skistodiaptomus is a genus of freshwater copepods in the family diaptomidae , found across north america .

the genus contains eight species , three of which are endemic to the united states and are listed on the iucn red list as vulnerable species ( vu ) or data deficient ( dd ) .

skistodiaptomus bogalusensis ( m. s. wilson & moore , 1953 ) skistodiaptomus carolinensis yeatman , 1986 skistodiaptomus mississippiensis ( marsh , 1894 ) skistodiaptomus oregonensis ( lilljeborg , 1889 ) skistodiaptomus pallidus ( herrick , 1879 ) skistodiaptomus pygmaeus ( pearse , 1906 ) skistodiaptomus reighardi ( marsh , 1895 ) skistodiaptomus sinuatus ( kincaid , 1953 )

skistodiaptomus

skistodiaptomus <EOT> dowell , k . m . 1997 . evidence for diapause in the freshwater copepod skistodiaptomus pallidus . american midland naturalist 137 ( 2 ) : 362 - 368 .

thum , r . a . and r . s . stemberger . 2006 . pure spatial and spatially structured environmental variables explain skistodiaptomus copepod range limits in the northeastern usa . canadian journal of fisheries and aquatic sciences 63 ( 6 ) : 1397 - 1404 .

table 1 . states with nonindigenous occurrences , the earliest and latest observations in each state , and the tally and names of hucs with observations† . names and dates are hyperlinked to their relevant specimen records . the list of references for all nonindigenous occurrences of skistodiaptomus pallidus are found here .

ghan , d . , j . d . mcphail , and k . d . hyatt . 1998 . the temporal - spatial pattern of vertical migration by the freshwater copepod skistodiaptomus oregonensis relative to predation risk . canadian j . of fisheries and aquatic sci . 55 : 1350 - 1363 .

chapman , m . a . , j . d . green , and t . g . northcote . 1985 . seasonal dynamics of skistodiaptomus pallidus herrick and other zooplankton populations in deer lake , s . w . british columbia . journal of plankton research 7 ( 6 ) : 867 - 876 .

duggan , i . c . , j . d . green , and d . f . burger . 2006 . first new zealand records of three non - indigenous zooplankton species : skistodiaptomus pallidus , sinodiaptomus valkanovi , and daphnia dentifera . new zealand journal of marine and freshwater research 40 : 561 - 569 .

potential : skistodiaptomus pallidus is an efficient omnivorous predator , with the ability to prey on preferred rotifers and microzooplankton from large distances . it also consumes algae and practices cannibalism , which may allow populations to persist when resource availability is low ( williamson and butler 1986 ; williamson and vanderploeg 1988 ) . it has also been known to attain very high densities in suitable habitats , reaching 10 , 000 per m3 in a lake erie marsh to unknown consequences ( krieger and klarer 1991 ) . skistodiaptomus pallidus became the primary calanoid copepod in a particularly eutrophic portion of lake tahoe , dominating two previously common species , leptodiaptomus tyrrelli and epischura nevadensis ( byron and saunders 1981 ) .

kipp , r . m . , a . j . benson , j . larson , t . h . makled , and a . fusaro , 2018 , skistodiaptomus pallidus herrick , 1879 : u . s . geological survey , nonindigenous aquatic species database , gainesville , fl , urltoken revision date : 5 / 2 / 2018 , access date : 7 / 9 / 2018

duggan , i . c . , neale , m . , robinson , k . v . , verburg , p . , & watson , n . t . n . ( 2014 ) . skistodiaptomus pallidus ( copepoda : diaptomidae ) establishment in new zealand natural lakes , and its effects on zooplankton community composition . aquatic invasions , 9 ( 2 ) , 195–202 . urltoken

after s . oregonensis molts to the first copepodid stage , it can filter feed on the same size class of particles ( 100 - 800ám│ ) as the adults ( mcqueen , 1970 ; comita and mcnett 1976 ) . the copepod prefers larger phytoplankon ( mcqueen , 1970 ) . richman ( 1966 ) showed that s . oregonensis exhibited it maximum ingestion rate in food concentrations of 25 000 - 50 000 cells ml‾╣ , ingestion rate declined dramatically . skistodiaptomus oregonensis may be adaped to feeding on large cells in high concentration , and would be at a feedin disadvange in lakes of very low productivity .

the north american calanoid copepod skistodiaptomus pallidus is an emerging invader globally , with non - indigenous populations recorded from constructed waters in new zealand , germany and mexico since 2000 . we examined the effects of s . pallidus establishment on the zooplankton community of a natural lake , lake kereta , where it was first recorded in late - 2008 , coincident with releases of domestically cultured grass carp ( ctenopharyngodon idella ) . although not present in any of our samples prior to august 2008 , s . pallidus was found in all samples collected in the subsequent five years . anosim indicated zooplankton community composition significantly differed between samples collected before and after s . pallidus invasion , whether the invader was included in the analysis or not . zooplankton species affected most greatly were the copepods calamoecia lucasi and mesocyclops sp . , which decreased in their relative importance , and the cladocerans bosmina meridionalis and daphnia galeata , which increased . rotifer species were relatively unaffected . as the length of grass carp released were > 6 . 5 cm , direct predatory effects by this species on the zooplankton community are unlikely . associated reductions in macrophyte biomass could explain increases in the relative abundances of planktonic cladocerans ( b . meridionalis and d . galeata ) . however , the effect of macrophyte reduction by grass carp on zooplankton communities is considered to be limited elsewhere , while the reduced macrophyte biomass cannot explain the decrease in relative abundance of the native planktonic calanoid copepod c . lucasi . competition between c . lucasi and s . pallidus is the most compelling explanation for the reduction in importance of the native calanoid copepod species . skistodiaptomus pallidus appears to have undergone a “boom - and - bust” cycle in lake kereta , increasing in relative abundance in the first three years following establishment , before declining in importance .

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first described in 1986 from lake ravenel , highlands , macon county , north carolina ( in the upper little tennessee watershed ) . has since also been found in ponds bordering the great smoky mountains national park .

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in the great lakes zooplankton literature , this sepcies was consistently referred to as diaptomus oregonensis until the subgenus name skisodiaptomus was elevated to genetric level .

in lake superior the females range from 1 . 2 - 1 . 4mm and average 1 . 33mm . males were somewhat smaller , ranging from 1 . 1 - 1 . 3mm and averaging 1 . 19mm .

the right antennule of adult male calanoids is geniculate and may possess a lateral projection on the antepenultimate segment . lateral spine on terminal segment of exopode of right 5th leg is subterminal in position .

s . oregonensis is present year round in the great lakes ( jahoda 1948 ; carter 1969 ; wilson and roff 1973 ; gannon 1975 ) . it is most abundant in the summer and fall but peaks have been observed from april to december . gannon ( 1972a , 1974 ) reported abundance peaks as early as february in green bay .

this species produces two generations each year in lake michigan . adults overwinter and produce a slow - growing spring generation that coincides with the phytoplankton maximum . this generation reproduces in the summer , and the offspring grow rapidly , maturing by fall overturn . most animals wait until spring to reproduce although gannon ( 1972a ) reported some winter reproduction in green bay . in lake superior , where abundance is low , s . oregonensis only produces one generation each year and overwinters as resting eggs ( selgeby 1974 )

sex rations of 1 : 1 are most common ( jahoda 1984 ; davis 1962 , 1968 ; torke 1975 ) , but females may outnumber males ( stewart 1974 ) . clutch size appears to be positively correlated with size of the females ( davis 1961 )

reproduction in the s . oregonensis is sexual . the male clasps the female with his modified first antennae and / or fifth legs , then transfers a packet of sperm , the spermatophore , from his genital pore to her genital segment . the sperms are stored in a seminal receptacle located in the female ' s genital segment . when the female releases eggs from her genital tract , they are fertilized by the stored sperm . female brood the eggs in one egg sac attached to the genital segment . after the first clutch of eggs has hatched , some females fertilize a second and third clutch utilizing more of the sperm stored from their first mating .

