papilio alexanor , the alexanor or southern swallowtail , is a butterfly species in the papilionidae or swallowtails .

the insect has a wingspan of 62 – 70 mm .

it flies from april to july in a single generation , in mountainous regions ( 1700 m ) of europe , asia minor , the balkans , and asia ( anatolia , iran , afghanistan , pakistan to west tian-shan ) .

papilio alexanor

these are nice photos of papilio alexanor ( southern swallowtail , südlicher schwalbenschwanz ) . papilio alexanor is a member of the
here papilio alexanor from greece ; collecting data : papilio alexanor eitschbergeri , male , bollino & sala , 1992 , pythagoreio , 100 m asl , samos island ( may , 1999 ) . papilio alexanor live in french provence : urltoken enjoy !
c . michael hogan changed the thumbnail image of
image of papilio alexanor
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biology and distribution of papilio alexanor esp . ( lepidoptera , papilionidae ) in central asia
kari pihlaviita added the finnish common name
välimerenritari
to
papilio alexanor esper 1800
.
biology and distribution of papilio alexanor esp . ( lepidoptera , papilionidae ) in central asia
biology and distribution of papilio alexanor esp . ( lepidoptera , papilionidae ) in central asia [ 1979 ]
jennifer hammock split the classifications by eol group on flickr from papilio alexanor esper 1800 to their own page .
c . michael hogan marked the classification from
irmng
as preferred for
papilio alexanor esper 1800
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biology and distribution of papilio alexanor esp . ( lepidoptera , papilionidae ) in central asia [ tadzhik ssr ]
synonyms : papilio asiatica ( moore , 1882 ) ; papilio machaon ( jordan , 1909 ) .
ussr - circa 1987 : a stamp printed in the ussr shows papilio alexanor , the southern swallowtail butterfly , circa 1987 .
bollino , m . and sala , g . , 2004 . synopsis of papilio alexanor esper . natura edizioni scientifiche , bologna .
on the top one pair of p . alexanor eitschbergeri ( samos is . )
i have found this informations in
synopsis of papilio alexanor
. the authors of this work , bollino & sala , think that specimens of
a chrysalis of alexanor . unusual in that it attached to rock by a silken girdle .
synonyms : papilio pammon ( linneaus , 1758 ) ; papilio romulus ( cramer , 1775 ) ; papilio pammon ( fabricius , 1775 ) ; papilio cyrus ( fabricius , 1793 ) ; papilio sakontala ( hewitson , 1864 ) ; papilio pammon ( wallace , 1865 ) ; papilio polytes ( butler , 1869 ) ; laertias romulus ( moore , 1881 ) ; papilio ceylanicus ( c . & r . felder , 1864 ) ; papilio walkeri ( jonson , 1879 ) ; papilio polytes ( leech , 1893 ) ; laertias pammon ( moore , 1903 ) ; papilio polytes ( talbot , 1939 ) seventeen subspecies . the pakistan form belongs to the subspecies romulus as shown above . this specimen was caught by g waldeck in karachi 1984 .
i can ' t help you with all the other alexanor ssp , but i have some in my collection .
a study of hybrids betweeen papilio xuthus and the p . polyxenes - machaon group
papilio phylogeny based on mitochondrial cytochrome oxidase i and ii genes . - pubmed - ncbi
the southern swallowtail ( papilio alexanor ) is a very nice butterfly of the palaearctic ecozone ( europe and asia ) . the distribution extend from the south of europe ( italy , greece ) to kazakhstan and pakistan .
sorry to have to tell you this , but the data must be wrong , p . alexanor does not occur in morocco .
hello , recently i have took photos of some of alexanors from my collection . thought it might be a good idea to post it here , too ( to tease somebody , e . g . panzerman ; d ) so , here we are . papilio alexanor destelensis : male . france : var . p . alexanor hazarajatica : male . turkmenistan : sw . kopet - dagh mts . p . alexanor hazarajatica : female . turkmenistan : sw . kopet - dagh mts . p . alexanor voldemar : male . uzbekistan : w . tian - shan mts . p . alexanor voldemar : female . uzbekistan : w . tian - shan mts . except for destelensis , all are collected by myself many years ago , in 1991 .
the taxonomy and identity of papilio nitra w . h . edwards in colorado ( papilionidae )
de freina , j . j . 1983 . papilio alexanor orientalis romanoff , 1894 : angaben zur biologie , verbreitung und zur frage der berechtigung dieses taxons ( lepidoptera , papilionidae ) . atalanta 14 ( 1 ) : 23 - 30 .
no previous molecular phylogenetic studies on papilio have resulted in a single robust or relatively comprehensive phylogenetic reconstruction . the improved resolution obtained here allowed us to revisit several inadequately resolved issues . in congruence with previous studies of papilio phylogeny ( aubert et al . , 1999 ; caterino et al . , 1999 ; reed and sperling , 1999 ) , our results in the ml and bayesian analyses on combined data show that the genus papilio comprises two lineages ( fig . 5 ) : one includes hancock ' s genera ( here treated as subgenera ) heraclides , pterourus , and chilasa and papilio alexanor , and the other consists of princeps , papilio ( sensu stricto ) , and the monotypic eleppone . a potential alternative basal topology , with heraclides as the first clade to diverge from the remainder of papilio sensu lato ( fig . 1 ) , suggests the existence of three major papilio lineages : heraclides , pterourus + chilasa + papilio alexanor , and princeps + papilio ( sensu stricto ) + eleppone . however , this branching pattern has weaker support in most analyses , and therefore we relied on the relationships obtained from the combined mp bootstrap , ml , and bayesian analyses .
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 .
p . alexanor : greece , peloponnesos , taygetos , kambos , 1999 . 05 . 19 , leg . puchmayr / kurz , coll . michael kurz
systematics of the papilio glaucus and p . troilus species groups ( lepidoptera : papiliondae ) : inferences from allozymes
p . alexanor atticus : greece , peloponnesos , taygetos , kambos , 1999 . 05 . 19 , leg . puchmayr / kurz , coll . michael kurz
p . alexanor atticus underside : greece , peloponnesos , taygetos , kambos , 1999 . 05 . 19 , leg . puchmayr / kurz , coll . michael kurz
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 .
