tellina tenuis , the thin tellin , is a species of marine bivalve mollusc in the family tellinidae .

it is found off the coasts of north west europe and in the mediterranean sea where it lives buried in sandy sediments .

bivalves are molluscs with a body compressed between two usually similar shell valves joined by an elastic ligament .

there are teeth at the edge of the shell and the animal has a muscular foot , gills , siphons , mouth and gut and is surrounded by a mantle inside the shell .

tellina tenuis

scientific synonyms and common names tellina tenuis da costa , 1778 tellina exigua poli , 1791 tellina polita pulteney , 1799 non tellina polita poli , 1795 non tellina polita spengler , 1798 tellina exigua var . alba costa o . g . , 1830 tellina exigua var . flavescens costa o . g . , 1830 tellina hyalina deshayes , 1835 non tellina hyalina gmelin , 1791 tellina exigua deshayes , 1835 non tellina exigua gmelin , 1791 tellina tenuis var . angusta philippi , 1836 non tellina angusta gmelin , 1791 macoma commutata monterosato , 1884 tellina exigua var . flavida monterosato , 1884 tellina exigua var . pudibunda monterosato , 1884 tellina exigua var . albida monterosato , 1884 tellina exigua var . major monterosato , 1884 tellina exigua var . minor monterosato , 1884 tellina exigua var . aurantia monterosato , 1884 tellina tenuis var . aita de gregorio , 1885 tellina tenuis var . browni de gregorio , 1885 tellina tenuis var . jeffreysi de gregorio , 1885 tellina tenuis var . brevior b . d . d . , 1886 tellina bourguignati locard , 1886 tellina tenuis var . maxima b . d . d . , 1886 tellina tenuis var . minuta b . d . d . , 1886 tellina carnaria sensu von born , 1780 non linné , 1758 tellina incarnata sensu chemnitz , 1782 non linné , 1758 tellina planata sensu pennant , 1777 non linné , 1758 tellina tenuis sensu auct . non da costa , 1778 tellina exilis sensu brusina , 1866 non lamarck , 1818 angulus tenuis
worms - world register of marine species - tellina tenuis var . angusta philippi , 1836
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tellina tenuis
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in kames bay the density of tellina tenuis falls in a progressive manner from l . w . m . to h . w . m .
distribution tellina tenuis was found in both periods in the western near - coastal zone . in addition , the species was also observed in . . .
the size - frequency curves and density of tellina tenuis in the other bays at l . w . m . correspond with those of half - tide in kames bay .
the present paper deals with the results of investigations into certain phases in the life - history of the bivalve mollusc tellina tenuis carried out during the autumn of 1926 and 1927 .
the tellina tenuis population in kames bay seems to be composed of four year - groups , one of which is almost unrepresented . collections from neighbouring bays indicate that older groups may be present .
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tellina crassa
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tellina pulchella
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tellina squalida
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dekker , r and beukema , j . j 1999 . relations of summer and winter temperatures with dynamics and growth of two bivalves , tellina tenuis and abra tenuis , on the northern edge of their intertidal distribution . journal of sea research , vol . 42 , issue . 3 , p . 207 .
holme , n . a . 1950 . population - dispersion in tellina tenuis da costa . journal of the marine biological association of the united kingdom , vol . 29 , issue . 02 , p . 267 .
barnett , peter r . o . 1985 . the effect of temperature on the growth of planktonic larvae of tellina tenuis da costa . journal of experimental marine biology and ecology , vol . 89 , issue . 1 , p . 1 .
ansell , alan d . and trevallion , ann 1967 . studies on tellina tenuis da costa i . seasonal growth and biochemical cycle . journal of experimental marine biology and ecology , vol . 1 , issue . 2 , p . 220 .
mcintyre , a . d . 1970 . the range of biomass in intertidal sand , with special reference to the bivalve tellina tenuis . journal of the marine biological association of the united kingdom , vol . 50 , issue . 03 , p . 561 .
the two closely related species , t . tenuis and t . fabula , are both plentiful in kames bay , but their range is not coincident .
buchanan , j . s . 1978 . cytological studies on a new species of rickettsia found in association with a phage in the digestive gland of the marine bivalve mollusc , tellina tenuis ( da costa ) . journal of fish diseases , vol . 1 , issue . 1 , p . 27 .
young tellina tenuis , passing the 2 - mm . sieve but retained by the 1 - mm . sieve , were found in kames bay in august , chiefly below l . w . m . , but in october plentifully distributed up to half - tide and in lesser numbers higher up the shore .
in kames bay tellina tenuis ranges from a little below h . w . m . to depths of about 4 fm . the maximum concentration of about 1000 per sq . m . is found at l . w . m . springs and the numbers decrease to zero at h . w . m . and 4 fm .
distribution tellina tenuis was found in both periods in the western near - coastal zone . in addition , the species was also observed in the area of the hinder banks , but only in the 1994 - 2001 period . the frequency of occurrence in both periods was relatively low and the species never reached high density levels ( maximum 30 ind . / m2 ) . [ details ]
habitat tellina tenuis occurs in a wide range of sediments : median grain size of 150 to 500 μm . the main restrictive factor appears to be the mud content : the species only occurs in sediments with a mud content < 20 % . the relative occurrence of 10 % in sediments with a mud content of 40 - 50 % is considered an outlier and hence seen as unreliable . [ details ]
the dispersion of tellina tenuis da costa in the laboratory was analysed by the clark and evans nearest - neighbour test and by the kolmogorov - smirnov one - sample test . the dispersion was random , with a slight tendency toward aggregation that was independent of density . in the field , the dispersion was analysed by the clark and evans nearest - neighbour test and by the x 2 approximation to fisher ' s coefficient of dispersion . the dispersion was again random , this time with a slight tendency toward spacing out that was independent of density . the tendency toward aggregation displayed in the laboratory was independent of the dispersion pattern shown at the start of the experiment , and also unaffected by the edge of the container . the apparent randomness suggests that t . tenuis is primarily a suspension feeder , but may be a deposit feeder under certain environmental conditions .
