elaeophora elaphi is a nematode parasite found in the blood vessels of the liver in red deer ( cervus elaphus ) in certain parts of spain .

the adult male measures 77 mm long and 549 µm wide , adult females are 91-109 mm long and 793-1049 µm wide , and microfilariae ( in utero ) are 225 µm long .

though adult e. elaphi induce lesions in the blood vessels , and appear to activate the local immune response , they seldom cause overt clinical symptoms in their hosts .

elaeophora elaphi

elaeophorosis in red deer caused by elaeophora elaphi : lesions of natural disease . - pubmed - ncbi
immunohistochemical characterization of hepatic lesions associated with elaeophora elaphi parasitism in red deer ( cervus elaphus ) .
elaeophora elaphi n . sp . ( filarioidea : onchocercidae ) parasite of the red deer ( cervus elaphus ) . with a key of species of the genus elaeophora .
elaeophora elaphi n . sp . ( filarioidea : onchocercidae ) parasite of the red deer ( cervus elaphus ) with a key of species of the genus elaeophora
immunohistochemical characterization of hepatic lesions associated with elaeophora elaphi parasitism in red deer ( cervus elaphus ) . - pubmed - ncbi
elaeophora elaphi n . sp . ( filarioidea : onchocercidae ) parasite of the red deer ( cervus elaphus ) . with a key of species of the genus elaeophora . - pubmed - ncbi
elaeophora elaphi - description and life cycle . . . the life cycle of e . . . however , the seasonal cycle of adult e . . . is similar to the seasonal abundance cycle of elaeophora schneideri . . .
e _ elaphi _ v1 _ 0 _ 4 , gca _ 000499685 . 1
elaeophorosis , caused by elaeophora elaphi , was observed in red deer ( cervus elaphus ) from toledo province ( spain ) for the first time . adult specimens of elaeophora elaphi were found in the hepatic vessels of nine of 151 red deer between october 1994 and september 1995 ; intensity of infection was two to 18 nematodes per host . adult nematodes were only found…
the nematode elaeophora elaphi is a nematode parasite found in the blood vessels of the liver in red deer ( cervus elaphus ) and a sheep ( ovis aries ) in certain parts of spain .
höfle u , vicente j , nagore d , hurtado a , peña a , de la fuente j , gortázar c ( 2004 ) the risk of translocating wildlife . pathogenic infection with theileria sp . and elaeophora elaphi in an imported red deer . vet parasitol 126 : 387–395
elaeophorosis , caused by elaeophora elaphi , was observed in red deer ( cervus elaphus ) from toledo province ( spain ) for the first time . adult specimens of elaeophora elaphi were found in the hepatic vessels of nine of 151 red deer between october 1994 and september 1995 ; intensity of infection was two to 18 nematodes per host . adult nematodes were only found during the period from fall through early spring . no differences were present between sex or age groups . parasites were not found in a limited sample from fallow deer ( dama dama ) . blood samples were negative for the presence of microfilariae .
we described a new species of nematode filarioid ( onchocercidae ) parasiting the hepatic vessels of the red deer cervus elaphus . this new species is characterized by the number and disposition of the papillae on the genital area of the male , and the presence and characteristics of an
area rugosa
situated just in front of the ventral impair precloacal papilla . we discussed this new species , giving a key to the identification of all the known species of the genus elaeophora .
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annales de parasitologie humaine et comparée . ( journal , magazine , 1923 ) [ worldcat . org ]
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# t . 19 , no 4 - 5 - 6 - : encyclopédie des sciences médico - biologiques . section - - parasitologie
t . 19 , no 4 - 5 - 6 - : encyclopédie des sciences médico - biologiques . section - - parasitologie
annales de parasitologie humaine et comparée . / ; paris : masson , [ 1923 ] - 1993 .
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the draft genome assembly was produced by the parasite genomic group at the wellcome trust sanger institute , in collaboration with antonio osuna ( universidad de granada ) , using illumina paired - end sequencing followed by an in - house genome assembly pipeline comprising various steps , including contig assembly , scaffolding , gap - filling and error - correction ( helminth genomes consortium , unpublished ) .
the gene predictions were made by the parasite genomics group at the wellcome trust sanger institute and wormbase , as part of the 50 helminth genomes initiative ( helminth genomes consortium , unpublished ) . an in - house pipeline was developed that used maker to generate high - quality annotations by integrating evidence from multiple sources : ab initio gene predictions from augustus , genemark - es , and snap ; projected annotation from c . elegans ( using genblastg ) and the taxonomically nearest reference helminth genome ( using ratt ) ; and ests , mrnas and proteins from related organisms aligned to the genome using blast , with refinement of alignments using exonerate .
under grant numbers bb / k020080 / 1 and bb / k020048 / 1 .
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ann parasitol hum comp . 1986 ; 61 ( 4 ) : 457 - 63 .
carrasco l 1 , fierro y , sánchez - castillejo jm , bautista mj , gómez - villamandos jc , sierra ma .
department of anatomy and comparative pathology , faculty of veterinary medicine , córdoba university , spain .
carrasco l 1 , gómez - villamandos jc , fierro y , sánchez - castillejo jm , bautista mj , pérez j .
departamento de anatomía y anatomía patológica comparadas , facultad de veterinaria , córdoba , spain .
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hernandez rodriguez , s . ; martinez gomez , f . ; gutierrez palomino , p .
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enter query sequence ( s ) in the text area . it automatically determines the format of the input . to allow this feature , certain conventions are required with regard to the input of identifiers . more . . .
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enter one or more queries in the top text box and one or more subject sequences in the lower text box . then use the blast button at the bottom of the page to align your sequences .
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enter coordinates for a subrange of the subject sequence . the blast search will apply only to the residues in the range . sequence coordinates are from 1 to the sequence length . the range includes the residue at the to coordinate . more . . .
select the sequence database to run searches against . no blast database contains all the sequences at ncbi . blast databases are organized by informational content ( nr , refseq , etc . ) or by sequencing technique ( wgs , est , etc . ) . more . . .
