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

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

margaritiferidae

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


the rhinoceros chameleon ( furcifer rhinoceratus ) is a species of chameleon that gets its common name from its horn-like nose which is most prominent in males .

it is endemic to dry forests in madagascar .

rhinoceros chameleon

today i want to write about an amazing looking reptile , the rhinoceros chameleon .
male rhinoceros chameleon ( furcifer rhinoceratus ) . ankarafantsika national park , north west madagascar .
both the male and female rhinoceros chameleon possess a dorsal crest . this is only present on the front half of the body , and , together with the lower casque , helps distinguish the male rhinoceros chameleon from the closely related furcifer antimena and furcifer labordi . the female rhinoceros chameleon can also be differentiated from these two species by the absence of a white line down the middle of the belly ( 2 ) .
endangered rhinoceros chameleon ( furcifer rhinoceratus ) resting on a branch in rain forest in ankarafantsika , madagascar . species is threatened ( designated vulnerable ) .
kauai : there was a single sighting of a veiled chameleon in 2004 . kisc responded but was unable to recover a chameleon .
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the rhinoceros chameleon , scientifically named furcifer rhinoceratus is one of the most distinctive of the genus furcifer . it’s named for its prominent scaly horn - like nose (
the rhinoceros chameleon is endemic to the dry deciduous forest region of western madagascar , although its exact distribution within this region is poorly understood ( 2 ) ( 4 ) .
little information is available on the reproductive behaviour of the rhinoceros chameleon . clutches of 4 to 11 eggs have been reported in captivity , with hatching occurring after 291 days , and the newly hatched rhinoceros chameleons weighing between 0 . 38 and 0 . 44 grams ( 2 ) .
rhinoceros chameleons are found in forests on the island of madagascar and are considered vulnerable to extinction due to habitat loss .
the rhinoceros chameleon inhabits dry deciduous forest ( 4 ) . little information is available on its precise habitat requirements , but it is reported to be commonly seen along roads and paths , as well as inhabiting the forest interior ( 2 ) .
this chameleon inhabits dry deciduous forest ( ramanamanjato and rabibisoa 2002 ) . it is not clear to what degree this species can survive in degraded habitats .
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offered on this website are simply our own and our readers opinions . every case must be treated on an individual basis . as always , my chameleon online encourages all chameleon owners to seek professional veterinary care . privacy policy terms & conditions
like many madagascan chameleons , the rhinoceros chameleon is threatened by habitat loss in the form of rapid deforestation ( 5 ) ( 6 ) . significant portions of forest habitat have already been cleared , and the remaining areas are often highly fragmented , as well as being critically threatened by logging , uncontrolled burning , and clearing for grazing and agriculture ( 4 ) .
a reptilian smoking gun : first record of invasive jackson’s chameleon ( chamaeleo jacksonii ) predation on native hawaiian species . brenden s . hollandsteven l . montgomery , vincent costello . urltoken
the name
chameleon
is derived from the greek words chamai ( on the ground , on the earth ) and leon ( lion ) so their name means
earth lion .
the meller ' s chameleon is the largest of the chameleons not native to madagascar . their stout bodies can grow to be up to two feet long and weigh more than a pound .
) . contrary to popular belief , a chameleon typically does not change colors to match its surroundings . instead , color is usually used to convey emotions , defend territories , and communicate with mates .
male cryptic or blue - legged chameleon ( calumma crypticum ) rests on a branch in the wilds of madagascar ( rain forest of ranomafana ) . incredible vibrant colors at night while sleeping . foliage , tree .
meller ' s distinguish themselves from their universally bizarre - looking cousins with a single small horn protruding from the front of their snouts . this and their size earn them the common name
giant one - horned chameleon .
the male rhinoceros chameleon is greyish or brownish in colour , with dark brown or black between the scales , and a white line down the flanks . the lips are also white , and the horn is often a bluish colour . although usually similar in colouration to the male , the female changes dramatically when carrying eggs ( gravid ) . in this state , the female turns an overall neon - purple colour , with black bands on the body , extending onto the tail , which is orange or red ( 2 ) .
this species is typically larger than a jackson chameleon . they also have a large casque ( sharkfin - like shield ) on their head , as compared to the three horns of a male jackson . this species is also a pest !
