oedignathus inermis is a species of king crab found off the pacific coasts of the united states and canada , from california to alaska , and disjunctly around the coasts of japan .

it is the only species in the genus oedignathus , and is sometimes called the granular claw crab , paxillose crab or tuberculate nestling lithode crab .

oedignathus

where are you likely to find oedignathus inermis ? how could you tell it apart from petrolisthes spp . ? what does it presumably eat ( two food types ) ?
security for granular claw crabs , oedignathus inermis , is an abandoned barnacle shell . they have to locate protection because their soft - shelled abdomens are vulnerable and nutritious . the specific epithet , inermis , means unarmed . some people call o . inermis soft - bellied crabs . they are much sought by predators .
these crabs are scientifically called oedignathus inermis . they are usually found in large numbers across the pacific coast of the usa , from california to alaska . a feature that differentiates them from other species is the large number of eminences which are visible on the planate chelipeds and leg areas . they usually reside underneath purple - colored algae .
shallow - water representatives of the hapalogastrinae ( oedignathus , hapalogaster , cryptolithodes ) and lithodinae ( paralithodes , lopholithodes ) have average egg diameters ranging from 0 . 63 to 1 . 15 mm and are significantly smaller ( p < 0 . 01 ) than those produced by deep - sea genera ( neolithodes , lithodes and paralomis ) .
( of oedignathus gilli benedict , 1895 ) mclaughlin , p . a . ; komai , t . ; lemaitre , r . ; listyo , r . ( 2010 ) . annotated checklist of anomuran decapod crustaceans of the world ( exclusive of the kiwaoidea and families chirostylidae and galatheidae of the galatheoidea . part i – lithodoidea , lomisoidea and paguroidea . the raffles bulletin of zoology . supplement no 23 , 5 - 107 . [ details ] available for editors [ request ]
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urltoken
mclaughlin et al . 2005 . common and scientific names of aquatic invertebrates from the united states and canada : crustaceans . american fisheries society special publication 31
williams , austin b . , lawrence g . abele , d . l . felder , h . h . hobbs , jr . , r . b . manning , et al .
disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
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mclaughlin , p . a . ; komai , t . ; lemaitre , r . ; listyo , r . ( 2010 ) . annotated checklist of anomuran decapod crustaceans of the world ( exclusive of the kiwaoidea and families chirostylidae and galatheidae of the galatheoidea . part i – lithodoidea , lomisoidea and paguroidea . the raffles bulletin of zoology . supplement no 23 , 5 - 107 . [ details ] available for editors [ request ]
liu j . y . [ ruiyu ] ( ed . ) . ( 2008 ) . checklist of marine biota of china seas . china science press . 1267 pp . ( look up in imis ) [ details ] available for editors [ request ]
( of hapalogaster inermis stimpson , 1860 ) mclaughlin , p . a . ; komai , t . ; lemaitre , r . ; listyo , r . ( 2010 ) . annotated checklist of anomuran decapod crustaceans of the world ( exclusive of the kiwaoidea and families chirostylidae and galatheidae of the galatheoidea . part i – lithodoidea , lomisoidea and paguroidea . the raffles bulletin of zoology . supplement no 23 , 5 - 107 . [ details ] available for editors [ request ]
( of hapalogaster brandtii schalfeew , 1892 ) mclaughlin , p . a . ; komai , t . ; lemaitre , r . ; listyo , r . ( 2010 ) . annotated checklist of anomuran decapod crustaceans of the world ( exclusive of the kiwaoidea and families chirostylidae and galatheidae of the galatheoidea . part i – lithodoidea , lomisoidea and paguroidea . the raffles bulletin of zoology . supplement no 23 , 5 - 107 . [ details ] available for editors [ request ]
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) . the first ( basal ) segment of the abdomen has calcified plates , as do the two terminal ones .
, and small spines but no large spines . note , however , that there are large spines on the anterior margins of leg 1 ( the
to 3 cm long and 2 . 5 cm wide in males , 2 cm wide in females ; wider posteriorly than anteriorly , brown with scales on the dorsal surface . it has white - centered orange granules and dark brown spots , but these colors may be masked by mud . may have white on the sternum . there are few if any
amchitka island , ak to monterey , ca ; eastern russia , japan , korea . mostly on the open coast . rarely seen in the san juan islands and is said to not to occur in puget sound ; rare in california .
these crabs are often found in pairs , and may be in such a tightly secluded space that they appear to be trapped . they feed by straining plankton from the water with their third
. captive individuals also catch worms and small crustaceans with their small claw and crush mussels with the large claw . predators include black oystercatchers .
the abdomen of this species is thick and soft . the basal segment and the two distal segments have some calcified plates , which are not evident in this view .
one claw is much larger than the other . the
palm
of the
the rostrum is short . the carapace has orange - red tubercles with a white spot in the middle .
granular claw crab found in an intertidal area of calvert island . note the granules covering the larger claw . photo by cody gold .
in : nishimura , s . ( ed . ) , guide to seashore animals of japan with color pictures and keys , vol . ii . hoikusha , osaka , 347 - 378 ( in japanese ) .
annotated checklist of anomuran decapod crustaceans of the world ( exclusive of the kiwaoidea and families chirostylidae and galatheidae of the galatheoidea ) . part i - lithodoidea , lomisoidea and paguroidea .
japanese crustacean decapods and stomatopods in color , vol . i . macrura , anomura and stomatopoda .
occurrence record in darwincore format ( elements of obis schema and some of dwc1 . 4 )
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king crabs are well - known due to their exceptionally big size and their ability to reside in cold waters . this article provides some basic facts about the various species of king crabs .
king crabs are one of the most hunted sea dwellers . there are different species of these creatures which are found in seas all around the globe . they are also known as ' stone crabs ' due to their appearance . they prefer to live in freezing cold waters , whereas other species of crabs are normally found in warmer waters . they are the largest amongst all kinds of crabs , and have a great commercial demand and importance . some of their species are used as food by many people due to their size and taste . japanese and american restaurants are famous for preparing king crab recipes . around 40 species of these crabs are known till now . the most common are red , blue , and golden king crabs , which are generally found in alaskan waters .
these crabs are usually hunted in alaska . their scientific name is paralithodes platypus . the ones which are caught in the pribilof islands are the largest among the blue king crabs . they have a brown - colored body with blue highlights on it . they have exceptionally big claws which seem really dangerous .
these crabs are also known as lithodes aequispinus , and are generally found in regions from the british columbia to the aleutian islands , and also japan . these are relatively smaller in size in comparison with other species . as their name suggests , they have a golden - colored outer covering .
these are also called golf - ball crabs , and are normally found at depths of about 10 - 75 meters . their shell is triangular in shape , and approximately seven centimeters in length . there are numerous spines and bristle - like structures present on the legs of these crabs .
they are scientifically known as lithodes couesi , and are smaller in size . they are found in large numbers , which lessens their commercial value .
fishing for king crabs is largely carried out in alaska . due to this reason , the government has implemented regulations on overfishing to save these king crabs from becoming extinct .