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psi - blast allows the user to build a pssm ( position - specific scoring matrix ) using the results of the first blastp run . )
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pseduocount parameter . if zero is specified , then the parameter is automatically determined through a minimum length description principle ( pmid 19088134 ) . a value of 30 is suggested in order to obtain the approximate behavior before the minimum length principle was implemented .
baneth g . , barta j . r . , shkap v . , martin d . s . , macintire d . k . , vincent - johnson n . 2000 . genetic and antigenic evidence supports the separation of
carreño r . a . , kissinger j . c . , mccutchan t . f . , barta j . r . 1997 . phylogenetic analysis of haemosporinid parasites ( apicomplexa : haemosporina ) and their co - evolution with vectors and intermediate hosts .
centeno - lima s . , do rosario v . , parreira r . , maia a . j . , freudenthal a . m . , nijhof a . m . , jonjegan f . 2003 . a fatal case of human babesiosis in portugal : molecular and phylogenetic analysis .
cortes a . , felger i . , beck h . p . 2003 . molecular parasitology of malaria in papua new guinea .
criado - fornelio a . , gutierrez - garcia l . , rodriguez - caabeiro f . , reus - garcia e . , roldan - soriano m . a . , diaz - sanchez m . a . 2000 . a parasitological survey of wild red foxes (
criado - fornelio a . , martínez - marcos a . , buling - saraña a . , barba - carretero j . c . 2003 . molecular studies on
criado - fornelio a . , gonzalez - del - rio m . a . , buling - saraña a . , barba - carretero j . c . 2004 . the “expanding universe” of piroplasms .
criado - fornelio a . , ruas j . l . , casado n . , farias n . , soares p . , muller g . , brum j . g . w . , berne m . e . a . , buling a . , barba - carretero j . c . 2006 . new molecular data on mammalian
criado - fornelio a . , rey - valeiron c . , buling a . , barba - carretero j . c . , jefferies r . , irwin p . 2007 . new advances in molecular epizootiology of canine hematic protozoa from venezuela , thailand and spain .
ewing s . a . , panciera r . j . 2003 . american canine hepatozoonosis .
felsenstein j . 1989 . phylip : phylogeny inference package ( version 3 . 2 ) .
galtier n . , gouy m . 1995 . inferring phylogenies from dna sequences of unequal base compositions .
garcía - sanmartín j . , aurtenetxe o . , barral m . , marco i . , lavin s . , garcía - pérez a . l . , hurtado a . 2007 . molecular detection and characterization of piroplasms infecting cervids and chamois in northern spain .
hall t . a . 1999 . bioedit : a user - friendly biological sequence alignment editor and analysis program for windows 95 / 98 / nt .
hofle u . , vicente j . , nagore d . , hurtado a . , peña a . , de la fuente j . , gortazar c . 2004 . the risks of translocating wildlife . pathogenic infection with
kimura m . 1980 . a simple method for estimating evolutional rates of base substitutions through comparative studies of nucleotide sequence .
kumar s . , tamura k . , nei m . 2004 . mega3 : integrated software for molecular evolutionary genetics analysis and sequence alignment .
spp . : pathological and partial 18s rrna sequence analysis from three brazilian dogs .
reichard m . v . , van den bussche r . a . , mewinkoth j . h . , hoover j . p . , kocan a . a . 2005 . a new species of
from pallas’ cats caught in mongolia and comments on the systematics and taxonomy of piroplasmids .
rubini a . s . , dos santos paduan k . , perez r . r . , ribolla p . e . , o’dwyer l . h . 2006 . molecular characterization of feline
thompson j . d . , higgings d . g . , gibson t . j . 1994 . clustal w : improving the sensitivity of progressive multiple sequence alignment through sequence weighting , positions - specific gap penalties and weight matrix choice .
van de peer y . , de wachter r . 1993 . treecon : a software package for the construction and drawing of evolutionary trees .
criado - fornelio , a . , buling , a . , casado , n . et al . acta parasit . ( 2009 ) 54 : 187 . urltoken
meningeal worm in deer from western nebraska , david w . oates , mauritz c . sterner , ed boyd
one hundred seventy - eight whitetailed deer ( odocoileus virginianus ) and 275 mule deer ( odocoileus hemionus ) collected from locker plants in the western 2 / 3 of nebraska ( usa ) in november 1997 were examined for the meningeal worm ( parelaphostrongylus tenuis ) . parelaphostrongylus tenuis was identified in 17 ( 10 % ) of 168 white - tailed deer and in one ( < 1 % ) of 273 mule deer . this is the first naturally occurring infection of p . tenuis recorded in a mule deer .
elaeophorosis in red deer from spain , mónica santín - durán , j . m . alunda , j . m . san miguel , eric p . hoberg , c . de la fuente
meningeal worm in free - ranging deer in nebraska , david w , oates , mauritz c . sterner , david j . steffen
the meningeal worm ( parelaphostrongylus tenuis ) was found in 22 ( 7 % ) of 300 white - tailed deer ( odocoileus virginianus ) ( 257 adults , 43 fawns ) examined from nebraska ( usa ) during november 1996 . none of 53 mule deer ( odocoileus hemionus ) ( 47 adults and 6 fawns ) examined were infected . twenty - two white - tailed deer from 18 counties in eastern nebraska were infected with parelaphostrongylus tenuis . this is the first record of p . tenuis in white - tailed deer from this state .
absence of tuberculosis in free - ranging deer in nebraska , david j . steffen , david w . oates , mauritz c . sterner , vickie l . cooper
lymph nodes from 271 white - tailed deer ( odocoileus virginianus ) and mule deer ( odocoileus hemionus ) in nebraska ( usa ) were examined microscopically for tuberculoid lesions . lymph nodes lesions in at least one node were found in 12 deer . lesions were examined with zeihl - neelson acid fast stains and by polymerase chain reactions using m . bovis specific probes . no evidence of tuberculosis was found . the small granulomatous lesions were likely caused by other bacteria .
when an appropriate fish host is selected , analysis of its parasites offers a useful , reliable , economical , telescoped indication or monitor of environmental health . the value of that information increases when corroborated by another non - parasitological technique . the analysis of parasites is not necessarily simple because not all hosts serve as good models and because the number of species , presence of specific species , intensity of infections , life histories of species , location of species in hosts , and host response for each parasitic species have to be addressed individually to assure usefulness of the tool . also , different anthropogenic contaminants act in a distinct manner . . .
helminth parasitism in martens ( martes americana ) and ermines ( mustela erminea ) from washington , with comments on the distribution of trichinella spiralis , eric p . hoberg , keith b . aubry , j . david brittell
helminth parasitism in martens ( martes americana ) and ermines ( mustela erminea ) from washington , with comments on the distribution of trichinella spiralis , eric p . hoberg , keith b . aubry , j . david brittell
helminths are reported for the first time from ermines ( mustela erminea ) and martens ( martes americana ) in washington , usa . among 22 adult ermines , 41 % were infected by one or more of five species ( taenia mustelae , alaria mustelae , molineus patens , m . mustelae , and trichinella spiralis ) . among 78 adult martens from three geographic localities , the prevalence was 83 % . nine species were identified ( mesocestoides sp . , t . mustelae and t . martis americana , e uryhelmis squamula , m . patens , baylisascaris devosi , physaloptera sp . , soboliphyme baturini , and t . spiralis ) . trichinella spiralis occurred with a maximum prevalence of 50 % in martens , but only occurred in 9 % of ermines . compression . . .
helminth parasites of northern spotted owls ( strix occidentalis caurina ) from oregon , eric p . hoberg , g . s . miller , e . wallner - pendleton , o . r . hedstrom
helminth parasites of northern spotted owls ( strix occidentalis caurina ) from oregon , eric p . hoberg , g . s . miller , e . wallner - pendleton , o . r . hedstrom
helminth parasites are reported for the first time from northern spotted owls . seventyone percent of a sample of strix occidentalis caurina from western oregon was infected . nematodes ( porrocaecum depressum , capillaria falconis , microtetrameres sp . and synhimantus hamatus ) were the most prevalent parasites although cestodes ( paruterina rauschi ) and acanthocephalans ( centrorhynchus conspectus ) were also represented . there was an association between components of this helminth fauna and the diet of spotted owls which is dominated by small rodents . the occurrence of p . rauschi rather than p . candelabraria in this geographic region and host - species may provide additional support for recognition of a parapatric distribution in . . .
