flabellina verrucosa , is a species of sea slug , an aeolid nudibranch , a marine gastropod mollusc in the family flabellinidae .

it is found on either side of the northern part of the atlantic ocean .

flabellina verrucosa

worms - world register of marine species - flabellina verrucosa ( m . sars , 1829 )
dear bill here are some photos of flabellina verrucosa from our recent trip to british columbia .
status in world register of marine species synonym of flabellina verrucosa ( m . sars , 1829 )
also one should be aware of bernard picton ' s flabellina browni , which really is similar to flabellina verrucosa with long and slender cerata , but which is differentiated on the coloration .
the relatively short spindle - shaped cerata , and their arrangement into ' clumps ' seems to be characteristic of flabellina verrucosa .
you will see that kåre ' s photos very much match the original descriptions of flabellina verrucosa ; except the arrangement of the cerata which more resembles odhner ' s flabellina borealis . but kåre ' s photos are flabellina verrucosa with short cerata . the problem is that flabellina verrucosa has a long list of synonyms , which include individuals with long and slender cerata , and in most cases it is the animals with long cerata that are illustrated on the net ( as in coryphella rufibranchialis ) .
hans - martin braun added the german common name
rotrückige fadenschnecke
to
flabellina verrucosa ( m . sars , 1829 )
.
scientific synonyms and common names eolidia verrucosa m . sars , 1829 eolis rufibranchialis johnston , 1832 eolidia embletoni johnston , 1835 eolis diversa couthouy , 1839 eolis mananensis stimpson , 1854 aeolis bostoniensis approximans mörch , 1857 coryphella robusta trinchese , 1874 coryphella rufibranchialis cocholata balch , 1908 eolis landsburgi sensu auct . non alder & hancock , 1846 coryphella gracilis bostoniensis ; lemche , 1936 coryphella verrucosa rufibranchialis ; odhner , 1939 coryphella verrucosa verrucosa ; odhner , 1939 coryphella verrucosa ( sars m . , 1829 ) : picton , 1995 flabellina verrucosa
of this one we can be fairly certain - flabellina verrucosa and its eggs on the well grazed bushy pink - mouthed hydroids ( ectopleura crocea ) .
the flabellina verrucosa were very abundant and were found feeding on tubularia up in the shallower water less than 40 feet deep . take care , bruce wight
concerning the correspondence about flabellina verrucosa with short cerata . these photos show the short cerata , with clear gaps in between . i am puzzled by this appearence - is it really a flabellina verrucosa ? i am bound to think otherwise . in particular look at the white lines on the rhinopores , and the length of them .
dear paul , terry gosliner has identified these as flabellina verrucosa . in the bottom right photo i have included an inset of the hydroid i presume they are looking for .
dear alan , as you can see in the messages from terry gosliner and chad sisson , your animals are flabellina verrucosa with a very interesting diet . best wishes , bill rudman
hi bill , i ' ve attached a picture of flabellina verrucosa showing the tail , which i find very distinctive , but isn ' t shown clearly in the pictures already on the forum .
i am pretty certain that the specimens that betsey photographed and that you tentatively identified as flabellina verrucosa is without a doubt correctly identified as f . verrucosa . the main external characteristics for differentiating f . verrucosa from flabellina gracilis is that the cerata are more distinctly arranged in rows in f . gracilis and there is a more prominent vestige of a notal brim in f . gracilis . also f . gracilis has a sort of notched head between the junction of the oral tentacles and more elongate anterior margins of the foot ' .
more feeding flabellina verrucosa . this time chomping down on the giant pink - mouth hydroid , tubularia indivisa . one specimen about to feed . the other image shows a specimen having completed its grazing on this hydroid .
so , what have we learned from these studies ? what features of a nematocyst are important in its selection by a nudibranch , specifically , by flabellina verrucosa ? consider these answers , then click here for explanations .
hi bill , here ' s a flabellina verrucosa balancing on a pink - mouth hydroid stalk . i ' ve assumed they ate the orange ones in my earlier photos , but this may show they like both varieties !
hi bill , here is another common british columbia aeolid , flabellina verrucosa . the dive site is dodd narrows , near nanaimo , on vancouver island . they are often found on this electric orange hydroid , garveia annulata .
thanks terry , it would be great to get some more photos showing the variation in f . verrucosa . bill rudman .
thanks bernard , very soon i won ' t have any excuse to misidentify any north atlantic flabellina . best wishes , bill rudman
greenwood & mariscal 1984 mar biol 80 : 35 ; drawing from kälker & schmekel 1976 zoomorph 86 : 41 for flabellina trilineata .
when i initially saw it i thought it was just a variation of f . verrucosa but wasn ' t sure . dr . larry harris , university of new hampshire has identified it as flabellina verrucosa , although a bit emaciated . he notes that the gonads are not fully developed . andy martinez , author of marinlife of the north atlantic concurs with dr . harris ' id .
more recent studies on nematocyst complement in flabellina verrucosa in the gulf of maine show that while it is a generalist predator of hydroids , it also eats jellyfish scyphistomae . about 70 % of flabellina ’s nematocysts come from the hydroids tubularia crocea and eudendrium spp . most of these are of a type known as heterotrichous microbasic euryteles , which are predominant in these species ( see drawing on left ) .
dear bill , alan shepard ' s photos look to me like pale specimens of flabellina verrucosa . this species is highly variable , especially in the color of the digestive gland that is visible through the cerata . it can range from pale pink to red to chocolate brown . the opaque white markings on the oral tentacles and the arrangement of the cerata appear to be characteristic of f . verrucosa .
hi bill , here is flabellina verrucosa from british columbia , canada . it is also present in north atlantic waters , i ' ve been told . it is very common on vancouver island . this is a fairly small animal , 1 to 3 cm . usually . marli .
dear bill , i checked with my advisor , larry harris , and he agreed that alan shepard ' s aeolids are flabellina verrucosa feeding on scyphistomae . note the white color variation and bright - white cnidosacs . also , they have unusually swollen anterior foot tentacles , probably from being stung by some potent nematocysts . we have documented f . verrucosa with a similar morphology and diet on floating docks in beverly , ma .
on this same dive i found placida dendritica ( first time in the cove ) , flabellina verrucosa , aeolidia papillosa , polycera dubia , acanthodoris pilosa and onchidoris muricata . not bad on a short 40 minute dive in 40f water . best wishes , alan shepard tolland , ct , usa
dear marli , animals such as flabellina verrucosa , which are found in the northern waters of both the pacific and atlantic oceans are called circum - boreal . you ' ll see photos on the forum of this species from the atlantic coast of usa . best wishes , bill rudman .
hi bill , further to your discussion on long and short cerata on flabellina verrucosa : i had never seen the short variety , when shortly thereafter found a small one whilst diving in howe sound near vancouver , british columbia . i have also never seen one in such an intense shade of crimson .
