the heliconiinae , commonly called heliconians or longwings , are a subfamily of the brush-footed butterflies ( family nymphalidae ) .

they can be divided into 45 – 50 genera and were sometimes treated as a separate family heliconiidae within the papilionoidea .

the colouration is predominantly reddish and black , and though of varying wing shape , the forewings are always elongated tipwards , hence the common name .

most longwings are found in the tropics , particularly in south america ; only the argynnini are quite diverse in the holarctic .

especially tropical species feed on poisonous plants , characteristically passifloraceae vines , as larvae , becoming poisonous themselves .

the adult butterflies announce their acquired toxicity with strong aposematic colours , warning off would-be predators .

there are several famous cases of batesian and müllerian mimicry both within this group and with other butterflies .

other commonly seen food plants are fabaceae ( which also contain several toxic species ) , and particularly among northerly species of violaceae .

heliconiinae

heliconiinae <EOT> a generic revision of the heliconiinae ( lepidoptera , nymphalidae ) . american museum novitates ; no . 1197

phylogenetic relationships of butterflies of the tribe acraeini ( lepidoptera , nymphalidae , heliconiinae ) and the evolution of host plant use .

phylogenetic relationships of butterflies of the tribe acraeini ( lepidoptera , nymphalidae , heliconiinae ) and the evolution of host plant use . - pubmed - ncbi

heliconiinae ( nymphalidae ) ; vane - wright & de jong , 2003 , zool . verh . leiden 343 : 229 ; [ wahlberg ] ; [ nl4a ] , 261

argynnini ( heliconiinae ) ; vane - wright & de jong , 2003 , zool . verh . leiden 343 : 235 ; [ wahlberg ] ; [ nl4a ] , 261

penz , c . , and d . peggie . 2003 . phylogenetic relationships among heliconiinae genera based on morphology ( lepidoptera : nymphalidae ) . systematic entomology 28 : 451 - 479 .

acraeini ( heliconiinae ) ; vane - wright & de jong , 2003 , zool . verh . leiden 343 : 236 ; [ wahlberg ] ; [ afrl ] ; [ nl4a ] , 262

penz , c . m . & p . djunijanti . 2003 . phylogenetic relationships among heliconiinae genera based on morphology ( lepidoptera : nymphalidae ) . systematic entomology 28 : 451 - 479 . [ links ]

penz , c . m . 1999 . higher level phylogeny for the passion - vine butterflies ( nymphalidae , heliconiinae ) based on early stage and adult morphology . zoological journal of the linnean society 127 : 277 - 344 . [ links ]

lamas , g . 2004 . heliconiinae , p . 262 - 274 . in : j . b . heppner ( ed . ) . atlas of neotropical lepidoptera . gainesville , association for tropical lepidoptera , scientific publishers , vol . 5a , 439p . [ links ]

paluch , m . ; m . m . casagrande & o . h . h . mielke . 2005 . comportamento de agregação noturna dos machos de actinote surima surima ( schaus ) ( lepidoptera , heliconiinae , acraeini ) . revista brasileira de zoologia 22 : 410 - 418 . [ links ]

freitas , a . v . l . ; r . b . francini & t . s . souza . 2009b . immature stages and natural history of the threatened butterfly actinote quadra ( nymphalidae : heliconiinae : acraeini ) . tropical lepidoptera research 19 ( 2 ) : 82 - 88 . [ links ]

freitas , a . v . l . ; l . a . kaminski ; r . g . matos & k . l . silva - brandão . 2009a . immature stages of the andean butterfly actinote rufina ( nymphalidae : heliconiinae : acraeini ) . tropical lepidoptera research 19 ( 1 ) : 18 - 21 . [ links ]

the subfamily heliconiinae is composed of a diverse group of species distributed widely in the new and old world tropics , and it also includes the largely holarctic fritillaries ( argynnini ) . many species in heliconiini and acraeini sequester and / or synthesize cyanogenic glucosides , which render them unpalatable to predators . as a result , they act as müllerian mimics or models for palatable batesian species in other groups .

francini , r . b . ; a . v . l . freitas & k . s . brown jr . 2005 . rediscovery of actinote zikani ( d ' almeida ) ( nymphalidae , heliconiinae , acraeini ) : natural history , population biology and conservation of an endangered butterfly in se brazil . journal of the lepidopterists ' society 59 ( 3 ) : 134 - 142 . [ links ]

silva - brandão , k . l . ; n . wahlberg ; r . b . francini ; a . m . l . azeredo espin ; k . s . brown jr ; m . paluch ; d . c . lees & a . v . l . freitas . 2008 . phylogenetic relationship of butterflies of the tribe acraeini ( lepidoptera , nymphalidae , heliconiinae ) and the evolution of host plant use . molecular phylogenetics and evolution 46 : 515 - 531 . [ links ]

silva - brandão , k . l . , wahlberg , n . , francini , r . b . , azeredo - espin , a . m . l . , brown , k . s . j . , paluch , m . , lees , d . c . & freitas , a . v . l . 2008 . phylogenetic relationships of butterflies of the tribe acraeini ( lepidoptera , nymphalidae , heliconiinae ) and the evolution of host plant use . molecular phylogenetics and evolution 46 , 515 - 531 .

a higher level phylogeny for the passion - vine butterflies ( nymphalidae , heliconiinae ) was generated by cladistic analysis of 146 morphological characters from all life stages . the 24 species studied were selected representatives of the ten currently accepted genera of the sub - tribe heliconiiti . analyses of only characters from larvae and pupae did not produce well resolved trees . however , some characters of the immature stages provided critical support for the monophyly of two clades . analysis of only adult characters yielded a tree that closely resembled that obtained from all data combined . the phylogeny here derived from the combined analysis of early stage and adult characters differed in topology from all previously proposed hypotheses , and supported the monophyly of all currently recognized genera . characters supporting each clade are described and illustrated , and various hypotheses of phylogenetic relatedness of passion - vine butterfly taxa are discussed .

the tribe acraeini ( nymphalidae , heliconiinae ) is believed to comprise between one and seven genera , with the greatest diversity in africa . the genera abananote , altinote , and actinote ( s . str . ) are distributed in the neotropics , while the genera acraea , bematistes , miyana , and pardopsis have a palaeotropical distribution . the monotypic pardopsis use herbaceous plants of the family violaceae , acraea and bematistes feed selectively on plants with cyanoglycosides belonging to many plant families , but preferentially to passifloraceae , and all neotropical species with a known life cycle feed on asteraceae only . here , a molecular phylogeny is proposed for the butterflies of the tribe acraeini based on sequences of coi , ef - 1alpha and wgl . both maximum parsimony and bayesian analyses showed that the tribe is monophyletic , once the genus pardopsis is excluded , since it appears to be related to argynnini . the existing genus acraea is a paraphyletic group with regard to the south american genera , and the species of acraea belonging to the group of " old world actinote " is the sister group of the neotropical genera . the monophyly of south american clade is strongly supported , suggesting a single colonization event of south america . the new world actinote ( s . str . ) is monophyletic , and sister to abananote + altinote ( polyphyletic ) . based on the present results it was possible to propose a scenario for the evolution in host plant use within acraeini , mainly concerning the use of asteraceae by the south american genera .

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the root of the current tree connects the organisms featured in this tree to their containing group and the rest of the tree of life . the basal branching point in the tree represents the ancestor of the other groups in the tree . this ancestor diversified over time into several descendent subgroups , which are represented as internal nodes and terminal taxa to the right .

you can click on the root to travel down the tree of life all the way to the root of all life , and you can click on the names of descendent subgroups to travel up the tree of life all the way to individual species .

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this neotropical forest genus is characterized by expanded androconial areas on the costal margin of the hindwing in adult males .

etymology : turner 1976 named this new genus after the chilenian poet pablo neruda , who won the nobel prize for literature in 1971 .

turner jrg . 1976 . adaptive radiation and convergence in subdivisions of the butterfly genus heliconius ( lepidoptera : nymphalidae ) . zool . j . linn . soc . 58 : 297 - 308 .

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each tol branch page provides a synopsis of the characteristics of a group of organisms representing a branch of the tree of life . the major distinction between a branch and a leaf of the tree of life is that each branch can be further subdivided into descendent branches , that is , subgroups representing distinct genetic lineages .

for a more detailed explanation of the different tol page types , have a look at the structure of the tree of life page .

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brower , a . v . z . 2000 . phylogenetic relationships among the nymphalidae ( lepidoptera ) , inferred from partial sequences of the wingless gene . proceedings of the royal society of london series b biological sciences 267 : 1201 - 1211 .

freitas , a . v . l . , and k . s . j . brown . 2004 . phylogeny of the nymphalidae ( lepidoptera : papilionoidea ) . systematic biology 53 : 363 - 383 .

wahlberg , n . , e . weingartner , and s . nylin . 2003 . towards a better understanding of the higher systematics of nymphalidae ( lepidoptera : papilionoidea ) . molecular phylogenetics and evolution 28 : 473 - 484 .

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actinote discrepans photographed in rio grande do sul , brazil , in april 2010 .