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moose ( alces alces ) are a culturally and economically valued species in minnesota , where the northeast population has decreased by 60 % since 2006 . the cause of the decline is currently unclear ; however , parasites , predation , and climate change have all been implicated . nematode parasites are important pathogens in north american moose , potentially causing severe disease and mortality . recent spread of rumenfilaria andersoni , a filarioid nematode of moose , has been documented in finnish cervids ; however , little is known about the epidemiology of this parasite in north america .
to investigate the prevalence and distribution of r . andersoni , 584 blood samples were collected from live - captured and dead animals and screened microscopically for the presence of microfilariae using a modified knott’s test . microfilariae were identified based on morphological characteristics . a subset of knott’s - positive animals was subjected to polymerase chain reaction ( pcr ) with filarioid - specific primers targeting the first internal transcribed spacer region ( its - 1 ) of the rrna gene cluster .
rumenfilaria microfilariae were present in 20 . 5 % of minnesota moose ( n = 352 ) , with slight fluctuations observed over four years . minnesota white - tailed deer ( odocoileus virginianus ) ( n = 2 ) and moose ( n = 44 ) from alaska , montana , washington , maine , and new hampshire also harbored r . andersoni , suggesting this parasite occurs widely throughout north american moose herds , and white - tailed deer can serve as a patent host . sequence analysis of cervid blood ( moose , n = 15 ; white - tailed deer , n = 1 ) confirmed the identity of r . andersoni and revealed the existence of two distinct clades . genetic comparisons of r . andersoni isolates from north america and semi - domesticated finnish reindeer found the two groups to be closely related , supporting previous hypotheses that r . andersoni was recently introduced into finland by the importation of deer from the united states .
to the best of our knowledge these observations represent the first report of r . andersoni within the contiguous united states and reveal this nematode as a common parasite of north american moose and white - tailed deer . although the implications of r . andersoni infection on moose health is unclear , increased awareness of this parasite will help prevent unintentional introduction of r . andersoni into naïve populations via the translocation of wild and captive cervids .
] . to prevent further loss of this species and natural resource , a better understanding of the minnesota moose herd’s overall health , as well as an understanding of the potential drivers of mortality , is urgently needed .
) . the species belongs to the family onchocercidae , which is a group of filarioid nematodes transmitted by hematophagous arthropods . although the exact details of the
life - cycle have yet to be elucidated , all adult filarioids of the family onchocercidae produce larval stages called microfilariae that reside in the circulatory system of the definitive cervid host . when an arthropod intermediate host ( vector ) obtains a blood meal and ingests the microfilariae , the microfilariae unsheathe , penetrate the vector’s gut wall , and develop to an infectious larval stage within the vector’s hemocoel . extrinsic development is completed when infectious larvae are inoculated into a definitive host by the vector during a subsequent blood meal .
the main objective of this study was to investigate the eco - epidemiology of r . andersoni in minnesota moose compared to other north american herds . to accomplish this , we explored the basic epidemiology of this parasite , including the identification of other patent host species , geographical distribution , prevalence of infection , and host demographics by collecting parasitological samples across spatial and temporal scales . we also performed the first genetic characterization and comparison of r . andersoni isolates to provide a basis for future population genetics studies . these data will contribute to a greater understanding of r . andersoni biology and provide baseline epidemiological data for future reference and research .
within the minnesota moose population , 352 blood samples were collected between 2012 and 2015 . blood was collected from either hunter - killed animals , opportunistic mortalities , or live - captured animals and placed into 5 - ml edta blood tubes . a limited number of blood samples were also obtained from hunter - harvested wild elk
from washington . whenever possible , the age , sex , and geographical location of the animal was noted . animals were classified as calves ( < 1 year of age ) , yearlings ( 1 to < 2 years of age ) , or adults ( ≥ 2 years old ) . blood samples ( 1 ml / animal ) were refrigerated and shipped to the molecular parasitology laboratory at the university of tennessee college of veterinary medicine to identify the presence of
prevalence of r . andersoni microfilariae ( rmf ) in blood samples drawn from free - ranging cervids . prevalence is defined as the percentage of samples that tested rmf - positive using a modified knott’s test . error bars represent 95 % confidence intervals . a prevalence of rmf in three species of minnesota cervids ( moose , n = 352 ; elk , n = 14 ; white - tailed ( wt ) deer , n = 36 ) . blood samples were collected over a four - year period for moose and a two - year period for elk and deer . fisher’s exact test ; p = 0 . 013 . b comparison of rmf prevalence in minnesota moose over time ( 2012 , n = 67 ; 2013 , n = 160 ; 2014 , n = 69 ; 2015 , n = 56 ) . fisher’s exact test ; p = 0 . 607 . c comparison of rmf prevalence in moose from several u . s . states ( mn , n = 352 ; nh , n = 16 ; me , n = 14 ; mt , n = 73 ; wa , n = 16 ; ak , n = 27 ) . fisher’s exact test ; p = 0 . 013 . d image of rmf from minnesota moose blood . sample was stained with methylene blue and viewed with a bright light microscope at 200× magnification . scale - bar : 20 μm
prevalence of r . andersoni within cervid populations was estimated for each cervid species based on the modified knott’s test results . blood samples were categorized as rmf positive based on the presence of microfilariae with morphological features consistent with r . andersoni identified via the modified knott’s test described above . animals with no microfilariae present or microfilariae morphologically distinct from r . andersoni were categorized as rmf negative . to determine if r . andersoni prevalence differed between geographical locations , a pearson’s χ 2 test or fisher’s exact test was used with bonferroni correction ( p ≤ 0 . 012 ) . the minnesota moose population was further analyzed by comparing prevalence between age class , sex , and sample year using the pearson’s χ 2 test or fisher’s exact test with bonferroni correction ( p ≤ 0 . 012 ) . statistical analyses were performed with spss software version 23 . 0 ( ibm corporation , armonk , ny , usa ) .