its egg hatch into small , active larva known as nauplii . the first nauplius stage ( ni ) is characterized by three pairs of appendages . the animals grow rapidly and molt to the second nauplius stage ( nii ) , which is slightly longer . the animals continue to grow and add appendages as they pass through six naupliar stages . the next molt is to the first copepodid stage ( ci ) . at this period the young species have the general body shape of the adult but are smaller and lack several of the swimming legs . growth , addition of swimming legs , and modification of the limbs continue at each molt until the adult ( cvi ) stage is reached . the animals then mate and produce the next

is one of the most widely distributed diaptomid species , occuring throughout the northern united states and southern ontario , canada ( marsh 1893 , 1929 ; carl 1940 ) . this species has been reported in all the great lakes , especially abundant in the warmer , southerly portions of the

and is of decreasing importance in the cooler , northerly regions ( robertson 1966 ) . now this species is rare in lake superior ( selgeby , 1975 ) and lake ontario , only appearing in fall and most common in lake erie ( davis 1961 , wright 1955 ) which peaks in abundance occur in september .

s . oregonensis always inhabits permanent water bodies , but occurs in lakes of different morphometric and tropic characteristics . it is common in lake michigan , but also occurs in quite shallow inland lakes and ponds . it frequently occurs with other calanoids , leptodiaptomus minutusis , l . ashlandi and l . sicilis , and occasionally with s . pallidus .

s . oregonensis is more abundant offshore than in the littoral zone ( stewart 1974 ) . during periods of thermal stratification , this species is usually found in the meta - and epilimnia , migrating toward the surface at night , but most densely concentrated near the thermocline during the day ( juday , 1903 ; langford , 1938 ; wells 1960 ) . in unstratified water masses and upwelling areas , distribution throughout the water column is more uniform ( wilson and roff 1973 ) . in teapot lake , ontario , cooley ( 1970 ) found that day - night differences in mean depth distribution were not significant for naupliar stages , but were significant for copepodid and adult stage , although these day - night distribution means only involved a depth difference of 1m or less , and some animals apparently did not migrate . birge ( 1895 ) observed no vertical movement for this species in lake mendota , dane co . diurnal migratory behavior is apparently weakly developed in this species , and the extent of migration may be influenced by age and individual behavior , and perhaps by temperature , light level , time of year and thermocline depth

s . oregonensis has been found in the stomach of severl species of fish including ass , crappie , carp , suckers , freshwater drum , trout - perch , yellow perch , and whitefish ( ewers 1933 ; wilson 1960 ) . comparing only a small percentage of the diet of some fish , it was found to be an important food item for small bloater ( wells and beeton 1963 ) .

balcer , korda , and dodson . 1984 . zooplankton of the great lakes : general morphology and ecology of the crustacean zooplankton , p . 8 - 11 . university of wisconsin pres . madison , wisconsin .

balcer , korda , and dodson . 1984 . zooplankton of the great lakes : life history and ecology of the major crustacean species , p . 91 - 93 . university of wisconsin pres . madison , wisconsin .

byron torke . 2001 . the distribution of calanoid copepods in the plankton of wisconsin lakes . hydrobiologia 453 / 454 : 351 - 365 , 2001 .

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the nonindigenous occurrences section of the nas species profiles has a new structure . the section is now dynamically updated from the nas database to ensure that it contains the most current and accurate information . occurrences are summarized in table 1 , alphabetically by state , with years of earliest and most recent observations , and the tally and names of drainages where the species was observed . the table contains hyperlinks to collections tables of specimens based on the states , years , and drainages selected . references to specimens that were not obtained through sighting reports and personal communications are found through the hyperlink in the table 1 caption or through the individual specimens linked in the collections tables .

the endopod of the 1st leg of this copepod species is bifurcate . unlike some other diaptomids , the antepenultimate segment of the right 1st antenna has no distinct appendage and is not produced into a blunt point in males . the male left 5th leg is shorter than the right , reaching to the end of or slightly past the 1st segment of the right exopod . there is a scythe - like inner process on the terminal exopod segment of the male left 5th leg . females have 3 urosomal segments , rounded metasomal wings with small sensilla , and a somewhat expanded genital segment ( pennak 1989 ; lesko et al . 2003 ) .

females are around 0 . 8–1 . 2 mm long while males are around 0 . 7–1 mm long ( geddes and cole 1981 ; torke 2001 ; lesko et al . 2003 ) .

is native to the north central plain states , northeast to new york , southern united states in the mississippi river basin , texas , and west to colorado ( pennak 1989 ; mills et al . 1993 ; torke 2001 ; thum and sternberger 2006 ) .

was first recorded from lake ontario in 1967 . in the 1970s , it was recorded from lake erie , lake st . clair , lake huron and the saginaw river . the occurrences in the lake huron drainage were discovered upon re - examining archived specimens from 1974 and 1975 ( mills et al . 1993 ; torke 2001 ; lesko et al . 2003 ) . unspecified locations in

( mclaughlin et al . , 2005 ) . collected from lake tahoe , on the

border in 1979 and several other california locations ( byron and saunders 1981 ) .

has expanded its range in north america westward ( duggan et al . 2006 ) . it has been introduced outside of north america , possibly via the aquarium trade .

* hucs are not listed for states where the observation ( s ) cannot be approximated to a huc ( e . g . state centroids or canadian provinces ) .

is typically found in waters with a ph range of 7 . 5–8 . 6 and a conductivity range of 77–660 μs cm

. this species prefers to dwell in cool waters < 12ºc and was found at 10 m in june and below 16 m in july , underneath the thermocline in belews lake , north carolina ( chapman et al . 1985 ; marcogliese and esch 1992 ; torke 2001 ; thum and stemberger 2006 ) .

produces some eggs that go through a diapause stage before they hatch and some that do not . breeding in general takes place from around march to november and females can produce up to around 20 eggs per brood . diapausing eggs produced between june and october hatch from december to june of the following year . when production of diapausing eggs occurs in the summer it may aid populations to avoid fish predation . these eggs have been known to reach densities of 105 per m

in the sediments of some habitats . such high densities may help mitigate the effects of a poor year in reproduction and recruitment . it may take around 7 weeks for resting stages to hatch . one study found that it takes around 66 days at 10ºc and 15 days at 25ºc for development to the adult stage . there are 2–5 generations per year in different parts of this species’ range ( geiling and campbell 1972 ; chapman et al . 1985 ; dowell 1997 ; torke 2001 ; lesko et al . 2003 ; wonham et al . 2005 ) .

feeds on phytoplankton , especially individual algae > 53 μm in size . it can also selectively and intensely prey on some rotifer species ( geiling and campbell 1972 ; williamson and butler 1986 ; torke 2001 ) .

could have been introduced accidentally in bait buckets , in fishing equipment , by recreational boaters , with hatchery stock from the mississippi river basin , or through dispersal ( mills et al . 1993 ; lesko et al . 2003 ; reid and hudson 2008 ) .

