urotrichini is a taxonomic tribe of the mole family , and consists of japanese and american shrew-moles .

the name " shrew-moles " refers to their morphological resemblance to shrews , while generally being thought of as " true moles " .

the species are the japanese shrew mole , true 's shrew mole and american shrew mole .

in japan , the word " himizu " ( ヒミズ ) may refer to both to the japanese shrew mole in particular and urotrichini in general ; when true 's shrew mole is distinguished from the general himizu forms , the feminine diminutive word " hime " is added to refer to the smaller size of that species .

although they are common in japan , their alpine habitats , small size , and secretive lifestyle makes them generally unknown except among some mountain people and researchers .

urotrichini

, subfamily uropsilinae ) and showed a phylogenetic history moderately different from that of shrew moles ( tribes neurotrichini and urotrichini , subfamily talpinae ) . members of the genus
) focused on less - saturated transversional substitutions , the depth of the calibration point seems too old ( 25–35 mya for the divergence between urotrichini and talpini ) , so that the estimated time scale may be overestimated . the estimates of kirihara et al . (
. . . another plesiomorphic character is the presence of a single and undivided mesostyle in the upper molars ( character 17 # 1 ; character 18 # 1 ) , a feature shared with uropsilus and the urotrichini . however , the teeth of t . minor do not show other specializations present in urotrichini , such as enlarged incisor alveoli and the broadened mandibular ramus ( barrow & macleod 2008 ; sansalone 2015 ) . the only explicit specializations that can be observed in t . minor are an enlarged p1 , which is morphologically different from the caniniform p1 exhibited by the highly fossorial moles ( see in example g . antiquus ; schwermann & martin 2012 ) , and slightly hypsodont molars . . . .
. . . another plesiomorphic character is the presence of a single and undivided mesostyle in the upper molars ( character 17 # 1 ; character 18 # 1 ) , a feature shared with uropsilus and the urotrichini . however , the teeth of t . minor do not show other specializations present in urotrichini , such as enlarged incisor alveoli and the broadened mandibular ramus ( barrow & macleod 2008 ; sansalone 2015 ) . the only explicit specializations that can be observed in t . minor are an enlarged p1 , which is morphologically different from the caniniform p1 exhibited by the highly fossorial moles ( see in example g . antiquus ; schwermann & martin 2012 ) , and slightly hypsodont molars . ziegler ( 2012 ) suggested this increased hypsodonty was an adaptation to drier environments . . . .
the chinese long - tailed mole ( scaptonyx fusicaudus ) closely resembles american ( neurotrichus gibbsii ) and japanese ( dymecodon pilirostris and urotrichus talpoides ) shrew moles in size , appearance , and ecological habits , yet it has traditionally been classified either together with ( viz subfamily urotrichinae ) or separately ( tribe scaptonychini ) from the latter genera ( tribe urotrichini sensu . . . [ show full abstract ]
urotrichini is a taxonomic tribe of the mole family , and consists of japanese and american shrew - moles . they belong to the old world moles and relatives branch of the mole family ( talpidae ) . there are only three species , each of which represents its own genus . the name ` shrew - moles ` refers to their morphological resemblance to shrews , while gene . . . . .
conflicting taxonomic classifications regarding the establishment and composition of subfamilies and tribes have been proposed for the talpid family ( hutchison 1968 ; yates 1984 ; mckenna and bell 1997 ; hutterer 2005 ) , with substantial confusion also surrounding the phylogenetic placement and taxonomic assignment of talpids in the fossil record ( ziegler 2003 , 2012 ; klietmann et al . 2015 ) . because of the robustness of the forelimb in some talpids , relatively plentiful numbers of humeri are preserved as fossils for some lineages ( e . g . , talpini , scalopini , urotrichini ) , and can provide insights into locomotory adaptations through geological time . however , the resolution of locomotory adaptations through time requires a well - resolved phylogeny , which is currently not available .
) . tribal divergences commenced during the priabonian stage ( 39 . 5–33 . 9 ma ) of the late eocene when scalopini diverged from other tribes at 37 . 0 ma ( 95 % ci = 39 . 5–35 . 3 ma ) , and continued through the rupelian ( 33 . 9–28 . 1 ma ) and chattian ( 28 . 1–23 . 03 ma ) stages of the oligocene when the three shrew mole tribes ( scaptonychini , urotrichini , neurotrichini ) diverged from each other between 30 and 27 ma ( 95 % ci = 34–23 ma ) . divergences between chinese and north american scalopins , between european and asian talpins , and between the two long - tailed mole lineages all occurred in the mid - miocene ( ca . 16 ma ; 95 % ci = 22–10 ma ) . finally , divergences between sister genera generally occurred between 18 and 7 my ( 95 % ci = 26–4 ma ) . the lineage - through - time ( ltt ) plot (
members of the mammalian family talpidae—the moles , shrew moles , and desmans—include some of the least studied mammals on earth . the 43 recognized living species reflect a surprisingly diverse set of lifestyles including terrestrial , semi - fossorial , semi - aquatic , fossorial , and aquatic - fossorial ( nowak 1999 ; hutterer 2005 ) . the evolution of adaptive specializations to these diverse environments produced substantial morphological variation , for example in skeletal anatomy ( freeman 1886 ) and sensory systems ( catania 2000 ) . however , increasingly derived “stepping - stone” phenotypic adaptations can be identified across a morphological gradient from the terrestrial shrew - like moles ( uropsilinae ) , to the semi - fossorial shrew moles / long - tailed moles ( neurotrichini , urotrichini , and scaptonychini ) , and further to the fossorial “true moles” ( talpini and scalopini ) . conversely , the more aquatic members of the family , the desmans ( desmanini ) and the aquatic - fossorial star - nosed mole ( condylurini ) , are among the most unique and morphologically isolated of living mammals .