prediction of safe levels of effluent discharge have in the past been largely based on the results of short term acute toxicity tests ( lc 5 0 s ) . such tests are usually carried out in unnatural experimental conditions , and in consequence the results are of only limited value . there is a need to enlarge the concept of the routine bioassay test to include quantitative measurement of the effects of pollutants on behaviour , physiology and metabolism . this paper describes a simple rapid bioassay test in which the effect of pollutants on the burrowing of the bivalve tellina tenuis has been used as a quantitative measure of pollutant effect .
description a thin , strongly flattened shell with an irregular oval shape . front edge broadly rounded , back edge tapering in a clear angle . the back is somewhat less acuminate than in the tellina fabula . the shell surface is practically smooth with only fine growth lines . measures up to 30 mm long . the colour varies from white with dark colour bands to pinkish red . [ details ]
the thin tellin is a species which regularly washes ashore . you usually find both valves still attached to one another . the connecting ligament between the two shell halves is namely very strong . the thin tellin is reasonably easy to find on the sand thanks to its pink to orange color . this colorful shellfish usually lives hidden in the sea bottom , where it ' s difficult to find despite its color . for food and oxygen , it extends its siphons out of the sand and filters the water . the thin tellin closely resembles the baltic tellin but is more delicate , flatter and lighter .
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ovate , posterior shortened , wedge - shaped and curves to the right when viewed from dorsal side . anterior gently sloping and broadly curved at anterior margin .
prominent mid - brown band extending less than a third the way to the posterior margin .
2 cardinals in each valve . in lv anterior bifid , posterior very weak . in rv posterior bifid . 2 laterals in rv , anterior short but prominent , posterior small , below nymph plate . 1 short posterior lateral in lv .
pallial sinus very deep , extends upwards and almost to anterior adductor and is confluent with the pallial line . cruciform muscle scars visible .
white / pink / orange / yellow external colouration more intense and covering more of the shell .
a burrower in sand and silty sand . lives from the mid intertidal to approximately 10m . widely distributed around all coasts .
but lacks the scissulate sculpture , is not so narrow and less acute posteriorly .
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m . j . de kluijver , s . s . ingalsuo & r . h . de bruyne
just behind the midline directed inwards and backwards , almost touching . approximately oval in outline with the
white , pink , rose , orange or yellow in colour and various shades of each , normally in irregular bands , exceptionally each valve may be differently coloured .
is transparent , glossy . inside of shell coloured like the outside but normally fainter .
a deep burrower in sand or mud . this habit has had effect in developing a powerful
inhabits fine sand generally from about the middle of the intertidal zone to a depth of a few metres . it sometimes occurs in very large populations , particularly near low water - mark , where it may be the most abundant macro - invertebrate , its numbers steadily decreasing up the beach , where the sands dry out appreciably at low tide , or as coarse sand is met . the depth to which the animal burrows depends on the state of the tide and may be 10 . 16 - 12 . 7 cm , when the tide is out but less at high tide .
haas , f . , 1926 . lamellibranchia . in grimpe , g . & wagner , e . : die tierwelt der nord - und ostsee .
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poppe , g . t . & y . goto , 1993 . european seashells . vol . ii . 221 pp . wiesbaden / verlag christa hemmen .
seaward , d . r . , 1990 . distribution of the marine molluscs of north west europe . nature conservancy council .
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philippi r . a . ( 1836 ) . enumeratio molluscorum siciliae cum viventium tum in tellure tertiaria fossilium , quae in itinere suo observavit . vol . 1 . schropp , berlin [ berolini ] : xiv + 267 p . , pl . 1 - 12 , available online at urltoken ; = article & id ; = 355 & itemid ; = 167 page ( s ) : 27 [ details ]
check list of european marine mollusca ( clemam ) . , available online at urltoken [ details ]
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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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da costa e . m . ( 1778 ) . historia naturalis testaceorum britanniae , or , the british conchology . london : millan , white , elmsley & robson . xii + 254 + viii pp . 17 pls . , available online at urltoken page ( s ) : 210 - 211 [ details ]
description a thin , strongly flattened shell with an irregular oval shape . front edge broadly rounded , back edge tapering in a clear . . .
long term trends in the macrobenthos of the belgian continental shelf ( macrobel ) .
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ardovini , r . ; cossignani , t . ( 2004 ) . west african seashells ( including azores , madeira and canary is . ) = conchiglie dell ' africa occidentale ( incluse azzorre , madeira e canarie ) . english - italian edition . l ' informatore piceno : ancona , italy . isbn 88 - 86070 - 11 - x . 319 pp . ( look up in imis ) page ( s ) : 52 [ details ]
zamouri - langar , n . ; chouba , l . ; ajjabi chebil , l . ; mrabet , r . ; el abed , a . ( 2011 ) . les coquillages bivalves des côtes tunisiennes . institut national des sciences et technologies de la mer : salammbô . isbn 978 - 9938 - 9512 - 0 - 2 . 128 pp . ( look up in imis ) [ details ]
backeljau , t . ( 1986 ) . lijst van de recente mariene mollusken van belgië [ list of the recent marine molluscs of belgium ] . koninklijk belgisch instituut voor natuurwetenschappen : brussels , belgium . 106 pp . ( look up in imis ) [ details ]
van zweeden , c . ; van asch , m . ; van den ende , d . ; troost , k . ( 2013 ) . het kokkelbestand in de nederlandse kustwateren in 2013 . imares wageningen report , c115 / 13 . imares wageningen ur : ijmuiden . 46 pp . ( look up in imis ) [ details ]
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in the exe estuary is shown to be uniformly distributed , indicating a significant degree of ‘over - dispersion’ .
area it is shown that fewer individuals than would be expected occur less than 1 in . from their nearest neighbour , and none occurs closer than 0·6 in .
when the population density was artificially increased on the shore the same characteristic spacing was found at a moderate density , but not at a rather higher density .
it is suggested that spacing is correlated with the foraging activities of the inhalent siphon on the soil surface .
very dense populations have been found by stephen in other areas , indicating that the size of the ‘territory’ does not limit density . his results show no evidence of the same phenomena as observed in the exe .
studies on the scottish marine fauna . additional observations on the fauna of the sandy and muddy areas of the tidal zone
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the size - frequency distribution shows a regular gradation from the lower to the higher levels . at l . w . m . and below individuals of small size predominate , while at h . w . m . they are proportionately few .