additionally , based on evidence from an ohio lake , it has been suggested that s . pallidus is an intermediate host for the parasitic worm tanaorhamphus longirostris , although study of this occurrence has been limited ( hubschman 1983 ) . documented evidence combined with its record of spread across the u . s . ( byron and saunders 1981 ) have led some recently colonized areas , like new zealand , to express concern over the potential effects of s . pallidus on native ecosystems ( duggan et al . 2006 ) .

there have been no recent records of this species in the great lakes . persistent populations of

may be lacking in the main water bodies of the great lakes , although individuals are probably washed into the lakes during floods . its native range could extend into tributaries and coastal wetlands within the basin ( mills et al . 1993 ; lesko et al . 2003 ; reid and hudson 2008 ) .

geddes , m . c . , and g . a . cole . 1981 . variation in sexual size differentiation in north american diaptomids ( copepoda : calanoida ) : does variation in the degree of dimorphism have ecological significance ? limnology and oceanography 26 ( 2 ) : 367 - 374 .

geiling , w . t . , and r . s . campbell . 1972 . the effect of temperature on the development rate of the major life stages of diaptomus pallidus herrick . limnology and oceanography 17 : 304 - 307 .

glmris . 2012 . appendix c : inventory of available controls for aquatic nuisance species of concern , chicago area waterway system . u . s . army corps of engineers .

hubschman , j . h . 1983 . diaptomus pallidus herrick , 1879 ( crustacea : copepoda ) as an intermediate host for tanaorhamphus longirostris ( van cleave , 1913 ) ( acanthocephala : neoechinorhynchidae ) . journal of parasitology 69 ( 5 ) : 930 - 932 .

krieger , k . a . , and d . m . klarer . 1991 . zooplankton dynamics in a great lakes coastal marsh . journal of great lakes research 17 : 255 - 269 .

lesko , l . t . , p . l . hudson , and m . a . chriscinske . 2003 . calanoid copepods of the laurentian great lakes . ann arbor , mi , online at urltoken

mclaughlin , p . l . , and 39 others . 2005 . common and scientific names of aquatic invertebrates from the united states and canada—crustaceans . american fisheries society special publication 31 , bethesda , maryland , american fisheries society , 545 p .

marcogliese , d . j . , and g . w . esch . 1992 . alterations of vertical distribution and migration of zooplankton in relation to temperature . american midland naturalist 128 ( 1 ) : 139 - 155 .

mills , e . l . , j . h . leach , j . t . carlton , and c . l . secor . 1993 . exotic species in the great lakes : a history of biotic crises and anthropogenic introductions . journal of great lakes research 19 ( 1 ) : 1 - 54 .

pennak , r . 1989 . fresh - water invertebrates of the unites states , 3rd ed . protozoa to mollusca . john wiley & sons , inc . , new york , new york state . 628 pp .

reid , j . w . , and p . l . hudson . 2008 . comment on “rate of species introductions in the great lakes via ship’s ballast water and sediments . canadian journal of fisheries and aquatic sciences 65 ( 3 ) : 549 - 553 .

torke , b . 2001 . the distribution of calanoid copepods in the plankton of wisconsin lakes . hydrobiologia 453 - 454 : 351 - 365 .

williamson , c . e . , and n . m . butler . 1986 . predation on rotifers by the suspension - feeding calanoid copepod diaptomus pallidus . limnology and oceanography 31 : 393 - 402 .

williamson , c . e . , and h . a . vanderploeg . 1988 . predatory suspension - feeding in diaptomus : prey defenses and the avoidance of cannibalism . bulletin of marine science 43 ( 3 ) : 561 - 572 .

wonham , m . j . , s . a . bailey , h . j . macisaac , and m . a . lewis . 2005 . modelling the invasion risk of diapausing organisms transported in ballast sediments . canadian journal of fisheries and aquatic sciences 62 : 2386 - 2398 .

kipp , r . m . , a . j . benson , j . larson , t . h . makled , and a . fusaro

this information is preliminary or provisional and is subject to revision . it is being provided to meet the need for timely best science . the information has not received final approval by the u . s . geological survey ( usgs ) and is provided on the condition that neither the usgs nor the u . s . government shall be held liable for any damages resulting from the authorized or unauthorized use of the information .

the data represented on this site vary in accuracy , scale , completeness , extent of coverage and origin . it is the user ' s responsibility to use these data consistent with their intended purpose and within stated limitations . we highly recommend reviewing metadata files prior to interpreting these data .

citation information : u . s . geological survey . [ 2018 ] . nonindigenous aquatic species database . gainesville , florida . accessed [ 7 / 9 / 2018 ] .

contact us if you are using data from this site for a publication to make sure the data are being used appropriately and for potential co - authorship if warranted . for queries involving fish , please contact pam fuller . for queries involving invertebrates , contact amy benson .

walter , t . c . & boxshall , g . ( 2018 ) . world of copepods database .

walter , chad . the world of copepods . , available online at urltoken [ details ]

webber , w . r . ; fenwick , g . d . ; bradford - grieve , j . m . ; eagar s . g . ; buckeridge , j . s . ; poore , g . c . b . ; dawson , e . w . ; watling , l . ; jones , j . b . ; wells , j . b . j . ; bruce , n . l . ; ahyong , s . t . ; larsen , k . ; chapman , m . a . ; olesen , j . ; ho , j . ; green , j . d . ; shiel , r . j . ; rocha , c . e . f . ; lörz , a . ; bird , g . j . ; charleston , w . a . ( 2010 ) . phylum arthropoda subphylum crustacea : shrimps , crabs , lobsters , barnacles , slaters , and kin , in : gordon , d . p . ( ed . ) ( 2010 ) . new zealand inventory of biodiversity : 2 . kingdom animalia : chaetognatha , ecdysozoa , ichnofossils . pp . 98 - 232 . [ details ]

( of leptodiaptomus pallidus ( light , 1938 ) ) walter , chad . the world of copepods . , available online at urltoken [ details ]

( of diaptomus pallidus herrick , 1879 ) walter , chad . the world of copepods . , available online at urltoken [ details ]

( of diaptomus reighardi marsh , 1895 ) walter , chad . the world of copepods . , available online at urltoken [ details ]

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s _ pygmaeus _ 5th _ leg . mov - virtual focus of the 5th leg ( 1 . 71 mb )

torke , b . 2001 . the distribution of calanoid copepods in the plankton of wisconsin lakes . hydrobiologia . 453 - 454 : 351 - 365 .

chow - fraser , p . and e . j . maly . 1988 . aspects of mating reproduction and co - occurrence in three freshwater calanoid copepods . freshwater biol . 19 : 95 - 108 .

havens , k . e . , n . d . yan , and w . keller . 1993 . lake acidification : effects on crustacean zooplanktonic populations . environ . sci . and technology . 27 : 1621 - 1624 .

hairston , n . g . and r . a . vanbrunt . 1994 . diapause dynamics of two diaptomid copepod species in a large lake . hydrobiologia . 292 - 293 : 209 - 218 .

bundy , m . h . and h . a . vanderploeg . 2002 . detection and capture of inert particles by calanoid copepods : the role of the feeding current . j . of plankton research . 24 : 215 - 223 .

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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .

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 .