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correspondence regarding this page should be directed to niklas wahlberg at and andrew v . z . brower at

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the nymphalidae is a family of several thousand species found in all zoogeographical regions of the world . most are medium or large in size , but the family is highly variable given that it also includes the satyrinae , a subfamily that has been designated as a family in its own right in earlier classifications .

the forelegs in both sexes are vestigial and useless for walking . in the male , there are typically only 2 tarsal joints and the legs have a brush - like appearance , resulting in a common name for this family - the " brush - footed butterflies " . the female foreleg has 4 tarsal joints which , when compared with the male , provides a mechanism of determining the sex of the adult . the midleg and hindleg are normal in both sexes and both tibia and tarsus may have spines .

the forewing always has 12 veins . given the variability of this family , there are no further distinguishing characteristics that apply to the family as a whole .

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silva - brandão kl 1 , wahlberg n , francini rb , azeredo - espin am , brown ks jr , paluch m , lees dc , freitas av .

depto . de zoologia , ib , universidade estadual de campinas , cp 6109 , cep 13083 - 970 , campinas , sp , brazil .

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american museum novitates novitates ( latin for " new acquaintances " ) , published continuously and numbered consecutively since 1921 , are short papers that contain descriptions of new forms and reports in zoology , paleontology , and geology . new numbers are published at irregular intervals .

department of library services american museum of natural history central park west at 79th st . , new york , ny 10024 © american museum of natural history , 2011

towards a better understanding of the higher systematics of nymphalidae ( lepidoptera : papilionoidea ) . - pubmed - ncbi

towards a better understanding of the higher systematics of nymphalidae ( lepidoptera : papilionoidea ) .

department of zoology , stockholm university , s - 106 91 stockholm , sweden . niklas . wahlberg @ urltoken

higher classification is mainly after [ wahlberg ] , november 2005 , except some clump / split of genera is not done yet .

the exact identification of these species is still unknown , but tentatively assumed to belong into this group .

checklist of afrotropical papilionoidea and hesperoidea ; compiled by mark c . williams , 7th ed . ( 2008 ) ( april 2007 ) ;

vane - wright & de jong , 2003 the butterflies of sulawesi : annotated checklist for a critical island faunda zool . verh . leiden 343 : 3 - 268 , pl . 1 - 16

if you have corrections , comments or information to add into these pages , just send mail to markku savela keep in mind that the taxonomic information is copied from various sources , and may include many inaccuracies . expert help is welcome .

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present address : department of biology , university of oregon , eugene or 97403 , u . s . a . e - mail : penz @ darkwing . uoregon . edu . from april 2000 : dept . invertebrate zoology , milwaukee public museum , 800 west wells street , milwaukee , wi 53233 , u . s . a .

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responsibility michael parsons . imprint san diego ; london : academic press , c1999 . physical description xvi , 736 p . , [ 172 ] p . of plates : ill . ( some col . ) , maps ; 30 cm .

zoologia ( curitiba , impr . ) vol . 28 no . 5 curitiba oct . 2011

i museo entomológico piedras blancas , comfenalco antioquia . medellín - colombia . e - mail : paduve12 @ urltoken ii grupo de entomología universidad de antioquia geua , universidad de antioquia . calle 67 , no . 53 - 108 , medellín - colombia , aa 1226 . e - mail : mwolff @ urltoken

the immature stages ( eggs , larvae and pupae ) , oviposition and larval behavior of altinote ozomene ( godart , 1819 ) are described here for the first time . larvae were reared from egg clutches collected from the host plants erato vulcanica ( klatt ) h . rob and munnozia senecionidis benth ( asteraceae ) . eggs were laid in groups on the undersides of leaves . the number of instars varied from five to eight within the same egg clutch , and the corresponding development time from larva to adult varied from 91 to 115 days . most ( 72 % ) larvae pupated during the sixth instar . the first four instars fed only on the leaf cuticle , whereas later instars consumed the whole leaf . larvae were gregarious during all instars but rested together only during the day in later instars , either hidden inside dry leaves , on the stem at the base of the host plants , or in the leaf litter . larvae showed similar morphology and behavior to those previously described for species of actinote hübner , 1819 from southeastern brazil and the andes .

key words : host plant ; immature behavior ; lepidoptera ; life cycle ; neotropical .

the nymphalid acraeini ( lepidoptera ) comprises between one and seven genera , depending on the taxonomy adopted , and occurs both in the neotropics and the old world , with the greatest diversity in africa ( silva - brandão et al . 2008 ) . three genera are found in the neotropics : actinote hübner , 1819 , abananote potts , 1943 and altinote potts , 1943 ( lamas 2004 ) . according to the most recent phylogeny of acraeini , which was proposed by silva - brandão et al . ( 2008 ) , altinote and abananote are embedded within actinote . several species , however , were not included in that study , including altinote ozomene ( godart , 1819 ) . given the current disarray in the systematics of acraeinae at the generic level , knowledge of the morphology and biology of the immature stages of the species in this subfamily could be of particular value in developing a stable classification ( harvey 1991 , penz 1999 , penz & peggie 2003 , freitas & brown 2004 , silva - brandão et al . 2008 ) .

most of the existing descriptions of the immature stages of neotropical acraeini involve members of actinote ( paluch et al . 1999 , 2001 , freitas et al . 2009a , b , 2010 ) . currently , no detailed descriptions exist for species of altinote , except for brief notes on altinote ozomene nox ( h . w . bates , 1864 ) ( sensu lamas 2004 , harvey 1983 , devries 1987 ) . the present paper thus provides the most complete description to date of the biology and morphology of the immature stages of an altinote species , focusing on al . ozomene .

the study was conducted between january 2006 and december 2009 in the parque ecológico piedras blancas nature reserve in the northeastern region of medellín , colombia ( 6º17 ' 40 . 68 " n , 75º30 ' 4 . 32 " w ) . the altitude at this site is 2350 m , and the average temperature is 15ºc . the area consists primarily of large pine plantations interspersed with small fragments of native forest dominated by quercus humboldtii bonpl ( fagaceae ) . observations of the behavior of the immature stages were conducted in the field . host plants were identified , collected and deposited in the herbarium of the universidad de antioquia ( hua ) .

eggs were collected in the field and taken to the laboratory where they were reared in plastic containers with host plant leaves . to determine the time spent in the embryonic phase ( from oviposition to eclosion of the eggs ) , five egg clusters were observed daily after oviposition . to determine the time spent in each larval instar , 4680 eggs from 39 egg clusters collected in the field were monitored daily from the hatching of larvae to the emergence of adults .

the samples of first instar larvae were placed in acetic acid for 48 hours , and the samples of other instars were fixed in kahle solution ( borror & delong 1971 ) and then transferred to 80 % ethyl alcohol . adults were mounted on entomological pins and deposited in the museo entomológico de piedras blancas ( mepb ) and in the laboratorio de colecciones entomológicas , universidad de antioquia ( ceua ) . in the following descriptions , t1 - t3 refer to thorax segments and a1a10 to abdominal segments . chaetotaxy of the first instar is described using the terminology of stehr ( 1987 ) . body length was measured at each stage using a millimeter rule under a stereomicroscope .

eggs of al . ozomene were collected from leaves of the asteraceae plants erato vulcanica ( klatt ) h . rob ( fig . 1 ) and munnozia senecionidis benth ( fig . 2 ; velez et al . 2008 ) . the former is a shrub , 4 m in height , with a 5 - cm diameter stem ; the latter has a prostrate habit of growth . both species are abundant in the study area , especially in open areas , scrub and forest edges and along stream margins , paths and tracks ( toro 2000 ) .

eggs were laid in clusters on the undersides of leaves . females typically alighted on the underside of the leaf around noon and remained there for over 2 . 5 hours , laying eggs ( fig . 3 ) . eggs and / or larvae were often found concentrated on plants within close proximity . eggs were sometimes laid on plants where eggs or larvae were already present ( up to two egg clutches were found on the same plant ) , generally on different leaves on large plants , and rarely on the same leaf . the number of eggs in each clutch varied from 100 to 380 ( n = 39 clutches ) . several clutches included four or five infertile eggs ( fig . 6 ) . larvae from clutches that contained fewer than 30 viable eggs ( either because oviposition was interrupted or because most of the eggs died from fungal attack ) died within the first three instars .

eggs from a single clutch hatched within 24 hours of one another . in 92 % ( 36 ) of the clutches , the number of instars varied within the same clutch from five to eight ( tab . i ) . in each of the other three clutches , all larvae showed the same number of instars : five in one clutch and six in the other two clutches . most of the larvae that reached adulthood ( 72 % , n = 349 ) pupated during the sixth instar , and 17 % pupated during the seventh instar ( tab . i ) .

the total developmental time was directly related to the number of molts : the time from egg to adult was 91 , 97 , 110 and 115 days for larvae that passed through five to eight instars , respectively ( tab . i ) . this is related to the fact that all instars are of similar duration ( 8 - 10 days ) , except for the final instar ( whether it is the fifth or the eighth ) , which typically lasts 13 - 14 days ( tab . i ) . however , a few larvae that passed through more than five instars pupated after only two or three days in their last instar . no relationship was observed between either the number of instars or the total developmental time and the sex of the adult butterfly . after emerging from pupae , the adults spend two or three days hanging , eliminating meconium , before they fly .