( of eolidia verrucosa sars m . , 1829 ) check list of european marine mollusca ( clemam ) . , available online at urltoken [ details ]
hi bill , here are some flabellina verrucosa from the broughton archipelago . they are eating tubularia crocea . these are the ' long cerata ' type . location : broughton archipelago , british columbia , canada . depth : 45 feet . , length : 10 - 12mm . , may 2003 . photos : marli wakeling
dear paul , this is flabellina verrucosa , about which we ' ve already said most things . this individual shows two of the features which i think are diagnostic for this species : the white line along the tail extending amongst the last few cerata and the narrow white line on the dorsal side of the oral tentacles .
if this is coryphella rufibranchialis then it is more correctly called flabellina verrucosa . there is a history of confusion in the names of this genus on both sides of the atlantic . i ' ll put up some comments when we get them definitively identified . i don ' t want to perpetuate the confusion by further misidentifications .
i may be wrong , but it doesn ' t look like f . verrucosa to me . hopefully some of our northern european colleagues will have some suggestions .
to accompany my message about salmon - gill flabellina from new england here are photos of the ones which are red or chocolate brown and are locally called red - gills .
dear betsey , thanks for the photos from maine . can you give me a little more detail on the locality please ? the brown colour form is a colour form of flabellina verrucosa . the colour in aeolids is often a result of the colour of their food which accumulates in the ducts of the digestive gland , which are in the cerata .
alan , thank you for the excellent photos , this is a relatively rare feeding type of f . verrucosa . all the best , chad sisson university of new hampshire zoology dept .
c . verrucosa has been reported on all depths down to 300 meters . sand or rocky substrate on current exposed locations seems to be preferred . if is usually encountered grazing on hydroids .
i ' m reasonably sure these are flabellina verrucosa , but would appreciate confirmation . they are fairly profuse at harper ' s ferry , a popular dive site on the west side of puget sound near port orchard . most are an inch long at most , so getting a good macro photo is a good challenge . the photos are of two different ones that we found on successive dives at this site .
hi , can anyone help id which flabellina these may be ? found at 40 - 45 feet in hakai pass - port hardy area , bc , canada . are they just variations on the same theme ?
flabellina iodinea showing brightly coloured ( to our eyes ) cerata with tips containing nematocyst - filled sacs . next to nothing is known about the how the sacs function in defense in these or other aeolid nudibranchs 3x
comparison from terry gosliner message below : ' the main external characteristics for differentiating f . verrucosa from flabellina gracilis is that the cerata are more distinctly arranged in rows in f . gracilis and there is a more prominent vestige of a notal brim in f . gracilis . also f . gracilis has a sort of notched head between the junction of the oral tentacles and more elongate anterior margins of the foot ' .
hi ! i found several individuals of this aeolid , looking a little like flabellina verrucosa , which is quite common along the norwegian coast . however , some details seem to differ : • shorter cerata , not organized in the traditional two rows . • a naked patch on the back . • some eggs found on the location , laid in circles . ( i am assuming they were laid by these individuals ) .
dear wendy , i am pretty sure that the upper right and lower left photos are of flabellina verrucosa , the white - tipped cerata , the white line on the oral tentacles and the white line down the ' tail ' all suggest that species . i am not so sure of the third photo [ lower right ] however and so if someone with local knowledge can comment i would be grateful . best wishes , bill rudman
dear bill , in response to kåre ' s recent message , here is a scan of the original illustrations by m . sars ( 1829 ) of eolidia verrucosa [ plate 2 , figs 1 - 4 ] .
although only about 0 . 5 % of flabellina ’s nematocysts come from scyphistomae ( see green dots in table ) , these polyps are selected over any others in choice tests , perhaps because of the toxicity of their nematocysts .
many species of aeolid nudibranchs , including flabellina trilineata shown in the photograph , sequester undischarged nematocysts , apparently for use in their own defense . the nematocysts are housed in special sacs or cnidosacs located in the tips of the cerata .
( of eolidia verrucosa sars m . , 1829 ) sars , m . ( 1829 ) . bidrag til söedyrenes naturhistorie . chr . dahl , bergen . 1 : 1 - 60 , plates 1 - 6 . , available online at urltoken page ( s ) : 9 [ details ]
of the red - gills there are elongated ones with a few growths on their backs [ lower left photo ] as well as the more common ones with completely covered backs [ other photos ] . we think these are f . verrucosa , f . pellucida , or f . gracilis .
references flabellina verrucosa ( m . sars , 1829 ) . in sea slug forum . australian museum , sydney . accessed 26 / 01 / 2016 . lamb , a . , and hanby , b . ( 2005 ) . marine life of the pacific northwest [ electronic version ] . madeira park , bc : harbour publishing . picton , b . e . & morrow , c . c . ( 2015 ) . coryphella rufibranchialis ( johnston , 1832 ) . encyclopedia of marine life of britain and ireland . accessed 26 / 01 / 2016 . authors and editors of page kelly fretwell ( 2016 ) .
these individuals were all photographed as a part in my project to elucidate the two forms ( long ceratal and short ceratal form ) relationship . the animals with long cerata were photographed at three meters depth at jordfall . the short ceratal form ( verrucosa form ) was photographed at yttre vattenholmen outside strömstad .
( of aeolidia verrucosa m . sars , 1829 ) 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 ]
dear alan , it certainly looks like a crustacean of some sort . certainly some glaucids , such as a austraeolis ornata and hermissenda crassicornis are known to eat the odd bit of carrion , but i haven ' t heard of species of flabellina doing this . it certainly seems an interesting observations for someone to follow up . best wishes , bill rudman
i just got a message from bernard picton who suggested the scyphistoma in chad ' s photo is probably aurelia aurita so i had another search of mcdonald & nybakken ' s list and found the following references to feeding on aurelia aurita ' polyps ' . clearly predation by f . verrucosa on the scyphistomae stage of aurelai can be an important factor in aurelia populations , at least in sweden .
note the conspicuous size difference between predator and prey . the left photo - pair shows aversive behaviour to f . iodinea . the right photo - pair shows the same predator 20min later attacking and eating another nematocyst - defended aeolid , hermissenda crassicornis . not all naive predators show the same aversive responses to flabellina ; about half of the 40 - or so pleurobranchaea tested in the study simply attack and eat up the prey
on the lower left photo the inset is showing the white sreak on the rhinophore which is one of the external identification points of this species . the colour and shape of the head and head tentacles are important in identifying species of flabellina but as your photos show , when these animals are busy looking for food they don ' t make it easy for you to photograph their important features . best wishes , bill rudman .