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parent taxon : talpinae according to o . g . bendukidze et al . 2009
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mammal species of the world : a taxonomic and geographic reference , 3rd ed . , vols . 1 & 2
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to elucidate the origins of the japanese mammalian fauna from the perspectives of biogeography and community ecology , i reviewed molecular phylogenetic and phylogeographic studies for all non - volant terrestrial mammals indigenous to the japanese archipelago ( 63 species ) , with a particular focus on obtaining reliable chronological data . the results of this review demonstrate that geological vicariance events in the tsugaru and korea ( tsushima ) straits can explain the distribution of many japanese mammals , in particular the hokkaido - endemic species with late pleistocene origins and the honshu–shikoku–kyushu - endemic species with middle pleistocene or earlier origins . phylogenetic relatedness also contributed to the observed patterns of distribution through the processes of competitive exclusion and species assortment , and abiotic environmental filtering was another important factor . later colonists of honshu–shikoku–kyushu , from northern hokkaido or the southern tsushima islands , were mostly excluded owing to the competitive dominance of earlier residents or environmental filtering . on the other hand , the fragmented distributions of some species with more ancient origins in both hokkaido and honshu–shikoku–kyushu may be a result of the competitive dominance of later migrants . ecological coexistence can be achieved by phylogenetically dispersed species , supporting the principle of species assortment . because almost all aspects of the mammalian faunal assembly in japan can be explained by geological events or community ecological processes , the japanese archipelago may be an ideal model island system in which to study the mechanisms of faunal assembly .
i am deeply grateful to masaharu motokawa and hiroshi kajihara for inviting me to contribute to their comprehensive book on japanese animals , and for their encouragement during the manuscript preparation process . i thank masaharu motokawa , hon - tsen yu , and an anonymous reviewer for providing valuable feedback on an earlier version of this manuscript . i also thank hitoshi suzuki and past and present members of his laboratory for their invaluable assistance with my research on the molecular phylogenetics and phylogeography of mammals .
here , i have reviewed molecular phylogenetic and phylogeographic studies of japanese mammals to obtain reliable estimates of the divergence time between the japanese and the continental species ( or lineage ) and the time to the most recent common ancestor ( mrca ) of the japanese mammals . where no chronological data were provided in the previous study , even though some dna sequences were determined and available in the dna database , i briefly calculated these times by using the lineage - specific evolutionary rate . it should be noted here that the most commonly used evolutionary rates proposed by brown et al . (
applied this rate to all the mammalian lineage divergence ) , so that the applications of these rates to other groups would be difficult because of the differences in the evolutionary rates among mammalian groups . for example , a corollary of the application of the evolutionary rate obtained from a slowly evolving lineage ( e . g . , ungulates ) into a rapidly evolving lineage ( e . g . , rodents ) is an overestimation of the divergence times on the rapidly evolving lineage .
) . these species could be classified into two subfamilies , soricinae and crocidurinae ( dubey et al .
) genes , these subfamilies were demonstrated to have diverged in the late early miocene [ ca . 20 mya ( million years ago ) ( fumagalli et al .
species in hokkaido established their lineages by different histories in spite of the similar distribution patterns . however , in ohdachi et al . (
) , the time scale for the lineage differentiations was not assessed based on their molecular data . hope et al . (
including the hokkaido linage was in the period of the late pleistocene ( < 0 . 13 mya ) . this time estimate is in agreement with the distribution pattern of
species evolution . however , because the evolutionary rate of hope et al . (
species might have fallen into circular argument . nevertheless , a similar evolutionary rate was suggested to be fit to the late quaternary phylogeography of various taxa from rodents to carnivorans around the bering straits ( hope et al .
) . therefore , i used the 11 % / myr ( million years ) divergence rate in this tentative time estimations . the average pairwise genetic distances between species endemic to the hsk region (
s , respectively ) were calculated to be 8 . 3 % and 8 . 8 % , respectively , thus suggesting that these lineages diverged 0 . 75 and 0 . 8 mya in the early to middle pleistocene , respectively . on the other hand , hokkaido lineages of
s were estimated to have diverged 0 . 07 and 0 . 56 mya , respectively . it is obvious to obtain the age estimate consistent with hope et al . (
because of the same assumption about the evolutionary rate . however , the middle pleistocene origin of the hokkaido lineage of
was inferred to have split 0 . 09 mya , implying that the lineage diversifications in hokkaido might have occurred in the late pleistocene . meanwhile , it was estimated that
, where the average pairwise distance between khabarovsk and hokkaido haplotypes and that among the hokkaido haplotypes both show 1 . 6 % corresponding to 0 . 15 mya based on the 11 % / myr divergence rate , suggesting that the hokkaido lineage diverged from the continental one in the final part of the middle pleistocene . regarding
, it was calculated that the divergence between the hokkaido and the continental lineages , that among three haplogroups in hokkaido , and that within each of the three haplogroups , occurred 0 . 07 , 0 . 05 , and 0 . 02 mya , respectively . all the events took place in the late pleistocene . the foregoing calculation suggests that the absence of
, whose origin in hokkaido is in the middle pleistocene , did not cross the tsugaru strait , when the land bridge was formed . probably , the lineage expansion would have occurred in the late pleistocene , as suggested in
at 3 . 5 mya ) . however , special caution must be given to the interpretation of their results . first ,
gene whose substitutions would be saturated in the time scale of their study , where the calibration point was set to approximately 20 mya , that is , at a level subjected to the saturation problem ( steppan et al .
, i suppose that their time estimates could be more realistic because they estimated divergence times by less - saturated nuclear protein - coding genes and obtained dates largely concordant with the fossil records ( he et al .
) estimated 1 . 25 mya , as already explained , 3 . 03 mya estimated for the split between
) could correspond to age about 0 . 83 mya by a very brief proportion calculation . furthermore , yuan et al . (
in taiwan is 10 . 5 % . if i use the 11 % / myr of hope et al . (
) , the time estimate for this divergence would be 0 . 95 mya , which is close to 0 . 83 mya , although slightly larger .