were selected for molecular analysis . dna was extracted using the zr fecal dna kit ( zymogen , irvine , ca , usa ) according to the manufacturer’s instructions . nuclease - free water served as the dna extraction control . rmf dna was amplified using a previously described semi - nested pcr protocol targeting the first internal transcribed spacer region ( its - 1 ) of the rrna gene cluster [
nematode ( ra - f3 ) isolated from a finnish reindeer and nuclease - free water served as the positive and negative pcr controls , respectively . the pcr products were separated by agarose gel electrophoresis and viewed under uv light . pcr amplicons around 600 bp were excised using a clean razor blade and the pcr product purified using the qiaquick gel extraction kit ( qiagen , valencia , ca , usa ) . the its - 1 pcr product was then cloned into a pgem - t easy vector ( promega , madison , wi , usa ) and transformed into competent dh5α
cells ( invitrogen , grand island , ny , usa ) via a 40 - second heat shock at 42 °c . transformed cells were cultivated in s . o . c . medium ( life technologies , grand island , ny , usa ) for 1 . 5 h at 30 °c with shaking . the transformed cells ( 100 μl ) were then plated on luria broth agar plates containing 1 μg / ml carbenicillin and 100 μl chromomax iptg / x - gal solution ( thermo fisher scientific , waltham , ma , usa ) as a top dressing . cultures were incubated 24–48 h at 30 °c ; single , white colonies were selected with a sterile toothpick and grown overnight in 5 ml luria broth with 1 μg / ml carbenicillin . cultures were centrifuged and the supernatant removed . plasmids were purified from the remaining cell pellet using the qiagen spin miniprep plasmid kit ( qiagen ) following the manufacturer’s instructions . to confirm the presence of the filarioid its - 1 pcr product insert , the plasmids were digested with ecor1 restriction enzyme ( thermo fisher scientific ) and examined via gel electrophoresis for multiple bands . plasmids containing an insert of approximately 600 bp were sequenced at the university of tennessee’s genomics core ( knoxville , tn , usa ) .
all 18s and its - 1 consensus sequence chromatograms were trimmed and edited by hand using sequencher 5 . 3 ( gene codes corporation , ann arbor , mi , usa ) . edited sequences were compared against the few known sequences for filarioid nematodes from cervid hosts in the ncbi genbank database . genetic data was also compared with sequences obtained from adult reference nematodes ( table
) , which had been identified morphologically and subjected to dna extraction and pcr amplification in our laboratory , as described above . alignment and construction of neighbor - joining trees of its - 1 and 18s filarioid worm sequences was performed using mega 6 . 0 [
adult nematodes were identified based on morphological characteristics . geographial origin and host species refer to the place and host from which the nematode was isolated . dna target refers to the targeted gene sequence ( 18s rrna or its - 1 ) that was amplified
prevalence of r . andersoni in free - ranging cervids of minnesota and other u . s . states
during 2012–2015 , rmf occurred in 20 . 5 % ( 95 % confidence interval [ ci ] : 16 . 3–24 . 7 % ;
= 0 . 104 ) . rmf were also detected in minnesota white - tailed deer , with an overall prevalence of 5 . 6 % ( 95 % ci : 0–13 . 1 % ;
) . over the 4 - year sampling period , rmf prevalence in the minnesota moose varied slightly , ranging from 22 . 5 % ( 95 % ci : 16 . 0–29 . 0 % ;
rmf were also observed in all other surveyed moose populations , including those in maine at 21 . 4 % ( 95 % ci : 0–42 . 9 % ;
= 14 ) , new hampshire at 25 . 0 % ( 95 % ci : 3 . 8–46 . 2 % ;
= 16 ) , and alaska at 40 . 7 % ( 95 % ci : 22 . 2–59 . 2 % ;
= 0 . 013 ) . moreover , we failed to observe rmf in any of the alaskan caribou blood samples or in mule or white - tailed deer from washington .
) . moose isolates varied in geographical origin , with five from montana , eight from minnesota , and two from maine . attempts to amplify additional isolates were not successful . sequences obtained from a morphologically confirmed
adult nematode ( ra - f3 ) isolated from a finnish reindeer served as the standard .
rumenfilaria andersoni its - 1 target dna sequences amplified from cervid blood . unless otherwise indicated , isolates were obtained from moose ( alces alces )
overall , the rmf its - 1 sequences were at - rich , with multiple sections of repeats with variations in length between isolates . phylogenetic analysis revealed that all of the its - 1 sequences obtained from the rmf - positive blood clustered with the ra - f3 standard , branching into two distinct
) . the minnesota isolates had representatives clustering into both clades , which we simply denoted as clades a and b . all montana isolates and six minnesota isolates clustered into clade a , which also contained the ra - f3 standard . clade b contained both maine isolates and two minnesota isolates . isolates of clades a and b had a mean difference of 0 . 038 ( se = 0 . 007 ) base substitutions per base pair . within clades , isolates had a mean difference of 0 . 015 ( clade a , se = 0 . 003 ; clade b , se = 0 . 004 ) base substitutions per base pair for both clades a and b .
phylogenetic analysis of its - 1 sequences obtained from rmf - positive blood samples from cervid hosts . sequences of 609 base pairs were aligned using clustalw , and the evolutionary history was inferred using the neighbor - joining method . evolutionary distances were computed using the kimura 2 - parameter method . the tree is drawn to scale . bootstrap values ( ×1000 ) greater than 50 % are shown above the branches . rmf isolates are marked with solid boxes ; its - 1 clades a and b are labelled . ra - f3 (
serve as reference standards . genbank accession numbers for all isolates are listed in tables
in addition to rmf , another morphologically distinct group of microfilariae was observed in 1 . 4 % ( 5 / 352 ) of minnesota moose . these microfilariae were characterized by blunt , rounded heads and long , thinly tapered tails , measuring between 285 and 315 μm long and 5–7 . 5 μm wide ( fig .
) . dual infection with rmf and non - rmf microfilariae was observed in a single minnesota moose . to identify this unknown filarioid , we attempted to sequence a portion of the 18s rrna gene using nematode - wide primers from the five positive minnesota moose blood samples [
] . only one blood sample was successfully pcr amplified . a comparison of 796 base pairs from the unknown filarioid and 18s sequences from our reference nematodes ( table
image of unidentified microfilaria observed in blood from minnesota moose . sample was stained with methylene blue and image taken under a bright light microscope at 200× magnification .
) and other known filarioid parasites of ungulates , with history inferred using the neighbor - joining method and evolutionary distances computed using the kimura 2 - parameter method . tree is drawn to scale . bootstrap values ( ×1000 ) are shown above branches . genbank accession numbers for all isolates are listed in tables
suggests the geographical range of this filarioid nematode is much more extensive than was previously appreciated . rmf were detected in all of the wild moose herds we sampled , including herds geographically isolated from one another ( fig .
and / or its preferred vector may be more highly adapted to subarctic climates , proliferating more easily in a colder environment . the recent rapid expansion of
] supports this hypothesis , but additional evidence is needed to substantiate this claim . further research on moose densities , vector distribution and competency , and distribution of other possible definitive hosts of
will be useful in understanding if the parasite is more adapted to subarctic climates or if it is strictly a host and / or vector density - dependent mechanism .
occurs in white - tailed deer , and deer can serve as a definitive host for the filarioid ( fig .