issg database : ecology of chamaeleo jacksonii . urltoken images : veiled chameleon , another popular pet and illegal animal to posess in hawaiʻi . urltoken two widespread introduced reptiles : ( top ) gold day dust gecko ; ( bottom ) brown anole
processes in ecology and evolution , natural and life history , and biodiversity conservation of reptiles and amphibians . research interests include african chameleon systematics and neuroanatomy , reproductive variation in north american lizards and frogs , and ecology of snakes and turtles in artificial habitats .
in some areas malagasy fear chameleons . they are also the subject of some well - known local proverbs including “manaova toy ny dian - tana jerena ny aloha , todihina ny afara , ” which translates to
like the chameleon , one eye on the future , one eye on the past
;
ratsy karaha kandrondro ,
meaning
ugly as a chameleon
;
mahatsidia vokon ' anjava kely izy fa mafoaka ,
a warning to walk carefully so as not to step on a brookesia , which would bring great misfortune .
jacksonʻs chameleon ( chamaeleo jacksonii ) : males are generally 10 - 12” long with a long , prehensile tail accounting for half of their length , and with three horns protruding from their forehead . females are generally a little shorter and never grow horns . this species is also a pest !
have a large casque ( sharkfin - like shield ) on their head ( note that the jackson’s chameleon , c . jacksonii , is a similar and more common alien species in hawaii , although male jackson’s have three horns on their forehead ) . have a fringe running from its mouth under its body .
other easily noted characteristics of chameleons include bulging eyes that move independently of one another , feet fixed in a grasping position , and the existence of horns or crests on the heads of many species . additionally , arboreal species have prehensile tails used for grasping objects when climbing and moving . finally , some species have long extensile tongues for catching insects or small vertebrates at a distance sometimes greater than the length of the chameleon .
monoceras : madagascar ( betsako , mojanga ) ; type locality : betsako bei mojunga , n . w . madagascar
note : tdwg regions are generated automatically from the text in the distribution field and not in every cases it works well . we are working on it .
furcifer monoceras is most likely a synonym of f . rhinoceratus ( f . glaw , pers . comm . )
andersson , lars gabriel 1910 . reptiles and batrachians from the north - west of madagascar collected by v . kaudern , 1906 - 1907 . arkiv för zoologi 7 ( 7 ) : 1 - 15 .
boettger , o . 1913 . reptilien und amphibien von madagascar , den inseln und dem festland ostafrikas . pp . 269 - 375 . in : voeltzkow , a . reise in ostafrika in den jahren 1903 - 1905 . wissenschaftliche ergebnisse . vol . 3 . systematische arbeiten . schweizerbart’ sche verlagsbuchhandlung , nägele und sproesser , stuttgart
boettger , oskar 1893 . katalog der reptilien - sammlung im museum der senckenbergischen naturforschenden gesellschaft in frankfurt am main . i . teil ( rhynchocephalen , schildkröten , krokodile , eidechsen , chamäleons ) . gebrüder knauer , frankfurt a . m . , x + 140 pp . - get paper here
brygoo , e . r . & c . a . domergue 1968 . les caméléos á rostre impair et rigide de l ' ouest de madagascar . validité des espéces chamaeleo labordi grandidier , 1872 , et c . antimena grandidier , 1872 . description d ' une espéce nouvelle c . angeli n . sp . et de la femelle de c . rhinoceratus gray , 184 mém . mus . nat . hist . nat . , paris a 52 ( 2 ) : 71 - 110
gardner cj , raxworthy cj , metcalfe k , raselimanana ap , smith rj , davies zg 2015 . comparing methods for prioritising protected areas for investment : a case study using madagascar’s dry forest reptiles . plos one 10 ( 7 ) : e0132803 , doi : 10 . 1371 / journal . pone . 0132803 - get paper here
glaw , f . & vences , m . 1994 . a fieldguide to the amphibians and reptiles of madagascar . vences & glaw verlag , köln ( isbn 3 - 929449 - 01 - 3 )
glaw , f . 2015 . taxonomic checklist of chameleons ( squamata : chamaeleonidae ) . vertebrate zoology 65 ( 2 ) : 167–246 - get paper here
glaw , f . & vences , m . 2001 . ein seltenes chamäleon aus madagaskar . datz 54 ( 7 ) : 16 - 19
gray , j . e . 1843 . description of two new species of reptiles from the collection made during the voyages of h . m . s . sulphur . ann . mag . nat . hist . ( 1 ) 11 : 46 - get paper here
gray , j . e . 1845 . catalogue of the specimens of lizards in the collection of the british museum . trustees of die british museum / edward newman , london : xxvii + 289 pp . - get paper here
