strepsodiscus is an extinct genus of very primitive fossil snail-like molluscs from the early part of the late cambrian ( dresbachian age ) of north america .

bouchet & rocroi ( 2005 ) divide the bellerophontoidea into 8 families listed paleozoic molluscs with isostrophically coiled shells of uncertain position within mollusca ( gastropoda or monoplacophora ) knight , et al. , 1960 included this genus within the cyrtolitidae , a paraphyletic or polyphyletic assemblage of proto-gastropods and tergomyan molluscs .

strepsodiscus

a new species of strepsodiscus knight ( gastropoda , bellerophontoi . . . : ingenta connect
taxon rank : family strepsodiscidae ( new , informal ) = genus strepsodiscus in knight , et al . , 1960 p . i 174 .
strepsodiscus is an extinct genus of very primitive fossil snail - like molluscs from the early part of the late cambrian ( dresbachian age ) of north america .
notes : knight , et al . , 1960 include strepsodiscus within the cyrtolitidae , an assemblage that includes both torted ( gastropod ) and untorted ( tergomyan ) forms and should be restricted to non - gastropods . for taxonomic purposes one might perhaps suggest transferring the genus strepsodiscus to its own ( presumably ) paraphyletic family (
strepsodiscidae
)
knight , et al . , 1960 included this genus within the cyrtolitidae , a paraphyletic or polyphyletic assemblage of proto - gastropods and tergomyan molluscs . strepsodiscus may be too primitive to be a true gastropod .
strepsodiscus austrinus sp . nov . ( gastropoda , bellerophontoidea ) from the lampazar formation at angosto de la quesera locality ( salta province , argentina ) is described . the new species is characterized by having a laterally compressed smooth shell . the spire is composed of three tangent whorls slightly coiled to the left , and the final 1 / 4 whorl free . strepsodiscus knight is reported in south america for the first time . the specimens studied were collected in association with trilobites of the parabolina ( neoparabolina ) frequens argentina zone ( uppermost cambrian ) .
the first gastropods are considered to have been exclusively marine , with the earliest representatives of the group appearing in the late cambrian ( chippewaella , strepsodiscus ) . early cambrian forms like helcionella and scenella are no longer considered gastropods , and the tiny coiled aldanella of earliest cambrian time is probably not even a mollusk .
strepsodiscus is an extinct genus of very primitive fossil snail - like molluscs from the early part of the late cambrian ( dresbachian age ) of north america . the coiled , slightly asymmetrical shells are about 3 cm in height . it is not known whether these are shells of gastropods ( sea snails ) or monoplacophorans , which are more primitive mollusks .
scientists think that gastropods , bivalves , and cephalopods developed in the cambrian period about 541 - 585 . 4 million years ago . gastropods , members of the class gastropoda , are colloquially known as snails and slugs , whether terrestrial , marine or freshwater . according to fossil records , the earliest gastropods lived in the ocean and possibly appeared during the cambrian period . from this , the mollusks of the genera strepsodiscus and chippewaella developed to become the most primitive snails .
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bouchet , p . ; rocroi , j . - p . ( ed . ) ( 2005 ) . classification and nomenclator of gastropod families . malacologia : international journal of malacology , 47 ( 1 - 2 ) . conchbooks : hackenheim . isbn 3 - 925919 - 72 - 4 . 397 pp .
part of : malacologia : international journal of malacology . institute of malacology : ann arbor . issn 0076 - 2997 , more
this is a vademecum for about 2 , 400 suprageneric names in recent and fossil gastropods , from the subtribe to the superfamily . for each name full bibliographic reference is given , with date of publication , type genus , nomenclatural availability and validity under the rules of the iczn . further 730 names of higher taxa are listed separately . altogether 611 valid families , 202 of them exclusively fossil , are recognised in the proposed classification .
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full reference : j . b . knight . 1948 . further new cambrian bellerophont gastropods . smithsonian miscellaneous collections 111 ( 3 ) : 1 - 6
see also horný 1963 , knight et al . 1960 , runnegar and jell 1976 , sepkoski 2002 , wagner 1999 and yochelson and nuelle 1985
full reference : c . d . walcott . 1879 . descriptions of new species of fossils from the calciferous formation . annual report of the new york museum of natural history 32 : 129 - 131
see also pinilla et al . 2008 , runnegar 1981 , walcott 1879 and walcott 1912
full reference : c . d . walcott . 1890 . description of new forms of upper cambrian fossils . proceedings of the united states national museum 13 : 267 - 279
distribution : found only at loc . 76 . hoyts quarry , 4 miles west of saratoga springs ( cambrian of new york )
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tropidodiscus curvilineatus ( conrad 1842 ) . length 3 cm . early devonian of laurentia ( new york state )
during the furongian ( late paiban age ) the first gastropods , torted bellerophontiforms , appear . according to the torted bellerophonts are here shown to be diphyletic . for the sake of convenience the order bellerophontida ( or superfamily bellerophontoidea , depending on your prefeernce ; in any case cladistic terminology does not use linnaean ranks ) is defined as the later clade to emerge . the early , paraphyletic grouping belongs to a separate grade , which includes subfamily tropodiscinae and a number of other species . were one to use a linnaean approach , and assuming the diphyly is valid , then these would represent a separate order , let us say the
tropodiscida
, which stand right at the beginning of gastropod evolution .
limpets , periwinkles , whelks , snails , slugs , torted bellerophonts , etc .
characteristics : $ torsion ; streptoneury ( as a result of torsion nervous system is twisted in a figure - 8 shape - reversed in advanced forms ) ; prosobranchy ( as a result of torsion gills in front of the heart - reversed in advanced forms ) ; primitively planispiral shell ; docoglossa - type radula .
note : using a functional definition , with gastropoda defined solely by the presence of torsion , this is the clade that represents the class as defined by cuvier . however some workers prefer to use crown group definitions ; i . e . the clade gastropoda as the last common ancestor of all extant gastropods . in that case this clade should be called
gastropodamorpha
or
gastropodiformes
to distinguish it from the
gastropoda proper
stratigraphic range : early furongian ( dresbachian [ runnegar _ and jell 1976 p . 121 ] = paiban age ) .
geographic distribution : laurentia [ = n . am - knight , et al . , 1960 p . i 174 ]
characteristics : $ anal emargination a deep angular sinus ; $ coiling slightly asymmetrical , with closely coiled early whorls protruding toward left , last whorl disjunct , all whorls sharply crested ; has posterior train ; growth lines are only surface features known [ knight , et al . , 1960 p . i 174 ] .
stratigraphic range : late furongian ( franconian - trempealeauan ( chalarostrepsis ) [ runnegar & jell ( 1976 ) ] = early or late dolgellian ) to early devonian ( tropidodiscus ) [ knight , et al . , 1960 p . i 179 ]
geographic distribution : cosmopolitan [ = n . am , s . am . , eu . , asia - knight , et al . , 1960 p . i 179 ]
characteristics : coil narrow , with wide umbilici ; $ slit deep ( evolved independently of bellerophontina ; commonly with a definite posterior train [ knight , et al . , 1960 p . 179 ] .
taxon rank : family tropidodiscidae knight , 1956 = subfamily tropidodiscinae in knight , et al . , 1960 p . i 179 = family tropidodiscidae in golikov & tarobogatov ( 1975 )
notes : according to wagner 1999 the tropidodiscid bellerophonts are the sister group of all other gastropods .
original material by m . alan kazlev ( creative commons ) this material may be freely used all other material © original authors or sources
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knight , j . b . , cox , l . r . , keen , a . m . , batten , r . l . , yochelson , e . l . , and robertson , r . ( 1960 ) . systematic descriptions ( archaeogastropoda ) . in moore , r . c . ( ed . ) treatise on invertebrate paleontology . part i . mollusca 1 , pp . 169 - 310 . geological society of america and kansas university press , colorado and kansas .
authors : pinilla , m . k . ; sabattini , n . m . ; tortello , m . f . ; aceñolaza , g . f .
neues jahrbuch für geologie und paläontologie continuously publishes current original contributions from all fields of geology , ever since its foundation in 1807 . all published contributions are in the english language .