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goniobranchus splendidus is a species of colourful sea slug , a dorid nudibranch , a marine gastropod mollusc in the family chromodorididae .

the specific epithet splendida means splendid in reference to the colouring , because this nudibranch has large red spots on a white background , a yellow line at the mantle edge , and magenta rhinophores .

goniobranchus splendidus

goniobranchus splendidus <EOT> goniobranchus splendidus , photographed by professor steve smith off nelson bay nsw in 2016 .

goniobranchus splendidus nudibranch ' s red spots vary across populations , but the yellow rim is present in all individuals .

wilson , ng , winters , ae & cheney , kl ( 2016 ) . tropical range extension for the temperate , endemic south - eastern australian nudibranch goniobranchus splendidus ( angas , 1864 ) . diversity 8 : 16

wilson ng , winters ae , cheney kl . tropical range extension for the temperate , endemic south - eastern australian nudibranch goniobranchus splendidus ( angas , 1864 ) . diversity . 2016 ; 8 ( 3 ) : 16 .

wilson , n . g . ; winters , a . e . ; cheney , k . l . tropical range extension for the temperate , endemic south - eastern australian nudibranch goniobranchus splendidus ( angas , 1864 ) . diversity 2016 , 8 , 16 .

wilson , nerida g . ; winters , anne e . ; cheney , karen l . 2016 . " tropical range extension for the temperate , endemic south - eastern australian nudibranch goniobranchus splendidus ( angas , 1864 ) . " diversity 8 , no . 3 : 16 .

epoxygoniolide - 1 ( 1 ) , possessing spiroepoxide lactone , enal , and masked dialdehyde functionalities , has been characterized from the conspicuously patterned mollusc goniobranchus splendidus . its relative configuration was investigated by spectroscopic analyses , molecular modeling , and density functional theory calculations . the biosynthesis of 1 may involve rearrangement of a diterpene . . . [ show full abstract ]

this study reports the isolation and characterisation of six new metabolites with ' gracilin ' - type carbon skeletons and of aplytandiene - 3 from the australian nudibranch goniobranchus splendidus . the structure of gracilin g is revised , and the c - 6 configuration deduced by comparison of calculated 3jc / h values with values measured using the exside pulse sequence . a lactone isolated from . . . [ show full abstract ]

using the gonibranchus splendidus in their study , they observed that the nudibranch displays a consistent yellow brim around a white body with red spots . it is found in the southern great barrier reef to new south wales . the colour and pattern of the red spots vary significantly across populations , but the yellow rim is the common factor .

three new norditerpenes ( 1 , 6 , and 7 ) and four diterpenes ( 2 - 5 ) with extensively rearranged carbon skeletons have been characterized from australian nudibranchs . the relative configuration of the cyclopropyl - containing verrielactone ( 1 ) from goniobranchus verrieri was suggested by spectroscopic analysis at 500 mhz informed by a combination of molecular modeling and dft calculations . the . . . [ show full abstract ]

t . 12 = ser . 3 : t . 4 ( 1864 ) - journal de conchyliologie . - biodiversity heritage library

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citation : johnson rf , gosliner tm ( 2012 ) traditional taxonomic groupings mask evolutionary history : a molecular phylogeny and new classification of the chromodorid nudibranchs . plos one 7 ( 4 ) : e33479 . urltoken

editor : jonathan h . badger , j . craig venter institute , united states of america

copyright : © 2012 johnson , gosliner . 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 .

funding : this work was supported by the california academy of sciences , the university of california santa cruz - department of ecology and evolutionary biology , the national science foundation deb 9978155 and deb 0329054 to tg , and an encyclopedia of life rubenstein fellowship to rj . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .

a . ‘phylogenetic scenario’ for the chromodorid genera modified from [ 5 ] , [ 26 ] . b . morphological phylogeny of generic representatives for the chromodorididae [ 13 ] . c . combined 16s and coi phylogram of the chromodorididae from [ 27 ] . d . combined 16s and coi phylogram of the chromodorididae and cadlinidae from [ 28 ] . rudman ' s ‘ chromodoris group’ in red , ‘ hypselodoris group’ in blue , cadlinella in yellow , diversidoris ( not included in [ 5 ] , [ 13 ] , [ 26 ] ) , cadlina in grey and other dorids in black .

new collections ( from this contribution and [ 28 ] in blue . genbank specimens in red . size of circle represents number of specimens collected in each region . specimen details in supplementary table s1 ) .

the goals of this contribution are : ( 1 ) generate a phylogeny that tests the species level relationships of the chromodorid nudibranchs and confirms the monophyly of the chromodorididae , ( 2 ) assess the phylogenetic validity of the chromodorid genera , and ( 3 ) propose a new classification for the chromodorid nudibranchs that reflects their relationships .

most of our samples were collected especially for molecular work and were preserved accordingly , either in 95 % etoh , sed buffer ( saturated nacl solution with edta and dmso ) or frozen . in addition to the specimens collected specifically for molecular study , we were also able to use museum material that was , either preserved in 70–75 % etoh or the original fixation method is unknown .

the cleaned , pcr products were copied and labeled with fluorescently dye - terminators ( big dye 3 . 1 abi ) in 10 µl reactions . each reaction contained 0 . 5–2 µl of cleaned pcr product , 1 . 63 µl of 5× reaction buffer , . 5 µl of primer ( 10 mm stock ) , 0 . 5 µl–0 . 75 µl of big dye and water to 10 µl . these reactions were run on a perkin elmer 9600 - geneamp pcr system or a biorad mycycler™ thermocycler ( software version 1 . 065 , bio - rad laboratories ) . the resulting labeled , single stranded dna was precipitated by addition of 2 . 5 µl of edta and sequential washing and pelleting in ( centrifuge details ) with 100 % and then 70 % etoh . the pelleted dna was denautured for two minutes at 94°c in 13–15 µl of hidi formamide ( applied biosystems ) . the denatured , labeled dna fragments were sequenced in both directions on the abi 3100 and 3130 genetic analyzer in the center for comparative genomics ( formerly the osher laboratory for molecular systematics ) at the california academy of sciences .

we assembled , edited and removed primer strands from forward and reverse strands for each gene fragment sequenced using sequencher ( ver . 4 . 7 . genecodes corporation ) and genious 3 . 0 - 5 . 3 . 3 ( biomatters ) . we aligned the coi sequences by eye and translated the base pair data into amino acids to using macclade 4 . 08 [ 46 ] to confirm alignment accuracy . we aligned 16s sequences with muscle [ 47 ] . we then further optimized the alignments by eye using both macclade [ 46 ] and genious 3 . 0 - 5 . 3 . 3 ( biomatters ) .

we tested for saturation or multiple substitutions at the same site by plotting the absolute number of transitions and transversions at each codon position ( 1 st , 2 nd , 3 rd ) for coi and at each base pair for 16s against both uncorrected p distance and log det using paup [ 48 ] and excel ( plots not shown ) .

sequence data for both genes was not obtained for every specimen we studied . we worked with two main data sets , because we wanted to test the effect of missing data on the resulting phylogeny : the two data sets were : 1 ) combined 16s and coi for specimens with sequence data for both genes , 2 ) all 16s and coi data for all specimens ( table s1 . ) both of these data sets were analyzed both including and excluding variable characters in the 16s alignment . for all of these analyses we used doris kerguelensis as the outgroup .