gray , j . e . 1865 . revision of the genera and species of chamaeleonidae , with the description of some new species . ann . mag . nat . hist . ( 3 ) 15 : 340 - 354 - get paper here
necas , p . 2012 . chamäleons der gattung furcifer . reptilia ( münster ) 17 ( 95 ) : 20 - 27 - get paper here
necas , petr 1999 . chameleons - nature ' s hidden jewels . edition chimaira , frankfurt ; 348 pp . ; isbn 3 - 930612 - 04 - 6 ( europe ) < br / > isbn 1 - 57524 - 137 - 4 ( usa , canada )
pawlik , k . 2012 . chamäleon - urlaub – eine madagaskar - reise . reptilia ( münster ) 17 ( 96 ) : 103 - 109 - get paper here
pianka , e . r . & vitt , l . j . 2003 . lizards - windows to the evolution of diversity . university of california press , berkeley , 347 pp . [ review in copeia 2004 : 955 ] - get paper here
rakotoarison , andolalao ; jesse erens , fanomezana m . ratsoavina , miguel vences 2015 . amphibian and reptile records from around the betsiboka delta area in north - western madagascar herpetology notes 8 : 535 - 543 - get paper here
schmidt , w . ; tamm , k . & wallikewitz , e . 2010 . chamäleons - drachen unserer zeit . natur und tier verlag , 328 pp . [ review in reptilia 101 : 64 , 2013 ] - get paper here
this database is maintained by peter uetz ( database content ) and jakob hallermann , zoological museum hamburg ( new species and updates ) .
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f . monoceras is considered a junior synonymn of f . rhinoceratus ( glaw and vences 2007 ) .
this species is uncommon . due to heavy human pressures on and the patchy distribution of remaining forest within its range , the population is likely to be both declining and severely fragmented .
this species was traded in small levels before the 1994 suspension on exports from madagascar . it is not currently reported illegally in international trade .
threats in the region include bush fires , cattle grazing , and deforestation for charcoal collection .
this species occurs in parc national ankarafantsika . this site needs to be managed to limit the intrusion of damaging human activities . research is needed to establish the extent of this species ' distribution in the west of its range , from which modern records are lacking , and to clarify the degree to which this species is able to tolerate habitat degradation and modification .
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many madagascan chameleons have also been heavily collected for the pet trade ( 6 ) . however , there are very few records of successful captive breeding of this species , and it is not reported to have been heavily exported in the past ( 2 ) .
however , very little is known about the ecology or conservation needs of this species , and it may therefore benefit from further research before any specific conservation measures , if necessary , can be put into place .
authenticated ( 07 / 03 / 11 ) by dr richard k . b . jenkins , madagasikara voakajy and durrell institute of conservation and ecology , university of kent . urltoken
deciduous forest forest consisting mainly of deciduous trees , which shed their leaves at the end of the growing season . dorsal relating to the back or top side of an animal . endemic a species or taxonomic group that is only found in one particular country or geographic area . territorial describes an animal , a pair of animals or a colony that occupies and defends an area .
gray , j . e . ( 1845 ) catalogue of the specimens of lizards in the collection of the british museum . british museum , london .
halliday , t . and adler , k . ( 2002 ) the new encyclopedia of reptiles and amphibians . oxford university press , oxford .
brady , l . d . and griffiths , r . a . ( 1999 ) status assessment of chameleons in madagascar . iucn species survival commission , iucn , gland , switzerland and cambridge , uk .
nature picture library 5a great george street bristol bs1 5rr united kingdom tel : + 44 ( 0 ) 117 911 4675 fax : + 44 ( 0 ) 117 911 4699 info @ urltoken http : / / www . urltoken
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jennifer hammock added an association between
madagascar dry deciduous forests habitat
and
mesitornis variegatus
.
jennifer hammock added an association between
madagascar dry deciduous forests habitat
and
xenopirostris damii
.
jennifer hammock added an association between
madagascar dry deciduous forests habitat
and
amaurornis olivieri
.
jennifer hammock added an association between
madagascar dry deciduous forests habitat
and
ardea humbloti
.
jennifer hammock added an association between
madagascar dry deciduous forests habitat
and
haliaeetus vociferoides
.