eggs ( figs 4 - 5 ) of al . ozomene are 0 . 6 - 0 . 8 mm in diameter , 0 . 9 - 1 . 1 in height ( n = 14 ) and barrel shaped , with a flat base adhered to the leaf and a flat apical region . the micropyle region resembles a rosette , with tiny ridges surrounded by a ring of polygonal cells , and a similar ring is present in the preapical area . slightly elevated longitudinal ridges run from the apex to the base , varying in number from 17 to 22 ( n = 16 ) and separated by slight grooves . poorly defined transverse ridges are also observed . eggs are ivory - colored from oviposition until the day preceding eclosion , when they become translucent , with the dark - colored head of the larva visible in the apical region . in many cases , the apical region of the egg was observed to be covered with a drop of viscous , transparent liquid throughout the embryonic period . eggs without this liquid were observed to develop a fungus .

first instar larvae ( figs 6 - 12 ) have a round , smooth head without scoli and with golden setae . the body is cylindrical , without scoli and with simple , dark brown setae arising from sclerotized brown pinacula standing out slightly from the skin . the head is black , and the body is ivory - colored immediately after hatching , becoming greenish - white the next day , once food is observable through the translucent skin . the prothoracic shield and anal shield are dark brown . the thorax becomes brown or chestnut and is darkest in t1 and fades through t3 . a1 is brown or dark yellowish ochre , and a2 is similar but has diffuse brown dorsolateral marks . the rest of abdomen is pale , clear yellow or greenish - yellow due to food showing through the skin .

chaetotaxy is illustrated in figures 10 - 12 . d1 and d2 are present in all body segments . d1 is posterior and dorsal to d2 in t1 and anterior and dorsal to d2 in the remaining segments . d1 is shorter in a1 - a7 than in a8 - a9 . sd2 is absent in a1 - a9 . l2 is absent in t2 , t3 and a9 , and l3 is absent in all segments . sv1 is present in all segments . sv2 is absent in a1 , a7 , a8 and a9 . sv1 and sv2 originate on the same sclerotized plate in a2 and are located on the lateral shield , outside of the prolegs , in a3 - a6 , with sv2 being anterior and shorter and sv1 lateral and longer . sv3 is found on t1 and a10 and sometimes on t2 and / or t3 . v1 is present on a1 , a2 and a10 . the coxae of the thoracic legs are surrounded by a laterally open , chitinous horseshoe - shaped plate with five setae : one anterior , one posterior and three setae on the internal mesal margin . exv1 is located very close to the midline in a10 , and the setae on both sides of the body originate on the same plate . the spiracles are circular and are larger on t1 and a8 , with protruding edges .

second instar larvae ( fig . 13 ) exhibit a shiny black head with thin pale setae , a translucent white clypeus , a brown labrum and a brown mandible . the thorax and a1 are generally brown , and the other abdominal segments are creamy yellow , with or without diffuse brown marks . on a few individuals , the abdomen is dark brown . from the second instar on , the larvae have ivory - colored infraspiracular bands that generally extend from t1 to a10 , and the subventral and ventral regions of the abdomen are creamy yellow . scoli are generally brown on the thorax and on a1 , and the abdominal infraspiracular scoli are creamy yellow . the other scoli vary from brown to creamy yellow . the scoli setae range from chestnut and creamy yellow to brown .

the head is smooth , without scoli , spines or chalazae . a pair of dorsal scoli is arranged linearly on each segment except for a10 , where it is absent . the t1 scoli are slightly longer than the remaining thoracic scoli , and the abdominal scoli are uniform in size and slightly smaller than those of t3 . from a1 to a8 , a second line of supraspiracular scoli appears below the dorsal scoli , each slightly anterior to a spiracle . one scolus is present adjacent to the anal shield on a10 , in line with the other supraspiracular scoli on the abdomen . in the lateral region of the thorax , one scolus appears between t1 and t2 and another between t2 and t3 ; both of these are in line with the abdominal spiracles . a third line of infraspiracular scoli that are posterior to the spiracles also appears on a1 to a8 . these scoli are the same size as the supraspiracular scoli and slightly shorter than the dorsal scoli on the abdomen .

the chitinous prothoracic shield is shiny black and almost oval in shape , with two setae that are longer than the dorsal scoli of the same segment . the anal shield is black , round and convex . chitinous plate on t1 - t3 and a9 , located below dorsal scolus : on t1 above spiracle , almost triangular in shape , with several setae , on t2 , t3 and a9 rounded in shape ; very small plate under these : on t1 anterior and opposite spiracle , almost square in shape , with several setae , on t2 and t3 , posterior to lateral scoli , rounded in shape . only the plates on t1 protrude from the skin . subventral , elliptic verruca with numerous setae are present above each thoracic leg , and these are arched in t2 and t3 . several ventral setae are clustered together on a1 , a2 , a7 and a8 , with one seta in each cluster that is longer than the others . the circular spiracles are the same size on t1 and a8 and larger on the remaining segments . the spiracle on t1 is slightly more ventrally located .

the chitinous plate surrounding the coxa of each thoracic leg is horseshoe - shaped , outwardly open , and of the same dark brown or gray color as the legs . prolegs exhibit dark grey plates on their lateral margins , with uniserial , biordinal crochets arranged in a mesal penellipse . two small grey verrucae are present in the region posterior to a10 under the anus , one of which is inferior and smaller .

the arrangement of scoli , plates , and verrucae in the third through eighth instars is the same as described for the second instar .

third instar larvae ( figs 14 - 15 ) are slightly variable in coloration , but the thorax is generally brown , the abdomen is brown or dark chestnut , and the thorax and a1 are darker than the abdomen . the abdomen is creamy yellow below the infraspiracular band . the scoli on the thorax and the back of the abdomen are dark brown , the abdominal supraspiracular and infraspiracular scoli are creamy yellow , and the supraspiracular scoli on a1 are typically brown . the spiracles on t1 and a8 are elliptical , equal in size and slightly larger than those on the rest of the abdomen , which are circular .

fourth instar larvae ( figs 16 - 18 ) exhibit a shiny black head and a dark brown , almost black body . the scoli are generally brown , although the infraspiracular scoli tend to be creamy yellow but are also occasionally brown . all of the body plates , including the prothoracic and anal shields , are shiny and black .

fifth to eighth instar larvae ( figs 19 - 21 ) are very similar to the fourth instar and to each other in color and morphology ( except in the shape of the spiracle ; see below ) , whether in their last instar or not . for example , all sixth instar larvae are similar to each other regardless of whether they then molt to a seventh instar larvae or whether they pupate . after the fifth instar , all body spiracles are elliptical in shape .

body length varies considerably within and between the fifth to eighth instars ( tab . ii ) . for example , in some cases , fifth instar larvae have the same body length as those of the sixth , seventh or eighth instars ; yet at the same time , sibling larvae at the same instar can exhibit differences in length of up to 10 mm . no relationship was found between the size of the larvae and the number of molts . the larvae pupated independently of their final size ( 25 - 40 mm ) . although it was not determined whether adult size is related to the size of the last instar larvae , it seems likely that the smaller pupating larvae become male butterflies , which are smaller than females .

pupae ( figs 23 - 24 ) are white , with thin , discontinuous black lines on the dorsal and lateral surfaces of the abdomen and a thick discontinuous band on both sides of the ventral midline that forms a barrel - like figure on each segment . a black outline is present on the proboscis , legs , antennae , eyes , labrum and wings . the clypeus and labrum are black . barely noticeable tiny black setae are distributed over the entire body , except the wing pads , antennae and legs . the cephalic region contains a pair of small protuberances , and the pronotum and mesonotum each possess a pair of aligned dorsal protuberances ; these proturbances decrease in size on the metanotum , where they appear only as small buttons . t2 exhibits a lateral protuberance at the base of the wings , crossed by a thick black line . lateral projections are present at the base of the wing pads . the spiracles on t1 are crescent - shaped and partially visible . the spiracles on a1 are hidden , those on a2 are hidden at the posterior end , and all others are elliptical and black . five pairs of identically shaped black dorsal spines without ramifications or setae appear on a2 to a6 , along the same line as the thoracic protuberances . a v - shaped marking that opens toward the anterior end appears at the base of each spine on a3 - a7 . posterior to this v - shaped marking , there is a transverse line and several supraspiracular lines . a cremaster formed by the ventral region of a9 and the dorsal and ventral regions of a10 is completely modified into a corrugated black structure .

larvae emerge at the preapical region of the egg ( fig . 5 ) and immediately feed on the chorion and on infertile eggs , if present . all activities including feeding , molting , resting and displacing are gregarious , from hatching to adulthood .