) clarified that there are four geographically separated phylogroups in the japanese archipelago , one in kyushu and the other three in honshu . the most southern kyushu lineage first diverged , then the chugoku lineage in the southwestern honshu , and in the last kinki lineage in the east of the chugoku region and the lineage in east and central honshu were separated . the time to the mrca of the four phylogroups was inferred to be 0 . 39 mya , and divergence in each phylogroup was estimated to have occurred within 0 . 059 mya , on the basis of the 1 . 36 % transversional susbtitutions at the third codon positions per million years calculated from setting the divergence time between soricinae and crocidurinae to be 20 mya , according to fumagali et al . (
) , in which the transversional susbtitutions at the third codon positions was checked to be not saturated . in addition , by using the 11 % / myr of hope et al . (
) where average pairwise genetic distances between haplotypes in kyushu and east - central clades is 3 . 0 % , the divergence time was calculated to be 0 . 27 mya , which is not inconsistent with the middle pleistocene origin of
as estimated earlier ( 0 . 39 mya ) . all these inferences suggest that
) . as already described , because the tsugaru strait was not considered to have had a land bridge in the late pleistocene , the absence of this species from hokkaido can be explained by the late pleistocene expansion of the honshu lineages from each refugium . the reason for the absence of this species from the shikoku islands is not clear .
( amami , okinawa , miyako , and yaeyama isl . , bp10 ; table
) genes , and estimated that the crocidurinae shrew species were inferred to have diversified on mainly palaearctic and oriental regions since the late miocene . one of major clades derived from the diversifications is the asian clade , where all the four
) . using the calibration point of 20 mya for the divergence between soricinae and crocidurinae , dubey et al . (
, whose phylogenetic affinity was not unambiguously inferred , was generated 5 . 42 mya during the earliest diversification among the asian clade .
in tsushima islands has not been dated in any of these previous studies , data used by motokawa et al . (
gene between the tsushima lineage and the most closely related cheju lineage was 1 . 2 % , suggesting that the divergence time is 0 . 1 mya on the basis of the 11 % / myr divergence rate of hope et al . (
including the japanese lineage have never been investigated . thus , the time scale cannot be evaluated here . concerning the other intraspecific genetic diversity of these crocidurinae species , it was suggested that there is a clear phylogeographic demarcation between eastern and western japanese lineages in
) . their average pairwise genetic distance is 3 . 7 % , indicating that evolutionary split between eastern and western linages occurred 0 . 34 mya based on the 11 % / myr divergence rate . it has also been known that individuals of
( zemlemrova et al . 2013 ) ] . no talpid species are observed in hokkaido . the biogeographic patterns for the talpid species are bp6 [ the lesser japanese shrew mole
) . their precise distributions are not considered to overlap except for the narrow geographic boundaries among them . in fact , the distribution of
predominantly occupies the western part of japan and possesses the same species as in the eurasian continent . these distribution patterns suggest the different evolutionary history among these mole and shrew mole species . previous molecular phylogenetic analyses have clarified that the asian fossorial moles formed a monophyletic lineage (
, i could not discuss the origin of this species hereafter ) . namely , among the japanese mole species ,
at the last . combined with the distributions of these moles in the japanese archipelago , these phylogenetic relationships suggest that the japanese mole species migrated from the eurasian continent following the branching order in the phylogeny , implying that the earlier the migration is , the more restricted the distributions are to montane regions , islands and small areas , and the northeastern region in honshu . on the basis of the fossil record ( 16 . 4–20 . 5 mya for the earliest talpini fossil in east asia and 4 . 0–4 . 3 mya for the oldest
lineage was estimated to be 17 . 1 mya in the analyses using the mitochondrial
( both eurasian and japanese lineages are included ) 5 . 7 mya . on the other hand , using the similar calibration standard based on the fossil records , he et al . (
split occurred 3 . 42 to 4 . 5 mya , thus indicating estimates younger than those in shinohara et al . (
) . this difference in the time estimates would be caused by the stronger effect of the mitochondrial genes in shinohara et al . (
branched off in this order 12 . 6–17 . 7 , 6 . 6–9 . 2 , and 2 . 5–3 . 4 mya , respectively ( tsuchiya et al .
branched off in this order 5 . 21 , 3 . 64 , 2 . 31 , and 1 . 20 mya , respectively . these time estimates are much younger than those in other studies . i suppose that the divergence times inferred by shinohara et al . (
) might be overestimates because they depended on the mitochondrial genes in some degree for the time scale , where the mitochondrial genes would not work well because of the saturation problem , and also adopted the calibration point based on the earliest record of talpini in east asia ( 16 . 4–20 . 5 mya ) , which might also be too old for the mitochondrial gene variations to be calibrated . although tsuchiya et al . (
) are in agreement with the geological and eustatic records and the migration ages of the proboscidean species into japan . kirihara et al . (
gene ( 4–6 % / myr ) , these two “species” diverged 0 . 3 to 0 . 5 mya ; this is a level of intraspecific variation ( kirihara et al .
endemic to the japanese archipelago , their origins have not been clearly understood . previous molecular phylogenetic analyses demonstrated that these two shrew mole species formed a clade ( shinohara et al .
) . however , no time scale has been estimated yet . furthermore , as no talpid species is closely related to one of these species ( that is , the clade consists of these two species only ) , it is not clear which species migrated into the japanese archipelago first . taking into account that
( average pairwise genetic distance was 11 . 4 % ; data were derived from shinohara et al .
show a very high extent of intraspecific variations corresponding to the interspecific difference in other mole and shrew mole species ( shinohara et al .
are 7 % , 10 . 6 % , and 6 . 9 % , respectively , on the basis of the data of shinohara et al . (
) , which could be calculated to be 1 . 17–1 . 75 mya , 1 . 77–2 . 65 mya , and 1 . 15–1 . 73 mya , respectively , following a 4–6 % / myr
) . assuming that the divergence between asian and barbary macaques occurred 3 . 0 mya , based on the fossil record and using a portion of the mitochondrial dna sequences (
0 . 65 to 0 . 73 mya and the differentiation among three japanese lineages took place 0 . 19 to 0 . 37 mya . on the other hand , tosi et al . (
) genes for the divergence time estimations postulating that the initial divergence of macaques occurred 5 . 5 mya , caused by the messinian salinity crisis . in this study , the time for the branching of
from the other closely related species above was estimated to be 1 . 0 mya in the y - chromosomal gene tree . although not estimated clearly , the mitochondrial gene tree suggested that
diverged in a time younger than 1 . 2 mya , which is consistent with the y - chromosomal gene estimation ( tosi et al .