to barcode of life ( 36 barcodes ) to biodiversity heritage library ( 13 publications ) to biodiversity heritage library ( 14 publications ) ( from synonym eolidia verrucosa sars m . , 1829 ) to biodiversity heritage library ( 44 publications ) ( from synonym coryphella rufibranchialis ( johnston , 1832 ) ) to clemam ( from synonym eolis mananensis stimpson , 1853 ) to clemam ( from synonym coryphella rufibranchialis cocholata balch , 1908 ) to clemam ( from synonym coryphella rufibranchialis ( johnston , 1832 ) ) to clemam ( from synonym eolis diversa couthouy , 1839 ) to clemam ( from synonym eolidia verrucosa sars m . , 1829 ) to clemam ( from synonym eolidia embletoni johnston , 1835 ) to clemam ( from synonym coryphella robusta trinchese , 1874 ) to clemam to encyclopedia of life to genbank ( 29 nucleotides ; 20 proteins ) to pesi ( from synonym coryphella robusta trinchese , 1874 ) to pesi to pesi ( from synonym eolidia embletoni johnston , 1835 ) to pesi ( from synonym eolis diversa couthouy , 1839 ) to pesi ( from synonym eolidia verrucosa sars m . , 1829 ) to pesi ( from synonym coryphella rufibranchialis cocholata balch , 1908 ) to pesi ( from synonym eolis mananensis stimpson , 1853 ) to pesi ( from synonym coryphella rufibranchialis ( johnston , 1832 ) ) to sea slug forum ( via archive . org ) to itis
in response to your question : i do not know of any research done involving this specific diet for f . verrucosa . i simply meant that these nudibranchs are rarely found associated with this food , but when they are , they have a very distinct color and morphology . i do not know if they switch to another diet once the scyphistomae have run out , but it wouldn ' t surprise me if they did take advantage of whatever is around .
your photos are useful reminder to me of how difficult it is for non - professionals to identify species they find or photograph . my first act when i find something i don ' t recognise is to dissect it and look at the radula to confirm what family or genus is belongs to , because the outside can sometimes be very misleading . i have just posted a page of photos of the radula of flabellina poenicia which illustrates a tooth shape which is very characteristic of the genus . looking at your three species i am not even sure they are all flabellinids . best wishes , bill rudman .
• hernroth , lars , & fredrik gröndahl . 1985 . the biology of aurelia aurita ( l . ) 3 . predation by coryphella verrucosa ( gastropoda , opisthobranchia ) , a major factor regulating the development of aurelia populations in the gullmar fjord , western sweden . ophelia , 24 ( 1 ) : 37 - 45 . • hernroth , l . , & f . gröndahl . 1985a . on the biology of aurelia aurita ( l . ) : 2 . major factors regulating the occurrence of ephyrae and young medusae in the gullmar fjord , western sweden . bulletin of marine science , 37 ( 2 ) : 567 - 576 .
description : currently considered to be a form of coryphella verrucosa . the body is translucent white in colour with opaque white pigment on the tips of the oral tentacles and in a broken line along the centre of the back which becomes continuous on the long tail . the cerata are numerous and arranged in clusters along the sides and back of the body , almost hiding the white centreline . they are filled with red or orange - red granular digestive gland . the cerata have a thin , sometimes incomplete , ring of white pigment below the tip . typically about 15mm - 25mm in length , but well - fed individuals may be larger .
dear kåre , this is an interesting find . you wouldn ' t by chance have have a photo showing the front of the foot more clearly ? in the smaller animal in the upper photo it looks like the front edge of the foot could have the front corners extended into pointed ' angles ' , but i am not sure . i first thought this might be a species of cuthona , but in that genus the anterior foot corners are rounded . the egg ribbon , if it does belong to these animals , could certainly be that of a flabellina . in your lower right photo the animal is eating what seems to be a large solitary hydroid - perhaps a tubularia .
the closest i can come to an identification is a painting of henning lemche ' s of an animal from jutland , denmark ( just & edmunds , 1985 ) , which the authors , suggest is an aberrant form of f . verrucosa . like your animals , the cerata are short and inflated , and are arranged in a pattern of triangles down the body with bare patches down the dorsal midline . even the colour is almost identical . unfortunately , just & edmunds only had lemche ' s painting , no accompanying notes could be found , and no anatomical information . they suggested it was probably ' unhealthy when painted ' which certainly doesn ' t seem to apply to your animals .
what would be interesting would be to see if they have any natural history differences . some species of flabellina change from feeding on one group of hydroids to another during the year so food differences may not be a simple thing to look at . another observation that might be useful would be to look at their egg masses . it is quite possible that their egg spirals might be of a different shape or even their eggs could be markedly different in size . if you would like a little project it would certainly be worth trying to photograph the egg ribbons of the two ' forms ' ideally as they are laying them . there may be no marked difference , but it would be exciting if there were , best wishes , bill rudman
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confusing ? it really is , and i think this group is in need of a careful revision .
• sars , michael . 1829 . bidrag til soedyrene naturhistorie , pt . 1 , : pp . 1 - 59 , pls . 1 - 6 . bergen .
one anatomical point which seems worth following up is the shape of the anterior foot corners . in both sars ' painting and kåre ' s photo the anterior corners of the foot are rounded . although none of the photos in the forum of the long cerata form show the anterior foot clearly , there are plenty of illustrations in the literature showing that it has distinctly angular or even tentacular foot corners [ thompson & brown , 1984 ] . to me the ceratal shape and arrangement and the anterior foot corners are pretty good arguments for some separating them .
• kuzirian , a . m . ( 1979 ) taxonomy and biology of four new england coryphellid nudibranchs . journal of molluscan studies , 45 : 239 - 261 .
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mate selection & copulation are dealt with in this section , while topics of egg - laying , embryonic development , hatching & larval life , settlement & metamorphosis , and ontogenetic development of behaviour are considered elsewhere .
this section on mate - selection & copulation is arranged alphabetically by genus . the genus hermissenda is considered here , while aeolidia , alderia , navanax & other cephalaspids , and aplysia & other anaspids are considered in their own sections .
done in oregon and california later are thought to be most likely accounts of mating . the reason for the misunderstanding owes in part to the extraordinary quickness of the actual copulatory act , which requires only 1 - 2sec . the author never observes copulation in
, in part because of a possible expectation of an activity that would last several minutes or even hours as is customary in other opisthobranchs . the complexity of interactions involved in one of the studies is shown in this computer - simulated flow
. be mindful as you go through it that the researcher is describing general behaviour in laboratory specimens with the understanding that it relates to agonism . with a bit of patience , and assuming that what is really being described is not agonism , but copulatory behaviour , the behavioural repertoire during mating can be noodled out to a few basic components . the line thicknesses in the diagram correspond with frequency of occurrence of any successive pair of behaviours .
begins with a touch of one individual by another , most commonly with tentacles or cerata . since
- ing , which is right - side to right - side orientation that places the gonopore openings of the 2 individuals opposite one another .