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simpson , r . : physical and biotic factors limiting the distribution and abundance of littoral molluscs on macquarie island ( sub antarctica ) . j . exp . mar . biol . ecol .
southward , a . j . : note on the temperature tolerance of some intertidal animals in relation to environmental temperatures and geographical distribution . j . mar . biol . ass . u . k .
speakman , j . n . and p . a . krenkel : quantification of the effects of rate of change on aquatic biota . wat . res .
tebble , n . : british bivalve seashells , 212 pp . london : trustees of the british museum ( n . h . ) 1966
k . v . singarajah , john moyse , e . w . knight - jones
richard w . eppley , robert w . holmes , john d . h . strickland


onthophagus is a genus of dung beetles in the onthophagini tribe of the wider scarab beetle family , scarabaeidae .

it is the most species-rich and widespread genus in the subfamily scarabaeinae ( the ' true ' dung beetles ) , with a global distribution .

onthophagus

habitus ( oblique and lateral views ) of onthophagus males ( holotypes , except 3–4 ) . 1–2 onthophagus abmisibilus 3–4 onthophagus catenatus , vicinity of mt hagen 5–6 onthophagus kokodanus kokodanus 7–8 onthophagus kokodanus hagenaltus .
this scarab could be confused with onthophagus of similar color and medium size ( more than 10 mm ) : onthophagus nigriventris and digitonthophagus gazella .
onthophagus males ( holotypes , except 13 , 15 , 18 ) . 9–11 , 21 onthophagus kokosquamatus ; 12 , 17 onthophagus abmisibilus ; 13 , 18 onthophagus catenatus , vicinity of mt hagen 14 , 15 , 19 onthophagus kokodanus kokodanus , 15 minor male e pt moresby 16 , 20 onthophagus k hagenaltus . 9–10 habitus ( oblique and lateral views ) 11–16 head and pronotum in dorsofrontal view 17–21 parameres in lateral view ( scales 1 mm ) .
onthophagus nigriventris male genitalia , caudal view ; photo by e . l . engasser
onthophagus nigriventris male genitalia , lateral view ; photo by e . l . engasser
onthophagus kokodanus hagenaltus subsp . n . – papua new guinea ( western highlands )
révision des onthophagus de l’archipel indo - néerlandais , avec description des espèces nouvelles .
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les onthophagus ( coleoptera , scarabaeidae ) des îles philippines
. –
all four species included in the onthophagus catenatus group are formally placed in the nominotypical subgenus .
onthophagus kokodanus kokodanus subsp . n . – papua new guinea ( e of port moresby )
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neue onthophagus - arten von neu - guinea and den benachbarten inseln
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hunt j , simmons lw . status - dependent selection in the dimorphic beetle onthophagus taurus .
four new taxa from new guinea are proposed in the dung beetle genus onthophagus latreille , 1802 , all in the operational group of onthophagus catenatus lansberge , 1883 . the group is discussed , defined , and the five taxa included are listed , keyed , and diagnosed . three new species are described : onthophagus abmisibilus ( from west new guinea , indonesia ) , onthophagus kokodanus , onthophagus kokosquamatus ( both from papua new guinea ) . one new species comprises a lowland and an upland subspecies : onthophagus kokodanus kokodanus and kokodanus hagenaltus ( both in papua new guinea ) .
phylogenetics and biogeography of the dung beetle genus onthophagus inferred from mitochondrial genomes . - pubmed - ncbi
taxonomy of papuasian onthophagus : twenty new species and their relatives ( coleoptera : scarabaeidae : scarabaeinae ) .
new guinea onthophagus : taxonomy of ten new small , unicolour species ( coleoptera : scarabaeidae : scarabaeinae ) .
new guinea onthophagus : taxonomy of ten small , unicolored new species ( coleoptera : scarabaeidae : scarabaeinae ) .
genital morphology and fertilization success in the dung beetle onthophagus taurus : an example of sexually selected male genitalia .
hunt j , simmons lw . maternal and paternal effects on offspring phenotype in the dung beetle onthophagus taurus .
diversity in the weapons of sexual selection : horn evolution in the beetle genus onthophagus ( coleoptera : scarabaeidae ) .
the mega - diverse subcosmopolitan genus onthophagus is currently represented by approximately 2 , 000 species worldwide ( scarabnet 2009 ) .
details of morphology of onthophagus . 22–26 onthophagus abmisibilus , holotype 27–32 onthophagus catenatus , male , vicinity of mt hagen 32 female , n pt moresby 22 , 27 , 32 head in dorsal view 23 , 28 pronotum in dorsal view 24 , 29 elytron in dorsal view 25 , 30 protibia , upper side 26 , 31 metatibia , underside . scales 1 mm .
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a new record of onthophagus ( sunenaga ) wallacei ( coleoptera , scarabaeidae ) from halmahera , maluku
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new guinea onthophagus : taxonomy of ten small , unicolored new species ( coleoptera : scarabaeidae : scarabaeinae ) . - pubmed - ncbi
this species could be confused with the similarly colored and medum - sized ( more than 10 mm ) digitonthophagus gazella and onthophagus sagittarius .