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dorsoventrally flattened , carapace as long as or wider than long ; carapace with linea anomurica ; outer orbital spines absent ; rostrum overreaching distal corneal margin , or not overreaching bases of corneas . eye cornea well developed ; ocular acicles absent .
bases widely separated ; crista dentata present ; accessory tooth present ; dactylus simple .
all of similar form ; 2 - 4 with basis and ischium fused ; dactyli of pereopods 2 to 3 simple . pereopod 3 about the same length as pereopod 2 ; pereopods 3 dactyli and propodi of right and left similar . pereopod 4 simple .
partially divided ; sternite of pereopod 5 reduced , contiguous with preceding sternite ; somite of pereopod 5 not fused with first abdominal somite , or somite of pereopod 5 fused with first abdominal somite .
3 - 5 absent ; none modified as gonopods . male with no other sexual modifications ; female with first pleopods paired and modified as gonopods . uropods absent .
cite this publication as : ' mclaughlin , p . , s . ahyong & j . k . lowry ( 2002 onwards ) . ' anomura : families . ' version : 2 october 2002 . urltoken ' .
the infraorder anomura has long captivated the attention of evolutionary biologists due to its impressive morphological diversity and ecological adaptations . to date , 2500 extant species have been described but phylogenetic relationships at high taxonomic levels remain unresolved . here , we reconstruct the evolutionary history—phylogeny , divergence times , character evolution and diversification—of this speciose clade . for this purpose , we sequenced two mitochondrial ( 16s and 12s ) and three nuclear ( h3 , 18s and 28s ) markers for 19 of the 20 extant families , using traditional sanger and next - generation 454 sequencing methods . molecular data were combined with 156 morphological characters in order to estimate the largest anomuran phylogeny to date . the anomuran fossil record allowed us to incorporate 31 fossils for divergence time analyses .
our findings are compared against current classifications and previous hypotheses of anomuran relationships . many families and genera appear to be poly - or paraphyletic suggesting a need for further taxonomic revisions at these levels . a divergence time analysis provides key insights into the origins of major lineages and events and the timing of morphological ( body form ) and ecological ( habitat ) transitions . living anomuran biodiversity is the product of 2 major changes in the tempo of diversification ; our initial insights suggest that the acquisition of a crab - like form did not act as a key innovation .
the infraorder anomura represents a highly diverse group of decapod crustaceans comprised of hermit crabs , mole crabs , king crabs , squat - lobsters and porcelain crabs . the fossil record contains representatives of nearly all extant families and spans the norian / rhaetian ( late triassic ) [
] and the common use of hermit crabs as pets in the aquarium trade . moreover , some species are threatened or endangered due to rarity in nature , e . g . , pylochelidae [
] . thus , improved understanding of these groups bears not only on appreciation of their diversity and ecology , but also strategies for their conservation .
] ] . early classifications from the 19th to the first half of the 20th centuries were based on adult morphological characters including mouthparts , antennae , gills , pleon type , and / or larval characteristics . these classifications often differed in higher - level composition and , in some cases , the infraordinal name ( e . g . anomura vs . anomala ) . since these studies , various researchers have proposed changes in the classification scheme [
] , many of which remain actively debated . more recently , molecular and / or morphological data have been used to reevaluate anomuran relationships [
one of the most debated evolutionary questions within anomura is phylogenetic relationships between hermit and king crabs . since the early 1800’s [ e . g . , [
] ] , studies have suggested king crabs and hermit crabs are close relatives , despite first appearances to the contrary . king crabs are among the largest arthropods and have a crab - like body shape , whereas hermit crabs are relatively small and depend on a shell for protection . despite glaring morphological differences as adults , an affinity between king crabs ( lithodoids ) and hermit crabs ( paguroids ) has been long suggested [
] . although most accept this claim , the evolutionary pathways and hypothesized ancestor of both groups has been debated for decades , with two major hypotheses being proposed . the first suggests that the lithodids (
) ( “hermit to king hypothesis” ) while the second suggests the opposite evolutionary pathway ( “king to hermit hypothesis” ) . here we revisit these hypotheses in light of new phylogenetic data to test the “hermit to king” / “king to hermit” evolutionary pathway .
additional controversy over anomuran relationships stems from apparently rampant examples of convergent and / or parallel evolution in body forms . anomurans span an impressive array of body configurations that include : 1 ) crab - like forms 2 ) squat - lobster forms 3 ) hermit crab forms with pleonal ( abdomen ) symmetry ( found in 1 hermit crab family ) and 4 ) hermit crab forms with pleonal asymmetry ( found in 4 hermit crab families ) . recent studies suggest that the acquisition of a crab - like body form , known as carcinization [ see , [
] , possibly impacting diversification rates within these lineages . for the first time , we explore diversification patterns in anomura and specifically test if carcinized lineages underwent unusually rapid diversification rates . if the emergence of the crab - like form promoted diversification we would expect the overall rate in carcinized lineages to be high compared to net of diversification across anomura . additionally , we test if the acquisition of different body forms ( i . e . , crab - like , squat - lobster - like , pleonal asymmetry and symmetry ( hermit ) ) arose once or multiple times during the emergence of the anomurans and reconstruct the evolutionary pathways of these transitions .