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any of several horned chameleons of the genera furcifer and chamaeleo ; specifically f . rhinoceratus of madagascar , the male of which has a prominent forward - projecting horn on the nose .
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these strange looking chameleons grow up to 27 cm long , with the males growing twice as big as the females .
the horns that give them their name are bigger in the males and they use them to fight with each other .
they spend most of their time in trees where they catch insects using their long tongues .
at breeding time , the females lay 4 - 11 eggs , which take almost a year to hatch .
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cites is an international agreement between governments , aimed to ensure that international trade in specimens of wild animals and plants does not threaten their survival .
the following bibliography has been generated by bringing together all references provided by our content partners . there may be duplication .
3 . gray , j . e . ( 1845 ) catalogue of the specimens of lizards in the collection of the british museum . british museum , london .
5 . halliday , t . and adler , k . ( 2002 ) the new encyclopedia of reptiles and amphibians . oxford university press , oxford .
6 . brady , l . d . and griffiths , r . a . ( 1999 ) status assessment of chameleons in madagascar . iucn species survival commission , iucn , gland , switzerland and cambridge , uk .
andersson , lars gabriel . 1910 . reptiles and batrachians from the north - west of madagascar collected by v . kaudern , 1906 - 1907 . arkiv för zoologi 7 ( 7 ) : 1 - 15 .
boettger , o . 1913 . reptilien und amphibien von madagascar , den inseln und dem festland ostafrikas . pp . 269 - 375 . in : voeltzkow , a . reise in ostafrika in den jahren 1903 - 1905 . wissenschaftliche ergebnisse . vol . 3 . systematische arbeiten . schweizerbart’ sche verlagsbuchhandlung , nägele und sproesser , stuttgart .
boettger , oskar . 1893 . katalog der reptilien - sammlung im museum der senckenbergischen naturforschenden gesellschaft in frankfurt am main . i . teil ( rhynchocephalen , schildkröten , krokodile , eidechsen , chamäleons ) . gebrüder knauer , frankfurt a . m . , x + 140 pp .
glaw , f . & vences , m . 1994 . a fieldguide to the amphibians and reptiles of madagascar . vences & glaw verlag , köln ( isbn 3 - 929449 - 01 - 3 ) .
glaw , f . & vences , m . 2001 . ein seltenes chamäleon aus madagaskar . datz 54 ( 7 ) : 16 - 19 .
gray , j . e . 1843 . description of two new species of reptiles from the collection made during the voyages of h . m . s . sulphur . ann . mag . nat . hist . ( 1 ) 11 : 46 .
gray , j . e . 1845 . catalogue of the specimens of lizards in the collection of the british museum . trustees of die british museum / edward newman , london : xxvii + 289 pp .
gray , j . e . 1865 . revision of the genera and species of chamaeleonidae , with the description of some new species . ann . mag . nat . hist . ( 3 ) 15 : 340 - 354 .
necas , petr . 1999 . chameleons - nature ' s hidden jewels . edition chimaira , frankfurt ; 348 pp . ; isbn 3 - 930612 - 04 - 6 ( europe ) isbn 1 - 57524 - 137 - 4 ( usa , canada ) .
pianka , e . r . & vitt , l . j . 2003 . lizards - windows to the evolution of diversity . university of california press , berkeley , 347 pp . [ review in copeia 2004 : 955 ] .
schmidt , w . ; tamm , k . & wallikewitz , e . 2010 . chamäleons - drachen unserer zeit . natur und tier verlag , münster , 328 pp .
regulatory status : injurious wildlife ( exhibit 5 , chapter 13 - 124 ) . it is illegal to transport these animals between islands or to commercially export to the mainland . penalties can include a fine of up to $ 200 , 000 and a possible prison sentence . the hawaii department of agriculture has an amnesty program allowing a person to turn in an illegal animal without prosecution
jackson’s chameleons are a popular pet that can escape the cage and establish feral populations in hawaiʻi . they are bright emerald green fading to a yellowish color on their undersides . males are generally 10 - 12” long with a long , prehensile tail accounting for half of their length , and with three horns protruding from their forehead . females are generally a little shorter and never grow horns . jackson’s chameleons are solitary creatures which spend most of their time in trees . chameleons move with a distinctive slow rocking motion .