, which we know is preparatory to copulation . however , because copulation is not in the author ' s mind , we have to choose a spot for it to happen . a possible spot for this is marked with a bright yellow blob on the flow diagram , its
- all familiar from research study 1 above . the author observes damage done by biting ( the jaws have serrated edges ) , describes “winners” and “losers” in the encounters , and notes that 26 % of the “winners” , presumably the ones that break away first , are the larger individuals of the pairs ( thus , 74 % of
winners
are smaller individuals ) . there are other behaviours shown in the diagram that may or may not play a role in copulation .
recall that the author believes the behaviour to be agonistic ; hence , the
physical
nature of this term . tentacle - touching is described by other authors as more of a
caressing
this is only one possible route from
sidling
that could lead to insemination . another shorter one would be
occurs with low frequency . another , more round - about route on the chart following from an hypothetical copulation after
, and this route also has the merit of large - dimension arrows . the reason for speculating on possible post -
) , and more withdrawals . this goes on for some time and the video switches to show full right side - to - right side touching (
sidling
) and perhaps even copulation . it is
how the author of research study 1 above could initially interpret these behaviours as agonism .
however , anthropomorphism is all too common in interpretation of animal behaviour . in this case , the terms
flagellate
,
sidle
,
winners
, and
losers
, are highly suggestive . is it possible that their initial selection may have prevented the author from standing back and viewing the behaviours more objectively ?
no other nudibranch species appears to exhibit it , nor is it a feature known to be particularly common in gastropods . the author remarks on never having seen agonistic behaviour in
in the field . as to the function of the putative agonistic behaviour , the author speculates that it may aid in food - getting and be a means of warding off predators . in another publication on the same subject the author tests what effect food deprivation has on the “agonistic” behaviour ? in these latter experiments , after 1 - 3d without food there is an increase in locomotory activity and an increase in
agonistic
interactions . also , with each progressive day the frequency of encounters involving biting goes up , suggesting to the author that hunger may be the cause of the “fighting”
a common behaviour in nudibranchs when food is scarce or lacking , however , is to copulate . this makes sense because most species of nudibranchs live seasonally and , when their prey items are in short supply , they spend more time copulating . . . not fighting
although the study may not be as relevant to “agonistic” behaviour as the author intended , the technique used may be useful for other types of experiments . in the present experiments , the author pinches off all cerata from several
with forceps , provides clam tissues as food , and waits until the cerata are regenerated . at this time the cnidosacs
of nematocysts . the author then compares the behaviour of these “nematocyst - free” animals with that of normal control animals and finds no difference in level of “agonistic” behaviour . the technique , although not working out in the way that the author intended could , nevertheless , be a useful way to produce nematocyst - free aeolids for other types of experiments , for example ,
to assess the defensive efficacy of the secondarily acquired nematocysts against fishes and other biting predators .
this is not a new idea , but appears not to have been done with west - coast nudibranchs .
the author is not able to check whether the cnidosacs were actually nematocyst - free , as all the animals die before this could be done . healthy
, a second author agrees with the first that the species does engage in aggressive interactions with conspecifics . however , what is missing in the first descriptions , acording to the second author , is information on the temporal aspects of the interaction , specifically the timing of events from first contact to onset of sidling , and then to cerata movement , lunging and biting , withdrawing , and ending . these events are observed in specimens of
from elkhorn slough , california and recorded on videotape for time - motion analysis . note in the data
that the mean duration of these activities is only about 80 seconds . the author comments that “although the animals are poised in copulation - like posture during sidling , it seems clear , as zack suggested , that these interactions are not copulatory since the time between initiation of sidling and cerata movement is on the average less than 10 seconds” . in discussing the results , the author provides 2 possible explanations for flagellation and sidling : 1 ) to assess the aggressive motivation and size of the other individual , or 2 ) to assess the willingness of the other individual to mate . the author concludes by suggesting that more observations are needed to clarify the matter .
the diagram shows the durations of 4 sets of activities from first contact : begin sidling , move cerata , withdraw , and end . each horizontal set includes overall range , standard deviation , and mean for each set of activities . the vertical blue dotted line shows the mean end time in seconds for 19 individual nudibranchs
, and that zack ' s original paper ( research studies 1 & 3 ) makes it clear that there may be other possible outcomes , including aggression .
because the 2 papers ( research studies 4 & 6 ) were submitted for review at the same time for publication in the same volume of veliger , out of courtesy and interest the editor allowed each set of authors to read the other’s submission and to make comments . the set of 4 articles in veliger is unique and makes for interesting reading
in the same journal as the foregoing publications ( research studies 4 & 5 ) is a paper entitled , “ hermissenda : agonistic behavior or mating behavior ? ” in this paper researchers at friday harbor laboratories , washington note that agonism is not seen in the field in h . crassicornis , and describe the following pre - copulatory behaviours in laboratory specimens : 1 ) one animal follows another , 2 ) front individual turns , 3 ) they meet head - to - head , 4 ) make tentacle / tentacle contact , 5 ) withdraw , 5 ) make tentacle / cerata contact , 6 ) withdraw , 7 ) make tentacle / foot contact , 8 ) advance , and 9 ) jockey back - and - forth . time required : 10 - 30min .
the individuals then align their bodies on the right sides ( 1 - 3min ) , evert penises , expand penis ends into bulbs , and simultaneously ejaculate sperm in muscular contractions . the sperm is emitted in mucus . time for copulatory ejaculation : 1 - 2sec . the authors observe no biting damage , but suggest that the function of pre - copulatory contact may be to reduce the probablility of being eaten . this seems to confirm that aggression occurs , sometimes leading to “cannibalism” , but the authors reiterate their belief that agonistic behaviour does not normally ocur in the field in
the authors conclude that zack’s observations are just of mating behaviour , and not anything else .
longley & longley 1981 veliger 24 : 230 ; longley & longley 1981 veliger 24 : 232 .
copulating . video and descrptive headings courtesy roger & alison longley , friday harbor laboratories .
in santa cruz , california provides further detail on copulation and efficacy of sperm transfer . first , the rapidity of the copulatory event ( about 4sec on average ) markedly decreases the chance of complete transfer of sperm . in
37 videotaped sequences of attempted copulation ( defined as 2 individuals with penises everted ) , almost half lead to failed sperm transfer in one or both individuals . interestingly , a frequent behaviour following sloppy sperm transfer is for an individual to twist its head backwards and consume all or part of the semen in the vicinity of the gonopore . if an individual is allowed to become
habitat : a common species in early spring at shallow exposed sites and deeper water which is exposed to tidal streams . the normal food is the hydroid tubularia indivisa but other smaller hydroids are eaten by juvenile specimens . the spawn consists of a thread which is laid in a smooth spiral like a clock spring .
distribution : a northern species in the british isles , occurring round scotland and in the irish sea south to the isle of man , but apparently absent from the western coasts of ireland and england . further distribution includes norway to the atlantic coast of america .
distribution map from nbn : interactive map : national biodiversity network mapping facility , data for uk .
picton , b . e . & morrow , c . c . ( 2016 ) . coryphella rufibranchialis ( johnston , 1832 ) . [ in ] encyclopedia of marine life of britain and ireland . urltoken accessed on 2018 - 07 - 09
discover a faster , simpler path to publishing in a high - quality journal . plos one promises fair , rigorous peer review , broad scope , and wide readership – a perfect fit for your research every time .
the evolution of gastropod body size in the deep sea is reexamined . using an extended and updated data set , and improved statistical methods , it is shown that some results of the previous study may have been artifactual , but that its central conclusion is robust . it is further shown that the effect is not restricted to a single gastropod clade , that its strength increases markedly with depth , but that it applies even in the mesopelagic zone .