divergence dating is a powerful tool used to estimate the timing and origins of diversity , morphological traits , habitat shifts , and diversification . although nearly all the family - level groups of anomura are represented in the fossil record , the discovery has not been as frequent as that of other decapod groups ( i . e . , true crabs , lobsters ) . two factors , variations in cuticular sclerotization and habitat preference , are likely responsible for the limited occurrence of anomuran fossils . many taxa are weakly calcified , whereas others possess well - calcified claws and poorly calcified carapaces and pleons . in addition , habitats currently occupied by anomurans , including freshwater , terrestrial , intertidal marine , deep marine , and hydrothermal vent areas are strongly underrepresented in the fossil record . despite these limitations , we incorporate 31 fossil calibrations to estimate the origin of lineages and major events during anomuran evolutionary history , including the transition of body forms and shift into freshwater and terrestrial environments .
here , we present the taxonomically broadest and largest dataset yet assembled . we combine sequences generated by traditional sanger and next - generation 454 sequencing methods with morphological characters , including 19 / 20 extant families and 137 species , to estimate phylogenetic relationships , character state evolution , divergence times , and diversification patterns among major lineages of this diverse clade of crustaceans . our comprehensive sampling , in combination with modern integrative approaches , allows us to present the most complete evolutionary picture for the infraorder anomura to date .
) . alternative outgroup sampling schemes did not affect internal relationships among anomura . the optimal models of evolution for each gene selected in modeltest were as follows : gtr + i + g 18s , 28s , h3 and tvm + i + g 12s , 16s . several sequences downloaded from genbank were excluded from the analysis due to contamination after a blast search and / or strange alignment results ( see additional file
an “n / a” not available indicates missing sequence data . new sequences are indicated as kfxxxxxx .
alternative outgroup selections did not affect internal anomuran relationships . with all outgroups included , brachyura was recovered as the sister taxon . the bayesian analysis from the combined molecular + morphology dataset recovers anomura as a monophyletic group with high support ( 100 = pp , figure
) . the majority of the nodes ( 86 % ) are recovered with very high support ( > 95 ) . three families are recovered as para - or polyphyletic ( diogenidae , paguridae , munididae ) . with the exception of three families ( blepharipodidae , kiwaidae , lomisidae ) each having a single representative , the remaining families were found to be monophyletic ( hippidae , albuneidae , munidopsidae , galatheidae , porcellanidae , parapaguridae , aeglidae , eumunididae , chirostylidae , lithodidae , hapalogastridae , pylochelidae , and coenobitidae ) with high support . blepharipodidae , hippidae , and albuneidae ( hippoidea ) group together with very high support ( 100 ) , being sister to the remaining 16 anomuran families . lomisidae , eumunididae , kiwaidae , and chirostylidae ( lomisoidea + chirostyloidea ) form a clade with high support ( 100 ) and are sister to aeglidae ( aegloidea ) . munidopsidae , galatheidae , munididae , and porcellanidae ( galatheoidea ) form a clade with high bayesian support ( 100 ) . within the galatheoidea , munididae is paraphyletic with the galatheids nested within the group . pylochelidae , parapaguridae , diogenidae , coenobitidae , paguridae , hapalogastridae , and lithodidae ( = paguroidea + lithodoidea ) form a statistically supported clade ( 97 ) . six of the seven anomuran superfamilies are monophyletic ( hippoidea , galatheoidea , aegloidea , lomisoidea [ monotypic ] , chirostyloidea , and lithodoidea ) . the remaining superfamily , paguroidea is found to be paraphyletic and includes the superfamily lithodoidea ( lithodidae + hapalogastridae ) . 11 genera were found to be poly - or paraphyletic (
combined bayesian phylogram based on molecular ( 3669 characters ) and morphological ( 156 characters ) data . vertical colored bars represent anomuran families , grey brackets represent superfamilies , and the black vertical line represents outgroups . bayesian posterior probabilities represented as percentages and > 70 % are noted above or below branches .
) is similar to our combined phylogeny , with most differences being found in placement and composition of paguroidea . unlike the combined phylogeny , which recovered paguroidea as paraphyletic , paguroidea was found to be polyphyletic . the family pylochelidae was recovered as polyphyletic according to molecular data but was monophyletic when morphology was added . parapaguridae was sister to a clade containing pylochelidae , aeglidae , lomisidae , eumunididae , kiwaidae , and chirostylidae , similar to the results of tsang et al . [
] based on nuclear protein coding genes . as in the combined phylogeny , coenobitidae is nested within the diogenidae , and lithodoidea nested within the paguroidea . within lithodoidea of the molecular–only phylogeny , hapalogastridae was found to be paraphyletic , with representatives of the genera
) end of the tree . however lithodoid relationships in the molecular - only phylogeny should be interpreted with caution as many were recovered with little to no support . in the combined phylogeny hapalogastridae was found to be a monophyletic and sister to lithodidae ( figure
some deep splits and short branches in the molecular - only phylogeny should be interpreted with caution , as support is low .
bayesian phylogram based on 5 genes 12s , 16s , 18s , 28s , h3 and 3669 characters . vertical colored bars represent anomuran families , grey brackets represent superfamilies , and the black vertical line represents outgroups . bayesian posterior probabilities represented as percentages and maximum likelihood bootstrap values are noted above or below branches .
] were tested using the shimodaira - hasegawa test ( s - h ) . three of the seven hypotheses were found to be significantly worse than our unconstrained topology (
= −68430 . 951016 ) . hypotheses that tested a “king to hermit” evolutionary pathway were all significantly worse than the alternative ( i . e . , “hermit to king” ) as recovered in our best ml tree (
a ) . analyses indicated that a crab - like ancestor gave rise to all extant anomuran lineages . in addition to the earliest branching clade , hippoidea , carcinization occurred independently three times during the evolution of the group , twice through squat lobster - like intermediaries ( squat intermediary = si on tree ) and once through an asymmetrical hermit crab - like ancestor ( asymmetrical hermit intermediary = ahi on tree ) ( figure
a ) . the squat lobster - like form arose once as an early branching lineage and gave rise to the crab - like clades , lomisidae and porcellanidae . within the hermit crab lineages , the symmetrical pleon arose once within the pylochelidae . the asymmetrical pleon arose once within the paguroidea , but was subsequently partially reverted to the ancestral symmetrical condition ( in males only ) within the crab - like lithodidae and hapalogastridae ( = lithodoidea , figure
b ) . in combination with divergence time results , we can make predictions about the timing of these events ( see discussion ) . maximum parsimony and maximum likelihood methods recovered similar ancestral state reconstructions for body form and habitat ( figure
ancestral state reconstruction analysis using maximum likelihood methods for body shape and habitat transition within anomura . colored taxa correspond to anomuran families as noted in legend . pie charts represent the likelihood of the ancestral state . ( a ) character states for body shape were defined as crab - like white , squat lobster blue , symmetrical hermit green and asymmetrical hermit black . ( b ) character states for habitat were defined as freshwater white , semi - terrestrial green , and marine black . subtrees are shown for the transition into freshwater ( aeglidae ) and semi - terrestrial habitats ( coenobitidae ) .