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a gastropod is any member of gastropoda , the largest and most diverse class of the phylum mollusca , with about 75 , 000 known living species . gastropods are also referred to as univalves since most have a single shell , or valve , which is characteristically coiled or spiraled , as in snails , limpets , abalones , cowries , whelks , and conches . but gastropoda is very diverse and many , such as slugs and sea slugs ( nudibranches ) , lack shells ; some even have shells with two halves , appearing as if bivalves .
this class is very successful and occupies almost every niche on earth . there are marine , freshwater , and terrestrial species , and they range in the ocean from the intertidal zone to the deepest trenches , on land from deserts to high mountains , and in all types of freshwater ecosystems . they include herbivores , carnivores , detritivores , and parasites . along the rocky shore of an estuary , for example , the many different types of gastropods may include those that scavenge for dead fish or other organic debris while others eat small algae or bacteria and still others at the top of the food chain eat mollusks and other gastropods lower on the food chain .
as with all mollusks , gastropods are characterized by having a true coelom , a fluid - filled and fully - lined body cavity ; a body divided into the three parts of head , visceral mass , and muscular foot ; and organ systems for circulation , respiration , digestion , excretion , nerve conduction , and reproduction ( towle 1989 ) . however , the gastropod body plan involves a torsion or twisting during larval development whereby the visceral mass twists 180 degrees in relation to the head , bringing the mantle cavity to the anterior of the animal . thus , the gills , and renal and anal openings are now near the front of the animal . gastropoda ( meaning
stomach - foot
) is typified by a large , ventral , muscular foot for locomotion , and a distinct head that has eyes and sensory tentacles .
the varied forms and colors of gastropod shells appeal to the inner aspect of humans ; even many gastropods without shells , such as nudibranches , can be brightly colored and beautiful . some gastropods , such as abalone , also serve as a food delicacy , and the abalone is a source of mother of pearl for decorative use .
gastropod classification follows a number of systems , but today is typically divided into two subclasses : orthogastropoda and eogastropoda ( ponder and lindberg 1997 ) .
the gastropod body plan is believed to be based on that of ancestral mollusks , with the difference that most of the body behind the head ( mantle , mantle cavity , visceral mass ) underwent torsion , twisting it 180 degrees , and bringing the mantle cavity , gills , and renal and anal openings forward near the head . some species are twisted counterclockwise and some clockwise . similarly , the shell is coiled asymmetrically , with reach coil beneath the preceding . the gastropod is able to bring its head into the mantle cavity when endangered ( towle 1989 ) .
gastropods typically have a well - defined head with two or four sensory tentacles , and a ventral foot , which gives them their name ( greek gaster , stomach , and poda , feet ) . the eyes that may be present at the tip of the tentacles range from simple ocelli that cannot project an image ( simply distinguishing light and dark ) , to more complex pit and even lens eyes ( götting 1994 ) . the larval shell of a gastropod is called a protoconch .
the feet of some gastropods are modified . the pteropods , or
sea butterflies ,
have a winglike flap , which allows them to swim on the surface of the sea ( towle 1989 ) .
gastropods have an open circulatory system , meaning that the blood is not enclosed entirely within blood vessels , but is released into spaces in tissues . it is collected in the gills or heart , and then pumped through a heart back into the blood - filled space , called a hemocoel or blood cavity .
most gastropods utilize for feeding a radula , which is a tongue - like ribbon of tissue with small teeth located back of the mouth , which act like a saw in scraping food from surfaces . the radula is usually adapted to the food that a species eats . aquatic snails use it to scrape algae and terrestrial snails to saw off leaves of garden plants ( towle 1989 ) . the simplest gastropods are the limpets and abalones , both herbivores that use their hard radulas to rasp at seaweeds on rocks . carnivorous oyster drills use the radula to bore through oyster shells to feed on the inner tissue . not all gastropods have radulas . marine gastropods include herbivores , detritus feeders , carnivores , and a few ciliary feeders in which the radula is reduced or absent .
most members have a shell , which is in one piece and typically coiled or spiraled that usually opens on the right hand side ( as viewed with the shell apex pointing upward ) . several species have an operculum that operates as a trapdoor to close the shell . this is usually made of a horn like material , but in some mollusks it is calcareous . in some members , the slugs , the shell is reduced or absent , and the body is streamlined so its torsion is relatively inconspicuous .
while the best - known gastropods are terrestrial , more than two thirds of all species live in a marine environment . many marine gastropods are burrowers and have siphons or tubes that extend from the mantle and sometimes the shell . these act as snorkels , enabling the animal to continue to draw in a water current containing oxygen and food into their bodies . the siphons are also used to detect prey from a distance . these gastropods breathe with gills , but some freshwater species and almost all terrestrial species have developed lungs . gastropods with lungs all belong to one group , pulmonata .
sea slugs are often flamboyantly colored , either as a warning if they are poisonous or to camouflage them on the corals and seaweeds on which many of the species are found . their gills are often in a form of feathery plumes on their backs , which gives rise to their other name , nudibranchs . nudibranchs with smooth or warty backs have no visible gill mechanisms and respiration may take place directly through the skin . a few of the sea slugs are herbivores and some are carnivores . many have distinct dietary preferences and regularly occur in association with certain species .
certain trail - like markings preserved in ancient sedimentary rocks are thought to have been made by gastropods crawling over the soft mud and sand . although these trails are of debatable origin , some of them do resemble the trails made by living gastropods today .
by the ordovician period , the gastropods were a varied group present in a range of aquatic habitats . commonly , fossil gastropods from the rocks of the early paleozoic era are too poorly preserved for accurate identification . still , the silurian genus poleumita contains 15 identified species . fossil gastropods are less common during the paleozoic era than bivalves .
most of the gastropods of the paleozoic era belong to primitive groups , a few of which still survive today . by the carboniferous period , many of the shapes we see in living gastropods can be matched in the fossil record , but despite these similarities in appearance the majority of these older forms are considered not to be directly related to living forms . it was during the mesozoic era that the ancestors of many of the living gastropods evolved .
one of the earliest known terrestrial ( land - dwelling ) gastropods is maturipupa which is found in the coal measures of the carboniferous period in europe , but relatives of the modern land snails are rare before the cretaceous period when the familiar helix first appeared .
in rocks of the mesozoic era , gastropods are slightly more common as fossils , their shell often well preserved . their fossils occur in beds that were deposited in both freshwater and marine environments . the
purbeck marble
of the jurassic period , and the
sussex marble
of the early cretaceous period , which both occur in southern england , are limestones containing the tightly packed remains of the pond snail viviparus .
rocks of the cenozoic era yield very large numbers of gastropod fossils , many of these fossils being closely related to modern living forms . the diversity of the gastropods increased markedly at the beginning of this era , along with that of the bivalves .
gastropod fossils may sometimes be confused with ammonites or other shelled cephalopods . an example of this is bellerophon from the limestones of the carboniferous period in europe , which may be mistaken for a cephalopod .
gastropods are one of the groups that record the changes in fauna caused by the advance and retreat of the ice sheets during the pleistocene epoch .
in their work , which has become a standard reference in the field , ponder and lindberg ( 1997 ) present the orthogastropoda as one of two subclasses of the gastropoda , the other subclass being the eogastropoda .
this subclass , which one could call the true snails , is defined most briefly as all those gastropods that are not members of patellogastropoda , the true limpets , or its ancestors . included are abalone , snails , whelks , cowries , sea slugs ( nudibranches ) , winkels , cones , and so forth , as well as keyhole limpets .
orthogastropods form a clade , supported by unambiguous synapomorphies . these synapomorphies ( a series of characteristics that appear in its members , but not in the other forms it diverged from ) are the identifying characteristics of the clade .
a flexoglossate radula ( with a flexible radular membrane ) . the radula is the snail ' s tongue , used as a rasping tool .
a single left hypobranchial gland ( on organ at gill , which releases secretions , such as the reddish dye tyrian purple ) .
true limpets are marine gastropods with flattened , cone - shaped shells in the order patellogastropoda in the subclass eogastropoda , the other subclass of gastropoda , along with orthogastropoda .
limpets live throughout the intertidal zone , from the high zone ( upper littoral ) to the shallow subtidal on the rocky coasts of most oceans . limpets can be commonly found attached to rocks , looking like little disks or bumps on the rock surface . they attach themselves to the substratum using pedal mucus and a muscular
foot ,
which enables them to remain attached through dangerous wave action and which also seals against the rock to protect from desiccation during low tide .
limpets forage by grazing on algae found on rock surfaces . they scrape films of algae from the rock with a radula . limpets move by rippling the muscles of their foot in a wave - like motion .
some limpets have a hole at the top , through which gaseous exchange can occur . most limpets are less than 3 inches ( 8 centimeters ) long , but a west mexican limpet grows to be 8 inches ( 20 centimeters ) .