within the first several hours after hatching , larvae form a close line such that each individual is in physical contact with its neighbors and begin to feed simultaneously , starting at the edge of the leaf ( fig . 9 ) . the first to fourth instars eat only the leaf cuticle ( figs . 9 and 16 ) and frequently consume less than half of the leaf before passing on to the next leaf . third and fourth instars rapidly devour the leaf cuticle ; the leaf then dries and folds and the two ends are joined with silk ( fig . 18 ) . the first instars weave silk threads on the leaf surface , forming a dense web , under which they move and deposit their feces ( figs . 9 and 18 ) or spend periods of rest or premolt .

beginning at the fifth or sixth instar , larvae rest during the day and become active in the early evening . beginning at 06 : 00 to 07 : 00 h , they climb down the stem , one after the other , and hide either among dry leaves attached to branches close to the ground , at the base of the stem , among cracks in the stem or under dry leaves on the ground near the plant ( fig . 22 ) . between 18 : 30 and 19 : 30 h , the larvae emerge from their hiding place and climb , single file , back up the stem of the same plant until they reach the leaves , where they form small groups to feed . in the later instars , the larvae feed simultaneously on the leaves from the edges ( fig . 19 ) and either eat the whole leaf or leave only the veins uneaten .

during the first four instars , larvae from different egg clutches do not mix ; each group remains on its own leaf . however , groups of older larvae may hide in the same places close to or on the ground , and these groups may mix when they climb back up the plant to feed . one or two later - instar larvae have been observed together with early - instar larvae on the same leaf . when larvae begin to pupate , they individually move three or four meters away from the host plant and attach themselves to the undersides of leaves on other plants at a minimum height of 60 cm above the ground .

larvae of the syrphid fly xanthandrus bucephalus ( wiedemann , 1830 ) ( diptera : syrphidae ) were observed preying on first - to third - instar al . ozomene larvae ( fig . 15 ) . this solitary predator apparently goes unnoticed inside the silk mass of larvae and feces . occasionally , spiders were also seen preying on larvae . larvae and pupae collected in the field were often found to be parasitized by different species of tachinidae ( diptera ) and braconidae ( hymenoptera ) . predation on adults was not observed .

during the first three or four instars , when one individual in a group is threatened , the whole group reacts simultaneously by raising their bodies , standing on the last pair of prolegs ( fig . 17 ) and / or moving the anterior half of the body rhythmically from side to side . older larvae entwine themselves together and fall freely from the leaf , suspended by a silk thread . when the pupae are disturbed , they move the first abdominal segments from one side to the other , forming an angle of up to 120º .

altinote ozomene adults were normally observed in open areas and at forest edges . males ( fig . 25 ) fly low , frequently alighting on the ground , and can be caught by hand . adults form small groups of four or five individuals while feeding on the ground or on avian ( duck ) excrement on the banks of a reservoir . on several occasions , adults were observed feeding on the ground together with males of gnathotriche mundina steinii ( dewitz , 1877 ) ( lepidoptera : nymphalidae ) , an unrelated species with a very similar wing pattern . females ( fig . 26 ) were observed less often than males .

altinote ozomene feed on asteraceae , as has also been observed of al . ozomene nox ( sensu lamas 2004 ; harvey 1983 , devries 1987 ) and species of actinote ( devries 1987 , paluch et al . 1999 , 2001 , 2005 , francini et al . 2004 , 2005 , francini & freitas 2010 , freitas et al . 2009a , b , 2010 ) . valencia et al . ( 2005 ) recorded clibadium sp . ( asteraceae ) as a host plant for al . ozomene in the western cordillera of the colombian andes .

as in all known species of actinote , al . ozomene lays egg clutches under leaves ( devries 1987 , paluch et al . 1999 , 2001 , 2005 , francini et al . 2004 , 2005 , francini & freitas 2010 , freitas et al . 2009a , b , 2010 ) . the maximum number of eggs per oviposition in al . ozomene is notably less than that in actinote pellenea pellenea hübner , 1821 ( francini & freitas 2010 ) and slightly less than that in actinote surima ( schaus , 1902 ) ( paluch et al . 1999 ) and ac . carycina jordan ( paluch et al . 2001 ; table iii ) . the number of eggs per oviposition in al . ozomene is likely a limiting factor for larval survival because first - instar larvae in groups of fewer than 20 died and larvae of the first four instars generally only survived if they were in large groups , which was probably due to a better response to leaf structural defenses . when the number of larvae is small , the silk web on which the larvae move is equally small ( pers . obs . ) , leaving the leaf trichomes exposed , which may impede larval displacement and anchorage on the leaf and therefore affect feeding , as suggested by fordyce & agrawal ( 2001 ) . in addition , oviposition in al . ozomene tends to occur in a grouped fashion , on closely spaced plants or on leaves of the same plants , as has been described for ac . pellenea pellenea ( francini & freitas 2010 ) . other factors , including habitat and plant characteristics , may also affect female preference during oviposition . although the general behavior of al . ozomene is similar to that described for species of actinote ( tab . iv ) , nocturnal gregarious feeding has only been recorded in al . ozomene , and hiding in dry leaves during later instars has only recently been described for ac . conspicua ( freitas et al . 2010 ) . however , in contrast to al . ozomene , larvae of ac . conspicua hide individually or in pairs and only during the last instar .

most species of actinote have univoltine or bivoltine life cycles ( paluch et al . 1999 , 2001 , 2005 , francini et al . 2005 , freitas et al . 2009a , b , 2010 ) . a multivoltine cycle is presently known for only ac . pellenea pellenea ( francini & freitas 2010 ) . since 2004 , we have found al . ozomene frequently throughout the year , at low densities , except during the second half of 2005 , when no adults or immature stages were found .

a variable number of instars , as observed here in al . ozomene , has never been recorded in actinote ( paluch et al . 1999 , 2001 , paluch et al . 2005 , francini et al . 2005 , freitas et al . 2009a , b , 2010 ) . however , our observations suggest that such variation may occur in other species of altinote ( unpubl . data ) , and further investigation of this phenomenon might reveal both why it occurs and whether it is restricted to this genus .

the immature stages of al . ozomene are similar to those of some species of actinote from brazil and the andes in several morphological features . these include the initial color , shape and size of the eggs , the body color and arrangement of the setae in the first instar , the absence of spines on the head , the arrangement of scoli in other instars , the shape of the pupa and the number and arrangement of its abdominal spines ( paluch et al . 1999 , 2001 , francini et al . 2005 , freitas et al . 2009a , b , 2010 ) . al . ozomene differs from the species of actinote in egg color during the days following oviposition ( the eggs become reddish in actinote ) and in the color pattern of the larvae after the first instar . additional studies are clearly needed to identify additional characters that vary within the tribe and thus might provide useful information for phylogenetic studies .

we thank eliana flórez , juan d . marín , diana grisales , andrés vélez , and marisol ortega for help with rearing the larvae in the laboratory and alvaro idarraga for the identification of the host plants . this study was funded by comfenalco - antioquia .

borror , d . j . & d . m . delong . 1971 . an introduction to the study of insects . new york , holt , rinehart and winston . 812p . [ links ]

brower , a . v . z . 2010 . neotropical actinote clade . version 13 february 2010 , the tree of life web project , available online at : urltoken [ accessed : 04 / iii / 2011 ] [ links ]

devries , p . j . 1987 . the butterflies of costa rica and their natural history . princeton , princeton academic press , 327p . [ links ]

francini , r . b . ; a . v . l . freitas & c . m . penz . 2004 . two new species of actinote ( lepidoptera , nymphalidae ) from southeastern brasil . zootaxa 719 : 1 - 10 . [ links ]

francini , r . b . & a . v . l . freitas . 2010 . aggregated oviposition in actinote pellenea pellenea hübner ( lepidoptera : nymphalidae ) . the journal of research on the lepidoptera 42 : 74 - 78 . [ links ]

freitas , a . v . l . & k . s . brown jr . 2004 . phylogeny of the nymphalidae ( lepidoptera ) . systematic biology 53 ( 3 ) : 363 - 383 . [ links ]

harvey , d . j . 1983 . actinote leucomelas , p . 679 - 680 . in : d . h . janzen ( ed . ) . costa rican natural history . chicago , university of chicago press , 816p . [ links ]

harvey , d . j . 1991 . higher classification of the nymphalidae , p . 225 - 273 . in : h . f . nijhout ( ed . ) . the development and evolution of butterfly wing patterns . washington , d . c , smithsonian series in comparative evolutionary biology , 297p . [ links ]

paluch , m . ; m . m . casagrande & o . h . h . mielke . 1999 . estágios imaturos de actinote surima ( schaus ) ( lepidoptera , nymphalidae , acraeinae ) . revista brasileira de zoologia , 16 ( supl . 2 ) : 129 - 140 . [ links ]

paluch , m . ; m . m . casagrande & o . h . h . mielke . 2001 . estágios imaturos de actinote carycina jordan ( lepidoptera , nymphalidae , acraeinae ) . revista brasileira de zoologia 18 ( 3 ) : 883 - 896 . [ links ]

stehr , f . w . 1987 . order lepidoptera , p . 288 - 596 . in : f . w . stehr ( ed . ) immature insects . dubuque , kendal / hunt , 750p . [ links ]

toro , j . l . 2000 . árboles y arbustos del parque regional arví . medellín , corantioquia , 281p . [ links ]

vélez , a . ; p . duque & m . wolff . 2008 . mariposas del parque ecológico piedras blancas . guía de campo . medellín , fondo editorial comfenalco antioquia , 204p . [ links ]

valencia , c . a . ; z . n . gil & l . m . constantino . 2005 . mariposas diurnas de la zona central cafetera colombiana . guía de campo . chinchiná , cenicafé , 244p . [ links ]

caixa postal 19020 81531 - 980 curitiba pr brasil tel . / fax : ( 55 41 ) 3266 - 6823 sbz @ urltoken

papilio doris linnaeus , 1771 ; mantissa plant . 2 : 356 ; tl : " surinami " [ surinam ]

crenis brylle hübner , 1821 ; index exot . lep . : [ 2 ]

e - mail : jshuey at tnc . org ; director of conservation science ; indiana office of the nature conservancy ;