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six new taxon [ sic ] of the subgenus indachoriusof the genus onthophagus ( coleoptera : scarabaeidae ) from borneo
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diversity in the weapons of sexual selection : horn evolution in the beetle genus onthophagus ( coleoptera : scarabaeidae ) . - pubmed - ncbi
moczek ap , emlen dj . male horn dimorphism in the scarab beetle , onthophagus taurus : do alternative reproductive tactics favour alternative phenotypes ?
huijbregts j ( 2012 ) . new guinea onthophagus : taxonomy of ten small , unicolored new species ( coleoptera : scarabaeidae : scarabaeinae ) .
what made you want to look up onthophagus ? please tell us where you read or heard it ( including the quote , if possible ) .
reiche , l . ( 1840 ) note sur deux espèces d ' onthophagus . revue et magasin de zoologie , pure et appliquée , 3 , 243–244 .
no reason is provided by smith ( 2009 : 43 ) for treating the senior name onthophagus curvicornis latreille 1812 as synonym of the junior name onthophagus incensus say 1835 . it may be that there is a homonymy , or that the senior name is for some reason unavailable , but further research would be required to determine this
simonnet f , moczek ap ( 2011 ) conservation and diversification of gene function during mouthpart development in onthophagus beetles . evol dev 13 : 280 - 289 . doi :
to cite this page : roof , j . 1999 .
onthophagus australis
( on - line ) , animal diversity web . accessed july 09 , 2018 at urltoken
google scholar ( 2016 ) [ digital resource with search terms :
digitonthophagus gazella
or
onthophagus gazella
] . available from urltoken [ accessed 25 august 2016 ] .
onthophagus species are boxed up in red . this picture is taken in the raffles museum of biodiversity research , national university of singapore , singapore . picture by m . y neo .
as o . semifex is not individually studied till date , many of its ecology habits are not well documented . however , their characteristics can be possibly inferred from the other onthophagus relatives .
the red arrows indicate the cupreous appearance on the onthophagus species . this picture is taken in the raffles musuem of biodiveristy research , national university of singapore , singapore . picture is by m . y . neo
details of morphology of onthophagus kokosquamatus ( male holotype ) 45 head in dorsal view 46 pronotum and head armature in dorsal view 47 elytron in dorsal view 48 protibia , upper side 49 metatibia , underside . scales 1 mm .
details of morphology of onthophagus ( holotypes , except 38 , 44 ) . 33–38 o . k . kokodanus , 38 female paratype , e pt moresby 39–44 onthophagus kokodanus hagenaltus 44 minor male paratype , mt hagen 33 , 38 , 39 , 44 head in dorsal view 34 , 40 pronotum and head armature in dorsal view 35 , 41 elytron in dorsal view 36 , 42 protibia , upper side 37 , 43 metatibia , underside . scales 1 mm .
scarab beetles of the genus onthophagus latreille north of mexico ( coleoptera : scarabaeidae ) howden h . f . , cartwright o . l . 1963 . proc . u . s . nat . mus . 114 : 1 - 135 .
krikken , j . & huijbregts , j . ( 2008 ) . distinguishing the sundaland species in the onthophagus ( parascatonomus ) aurifex group ( coleoptera : scarabaeidae : scarabaeinae ) . tijdschrift voor entomologie . pp . 151 : 173 - 185 .
emlen , d . j . & nijhout , f . h . ( 1999 ) . hormal control of male horn length dimorphism in the dung beetle onthophagus taurus ( coleoptera : scarabaeidae ) . journal of insect physiology . pp . 45 : 45 - 53 .
onthophagus fracticornis ( preyssler , 1790 ) : tronquet ( 2014 ) : 387 . [ statut pour la france métropolitaine ] tronquet , m . [ coord . ] 2014 . catalogue des coléoptères de france . revue de l’association roussillonnaise d’entomologie , 23 ( supplément ) : 1 - 1052 .
your discussions , opinions and reviews for this site will be greatly appreciated . if you have any new information on onthophagus semifex that you wish to be added in , do leave your reference material and your contact in the discussion box below . your request will be attended as soon as possible .
onthophagus is the most diverse tunneling dung beetle genus , with over 2 , 000 species described [ 32 ] . in the united states alone , there are at least 35 native onthophagus spp . [ 32 ] and at least 5 more introduced , naturalized , non - native species [ 33 – 35 ] . all of these species are known to be dung specialists . onthophagus taurus , the bull - headed dung beetle , is one of the most abundant dung beetles specialized on cattle dung . it is a native of the mediterranean , was first found in florida in 1974 [ 36 ] , and has increased its distribution throughout the u . s . ever since [ 37 ] . although native dung beetles such as o . hecate and o . pennsylvanicus are also found in agricultural habitats , o . taurus is the most abundant beetle in these settings . however , o . taurus is uncommon in non - agricultural ecosystems [ 37 ] .
citation : estes am , hearn dj , snell - rood ec , feindler m , feeser k , abebe t , et al . ( 2013 ) brood ball - mediated transmission of microbiome members in the dung beetle , onthophagus taurus ( coleoptera : scarabaeidae ) . plos one 8 ( 11 ) : e79061 . urltoken
this dataset contains the digitized treatments in plazi based on the original journal article krikken , j . , huijbregts , j . ( 2013 ) : new guinea onthophagus : taxonomy of ten small , unicolored new species ( coleoptera : scarabaeidae : scarabaeinae ) . zootaxa 3619 ( 5 ) : 501 - 525 , doi :
bianchin , i . , alves , r . g . o . & koller , w . w . ( 1998 ) efeito de carrapaticidas / inseticidas “pour - on” sobre adultos do besouro coprófago africano onthophagus gazella fabr . ( coleoptera : scarabaeidae ) . anais da sociedade entomológica do brasil , 27 , 275–279 . urltoken
the presence of an isolated conical protrusion right in front of the horns is shared with other new guinea species , like onthophagus heurni gillet , 1930 and joliveti paulian , 1973 , which may be confusing . the males of both these species , however , have on their horns , which are more or less erect , a distinct basal - internal lobe or denticle , and their eyes are narrow .
there are currently six onthophagus species that can still be found in the uk ( extant species ) and two that are now thought to be extinct . in many species , males have a noticeably pointed horn on the posterior margin of the head . the beetles can also be sexed by looking at the abdomen – in females the pygidium is uniform in width and , in males it is narrower in the middle ( see image below ) .
based on the nutritionally incomplete diet dung provides , the nesting behavior of the female , and the larval feeding behavior , we hypothesize that a vertically transmitted microbiome is present that is transmitted through the brood ball in the dung beetle , onthophagus taurus . here we use a sterile rearing technique along with culture based and molecular methods to assess the potential for microbiome transmission and inheritance of the bull headed dung beetle , o . taurus , through the brood ball .