the replication of the island rule in a distant taxonomic group and a partially analogous ecological situation could help to uncover the causes of the patterns observed—which are currently much disputed . the gastropod pattern is evident at intermediate depths , and so cannot be attributed to the unique features of abyssal ecology .
citation : welch jj ( 2010 ) the “island rule” and deep - sea gastropods : re - examining the evidence . plos one 5 ( 1 ) : e8776 . urltoken
copyright : © 2010 john j . welch . 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 : some preliminary work undertaken when jw was funded by bbsrc grant do17750 awarded to andrew rambaut . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
here , i re - examine whether deep - sea gastropods manifest the island rule , making use of the improved statistical methods , and data collated from the recently updated malacolog database [ 24 ] , which has been both expanded , and revised to reflect advances in gastropod systematics [ 25 ] . it is found that the central conclusion of mcclain et al . [ 12 ] is robust , and that gastropod colonists of the deep - sea benthos do indeed exhibit island - rule - like evolution .
part a shows how different tests of the ‘island rule’ can give qualitatively different results . “deep - sea” species were defined as those with a depth range midpoint > 200m , and all other species defined as “shallow - water” . the ordinary - least - squares regression ( dashed line ) differs significantly from the 1∶1 line of the null ( dotted line ) , but the standardized - major - axis regression ( solid line ) shows no significant departure . part b shows a less ambiguous case : “deep - sea” species are those never observed above 400m , and “shallow - water” species those never observed below 200m ; body sizes are within - genus means , taking equal numbers of deep - and shallow - water species in each genus .
the body sizes of deep - sea gastropods are plotted against those of their shallow - water congeners . “shallow - water” species were never observed below 200m , and “deep - sea” species never observed above depths of a : 200m , b : 400m and c : 600m . separate standardized - major - axis regression lines are shown for the neogastropoda ( black points ) and all other groups ( grey points ) . the dotted line is the 1∶1 expected under the null . genera with fewer than two deep and two shallow species were excluded .
we are therefore still far from understanding the causes of the patterns observed – and particularly the roles of inter - and intra - specific competition [ 3 ] , [ 4 ] , [ 11 ] , [ 12 ] . a detailed clarification of where the pattern does and does not hold will be an important step toward achieving this goal [ 4 ] , [ 12 ] , [ 19 ] , [ 20 ] .
many thanks are due to andrew rambaut for providing a script to mine the malacolog database . many thanks also to lucy weinert , nicolas bierne , gary rosenberg , shai meiri . simon joly and an anonymous reviewer , who all greatly improved the manuscript with their comments and advice .
conceived and designed the experiments : jw . performed the experiments : jw . analyzed the data : jw . wrote the paper : jw .
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( of coryphella robusta trinchese , 1874 ) trinchese s . ( 1874 ) . intorno ai generi hermaeina e acanthopsole . memorie dell ' accademia delle scienze dell ' istituto di bologna ( 3 ) 5 : 73 - 80 pl . 1 [ details ]
( of eolidia embletoni johnston , 1835 ) johnston g . ( 1835 ) . introduction to the natural history of molluscous animals . magazine of natural history 8 : 71 - 81 [ details ]
( of eolis diversa couthouy , 1839 ) couthouy j . p . ( 1839 ) . monograph on the family osteodesmacea of deshayes , with remarks on two species of patelloidea , and descriptions of new species of marine shells , a species of anculotus , and one of eolis . boston journal of natural history , 2 : 129 - 189 , pl . 4 . , available online at urltoken page ( s ) : 187 - 189 ; pl . 4 , fig . 9 [ sic , 14 ] [ details ]
( of eolis mananensis stimpson , 1853 ) stimpson , w . ( 1853 ) . synopsis of the marine invertebrata of grand manan : or the region about the mouth of the bay of fundy , new brunswick . smithsonian contributions to knowledge . 6 : 1 - 66 , pls 1 - 3 . , available online at urltoken [ details ]
howson , c . m . ; picton , b . e . ( 1997 ) . the species directory of the marine fauna and flora of the british isles and surrounding seas . ulster museum publication , 276 . the ulster museum : belfast , uk . isbn 0 - 948150 - 06 - 8 . vi , 508 ( + cd - rom ) pp . ( look up in imis ) [ details ] available for editors [ request ]
turgeon , d . ; quinn , j . f . ; bogan , a . e . ; coan , e . v . ; hochberg , f . g . ; lyons , w . g . ; mikkelsen , p . m . ; neves , r . j . ; roper , c . f . e . ; rosenberg , g . ; roth , b . ; scheltema , a . ; thompson , f . g . ; vecchione , m . ; williams , j . d . ( 1998 ) . common and scientific names of aquatic invertebrates from the united states and canada : mollusks . 2nd ed . american fisheries society special publication , 26 . american fisheries society : bethesda , md ( usa ) . isbn 1 - 888569 - 01 - 8 . ix , 526 + cd - rom pp . ( look up in imis ) [ details ]
linkletter , l . e . ( 1977 ) . a checklist of marine fauna and flora of the bay of fundy . huntsman marine laboratory , st . andrews , n . b . 68 : p . [ details ]
pollock , l . w . ( 1998 ) . a practical guide to the marine animals of northeastern north america . rutgers university press . new brunswick , new jersey & london . 367 pp . , available online at urltoken [ details ]
rosenberg , g . 2004 . malacolog version 3 . 3 . 2 : western atlantic gastropod database . the academy of natural sciences , philadelphia , pa . , available online at urltoken [ details ]
abbott r . t . ( 1974 ) . american seashells . the marine mollusca of the atlantic and pacific coast of north america . ed . 2 . van nostrand , new york . 663 pp . , 24 pls . [ october 1974 ] . ( look up in imis ) [ details ]
trott , t . j . ( 2004 ) . cobscook bay inventory : a historical checklist of marine invertebrates spanning 162 years . northeastern naturalist . 11 , 261 - 324 . , available online at urltoken [ details ] available for editors [ request ]
integrated taxonomic information system ( itis ) . , available online at urltoken [ details ]
check list of european marine mollusca ( clemam ) . , available online at urltoken [ details ]
( of eolidia embletoni johnston , 1835 ) check list of european marine mollusca ( clemam ) . , available online at urltoken [ details ]
( of eolis diversa couthouy , 1839 ) check list of european marine mollusca ( clemam ) . , available online at urltoken [ details ]
( of coryphella robusta trinchese , 1874 ) check list of european marine mollusca ( clemam ) . , available online at urltoken [ details ]
( of coryphella rufibranchialis ( johnston , 1832 ) ) check list of european marine mollusca ( clemam ) . , available online at urltoken [ details ]
( of coryphella rufibranchialis cocholata balch , 1908 ) check list of european marine mollusca ( clemam ) . , available online at urltoken [ details ]
( of eolis mananensis stimpson , 1853 ) check list of european marine mollusca ( clemam ) . , available online at urltoken [ details ]
the translucent white body may reach 6 cm in extreme cases , but 2 - 3 cm is common . often , there is a white line running along the back , but it tends to be covered by the cerata . the cerata are arranged in rows and come in a wide range of colors , but orange , red and brown are perhaps most frequent . there may be a pink tint on the tentacles . the rhinophores ( head tentacles ) are wrinkled and usually have a white pigmented line .
the original identification of the species was done by sars in 1829 . he drew a specimen with short , ice - cone - shaped , white - tipped cerata . studies have later proven that


even stevens ( 1957 – 1975 ) was a thoroughbred racehorse that won both the caulfield and melbourne cups in australia in 1962 .