limpets found on sheltered shores ( limpets that are less frequently in contact with wave action , and thus less frequently in contact with water ) have a greater risk of desiccation due to the effects of sunlight , water evaporation , and the wind . to avoid drying out they will clamp to the rock they inhabit , minimizing water - loss from the rim around their base . as this occurs chemicals are released that promote the vertical growth of the limpet ' s shell .
some species of limpets exhibit homing behavior , returning to the same spot on the rock , known as a
home scar ,
just before the tide recedes . in such species , the shape of their shell often grows to precisely match the contours of the rock surrounding the scar . this behavior presumably allows them to form a better seal to the rock and may help protect from either predation or desiccation . it is still unclear how limpets find their way back to the same spot each time , but it is thought that they follow a mucus trail left as they move , and utilize their tentacles .
other species , notably lottia gigantea seem to
garden
a patch of algae around their home scar . they will aggressively push other organisms out of this patch by ramming with their shell , thereby allowing their patch of algae to grow for their own grazing . also , where the limpets eat the algae off of bare rocks , it causes places where other organisms can grow and thrive .
limpets are preyed upon by a variety of organisms including starfish , shore - birds , fish , seals , and humans . limpets exhibit a variety of defenses , such as fleeing or clamping their shells against the substratum . the defense response can be determined by the type of predator , which can often be detected chemically by the limpet .
limpets can be long lived , with tagged specimens surviving for more than ten years .
the taxonomy of the gastropoda is under constant revision , but more and more of the old taxonomy is being abandoned . in a sense , we can speak of a taxonomic jungle when we go down to the lower taxonomic levels . the taxonomy of the gastropoda can be different from author to author . with the arrival of dna - sequencing , further revisions of the higher taxonomic levels are to be expected in the near future .
according to the newest insights ( ponder & lindberg , 1997 ) , the taxonomy of the gastropoda should be rewritten in terms of strictly monophyletic groups . they recognize the subclasses of orthogastropoda and eogastropoda .
integrating new findings into a working taxonomy will be a true challenge in the coming years . at present , it is impossible to give a classification of the gastropoda that has consistent ranks and also reflects current usage . convergent evolution , observed at especially high frequency in the gastropods , may account for the observed differences between phylogenies obtained from morphological data and more recent studies based on gene sequences .
other extant classes of the mollusca are bivalvia , scaphopoda , aplacophora , polyplacophora , monoplacophora , and cephalopoda .
breen , p . a . 1971 . “homing behavior and population regulation in the limpet acmaea ( collisella ) digitalis . ” veliger 14 : 177 - 183 .
götting , k . - j . 1994 . “schnecken . ” in u . becker , s . ganter , c . just , and r . sauermost , lexikon der biologie . heidelberg : spektrum akademischer verlag . isbn 3860251562 .
jeffery , p . 2001 . suprageneric classification of class gastropoda . london : the natural history museum .
ponder , w . f . , and d . r . lindberg . 1997 . “towards a phylogeny of gastropod molluscs : an analysis using morphological characters . ” zoological journal of the linnean society 119 : 83 - 2651 .
towle , a . 1989 . modern biology . austin , tx : holt , rinehart , and winston . isbn 0030139198 .
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full reference : r . p . whitfield . 1878 . preliminary descriptions of new species of fossils from the lower geological formations of wisconsin . annual report of the wisconsin geological survey for 1877 : 50 - 89
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nomenclator zoologicus . a list of the names of genera and subgenera in zoology from the tenth edition of linnaeus , 1758 to the end of 2004 . digitised by ubio from vols . 1 - 9 of neave ( ed . ) , 1939 - 1996 plus supplementary digital - only volume . urltoken ( as at 2006 ) .
the evolution of snails is still not clear enough , and their taxonomy or scientific classification keep changing over time . out of the approximately 5 million species of invertebrate animals ( those that do not have an internal skeleton or backbone ) , about 50 , 000 - 85 , 000 are mollusks that have adapted to both aquatic habitats and terrestrial habitats .
about 50 , 000 - 85 , 000 are mollusks that have adapted to both aquatic habitats and terrestrial habitats .
in the ordovician period , mollusks underwent great diversification and cephalopods , bivalves and gastropods multiplied in number and species in aquatic habitats . they still did not dominate the land , but they were colonizing other places to live .
during the paleozoic era , gastropods were abundant in the sea . some groups of them still exist and show characteristics of that period . from the mesozoic era , marine snails began to evolve and developed the features that are present in current snails . fossils of mesozoic species that are in good condition show reasonable resemblances that make them easily identifiable as snails .
the clade stylommatophora , which includes land snails and slugs , appeared about 150 million years ago , probably in the cretaceous period . this group diversified , as result of adaptive radiation , and gave origin to numerous families of snails and slugs , some of which persist in the sites where their fossils arose . by the way , adaptive radiation is a biological process through which living beings rapidly diversify , that is , new forms of a family or group appear that adapt to a particular environment , called a niche .
after stylommatophora , the groups basommatophora , orthurethra , and sigmurethra were developed . thus , the number of land snail species grew , and they occupied several habitats or places to live . the oldest registered terrestrial gastropod is maturipupa , which lived in europe during the carboniferous . the species of the genus helix , from which the garden snail is perhaps the best known , flourished in the cretaceous period . in the cenozoic , the gastropods once again had a period of significant diversification .
land snails are currently abundant animals ( with some exceptions that are in danger of extinction ) and are present in most of the habitats of the planet .
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vulpes riffautae is an extinct species of fox from the late miocene of chad ( approximately 7 ma ) .

fossils of v. riffautae potentially represent the earliest record of the dog family , canidae , in the old world .

v. riffautae was intermediate in size between rüppell 's fox ( vulpes rueppellii ) and the fennec fox ( v. zerda ) .

vulpes riffautae

the oldest african fox ( vulpes riffautae n . sp . , canidae , carnivora ) recovered in late miocene deposits of the djurab desert , chad
the oldest african fox ( vulpes riffautae n . sp . , canidae , carnivora ) recovered in late miocene deposits of the djurab desert , chad | springerlink
vulpes riffautae is an early fox species known by fossils found the african country of chad , which date back to the miocene around 7 mya . it was probably the earliest representative of the dog family anywhere in the old world ( dogs evolved in north america ) .
though debatable , the earliest fox is thought to be metalopex macconnelli a californian fox dating to about 10 . 5 ma . how the fox dispersed into africa from there , is a major focus of palaeontological debate . vulpes riffautae is the earliest representative of foxes in africa about 8 ma .
the latest paper to be published about the remains of malapa draws our attention to a little known remains of fox fossils . the three specimens were compared to both fossil and extant remains of fox . the malapa fox is sufficiently different from the comparative sample to warrant a new species for the malapa fox – vulpes skinneri .
these remains were attributed to vulpes cf . v . chama when they were first described in another paper in 2010 . time passes and research continues to a point where the palaeontological team now believe there is considerable shape differences in the bones and teeth to suggest a new species . the malapa fox remains were compared to a wide variety of museum collections throughout the world , including the natural history museums of new york , south africa and stockholm .
we report on the oldest fox ( canidae ) ever found in africa . it is dated to 7 ma based on the degree of evolution of the whole fauna . it belongs to a new species . its overall size and some morphological characteristics distinguish the chadian specimen from all the other foxes . the presence of vulpes and of the genus eucyon in slightly younger african locality , as well as in southwestern europe in the late miocene , may indicate that canids migrated in europe from africa through a trans - mediterranean route .
berta a ( 1987 ) origin , diversification , and zoogeography of the south american canidae . fieldiana geol 39 : 455–471
, a new ictithere ( mammalia , carnivora , hyaenidae ) from the late miocene of toros - menalla , chad . comptes rendus palevol 4 : 671–679
bowdich te ( 1821 ) an analysis of the natural classification of mammalia for the use of students and travellers . j . smith , paris , pp 1–115
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the palaeoanthropological community got excited with the discovery of new hominin fossils at the site of malapa , in south africa , over two years ago .
the excited was linked to the possibilities of what the two individuals could reveal about the origins of humanity . the refreshing aspect here is the scientific team’s endeavour to publish papers on the other fossil remains that found their way into the malapa cave system 1 . 977 million years ago .
one would imagine europe being the missing link in the chain from the americas and africa . from then on fossil fox remains have been uncovered in the eastern and southern regions of the african continent . by the pleistocene nyctereutes terblanche was roaming the now famous “cradle of humankind” in south africa . the scottish palaeontologist robert broom referred to this species as a jackal , but it is now generally considered a racoon dog .