established . there is conflicting information regarding the arrival of this species to hawaii . markin and yoshioka ( 1998 ) reported that onthophagus sagittarius was purposely released on oahu in 1957 and 1958 . however , harris et al . ( 1982 ) stated that the species was accidentally introduced . regardless , this scarab is now established on both molokai and oahu , where it is one of the most commonly encountered dung beetles ( nishida , 2002 ; harris et al . , 1982 ) .
many of the species on earth have yet to be described or documented , thus they are still unknown and not recognized for their ecological influences on nature and man . luckily for onthophagus semifex , several aspects of its biology are described in the year 2008 , though it is still relatively incomplete and insufficient as compared to many other renowned animals . but to be able to gain an official name , this little dung beetle is fortunate enough to enter the encyclopedia of life before its extinction .
o . semifex , similar to other onthophagus tropical dung beetles , may locate their food resource by smell . they fly in a zig - zag manner , usually one meter or less above ground and land close to the source of the odour . they are also known to be tunnellers , which they bury the dung in preformed burrows . they transport their food fast owing to resource competition by other dung beetles , necrophagous beetles and flies ( hanski , 1989 ; philips et al . , 2004 ) .
like related onthophagus species , this nocturnal scarab is a dung tunneler , with the female creating a burrow under or near a fresh dung source ( simmons and emlen , 2008 ) . the burrow is then provisioned with dung in the form of brood balls . each ball is impregnated with an egg ; larval development occurs within the brood ball . this species is confined to tropical areas that experience warm , wet summers , and annual rainfall over 800 mm ( 31 . 5 in ) ( edwards , 2007 ) .
till date , there are currently eight dung beetle representatives belonging to the o . aurifex species group , under the genus onthophagus of latreille , 1802 . o . aurifex was made the ‘type’ of this group . furthermore the group has been divided into two complexes - the o . aurifex complex and the o . sarawacus complex in a recent article published by tarasov et al . ( 2010 ) . in this , the authors classified four representatives in the o . aurifex complex , which are characterized by their shiny metallic upper - side body .
figures 1 – 8 . habitus ( oblique view ) of onthophagus species , with indication of body length . 1 , o . acerus , holotype , , 6 mm ; 2 , o . aceroides , holotype , 5 mm ; 3 , o . baiyericus , holotype , 5 mm ; 4 , o . mimikanus , female holotype , 6 mm ; 5 , o . kokopygus , holotype , 3 mm ; 6 , o . daymanus , holotype , 4 . 5 mm ; 7 , o . kokocellosus , holotype , 4 . 5 mm ; 8 , o . bituberoculus , female paratype , timika , 7 mm .
total body length 10 . 0–13 . 0 mm ( 0 . 39–0 . 52 in ) . body shape oval ; may be caked in dung . color dark brown ; elytra pale brown . medium - sized onthophagus , more than 10 mm . clypeal apex rounded to sinuate ; not strongly produced or reflexed in either sex . head of male with paired tusk - like horns on the clypeus ; female with single horn on the frons . ocular canthus not completely dividing eye . pronotum with anterior angle rounded . pronotum of male with broad , hump - like process ; female with spine - like process . front tibia of male and female similar . scutellum absent .
diagnosis . clypeal apex bidentate ; major males have their denticles set on a reflexed lobe ( emargination between denticles not going down to base of lobe ) , this lobe on either side delimited by a sinuate border ( contrary to o . dissidentatus ) . clypeus and frons in both sexes without any further protrusions ( contrary to o . dissidentatus ) . onthophagus kokodentatus usually shiny black ( occasionally with metallic lustre ) ; most of dorsum distinctly , finely punctate , punctation varying in strength . eye foramina narrow . pronotum entirely evenly convex , lateral and basal borders evenly , widely rounded . proximal serrate section of protibia long ( 8 – 10 very small denticles ) . pygidium simply punctate . body length usually 3 . 5 – 4 . 5 mm .
adults fly into the dung pat to feed and mate . beneath the dung pat , the juvenile life stages of onthophagus taurus are isolated in the brood chamber of the brood balls constructed by the female beetles in tunnels . females lay several brood balls in each tunnel that would all be at the same developmental stage . however , for illustrative purposes all life stages are represented in one tunnel . these stages include the : ( a ) egg , ( b ) 1 st larval instar , ( c ) 2 nd larval instar , ( d ) 3 rd larval instar , ( e ) pupa , and ( f ) an eclosing adult beetle that is tunneling toward the surface . the brood ball chamber is larger with each successive life stage as the larva feeds on the chamber walls within the brood ball . the top inset shows ( g ) the fecal pedestal the egg is positioned upon in brood ball . the bottom inset shows ( h ) the larval instar feeding on the walls of the brood ball chamber .
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37 in our area , 130 in the new world , > 1500 spp . total
the scarabaeoid beetles of nebraska brett c . ratcliffe & m . j . paulsen . 2008 . university of nebraska state museum , vol 22 , 570 pp .
peterson field guides : beetles richard e . white . 1983 . houghton mifflin company .
scarab beetles ( coleoptera : scarabaeidae ) of south carolina phillip j . harpootlian . 2001 . clemson university public service .
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this species tends to be found in locations with a slightly heavier soil type i . e . some clay content . the brood burrows are vertically constructed with no side chambers and the brood balls are stacked one on top of the other .
distribution : south to central england and wales , absent from north england and scotland .
this beetle was formerly found in good numbers in hampshire but has declined massively in this area . it is now most frequently found in the mendip hills in somerset but , even here it remains rare .
distribution : south england , less common central england and wales . absent from scotland .
this species prefers alluvial soils and it is therefore most often found on riverside pastures and flood plains .
this species was widespread but is now rare . it is a sandy soils specialist – most often found on sand dunes .
a large and very distinctive looking beetle . the males have two long curved horns which are lacking in the female .
distribution : the last mainland record for this species was in the 1800’s . it is still present in the channel islands .