] . the authors observed 22 % of the deer surveyed in finland were rmf - positive , higher than the 5 . 6 % prevalence value observed in deer from minnesota . as mentioned above , it is possible the parasite is better adapted to subarctic environments , thus explaining the lower parasite prevalence at the more southern latitude ; however , a larger sample size along a latitudinal gradient would be needed to properly address that particular hypothesis .
in addition to white - tailed deer , our survey included specimens from caribou , elk , and mule deer from various geographical locales . we were unable to find evidence of
in our alaskan caribou specimens may be due to an insufficient sample size rather than a lack of host - parasite co - adaptation . additionally , the number of elk and mule deer surveyed may have been insufficient , resulting in no rmf detected in the modified knott’s test ; however , it is also possible these species are not suitable hosts for the nematode . further sampling will better characterize
in their mammalian hosts . timing of sample collection could significantly influence estimated prevalence values , potentially causing early - stage infections to be missed or causing false negatives with the synchronous influx of microfilariae into general circulation that coincides with circadian rhythms [
] . furthermore , the modified knott’s test is unable to detect infections with nematodes of a single sex due to the female filarioid being unable to mate and produce progeny . at this time , the only alternative method to estimate prevalence would be to grossly examine host carcasses for the presence of adult
, presumably in the lymphatic vessels of the rumen ; however , in addition to being a laborious and tedious process , fresh wildlife carcasses can be difficult to obtain .
nematodes , we compared its - 1 sequences obtained from rmf - positive blood samples ( fig .
in north america , with all montana isolates associating with clade a and all maine isolates falling into clade b . interestingly , minnesota isolates have representatives in both clades . possible reasons for the mixed clades in minnesota include potential overlapping rmf populations or previous cervid translocation events . it is also possible multiple paratypes of
populations exist ; however , a dual dna - based and adult morphological study will be needed to identify if there is a distinct relationship between caudal papillae phenotypes and phylogenetic assortment . furthermore , studies comparing
reference nematode’s ( ra - f3 ) its - 1 sequence clustered into clade a with isolates from minnesota and montana ( fig .
resulted from the introduction of non - native white - tailed deer from north america , specifically the u . s . state of minnesota . five deer , one male and four females , were imported into southern finland in 1935 as a gift from finnish immigrants from northern minnesota [
nematodes’ ancestors originated from minnesota , we would expect the finnish its - 1 sequences to be similar to those of north american specimens , especially those of minnesota ; conversely , we would expect significant genetic variation if the finnish population had an extended history of geographical distribution and isolation in fennoscandia . our data demonstrate a lack of divergence between these isolates , indicating that a more recent , anthropogenically - driven introduction of the parasite occurred , supporting the minnesota theory , rather than an introduction coinciding with the geographical colonization by moose from central europe and russia after the last glaciation , approximately 10 , 000 years ago [
in addition to the presence of rmf , our data revealed another filarioid circulating within the minnesota moose herd ( fig .
is likely widespread amongst north american moose herds . this parasite can cause a wide range of disease in cervids . mild fibrin formation on serosal surfaces has been reported in
infections within moose populations have not been studied . thus , it is unknown the impact
and declining populations of moose . no significant difference in rmf prevalence was observed between the minnesota moose herd , which is exhibiting a severe population decline [
does not appear to have an obvious association with declining moose populations . at this time , it is still unknown what impact , if any ,
infection may have on the health of the moose host . laaksonen et al . , in 2010 , observed macroscopic inflammatory changes within the ruminal lymphatic vessels of infected finnish reindeer [
] ; however , there have yet to be any reports of pathological changes associated with moose . given the high prevalence of rmf in moose and the high prevalence and intensity of rmf in finnish reindeer , future studies on potential subclinical and clinical disease is warranted . it is reasonable to hypothesize that infection with
has a metabolic cost , and it is possible that heavy worm burdens or systemic microfilaremia would result in adverse health effects , potentially rendering the host more susceptible to other infectious agents or poor body condition , but future studies will be required to assess the validity of these hypotheses .
is a nematode widespread throughout moose herds of north america . in addition to moose , white - tailed deer appears to be a natural , definitive host of this parasite . recognizing the geographical distribution and host range of this filarioid is especially important for preventing the introduction of
into naïve populations by translocation of animals by state conservation agencies or commercial hunting businesses . our genetic comparison of
infection can lead to clinical or subclinical disease . continued efforts to document this parasite in cervid hosts will help to provide clarity on this topic .
reference nematodes were generously donated by kimberlee beckmen at the alaskan department of wildlife , fish and game and john henningsen at the wyoming state veterinary laboratory . the authors also thank nick decesare of the montana fish and wildlife department , ella rowan and kristin mansfield of the washington state department of fish and wildlife , lee kantar of maine department of inland fisheries and wildlife , anne lichtenwalner of the university of maine , and henry jones of the university of new hampshire for providing cervid blood samples for analysis . we thank ann reed at the university of tennessee institute of agriculture for statistical consultation . we also thank roger moon of the department of entomology at the university of minnesota , for his helpful comments while preparing this manuscript , and misty bailey ( funded by the university of tennessee college of veterinary medicine ) for technical editing of the manuscript . this work was partially supported by the minnesota department of natural resources section of wildlife and the university of tennessee department of biomedical and diagnostic sciences . finnish samples were collected as part of the project “reindeer health in the changing environment , ” funded by the finnish ministry of agriculture and forestry ( makera ) and the eurgnegvec cost action td1303 . fellowship funding for cg was provided by the university of tennessee department of microbiology .
this work was partially supported by the minnesota department of natural resources section of wildlife and the university of tennessee department of biomedical and diagnostic sciences . finnish samples were collected as part of the project “reindeer health in the changing environment , ” funded by the finnish ministry of agriculture and forestry ( makera ) and the eurgnegvec cost action td1303 . fellowship funding for graduate student ( cg ) was provided by the university of tennessee department of microbiology .
the datasets supporting the conclusions of this article have been included within the article . genetic sequence data has been submitted to the genbank ( national center for biotechnology information ) database under accession numbers kt020850 , kt031392–kt031393 ; kt873719–kt873733 ; kt878970–kt878979l ; and ku757075–ku757077 .
cg and rg conceived and designed the study and drafted the manuscript . cg and je performed the modified knott’s tests and molecular analysis of reference nematodes . cg completed all genetic analysis of rmf isolates . mc and eh orchestrated and oversaw collection of minnesota cervid samples and drafting of the manuscript . ao and sl collected finnish samples and were active in writing the manuscript . all authors were involved in analyses of data , and reviewed and approved the final manuscript .
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) , which permits unrestricted use , distribution , and reproduction in any medium , provided you give appropriate credit to the original author ( s ) and the source , provide a link to the creative commons license , and indicate if changes were made . the creative commons public domain dedication waiver (
) applies to the data made available in this article , unless otherwise stated .
decesare nj , smucker td , garrott ra , gude ja . moose management in montana . alces . 2014 ; 50 : 35–51 .