) . all parameters reached convergence for individual runs . beast estimated the divergence of the anomurans from the true crabs , brachyura , to be in the permian ( ~ 259 ( 224–296 ) mya , figure
, square g ) . the origin of the asymmetrical hermit crab lineages followed soon after in the pliensbachian , early jurassic ( ~ 187 mya , square h ) . two hermit crab families were recovered as non - monophyletic assemblages ( diogenidae , paguridae ) , which resulted in multiple timing of origins for these families . parapaguridae split from one clade of diogenidae (
) in the bathonian , middle jurassic ( ~ 167 mya , square i ) , while the family coenobitidae is found nested within a slightly older clade of diogenidae ( ~ 173 mya , square j ) , which includes most present day genera . paguridae is not monophyletic , because of the internally nested lithodidae and haplogastridae . the most recent common ancestor of the pagurid + lithodid + hapalogastrid clade was placed in the late cretaceous ( cenomanian , ~ 98 mya , square k ) with lithodidae and hapalogastridae splitting from one another around 18 mya ( burdigalian , miocene ) .
) . divergence time estimates ( my ) are noted adjacent to their respective nodes and blue nodal bars correspond to the 95 % highest posterior density regions . geological periods are superimposed onto the phylogeny and listed as follows : d , devonian ; c , carboniferous ; p , permian ; tr , triassic ; j , jurassic ; k , cretaceous ; t , tertiary . colored taxa correspond to anomuran families as noted in the legend . green boxes indicate a diversification shift .
] was used to detect whether any clade within the anomuran tree was best explained by independent diversification models , and to specifically address whether acquisition of the crab - like form resulted in an increase of diversification rates . the background tempo of diversification across the anomuran tree is characterized by a speciation rate
) . a slow speciation rate is detected in the lineage leading to the monotypic and carcinized family lomisidae , and an increase rate occurred in the squat - lobster family chirostylidae . the ancient but species - depauperate branch leading to the monotypic family lomisidae was optimally modelled separately with maximum likelihood estimate of
of 0 . 054182 ( rate acceleration ) . all three resulting clade - specific diversification models were optimally fit as yule models ( aic = 339 . 3032 ) .
] to resolve phylogenetic relationships . these studies have dramatically increased our understanding of anomuran relationships and resulted in several major changes within higher - level classification [
) . as mentioned previously , anomurans have undergone dramatic changes in higher - level classification based on recent phylogenetic studies . galatheoidea has been revised recently to exclude aeglidae , kiwaidae , and chirostylidae [
] . with the recent revision of galatheoidea , all superfamilies were recovered as monophyletic ( i . e . , hippoidea , aegloidea , lomisoidea , chirostyloidea , galatheoidea , lithodoidea ) , except for paguroidea ( figures
] for review of literature ] , based on morphological characters including mouthparts , gills , and pleonal characters . however , the evolutionary pathways of the two groups continue to be debated ( see also “hermit to king , king to hermit evolutionary hypotheses” ) with all recent evidence pointing to a “hermit to king” hypothesis .
) . galatheidae was found nested inside munididae , but alternative topologies that recovered munididae as monophyletic were not significantly worse than our best estimate ( see results ) . deeper sampling within both families is needed to resolve family and genus level relationships . the families galatheidae , munidopsidae , and porcellanidae were all recovered as monophyletic with high support ( figure
] . alternative hypotheses proposing the monophyly of these families ( i . e . , diogenidae , paguridae ) were rejected using s - h tests , confirming our findings ( see results ) . coenobitidae ( semi - terrestrial hermit crab ) was deeply nested within diogenidae ( left - handed hermit crabs ) while
] , which he collectively called the paguristinen . the families pylochelidae and hapalogastridae were found to be polyphyletic in the molecular analysis ( figure
generic relationships within anomura seem to be much less resolved than superfamily and family level relationships . we found several genera to be poly - or paraphyletic ( i . e . ,
] . most instances of non - monophyly occur within highly speciose genera ( i . e . ,
, suggesting deeper sampling and continued research needs to be undertaken on these groups .
although past studies have shown an affinity between paguridae ( hermit crabs ) and lithodidae ( king crabs ) , the evolutionary pathways and ancestry of these anomuran lineages have been debated for the past two centuries . the traditional and prevalent hypothesis posits that lithodids are free - living hermit crabs that abandoned shell use and underwent a series of morphological changes ( carcinization ) resulting in a crab - like form . it has been argued that the asymmetry of the lithodid female pleon , in particular , is evidence of asymmetrical hermit crab ancestry . boas [
] similarly derived the lithodids from the pagurids , agreeing with boas on the structural pleonal similarities between these two groups . however , bouvier also proposed a series of gradual and linear progressive stages in the transformation of the pagurid pleon , starting from a pagurid precursor to various genera of hapalogastrids (
) . in modern times , this concept of pagurid and lithodid evolution was brought to attention when cunningham et al . [
] supported this same evolutionary view of pagurid and lithodid evolution . recently , a study that examines the hemolymph vascular system in hermit and king crabs found close similarities in arterial systems of the dorsal cephalothorax [
] . based on observations of the complex changes in pleonal tergites from megalopa to juvenile crab stages , these studies demonstrated that adult lithodid pleonal tergite structure in several species was the result of decalcification and sundering , not secondary calcification and fusion as had been proposed by bouvier .