) and also is not contradicted by the estimates of hayasaka et al . (
was estimated to be 0 . 38 to 0 . 42 mya ( chu et al .
into the eastern marginal islands of the eurasian continent . although the exact time of the migration has not been fixed yet , the last estimate by chu et al . (
experienced a rapid expansion 0 . 16 to 1 . 00 mya ( marmi et al .
with more individual samples from the northernmost shimokita peninsula ( located in the northern part of aomori prefecture in northernmost honshu ) to the southernmost yakushima islands ( located in the south of the kyushu islands ) by the control region sequences , demonstrated that there are major eastern and western lineages in the japanese archipelago , and suggested that the eastern lineage experienced the recent northward demographic expansion from a refugium in the post - glacial periods in the late quaternary .
) , in which almost all the species are observed in europe and africa . the japanese dormouse
is the only species within gliridae that inhabits east asia , it is extremely difficult to infer the origin of this species in japan by only investigating the extant species . nevertheless , nunome et al . (
( 25 mya ) that their divergence occurred 24 to 30 mya under the background of major glirid diversifications during a relatively warmer climate period from the early oligocene ( 34 mya ) to the mid - miocene climatic optimum ( 15–17 mya ) . these time estimates suggest that the origin of the japanese dormouse may have predated the formation of the japanese archipelago ( about 15 mya ; neall and trewick
) . additionally , it could be argued that the past 15 million years have seen declines of east asian dormice because of the climate change into colder and drier environments , which the dormice do not favor ( zachos et al .
possesses nine geographic lineages that diversified 3 to 5 mya . yasuda et al . (
) concluded that the divergences into these local populations occurred within the japanese archipelago . furthermore , they showed that the extent of the genetic divergence among the geographically separated populations corresponds to that of interspecific variations in other mammalian groups . thus , they also suggested that each local population can be regarded as a cryptic species .
) . there are three types of ecological lifestyles among squirrels [ arboreal ( tree squirrels ) , terrestrial ( ground squirrels ) , and gliding ( flying squirrels ) locomotors ] . recent molecular phylogenetic analyses of the multiple nuclear and mitochondrial genetic loci have determined five major lineages corresponding to subfamilial level and suggested that the arboreal tree squirrel is the ancestral type that would have had a key role in the major squirrel diversifications and subsequent derivation of the terrestrial and gliding squirrel lineages ( mercer and roth
) , which are assigned to two of these major clades , a clade of arboreal and gliding squirrels ( sciurinae in steppan et al .
) and a clade of mostly ground squirrels ( xerinae in steppan et al .
( terrestrial , but , in fact , intermediate between terrestrial and arboreal ) ( steppan et al .
the japanese sciurid species show clear distributional trends . namely , congeneric species do not share the same distribution although species in the different genera do coexist in the same region (
in the hsk region ) . hence , it follows that there are only two biogeographic patterns for the squirrels on the japanese archipelago , bp2 ( table
species in the family soricidae . although some phylogenetic and phylogeographic studies have been known for
) , few of these examined time scales for the lineage differentiations or demographic events . oshida and masuda (
to be 4 . 0 to 5 . 2 mya based on the 0 . 5 % transversional substitutions / myr of the
) . this value would be an overestimate , probably because of the use of the underestimated substitution rate ( from the application of the slowly evolving ungulate rate into the rapidly evolving rodent time scale ) . later , oshida et al . (
gene considering that the divergence rate was suggested to be variable among rodents within the range of 5–10 % / myr [ see brunhoff et al . (
) for the rationale ] . if i adopt the 5–10 % / myr of divergence rate for the case of
was inferred to be 4 . 9–6 . 8 % , the divergence time between these species is 0 . 49–1 . 36 mya ( 4 . 9–6 . 8 % divided by 5–10 % ) , which is more consistent with the palaeontological evidence suggesting that the lineage of
lineage migrated into the japanese archipelago via northern or southern routes is not easy , the origin from the korean peninsula , the southern route , may be plausible because their absence in hokkaido can more easily be explained . however , even if it is the case , it is difficult to understand why the ancestral lineage did not reach hokkaido despite the multiple chances of migration via the land bridges formed in the tsugaru strait since the early to middle pleistocene period . probably , the population expansion might have occurred in the late pleistocene as in
is not clear because there have been no molecular chronological studies comparing the continental and hokkaido lineages . if i calculate their divergence time on the basis of the
; hokkaido , korean , and transbaikalian individuals ) , it was inferred that the hokkaido lineage diverged from the monophyletic korean and transbaikalian lineage 0 . 18 to 0 . 36 mya in the middle pleistocene , where the average between genetic distances of the hokkaido / korea ( 1 . 3 % ) and the hokkaido / transbaikalia ( 2 . 3 % ) was divided by the 5–10 % / myr divergence rate ( oshida et al .
from europe to east asia and suggested the recent rapid population expansion , although no time frame was provided . my tentative hypothesis is the middle pleistocene origin for the hokkaido lineage of
. however , more rigorous phylogeographic studies are needed for future precise time estimations .
, the hokkaido population was clarified as a monophyletic lineage ( oshida et al .