the sequenced coi fragment is 658 base pairs ( bp ) long . the edited 16s sequences are 531 bp long . the combined data sets with gaps introduced for alignment are 1189 base pairs long . all sequences are available from genbank coi ( jq727822–jq727914 ) , 16s ( jq727689–jq727821 ) and aligned data matrices are available upon request from the corresponding author . excluded variable 16s regions are identified as character sets in all nexus files . saturation was not found in the 16s fragment or the first or second positions of the coi fragment . there is slight saturation in the third position transitions in the coi data set ( not shown ) . the third positions were included in the bayesian analysis as the partitioning allows the parameters of this position to be estimated separately and the inclusion of the third positions did not change the resulting trees . the recommended model of evolution ( aic form mr . model test ) was used to set parameters in mr . bayes for each partition . the resulting best - fit model of evolution for each partition using the aic selection from mr . modeltest ver . 2 [ 78 ] were coi 1 st : gtr + g , coi 2nd : trn + i + g , coi 3 rd : gtr + i + g and 16s : gtr + i + g . these models correspond to the following settings in mr . bayes ; all partitions set to nst = 6 and rates = invgamma except for the coi second codon position partition which was set rates = gamma .

the figured trees are the resulting consensus phylograms from the bayesian analyses ( figures s1 , s2 ) . all posterior probabilities are shown above the branches on the bayesian phylograms . tree topology was not altered with the inclusion or exclusion or the 16s fragment ' s variable regions ( see figure s2 for comparison of trees with and without variable regions ) . the resulting phylogenetic hypotheses for each dataset are summarized below . we will discuss relationships in terms of posterior probabilities .

coi and 16s combined analysis : including only specimens with data for both genes ( figure s1 ) .

this data set included 244 individual chromodorids , representing 157 species , four species of actinocyclidae , four other dorids . the outgroup was doris kerguelensis .

tree is the same bayesian phylogram as figured in s3a . all specimens , both genes and all characters included .

tree is the same bayesian phylogram as figured in s3a . all specimens , both genes and all characters included . blue = indo - pacific , red = atlantic and caribbean , gold = eastern pacific , green = sister group , black = outgroups . dark grey = solely eastern pacific and atlantic clades . light grey = primarily indo - pacific clades with eastern pacific members .

the two species of cadlinella included here , cadlinella ornatissima and cadlinellla subornatissima form a clade and are sister to the rest of the chromodorid species ( pp = 1 . 00 ) . these findings support previous results [ 27 ] , [ 28 ] and rudman ' s evolutionary scenario [ 5 ] , [ 26 ] . the widespread indo - pacific genus , cadlinella is an enigmatic taxon . it has at different times been considered it own separate family [ 64 ] , a part of the cadlininae [ 65 ] and a member of the chromodoridinae / chromodorididae [ 26 ] , [ 66 ] .

cadlina burnayi ortea , 1988 [ 68 ] = t . nobilis [ 69 ]

the only two species of tyrinna : t . evelinae and t . nobilis are included here . tyrinna is always monophyletic ( pp = 1 . 00 ) . after the split from cadlinella , this clade is poorly - supported as the sister group to the main group of chromodorids ( pp = 0 . 83 ) . rudman [ 26 ] suggested that tyrinna , cadlinella and cadlina form a basal grade of primitive chromodorids . cadlina had been shown not to be a chromodorid [ 28 ] , but our results support rudman ' s suggestion that tyrinna and cadlinella are basal to the rest of the chromodorids . muniain et al [ 70 ] and schrödl and millen [ 71 ] extensively reviewed the morphology of the two species in this clade .

verconia verconis is well supported as part of a clade that includes n . haliclona , n . laboutei , n . romeri and n . simplex ( pp = 1 . 00 ) . noumea varians , n . purpurea and n . norba form a well - supported clade ( pp = 1 . 00 ) that is not part of a name bearing clade , but is one branch of the polytomy that includes the ‘noumea sensu stricto’ and the branch leading to the rest of the family ( pp = 0 . 88 ) . the monotypic genus verconia is nested within the noumea clade as suggested by rudman [ 75 ] and weakly supported as the sister species to another south australian species , n . haliclona , as found in the preliminary results shown by turner & wilson [ 27 ] .

type species : doris xantholeuca ehrenberg , 1831 [ 76 ] = g . pallida ( by subsequent designation )

the glossodoris clade ( pp = 1 . 00 ) includes species g . pallida and g . rufomargninata . in an important , but often overlooked detailed examination of the relationships of the species classified in the genus glossodoris , rudman identified five subgroups of this genus based on morphology [ 77 ] . the species in this glossodoris clade were considered by rudman [ 77 ] to be members of the ‘ glossodoris pallida subgroup’ . this clade also includes two species he did not include in any subgroup , g . cincta and g . hikuerenesis .

lissodoris odhner , 1934 [ 80 ] . type species : l . mollis odhner , 1934 ( = c . aureomarginata cheeseman , 1881 [ 86 ] ( by monotypy )

casella h . & a . adams , 1858 : 57 [ 84 ] . type species : c . gouldii h . & a . adams , 1858 [ 84 ] ( by monotypy )

species included in the glossodoris atromarginata subgroup [ 77 ] are recovered in this clade , with the addition of g . sedna and digidentis kulonba ( pp = 0 . 95 ) .

this name will be used for all eastern pacific and atlantic species of chromodoris and glossodoris ( except glossodoris sedna ) . these species form a polytomy including glossodoris baumanni and three clades of atlantic and eastern pacific chromodorids .

chromodoris clenchi , c . norrisi and c . sphoni ( pp = 1 . 00 )

chromodoris krohni , c . luteorosea and c . purpurea ( pp = 0 . 78 )

these exclusively eastern pacific and atlantic clades do not form a monophyletic group , but we will provisionally name all of these species ‘felimida’ . this is the most conservative choice , the choice that requires the fewest name changes and is the least disruptive pending further information and broader taxon sampling .

the ardeadoris clade includes both species of ardeadoris : a . egretta and a . scottjohnsoni , five species of glossodoris ( g . averni , g . pullata , g . rubroannulata , g . tomsmithi and glossodoris undaurum ) and noumea angustolutea ( pp = 1 . 00 ) . . based on their analysis , turner and wilson [ 27 ] suggested that with more sampling it would be come clear if ardeadoris should be synonmized with glossodoris . by sampling more broadly within the family , we found the converse . four species of glossodoris and noumea angustolutea need to be included within ardeardoris because they are strongly supported as part of the clade including ardeadoris egretta and not the type species of glossodoris . three of the species , g . averni , g . undaurum and g . rubroannulata , found in this clade were part of rudman ' s glossodoris sedna subgroup [ 77 ]

this clade includes all of the black - lined species of chromodoris and chromodoris aspersa ( pp = 1 . 00 ) . this clade was identified by , both wilson & lee [ 17 ] and turner & wilson [ 27 ] , as the planar spawning or black - lined chromodoris clade . all of the members of this clade lay flat egg masses .

type species : diversidoris aurantionodulosa rudman , 1987 [ 89 ] ( by original designation ) .

the diversidoris includes , diversidoris aurantionodulosa , two yellow species of noumea , n . crocea and n . flava , and a new species from moorea , french polynesia - chromodoridae biocode 2937 ( pp = 0 . 95 ) .

the miamirinae clade includes all of the species currently classified as ceratosoma , durvilledoris , hypselodoris , mexichromis , pectenodoris , risbecia , thorunna and two species of digidentis ( pp = 1 . 00 )

this clade was first predicted by rudman [ 26 ] based on morphological similarities and then confirmed by rudman & berquist ' s [ 5 ] finding that all of the species in this clade feed exclusively on sponges of the family dysideaidae , although they assumed all of the genera to be monophyletic . miamirinae bergh 1891 is the oldest appropriate and available subfamily or family name for this clade . the remaining six genera ; miamira , ceratosoma , felimare , mexichromis , thorunna and hypselodoris make up the miamirinae .