] . with the largest number of taxa and most robust molecular / morphological dataset ever used in a phylogenetic study of anomurans , our study once again shows lithodoidea to be nested within paguridae . moreover , our conclusions are consistent with the fossil record , which suggest hermits are much older ( jurassic ) than king crabs ( miocene , table
< 0 . 05 ) ( i . e . , “hermit to king” ) ( see results ) .
while there is undeniable evidence of a close relationship between hermits and king crabs , it is less clear how morphological changes associated with carcinization may have proceeded within the lithodoidea . a recent study comparing hermit and king crab circulatory systems identified several vascular changes that occurred as the result of carcinization , arguing for more comparative studies that look at morphology ( both internal and external ) and development [
] . however , only with a clear phylogenetic hypothesis can many of these studies be correctly interpreted . recent molecular or combined morphological - molecular phylogenies recover conflicting evolutionary relationships , although only three lithodoid genera ( and not always the same , or excluding hapalogastridae ) have been used in previous analyses [
) shows the less carcinized and less calcified hapalogastridae as sister to lithodidae , in agreement with virtually every study since bouvier’s in the 19th century . but within lithodidae , and in contrast to bouvier’s linear hypothesis , our study places
) . it thus appears that the process of heavy calcification may have appeared at least twice in lithodid lineages . more lithodoid genera / species are needed to examine the process of carcinization within the lithodoidea and to properly test bouvier’s and boas’ earlier hypotheses ( explaining the transition of a shell - dwelling hermit crab to a fully calcified lithodid crab ) . in conclusion , while recent , modern studies , including ours , overwhelmingly and clearly support a “hermit to king” evolutionary scenario , it is also clear that the evolutionary process and concomitant morphological changes ( particularly in pleonal tergal plates and pleopods ) that occurred within the lithodoidea to produce the various degrees of crab - like forms in that family , is at best poorly understood .
” could be the most likely candidate for lithodoid ancestry . the close relationship between
- like hermit crab as the precursor to the crab - like lithodoids . all species of
are tube - dwellers , not shell - dwellers , and show pleonal asymmetry only in having unpaired pleopods . the genus is relatively small in size compared to the typically large - sized lithodoids with a distribution across both sides of the north pacific , from the sea of japan to puget sound and the straits of juan de fuca , washington [
) in similar areas . future studies with increased sampling within these groups will shed light into the evolutionary pathway of lithodoids from paguroid ( possibly
] . although this date is considerably older than the hippoid fossil record , closely related extinct forms extend into the triassic and present day hippoidea are found in substrates underrepresented in the fossil record . the superfamily hippoidea containing blepharipodidae , hippidae , and albuneidae , has been described as being similar to primitive brachyurans [
a ) . the next radiation occurred in the late triassic , giving rise to the squat - lobsters and crab - like superfamilies chirostyloidea and galatheoidea , aegloidea , lomisoidea , and the hermit crab and crab - like superfamilies paguroidea and lithodoidea . our results suggest these superfamily clades were derived from a squat - lobster - like ancestor approximately ~ 205 mya ( figures
with a possibly squat - lobster - like body form , dates back to the late triassic ( ~ 201 . 6 - 228 mya ) and has strong morphological affinity with the superfamilies chirostyloidea and galatheoidea . this fossil was found as part of a biotic assemblage suggesting that
around 137 mya a squat - lobster like ancestor gave rise to a unique superfamily of anomurans , aegloidea . aegloid crabs represent the only freshwater anomuran family and can be found in caves , lakes , and streams throughout the neotropical region of south america [
] . in combination with our divergence time analyses , we hypothesize that the complete transition in freshwater occurred sometime between the late cretaceous and miocene . this transition appears to have allowed for rapid diversification approximately 13 mya ( 20–7 . 4 mya ) .
from approximately 180 mya to 147 mya , the families of galatheoidea radiated and diversified . these include the squat lobsters families munidopsidae , munididae and galatheidae , and the porcelain crab family porcellanidae . the porcellanids diverged in the middle jurassic ( ~ 172 mya ) from squat - lobster like ancestors , but a crab - like body form evolved by the tithonian ( ~ 151 - 145 . 5 mya ) based on fossil evidence and ancestral reconstruction analyses . this was the first occurrence of carcinization from a squat - lobster or hermit - like ancestor within anomura ( figures
] suggested porcellanid crabs were derived galatheids despite the differences in body shape and form , and this is consistent with our current evolutionary hypothesis .
lomisoidea and chirostyloidea diverged around 122 mya from a squat - lobster like ancestor . this body form was retained within the chirostyloids and underwent further carcinization , attaining a crab - like form in the monotypic lomisidae , endemic to australia .