[ maps ] warning ! the maps are automatically generated from the textual information , and the process does not always produce acceptable result ; see about maps for more info .

weymer , 1891 exotische lepidopteren . v stettin ent . ztg 51 ( 7 - 12 ) : 280 - 294

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the grey-lined hawk ( buteo nitidus ) is a smallish raptor found in open country and forest edges .

it is sometimes placed in the genus asturina as asturina nitida .

the species has been split by the aou from the grey hawk .

the grey-lined hawk is found from southern costa rica to argentina .

the grey-lined hawk is 46 – 61 cm ( 18 – 24 in ) in length and weighs 475 g ( 16.8 oz ) average .

the adult has a pale grey body , the tail is black with three white bands and the legs are orange .

it has fine white barring on the upper parts .

immature birds have dark brown upperparts , a pale-banded brown tail , brown-spotted white underparts and a brown streaked buff head and neck .

this species is quite short-winged , and has a fast agile flight for a buteo .

it feeds mainly on lizards and snakes , but will also take small mammals , birds and frogs .

it usually sits on an open high perch from which it swoops on its prey , but will also hunt from a low glide .

the nest is of sticks and built high in a tree .

the usual clutch is one to three , usually two white to pale blue eggs .

the young take about 6 weeks to fledging .

grey - lined hawk

grey - lined hawk <EOT> photo of adult grey - lined hawk perched in dead tree . note barred back and nape , compared with unbarred grey back and nape of grey hawk .

grey - lined hawk ( buteo nitidus ) is a species of bird in the accipitridae family .

38–46 cm ; male 350–497 g , female 320–592 g ; wingspan 75–94 cm . grey overall , with fine dark grey barring above and bolder white barring below ; tail . . .

38 - 46 cm . a small pale grey hawk with fairly broad wings , extensively barred dark grey . the tail has a single solid black sub - terminal band . fast flapping flight with short glides , soars on flat wings . voice . a high , clear , drawn out disyllabic whistle .

bierregaard , r . o . , jr , boesman , p . & marks , j . s . ( 2018 ) . grey - lined hawk ( buteo nitidus ) . in : del hoyo , j . , elliott , a . , sargatal , j . , christie , d . a . & de juana , e . ( eds . ) . handbook of the birds of the world alive . lynx edicions , barcelona . ( retrieved from urltoken on 9 july 2018 ) .

eitniear , j . , s . mcgehee , w . waddell . 1990 . gray hawk mobbed by olive - throated parakeets .

glinski , r . 1988 . gray hawk . pp . 83 - 86 in r glinski , b pendleton , m moss , m lefranc , jr . , b millsap , eds .

grey - lined hawk , buteo nitidus , parque natural tayrona , birds caribe birding in santa marta many endemics ( about 25 ) and specialties occur here including santa marta antpitta , band - tailed , santa marta wood wren , bangs wood wren , white - tipped quetzal , santa marta sabrewing , santa marta wren , blue bearded helmetcrest , white - tailed starfrontlet , santa marta warbler , santa marta parakeet , santa marta mountain - tanager , yellow - crowned whitestart , santa marta bush - tyrant , santa marta rufous antpitta , brown - rumped tapaculo , rusty - headed spinetail , santa marta brush - finch , santa marta blossomcrown , santa marta foliage - gleaner , golden - breasted fruiteater , yellow - legged and black - hooded thrushes , black - headed tanager –furthermore an undescribed species of santa marta screech - owl , rusty - breasted antpitta , santa marta tapaculo , black backed thornbill , .

apparently some gray hawk pairs stay together after the breeding season is over . the gray hawks that overwintered in arizona were a pair . glinski called them out by mimicking their territorial breeding calls , and they responded in kind ( glinski , 1998 ) .

this species and b . plagiatus sometimes placed in asturina , partly because of distinctive moult pattern . probably closest to b . ridgwayi and b . lineatus ; previously considered to form a clade with these and rupornis magnirostris , but latter found to be basal to all buteos # r # r # r , and this group may be more closely allied to leucopternis than to buteo . until recently , commonly considered conspecific with b . plagiatus , which differs in its plain vs barred grey crown , nape , back and wings ( 3 ) ; uppertail coverts white vs grey with white tips , resulting in bold vs narrow white band on base of uppertail ( 2 ) ; and longer call , without sudden drop in pitch in middle of note ( 2 , perhaps 3 ) # r . n race blakei previously known as costaricensis , but latter name invalid , as preoccupied ( by b . jamaicensis costaricensis ) . three subspecies usually recognized .

there are no documented cases of predation of gray hawks , but like other mid - sized hawks , they may be vulnerable to larger birds of prey . raptor nestlings in the tropics may on rare occasions succumb to predation from arboreal hunters such as monkeys , coatis , snakes , and other birds , so it is possible that gray hawk nestlings may as well ( emmons and feer , 1997 ) .

on their wintering range , other factors could lead to gray hawk decline . glinski banded seven nestlings in arizona that were recovered in northern sinaloa state in mexico , and six of these had been shot . the greatest threat on these wintering grounds , however , is habitat loss . there , the gray hawks’ thornscrub habitat is being widely cleared for agriculture . gray hawks continue to persist along the living fence rows of trees that divide the agricultural fields , but it is unclear how many hawks this disrupted habitat can support , nor , ultimately , what effects this might have on arizona’s breeding population . little is known about demographics or conservation throughout the rest of the gray hawk ' s range ( glinski , 1988 , 1998 ; nabhan and sheridan , 1977 ; tellman et al . , 1997 ; urls below ) .

gray hawks are important predators of smaller lizards and snakes throughout their range , though cryptic coloration , speed , and good hiding places can help some reptile prey evade capture . gray hawks occasionally prey upon small birds . smaller birds will often deter predation by mobbing birds of prey to drive them away . olive - throated parakeets have been seen mobbing a gray hawk to force it to leave the area ( eitniear et al . , 1990 ) .

throughout their range , gray hawks inhabit woodlands and arid deciduous forests . in the tropics they prefer dry second growth forest and thorn scrub . they tend to select patchy open forest , forest edges , and savanna trees . it is not uncommon to find them on agricultural fields . in denser woodland they tend to keep high in the forest canopy . gray hawk northern breeding range is found in deciduous cottonwood - willow forests and mesquite bosques along riparian corridors or in evergreen oak woodlands ( glinski , 1998 ; stiles and skutch , 1989 ) .

since the range of gray hawks in the united states is so limited , many birdwatchers come to arizona and texas to see them . bird watching has become increasingly important to the economy of southeastern arizona . just as the gray hawk ' s distribution barely extends into southern arizona , so do the ranges of many other tropical and subtropical bird species . several migratory species also use the area’s river corridors as they pass north from mexico . over 400 bird species draw birdwatchers from around the world . their visitation has been an economic boon to the area . in 2001 , visitors to two well - known birding spots in southeastern arizona - ramsey canyon preserve and the san pedro riparian national conservation area - spent an estimated $ 10 . 1 million to $ 16 . 9 million . about a third of the gray hawk nest sites in arizona have been identified in the san pedro ncr . while gray hawks are not the sole draw to the area , the numbers clearly illustrate the impact that conservation has on regional economies . gray hawks , as well as people , stand to benefit from this perceived economic value . ( bibles , 1999 ; relly , 2002 ; viers , 2000 ) .

bibles ( 1999 ) also revealed that gray hawks prefer home ranges with taller trees and more open understory , probably because these enable them to observe their cryptic prey more easily . the increased flight space may also facilitate greater capture success . mesquite bosques are the primary foraging areas of gray hawks breeding in arizona because they have these characteristics . the amount of mesquite bosque seems to be the main factor that determines habitat quality in gray hawk nesting range in arizona . ( amadon and phillips , 1939 ; bibles , 1999 ; glinski , 1988 ; glinski and millsap , 1987 ; gurrola - hidalgo and chavez , 1996 ; stensrude , 1965 ) .

gray hawks from northern populations arrive in arizona around mid - march to breed . nest site fidelity is high , and gray hawk territories remain fairly constant from year to year . pairs begin building the nest right after courtship , but they partition the work . the male builds the foundation , and the female shapes most of the bowl out of green , leafy twigs from the nest tree or neighboring trees . northerly - breeding gray hawks primarily nest in cottonwood trees , but they will also nest in willow , ash , oak , hackberry , and mesquites . they choose nest sites in the upper third of the tree , usually in branches away from the trunk . nests are crow - sized , about 60 cm across . the female usually lays two eggs in early may , although studies of nest productivity in arizona have recorded nests with four or even eight eggs per clutch with a mean of 1 . 2 or 1 . 1 young per occupied breeding site . the eggs are white to pale blue and rarely marked . only females incubate the eggs . incubation lasts about 33 days , during which the male captures food for the female . after hatching , the young stay in the nest for about six weeks .