the miamira clade includes the following species ( as currently classified ) ceratosoma alleni , ceratosoma magnificum , ceratosoma miamiranum , ceratosoma sinuatum . miamira is part of a grade with ceratosoma . the morphological phylogeny of species of ceratosoma and classified as miamira and orodoris , that was used as justification for their synonomy , predicted a sister group relationship between species of miamira and ceratosoma alleni [ 93 ] . our results confirm that c . alleni is more closely related to species of miamira , but do not find support for synonymy of miamira and ceratosoma . although , it is possible this relationship will be recovered with further sampling and by including molecular markers that will help resolve basal branches on the phylogeny .

the ceratosoma clade includes c . amoenum , c . gracillimum , c . ingozi , c . tenue , c . trilobatum and a new species . ( pp = 1 . 00 )

the felimare clade includes all eastern pacific , atlantic and mediterranean species of hypselodoris and two species of mexichromis , m . porterae and m . kempfi from the eastern pacific and caribbean respectively ( pp = 1 . 00 ) . both gosliner and johnson [ 13 ] and alejandrino and valdés [ 31 ] hypothesized a sister group relationship between the indo - pacific and eastern pacific / atlantic species of hypselodoris . turner and wilson [ 27 ] did not recover that relationship , but instead found the same relationships shown here .

this clade includes the type species of mexichromis , m . antonii , known only from the eastern pacific and the three included species of durvilledoris , d . lemniscata , d . pusilla and d . similaris , the two described species of pectenodoris , p . aurora and p . trilineata and all of the indo - pacific species currently considered mexichromis , m . festiva , m . macropus , m . mariei and m . mutituberculata ( pp = 1 . 00 ) . there are two well - supported clades within the mexichromis clade . the clade including mexichromis antonii and the species of durvilledoris is sister to the clade including pectenodoris and indo - pacific mexichromis . these clades could be given two names , but it is much less disruptive and confusing to maintain the name mexichromis for all clade members . the clade including p . aurora and p . trilineata can be called the ‘ pectenodoris ’ clade of mexichromis .

the thorunna clade includes all species of thorunna and two species of digidentis , d . arbutus and d . perplexa . all of species currently classified as thorunna are found in the indo - pacific and the species of digidentis are limited to southern australia . as suggested by rudman [ 26 ] , the only species within thorunna with mantle glands , t . australis and the species of digidentis ( all of which have mantle glands ) form a clade .

this clade includes all of the indo - pacific species of hypeslodoris and risbecia ( pp = 1 . 00 ) .

species of risbecia s . s forms a well - supported clade nested within hypselodoris and can be referred to as the risbecia clade of hypselodoris . risbecia aplogema is not part of this risbecia clade and was previously considered a species of hypselodoris . including all of the members of the risbecia and hypselodoris bullocki clade in risbecia is not an option because this would render hypsleodoris paraphyletic . the second clade includes , h . bennetti , h , maritima , h . bertschi , h . paulinae , h . kaname , h . bollandi , h . obscura , h . infucata , h . zephrya and one or two new species . the third clade includes h . reidi , h . krakatoa , h . jacksoni and one new species . this clade was also recovered in gosliner & johnson ' s [ 13 ] morphological phylogeny of hypselodoris .

in future contribtuions , we will work out synapomorphies for the clades identified here , but because of the amount of homoplasy and number of incomplete descriptions , this is a huge undertaking and not appropriate here .

in summary , with the most comprehensive sampling of chromodorid species to date , we confirmed that the chromodorids are monophyletic and are sister to the monophyletic actinocyclids . we also found that the majority , 12 / 14 non - monotypic traditional genera , were not monophyletic or make another clade paraphyletc . seven traditional genera , ceratosoma , chromodoris , digidentis , glossodoris , hypselodoris , mexichromis , noumea are non - monophyletic and three ( durvilledoris , pectenodoris , risbecia ) are monophyletic but render another genus paraphyletic . both ardeadoris and thorunna are made paraphyletic by nested members of other genera ( noumea , glossodoris and digidentis ) . the two monotypic genera , diversidoris and verconia are nested within clades . only tyrinna and cadlinella are monophyletic and without disruption to any other clades ( figure 3 , s1 , s2 ) . the classification proposed here and discussed at length above renames clades and is more consisitent with evolutionary history ( figure s3 ) .

the most speciose chromodorid genera : chromodoris , glossodoris and hypselodoris were originally created to describe indo - pacific species . it wasn ' t until some time after these names were created that previously described , similar , brightly colored cryptobranch dorid species found in the eastern pacific , western atlantic and mediterranean were added to these genera [ 1 ] , [ 26 ] , [ 65 ] , [ 95 ] , [ 102 ] , [ 103 ] . in mexichromis the opposite is true . the type species , mexichromis antonii , was described from the eastern pacific and indo - pacific species were included later included in this genus [ 26 ] , [ 95 ] . other eastern pacific “ mexichromis ” are shown here to belong to felimare .

this new classification clarifies our view of biogeographic patterns in the chromodorid nudibranchs . instead of taxonomy obscuring patterns of diversification in this group , this taxonomy reflects and reinforces evolutionary history . it gives us a much better framework for exploring evolutionary questions .

the electronic version of this document does not represent a published work according to the international code of zoological nomenclature ( iczn ) , and hence the nomenclatural acts contained in the electronic version are not available under that code from the electronic edition . therefore , a separate edition of this document was produced by a method that assures numerous identical and durable copies , and those copies were simultaneously obtainable ( from the publication date noted on the first page of this article ) for the purpose of providing a public and permanent scientific record , in accordance with article 8 . 1 of the code . the separate print - only edition is available on request from plos by sending a request to plos one , public library of science , 1160 battery street , suite 100 , san francisco , ca 94111 , usa along with a check for $ 10 ( to cover printing and postage ) payable to “public library of science” .

table of specimens used in this study . specimens used in this study listed by family . the names in this table reflect current classification not proposed classification ( new names are listed in the text ) . abbreviations are as follows : casiz = california academy of sciences , sam = south australian museum , wam = western australian museum , am = australian museum , zsm = zoologische staatssammlung münchen , sio - bic = scripps institute of oceanography , biocode = moorea biocode project .

new classification of the chromodorididae with synonyms . generic names and type species in bold and the most recent genus membership follows . listing order follows phylogeny .

bayesian consensus phylogram including all specimens with data for both genes . posterior probabilities are listed above branches . doris kerguelensis is the outgroup . this phylogram is the consensus of 50 , 000 , 000 generations with trees sampled every 1000 generations with a burnin of 25 % . data was partitioned by gene and by codon position .

bayesian consensus phylograms including all specimens . a . phylogram resulting from the inclusion of all characters b . phylogram resulting from excluding hard to align characters . doris kerguelensis is the outgroup . these phylograms are the consensuses of 50 , 000 , 000 generations with trees sampled every 1000 generations with a burnin of 25 % . data was partitioned by gene and by codon position . dotted lined indicate areas of disagreement .