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carstensen m , hildebrand ec , plattner d , dexter mh , jennelle c , wright rg . determining cause - specific mortality of adult moose in northeast minnesota . in : cornicelli m , carstensen m , grund m , larsen m , lawrence j , editors . summaries of wildlife research findings . st . paul : minnesota department of natural resources ; 2015 . p . 161–71 .
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fragum unedo is a species of cockle , a marine bivalve mollusc in the family cardiidae , commonly known as the pacific strawberry cockle .

it is found in tropical seas in the indo-pacific region and the empty shells are prized for use in decorative crafts .

fragum unedo

pacific strawberry cockle ( fragum unedo ) from sealife base : technical fact sheet .
ultrastructural differences between the anterior and posterior adductors of the strawberry cockle , fragum unedo .
ultrastructural differences between the anterior and posterior adductors of the strawberry cockle , fragum unedo . - pubmed - ncbi
fragum fragum is a photosymbiotic species intermediate between corculum and fragum unedo . semi - transparent posterior areas ( pigment and windows ) exist to allow light penetration . as well , siphonal tentacles are exposed to light on posterior shell surface and contain photosymbionts .
family : bivalvia born : 1758 , linne genus and description : fragum unedo , 26 . 5 & 31 mm , f + + / f + + + , set of 2 specimens origin : collected by a local fishermen by nets off panglao island , bohol philippines , march 2013 .
pacific strawberry cockle ( fragum unedo ) in the bivalves section by j . m . poutiers in the fao species identification guide for fishery purposes : the living marine resources of the western central pacific volume 1 : seaweeds , corals , bivalves and gastropods on the food and agriculture organization of the united nations ( fao ) website .
this is the type species of the genus fragum . kirkendale ( 2009 ) has shown ( and others have confirmed e . g . herrera et al . 2015 ) that fragum is paraphyletic and includes corculum and lunulicardia , the two other wholly photosymbiotic lineages of cardiids . the phylogeny of members of the fraginae subfamily is in a state of flux and the taxonomy of the group is in need of attention .
fragum unedo is commonly found in the indo - pacific region . prior literature shows that specimens have been found in sri lanka , southern japan , the pacific islands as well as the northern and eastern coasts of australia . ( 12 . 87n , 93e , 41 . 23n , 142 . 55e , 35 . 9s , 159 . 08e ) ( e . o . l . 2012 ) .
strawberry cockle fragum unedo family cardiidae updated oct 2016 where seen ? this pretty clam is not often seen , usually on sandy shores near reefs . elsewhere , they are shallow burrowers in sandy bottoms , often occuring in dense populations . features : 4 - 6cm . the sturdy two - part shell is heavy , squarish with strong ribs marked with little red beads . human uses : elsewhere , it is used in decorative shellcraft and may be eaten by coastal dwellers .
qld , nt and wa in australia , but widely distributed in the indo - west pacific . wilson and stevenson 1977 note that placement of red scales appears different than in f . unedo from the pacific ocean and philippines .
kirkendale ( 2009 ) has shown ( and others have confirmed e . g . herrera et al . 2015 ) that fragum is paraphyletic and includes corculum and lunulicardia , the two other wholly photosymbiotic lineages of cardiids . the phylogeny of members of the fraginae subfamily is in a state of flux and the taxonomy of the group is in need of attention
( of cardium unedo linnaeus , 1758 ) linnaeus , c . ( 1758 ) . systema naturae per regna tria naturae , secundum classes , ordines , genera , species , cum characteribus , differentiis , synonymis , locis . editio decima , reformata . laurentius salvius : holmiae . ii , 824 pp . , available online at urltoken [ details ]
adductors of fragum unedo were observed ultrastructurally and their muscle cells were classified according to the statistically analyzed diameter of their thick myofilaments . two types of smooth muscle cells were observed in the opaque portion of the anterior adductor : a - type cells containing thick myofilaments of about 46 nm in diameter and b - type cells having 62 nm thick myofilaments . the posterior adductor was also composed of two kinds of cells : the b - type cell , which had thick myofilaments of about 67 nm in diameter , and the c - type , containing thick myofilaments of 90 nm . two types of oblique - striated cells were commonly recognized in the translucent portions of anterior and posterior adductors . our observations thus indicate that the posterior adductor generally consists of cells which have thicker myofilaments than the ones of the anterior adductor .
as f . unedo burrows just below the surface ( habitat ) they are still susceptible to predation . in northern australian cockle - bed , starfish and rays are important ecological predators of cardiids ( rudman 1998 ) . the locomotory pattern of jumping ( locomotion ) is used as a means to escape from predatory species , displacing the cardiid a few centimetres from their prior location .
members of the cardiidae family , like many other marine bivalves are gonochoristic , meaning individuals are either male or female . by means of broadcast spawning they are able to reproduce sexually ( hickman et al . 2011 ) , utilizing processes of external fertilisation ( ruppert , fox and barnes 2004 ; wilbur and yonge 1964 ) . no research has been conducted into the seasonal spawning patterns of f . unedo individuals . however , research conducted by kandeel et al . ( 2013 ) on cerastoderma glaucum , a cardiid cockle , has found that spawning events maybe be dependent on temperature and lunar cycles . c . glaucum are known to have an annual pattern of four spawning events ( kandeel et al . 2013 ) , it is possible that f . unedo may follow a similar trend .
like many other cardiids , f . unedo individuals spend most of their adult lives as a sedentary ( soo & todd 2014 ) , suspension feeder ( respiration and feeding ) . however , f . unedo individuals are not stuck in a single place for their entire lives , as they have the capability to locomote and burrow into sediments , using a muscular foot ( locomotion ) . the suspension feeding lifestyle imposes constraints upon the bivalves’ lifestyle , generally restricting it to aquatic environments ( habitat ) . however , during seasonal changes in tidal levels ( underwood 1981 ; tsimplis & woodworth 1994 ) , expose the intertidal reef zones . like other intertidal bivalves , f . undeo have developed the ability to survive without access to water for periods of time ( up to 8h at times ) . they accomplish this via aerial respiration , which is able to take place across the exposed posterior mantle region .
it is assumed that , f . unedo have a typical bivalve open circulatory system . with a three chambered heart , consisting of a single ventricle fed by a pair of lateral atria ( rudman 1998 ) , situated in a pericardial chamber . the heart is used to pump recently oxygenated blood , oxygenated haemoglobin is then transported to the rest of the body through blood ( ruppert , fox and barnes 2004 ; hickman et al . 2011 ) . while veins return de - oxygenated blood back to the gills and eventually the heart .
it is assumed that , the excretory system of f . unedo is similar to many other bivalves . made up of a complex , were the heart and kidney have become a single organ . the excretory system is separated from the circulatory system by the pericardium ( wilbur and yonge 1964 ) . the pericardium contains pericardial glands , which aid in the nutrient filtration process . the system contains two nephridium , that act as pseudo - kidneys and concentrate waste products ( rudman 1998 ) . each nephridium empties into the exhalant siphon chamber via their respective nephridiopore ( ruppert , fox and barnes 2004 ) .
tan siong kiat and henrietta p . m . woo , 2010 preliminary checklist of the molluscs of singapore ( pdf ) , lee kong chian natural history museum , national university of singapore .
tan , k . s . & l . m . chou , 2000 . a guide to the common seashells of singapore . singapore science centre . 160 pp .
abbott , r . tucker , 1991 . seashells of south east asia . graham brash , singapore . 145 pp .