) range from the amazon basin in south america into the southwestern united states . they are migratory in the northern part of their range , arriving in southern arizona and extreme south texas in the spring to breed , and occasionally entering new mexico . these northern members of the species generally depart in mid - october to overwinter in mexico , although they can be found in south texas year - round , and rare records exist of winter residents in arizona . further south in their range , gray hawks are non - migratory ( glinski , 1998 ; kaufman , 2000 ; stiles and skutch , 1989 ; terres , 1980 ) .

gray hawks are medium - sized , woodland buteos , with shorter wings and longer tails than typical buteos . adult birds have a slate - gray back , finely barred gray and white underparts , a black tail with two or three white bands , and a white rump . juveniles have dark brown backs and buff - streaked underparts , brownish - gray tails with five to nine narrow black bars , and a dark brown eye stripe . both adult and immature birds have dark gray or black beaks , brown irises , and yellow ceres and legs . males are smaller than females . gray hawks fly with accipiter - like movements , alternately flapping and gliding gracefully . their flight pattern and gray color is similar to that of northern goshawks , which led to their other common name , “mexican goshawks . ” northern goshawks , however , have a white eyebrow and lack the tail barring distinctive of gray hawks ( glinski , 1998 ; kaufman , 2000 ; stiles and skutch , 1989 ; terres , 1980 ; wheeler and clark , 1995 ) .

arizona is the northernmost stronghold of breeding gray hawks , with nearly 80 known nest sites , most of which are protected in nature preserves or conservation easements . six pairs also nest regularly along the rio grande river in south texas . there is one reported incidence of nesting in new mexico . further south in their range , gray hawks are non - migratory and nest from december through may in tall , evergreen trees ( bibles , 1999 ; brandt , 1951 ; glinski , 1988 ; glinski , 1998 ; hubbard , 1974 ; stiles and skutch , 1989 ; terres , 1980 ) .

females alone provide incubation , but the male feeds her . the male provides most of the food for the first two weeks post - hatching , after which the female begins to hunt as well . it is not known how quickly the young can hunt for themselves once they leave the nest .

gray hawks are swift , agile fliers that can actively pursue prey by maneuvering through trees . they perch in the forest understory , locate prey in the trees or on the ground with their keen eyesight , and make short dashes to capture it whit their talons . they take reptiles , small mammals , birds , and some insects . in arizona , glinski ( 1988 ) studied food types delivered to nestlings and found that the diet was composed predominantly of terrestrial and arboreal lizards ( 74 % ) , garter snakes ( 5 % ) , nestling and adult birds ( 11 % ) , and mammals ( 10 % ) . bibles’ ( 1999 ) study of nest productivity in arizona revealed similar findings , with reptiles comprising 68 . 6 % of prey delivered to nestlings ( all were lizards but for one snake ) , mammals 19 . 6 % ( rodents and one rabbit ) , birds 9 . 8 % , and amphibians 2 % ( a toad ) .

gray hawks have no special conservation status , although their limited numbers at the northern extreme of their range have led management agencies in arizona to regard them as “sensitive species” and in texas to consider them “threatened . ” arizona breeding populations seem to be holding steady , with reproduction balancing the losses from mortality . fortunately , most of the 80 nest sites in arizona today are located on protected lands . concerned about how human demand for ground and surface water has seriously reduced desert riparian areas in the southwestern united states , private landowners and public and private conservation organizations have worked to set aside the remaining riparian corridors in southern arizona . further protection of the cottonwood - willow forests and mesquite bosques they support will demand protection of the waters that sustain them . if land managers succeed , this will be good news for gray hawks and the myriad other species that use these areas . their only threat then would be recreational disturbance . nesting gray hawks are sensitive to human activity near their nests , and many of the protected areas are frequented by recreational groups . something as innocuous as a family picnic unknowingly staged near a nest tree can cause gray hawks to abandon their nest .

robin kropp ( author ) , university of arizona , jorge schondube ( editor ) , university of arizona .

living in the nearctic biogeographic province , the northern part of the new world . this includes greenland , the canadian arctic islands , and all of the north american as far south as the highlands of central mexico .

living in the southern part of the new world . in other words , central and south america .

young are born in a relatively underdeveloped state ; they are unable to feed or care for themselves or locomote independently for a period of time after birth / hatching . in birds , naked and helpless after hatching .

having body symmetry such that the animal can be divided in one plane into two mirror - image halves . animals with bilateral symmetry have dorsal and ventral sides , as well as anterior and posterior ends . synapomorphy of the bilateria .

humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals . ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals .

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature . endothermy is a synapomorphy of the mammalia , although it may have arisen in a ( now extinct ) synapsid ancestor ; the fossil record does not distinguish these possibilities . convergent in birds .

forest biomes are dominated by trees , otherwise forest biomes can vary widely in amount of precipitation and seasonality .

offspring are produced in more than one group ( litters , clutches , etc . ) and across multiple seasons ( or other periods hospitable to reproduction ) . iteroparous animals must , by definition , survive over multiple seasons ( or periodic condition changes ) .

the area in which the animal is naturally found , the region in which it is endemic .

reproduction in which eggs are released by the female ; development of offspring occurs outside the mother ' s body .

referring to something living or located adjacent to a waterbody ( usually , but not always , a river or stream ) .

the region of the earth that surrounds the equator , from 23 . 5 degrees north to 23 . 5 degrees south .

" texas parks and wildlife , texas threatened and endangered birds " ( on - line ) . accessed april 12 , 2002 at urltoken .

" usda forest service , coronado national forest , santa catalina ranger district , tucson , arizona , usa . threatened , endangered , and sensitive species " ( on - line ) . accessed april 12 , 2002 at urltoken .

amadon , d . , a . phillips . 1939 . notes on the mexican goshawk .

. albuquerque , new mexico : unpublished report to office of endangered species , u . s . fish and wildlife service .

gurrola - hidalgo , m . , n . chavez . 1996 . serpentes : lampropeltis triangulum nelsoni ( milk snake ) . predation . .

nabhan , g . , t . sheridan . 1977 . living fencerows of the rio san miguel , sonora , mexico : traditional technology for floodplain management .

relly , j . 2002 . " arizona daily star : birding ' s big bucks . february 7 , 2002 " ( on - line ) . accessed april 12 , 2002 at urltoken .

stensrude , c . 1965 . observations on a pair of gray hawks in southern arizona .

arizona’s changing rivers : how people have affected the rivers . water resources research center issue paper 19

viers , j . 2000 . " economic development institute 2000 abstracts - southeastern arizona birdng trail " ( on - line ) . accessed april 12 , 2002 at urltoken .

. london , san diego : academic press , harcourt brace and co . .

to cite this page : kropp , r . 2002 . " asturina nitida " ( on - line ) , animal diversity web . accessed july 09 , 2018 at urltoken

disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .

this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .

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del hoyo , j . , collar , n . j . , christie , d . a . , elliott , a . and fishpool , l . d . c . 2014 . hbw and birdlife international illustrated checklist of the birds of the world . volume 1 : non - passerines . lynx edicions birdlife international , barcelona , spain and cambridge , uk .

buteo nitidus and b . plagiatus ( del hoyo and collar 2014 ) were previously lumped as b . nitidus following aou ( 2006 ) , the gender agreement of which follows david and gosselin ( 2002a ) , and before then were placed in the genus asturina following aou ( 1998 ) , and before then were split as a . nitida and a . plagiata following sibley and monroe ( 1990 , 1993 ) .

justification : this species has an extremely large range , and hence does not approach the thresholds for vulnerable under the range size criterion ( extent of occurrence < 20 , 000 km 2 combined with a declining or fluctuating range size , habitat extent / quality , or population size and a small number of locations or severe fragmentation ) . despite the fact that the population trend appears to be decreasing , the decline is not believed to be sufficiently rapid to approach the thresholds for vulnerable under the population trend criterion ( > 30 % decline over ten years or three generations ) . the population size has not been quantified , but it is not believed to approach the thresholds for vulnerable under the population size criterion ( < 10 , 000 mature individuals with a continuing decline estimated to be > 10 % in ten years or three generations , or with a specified population structure ) . for these reasons the species is evaluated as least concern .

this species has a large range from central america to amazonia and south into argentina . occurs from southwest costa rica south through panama and colombia to west ecuador on the west of the andes and on the east of the andes from east colombia , venezuela and guianas through east ecuador , northeast peru and much of brazil to north and east bolivia , paraguay and north argentina . the species is also found on trinidad .

described in parts of very large range as locally common and tolerant of open and secondary forest ; typically absent from closed humid forest . population likely to be large ( ferguson - lees & christie 2001 ) . trend justification : this species is suspected to lose 21 . 3 - 28 . 7 % of suitable habitat within its distribution over three generations ( 22 years ) based on a model of amazonian deforestation ( soares - filho et al . 2006 , bird et al . 2011 ) . it is suspected to decline by < 25 % over three generations as it is not highly forest - dependent , favouring open country and forest edges .

occurs in very wide variety of forest habitats from gallery and open woodland through to fairly open savannah with clumps of trees but is not typically found in dense humid forest . found from sea level up to 800 m , occasionally as high as 1 , 300 m . feeds on reptiles , amphibians , insects , small mammals and birds .