he was ridden in both cups by his regular rider les coles .

even stevens ( horse )

melbourne cup 1962 national jockeys ' trust even stevens l . coles t . j . smith
even stevens ( nz ) ch . h , 1957 { 12 - f } dp = 14 - 4 - 2 - 0 - 6 ( 26 ) di = 2 . 71 cd = 0 . 77 - ? starts , 8 wins , 2 places , 1 shows career earnings : 43 , 895 pounds
the first anniversary of the settlement of wellington in january 1841 included a hurdle race on the third day of the celebrations . it was won by henry petre riding his own horse , calmuc tartar . a jockey club was formed for the meeting but it lapsed after a few years . the first formal meeting was held at petone beach on 20 october 1842 , when the imported horse figaro beat calmuc tartar in a 10 - guinea sweepstake run in heats over a mile and a half . racing later took place at hutt park and burnham water ( the site of a former miramar lagoon ) . the latter was probably the first racecourse in new zealand and had a grandstand .
the growth of racing in the post - war years was even more marked than in the early 1920s . race permits were not restored immediately the war finished but over the two years following . the conference pressed for more permits , but these were not granted until after the report of the 1946 royal commission on gaming and racing had been considered by parliament and the gaming amendment act passed in 1949 .
it could be expected that , in a colony settled by predominantly british people , horse racing in some form or other would soon begin . horses were a valuable necessity in the colonies . the first horses to be landed in new zealand were probably those brought from australia by the rev . samuel marsden to rangihoua in the bay of islands on 23 december 1814 from the ship active . they were from new south wales , the gift of governor macquarie to the maoris . horses from new south wales were to have an important place in the establishment of thoroughbred breeding in new zealand . there are few records of the very early importations . horses came with the military garrisons and it is recorded that the first horses arrived in wellington on 2 march 1840 . the first acknowledged thoroughbred horse , figaro , landed in wellington . he was bred by t . icely , of cooming , new south wales , a celebrated breeder of the time .
there was a meeting in wanganui on 28 december 1848 and there , too , the officers of the militia had a hand in starting the sport . the first races in dunedin were held on 23 march 1849 as part of the anniversary day celebrations , with the eccentric dr manning a prominent owner . following the gold discoveries in the province , otago was , for a time , the strongest racing centre in new zealand . the celebrations on the first anniversary of the canterbury settlement on 16 december 1851 included four horse races over a course in hagley park facing the road running from the riccarton hotel to the fendalton bridge . the course was still in its native tussock . there was also early racing on the west coast and in taranaki ; but not in hawke ' s bay ( a district later to play an important part in new zealand racing ) until 1 january 1857 .
this information was published in 1966 in an encyclopaedia of new zealand , edited by a . h . mclintock . it has not been corrected and will not be updated .
a hurdle race was run on the first anniversary of nelson “through fern and flax , up hill and down hill” . nelson first placed racing on a sound footing . there was a good course , with thoroughbreds imported for racing and breeding , and horses trained and brought out to race in something like “condition” . the course was at stoke , 4 miles from nelson . it was first used on 3 february 1845 .
the early race meetings in the colony were controlled by local committees elected for the meeting only , generally at a public meeting of interested citizens . those elected made the arrangements , drew up the rules , and appointed the officials . in the larger towns the establishment of a racing club generally followed . these local clubs had their own locally varying rules , but based in common on those of the english jockey club . until the late 1860s each club was a separate identity , with little coordination because of the difficulties of travel and communication . consequently , disqualifications imposed by one club would not apply at another . the first attempts to introduce some form of unified control were made by the metropolitan clubs , a rather grandiose title for the times . it is not clear how certain clubs came to be so designated and to assume a limited control over the racing within their districts . but the metropolitan clubs of the 1860s and early 1870s did correspond with the main towns of the provincial districts . it is possible that there was some direction given from the colonial secretary at wellington , since at that time permits for race meetings were issued by his office , and programmes in each district were approved by the resident magistrate . in early minute books there are instances of the resident magistrate referring programmes back to the metropolitan club because they had not first had that body ' s approval . so probably the sheer need for a responsible body to give guidance on those matters and to settle disputes forced the metropolitan clubs to act as a miniature jockey club .
after racing had been established for 30 years , the metropolitan clubs realised the need for some governing body to obtain uniform rules of racing and a uniform scale of weights . the first recorded move was made by the canterbury jockey club in 1875 and , on 11 november 1876 , during the course of the canterbury jockey club race meetings , a meeting of delegates resolved “that it was desirable to establish a new zealand jockey club , to frame rules and make a scale of weights to be used by all clubs running under the rules” . there was obviously some dissatisfaction at the time , for in 1877 the canterbury jockey club resolved to recognise only the dunedin , wellington , auckland , and hawke ' s bay clubs . this made wanganui and taranaki hostile .
the first truly constructive move came from the hawke ' s bay jockey club which , on 12 july 1883 , decided to set up a subcommittee , consisting of captain w . r . russell ( later to be the first president of the racing conference ) , r . u . burke , and c . b . winter ( the mover of the proposal ) , “to consider the establishment of a new zealand racing association , and the drafting of rules for same , and that the matter be submitted to the clubs already mentioned and the taranaki and wanganui jockey clubs , which were to be the metropolitan clubs for the proposed districts the colony would be divided into” . the proposal also suggested the monthly publication of a racing calendar , the registration of colours , and a turf register .
the only record of this meeting is a letter addressed by the chairman to the colonial secretary . the letter declared that the purpose of the meeting was to help racing as a whole , pointed out certain advantages to the country in using the totalisator confined within its lawful limits , and suggested ways of controlling the totalisator and of encouraging the breeding industry . the colonial secretary ' s reply pointed out to the chairman that legislation would be necessary to give metropolitan clubs a defined and legal status under the gaming and lotteries act before the suggestions could be acted on . the colonial secretary proposed that the delegates ' recommendations could best be carried out by formulating them in a bill , which might then be introduced into either house of the legislature by some member interested in racing . no bill has ever been introduced ; the authority of the racing conference is still not enforced by statute .