) , and the divergence from the eurasian lineages was estimated to have taken place 0 . 2 to 0 . 4 mya based on the 5–10 % / myr divergence rate for the
, no studies have been conducted on the intraspecific genetic diversity . comparison of the
) enabled me to estimate that the average sequence difference between them was 12 . 3 % , implying that they diverged 1 . 2 to 2 . 5 mya on the basis of the 5–10 % / myr divergence rate as above ( oshida et al .
are mainly distributed in the southern parts of the eurasian continent rather than the northern eurasian regions . among them ,
, endemic to japan , is the most divergent lineage within the genus ( oshida et al .
lineage took place around the late miocene period ( about 8–10 mya ) . li et al . ( 2013 ) also examined the
gene and obtained a much older time estimate ( 12 . 5 mya ) for the origin of the
( 0 . 6–1 . 3 mya ) as the calibration points . however , these time values might be results of overestimations : first , because of using the underestimated evolutionary rate of irwin et al . (
gene to estimate reliable divergence times because of the severe saturation effect as has been noted . if i apply the 5–10 % / myr divergence rate of
and the other congeneric species was estimated to have occurred 1 . 39 to 3 . 06 mya , which is more similar to or slightly more ancient than the date of the origin of similarly the hsk - endemic sciurid species ,
) . the phylogeographic analyses of the mitochondrial d - loop region by oshida et al . (
in the japanese archipelago and suggested that the kyushu is the ancestral region for the diversification of this species because the kyushu population has the most divergent or ancestral haplotypes and two of three major lineages . combined with the presence of the closely related
. the divergence among the three major lineages was estimated to have occurred 0 . 4 to 1 . 0 mya based on the evolutionary rate of the human d - loop sequences ( 8 . 6 % / million years ; vigilant et al .
gene and found that there are five distinct phylogroups ( kyushu , southwest , southeast , central , and north ) , including the three major lineages of oshida et al . (
) . using 5–10 % / myr for the divergence rate , they estimated that diversifications among five major lineages occurred 0 . 09 to 0 . 24 mya . these estimates are in agreement with the divergence time between
and the closest species ( 1 . 39–3 . 06 mya ) . additionally , oshida et al . (
, in which the other species are all found in the american continent . there have been few phylogeographic studies including the hokkaido lineage of
) examined one individual from hokkaido together with many samples from russia , china , and korea . they showed that there are at least three divergent lineages in this species , northern eurasian , korean , and central chinese lineages ( but the validity of the last one is still open to question because this is based on pet shop samples with unknown origin ) . the japanese chipmunk is included in the northern eurasian lineage . although their approach for the divergence time estimation is arbitrary because they forced the divergence among these major lineages to have occurred in the onset of the middle pleistocene period without clear explanations for the calibration procedure , the differentiations among lineages within the northern eurasian clade , including the lineage of hokkaido , was calculated to have transpired 0 . 11 to 0 . 47 mya in the middle to late pleistocene period . here again , if i apply the 5–10 % / myr rule to the data of obolenskaya et al . (
) , the divergence time among the northern eurasian lineages was estimated to be 0 . 10 to 0 . 19 mya ( average genetic difference among 90 individuals within the northern eurasian clade , 0 . 95 % , divided by 5–10 % ) , which is congruent with the estimate of obolenskaya et al . (
, all belonging to the subfamily murinae . the murinae is the most diversified subfamily within the muridae , and the interrelationships among the major lineages ( genera ) within this subfamily have remained to be elucidated because of the rapid diversifications during a relatively short evolutionary time span ( sato and suzuki
lineage endemic to nepal , the lineage mainly occupying the east asia , and the lineage mainly observed in europe . four species are found in japan , the striped field mouse
divergence at 12 mya . because more closely related species are missing in continental asia , it is supposed that the formation of the japanese archipelago might have affected the generation of such highly divergent endemic species . because the time estimates for the generation of
( 5 . 9 mya ) correspond to the late miocene age that saw the vegetation changes promoted by the environmental shift to a colder and drier climate ( cerling et al .
from all the major islands and neighboring small islands , and estimated , with the evolutionary rate of 2 . 4 % / lineage / myr ( suzuki et al .
separating central ( honshu , shikoku , and kyushu islands ) and peripheral ( hokkaido , sado , izu , and satsunan islands ) lineages . tomozawa and suzuki (
) for the central / peripheral trend , although the hokkaido and sado lineages were included in the central lineage , the satsunan lineages were observed in both lineages , and the oki and tsushima lineages were added to the peripheral lineage . tomozawa and suzuki (
) discussed that allopatric fragmentation might have caused the generation of the central and peripheral lineages , and the recent population expansion at 0 . 14 mya ( based on 2 . 4 % / lineage / myr as earlier ) formed the current distribution pattern of the central lineage , where the male - biased dispersal might replace the nuclear genome of hokkaido , sado , and a part of satsunan populations with the peripheral mtdna types that remained because of female philopatry . it should be noted here that although all the dating estimates are fundamentally based on the frequently used divergence time between
( 12 mya ) , the recent palaeontological ( 11 . 0–12 . 3 mya ; benton and donoghue
) and molecular phylogenetic ( 8 . 6–10 . 3 mya ; steppan et al .
) studies have gradually supported younger dates than 12 mya . therefore , the real divergence and population expansion times could also be more recent than obtained here based on the 12 mya for the
and clarified the monophyly of the hokkaido population to the continental one . the estimate for the divergence time of the hokkaido lineage from the continental equivalent in the most recent study was 0 . 1 mya in the late pleistocene , based on fossil calibrations that the divergence between
) . both these studies indicated that the population from siberia to the russian far east harbors a higher extent of genetic diversity , therefore implying the existence of refugia in the glacial periods in the pleistocene . probably the hokkaido lineage is a result of the population expansion from the refugial region in far east asia ( serizawa et al .
) and mainly favors open environments ( e . g . , grasslands ; ohdachi et al .
) , which is suggestive of the recent rapid population expansion across wide areas in eurasia . yasuda et al . (
) estimated on the basis of 2 . 4 % / lineage / myr ( suzuki et al .
) that the divergence between the japan / korea and european lineages occurred 0 . 08 mya in the late pleistocene and also suggested that the population expansion within the japanese archipelago occurred 0 . 03 mya . such a recent establishment of this species in japan is in agreement with the lack of any fossil records of this species in japan ( ohdachi et al .