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( todd , 1915 ) – e bolivia and c brazil ( mato grosso to piauí and rio de janeiro ) s to paraguay and n argentina ( s to tucumán and n santa fe ) .

there are two primary vocalizations : a series of 3–8 piping notes used primarily during the . . .

adaptable , found in lowland tropical to subtropical zones in rain forest edge , disturbed forest and . . .

not globally threatened ( least concern ) . cites ii . no hard data on numbers or population trends , but generally considered widespread , locally common to relatively numerous , . . .

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previously incorporated several other genera , including e . g . rupornis , morphnarchus , pseudastur , as well as leucopternis and buteogallus . present arrangement based largely on recent molecular studies # r # r # r , but further modification likely .

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hbw alive contains information on descriptive notes , voice , habitat , food and feeding , breeding , movements , status and conservation plus a list of bibliographical references for this species account .

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josep del hoyo , thore noernberg , pieter de groot boersma , greg baker , jacob . wijpkema , david ascanio , desmond allen , anthony laven , helbert e . noventa .

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recommended citation birdlife international ( 2018 ) species factsheet : buteo nitidus . downloaded from urltoken on 09 / 07 / 2018 . recommended citation for factsheets for more than one species : birdlife international ( 2018 ) iucn red list for birds . downloaded from urltoken on 09 / 07 / 2018 .

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myavibase allows you to create and manage your own lifelists , and produce useful reports to help you plan your next birding excursion .

there are more than 12 , 000 regional checklists in avibase , offered in 9 different taxonomies , including synonyms more than 175 languages . each checklist can be viewed with photos shared by the birding community , and also printed as pdf checklists for field use .

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howard and moore 4th edition ( incl . corrigenda vol . 1 - 2 ) :

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erebia cyclopius is a species of butterfly of the satyrinae subfamily in the family nymphalidae .

it is found from the ural to siberia , northern mongolia , northern china and north korea .

the habitat consists of forest edges , flowery meadows and sparse larch forests .

the wingspan is 46 – 62 mm .

adults are on wing from june to july .

the species overwinters in the larval stage .

erebia cyclopius

erebia cyclopius <EOT> erebia cyclopius ( eversmann , 1844 ) = erebia cyclopia = cyclopius ( eversmann , 1844 ) .

buy butterflies for sale satyridae : erebia cyclopius set 10 from siberia online . worldwide shipping ! beautiful insect butterflies for sale satyridae : erebia cyclopius set 10 for sale at the bugmaniac , one of the world ' s largest dealers of preserved dried insects .

( 2008 ) : a checklist of the satyrine genus erebia ( lepidoptera ) ( 1758–2006 ) .

{ author1 , author2 . . . } , ( n . d . ) . erebia cyclopius eversmann , 1844 . [ online ] india biodiversity portal , species page : { name of species field } available at : urltoken [ accessed date jul 9 , 2018 ] .

first of the three erebia plates in the 1915 macrolepidoptera of the world , edited by adalbert seitz . this work was published near the height of taxonomic confusion about these butterflies .

the yellow - banded ringlet ( erebia flavofasciata ) is a member of the satyrinae subfamily of nymphalidae . it is a high mountain butterfly found in a small area of the alps in switzerland and italy .

erebia is a holarctic genus of brush - footed butterflies , family nymphalidae . most of the about 90–100 species ( see also below ) are dark brown or black in color , with reddish brown to orange or more rarely yellowish wing blotches or bands . these usually bear black spots within , which sometimes have white center spots .

this genus has found it easy to adapt to arid and especially cold conditions . most of its members are associated with high - altitude lands , forest clearings or high latitude and tundra . erebia species are frequent in the alps , rocky mountains , subarctic and even arctic regions , and the cooler parts of central asia . in fact , the north american term for these butterflies is " alpines " . eurasian species are collectively known as " ringlets " or " arguses " . however , none of these terms is used exclusively for this genus .

a peculiar case is the colorado alpine , the nominate subspecies of erebia callias . this isolated rocky mountains population has been lumped with the siberian brassy ringlets as they are almost alike morphologically . though one might suspect stronger differentiation and perhaps marked cryptic speciation across the wide range , the rocky mountains population is apparently a very recent isolate . its ancestors apparently crossed over the bering strait at the end of the wisconsinian glaciation , about 15 . 000 - 10 . 000 years ago . thus , brassy ringlets are present on the north american continent quite exactly for the same length of time as a significant human population .

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· type locality . " . . . in provincia irkuzkiensi " [ irkutsk region , russia ] .

· range . from the n . urals across siberia , n . mongolia and n . china to n . korea .

· distribution and variation . the s . urals , central and s . siberia , the altai and sayan mts . , transbaikalia - the nominate subspecies ; the amur and ussuri regions - ssp . aporia schawerda , 1919 ; sakhalin - yoshikurana kishida et nakamura , 1941 .

· habitat and biology . forest edges , flowery meadows , sparse larch forests , in the mountains up to 2 , 000 m a . s . l . flight period : june - july . larvae hibernating .

· similar species . e . wanga : unh without bluish postdiscal band . e . edda : upf double eye - spot diffusely bordered with reddish colour ; unh without bluish postdiscal band .

photo and text : guide to the butterflies of russia and adjacent territories volume 1 . pensoft , sofia - moscow . 1997

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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .

disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .

this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .

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copyright © andre gorodinski . the insects from the palaearctic region . web design by mrs . l . gorodinski . special thanks to mr . a . malinin for technical support website .

the wingspan is 46–62 mm . adults are on wing from june to july .

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regions of the russian federation : gorno - altai , trans - baikal , krasnoyarsk , nizhny - amur , prealtay , of baikal , pribaikalskiy , primorye , sakhalin , the north - yenisei north okhotsk , mid - amur , average okhotsk , mid - ural , sredneobskaya , tuva , south - uralsky , south yakutia .

[ 10 ] de jong , y . s . d . m . ( ed . ) ( 2011 ) fauna europaea version 2 . 4 ( faunaeur . org )

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curators : konstantin efetov , vasiliy feoktistov , svyatoslav knyazev , evgeny komarov , stan korb , alexander zhakov .

species catalog enables to sort by characteristics such as expansion , flight time , etc . .

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a diverse and widespread holarctic genus , with some species distributed in both western north america and eastern eurasia .

phylogenetic relationships among the tyndarus group are based on albre et al . ' s ( 2008 ) study of mitochondrial dna sequences . the remainder of this diverse genus awaits phylogenetic study .

larval host plant is unknown . a swiss subspecies , e . flavofasciata warreni , has been proposed .

( i . e . , without discussing synonymy and relationships ) was published in 2008 .

. but things hardly improved as more and more of the diversity of these butterflies came to note . in

. in the 19th and early 20th century the alps were a popular destination for butterfly collectors and specimens of alpine butterflies were very profitable for dealers . the dealers , mostly german , not only sold specimens , but were

this , together with the then - popular , even obsessive study of variation by entomologists—examples are james william tutt , george wheeler , felix bryk and brisbane charles somerville warren —led to very many names being applied to what may be or much more likely may not be biological species or subspecies . a further problem is the use of the term " variety " . authors of that time used this for an individual variant , a group of individuals morphologically but not otherwise related , seasonal forms , temperature - related forms , or geographic races ; it was later usually taken to mean the last subspecies though this is often suspected to have been premature .

studies add to the available data , it is becoming clear that most " varieties " that have at least been commonly considered subspecies in the latter 20th century are indeed lineages distinct enough to warrant some formal degree of recognition . another result of recent research is confirmation of the theory that this genus contains many glacial

species is given variously around 90 - 100 , as developments happen so fast that it is hard for authors to remain up to date regarding the newest changes .

brower ( 2006 ) , albre et al . ( 2008 ) , and see savela ( 2008 ) for more sources

jim p . brock and k . kaufman . kaufman field guide to butterflies of north america , new york , ny : houghton mifflin , 2003 .

( lepidoptera , rhopalocera , nymphalidae , satyrinae ) species group combining coxii and nd5 mitochondrial genes : a case study of a recent radiation .