] . in our combined analysis , the hermit crab families , pylochelidae , parapaguridae , diogenidae , coenobitidae , and paguridae , formed a monophyletic group with the inclusion of lithodidae or king crabs , and hapalogastridae . we estimated these families arose early in the evolution of anomura , approximately 205 mya . the symmetrical hermit crabs , pylochelidae , are unique with most having complete body symmetry and in utilizing broken gastropod shells , siboglinid tubes , and coral pieces for shelter and protection , in contrast to other hermit groups that commonly use coiled gastropod shells [
) . this is consistent with our divergence time analysis , which recovers these families as early branching lineages . diogenidae , coenobitidae , and paguridae typically possess an asymmetrical pleon accompanied by an enlarged right or left chela . according to our combined analysis , pleonal asymmetry in hermits appears to have been derived once in the evolution of the anomurans , most probably between 200–187 mya . this contrasts with the results obtained by tsang et al . [
] , who proposed that the pleonal asymmetry evolved independently in two different hermit crab lineages , once in parapaguridae , and a second time in diogenidae , coenobitidae , and paguridae . these contrasting differences are the result of incongruent phylogenies based on total evidence ( molecular + morphology , this paper ) and molecular only approaches [
] . note , however , that our molecular - only analyses recover similar results to those of tsang et al . [
) . carcinization occurred for the third time in the crab - like superfamily lithodoidea between 29–18 mya from an asymmetrical hermit - like ancestor . this estimation is consistent with other timing estimates of king crab carcinization [
the crab - like body form was recovered in our study as the ancestral state for all the anomurans . in our study , all alternative body forms were present ( crab - like , squat lobster , symmetrical hermit , and asymmetrical hermit ) early in the divergence of the anomurans . from these ancestral character states , carcinization occurred independently three times during the evolution of anomura , once in the lithodoidea through an asymmetrical hermit intermediate , and twice in lomisidae and porcellanidae through squat lobster intermediates ( see ahi and si , figure
] . however , our tree differs significantly from tsang et al . ’s study [
] in the deep ancestral origins of carcinization . tsang et al . ’s hypothesis suggests a symmetrical hermit crab - like ancestor predated the squat lobster and asymmetrical intermediaries , whereas we recovered a crab - like ancestor to predate these intermediaries . we acknowledge that our analysis recovers two deep nodes that are unresolved , however symmetrical reconstruction at these nodes seems unlikely ( figure
a ) . it must also be noted that the most recent common ancestor of anomura is unresolved in the tsang et al . analysis , although it appears to be a crab - like or symmetrical hermit ancestor . the major differences in the two analyses stems from the differences in phylogeny and more specifically the monophyly ( our study ) or polyphyly [
] of paguroidea and families therein ( i . e . , pylochelidae ) . there is agreement with tsang et al . in the sister group relationship between paguridae and lithodoidea , although tsang et al . used only four lithodid genera ( vs . eight in our study ) and did not include representatives of hapalogastridae . in addition , both studies provide strong evidence for the intermediary ancestors directly predating carcinization across anomura ( twice through squat lobster ( si ) and once through asymmetrical hermit ( asi ) , figure
the multiple cases of carcinization among the anomurans have been noted since the early 1900s . borradaile ( 1916 ) was the first to propose the term carcinization to explain the crab - like aspects of the hermit crab
and the tendency of anomurans to achieve this form , a phenomenon unique to anomura not evident in other decapod lineages ( e . g . , lobsters , shrimp ) . the emergence of the crab - like form is not ‘evenly distributed’ across our phylogeny , first occurring in the older lineages porcellanidae and lomisidae and only more recently within the superfamily lithodoidea . some questions naturally arise . why did carcinization occur independently three times during the evolution of the anomura ? why did the presumably shell - dwelling asymmetrical hermit crab ancestors of lithodid king crabs forsake the use of shells for protection , which already provided them with survival advantage ? morrison et al . [
] suggest that the crab - like form might represent a key innovation that is associated with an evolutionary advantage , possibly due to the greater mobility and agility provided by this morphology . this seems to be evident within the true crabs , or brachyura , which dominate decapods in terms of species richness [ > 6 , 559 species ; 34 ] and have thrived in marine , freshwater , and terrestrial environments . although diversification seems to be low in the crab - like anomurans when compared to the brachyurans , fossil evidence and divergence time analyses suggest crab - like anomurans are much younger when compared to the closely related true crabs ( table
) . furthermore , the crab - like porcellanids are one of the oldest ( ~ 172 mya , mrca = 139 mya ) and most diverse families of anomurans [ ~ 247 species , 22 ] . lithodids represent an even younger lineage , originating ~ 18 mya , but comprising over 100 extant species . it is plausible that a crab - like form may hold some evolutionary advantage when considering age and diversification within anomura , although this does not seem to hold true for all groups that underwent the crab - like transition ( i . e . , monotypic family lomisidae ) . a second hypothesis explains the possible advantage of carcinization from a hermit - like ancestor . previous studies have suggested a free - living body form may have a selective advantage in obtaining food resources when unconstrained by a gastropod shell [
the extraordinary morphological and ecological diversity of anomurans has long fascinated evolutionary biologists . previous studies covering a wide range of faunas have shown how morphological or ecological factors may influence the course of subsequent evolutionary diversification [
our analysis reports the pattern of diversification in anomura to be characterized by a low net rate of diversification , with two major changes in the rates of speciation along its evolutionary history . the initial diversification of the group during the late permian was characterized by slow rates of diversification and it was not concomitant with major family radiations , which took place from the jurassic onwards .
) and currently occupy deep - sea habitats suggests that similar geological and environmental changes may also have driven major diversification within the munididae , which shifted habitats at some point because the jurassic forms are nearly all coral - reef associated . currently , the family chirostylidae accounts for 7 % of all anomuran species , but the true diversity is underestimated and about 100 new species are in hand of taxonomists [
] . clearly , a more accurate phylogenetic framework is needed to interpret in detail the exceptionally high speciation rates reported here .
the monotypic family lomisidae showed a strikingly lower rate than the overall tempo of diversification in anomura .
is anomalous in its prolonged persistence despite an inferred speciation rate of zero ( as recovered by the medusa analysis , see results ) . these taxa , old lineages with few extant species , have been reported in several invertebrates and vertebrates [
] , suggesting that extremely low rates of diversification characterize these groups . high extinction rates could also account for this pattern ; however , we report that a pure - birth yule model best explains our data . under a high - extinction scenario we would expect to see an overabundance of more recently arisen species that simply have not yet gone extinct ; such a pattern is not observed in our phylogeny .
our analysis failed to identify a correlation between the timing of branching events ( speciation ) and the evolutionary history of carcinized lineages , which suggests that the acquisition of a crab - like form did not play a major role in shaping extant anomuran biodiversity . however , a major limitation of the medusa approach is that rate shifts cannot be assigned below the level of phylogenetic resolution [
our study included extant representatives from 19 families , 77 genera , and 137 species of anomurans . the exceptionally rare family pylojaquesidae is excluded for lack of molecular grade tissue samples . a total of 345 sequences from 76 of 144 anomuran specimens were new to this study , while sequences for all five genes from 68 taxa were obtained from genbank . newly included specimens were collected on cruise and field expeditions , from collaborators , or from the university of louisiana at lafayette zoological ( ullz ) collection of molecular grade specimen and tissue samples ( table
) spanning several decapod lineages . different outgroups were included / excluded to explore sister relationships to anomura and the impact of outgroup selection on anomuran relationships . they consist of representatives from infraorders brachyura ( 5 ) , axiidea ( 4 ) , gebiidea ( 3 ) , caridea ( 4 ) , and suborder dendrobranchiata ( 2 ) .