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1913 ) , which shows a distribution restricted to tropical east african highlands and cool temperate highlands of south africa .
tunnelling species are predominantly from cattle dung with a few records from buffalo , giant forest hog , zebra , and elephant dung ( australian csiro 1970 - 1986 , unpublished records ) . it has been recorded entirely from finer - grained soils , especially lateritic sandy clay loam , but occasionally also on black cotton clay or sandy loam . all records are from open vegetation , particularly extensive highland pastures and grassland , but it has occasionally been found in pastures within forest or woodland clearings .
some environmental characteristics for 48 locality records are as follows : altitude : mean : 1 , 929 ± 357 ( s . d . ) , range : 812 - 2 , 509 m ; annual rainfall : mean : 1 , 004 ± 254 ( s . d . ) , range : 538 - 1 , 623 mm ; annual temperature : mean : 17 . 0 ± 2 . 3 ( s . d . ) , range : 13 . 9 - 23 . 4 o c ( max . + min . / 2 ) .
although this species is clustered around forested highlands , primarily in east africa , it has been frequently collected on cattle dung in open vegetation , which suggests that it is not affected by any major threats to forest habitats .
there are no species - specific conservation actions in place for this species , and it is unlikely that any actions are required at present , as it is frequently recorded in the open vegetation of farming areas .
to make use of this information , please check the < terms of use > .
east africa . this species is native to moist highlands in eastern africa . it also was introduced to australia ( tyndale - biscoe , 1990 ) .
none . this species feeds on dung as both an adult and larva . there are no records of this beetle feeding on live plant tissues .
( tyndale - biscoe , 1990 ) : adults of this diurnal species live 2–4 months . during that time , adults actively fly in search of fresh dung . females create oval - shaped brood balls in burrows constructed under or near dung . development from egg to adult requires 4–7 weeks . there are multiple generations per year .
none . this species recycles dung and is beneficial for ranching and farming in hawaii . primarily being a dung feeder , this species has never been recorded damaging crop or ornamental plants . additionally , this species is not a threat to native dung beetles because none occur in hawaii or guam .
established . in hawaii , this species was imported in 1975 to big island and maui to combat the horn fly ( haematobia irritans ) , a biting pest of livestock ( markin and yoshioka , 1998 ) . it is established in the highlands of both islands , being rare on maui ( krushelnycky et al . , 2007 ) but more common on big island ( markin and yoshioka , 1998 ) .
not established or recorded . there are no records of this species from guam .
major males are readily separated by examining the head armature ( o . nigriventris lacking horns on head versus d . gazella with 2 short , upward curving horns and o . sagittarius with 2 tusk - like horns ) .
minor males and females can be separated by examining base of the head ( o . nigriventris with a sinuate ridge versus d . gazella with straight , transverse ridge versus o . sagittarius with single horn ) .
southeastern asia . this species is native to southeastern asia , where it has been recorded from malaysia , indochina ( hawaiian entomological society , 1964 ) , india ( chandra , 2000 ) , and sri lanka ( edwards , 2007 ) . this species was introduced to australia ( edwards , 2007 ) .
none . this species recycles dung and is beneficial for ranching and farming in hawaii . primarily being a dung feeder , this species has never been recorded damaging crop or ornamental plants . additionally , this scarab is not a threat to native dung beetles because none occur in hawaii or guam .
major males of these species can quickly be distinguished by examination of the head armature ( o . sagittarius with two tusk - like horns on the clypeus versus o . nigriventris without horns or ridges , d . gazella with two short , upward curving horns at the base of the head ) .
females and can be separated by examining the base of the head ( o . sagittarius with a single horn o . nigriventris with a sinuate ridge , d . gazella with a straight , transverse ridge ) .
the range of the native dung beetle is over most of southeastern australia . it can be found from southern queensland to tasmania and south australia .
the most distinctive feature is the segmented antennae . this antennae forms a club containing three to seven leaves . these leaves can be expanded or folded together to form one compact club . the beetle is brightly colored . the surface of the dung beetle ' s head varies , but it is short , broad , and partially deflexed . the eyes are oval and prominent . the wings are large and well - developed .
native dung beetles emerge mainly during late summer into fall . for the first several weeks they are in a
maturation feeding stage
. a majority of beetles are in the adult stage during the winter . ocassionally eggs are laid in the fall , which then emerge in spring .
it is unclear what the exact function of food balls is in the life cycle of the native dung beetle . while most beetles store dung in food chambers before reproduction , the native dung beetle forms food balls during reproduction . this leads to
being the only beetle found in dung when the first population of bush flies begin oviposition in the spring .
living in australia , new zealand , tasmania , new guinea and associated islands .
having body symmetry such that the animal can be divided in one plane into two mirror - image halves . animals with bilateral symmetry have dorsal and ventral sides , as well as anterior and posterior ends . synapomorphy of the bilateria .
the area in which the animal is naturally found , the region in which it is endemic .
a terrestrial biome . savannas are grasslands with scattered individual trees that do not form a closed canopy . extensive savannas are found in parts of subtropical and tropical africa and south america , and in australia .
a grassland with scattered trees or scattered clumps of trees , a type of community intermediate between grassland and forest . see also tropical savanna and grassland biome .
a terrestrial biome found in temperate latitudes ( > 23 . 5° n or s latitude ) . vegetation is made up mostly of grasses , the height and species diversity of which depend largely on the amount of moisture available . fire and grazing are important in the long - term maintenance of grasslands .
the beetles of the united states . ross arnett . the american entomological institute . loudonville , new york . 1968 .
phenology of o . australis . marina tyndale - biscoe and josephine walker . australian journal of zoology . volume 40 ( 3 ) .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
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breeschoten t 1 , doorenweerd c 2 , tarasov s 3 , vogler ap 4 .
department of life sciences , natural history museum , cromwell road , london sw7 5bd , united kingdom ; naturalis biodiversity center , darwinweg 2 , 2333 cr leiden , the netherlands ; institute biology leiden , leiden university , sylviusweg 72 , 2333 be leiden , the netherlands . electronic address : thijmen . breeschoten @ gmail . com .
naturalis biodiversity center , darwinweg 2 , 2333 cr leiden , the netherlands ; institute for biodiversity and ecosystem dynamics , university of amsterdam , science park 904 , 1098 xh amsterdam , the netherlands .
national institute for mathematical and biological synthesis , university of tennessee , 1122 volunteer blvd , ste . 106 , knoxville , tn 37996 , usa .
department of life sciences , natural history museum , cromwell road , london sw7 5bd , united kingdom ; department of life sciences , silwood park campus , imperial college london , buckhurst road , ascot sl5 7py , united kingdom .