chameleons’ ability to thrive in a variety of forest environments where they are yet another predator of hawaiʻi’s native insects , spiders , and snails , making them a less welcome guest in hawaiʻi . chameleons are also a potential prey base for the brown tree snake and other snakes , which increases the likelihood of this alien species establishing populations if ever introduced .
introduced from kenya and tanzania to hawaiʻi in 1972 , this popular pet was inte ntionally released in kāneʻohe and now has established free ranging populations on the islands of hawaiʻi , maui , and oʻahu . they have not become established on kauaʻi - report any sightings on kauaʻi !
diet of the invasive lizard chamaeleo jacksonii ( squamata : chamaeleonidae ) at a wet - forest site in hawai‘i . fred kraus and david preston : urltoken
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madagascar is home to about half the world ' s 150 or so species of chameleons , including both subfamilies , typical chameleons ( chamaeleoninae ) and dwarf chameleons ( brookesiinae ) .
chameleons are diurnal , solitary , and often aggressive towards members of their own species ( marked by rapid color change and aggressive posturing ) . they are opportunistic hunters that wait for prey to pass within range of their long tongues . chameleons have a bizarre way of moving in which they slowly rock back and forth between each step taken , often in time with the movement of nearby leaves being blown by the wind . most chameleons lay eggs .
this portable guide offers a full survey of all madagascar ' s mammals , both endemic and introduced , including many newly identified species . with vivid color photographs , line illustrations , and maps , mammals of madagascar : a complete guide
home | photos index | search | about | contact unless otherwise noted , all content and images are the property of rhett butler , content copyright 2004 - 2012 . all rights reserved .
this reptile ' s tongue is as long as its body ( up to 55 cm ) . its tip is muscular and sticky . it projects it at an average speed of 21km / hr !
it only takes a few seconds for its color to change . the color varies according to stress , fear , light , etc . it can ' t imitate a complex design such as a flowery carpet or a checkerboard .
its eyes pivot independently from each other , allowing it to look in two opposite directions at the same time . it can therefore keep watch and capture its prey with 99 % precision .
being very territorial , it does not tolerate any same sex individuals closer than two meters . facing an adversary , it can inflate its body and open its mouth while hissing and growling . it can inflict painful bites when fighting .
a popular pet , over 15 , 000 individuals were captured annually in nature . since 1999 , this number has been restricted to 2 , 000 , but logging and agricultural exploitation are fragmenting and rarefying its habitat .
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what a great time . would definitely go again . the zoo care takers are amazing at what they do and so friendly . i would recommend if you have never gone it ' s a bucket list . oh and don ' t forget to go and pet the sting rays .
our whole family attended yesterday . it was absolutely spectacular . we had a fantastic time ! thanks for the memories !
amazing place to take the family ! lots of animals to see , amazing water park and fun rides . great packages available with hotels close by for out of town visitors . highly recommend !
it ' s the 3rd year in a row that my family and i visit this zoo . even though it was during the busiest day they had this year , we were able to visit everything and fully enjoy the experience . don ' t forget to bring your bathing suit cause they have an amazing wave pool !
amazing place . perfect for the whole family . they constantly improve their environments and animals collection .
a good place for family outing with zoo , water park and rides all at one place . all the attractions are conveniently placed very close to each other . food court is good too and you can also bring in your lunch .
we had an excellent time ! we have not been in over 10 years and are so glad we went ! the park is clean . the food was well priced for a theme park . we loved the dinozoo ! we have terrible french speaking abilities but appreciated your courteous and friendly staff , we felt welcome !
i had an amazing time here with my family ! my kids absolutely loved it . amazing experience : )
we love going to the zoo . ( … ) most of the animals were out and the grandkids were thrilled . i ' d never been to the dinozoo before and i thought it was pretty cool especially with the movement and sounds . the design is great in that it ' s seperated and all goes back to the center .
first time visiting here on our annual summer week in québec . we went for the 2 day family pass , and had an amazing time . the 2 days allowed us to take our time around the park , and gave plenty of time as well for cooling off at the amazoo . we ' ll definitely come back !
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the chameleons ' eyes can rotate and focus separately on 180 - degree arcs , which gives them a full 360 - degree field of vision . their tongue can reach its prey in just 0 . 07 seconds , with acceleration reaching over 41 g ' s of force .
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