from 1887 to 1891 metropolitan club representatives met every year and sometimes twice a year , with some meetings being attended by representatives of the greymouth , nelson , and marlborough clubs . many of the early meetings were held in parliament buildings , and several of those early delegates were members of either the house of representatives or the legislative council . the hon . g . mclean , m . l . c . , chaired the first meeting and , except for the hon . j . d . ormond , the hon . w . r . russell ( who became a member of the house of representatives ) chaired all the other early meetings . f . d . luckie , of hawke ' s bay , became the conference secretary . the hon . e . mitchelson , the hon . o . samuel , dr earle , freeman r . jackson , r . h . nolan , francis henry dillon bell , and , above all , sir george clifford , were prominent in helping the racing conference to gain its high standing as quickly as it did .
in 1897 r . h . nolan succeeded in having adopted his scheme for appeals against decisions of metropolitan clubs ' committees being heard by three appeal judges appointed by the president . this system was acclaimed at the time and has stood the test of the present day . the first appeal was heard in 1898 , the judges being dr earle , geo . hunter , and nolan .
in 1899 it had been decided that country clubs be allowed two representatives on metropolitan committees . the registration of all racing clubs was enforced in 1900 . a most important step for owners was the institution of accident fees in 1903 . an accident fund for trainers , jockeys , and stablehands was established to relieve owners of their liability under the statutory provisions of the employers ' liability and like acts . the jockeys ' and trainers ' provident funds controlled by each metropolitan committee continued , but on a gradually changing basis .
when the first world war broke out , the racing conference was pressed first to stop all racing , and then to reduce the number of race days . following the lead given in england , racing continued , but in 1917 a special committee agreed with the government to reduce race days by a third . the racing clubs were soon active contributors to the various war funds , and a number of courses were taken over for military purposes , notably wellington ( trentham ) , wairarapa , and manawatu .
in 1921 the appointment of racecourse inspectors was agreed to , the first appointments being a . ward , r . g . black , f . cullen , and j . torrance . all were ex - police officers . through the work of the racecourse inspectors ( who cooperate closely with the police ) , the racecourses in new zealand are kept remarkably free of undesirables and prohibited persons . the independent licensing of trainers and jockeys by each metropolitan district committee also ended in 1921 , when a licensing committee was appointed and all licences issued by the racing conference . the southland metropolitan committee was created in 1925 , all clubs in the southland province being separated from the otago district .
from 1898 the affairs of the racing conference had been administered from christchurch . with the growth of racing after the first world war it was decided that the headquarters should be more central . in 1930 they were moved to wellington . the constitution of the conference was altered in 1928 when the executive committee was formed . this first consisted of the president , vice - president , and six representatives , but in 1929 this was changed to include one representative of each metropolitan district . the licensing committee and the dates committee were abolished in 1933 and their duties taken over by the executive committee .
racing fell off during the depression of the 1930s . the racing conference was faced with many difficulties because of the plight of some of the smaller country clubs . some became defunct and their permits were taken up by other clubs . race days were again reduced during the second world war and many racecourses were taken over by the military . the restricted racing and lack of transport raised many problems , as the number of horses in training was not reduced , nor was there any loss of interest in race meetings . these were often held under great difficulties because of the military occupation of the courses being raced on . in the later years of the war the executive committee sanctioned race meetings to raise patriotic funds .
the conference considerably advanced steps in the interests of racing and the national bloodstock breeding industry when it set up its “dope detection” scheme in 1953 . racing expanded quickly in the auckland province ( especially in the waikato district ) with the great increase of population there . this led to the formation of the waikato metropolitan district in 1949 by the division of the auckland metropolitan district , then by far the largest and strongest in new zealand . there have been no changes in the metropolitan districts since . in 1962 the conference for the first time had its own building , in farish street , wellington .
thus racing in new zealand is now controlled by the new zealand racing conference , which , consistent with its origin , is an association of the clubs registered under its rules . the racing conference does not run race meetings , as it is a purely legislative and administrative body . the year - to - year administration is done by an executive committee elected annually and comprising the president and vice - president , ex - officio , and one representative of each of the 10 metropolitan racing districts of new zealand . delegates of racing clubs meet annually in wellington in july , when legislative matters are dealt with by way of remits from the executive committee , district committees , or clubs . the racing conference registers all horses , issues all licences , administers the general trust fund ( accident fund ) , and publishes the new zealand stud book and the new zealand racing calendar . the secretary is the principal executive officer , and a staff of stipendiary stewards and racecourse inspectors attend all race meetings .
in each metropolitan racing district a district committee generally supervises the racing in its district whether the meetings are run by totalisator , non - totalisator , or sports clubs . each district committee comprises one representative of each district totalisator club and an equal number of representatives of the senior club , which is known as the metropolitan club . each district committee must approve all programmes in its district and consider all applications for dates and licences before they are submitted with recommendations to the racing conference . it also hears all appeals against the decisions of racing club judicial committees and reviews all penalties imposed by the latter .
there are 71 racing clubs and 17 hunt clubs authorised to use the totalisator and a total of 259 days allocated to racing . all racing clubs are non - proprietary , and the committee and the stewards of each are elected from the club members . each racing club runs its own affairs and its own race meeting . on race day , the committee or the stewards ( or sometimes both according to the club ' s constitution ) control the meeting , and all judicial matters are dealt with by a judicial committee appointed by the club . this committee must investigate all matters submitted to it by a stipendiary steward or racecourse inspector , neither of whom has any judicial powers . most racing clubs own their own racecourses . hunt clubs do not , and they are not individually represented on a district committee .
how to cite this page : ' history - establishment and administration ' , from an encyclopaedia of new zealand , edited by a . h . mclintock , originally published in 1966 . te ara - the encyclopedia of new zealand url : urltoken ( accessed 10 jul 2018 )
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the new zealand thoroughbred industry is one of the most successful in the world . in 2010 - 11 , the industry produced over 4000 foals and exported 1600 horses at an estimated value of $ 150 million . so , what is the secret of new zealand ' s remarkable success as a thoroughbred breeding nation ? learn more ›
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fertility rate : this is worked out as the total number of foals ( live , dead or slipped ) as a percentage of total mares covered less exported , not returned , dead or indeterminate results .
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emeraldella is a genus of arthropod known from the middle cambrian burgess shale .

21 specimens of emeraldella are known from the greater phyllopod bed , where they comprise < 0.1 % of the community .