the species compared included v . pattisoni and v . chama , with the malapa fox showing more similarities with the extant ( living ) foxes of south africa . most species of organism on the planet has a holotype , which is a specimen that other skeletal remains can be compared to . it basically acts as a standard for a species . the team of scientists realised that the holotype of v . pattisoni is likely to be a juvenile as evidence from the crowded cheek teeth .
the malapa fox exhibited differences in dental morphology to those of the above mentioned two fossil species and so the team of palaeontologists considered it appropriate to propose a new species – v . skinneri .
professor john d . skinner was director of the university of pretoria mammal research institute and so his name was appropriate from the malapa fox .
charles hails from longford in the midlands of ireland . from an early age , he developed an interest in general knowledge which morphed into archaeology . he graduated with a b . sc . in applied archaeology at the institute of technology sligo , in 2011 . his dissertation involved a class of neolithic monument called an earthen embanked enclosure - the irish equivalent of a henge . one module - world archaeology - would determine his future however . after being introduced to australopithecines , he was hooked ever since . in 2012 , he graduated with an m . sc . in palaeoanthropology and palaeolithic archaeology at the ucl institute of archaeology and department of anthropology . his masters dissertation focused upon specie - level signatures of mandibular 2nd molars around 2 million years of age . now out of academia , he hopes to return and pursue a phd . he maintains the cennathis youtube channel , blog , podcast , twitter and facebook pages . he currently lives in the city of london , uk .
discoveries by archaeology team give new clues on life in ancient jewish . . .
organic components of levantine rock art investigated for the first time with . . .
heritagedaily is a publishing platform with a focus on archaeology , anthropology , palaeontology and general science .


papilio ( chilasa ) epycides , the lesser mime , is a swallowtail butterfly found in india and parts of south-east asia .

the butterfly belongs to the mime ( chilasa ) subgenus or the black-bodied swallowtails .

it is a mimic of a common indian danainae , the glassy tiger butterfly .

papilio epycides

vijay barve marked
papilio epycides
as trusted on the
papilio ( chilasa ) epycides hewitson , 1862
page .
you selected papilio epycides curiatius fruhstorfer , 1901 . this is a synonym for :
no one has contributed data records for papilio epycides melanoleucus yet . learn how to contribute .
papilio epycides occurs in india , nepal , myanmar , thailand , laos , cambodia and china .
here you can see some photos of chilasa epycides ( lesser mime ) ( male above / female below ) . chilasa epycides is a member of the
[ thailand ] papilio epycides epycides ; godfrey , 1927 : 119 . ( north ) papilio epycides epycides ; godfrey , 1930 : 209 . ( chiang mai ) chilasa epycides epycides ; pinratana , 1977 : 22 , fig . p16 , ♂ , ♂ ( un ) . ( north ) chilasa epycides epycides ; pinratana , 1992 : 18 , 40 , pl . 19 , fig . 17 , ♂ . ( chiang mai ) chilasa epycides epycides ; ek - amnuay , [ 2007 ] : 98 , pl . 21 , fig . p44 , ♂ , ♂ ( un ) , ♀ . ( wiang pa pao ) chilasa epycides hypochra ; kimura et al . , 2011 : 119 , fig . ♂ , ♂ ( un ) , ♀ , ♀ ( un ) . ( doi pha hom pok / chiang dao / doi suthep ) papilio epycides hypochra ; ek - amnuay , 2012 : 98 , pl . 21 , fig . p41 , ♂ , ♂ ( un ) , ♀ . ( wiang pa pao / mae hong son ) [ laos ] papilio epycides f . epycides ; dubois & vitalis de salvaza , 1921 : 9 . ( laos ) chilasa epycides epycides ; osada , uémura & uehara , 1999 : 200 , pl . 6 , figs . ♂ , ♂ , ♂ , ♀ . ( phong saly / lak sao ) papilio epycides hypochra ; cotton & racheli , [ 2007 ] : 370 - 371 . ( xam neua : phou phan / xaysomboun : nam cha / lak sao ) chilasa epycides hypochra ; nakamura & wakahara , 2012 ; 41 . [ vietnam ] papilio epycides epycides ; metaye , 1957 : 81 . ( north ) chilasa epycides epycides ; monastyrskii , 2007 : 40 - 41 , pl . 8 , figs . 2a♂ , 2b♂ ( dark form ) . ( north : hoang lien ; van ban ; pia oac ; hoa binh / central : vu quang ; huong son ; kon ka kinh )
the genus includes a number of well - known north american species such as the western tiger swallowtail ( papilio rutulus ) . familiar species in asia include the mormons ( papilio polytes , papilio polymnestor , papilio memnon , and papilio deiphobus ) , the orchard and ulysses swallowtails in australia ( papilio aegeus , papilio ulysses , respectively ) and the citrus swallowtail of africa ( papilio demodocus ) .
this species mimes other uneatable butterflies . therefore chilasa epycides is also portected against preditors .
in common with other members of the genus , epycides is palatable to insectivorous birds . in order to protect itself from predation it has evolved its appearance to become superficially very similar to an unpalatable species . papilio epycides is a mimic of the tiger butterflies parantica and tirumala .
papilio epycides camilla rousseau - decelle , 1947 : bull . soc . ent . fr . 51 : 128 - 129 . tl . hoa binh , tonkin . ( ♂ )
jennifer hammock split the classifications by mcz resource from chilasa epycides ( hewitson , 1862 ) to their own page .
on : papilio epycides hypochra jordan , 1909 od : in seitz , grossschmett . erde . 9 : 41 . tl : shan states , burma . distribution : myanmar , n . thailand , laos , vietnam .
a study of hybrids betweeen papilio xuthus and the p . polyxenes - machaon group
vijay barve marked the common name
lesser mime
in an unknown language from
chilasa epycides ( hewitson , 1862 )
as trusted .
here are about 60 species of papilio in the oriental region , and 210 species worldwide .
the taxonomy and identity of papilio nitra w . h . edwards in colorado ( papilionidae )
{ author1 , author2 . . . } , ( n . d . ) . papilio epycides hewitson , 1869 . [ online ] india biodiversity portal , species page : { name of species field } available at : urltoken [ accessed date jul 10 , 2018 ] .
the fore wings are black . many yellowish stripes and spots dominate the wing . the underside of chilasa epycides is very similar to the upside . the basic colour is brown .
chilasa epycides is a butterfly from the australasia / indomalaya ( australia ) ecozone . you can observe this butterfly from the north of india until the south of china and malaysia .
systematics of the papilio glaucus and p . troilus species groups ( lepidoptera : papiliondae ) : inferences from allozymes
the tawny mime papilio ( chilasa ) agestor is a swallowtail butterfly , native to india and widely found across asia . the butterfly belongs to the mime subgenus , chilasa , of the genus papilio or the black - bodied swallowtails .
evolution of the mimetic african swallowtail butterfly papilio dardanus : molecular data confirm relationships with p . phorcas and p . constantinus
the hind wings of chilasa epycides are black . many yellowish stripes and spots dominate the wing . at the inner edge there is a red eye . the underside is very similar to the upside . the basic colour is brown .
the inferred pattern of phylogenetic relationships within papilio as found here at least partly conflicts with these previously proposed vicariance hypotheses . the major conflict stems from the origin of papilio alexanor , subgenus chilasa , and subgenus pterourus from a common ancestor and the distant relationship between new world branches of papilio and the old world p . ( eleppone ) anactus . the origin of chilasa from the common ancestor with eleppone via gondwanaland to australia and asia , as was suggested based on a traditional view of papilio phylogeny ( scriber , 1995b ) , is not supported by our results .
a second lineage of old world princeps - like swallowtails underlies extant asian sinoprinceps and holarctic papilio ( sensu stricto ) , which apparently diverged around 33–36 mya . speciation in papilio ( sensu stricto ) has been attributed to several subsequent dispersals across beringia before and during the pleistocene ( sperling , 1987 ) .