( of cardium cruentum perry , 1811 ) perry , g . ( 1811 ) . conchology , or the natural history of shells : containing a new arrangement of the genera and species , illustrated by coloured engravings executed from the natural specimens , and including the latest discoveries . 1 - 4 , plates 1 - 61 . london . , available online at urltoken [ details ]
( of hemicardium tegulatum dautzenberg , 1900 ) dautzenberg , ph . ( 1900 ) . description d ' une espèce nouvelle appartenant au genre hemicardium . j . conchyliol . xlviii : 5 - 8 , plate i ( look up in imis ) [ details ]
poorten , j . j . ter , 2005 . outline of a systematic index - recent cardiidae ( lamarck , 1809 ) . visaya net . ( updated 2009 for worms ) , available online at urltoken [ details ]
poorten , j . j . ter , 2009 . the cardiidae of the panglao marine biodiversity project 2004 and the panglao 2005 deep - sea cruise with descriptions of four new species ( bivalvia ) . vita malacologica 8 : 9 - 96 [ 21 november 2009 ] . available online at urltoken available online at urltoken [ details ]
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 cardium cruentum perry , 1811 ) poorten , j . j . ter , 2005 . outline of a systematic index - recent cardiidae ( lamarck , 1809 ) . visaya net . ( updated 2009 for worms ) , available online at urltoken [ details ]
( of hemicardium tegulatum dautzenberg , 1900 ) poorten , j . j . ter , 2005 . outline of a systematic index - recent cardiidae ( lamarck , 1809 ) . visaya net . ( updated 2009 for worms ) , available online at urltoken [ details ]
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) . the two domed valves are equivalve , valves that mirror each other in both size and shape . however , each valve is asymmetrical . the valves are usually white or cream in colour , with strawberry - red scales ( fig . 2 ) . when in a still environment two siphons are able to be observed , an inhalant siphon (
individuals usually grow to lengths of 40mm , but when conditions are good they are able to grow to maximum lengths of 65mm .
shell with a scanning electronic microscope . they found that the shell was a bioceramic composite . made up of long and thin aragonite , crystal form of calcium carbonate caco3 , sheets and layers of collagen protein . the aragonite sheets were found to have a herringbone structure , which gives the shells added strength ( chen , peng and wu 2007 ) .
lateral view , the shell is of rhomboidal shaped ( fig . 3 ) .
dorsal view , the prosogyrate umbo aids to form the major characteristic heart shape ( fig . 4 ) .
the shells have a short heterodont , cyclodont hinge , and an external parivincular ligament .
the hinge of each valve usually has two cardinal teeth ( fig . 7 ) :
no research has been conducted into embryonic or larval development . the most educated assumption , is that they have similar embryonic and larval development to most other heterodont bivalves .
after the egg has been externally fertilized , the embryo undergoes the process of spiral cleavage ( hickman et al . 2011 ; wilbur and yonge 1964 ) . upon completion of embryonic cleavage , the embryo develops into a free swimming trochophore larvae ( ruppert , fox and barnes 2004 ) . this simple planktonic larval form , is dispersal utilizing ocean currents .
the trochophore larvae develops into the , longer lived , veliger larvae , unique to the molluscan phyla . during the veliger stage the mantle , shell and foot of the mollusc begin to develop ( hickman et al . 2011 ) . after successful feeding and dispersal , of the veliger larvae in the pelagic zone , the veliger larval form is lost as the bivalve develops into their adult forms (
savazzi ( 1985 ) states that members of the cardiidae family , display a variety of locomotory patterns that utilize the molluscs well developed l - shaped foot ( fig . 11 ) . these include , burrowing , jumping , ploughing and emerging ( fig . 12 ) .
to loosen the sediment , cardiids eject water into the substrate at the beginning of each burrowing sequence ( wilbur and yonge 1964 ) allowing for easier burrowing . the burrowing process consists of a small number of burrowing sequences ( savazzi 1985 ) in which the shell rocks forwards and backwards ( fig . 13 ) , while being pulled downward by the foot ( ruppert , fox and barnes 2004 ) .
jumping involves the rapid extension of the muscular foot , propelling the shell backwards . it is not uncommon for several jumps to occur in response ( savazzi 1985 ) .
ploughing , horizontal movement through the sediment , is the process of repeated burrowing sequences , with the hinge line horizontal and the foot pointing forward ( savazzi 1985 ) .
laboratory tests completed by savazzi ( 1985 ) found that movement involved in emerging from substrate occurred in a single movement , displacing the cardiid forward . emerging was often a result of insufficient oxygenation ( savazzi 1985 ) .
respire through the use of gas exchange which primarily occurs via the gills . additionally , when exposed from water , aerial respiration is possible through oxygen diffusion through the mantle ( kawaguti 1983 ) .
most heterodont bivalves are suspension feeders ( fig . 14 ) , pumping water and plankton in through an inhalant siphon ( ruppert , fox and barnes 2004 ) . large ctenidia are used to filter water and food particles . within ctenidia , water is moved using specialised cilia , that create current , causing the oxygenated and plankton filled waters to pump past the thin gill filaments of the demibranchs ( wilbur and yonge 1964 ) . this allows nutrients and gasses to diffuse across the cells . the filtered water is then pumped out through the exhalent siphon ( ruppert , fox and barnes 2004 ) .
have a similar nervous system to other bivalves . where a reduction of the head has resulted in limited cephalisation . instead of having a single large brain , bivalves have four separated ganglia ( wilbur and yonge 1964 ) :
cerebropleural ganglia , aids in the coordination of the muscular foot and the adductor muscle . pedal ganglia , provide motor control over the anterior pedal retractor muscles , byssal retractor muscles and the muscular foot . visceral ganglia , are responsible for the mantle , siphons , gills , heart , nephridia and gonads . siphonal ganglia , receives sensory information obtained by the siphons ( ruppert , fox and barnes 2004 ) .
as the head is located internally within the bivalves’ shell , it is unable to collect sensory information , and this task has been passed onto the foot and siphons ( rudman 1998 ) .
evolutionally bivalves undergone a complete loss of their radula , and developed two shells ( valves ) that are bilaterally symmetrical , with their hinge as the axial point . balvalia can be grouped into three major clades based on their morphological characteristics ( ruppert , fox and barnes 2004 ) :
consists of most extant bivalves , they are filter feeders with complex filibranch gills .