our morphological data matrix consisted of 156 characters and 154 species ( including outgroups ) and was constructed in macclade 4 . 0 ( see additional files
missing data were scored as unknown ( ? ) and polymorphisms were scored as such rather than assuming a plesiomorphic state . just as alignment gaps in molecular data have been variously treated as a fifth position or as missing in different studies , inapplicable character states in the morphological data may be scored as missing or as an additional character state , ‘inapplicable’ [
total genomic dna was extracted from the pleon or gills using the qiagendneasy® blood and tissue kit cat . no . 69582 . two partial mitochondrial genes , 16s and 12s , were amplified by pcr using the following primers , respectively : l2 / l9 & 16s1472 or 16sf & 16s1472 [ ~ 580 bps , [
] ] . the nuclear large subunit 28s rrna was amplified in sections by 1 . 3 f / 4b , 3 . 25 / 4 . 4b , sa / 5b , and 4 . 8 / 6b [ ~ 2200 bps , [
] ] or by 1 f / 2 . 9 , 0 . 7 / bi , 2 . 0 / 9r [
] ] . the majority of target gene regions were obtained through traditional sanger sequencing and data for seven taxa were obtained through next - generation 454 sequencing ( see below ) .
pcr amplifications were performed in 25 μl volumes containing 1 μl of taq polymerase hotmaster or redtaq , pcr buffer , 2 . 5 mm of deoxyribonucleotide triphosphate mix dntps , 0 . 5 μm forward and reverse primer , and extracted dna . the thermal profile used an initial denaturation for 1 min at 94°c followed by 35–40 cycles of 30 sec at 94°c , 45 sec at 45 - 60°c depending on gene region , 1 min at 72º and a final extension of 10 min at 72°c . pcr products were purified using plate filters prepease™ pcr purification 96 - well plate kit , usb corporation and sequenced with abi bigdye® terminator mix ( applied biosystems , foster city , ca , usa ) . cycle sequencing reactions were performed in an applied biosystems 9800 fast thermal cycler ( applied biosystems , foster city , ca , usa ) , and sequencing products were run forward and reverse on an abi 3730xl dna analyzer 96 - capillary automated sequencer in the brigham young university ( byu ) sequencing center .
] . the process required a two - step pcr to prepare selected dna regions for targeted / directed sequencing . the first pcr used a locus specific primer ( e . g . , 16s , 12s , etc . ) with a 22 bp adapter . these amplicons were cleaned using plate filters prepease™ pcr purification 96 - well plate kit , usb corporation . one μl of cleaned pcr product was used as template for the second pcr . pcr ii incorporated a 10 bp barcode multiplex identifier , mid , 4 bp key , and a 21 bp 454 titanium primer . samples were again cleaned using the millipore system and subsequently combined in emulsion pcr and sequenced via 454 gs flx titanium pyrosequencing technology ( roche ) at the byu sequencing center . the bioinformatic pipeline , barcodecruncher , allowed us to exclude short reads , trim adapters , identify contamination , parse barcoded sequences , and assembly consensus sequences for phylogenetic reconstruction [ for full description of methods see [
sequences were cleaned and assembled using sequencher 4 . 9 ( genecodes , ann arbor , mi , usa ) . to check for pseudogenes , we followed suggestions by song et al . ( 2008 ) , which included extracting dna from tissue with high amounts of mitochondrial gill tissue , translating protein - coding sequences h3 to check for indels and stop codons , comparing sequences among closely - related species , and building individual gene trees to ensure similar topologies [
] . comparing gene trees and blast searches helped identify contamination . two datasets were assembled : 1 ) molecular dataset including all 5 gene regions 2 ) combined dataset including molecular + morphological data .
individual gene alignments were performed using mafft , implementing the “e - ins - i” option . for non - protein coding genes 12s , 16s , 18s , 28s , gblocks v0 . 91b were used to exclude regions of the alignment with questionable positional homology [


synanthedon pini , the pitch mass borer , is a moth of the family sesiidae .

the pitch mass borer occurs on spruce and pine in eastern north america .

it does not kill trees , but the pitch-filled larval tunnels in the wood cause defects in the lumber .

synanthedon pini

species synanthedon pini - pitch mass borer - hodges # 2585 - bugguide . net
© william taft and mich . st . univ . , moth photographers group · 2 synanthedon pini , male
all are in one genus - synanthedon sp . single eggs are laid on the bark of the pines . larvae of some species tunnel into the cambium area soon after hatching . they make a hole in which they feed . sap oozes out of these pits / holes . the pitch mass borer
duckworth , w . d . & t . d . eichlin 1973 . new species of clearwing moths ( lepidoptera : sesiidae ) from north america . proceedings of the washington entomological society 75 ( 2 ) : 150 - 159 melittia calabaza , synanthedon arkansasensis , s . canadensis , s . dominicki , s . engelhardti
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larva - body white with a light brown head . ( kellicott , 1881 ) .
the males are strongly attracted to zzoh / ezoh 50 : 50 pheromone lures and are on the wing starting in late june in michigan .
, pinaceae ) . larvae typically bore in large trees below a broken branch or scar as high as 40 ' up the trunk ( kellicott , 1881 ) .
it is found rarely in insect collections and museums . trees with active larvae have pitch mass swellings on the tree trunk with a white , powdery appearance .
beuttenmüller , w . 1901 . monograph of the sesiidae of america , north of mexico . memoirs of the american museum of natural history 1 ( 6 ) :
engelhardt , g . p . 1946 . the north american clear - wing moths of the family aegeriidae . united states national museum bulletin 190 :
kellicott , d . s . 1881 . observations of several species of ageriadae inhabiting the vicinity of buffalo , n . y . . the canadian entomologist 13 ( 1 ) : 5 - 7
dictionary of word roots and combining forms donald j . borror . 1960 . mayfield publishing company .
the north american clear - wing moths of the family aegeriidae . george p . engelhardt . 1946 . united states national museum bulletin 190 : 1 - 222 , pl . 1 - 32 .
monograph of the sesiidae of america , north of mexico . william beutenmüller . 1901 . memoirs of the american museum of natural history 1 ( 6 ) : 218 - 352 , pl . 29 - 36 .