. the first description was in 1864 by hewitson . the wingspan is about 8 – 9 cm . this butterfly is a member of the family papilionidae . the basic colour of chilasa epycides is black . it has many yellowish stripes and spots on the wings .
age estimates ( ±sd ) in millions of years of internal nodes in papilio tree using calibration points shown in figure 4 and model - corrected branch lengths .
extensive metallic iridescent scales have evolved independently in three different lineages of papilio : achillides , known as gloss - papilios ; the p . nireus group from african princeps ; and papilio troilus , a mimic of battus philenor . this trait appears convergent with iridescent scales in some troidini ( e . g . , trogonoptera ) .
many other morphological characteristics of papilio may have polyphyletic origins ( fig . 4 ) . some of these have been used as significant taxonomic characters justifying papilio subdivisions . for example , a granular female secretion attached to the egg surface , which is well known in troidini , is also reported from a number of species of papilio ( igarashi , 1984 ) . in the context of our phylogeny there is no consistent pattern for this character among species groups , although it had been hypothesized previously as a synapomorphy of heraclides + chilasa ( igarashi , 1984 ) . this character instead appears to be either symplesiomorphic or derived in two separate lineages ( chilasa and heraclides ) in papilio .
reconstructed ancestral states and major gains and losses ( excluding character changes in individual terminal taxa ) for selected characters in papilio using pachliopta neptunus ( troidini ) as its sister taxon .
now included in the genus papilio , are the former genera : achillides , eleppone , druryia , heraclides ( giant swallowtails ) , menelaides , princeps , pterourus ( tiger swallowtails ) , and sinoprinceps .
comparison of ae ' s ( 1979 ) differentiation index versus coi - coii uncorrected percentage nucleotide sequence divergence for papilio species pairs from the same species groups , different species groups , and different subgenera .
[ vietnam ] chilasa imitata ; monastyrskii , 2007 : 41 - 42 , pl . 8 , figs . 3a♂ , 3b♀ . ( central : bi doup / nui ba ; hon ba ) chilasa epycides imitata ; miyazaki , saito & saito , 2007c : 36 - 37 , fig . pa - 36♂ . ( lam dong : dambri )
only two species represented the subgenus chilasa in our study . one important species group , elwesi , could not be sampled . nevertheless , monophyly of chilasa sensu stricto , represented by c . epycides and c . clytia , has strong support . contrary to hancock ' s conclusions , chilasa appears to be the sister taxon of pterourus . although support for this particular result is only moderate , chilasa clearly belongs to the same lineage as pterourus and papilio alexanor , as indicated by strong bootstrap support in the combined ml tree and the high posterior probability of the corresponding clade in the bayesian phylogeny .
phylogenetic relationships of species groups and major subdivisions of papilio sensu lato : left : tree according to hancock ( 1983 ) ; right : tree based on data presented here . roman numerals correspond to hancock ' s genera : i = pterourus ; ii = heraclides ; iii = eleppone ; iv = chilasa ; v = papilio ; vi = princeps . letters refer to hancock ' s subgenera : a = pyrrhosticta ; b = pterourus ; c = sinoprinceps ; d = druryia ; e = princeps ; f = menelaides . single asterisk indicates inclusion of papilio alexanor as per hancock ( 1983 ) . double asterisks indicate exclusion of p . alexanor based on our data .
— taxobox name = tawny mime image width = 200px regnum = animalia phylum = arthropoda classis = insecta ordo = lepidoptera familia = papilionidae genus = papilio subgenus = chilasa species = c . agestor binomial = chilasa agestor binomial authority …
older classifications of the swallowtails tended to use a large number of rather small genera . more recent classifications have been more conservative , and as a result a number of former genera are now absorbed within papilio . the genus as recognized by modern systems has about 200 members . the genus chilasa is regarded as a subgenus of papilio by some workers , as are the baggy - tailed swallowtails ( agehana ) , although the latter taxon is usually considered a subgenus of chilasa .
paup * 4 . 0b8 - b10 ( swofford , 1998 ) was used for all parsimony , ml , bootstrap , and decay analyses . to get an accurate estimation of phylogenetic relationships in papilio sensu lato , we followed the following strategies .
papilio memnon and p . rumanzovia are sister species with allopatric distributions ( collins and morris , 1985 ) . papilio memnon occurs widely in southeast asia , from india to southern japan and indonesia , but not in the philippines . papilio rumanzovia has a more restricted geographic range and occurs in the philippines ( except palawan ) and on batu , talaud , and sangihe islands . around 5 mya , the philippine platform approached its present position in the immediate vicinity of east malaysia and indonesia ( hall , 1996 , 2001 ) , allowing the ancestral species to expand its range and eventually evolve into two distinct species . based on this assumption , we constrained the maximum age of the p . memnon – p . rumanzovia split at 5 my .
successful hand - pairing of swallowtail butterflies by clarke ( 1952 ) gave rise to studies of experimental hybridization in papilio ( e . g . , clarke and sheppard , 1955 , 1956 , 1957 ; ae , 1960 , 1962 , 1990 ; remington , 1960 ) . data on egg viability , sex ratio in hybrid progeny , and fertility of f 1 ' s were summarized in the form of a biological incompatibility index used to evaluate papilio relationships ( ae , 1979 , 1995 ; scriber , 1995a ) .
average values of uncorrected percentage sequence divergence in two mitochondrial genes and one nuclear gene compared at different taxonomic levels within the papilionidae . three putative genera in papilio represent heraclides , pterourus + chilasa + papilio alexanor , and princeps + eleppone + princeps sensu stricto . twelve putative genera constitute all major lineages from figure 1 as a , b , c , d , e , f , g , h + i , j , k , l , and m . solid bars indicate 95 % confidence interval ; dashed bars show ranges of values .
in their laval form , members of papilio typically feed upon plants of the rutaceae including common ornamental and agriculturally important species such as citrus spp . , murraya spp . , choisya spp . and calodendrum spp . caterpillars sequester terpenoids from their diet to produce a foul smelling oil used in defence .
maximum parsimony tree from combined equally weighted data . numbers above branches indicate bootstrap support ( values > 50 shown ) . bremer support ( decay index ) is given under branches for nodes within papilio sensu lato . circled letters label subclades and nodes to indicate terminal nodes on trees in figure 2 .
evgueni v . zakharov , michael s . caterino , felix a . h . sperling ; molecular phylogeny , historical biogeography , and divergence time estimates for swallowtail butterflies of the genus papilio ( lepidoptera : papilionidae ) , systematic biology , volume 53 , issue 2 , 1 april 2004 , pages 193–215 , urltoken
mitochondrial ( coi - coii ) plus nuclear ( ef - 1 α ) dna sequence data have provided considerable new resolution for the phylogeny of papilio , a large and important genus for research in ecology , genetics , and evolutionary and conservation biology . this phylogeny provides a number of insights into the evolutionary history of the group .
our most significant taxonomic finding is the strong placement of papilio esperanza . several other researchers , although recognizing some uncertainty , concluded that this species was a member of heraclides ( hancock , 1983 ) . we find strong support for the countervailing position , that the species is instead related to ( and in our analysis basal to ) the troilus species group ( see collins and morris , 1985 ) .
based on the possible sister relationship of papilio and meandrusa , whose larvae develop on lauraceae ( igarashi , 1984 ) , it was hypothesized that use of these plants should be regarded as a primitive character ( aubert et al . , 1999 ) . based on this assumption , there were at least two shifts to rutaceae , once in heraclides and once in the old world lineage of papilio , with subsequent shifts in the p . machaon group to apiaceae and in some african princeps back to lauraceae or close relatives ( ackery et al . , 1995 ) . however , the reliability of morphological characters that were used to justify the affinity of meandrusa to the tribe papilionini is questionable ( haüser , 1993 ) , and molecular evidence for this relationship ( aubert et al . , 1999 ) is tentative .
heuristic searches performed for three data sets under estimated parameters produced the trees shown in figure 2 . analyses based on only mtdna data produced a tree with the same relationships as illustrated in the mp bootstrap consensus tree in princeps and papilio sensu stricto but with alternative groupings in the heraclides + pterourus + chilasa clade . analysis from the ef - 1 α data set resulted in a tree with several polytomies .
both independent analyses for coi - coii reached stationarity well before 300 , 000 cycles . we discarded 3 , 000 trees as burn - in samples and computed the consensus tree from the remaining 14 , 000 trees . the branching pattern is identical to that of the ml tree and resembles the mp tree based on only mtdna data ( fig . 2 ) , where heraclides ( see fig . 1 ) is a sister group to the rest of papilio . also , as in the ml tree , pterourus appeared to be paraphyletic with respect to papilio alexanor + chilasa , but there is low support for this relationship . many nodes on the tree are well supported , and the overall posterior probability of the tree was 0 . 93 . of the 53 total nodes , 32 had a significance level of 1 . 0 and 42 had support > 0 . 9 .