) that the korea ( tsushima ) strait was not available by land bridges in this age . i discussed the possible process of the origin of
in the text ( oversea dispersal hypothesis ) . here again , also note that dating with the 2 . 4 % / lineage / myr rate of suzuki et al . (
gene lineage that is not found in the other populations in southeast asia and concluded that it was established by a natural dispersal instead of an anthropogenic effect . on the other hand , shimada et al . (
in ryukyu islands was shown to be closely related to the laos lineage within a strongly supported clade of the southern southeast asian clade and did not show close affinity with the geographically close taiwan lineage . because of this ambiguous status , i did not consider this species in discussions .
) , and the divergence time between these species was estimated to be 6 . 5–8 . 1 mya by nuclear gene analyses (
) suggested with rdna - rflp data that the lineages in amami–oshima and tokunoshima diverged 1 . 2 to 2 . 3 mya based on the previously estimated evolutionary rate of 1–2 % / myr , whereas the time was 4 . 4 mya based on a partial
gene data although it depended on the evolutionary rate of brown et al . (
was 5 . 4 % , which could be calculated to be 2 . 58 and 1 . 13 mya , respectively , on the basis of the 2 . 4 % / lineage / myr substitution rate ( suzuki et al .
split , which is not consistent with the current knowledge as already noted ( 8 . 6–10 . 3 mya , steppan et al .
) . if i adopt the average 10 . 55 mya { ( [ 8 . 6 + 10 . 3 ] / 2 + [ 11 + 12 . 3 ] / 2 ) / 2 } , 20–30 % means 2 . 11–3 . 17 mya for the divergence of the lineage for
) that the ryukyu islands were connected multiple times to the eurasian continent in two stages , the former of which was 1 . 3 to 1 . 6 mya .
) for the genus name , but basically adopted the common names and species numbers in ohdachi et al . (
) . the origins of these three species in hokkaido are considered independent of each other . kohli et al . (
) . they found that there are distinct phylogroups that would have been derived from at least three refugia located in western eurasia , central eurasia , and beringia . the divergences among these phylogroups were estimated to have occurred during the last 0 . 1 million years in the late pleistocene . it was also suggested that the monophyletic hokkaido lineage was possibly originated by the expansion from a northern refugium , beringia . if the establishment of this species in hokkaido originally occurred in the late pleistocene , it can explain the absence of this species in more southern japanese islands because of the same reason for
at 2 . 5 mya ) that the monophyletic hokkaido lineage diverged from the clade mainly composed of the far east russian and sakhalin lineages 0 . 27 mya and experienced the population expansion 0 . 04–0 . 05 mya in hokkaido . although abramson et al . (
) did not provide intra - lineage dating information , the most recent common ancestor of the hokkaido individuals could be estimated to be present in the late pleistocene ( < 0 . 13 mya ) , inferred from the branch length of the phylogeny that they provided . therefore , the absence of
) was estimated to have occurred in the middle pleistocene ( 0 . 81 mya ; abramson et al .
) . based on the results of the mitochondrial dna control region , kawai et al . (
) detected four phylogroups from the several fragmented populations in hokkaido and concluded that such phylogroups were shaped by the genetic divergences across several refugia formed in the glacial periods since the middle pleistocene . this conclusion is because in their study the time to the mrca of the
phylogroups was estimated to be 0 . 12–0 . 58 mya in the middle pleistocene based on the “mutation” rate of 3 . 6 % / myr ( based on the divergence time of 7–8 mya between
) and 17 % / myr ( based on the divergence time of 1 . 8 mya for the siberian and the nearctic brown lemmings ; fedorov and stenseth
) estimated for the arvicoline control region diversity . however , despite that these rates were originally proposed as “divergence” rates considering two descendant lineages , kawai et al . (
) adopted them as per - lineage mutation rates . therefore , the foregoing estimates ( 0 . 12–0 . 58 mya ) should be corrected to 0 . 24–1 . 16 mya . the corrected estimates also largely include the middle pleistocene period and are not inconsistent with the divergence of the
lineage as estimated above ( 0 . 81 mya ) . it is not clear why
did not reach more southern japanese islands despite the presence of the land bridge in the tsugaru strait in the middle pleistocene , but a community ecological mechanism may explain the absence in honshu and more southern islands .
) . the former was mainly found in the eastern honshu and the latter in the western honshu , shikoku , and kyushu , while their distributions are partly overlapped in the central honshu . their ancestral common lineage was inferred to have diverged 0 . 9 to 2 . 3 mya from the clade composed of
) . time scales for the differentiations have not been fully assessed to date in a reliable manner . assuming that the time to the mrca of the red - backed voles was 1 . 8 mya , luo et al . (
) estimated with the fossil calibration ( 2 . 6 mya for the time to the mrca of
0 . 50 mya . these time estimates might be too recent to be regarded as those for valid species differentiation . iwasa and suzuki (
species in honshu by morphological criteria only . probably future taxonomic revision would be needed on the basis of more rigorous morphological and molecular phylogenetic studies for the
species in the hsk region . alternatively , it is also probable that the ancestral polymorphisms of both species have been retained since the divergence between
. female philopatry might have influenced the maintenance of such divergent lineage in the mitochondrial gene . iwasa and suzuki (
gene , which is male specific and irrelevant for the female philopatry . also in this case , all the populations may be assigned to one
) and includes mainly herbivorous small rodents adapted to grasslands , taiga , steppe , and tundra . there is only one
) . there have been few studies that examined chronological aspects of this species based on molecular data . bannikova et al . (
gene and the time constraint of 2 . 2 mya for the radiation of the basal
diverged from the aforementioned closely related species 0 . 95 mya . the time estimate is similar to that of the
species in hsk . in the fossil evidence , this species was found since the middle pleistocene ( kawamura et al .
could have expanded to all the japanese islands , hokkaido , honshu , shikoku , and kyushu . the absence of this species from hokkaido and shikoku requires some explanation in light of phylogeography or ecology . kaneko (
and species in the same guild would shed more light on the reasons for the peculiar distribution pattern of this species .