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eastern white , scots and jack pines as well as white , norway and colorado blue spruce .
pine pitch moth adults resemble yellowjackets . the adults are clear wing moths . there are many types of pitch moths .
large volumes of pitch ooze from the holes made in the trunk by these insects .
attacks austrian , eastern white , scots and jack pines as well as white , norway and colorado blue spruce . adults are only seen during the summer and the insect can take up to three years to complete its life cycle .
check with your local land grant university ( cooperative ) extension service for recommended insecticide .
the home , yard & garden pest guide ( c1391 ) provides is written for homeowners and other residents and provides nonchemical and current chemical recommendations for controlling pests associated with trees , shrubs , turf , flowers , groundcovers , vegetables , fruit , and houses . in addition , you ' ll find detailed information about integrated pest management , pesticide safety , and pesticide application and calibration techniques . this publication may be purchased at your local university of illinois extension unit office , or by calling 800 - 345 - 6087 , or by placing an order online ( search for
c1391
) . visit site > >
the illinois commercial landscape and turfgrass pest management handbook ( iclt ) is written for professional applicators and provides nonchemical and current chemical recommendations as well as application timing information for all major pests of turf , woody ornamentals and herbaceous ornamentals . this publication may be purchased at your local university of illinois extension unit office , or by calling 800 - 345 - 6087 , or by placing an order online ( search for
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) . visit site > >
a free plant , weed , insect and disease identification service available through your local university of illinois extension office . center educators or state specialists review & respond to information and digital images submitted by local extension office personnel . some samples may require further examination or culture work ( nominal fee involved ) at the u of il plant clinic . visit site > >
services include plant and insect identification , diagnosis of disease , insect , weed and chemical injury ( chemical injury on field crops only ) , nematode assays , and help with nutrient related problems , as well as recommendations involving these diagnoses . microscopic examinations , laboratory culturing , virus assays , and nematode assays are some of the techniques used in the clinic . visit site > >
eastern white pine , scots pine , austrian pine , jack pine and red pine . as well as white spruce , norway spruce and blue colorado spruce trees .
the larvae tunnel into the inner bark on trunks and limbs of host trees . this causes a cavity .
the clearwing moth looks similar to yellow jacket wasps . it lays its eggs in midsummer in pruning cuts and on the bark of the trunk or limbs . the larvae tunnel into the inner bark to feed on resin from the damaged tissue . the mature larvae reach a length of about 25mm . their body colour is usually near white to pink depending on which tree they are feeding on . their brown head is smaller than their prothorax . large amounts of pitch and frass form at the point of entry . pupation occurs within the pitch mass from late may to june . the clearwing moths appear from the middle to the end of june , although others can emerge in july and august depending on locations . two or three years are required to complete a life cycle .
larvae and pupa are found under the pitch masses . they can be removed and killed .
do not prune trees during the egg laying period of the pitch mass moth . ( midsummer )
complete this form for a certified arborist to visit your property and provide recommendations and an estimate .
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hodges , r . w . et al . , eds . 1983 . check list of the lepidoptera of america north of mexico . e . w . classey limited and the wedge entomological research foundation , london . 284 pp .
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ridgely , r . s . , t . f . allnutt , t . brooks , d . k . mcnicol , d . w . mehlman , b . e . young , and j . r . zook . 2003 . digital distribution maps of the birds of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
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patterson , b . d . , g . ceballos , w . sechrest , m . f . tognelli , t . brooks , l . luna , p . ortega , i . salazar , and b . e . young . 2003 . digital distribution maps of the mammals of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
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adults are wasp - like clearwing moths with a mostly orange head , grey hairs on the face , a brown - black thorax , clear hind wings , opaque brown forewings with ( usually ) some orange scales and a mostly blue - black abdomen with an orange 4th segment .
larvae are white to pink in colour and can vary depending on the host tree . they have a head capsule that is smaller than the prothorax . when compared to the
eggs are laid on the bark of trunks or branches . females prefer fresh wound sites ( e . g . recently pruned branches ) if available .
larvae bore into the inner bark and sap wood . tunneling in this area causes copious amounts of sap to flow out of the wound .
female moths deposit eggs in june to july on the trunk or larger branches , preferably at a wound site . larvae tunnel into the sapwood behind the bark , causing copious amounts of sap to flow out of the infested area . it takes 2 - 3 years for the larvae to mature . larvae overwinter within the pitch mass . there is only one generation every 2 - 3 years . adults emerge in spring to early summer .
johnson , w . t . and lyon , h . h . 2003 . insects that feed on trees and shrubs . second edition . cornell university press , ithaca , new york . 560 pp .
eichlin , t . d . , duckworth , w . d . 1988 . the moths of america north of mexico , sesioidea , sesiidae , fascicle , 5 . 1 . the wedge entomological research foundation .
contributed by maury j . heiman on 18 june , 2013 - 12 : 00am
contributed by maury j . heiman on 25 may , 2013 - 7 : 53pm
contributed by maury j . heiman on 1 august , 2013 - 10 : 45pm
memoirs of the american museum of natural history 1 ( 6 ) : 218 - 352 , pl . 29 - 36 , 1901
beutenmüller , w . 1901 . monograph of the sesiidae of america , north of mexico . memoirs of the american museum of natural history 1 ( 6 ) : 218 - 352 , pl . 29 - 36
brown , l . n . & r . f . mizell , iii 1993 . the clearwing borers of florida ( lepidoptera : sesiidae ) . tropical lepidoptera 4 ( 4 ) : 1 - 21 ( pdf )
eichlin , t . d . 1992 . clearwing moths of baja california , mexico ( lepidoptera : sesiidae ) . tropical lepidoptera 3 ( 2 ) : 135 - 150 ( pdf ) andrewsi bibionipennis erici faulkneri gilberti gloriosa hennei magnifica mexicanus palmii ( neumoegen ) palmii ( beutenmüller ) polygoni resplendens robiniae snellingi
united states national museum bulletin 190 : 1 - 222 , pl . 1 - 32 , 1946
engelhardt , g . p . 1946 . the north american clear - wing moths of the family aegeriidae . united states national museum bulletin 190 : 1 - 222 , pl . 17 - 32 species index food - plant index