phylogenetic tree inferred by ml from the combined data set with a reduced number of outgroups . the identical topology with similar branch lengths resulted from bayesian analysis of the same data . numbers above branches indicate bootstrap support counted in minimum evolution analysis under the same model of substitution used in ml analysis ( values > 50 shown ) . numbers under the nodes indicate bayesian posterior probabilities . vertical lines beside taxon names correspond to papilio sections sensu munroe ( 1961 ) .
bayesian analyses performed with two independent runs on the ef - 1 α data subset yielded only 2 , 000 burn - in trees per run . a consensus tree was constructed from the remaining 16 , 000 trees and resulted in a resolution very similar to that of the tree from the corresponding partitioned ml analysis ( fig . 2 ) . the overall probability of the tree was only 0 . 85 , and many nodes reflecting phylogenetic relationships between major lineages in the princeps + papilio ( sensu stricto ) clade had lower posterior probabilities .
character evolution optimized on the phylogeny of papilio is illustrated in figure 4 . based on the inferred phylogeny , the available morphological and ecological data , and pachliopta neptunus ( a member of troidini ) as the sister taxon , mp reconstructions indicated that the ancestral states for the selected characters were larval feeding on rutaceae , nonmimetic wing pattern , lack of iridescent patches on wings , lack of female secretion deposits on egg surface , straight larval minute body hairs , one row of crochets on larval prolegs , and lack of a silk pad on the larval resting site ( table 5 ) .
larvae of pachliopta neptunus and all other troidini , a generally accepted sister group of papilionini , develop on species of aristolochiaceae , whereas the primary hosts for leptocircini ( the sister group for troidini and papilionini ) are annonaceae . some species of leptocircini also develop on lauraceae , hernandiaceae , and rosaceae . however , according to a recently published molecular phylogeny of the genus graphium ( makita et al . , 2003 ) , basal lineages of leptocircini are those whose larva develop on hernandiaceae and rosaceae . thus , none of the presumed close relatives of the genus papilio indicate lauraceae as the ancestral host plants .
to evaluate the effect of different outgroup and ingroup combinations , we performed several mp analyses for the combined data set : ( 1 ) with pieris napi as an outgroup , and the 70 remaining taxa as the ingroup ; ( 2 ) with pieris napi excluded from the analysis , two sequences from baronia brevicornis as an outgroup , and the 68 remaining taxa in the ingroup ; and ( 3 ) with only papilio sensu lato in the ingroup , two outgroups ( pachliopta neptunus and eurytides marcellus ) used in previous studies ( caterino and sperling , 1999 ; reed and sperling , 1999 ) , and all other taxa excluded from the analyses .
previous authors have outlined relatively concrete biogeographic scenarios for papilionini ( munroe , 1961 ; hancock , 1983 ; miller , 1987 ; scriber , 1995b ) . the ancestor of the papilionini , from a center of origin in eastern north america , is considered to have dispersed first to the palearctic and later to south america before the end of the cretaceous , leaving pterourus in north america . the palearctic ancestor spread to china ( and subsequently to southeast asia , africa , and australia ) as princeps , leaving papilio sensu stricto in western asia . the south american ancestor spread via gondwanaland to australia as eleppone and from there to southeast asia as chilasa , leaving heraclides in south america .
modeltest supported use of the gtr model ( lanave et al . , 1984 ) , with invariable sites ( i ) and gamma - distributed rates ( γ ) , as the best fit for all data sets ( including the combined data set ) except for a reduced ef - 1 α data set ( papilio sensu lato plus two outgroup taxa ) , for which the best model was trn ( tamura and nei , 1993 ) + i + γ . estimated parameters used in ml analyses are given in table 3 . the c - t substitution rate was substantially higher than other substitution types , and substitution rates for a - c and c - g were 2–2 . 5 times higher for the 71 - taxon data set within the coi - coii partition .
london : taylor and francis ; calcutta and simla , thacker , spink , & co . ; [ etc . , etc . ]
evidence reported by james - hixon for item butterflies02bingiala on november 18 , 2006 : no visible notice of copyright ; stated date is 1907 .
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hewitson , 1862 – lesser mime . kunte , k . , s . sondhi , and p . roy ( chief editors ) .
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the body ( abdomen ) is black . thorax and head are also black . all parts of the body have white spots .
the first description of this butterfly was in 1864 by hewitson . there are five subspecies .
cites : ( convention on international trade in endangered species of wild fauna and flora ) : - no entry - ( as at 12 . 02 . 2008 )
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this species is found in rainforest at elevations between sea level and about 800m .
males commonly imbibe mineralised moisture from wet ground . they are sometimes encountered singly , but more often several will be present in mixed aggregations .
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♂ , 1992 . 03 . 05 . wiang pa pao , chiang rai , thailand .
♀ , 1987 . 02 . 21 . doi suthep , chiang mai , thailand .
♂ , 1990 . 02 . 06 . doi suthep , chiang mai , thailand .
♂ , 1992 . 01 . 30 . ban ten doi , mae hong son , thailand .
♂ ab , 1990 . 02 . - - . doi pha hom pok , chiang mai , thailand .
♀ , 1989 . 02 . 26 . chiang dao , chiang mai , thailand .
♂ , 2006 . 02 . 27 . nam cha , xaysomboun , laos . ( photo by a . m . cotton )
♂ , 2006 . 02 . 14 . nam phao , lak sao , laos . ( photo by a . m . cotton )
♂ , 2007 . 02 . 19 . nam phao , lak sao , laos . ( photo by a . m . cotton )
♂ , 2006 . 03 . - - sapa , vietnam . ( photo by a . m . cotton )
♀ , 2006 . 03 . 27 . phu phan , xam neua , laos . ( photo by a . m . cotton )
♀ , 2007 . 03 . 05 . nam phao , lak sao , laos . ( photo by a . m . cotton )
♂ , 2006 . 03 . 05 . nam phao , lak sao , laos . ( photo by a . m . cotton )
♂ , 2007 . 02 . 20 . nam phao , lak sao , laos . ( photo by a . m . cotton )
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urltoken
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urn : lsid : catalogueoflife . org : taxon : 322c1faa - 4e2c - 11e8 - 9ed0 - fa163e792e6e : col20180626
urn : lsid : catalogueoflife . org : taxon : 322c25e1 - 4e2c - 11e8 - 9ed0 - fa163e792e6e : col20180626
urn : lsid : catalogueoflife . org : taxon : 322c277b - 4e2c - 11e8 - 9ed0 - fa163e792e6e : col20180626
urn : lsid : catalogueoflife . org : taxon : 322dd07d - 4e2c - 11e8 - 9ed0 - fa163e792e6e : col20180626
urn : lsid : catalogueoflife . org : taxon : 322de4fe - 4e2c - 11e8 - 9ed0 - fa163e792e6e : col20180626
urn : lsid : catalogueoflife . org : taxon : 322de75d - 4e2c - 11e8 - 9ed0 - fa163e792e6e : col20180626
urn : lsid : catalogueoflife . org : taxon : 33051f4f - 4e2c - 11e8 - 9ed0 - fa163e792e6e : col20180626
häuser c . , holstein j . , kroupa a . s . , steiner a . & turiault m . ( eds ) . ( 2018 ) . globis ( gart ) : global butterfly information system ( version sep 2013 ) . in : roskov y . , abucay l . , orrell t . , nicolson d . , bailly n . , kirk p . m . , bourgoin t . , dewalt r . e . , decock w . , de wever a . , nieukerken e . van , zarucchi j . , penev l . , eds . ( 2018 ) . species 2000 & itis catalogue of life , 30th june 2018 . digital resource at urltoken species 2000 : naturalis , leiden , the netherlands . issn 2405 - 8858 .
urn : lsid : catalogueoflife . org : taxon : 40902927 - 4e2c - 11e8 - 9ed0 - fa163e792e6e : col20180626
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is a high - altitude butterfly found in india . it is . . .
linnaeus , 1758 – common mime . kunte , k . , s . sondhi , and p . roy ( chief editors ) .