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towson university , department of biological sciences , baltimore , maryland , united states of america , j . craig venter institute , inc . , plant genomics , rockville , maryland , united states of america ,
current address : department of ecology , evolution , and behavior , university of minnesota , st . paul , minnesota , united states of america
contributed equally to this work with : julie c . dunning hotopp , armin p . moczek
copyright : © 2013 estes et al . this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
funding : this research was supported in part through funds from : the towson university fisher college undergraduate research awards ( kf , ta ) , a towson university undergraduate research award ( kf ) , orchid society of arizona funding ( ame ) , nsf reu dbi0552654 ( mf ) , nsf ios 0445661 ( am ) , nsf ios 0718522 ( am ) , and state of maryland internal funds ( jdh ) . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
although it may seem surprising that such diverse insects have radiated onto a nutritionally - poor resource , mutualistic symbioses between bacteria and their eukaryotic hosts allow animals to feed on a diversity of diets that would otherwise be inaccessible to the host . such beneficial endosymbionts can provide essential amino acids and vitamins lacking in the host diet , or they can synthesize novel enzymes , such as cellulases and hydrolases , to degrade otherwise indigestible materials like cellulose , lignin , and chitin [ 9 , 10 ] . these mutualisms are seen in animals ranging from cellulose feeding vertebrates to wood - , sap - , and blood - feeding insects [ 9 ] . in insects , mutualistic endosymbionts frequently supplement the host with essential amino acids and vitamins missing from their food source [ 10 , 11 ] . this supplementation may be provided primarily by one symbiont species , as in the aphid - buchnera system [ 12 , 13 ] , or by a community of symbionts , such as in termites [ 14 , 15 ] .
no matter the number of beneficial endosymbionts , the faithful transmission of these specific mutualists from parent to offspring is essential for offspring survival . there are two broad categories of transmission : vertical transmission , where symbionts are acquired from the parent , and horizontal transmission , where they are not [ 9 , 16 ] . in the dung beetle system , horizontal transmission could be from many sources including adult siblings , other insect taxa including other species of dung beetles , and host dung . vertical transmission is the dominant transmission type in evolutionarily stable , obligate insect - bacterial nutritional mutualisms . vertically transmitted endosymbionts are frequently transmitted transovarially , though alternate methods of transmission from parents to offspring are known [ 9 , 16 ] . these include endosymbiont transmission through proctodeal trophallaxis [ 17 ] , milk glands [ 18 , 19 ] , coprophagy [ 20 ] , egg smearing [ 21 , 22 ] , capsules [ 23 ] , and brood chambers [ 24 ] .
the clovr pipeline with chimera check [ 51 ] was run on 275 sequences for a 1425 bp amplicon of the 16s rrna gene . clovr uses scripts from qiime [ 52 ] to cluster reads into operational taxonomic units ( otus ) , select representative sequences from each otu , and use the ribosomal database naïve bayesian classifier ( 0 . 5 confidence interval ) to assign each otu to a known taxon [ 53 ] . the nucleotide sequence identity within an otu is 95 % . otu’s ( n = 22 ) that had less than an 80 % confidence with rdp were not classified [ 54 , 55 ] . clovr uses scripts from mothur [ 56 ] to plot rarefaction curves of richness and diversity , which were calculated every five sequences [ 53 , 57 ] .
to assess the relatedness of 16s rrna sequences , sequences were aligned with muscle v3 . 7 [
] , visualized and trimmed with bioedit v7 . 1 . 3 . 0 [
substitution model , random seeds of 12345 , and 100 bootstraps . the best tree as ascertained by raxml is shown and is decorated with symbols denoting the key characteristics of the sample . percent similarity was determined between all sequences (
to compare the microbiome community composition of the cultured isolates to the microbiome community found using molecular methods , all cultured isolates were screened using previously described degenerate primer sets designed for 10 single copy loci . the
] amplified the majority of the cultured isolates . cultured isolates that amplified with the
amplicons sequenced from cultured bacteria isolated from the dung beetle larval digestive system . for the constrained analysis , the concatenated
from cultured isolates : kf193370 - kf193385 ) . phylogenetic trees and amino acid alignments used for phylogenetic inference have been deposited in dryad
in order to assess the transmission pattern of the microbiome , dung beetles needed to be both surface sterilized and reared aseptically . beetle offspring were surface sterilized by washing with dilute bleach and detergent followed by two water rinses . no 16s rrna pcr amplicons were observed for dna extracted from the rinse water from surface sterilized beetles ( figure s1 , lane rw ) . under the same conditions , dna extracted from bactrocera oleae , the olive fruit fly , which is known to have bacterial symbionts that amplify with the primer pair 10f - 1507r [ 21 , 62 ] yielded the ~ 1500 bp amplicon and served as a positive control ( figure s1 , lane + ) . the negative control with sterile water as the template did not produce an amplicon ( figure s1 , lane - ) . this experiment was repeated two additional times with the same results .
to ensure that bacteria were not living or that exogenous dna could not be amplified from the dung , dna was extracted from both frozen and autoclaved dung . the ~ 1500 - bp 16s rrna amplicon could not be obtained from either the frozen or autoclaved dung ( figure s1 , lane fd and ad ) . when the bactrocera oleae positive control was added to these same negative control samples , a 1490 bp product was formed demonstrating that the lack of amplification was not due to the differential presence of inhibitors of the pcr reaction ( figure s1 , lane ad + and ad + + ) . no amplicons were formed from the dna of the autoclaved water or autoclaved soil ( figure s1 , lanes aw and as ) . amplicons were produced in fresh , unautoclaved , unfrozen dung ( figure s1 , lane ug ) and the dna extracted from the homogenized digestive system of the 3 rd larval instar ( figure s1 , lane gut ) . this experiment was repeated two additional times with the same results .