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upper : shiprock , port hacking , sydney , new south wales , 3m , march 1982 . lower : shiprock , port hacking , sydney , new south wales , december 1982 , on food sponge . photos : bill rudman .

there are many red and orange - spotted species of chromodorid in new south wales and southeastern australia . i have discussed this example of mimicry on a separate page .

references : • rudman , w . b . ( 1983a ) the chromodorididae ( opisthobranchia : mollusca ) of the indo - west pacific : chromodoris splendida , c . aspersa and hypselodoris placida colour groups . zoological journal of the linnean society 78 : 105 - 173 . • rudman , w . b . ( 1991 ) purpose in pattern : the evolution of colour in chromodorid nudibranchs . journal of molluscan studies , 57 , ( t . e . thompson memorial issue ) : 5 - 21 .

it usually takes about 3 months before the first slugs show up and they are always austraeolis ornata and jorunna sp 1 . plocamopherus imperialis then arrives on the main sea wall and dendrodoris denisoni and dendrodoris fumata show up further upstream . sometimes aplysia spp also arrive . it is always after these species establish that chromodorids appear about half a kilometre up river and always on the same rock wall . chromodoris geometrica , chromodoris daphne , chromodoris cf reticulata and hypselodoris obscura all appear about the same time and what is curious is that they are often large animals with almost no juveniles seen , and i have always had a thorough search before hand .

if no slugs survive after a flood ( chromodorids definitely dont ) how do these species re - establish . the odd animal may come in on weed but the regularity of their arrival seems to indicate that they must have at least a short planktonic stage and can only establish once their food source is present . i have no qualification to make any assumption but i believe that my observation is correct so i would appreciate your opinion

we basically need to know a lot more about the lifecycles of the nudibranchs and of the food organisms - algae , sponges , hydroids , bryozoans etc - before we can say much more about how nudibranchs re - establish themselves at certain places after catastrophes . it is a very interesting - if very complicated topic .

dear bill , in answer to your question : " i wondered if two sponge species were interwoven , but i suspect the bluish regions are paler parts of the chelonaplysilla - perhaps where the slug has been feeding . i would be interested in your comments . do you perhaps have a photo showing the join between blue and darker regions more clearly ? " we have another photo of each nudibranch and sponge in the previous message . the two we sent in showed the mouth everted and indicated feeding better than the ones we didn ' t send . here are the two other photos . we think what you are seeing is not two sponges but the hills and hollows of the sponge making shadows and light . we will keep looking for chromodoris daphne feeding and see if we can come up with some photos that clear up the puzzle .

locality : the pipeline , nelson bay , port stephens and fly point , port stephens - great lakes marine park , port stephens , 5metres , new south wales , australia , pacific ocean , 09 january 2009 & 10 january 2009 , sandy silty bottom with sponges , ascidians , gorgonias , soft corals , hydroids and seaweed . length : 30 mm . photographer : leanne and david atkinson .

i think these extra photos have cleared up the puzzle quite well . some sponges change colour in different parts of the same colony - certainly parts that are shaded are often much paler or almost colourlesss . it seems as though chelonaplysilla violacea can range in colour from deep violet to dull blue to a reddish colour and also to a pale straw colour . one of the distinguishing features of chelonaplysilla is its incorporation of small sand grains into its tissue which can usually be seen in c . violacea , in the granular appearance of the honeycomb - like surface of the colony . interestingly your lower photo shows part of the honeycomb layer [ c ] being hidden by a smooth layer with raised conical papillae more typical of the related genera darwinella . if i had seen the smooth section only i wouldn ' t have thought of it as chelonaplysilla .

as i have often said , sponges are notoriously difficult to identify externally , and one of the reasons for this is that we don ' t have a good idea of the variability of living colonies . although this is not a ' sponge forum ' our growing collection of ' nudibranch food ' photos is increasing our knowledge in both fields of biology .

hi bill and everyone ! so chromodoris daphne is the hot topic . . . i won ' t go into feeding but will go into egg masses and mating . this species is fairly common here and is typically a botton dweller . in all the dive we have seen it , it has been crawling along the silty sandy bottom . it has been found intertidally and at the mooloolaba rock wall / ledges , which is a shore dive and old woman island and the local reefs . i collected a couple to see if they would lay eggs and walla . . . two animals laid egg masses together at the same time and then went right back to mating ! i could not believe my eyes .

locality : mooloolaba , sunshine coast , intertidal down to 20 m , sthn queensland , australia , pacific ocean , 20 january 2009 . length : 15 - 25 mm . photographer : gary cobb .

hi bill , we found these chromodoris daphne feeding on a black sponge on two recent dives , one at fly point and the other at the pipeline . these dive sites are at opposite ends of nelson bay beach . there is some discussion on where to find these nudibranchs in an earlier message by dez paros . we usually find these nudibranchs in less than 12 metres in the zone where there is a mixture of seaweed and sponges .

locality : the pipeline , nelson bay , port stephens and fly point , port stephens - great lakes marine park , port stephens , 5 metres , new south wales , australia , pacific ocean , 09 january 2009 & 10 january 2009 , sandy bottom sponges , bryozoans , soft corals , hydroids and ascidians . length : 30 mm . photographer : leanne and david atkinson .

to accompany leanne & david atkinson ' s message [ # 18549 ] , here is another record of chromodoris daphne feeding .

locality : shiprock , port hacking , sydney , new south wales , australia . 7 m . 20 december 1982 , many specimens , 20 - 35 mm long on food sponge . coll : g . avern . am c137068 . photos : bill rudman .

many specimens were found , all on a large empty beer bottle which was completely covered in a colony of the purple darwinellid sponge chelonaplysilla violacea .

hi bill , we came across this chromodoris daphne we think was feeding and thought it might be of interest .

locality : little beach , fly point marine reserve port stephens , 8 metres , nsw australia , pacific , 12th november 2006 , sandy bottom scattered sponges , kelp , ascidians soft corals and gorgonias . length : 40 mm . photographer : leanne & david atkinson .

the upper shot shows the sponge , the damage where feeding has occurred and a mucus trail to the chromodoris daphne . the lower shot quite clearly shows the mouth everted . water temperature was 19 degrees celcius . there didn ' t seem to be any other messages or information about its food on the site . regards , leanne & david atkinson

dear leanne & david , this is a very valuable addition to our knowledge . i have added a couple of closeups to show the damage and sponge structure in more detail . the sponge is a darwinellid , and i suspect a species of chelonaplysilla but will need to check with prof . bergquist . your mention about no feeding information on the site for this species surprised me as i thought i had photos of it on a discarded beer bottle which was covered in the purple sponge chelonaplysilla violacea . on checking i have only a very cursory reference on the fact sheet so i have prepared a fuller message to accompany your record .

it ' s very good to have some confirmatory observations - and interesting from the sponge point of view as well because we still have a lot to learn about how many species of chelonaplysilla there are .

dear bill , we were thrilled to find this chromodoris daphne laying eggs on our last dive . sadly we were still learning to use our newly housed digital camera and badly overexposed the shots showing the eggs coming out . it was definitely laying these eggs and not just crawling over them . locality : fly point marine reserve port stephens , 5 metres , new south wales , australia , pacific ocean , 18 . 03 . 2006 , sandy bottom seaweed with scattered sponges and kelp . length : 25 mm . photographer : leanne & david atkinson .


charopa is a genus of air-breathing land snails , terrestrial pulmonate gastropod mollusks in the family charopidae .

charopa is the type genus of the family charopidae .

charopa

charopa <EOT> worms - world register of marine species - charopa e . von martens , 1860

karin mahlfeld added text to " brief summary " on " charopa coma ( gray 1843 ) " .

significant find : the ' charopa lafargei ' was found in the northernmost part of gunung kanthan in ipoh .