) . in japan , 3 species are present and their distributions are geographically partitioned . the mountain hare
from the other genera was 9 . 44 mya in the total data analysis and 8 . 63 mya in the nuclear gene data analysis on the basis of the palaeontological records ( leporidae–ochotonidae split , 20–40 mya ; the origin of the modern leporid , 12–20 mya ; the divergence of the genus
, 4–6 mya ; the oldest divergence time of the ingroup , 60 mya ) . probably the former value ( 9 . 44 mya ) would be overestimated because the mitochondrial genes were included in the examined data matrix . the latter estimate ( 8 . 63 mya ) is not contradicted by the late miocene origin of the murine rodent genus
, also endemic to the ryukyu region ( 6 . 5–8 . 0 mya ; sato and suzuki
was not resolved on the earlier radiation among different leporid genera . assuming the molecular clock of the
from each sister lineage were estimated to be 12–16 mya , 4–5 mya , and 0 . 5–0 . 6 mya , respectively ( yamada et al .
lineage was 3 . 62 mya , which is not in disagreement with the estimate within the pliocene in yamada et al . (
( 12–16 mya ) is much earlier than that of matthee et al . (
) ( 8 . 63 mya ) , which mainly used nuclear genes . the difference would be attributed to the distinct properties between mitochondrial and nuclear genes for dating divergences , as repeatedly discussed earlier . therefore , the divergence times of yamada et al . (
) might be somewhat overestimated because of the exclusive use of the mitochondrial genes . if i conduct a simple calculation that 8 . 63 mya is 62 % of 14 mya ( average between 12 and 16 myas ) and apply this proportion to the other estimates of yamada et al . (
are calculated to be 2 . 48–3 . 10 mya and 0 . 31–0 . 37 mya , respectively .
) detected major two lineages ( northern and southern lineages ) and suggested several regions for refugia during the pleistocene glacial periods ( kanto , chubu , shikoku , and kyushu ) , where the genetic diversity was estimated to be higher . these two major clades were inferred to have diverged 1 . 2 mya following the lineage - specific evolutionary rate of 1 . 4 % / lineage / myr ( therefore , 2 . 8 % divergence rate ) on the basis of the assumption that
gene also indicated a similar but a little younger divergence time between northern and southern lineages ( 1 . 07 mya ; nunome et al .


cuspivolva tigris , known commonly as the tiger egg cowry , is a species of sea snail , a marine gastropod mollusk in the family ovulidae , the ovulids , cowry allies or false cowries .

cuspivolva tigris

[ cuspivolva tigris , syn . : c . renovata , crenavolva tigris , ovulum dentatum , primovula tigris ]
what type of species is cuspivolva tigris ? below , you will find the taxonomic groups the cuspivolva tigris species belongs to .
which photographers have photos of cuspivolva tigris species ? below , you will find the list of underwater photographers and their photos of the marine species cuspivolva tigris .
how to identify cuspivolva tigris marine species ? below , you will find the list of main identification criteria and physical characteristics of marine species cuspivolva tigris . for each identification criteria , the corresponding physical characteristics of marine species cuspivolva tigris are marked in green .
cuspivolva tigris orange - ovulidae - philippines sea shell - 9 . 6mm - lot 5
where is cuspivolva tigris found in the world ? below , you will find the list and a world map of the geographic distribution where the marine species cuspivolva tigris can be found .
tiger egg cowrie ( cuspivolva tigris ) is . . . - nomad divers , bangka | facebook
cuspivolva tigris orange - ovulidae - philippines sea shell - 9 . 6mm - lot 5 on ebid australia | 126313157
cuspivolva renovata iredale , t . , 1930 : c indo - w pacific ( nomen novum , nomen dubium )
( of primovula tigris yamamoto , 1971 ) lorenz f . & fehse d . ( 2009 ) the living ovulidae . a manual of the families of allied cowries : ovulidae , pediculariidae and eocypraeidae . hackenheim : conchbooks . [ details ]
( of crenavolva tigris ( yamamoto , 1971 ) ) lorenz f . & fehse d . ( 2009 ) the living ovulidae . a manual of the families of allied cowries : ovulidae , pediculariidae and eocypraeidae . hackenheim : conchbooks . [ details ]
( of primovula tigris yamamoto , 1971 ) cate c . n . ( 1973 ) . a systematic revision of the recent cypraeid family ovulidae . the veliger . 15 ( supplement ) : 1 - 117 . , available online at urltoken [ details ]
( of crenavolva tigris ( yamamoto , 1971 ) ) cate c . n . ( 1973 ) . a systematic revision of the recent cypraeid family ovulidae . the veliger . 15 ( supplement ) : 1 - 117 . , available online at urltoken [ details ]
lorenz f . & fehse d . ( 2009 ) the living ovulidae . a manual of the families of allied cowries : ovulidae , pediculariidae and eocypraeidae . hackenheim : conchbooks . [ details ]
my favourite cowrie is the wonderful tiger egg cowrie . it’s taken me 4 years in indonesian to finally find one . another great find in ambon .
like all the cowries , the amazing patterns and colours are not actually on their shells . the colours and patterns you see come from the cowrie’s mantle shirt , which is the top of the cowrie’s foot . it extends out and up to completely cover the shell . this is why it is so important not to collect these creatures , especially if you think it would look so lovely on your desk… . not only should we not be taking anything from the ocean , but in this case you would be left with a white shell .
you can see the tiger cowrie is feeding on particles collecting on the sponge it is on . the tube - like mouth is called the proboscis and is fascinating to watch as it seems to be gulping down its meal .
they are about 1 . 5cm ( 5 / 8” ) long and are very rare .