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click on the links above or use the thumbnails below to access full size images and detailed articles about the lifecycle , habitats and biology of each species . alternatively browse using the scientific names via the oriental region
♂ , 2007 . 02 . 11 . dambri , lam dong , vietnam . ( photo by k . saito )
on : chilasa imitata monastyrskii & devyatkin , 2003 od : atalanta 34 ( 1 / 2 ) : 77 - 78 , pl . 5 , figs . 1♂ , 2♂ ( un ) . tl : c . vietnam . ( bmnh ) distribution : c . vietnam .
holotype ♂ , bi doup - nui ba nature reserve , dong mong , lam dong , c . vietnam . 2002 . 04 . 16 ( bmnh ) paratypes 4♂ bi doup - nui ba nature reserve , dong mong , lam dong , c . vietnam . 2002 . 04 . 16 ( msu ) paratype 1♀ , bi doup - nui ba nature reserve , dong mong , lam dong , c . vietnam . 2002 . 04 . 05 paratype 1♀ , dien khanh , khanh hoa , c . vietnam . 2003 . 04 . 06
atalanta ( augst 2003 ) 34 ( 1 / 2 ) : 75 - 109 , 5 col . pls .
linnaeus , 1758 – paris peacock . kunte , k . , s . sondhi , and p . roy ( chief editors ) .
the butterflies of north america , james a . scott , isbn 0 - 8047 - 1205 - 0 , 1986
2 st ( 1 ♂ , 1 ♀ , ex coll . l . de nicéville , coll . h . fruhstorfer )
cramer , 1775 – spangle . kunte , k . , s . sondhi , and p . roy ( chief editors ) .
kunte , k . , s . sondhi , and p . roy ( chief editors ) .
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felder & felder , 1864 – redbreast . kunte , k . , s . sondhi , and p . roy ( chief editors ) .
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the utility of dna sequence comparisons for inferring phylogenetic relationships has been demonstrated thoroughly ( e . g . , hasegawa and yano , 1984 ; gielly and taberlet , 1994 ; simon et al . , 1994 ) . however , their use for dating divergences has been much more controversial . early hopes for a molecular clock ( zuckerkandl and pauling , 1962 , 1965 ) via evolution in accordance with neutral theory ( kimura , 1983 ) have generally not been satisfied . rather , most studies have revealed substantial variation in rates of evolution across genes and lineages ( britten , 1986 ; gillespie , 1986 ; avise , 1994 ; li , 1997 ; pawlowski et al . , 1997 ; page and holmes , 1998 ; hebert et al . , 2002 ; soltis et al . , 2002 ) , and methods of divergence time estimation must account for this variation ( rambaut and bromham , 1998 ; thorne et al . , 1998 ; sanderson , 2002 ) .
note : species groups shown in bold are represented in this study ( by the species listed in brackets ) .
sequences of coi - coii genes were aligned against the published sequence from drosophila yakuba ( clary and wolstenholme , 1985 ) with multiple sequence alignment using clustalx 1 . 81 ( thompson et al . , 1997 ) with the default settings ( gap opening = 10 , gap extension = 0 . 20 ) followed by adjustment by eye . ef - 1 α gene sequences did not contain any introns and were aligned against sequence from bombyx mori ( kamiie et al . , 1993 ) by eye with the aid of seal 2 . 0 ( rambault , 2002 ) . to determine codon positions , we used macclade 4 . 0 ( maddison and maddison , 2000 ) .
to obtain sds for estimated divergence times , the data set was bootstrapped 100 times using the seqboot module from phylip 3 . 6 ( felsenstein , 1989 ) , and branch lengths were reestimated for each node under the constrained initial topology in paup * . the dating analyses were then repeated for each tree , and node statistics were summarized using the program r8s .
the next calibration point was selected for another species with a highly restricted distribution , p . hospiton , which is endemic to the islands corsica and sardinia . the separation to the corsica–sardinia microplate from the iberian peninsula is consistently dated at approximately 29 mya ( alvarez , 1972 ) , although this date seems to be too old and rather implausible to explain endemism of p . hospiton . a more recent event , recession of the mediterranean sea for about 1500 years and successive reisolation of the islands by the waters from the atlantic ocean , took place about 5 mya ( hsu , 1972 ) and provides a more reasonable date to constrain the split between p . hospiton and p . machaon .
to obtain estimates for a possible range of divergence dates , the selected calibration points were applied in different combinations . we fixed the time of the tree root at 82 . 5 mya ( first ) and 89 . 1 mya ( second ) , with no other constraints enforced . then we applied all discussed constraints for internal nodes without constraining the time of the root . we then applied all age constraints at the same time .
alignment of ef - 1 α sequences was unambiguous because of the absence of indels . total length of the aligned ef - 1 α region was 995 bp . some length differences were found in the mtdna sequence , primarily in the coi initiation region ( caterino and sperling , 1999 ) . seventeen sites here were deleted from the alignment , including six nucleotides corresponding to the first two codon positions of the coi gene ( clary and wolstenholme , 1985 ; caterino and sperling , 1999 ) . as caterino and sperling ( 1999 ) noted , a few taxa demonstrated 1 - bp indels ( mostly phylogenetically uninformative ) in the trna - leucine gene . there were also a 3 - bp insertion immediately following the coi termination codon in p . zelicaon and an insertion of one codon ( aat : arginine ) in p . dardanus between positions equivalent to 3 , 471 and 3 , 472 of drosophila yakuba . we found that p . oribazus had a 3 - bp insertion ( aaa : lysine ) in the same position as in p . dardanus . the final aligned sequences included 2 , 293 nucleotides for the mtdna partition and 3 , 288 nucleotides in the combined data set . previously published ef - 1 α sequence for atrophaneura alcinous ( caterino et al . , 2001 ) was determined to be incorrect at the 5′ end and has been updated for this study and in genbank .
no base composition heterogeneity was found for any gene partition among ingroup taxa ( coi : χ 2 = 61 . 6 , df = 159 , p = 1 . 0 ; coii : χ 2 = 56 . 6 , df = 159 , p = 1 . 0 ; ef - 1 α : χ 2 = 30 . 7 , df = 159 , p = 1 . 0 ) and across all species including outgroup taxa ( coi : χ 2 = 103 . 2 , df = 213 , p = 1 . 0 ; coii : χ 2 = 74 . 6 , df = 213 , p = 1 . 0 ; ef - 1 α : χ 2 = 124 . 9 , df = 213 , p = 0 . 99 ) . however , after all noninformative characters were excluded , base composition heterogeneity was revealed in the coi partition both for ingroup only ( χ 2 = 237 . 9 , df = 159 , p < 0 . 005 ) and for the full set of taxa ( χ 2 = 390 . 7 , df = 213 , p < 0 . 005 ) . however , analysis of the corresponding mp tree did not reveal any grouping of species with similar nucleotide frequencies . analysis of partitioned and combined data using logdet distances resulted in trees with topologies almost identical to those of the corresponding ml trees .
based on the results of the ild test , partitions of the data into coi , coii , and ef - 1 α were homogeneous ( sum of gene tree length = 8 , 407 ; p = 0 . 114 ) . coi and coii gene partitions alone were homogeneous ( sum of gene tree length = 6 , 789 ; p = 0 . 492 ) and partitions between coi and coii together and ef - 1 α were also homogeneous ( sum of tree lengths = 8 , 505 ; p = 0 . 774 ) . we analyzed both partitioned and combined data to gain insight into any distinctively different phylogenetic results due to data partitions and to obtain a phylogenetic reconstruction based on a maximum number of informative characters . in all cases , when all the sequences were combined for the analysis resolution and node support in the tree improved markedly . many other studies have demonstrated improved resolution and increased bootstrap supports in combined analyses ( e . g . , baker and desalle , 1997 ; vogler and welsh , 1997 ; crespi et al . , 1998 ; remsen and desalle , 1998 ; klompen et al . , 2000 ; chapco et al . , 2001 ) .
the total number of informative characters for the combined data set with all outgroups was 1 , 129 ( 34 . 3 % ) , with 847 sites in the mtdna partition and 282 sites in the ef - 1 α data set . number of informative characters was greatest for third - codon positions ( 601 sites [ 71 % of all informative characters ] for coi - coii and 260 sites [ 92 . 2 % ] for ef - 1 α ) . with all outgroups excluded , number of informative characters was reduced to 933 ( 28 . 4 % ) , with 725 informative sites in the mtdna partition and 208 sites in the ef - 1 α data set . proportion of informative characters for third - codon positions increased and was 75 % ( 547 sites ) of all informative characters for coi - coii and 94 % ( 196 sites ) for ef - 1 α .