species charopa novoguineensis l . soós , 1911 accepted as pilsbrycharopa kobelti ( boettger , 1908 ) ( junior synonym )

species charopa descendens i . rensch , 1937 accepted as sinployea descendens ( i . rensch , 1937 ) ( original combination )

species charopa novopommerana i . rensch , 1937 accepted as sinployea novopommerana ( i . rensch , 1937 ) ( original combination )

species endodonta ( charopa ) vortex r . murdoch , 1897 accepted as geminoropa vortex ( r . murdoch , 1897 ) ( original combination )

species charopa nigrofusca e . a . smith , 1896 accepted as pilsbrycharopa nigrofusca ( e . a . smith , 1896 ) ( original combination )

species charopa rotumana e . a . smith , 1897 accepted as sinployea rotumana ( e . a . smith , 1897 ) ( original combination )

“we named this species charopa lafargei after lafarge , whose declared goals for biodiversity include minimising and avoiding damage to important habitats , ” the report said .

found by two malaysians in 2011 , charopa lafargei is only 1 . 4mm tall and takes its name after lafarge , which is currently mining the limestone hill gunung kanthan .

heliosia charopa turner , 1904 ( arctiidae : lithosiinae ) , female - qld , townsville , 11 . oct . 1901 , f . p . dodd leg . ( anic ) .

arina charopa ( blr ) ' 15 corbeil tournament all around / tournoi de corbeil concours général 15 . 533 ( diff : 7 . 6 / ex : 7 . 933 ) rank : 9th

“these adapted species often occur confined to a single hill or a small group of hills , and are found nowhere else in the world . charopa lafargei may be such a species , ” he said .

vermeulen , j . j . and marzuki , m . e . 2014 . ' charopa ' lafargei ( gastropoda , pulmonata , charopidae ) , a new , presumed narrowly endemic species from peninsular malaysia . basteria 78 ( 1 - 3 ) : 31 - 34 .

‘charopa’ lafargei spec . nov . is described . it is apparently restricted to the isolated limestone hill gunung kanthan in peninsular malaysia . the hill is scheduled for quarrying by the lafarge malaysia cement company , which has a high likelihood of resulting in the extinction of the species .

( of patula ( charopa ) e . von martens , 1860 ) gottschick , f . ( 1911 ) . aus dem tertiärbecken von steinheim a . a . jahreshefte des vereins für vaterländische naturkunde in württemberg . 67 , 496 - 534 . , available online at urltoken page ( s ) : 501 [ details ]

obis indo - pacific molluscan database . , available online at urltoken [ details ]

( of simplicaria mousson , 1890 ) bank , r . ( 2017 ) . classification of the recent terrestrial gastropoda of the world . last update : july 16th , 2017 . [ details ]

a leg injury ruled her out of the 2016 world cup event in minsk , belarus . ( beatricevivaldi . com , 20 may 2016 ) she injured her leg in january 2015 . ( passion , beauty and grace are rhythmic gymnastics facebook page , 19 jan 2015 )

winning a silver medal in the team event at the 2014 world championships in izmir , turkey . ( ctv . by , 10 aug 2014 ; sportpanorama . by , 28 sep 2014 )

north , south , chatham and stewart islands ( with distributional gap from southern westland across to mid - canterbury , most of otago and southland and western fiordland ) .

litter dweller in a wide range of habitats including drier , open and more modified habitats .

distribution north , south , chatham and stewart islands ( with distributional gap from . . .

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3 . 5 miles sw of sungei siput , 4°46’53 . 58”n , 101°7’15 . 99”e ( leg . m . e .

ery on the last whorl . radial ribs predominant , c . 30 on the

of whorls up to 4 1 / 2 , height aperture c . 0 . 5 mm ; width aper -

list of threatened species . version 2013 . 2 . < www . iucnredlist . org > .

, 2014 . lafarge biodiversity strategy : 1 - 10 . lafarge , paris .

iucn 2013 . iucn red list of threatened species . version 2013 . 2 .

inventorise the rich mollusc faunas of varied depositional environments of nw borneo , covering miocene to recent environments ranging from estuarine to mid shelf .

notes on the non - marine molluscs of the island of borneo 5 . the genus diplommatina ( gastropoda proso . . .

notes on the non - marine molluscs of the island of borneo 8 . the genus arinia ; additions to the gener . . .

notes on the non - marine molluscs of the island of borneo 6 . the genus opisthostoma ( gastropoda proso . . .

a new land snail , arinia ( notharinia ) micro ( caenogastropoda : cyclophoroidea : diplommatinidae ) , from . . .

a new snail species of the family diplommatinidae is described from perak , western peninsular malaysia . arinia ( notharinia ) micro , new species , differs from known congeners by its remarkably smaller shell . it is among the smallest land snails in the world with a mean shell height of 0 . 85 mm and mean shell width of 0 . 35 mm . the shell has tight radial ribbed whorls ( except for the smooth . . . [ show full abstract ]

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apparently this species is restricted to the isolated limestone hill gunung kanthan in peninsular malaysia . the hill lies close to the limestone quarry 3 . 5 miles southwest of sungei siput ( 4°46’53 . 58”n , 101°7’15 . 99” ) .

there are no data on population trends , as the species was recently described .

found on leaf litter at the base of a limestone cliff at the northern extremity of the hill .

the primary limestone forest in this area was previously in proximity to a nearby hindu temple , providing some influence on forest management but the cement company lafarge malaysia is currently in negotiation with the various temples around the hill to use the habitats for quarrying hence the continued presence of the forests and adjacent temples is not guaranteed , as when quarrying starts , the forests will be logged , and hence the habitats will be lost , and the adjacent regions will suffer from dust pollution . the forest habitat is also being possibly being degraded by the presence adventitious oil palm seedlings from the adjacent plantations which may continue to impact the original forest habitat .

there are no species - specific conservation actions in place for the mollusc . the changes in land management may adversely affect the habitats of this species .

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ipoh : a new species of snail has been discovered in perak and named after the cement company quarrying the hill it was found on .

published in a scientific journal by the netherlands malacological society on aug 17 , the report said the snail was found on the hill’s northern region .

“on a species level , it is uniquely identified among west malaysian charopidae by its conical shell and high , lamella - shaped radial ribs , ” the report , co - authored by dutch taxonomist jaap van vermuelen and malaysian mohammad effendi marzuki , said .

it added that the snail was found on leaf litter at the base of a limestone cliff and termed as “presumed narrowly endemic” , meaning that it was found only in a small area and that more surveys needed to be done to be sure .

speaking to the star via e - mail , vermuelen said the snail was discovered by mohammad and liew thor seng , a biologist currently based at universiti malaysia sabah .

description : shell minute , spire flat or slightly sunken . protoconch smooth , about 1¼ whorls , first ½ whorl sloping downwards . teleoconch of about 3½ rounded whorls , smooth apart from fine , dense axial growth ridges . aperture nearly circular , outer lip simple , thin . umbilicus widely open . shell transparent becoming opaque white with age .

distribution : endemic to australia : port stephens , nsw , southwards and around southern australia , to shark bay , wa , including tasmania .

habitat : specimens are from intertidal and shallow subtidal rock and algal washings , and from beach washup . common .

remarks : the anatomy of this species was described in detail by ponder ( 1990 ) .

figs . 1–3 : st helens point , tasmania ( c . 213600 ) . the dried animal is inside the shell , and the operculum is visible in the aperture .

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