, was an important economic subsistence species for aboriginal communities in the western torres strait , before modern times .
has not yet been assessed for the iucn red list , and the current status of the species in unknown . however , like most other animals with a calcium carbonate , strawberry cockles are becoming more and more disadvantaged as the oceans become more acidic .
atlas of living australia . distribution map ( n . d . ) retrieved from
, new york , ny : mcgraw - hill companies , pp . 352 - 358 .
linnaeus , c 1758 , “system naturae” , regnum animale . cura societatis zoologicae germanicae ( 10th edi ) ,
ohno , t , katoh , t and yamasu , t 1995 . “the origin of algal - bivalve photo - symbiosis” ,
poutiers , jm 1998 , “bivalves . acephala , lamellibranchia , pelecypoda” , pp . 123 - 362 . in carpenter , ke and
, vol . 1 , seaweeds , corals , bivalves , and gastropods . rome ,
, belmont , calif : thomson - brooks / cole , pp . 367 - 403 .
acanthocardia tuberculate ) ” , stuttgarter beitr . naturk . a ( biol . ) , vol . 353 , pp . 1 - 12 .
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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
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shell solid and inequilateral , umbonal keel rounded , subquadrate to trapezoidal with strongly digitate posterior margin . no strong keel . strong broad , flat radial ribs ranging from 23 to 31 , mean 27 . lunule narrow , smooth with raised dorsal margin . escutcheon broad , smooth . exterior white , ribs with irregular , transverse red scales , sometimes with pale brown blotches ; interior white . it is distinct from all other fragines by the prominent red scales on the ribs of the shell . shell length and width to around 5 cm while depth to above 6 cm .
this is the most heavily bodied , heavily shelled species of the subfamily fraginae . in contrast to corculum that has a very light , thin and transparent shell ( in areas ) , this species also participates in photosymbiosis but in a very different manner . soft anatomy is not contained within the shell but is exposed as highly distinct leaf - like extensions of the posterior siphonal area , similar to but unique from the hypertrophied mantle of giant clams . these are spread out on the surface of the sediment and function to greatly increase surface area for photosymbionts that are contained within the exposed tissue .
all cockles or cardiids have short siphons . because of this morphological constraint , infaunal cardiids live close to the sediment water interface in order to filter water . this could be considered a preadaptation for a photosymbiotic lifestyle , as it also facilitates optimal light penetration into the soft tissues in shell interior where photosymbionts are housed .
rights we support the open release of data and information about our collections . text content on this page is licensed under a creative commons attribution 4 . 0 international license . image content on this page is copyright wa museum .
the great barrier reef and adjacent isles : a comprehensive survey for visitor , naturalist and photographer .
hylleberg , j . ( 2004 ) . lexical approach to cardiacea . records of taxa . illustrated and annotated records of living and fossil shells , with emphasis on the families cardiidae and lymnocardiidae ( mollusca : bivalvia ) . part 3 : n - z .
kirkendale , l . ( 2009 ) . their day in the sun : molecular phylogenetics and origin of photosymbiosis in the ‘other’ group of photosymbiotic marine bivalves ( cardiidae : fraginae ) .
brisbane , queensland : jacaranda press . nielsen , c . ( 1976 ) . checklist of bivalves from pmbc beach and reef flat , phuket , thailand .
oostingh , c . h . ( 1925 ) . report on a collection of recent shells from obi and halmahera , molluccas .
poorten , j . j . ter . ( 2009 ) . the cardiidae of the panglao marine biodiversity project 2004 and the panglao 2005 deep - sea cruise with descriptions of four new species ( bivalvia ) . vita malacologica , 8 , 9 - 96 .
persselin , s . l . ( 1998 ) . the evolution of shell windows within the fraginae ( bivalvia : cardiidae ) and the origin of algal symbiosis in cardiids . m . s . thesis in biology : university of guam .
reeve , l . a . ( ed . ) . conchologica iconica . london : reeve & bentham vol . 2 ( as
, report on the zoological collections made in the indo - pacific ocean during the voyage of the h . m . s . ' alert ' 1881 - 2 . london , british museum trustees , printed by taylor , & francis . [ part i . collections of melanesia , pp . 34 - 116 ; part ii . collections from the western indian ocean , pp . 487 - 508 ; explanation of plates pp . 657 - 659 ] .
tan , s . k . , & low , m . e . y . ( 2014 ) . checklist of the mollusca of cocos ( keeling ) / christmas island ecoregion . raffles bulletin of zoology , suppl . , 30 : 313 - 375 .
wilson , b . r . , & stevenson , s . e . ( 1977 ) . cardiidae ( mollusca , bivalvia ) of western australia .
willan , r . c . , bryce , c . , & slack - smith , s . m . ( 2015 ) . kimberley marine biota . historical data : molluscs .
shell medium , inequilateral , shell notably higher than long , sharply pointed ventrally with strong keel . posterior flattened and narrow , steep . sculpture of strong radial ribs circa 32 range 27 - 37 ( wam material ) , surfaces broad and flat . ribs flattened with close set transverse curved scales , becoming coarser anteriorly . lateral teeth extremely inequidistant from cardinals . shell cream , rugae or nodules on ribs often yellowish . interior color white , but with often yellow - orange flares radiating from umbonal cavity .
intertidal to shallow subtidal insandy areas . generally in sheltered areas . in reefal habitats can be found from sandy areas through to seagrass bed perimeters and rubble .
qld , nt and wa in australia , but widely distributed in the indo - west pacific .
hedley , c . ( 1899 ) . the mollusca of funafuti . part 2 . pelecypoda and brachiopoda .
hylleberg , j . ( 2004 ) . lexical approach to cardiacea . records of taxa . illustrated and annotated records of living and fossil shells , with emphasis on the families cardiidae and lymnocardiidae ( mollusca : bivalvia ) . part 2 : a - m .
. vol . 1 . bathhurst , new south wales : crawford house press .
maes , v . o . ( 1967 ) . the littoral marine mollusks of cocos - keeling islands ( indian ocean ) .
persselin , s . l . ( 1998 ) . the evolution of shell windows within the fraginae ( bivalvia : cardiidae ) and the origin of algal symbiosis in cardiids . m . s . thesis in biology , university of guam .
reeve , l . a . ( ed . ) . conchologica iconica , vol . 2 . london :
tan , s . k . & low , m . e . y . ( 2014 ) . checklist of the mollusca of cocos ( keeling ) / christmas island ecoregion . raffles bulletin of zoology , suppl . , 30 , 313 - 375 .
tantanasiriwong , r . ( 1979 ) . a checklist of marine bivalves from phuket ( area ) .
warning : the ncbi web site requires javascript to function . more . . .
department of biology , faculty of science , shimane university , matsue , japan .
these keywords were added by machine and not by the authors . this process is experimental and the keywords may be updated as the learning algorithm improves .
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