otus were assigned using the clovr pipeline with a 95 % threshold as described in the methods for the 275 16s rrna sequences obtained . predicted chimeric sequences ( n = 4 ) as well as sequences shorter than 470 bp ( n = 6 ) were discarded . from the 265 remaining sequences , 71 otus were identified across 19 individuals . of those 71 otus , 49 otus were classified to the family level , 4 to the phylum level , and 4 could not be classified further than “root” ( figures 3 , 4 , and 5 ) . the most abundant genus found across all samples was enterobacter ( figures 3 , 4 , and 5 ) .
the relative abundance of each taxon is shown for each sample . enterobacteriaceae ( bright blue ) were found in 95 % of the samples , in all populations , and all life stages . other major components include bacteria in the comamonadaceae ( green ) and pseudomonadaceae ( bluish purple ) . sample names are abbreviated as follows : 1l ( 1 st instar larva ) , 2l ( 2 nd instar larva ) , 3l ( 3 rd instar larva ) , pu ( pupa ) , ci ( cultured isolates ) , fp ( female parent ) , in ( indiana population ) , and nc ( north carolina population ) . the number after the life stage indicates different individuals sampled .
bacterial otus identified in qiime are arranged in a bulls - eye configuration with the most commonly occurring group in the yellow center , those occurring in two or more individuals in the blue circle , and those genera occurring only one time in the outermost rim . the lowest common ancestor qiime classification is given and may be at the genus , family , or class level and as such may overlap . enterobacter was found in offspring of all parental females and in 68 % of the individuals . samples denoted with a star are those otus that were found using both culture - dependent and - independent techniques .
an unrooted phylogenomic tree based on 287 loci from 1095 sequenced bacterial genomes was made as a skeletal tree on which was placed the lepa and gyrb sequences from 16 bacterial cultures isolated from the digestive system of four different 3 rd instar larvae from female parent 2 from the north carolina population . this analysis placed the isolates sister to ( a ) providencia stuartii , ( b ) enterobacter cloacae , ( c ) pusillimonas , ( d ) pedobacter heparinus , and ( e ) lysinibacillus sphaericus .
several offspring and their female parents have bacteria with 100 % identical 16s rrna sequencing , suggesting vertical transmission through the brood ball and its contents ( figure s3 and table s3 ) . for example in the north carolina samples , one sequence was recovered five times from the female parent , four times from the 2 nd instar larva , and five times from the 3 rd instar larvae ( table s3 ) . additionally , otu’s classified as alcaligenes and pasteurella are similar between one of the female parents and the cultured isolates from a different female parent . deeper sequencing of individuals may reveal more shared bacterial otus .
rarefaction curves of the microbial community in several larvae and the in parental female beetle suggests that additional sampling was needed to identify additional taxa in the microbiome ( figure s4 ) . the rarefaction curves of the nc parental female beetle ( 35 sequenced clones ) and of a second larval instar ( 20 sequenced clones ) suggests that microbiome taxonomic sampling to be close to saturation ( figure s4 ) . the highest bacterial taxonomic richness was isolated from the larval samples and the in parental female , and lower richness was sampled in pupae and in the nc parental female . however , different primer sets and deeper sampling may amplify a wider diversity of bacterial taxa .
these initial results suggest that deeper sequencing of individuals may reveal more shared bacterial otus . the primers used are biased for amplifying γ - proteobacteria , thus additional taxa may be identified using different primers . lastly , some bacteria from the dung also may have a role in the microbiome of beetles in the wild . while , it is not a subject of this study such studies may be fruitful .
collectively , the sterile rearing conditions , overlap of the microbiome between female parent and offspring , specialized dung processing behavior , and bacterial colonies found in the matrix on the brood ball chamber walls , but absent from other locations in the brood ball , suggest that the o . taurus microbiome is maternally transmitted via the brood ball .
the transmission from mother through the brood ball to offspring may be essential for provisioning specific beetle endosymbionts to the offspring since dung beetles develop in an environment rich with other microbes that were excreted from the digestive tract of the mammalian host , as well as bacteria from the soil . thus , the brood ball provides offspring not only with a safe refuge and food , but it is likely that the female parent additionally provisions a microbiome alongside the egg . however , future experimental validation is needed to further test this hypothesis .
vertical transmission via the brood ball may favor organisms that are culturable on standard microbiological media , as the bacteria must persist in an external environment prior to ingestion by the larvae . our analyses based on separate loci for the cloning versus culture - based assays reveals shared bacterial taxa , suggesting a portion of the microbiome is culturable . this is in contrast to relative unculturability of the majority of vertically transmitted endosymbionts that have been studied thus far in insects .
our data suggest that the dung beetle , o . taurus , has endosymbionts that are vertically transmitted in a unique method – via a brood ball . several of the endosymbionts are culturable . due to the manipulability of the system , the dung beetle symbiosis may be an ideal system to understand how host and microbes co - operate , and perhaps co - evolve , to thrive on nutritionally unbalanced diets . additionally , these data suggest that the essential ecosystem function of dung beetles , the degradation of cellulose - rich dung , may be due to communities of bacterial endosymbionts . whether other dung feeding insects have a specific a vertically transmitted microbiome warrants future studies .
a more detailed version of figure 6 that includes the species or strain names of the taxa used to build the tree .
dendrogram illustrating sequence identity between 16s rrna sequences relative to the samples . abbreviations are the same as in figure 4 .
rarefaction curves for selected individuals where the frequency with which diversity is calculated every five sequences .
accession numbers and ncbi taxonomy of the bacterial genomes used for the phylotyping analysis .
sequence identity matrix for 16s rrna samples . cells labeled id are where the sequence is compared to itself . cells labeled na were those where the sequence was not long enough for comparison .
conceived and designed the experiments : ame ecsr djh apm . performed the experiments : ame ecsr mf kf ta djh . analyzed the data : ame djh jcdh . contributed reagents / materials / analysis tools : ame jcdh apm . wrote the manuscript : ame djh jcdh . developed pipeline for phylotyping analysess : djh . collected females from the field : apm ecsr . provided beetle sterile rearing facilities and personnel : apm .
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