emeraldella

other deposits : emeraldella sp ? from the marjum formation , house range , utah , usa .
emeraldella brocki is very rare in the walcott quarry ( less than 0 . 01 % of the community , caron and jackson , 2008 ) .
emeraldella is a rare arthropod of relatively large body size that belongs with the trilobite - like arthropods , artiopoda . e . brutoni n . sp . from the wheeler formation of west - central utah is the second species described and marks the first confirmed occurrence of emeraldella outside the burgess shale of british columbia . an articulated , flagelliform telson , similar to that of the burgess shale taxon molaria , is recognized in emeraldella . evidence for the presence of lamellae on the exopods of molaria is presented , supporting affinity of that taxon with artiopoda . a close relationship between emeraldella and molaria is tentatively suggested , based on the morphology of tergites and telson .
emeraldella brocki was first described by walcott ( 1912 ) . bruton and whittington ( 1983 ) restudied the material in detail , clarifying many aspects of the animal’s morphology . one possible specimen of emeraldella has also been described from the marjum formation in utah ( briggs and robison , 1984 ) . further work examining the phylogenetic placement of emeraldella and the arachnomorphs has been conducted by hou and bergström ( 1997 ) , wills et al . ( 1998 ) , edgecombe and ramsköld ( 1999 ) , budd ( 2002 ) , cotton and braddy ( 2004 ) , scholtz and edgecombe ( 2006 ) and hendricks and lieberman ( 2008 ) .
emeraldella – from emerald lake , peak , pass , river and glacier north of burgess pass , british columbia , canada . emerald lake was named by guide tom wilson in 1882 for the remarkable deep green colour of the water .
bruton , d . l . and h . b . whittington . 1983 . emeraldella and leanchoilia , two arthropods from the burgess shale , middle cambrian , british columbia . philosophical transactions of the royal society of london b , 300 : 553 - 582 .
emeraldella bella is a problematic animal that also occurs in the burgess shale that is usually considered a primitive soft - bodied arthropod of uncertain class . research has also suggested that the animal belongs to a clade related to chelicerata that includes the spiders and scorpions ( araneida ) , horseshoe crabs ( xiphosura ) and the extinct sea scorpions ( eurypterida ) . this specimen is exceptional and rare in the marjum formation of utah and is shown here with both part and counterpart . emeraldella also has some similarities to parapaleomerus sinensis from the chengjiang biota .
description : this unusual specimen has trilobitomorph features , and has as yet not been described in the literature . it is thought to have affinities with the burgess shale genus emeraldella . notice that the pointed telson is present , a feature not always preserved with either the trilobitomorphs or the aglaspidids of the region . notice too the strong segmentation evident in this part / counterpart example .
emeraldella is of uncertain phylogenetic affinity due to the paucity of specimens . it was previously placed in the arachnomorphs , as closely allied either with the chelicerates ( wills et al . 1998 ; cotton and braddy , 2004 ; hendricks and lieberman , 2008 ) or the trilobites and lamellipedians ( hou and bergström , 1997 ; edgecombe and ramsköld , 1999 ; scholtz and edgecombe , 2006 ) , but it has also been considered as a stem - lineage euarthropod ( budd , 2002 ) .
the trunk of emeraldella has eleven broad segments with curved , smooth margins . each segment has a pair of biramous limbs similar to the ones of the head . behind the trunk segments are two cylindrical body tergites and a long , tapering posterior spine . a dark band running the length of the trunk and into the base of the posterior spine may be the alimentary canal . in the head region , the alimentary canal is u - shaped as it leads forward and upwards from the backward - facing mouth .
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brocki – for reginald walter brock , director of the geological survey of canada from 1907 to 1914 .
lectotype – usnm 57702 in the national museum of natural history , smithsonian institution , washington , dc , usa .
middle cambrian , bathyuriscus - elrathina zone ( approximately 505 million years ago ) .
the body consists of a semicircular head shield , segmented trunk and elongated posterior spine , with total body length ( excluding spine and antennae ) ranging between 1 . 1 cm and 6 . 5 cm . with antennae and spine the entire animal would have reached up to 15 cm in length . the body is convex in cross - section and tapers along the posterior half of the trunk . the head shield is smooth , with no evidence of eyes . a pair of long , flexible antennae consisting of over 110 short segments with bristled junctions is attached to the ventral surface at the front of the head . the mouth is ventral and faces backwards . behind the antennae are five pairs of biramous limbs with a segmented inner branch and a lobed outer branch . the inner branch has six podomeres , including the gnathobase ( a robust and spiny basal podomere or segment used for crushing food items ) , four adjacent podomeres that also bear spines , and a slender terminal podomere armed with three sharp claws . the outer branch of the biramous limb is broad and has three main lobes with filaments and blades .
the inner branches of the biramous limbs were likely used for walking on the sea floor , especially the middle eight or nine limbs , which were longer than the posterior limbs . spines on the inner margin of the walking limbs could have been used to grasp soft prey items , and the terminal claws would push food towards the ventral gnathobases . these strong spiny plates would then shred the food and pass it along the underside of the body towards the mouth . the antennae were used to explore the environment and search for live prey or carcasses , perhaps by ploughing through the soft sediment . while the head was tilted down in the search for food , the posterior segments of the body and the posterior spine may have flexed upwards for balance . the outer limb lobes likely served as gills for respiration . the animal might have been capable of short bursts of swimming , using its broad outer limb branches to propel itself through the water using a wave - like motion .
briggs , d . e . g . and r . a . robison . 1984 . exceptionally preserved non - trilobite arthropods and anomalocaris from the middle cambrian of utah . the university of kansas paleontological contributions , 111 : 1 - 24 .
budd , g . e . 2002 . a palaeontological solution to the arthropod head problem . nature , 417 : 271 - 275 .
caron , j . - b . and d . a . jackson . 2008 . paleoecology of the greater phyllopod bed community , burgess shale . palaeogeography , palaeoclimatology , palaeoecology , 258 : 222 - 256 .
cotton , t . j . and s . j . braddy . 2004 . the phylogeny of arachnomorph arthropods and the origin of the chelicerata . transactions of the royal society of edinburgh - earth sciences , 94 : 169 - 193 .
edgecombe , g . d . and l . ramsköld . 1999 . relationships of cambrian arachnata and the systematic position of trilobita . journal of paleontology , 73 : 263 - 287 .
hendricks , j . r . and b . s . lieberman . 2008 . phylogenetic insights into the cambrian radiation of arachnomorph arthropods . journal of paleontology , 82 : 585 - 594 .
hou , x . and j . bergström . 1997 . arthropods of the lower cambrian chengjiang fauna , southwest china . fossils and strata , 45 : 1 - 116 .
scholtz , g . and g . d . edgecombe . 2006 . the evolution of arthropod heads : reconciling morphological , developmental and palaeontological evidence . development genes and evolution , 216 : 395 - 415 .
walcott , c . d . 1912 . middle cambrian branchiopoda , malacostraca , trilobita and merostomata . smithsonian miscellaneous collections , 57 : 145 - 228 .
wills , m . a . , d . e . g . briggs , r . a . fortey , m . wilkinson , and p . h . sneath . 1998 . an arthropod phylogeny based on fossil and recent taxa . pp . 33 - 105 . in g . d . edgecombe ( ed . ) , arthropod fossils and phylogeny . columbia university press , new york .
hou & bergström , 1997 : arthropods of the lower cambrian chengjiang fauna , southwest china . - - fossils & strata , number 45 , 22nd december 1997 , pp . 1 - 116 briggs d . e . g . , and r . a . robison . 1984 . exceptionally preserved non trilobite arthropods and anomalocaris from the middle cambrian of utah . university of kansas paleontological contributions , paper 111 : 1 - 24 .
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parent taxon : arachnomorpha according to d . e . g . briggs et al . 2008
department of earth sciences , university of cambridge , sedgwick museum , downing street , cambridge cb2 3eq , u . k .
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