. version of 2006 - nov - 28 . retrieved 2008 - aug - 11 .

. version of 2008 - feb - 04 . retrieved 2008 - aug - 11 .

. version of 2008 - mar - 15 . retrieved 2008 - aug - 11 .

– , and in many cases certainly close relatives . a notable trait of their genus is an ability to adapt well to cold and somewhat

brassy ringlets are mid - sized members of their genus , with a length of 17–22 mm ( roughly two - thirds to one inch ) .

- colored patch that is lighter than the blackish - brown upperwings . in some , a third black spot , much smaller and without the white dot , is present at the opposite end of the reddish patch . the hindwings have no black spots in many , but in some

there are a few ( 2 - 4 or so ) black dots , usually without white in the center , paralleling the outer margin . if hindwing spots are present , they are sometimes surrounded by a lighter brown field like the forewing spots , sometimes not . the wing undersides are

colored , usually in greyish - brown and often with a noticeable band arching through the central hindwing , the rest of which has a silvery sheen which makes these species quite recognizable in flight .

the eyespots and the forewing patch surrounding them are found on the forewing undersides also ; if a hindwind pattern is present , it may or may not show up on the under hindwings either whole or in parts . the common name derives from the

as it seems , the origin of this group is perhaps south of the central asian or more likely in the balkans region . probably around 1 million years ago during the pre - pastonian stage , the original population expanded north . during an interstadial , the southern montane metapopulation and the one to the north which ranged across the eurasian taiga split .

two major southern populations were established some 800 . 000 - 700 . 000 years ago during the pastonian stage , when the habitat belt moved uphill , cutting off gene flow between major mountain ranges . coincident with the günz - mindel interglacial , about 600 . 000 - 500 . 000 years ago some more distinct local populations diverged in the south .

meanwhile , the northern population had been spreading across europe to the pyrenees and deep into siberia . with the ice – which had then been covering much of northern and central europe – retreating at the end of the elsterian stage between 400 . 000 - 300 . 000 years ago , the taiga population and their relatives in the mountains of central and western europe became isolated for good . finally , at the end of the riss / saale glaciation about 130 . 000 years ago , alpine and siberian populations fragmented further . the former stayed rather distinct while more subsequent gene flow occurred between the latter ; this difference is probably an effect of habitat topography .

, the molecualr and morphological data are still more in favor of retaining a single species .

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size : 160 x 230 mm wide extend : 544 pp , 10000 colour illustrations , more then 700 maps .

n . africa ( from morocco to egypt ) , asia minor and near east .

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2 probably this species photographed on the dry hill north of markakol and 4 + almost certainly this species on the way up the alatay pass . 2 / 14

subspecies ‘alpherakyi’ . seen above cim bulak , almaty and 2 + between kamenka and ust - kamenogorsk . 2 / 14

probably widespread but only identified with certainty on 4 days . certainly there were other colias species present that went unidentified

noted in the kalbinskiy hills and 1 on the alatay pass . 3 / 14

at times the commonest of the wood whites at markakol . distinctive with pronounced ‘cross - bars’ on the hindwings and slightly more attentuated forewings . 3 / 14

1 on the marble pass and 3 + on the dry hill north of markakol . 2 / 14

noted between ust - kamenogorsk and zaysan and several at the yourta service area there on the way back . 2 / 14

the commonest blue in meadows between markakol and the burkhat pass . 2 / 14

common on the dry hill north of markakol and on the alatay pass . 2 / 14

singles on the alatay pass and on the northward tributary of the buktarma below the climb to rakhmanovskie klyuchi . 2 / 14

1 in the kalbinskiy hills and also noted at the bee farm near kamenka . 2 / 14

seen on the alatay pass . like the next species but with no orange areas on the inner part of wing

singles at markakol and by the north tributary of the bukhtarma river below the rise to rakhmanovskie klyuchi . 2 / 14 . a very distinctive ringlet

singles on the alatay pass and in a bog by the lake at rakhmanovskie klyuchi . 2 / 14


amphizoa lecontei is a species of aquatic beetle .

adults have a body length between 11-16mm .

its elytron has a distinct carina on fifth interval .

found in western north america , especially in the rocky mountains .

its common name is " trout-stream beetle " .

its synonym is amphizoa carinata .

amphizoa lecontei

amphizoa lecontei <EOT> html public " - / / w3c / / dtd html 4 . 01 transitional / / en " " urltoken "

proceedings of the california academy of sciences , vol . 44 , no . 6

disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .

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upcoming events 2018 bugguide gathering in virginia july 27 - 29 : registration and discussion photos of insects and people from the 2015 gathering in wisconsin , july 10 - 12 photos of insects and people from the 2014 gathering in virginia , june 4 - 7 . photos of insects and people from the 2013 gathering in arizona , july 25 - 28 photos of insects and people from the 2012 gathering in alabama photos of insects and people from the 2011 gathering in iowa photos from the 2010 workshop in grinnell , iowa photos from the 2009 gathering in washington

contributed by phillip harpootlian on 18 december , 2008 - 12 : 02pm additional contributions by mike quinn , t . loh last updated 5 april , 2016 - 6 : 24pm

disclaimer : dedicated naturalists volunteer their time and resources here to provide this service . we strive to provide accurate information , but we are mostly just amateurs attempting to make sense of a diverse natural world . if you need expert professional advice , contact your local extension office .

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new evidence on the phylogeny and biogeography of the amphizoidae : discovery of a new species from china ( coleoptera ) .

citation : peiyu , y . , and n . e . stork . 1991 . new evidence onthe phylogeny and biogeography of the amphizoidae : discovery of a new species from china ( coleoptera ) . syst . entomol . 16 ( 2 ) : 253 - 256 .

citation : kavanaugh , d . h . , 1986 . a systematic review of amphizoid beetles ( amphizoidae : coleoptera ) and their phylogenetic relationships to other adephaga . proc . california acad . sci . 44 ( 6 ) : 67 - 109 .

a treatise on the western hemisphere caraboidea ( coleoptera ) : their classification , distributions , and ways of life . vol . iii

pensoft series faunistica 99 . pensoft publishers , sofia - moscow . 412 pp . , 2011

full title : a treatise on the western hemisphere caraboidea ( coleoptera ) : their classification , distributions , and ways of life . volume iii ( carabidae – loxomeriformes , melaeniformes )

pensoft series faunistica 66 . pensoft publishers , sofia - moscow . 323 pp . , 2007

full title : a treatise on the western hemisphere caraboidea ( coleoptera ) : their classification , distributions , and ways of life . volume i ( trachypachidae , carabidae - nebriiformes 1 ) .

checklist of the water beetles of florida this checklist registers species known to occur in florida , based on literature citations and material examined by the author . it also includes taxa which may occur in florida ; many of these taxa occur on the u . s . southeastern coastal plain but have not been positively identified from florida . note also that some literature records may be considered doubtful ; some species recorded in earlier literature but misidentified are not listed . only important synonyms pertaining to florida water beetles are listed . in general , only aquatic or semi - aquatic species are listed . the families , genera and species are listed in alphabetic order . undescribed taxa are assigned letter or number designators .

by nilsson , a . n . and b . j . van vondel . 2005 .

limited preview anders n . nilsson , bernhard j . van vondel . 2005 . world catalogue of insects . vol . 7 : amphizoidae , aspidytidae , haliplidae , noteridae and paelobiidae . ( coleoptera : adephaga ) . apollo books , vester - skerninge , denmark . 171 pp . world catalogue of insects is an initiative aiming at compiling worldscale , authoritative catalogues of monophyletic insect taxa . volumes in this series will contain standard nomenclatoral information on all names pertaining to the taxon treated , including type locality and distribution to the extent this is relevant .

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medium - sized ( 11 . 7 - 14 . 0mm ) ; body moderately broad and dark brown or dull black ; pronotum with coarse , sparse puntures ; elytra finely and densely pitted ; elytral striae completely and faintly impressed ; coarsely punctate ; elytron flattened medially with blunt but distint carina on 5th interval ; area medial to carina elevated and flat ; area lateral to carina slightly concave . ( see also identification of genus ) .

extends from southern yukon territory , south along rocky mountains through alberta and british columbia to chuska mountains of northeast arizona and the sangre de cristo range of northern new mexico ; eastward from wallowa mountains of northeastern oregon and independence mountains of northeastern nevada to the bighorn mountains of north - central wyoming and the front range of central colorado .

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confused by a class within a class or an order within an order ? please see our brief essay .

to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .

disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .

this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .

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the phylogenetic propositions of baehr ( 1979 ) , and kavanaugh ( 1986 ) , are not consistent with the interpretation provided herein , whereas the conclusions of burmeister ( 1976 ) , ruhnau ( 1986 ) and beutel ( 1986 ) seem to correlate well . therefore , it is suggested that the amphizoidae are the sistergroup of hygrobiidae + dytiscidae .

present address : institut für zoologie , rwth aachen , kopernikusstrasse 16 , d - 5100 aachen , federal republic of germany .

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