γ = α , estimated shape parameter ; i = proportion of invariable sites .
the ml analysis for combined data revealed a single tree with a negative log - likelihood score ( –ln l ) of 31023 . 619 that was identical to the combined mp bootstrap consensus tree ( fig . 2 ) . in the expanded ml tree for combined data that shows all species ( fig . 3 ) , grouping within the major terminal subclades ( corresponding to species groups and subgenera ) is usually congruent with the relationships inferred by the mp combined data tree ( fig . 1 ) . however , relationships among many major clades remain weakly supported .
three independent analyses for the combined data set converged on similar log - likelihood scores and reached stationarity before generation 300 , 000 . for the coi - coii data , the initial 3 , 000 trees from each analysis were discarded . a consensus tree was constructed from the combined set of 21 , 000 trees ( fig . 3 ) . the branching pattern of the tree is completely identical to that of the mp bootstrap consensus and ml trees from combined analyses . the average posterior probability for the inferred phylogeny was 0 . 97 . thirty - eight ingroup nodes ( of 53 ) had posterior probabilities of 1 . 0 , and 46 nodes were supported with significance level > 0 . 9 . bayesian analyses supplied higher values for posterior probabilities compared with bootstrap supports from ml and mp analyses .
likelihood ratio tests rejected clocklike behavior of sequences ( p < 0 . 001 ) for combined and partitioned ( coi - coii vs . ef - 1 α ) data and also for full ( 71 taxa ) and reduced ( 56 taxa ) taxon sets . tests for all six permutations of partitions ( three ) versus taxon sets ( two ) gave p values of ≪ 0 . 001 . application of the selected calibration points in the penalized likelihood procedure provided us with a range of dates ( with sds ) . initial results were obtained with the default settings for dating analyses in the r8s program , with the cross - validation function enforced . the rate smoothing parameter with the lowest cross - validation scores was selected , and the dating procedure was then repeated . age estimates ( with sds ) for all internal nodes numbered in figure 4 are shown in table 4 .
no age estimation could be obtained for nodes that were collapsed in partitioned phylogenies .
estimates of marginal ancestral state probabilities are given as character state ( negative log likelihood ; proportion of a total likelihood ) .
another significant result is the strongly supported placement of p . anactus at the base of part of the princeps lineage . this species had previously been placed in its own subgenus , close to chilasa ( in the other major clade of papilionini ) ( hancock , 1979 ) . the placement was based in part on shared mimicry , although the specific mimetic patterns differ as do larval morphology and host plants . larvae of chilasa develop on lauraceae and magnoliaceae , whereas those of p . anactus , like members of princeps , feed on rutaceae ( hancock , 1979 ) .
species with curved minute hairs on the body of the last instar larvae form a monophyletic group , as was suggested by igarashi ( 1984 ) , with two exceptions : the p . demoleus group and p . nobilis . more species of princeps should be studied to clarify the evolution of this trait .
mitochondrial dna sequence variation and phylogeography of oceanic insects ( hemiptera : gerridae : halobates spp . )
paup * : phylogenetic analysis using parsimony ( * and other methods ) , version 4 . 0
molecular phylogeny of butterflies parnassius glacialis and p . stubbendorfii at various localities in east asia
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, moore . — very closely resembles the typical form but can always be distinguished by the presence of a complete series of postdiscal bluish - grey spots on the upperside of the hind wing . it is generally also smaller in both sexes than typical
, and on the upperside of the bind wing the groundcolour in the anterior interspaces is broadly centred along the middle with very dark brown , almost black . the female frequently has the ground - colour of the hind wing a much brighter chestnut ( almost ochraceous ) than in any specimen of female of
the butterfly is found in pakistan from west kashmir to sikkim and right across from assam to myanmar ( including nepal , bhutan and bangladesh ) , thailand , vietnam , laos , kampuchea , central and southern china and peninsular malaysia .
never common but not known to be threatened across most of its range . extremely local . considered to be vulnerable in peninsular malaysia .
this butterfly frequents woods in hilly regions and generally flies between 4000 to 8500 ft in the himalayas and assam .
the tawny mime flies slowly in the sunshine in glades and clearings of oak ( quercus ) forests usuually 10 to 12 feet above the ground . it is known to patrol a beat and respond aggressively to passing butterflies and insects for long periods in a locality . it does not visit flowers .
the female lays her eggs on the young leaves of machilus odorattisimus trees , natural order lauraceae , about the end of april . the larva is at first of a reddish colour but very soon turns black and white , and lies on the upper surface of a leaf where it greatly and protectively resembles a bird ' s droppings .
adult larva : brown with two subdorsal and two lateral rows of fleshy - pointed tubercles , each with a spot of red at its base ; anterior , middle and posterior lateral patches of dull ochraceous , the latter two meeting on the dorsum ; the rest of tho larva spotted with black and red .
pupa dark brown with lines of dull ochraceous , roughened , with the anal end truncate .
( described from plate to mackinnon & de niceville ' s paper . )
^ bingham , c . t . 1907 . fauna of british india . butterflies . volume 2
collins , n . m . , morris , m . g . ( 1985 ) threatened swallowtail butterflies of the world . iucn . isbn 2 - 88032 - 603 - 6
gay t . , kehimkar i . & punetha j . c . , ( 1992 ) common butterflies of india . wwf - india and oxford university press . mumbai . india .
haribal , meena . , ( 1992 ) the butterflies of sikkim himalaya . sikkim nature conservation foundation , gangtok
kunte , krushnamegh ( 2000 ) butterflies of peninsular india . universities press , hyderabad . isbn 81 - 7371 - 354 - 5
wynter - blyth , m . a . , ( 1957 ) butterflies of the indian region , bombay natural history society , mumbai , india .
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= = reprezentanții principali ai genului = = * accept * armored saint * black sabbath * budgie * deep purple * diamond head * dio * iron maiden * judas priest * king diamond * led zeppelin * manowar * motörhead * ozzy osbourne * queensrÿche * raven * rainbow * saxon * scorpions * tank * u . d . o .
= = key figures and important bands of the genre = = the references testify the notability of the following bands in the genre : * accept * armored saint * black sabbath * budgie * deep purple * diamond head * dio * iron maiden * judas priest * king diamond * led zeppelin * manowar * motörhead * ozzy osbourne * queensrÿche * rainbow * raven * saxon * scorpions * tank * u . d . o .
marginalia — devil lore in ' the raven '
from poe studies vol .
revue neurologique
, 1964 , 110 , 553 - 57 ( english translation in critchley ,
the divine banquet of the brain and other essays
, new york : raven press , 1979 , pp .
two verse masterworks : ' the raven ' and ' ulalume '
, collected in
the cambridge companion to edgar allan poe
, edited by kevin j . hayes .
de la nord la sud : dealul lui taylor , dealul stansbury , dealul lui marye și dealul willis .
( although popularly known as marye ' s heights , the ridge was composed of several hills separated by ravines , from north to south : taylor ' s hill , stansbury hill , marye ' s hill , and willis hill .
*
jewish and christian apocalypses
( 1914 ) schweich lectures of the british academy 1913 *
the religion of the manichees
( 1925 ) donnellan lectures 1924 *
palestine in general history
( 1929 ) schweich lectures 1926 , with theodore h . robinson , j . w . hunkin *
christian worship
( 1930 ) *
the church of today part 2
( 1930 ) with p . gardner - smith and c . e . raven , the christian religion and its origin and progress , volume 3 *
jesus christ : an historical outline
( 1932 ) *
church and gnosis : a study of christian thought and speculation in the second century
( 1932 ) *
franciscan essays ii
( 1932 ) with h . e . goad and a . g . little *
the dura fragment of tatian
.
* digitalis , raven :
goth craft : the magickal side of dark culture
( 2007 : llewellyn worldwide ) —includes a lengthy explanation of gothic history , music , fashion , and proposes a link between mystic / magical spirituality and dark subcultures .
* povestirea apare pe albumul dublu
the raven
( 2003 ) al lui lou reed ' s .
* the story features as part of lou reed ' s 2003 double album
the raven
.
2 , december 1972 disponibil online * hirsch , david h .
the raven and the nightingale
as collected in
poe and his times : the artist and his milieu
, edited by benjamin franklin fisher iv .
2 , december 1972 available online * hirsch , david h .
the raven and the nightingale
as collected in
poe and his times : the artist and his milieu
, edited by benjamin franklin fisher iv .
= = rezumat = = în timp ce galleria ( raven - symoné ) este plecată în anglia pentru a urma cursurile universității cambridge , chanel ( adrienne bailon ) , dorinda ( sabrina bryan ) și aquanette ( kiely williams ) sunt invitate să joace într - un film indian , la bollywood .
= = plot = = with galleria at the university of cambridge in england , chanel ( adrienne bailon ) , dorinda ( sabrina bryan ) , and aqua ( kiely williams ) are cast in the lavish new bollywood movie
namaste bombay
.
credits - computer translations are provided by a combination of our statistical machine translator , google , microsoft , systran and worldlingo .
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