pebblesnail” be used as the common name for f . fresti .
shells , opercula and radula , f . umpquaensis , sp . n . a holotype , usnm 1144535 b shell , usnm 1144736 c shell , usnm 1144371 d , e opercula ( outer , inner sides ) , usnm 114535 f portion of radular ribbon , usnm 1422224 g central teeth , usnm 1422224 h lateral teeth , usnm 1422224 . scale bars : a–e , 1 . 0 mm ; f , 100 µm ; g–h , 10 µm .
fluminicola virens . – pilsbry 1899 : 122 - 123 ( in part ) . henderson 1929 : 169 ( in part ) . burch and tottenham 1980 : 102 ( in part ) .
holotype , usnm 1144535 , umpqua river at bunch bar access ( county ) , west of or38 ( umpqua highway ) , douglas county , oregon , 43 . 6462°n , 123 . 6670°w , 9 / 27 / 1998 , terrence j . frest and edward j . johannes . paratypes , usnm 1422224 ( a large series of dry shells and alcohol - preserved specimens ) , from same lot .
a large fluminicola ( maximum shell height , 9 . 5 mm ) having a conical shell with eroded spire and a broad , little tapered penis . differs from closely related f . gustafsoni and f . virens in its reniform - shaped bursa copulatrix and in its mtdna sequences . further differentiated from f . gustafsoni by its more elongate shell , longer outer wing of the lateral radular teeth , and smaller seminal receptacle ; and from f . virens by its more convex shell whorls and longer bursal duct .
) conical , spire usually eroded , whorls > 3 . 5 . teleoconch whorls medium convex , narrowly shouldered . aperture ovate , slightly angled above ; inner lip complete , thickened . outer lip thin , prosocline . umbilicus absent . shell white , periostracum tan or olive . shell measurements and whorl count data are summarized in table
) as for genus ; dorsal edge of central teeth concave , lateral cusps four–five , basal cusp two–four . lateral teeth having three–six cusps on inner side and four–six cusps on outer side ; length of outer wing 180–200 % length of cutting edge . inner marginal teeth with 13–18 cusps , outer marginal teeth with 11–17 cusps . radula data are from
head - foot rather lightly pigmented , cephalic tentacles with central brown longitudinal stripe along length . pallial roof , visceral coil dark brown , almost black . penis having dense core of internal black pigment along penial duct . ctenidial filaments 34–36 (