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english - german online dictionary developed to help you share your knowledge with others . more information ! contains translations by tu chemnitz and mr honey ' s business dictionary ( german - english ) . thanks on that account ! links to this dictionary or to single translations are very welcome ! questions and answers
version 43 . 0 went live 11 / 6 / 2018 - i hope that the majority of issues have been fixed . my email address is on the home page if you see anything wrong .
ovulum dentatum sowerby , g . b . ii in adams , a . & l . a . reeve , 1848 : singapore - taiwan ( preoccupied )
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( g . b . sowerby ii in a . adams & reeve , 1848 )
delonovolva formosa ( g . b . sowerby ii in a . adams & reeve , 1848 )
ovulum formosum g . b . sowerby ii in a . adams & reeve , 1848
primovula ( delonovula ) formosa ( g . b . sowerby ii in a . adams & reeve , 1848 )
the living ovulidae : a manual of the families of allied cowries : ovulidae , pediculariidae and eocypraeidae .
in : okutani , t . ( ed . ) , marine mollusks in japan . tokai university press , tokyo , 211 - 223 ( in japanese ) .
accessed through : world register of marine species at urltoken on 2014 - 10 - 03 .
a new species of primovula from japan , parasitic to a gorgonid ( gastropoda : ovulidae ) .
publications of the seto marine biological laboratory , 19 ( 4 ) , 191 - 195 .
occurrence record in darwincore format ( elements of obis schema and some of dwc1 . 4 )
uncommon to rare in the solitary islands marine park ( simp ) . recorded at south solitary island and korff ' s islet , just outside the simp . recorded from japan , indonesia and australia .
found from depths of 18 - 70m , this ovulid feeds only on the gorgonian coral euplexaura sp .
the mantle of this ovulid is a delicate orange with black stripes bordered with white . the foot is orange with black spots . the siphon is orange and the tentacles are orange tipped with black and white . the shell is uniformly yellowish orange and grows to 15mm .
all information was correct at the time of publication and is subject to change without notice . copyright 2014 , solitary islands underwater research group inc . ( abn : 38 104 639 980 ) - all rights reserved .
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i sold 4 more of my pin badges tonight , so over 100 sales now . the same newbie who purchase 4 of the 4th february has come back for 4 more . i ' ll have to get busy and list some more soon .
42 created tue 10 jul 2018 05 : 23 : 17 ( aest ) . copyright © 1999 - 2018 ebid ltd
the living ovulidae . a manual of the families of allied cowries : ovulidae , pediculariidae and eocypraeidae .
c . n . cate & azuma in c . n . cate , 1973
updated on 9 july 2018 with 703 different species : 404 gastropods , 34 worms , 34 cephalopods , 42 frogfishes and scorpionfishes , 50 pipefishes and seahorses , 139 crustaceans . database updated on a daily basis as a result of encounter logbook compiled by guests and guides .
note : si incontrano esemplari marrone chiaro con bordo dei parapodia marrone / grigio ed esemplari marrone scuro col bordo dei parapodia bluastro .
note : notare il peduncolo arancione sul fianco , parziale estrazione del membro riproduttivo .
note : si differenzia da c . strigata per la presenza di arancione fra le righe longitudinali nere e inoltre bordo del mantello , rinofori e ciuffo branchiale sono più scuri .
note : già glossodoris , è simile a d . sibogae da cui si differenzia per colletto dei rinofori bianco e per tonalità bianco / nero piuttosto che giallo / nero .
note : già glossodoris , è simile a d . atromarginata da cui si differenzia per colletto dei rinofori nero e per tonalità giallo / nero piuttosto che bianco / nero .
note : avvistati fra le uova deposte da altri nudibranchi , spesso fra quelle rosse deposte da hexabranchus sanguineus .
note : stadio adulto , con colore del corpo a predominante marrone e punte dei cerata arancioni .
note : stadio semi - adulto , il corpo è ancora poco marrone e le estremità dei cerata non sono ancora arancioni .
note : stadio giovanile , l ' animale è quasi solamente bianco / azzurro .
note : inusualmente si incontrano esemplari dalle tinta scolorite che possono raggiungere anche gli 80 mm .
note : nel ciuffo branchiale sovente si intravvedono un paio di sacche ovigere deposte da un parassita ( colore arancione ) .
note : nel ciuffo branchiale sovente si intravvedono un paio di sacche ovigere deposte da un parassita ( colore arancio ) .
note : si ciba di altri nudibranchi , anche di pari o maggiori dimensioni come hypselodoris apolegma , aggredendo ciuffo branchiale o rinofori .
note : da non confondere con h . malesso , in cui le righe arancioni sono più rade ed evidenti , oltre ad essere longitudinali ai cerata .
note : simile a h . batangas ( notare le differenze nello spessore e disposizione delle righe arancioni che caratterizzano il corpo ) .
note : simile a h . bullockii , la differenza più evidente è nel bianco bordo del mantello , per h . bullockii sottile e netto , per h . apolegma sfuma .
note : se ne incontrano due varianti di colore , talvolta anche in accoppiamento , una blu - viola e l ' altra bianco - avorio .
note : molto simile ad alcuni esemplari di h . kanga , questo ha una sola riga rossa per ciascun rametto del ciuffo branchiale ( h . kanga ne ha due ) .
note : alcuni esemplari sono molto simili a h . infucata , ma questo ha due righe rosse per ciascun rametto del ciuffo branchiale ( h . infucata ne ha due ) .
note : se ne contano diverse varianti , sia come colori che come disegni del corpo .
note : molto simile a h . maculosa , ma è considerata specie a sè .
note : molto simile a h . marittima , ma è considerata specie a sè .
note : simile a h . apolegma e a h . bullockii , ma è considerata specie a sè . notare il bordo del mantello , largo e non sfumato .
note : molto simile a hypselodoris sp . 20 , il ciuffo branchiale ha diverso colore .
note : molto simile a hypselodoris sp . 19 , il ciuffo branchiale ha diverso colore .
note : nel ciuffo branchiale sovente si intravvedono un paio di sacche ovigere deposte da un parassita ( colore viola ) .
note : può avere rinofori e ciuffo branchiale da blu a viola ed avere o meno le macchie arancioni lungo il bordo del mantello .
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