papilio alexanor has been proposed by esper ( 1800 ) as replacement name for papilio polidamas lezzolo in prunner ( 1798 : 69 ) , which is a junior primary homonym of papilio polydamas ( = battus polydamas ) linné , 1758 :
es hat ihn bereits hr . graf excoffier de lezzolo , nach obiger anzeige , unter den nahmen des p . polidamas beschrieben . da dieser schon an einen ausländer vergeben war ; so hatte ich eine aenderung anzugehen , und die vorgesetzte benennung gewählt .
( esper 1800 ) .
evolution of the mimetic african swallowtail butterfly papilio dardanus : molecular data confirm relationships with p . phorcas and p . constantinus
hello please , what subspecies of p . alexanor are from croatia ? its very rare there . if someone have it can you put pictures please . thank you very much !
papilio is relatively old and is a highly complex and diverse taxonomic unit . many characteristics of papilio have a polyphyletic origin , with multiple gains and losses of particular morphological or ecological traits . although no definitive morphological synapomorphies for generic subdivisions are broadly recognized , the evolutionary age and degree of genetic divergence between monophyletic groups within papilio would support elevation of at least five taxa , heraclides , pterourus , chilasa , papilio , and eleppone , to generic rank . a new genus name would also be required for p . alexanor . further studies are required to clarify the composition of princeps ( sensu hancock , 1983 ) , which in contrast to traditional taxonomy is paraphyletic and seems to include at least three divergent groups . however , we refrain from offering formal taxonomic conclusions until evolutionary patterns are clarified more consistently across papilio .
the yellow swallowtails papilio machaon linné , 1758 , papilio saharae oberthür , 1879 and papilio hospiton guenée , 1839 can be distinguished easily by the black wing markings , especially by the lack of a black transverse fascia in the basal half of the hindwing . the similarly marked iphiclides podalirius linné , 1758 has the ground colour much lighter creamy - white , with much longer tails on the hindwings .
. the first description was in 1800 by esper . with a wingspan of 7 . 0 – 8 . 0 cm the southern swallowtail is a small member of the family papilionidae . the butterfly is yellowish with typical black tiger - strips . hind wings have long tails . papilio alexanor has a small area of distribution and is rare .
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 ) .
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 .
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 .
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 .
extrait du documentaire de 25 minutes . dans le sud - est de la france vit l’alexanor ( papilio alexanor esper , 1799 ) , famille de papilionidae très localisé en région paca . l ' alexanor est un grand papillon avec une envergure de 62 à 70 mm et figure parmi une espèce protégée en france . il vole en une génération entre fin - mai et mi - juillet et fréquente les sites calcaires chauds , secs , caillouteux ou rocheux de 600 à 1500 m . la femelle dépose ses œufs isolément sur les ombelles de sa plante - hôte principale le ptychotis saxifraga , avec une préférence pour celles en bouton ou en début de floraison . a terme , la chenille quitte la plante - hôte et part à la recherche d’un site approprié pour préparer sa métamorphose . les chrysalides hibernent immobiles jusqu’au printemps suivant . remerciements roger gibbons pour plus d ' information visitez le site web filming varwild urltoken urltoken
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 .
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 .
thanks for the compliments ! it might be a good idea to put here a photos of all the described sspp . of alexanor . there is a thread on this topic at the old forum , of course . though it would be handy to have everything gathered at one place , imho alexei
species of the genus papilio are highly diverse in adult wing pattern and immature stage morphology . they also have highly varied larval host plant utilization , ranging from restricted use of one or two host species to broad polyphagy of up to eight different plant families ( see bossart and scriber , 1995 ) . however , about 80 % of species of papilio ( eleppone and almost all species in heraclides and princeps ) are primarily rutaceae feeders . the main hosts of most papilio ( sensu stricto ) species are apiaceae . larvae of chilasa usually develop on lauraceae . basal species of pterourus also feed on members of the families lauraceae and magnoliaceae , but some species utilize rosaceae and other hosts ( see tyler et al . , 1994 ; scriber et al . , 1995 ) . the lack of a stable papilio phylogeny and its relationships with other swallowtails hindered previous attempts to resolve host use evolution and to determine ancestral host plant relationships ( miller , 1987 ) .
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 .
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 .
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 .
a summary of major classifications of papilio is presented in table 1 . munroe ( 1961 ) divided papilio into five sections , which he did not designate as subgenera because of the lack of any simple system of adult characters . hancock ( 1983 ) , in his explicitly cladistic estimation of relationships within papilio , recognized six genera based on phylogenetic evidence and their inferred evolutionary antiquity , but that phylogeny still suffered from lack of character justification . another classification was proposed by igarashi ( 1984 ) based on morphology of immature stages , but this work was not complete in representing all of hancock ' s genera . igarashi recognized seven genera , with numerous discrepancies between this treatment and that of hancock ( 1983 ) . hancock ' s classification was criticized by miller ( 1987 ) , who did not find justification for elevating papilio subdivisions to the generic level . however , some elevation is seen in the latest widely available checklist of swallowtail butterflies ( haüser et al . , 2002 ) , where chilasa is treated as a genus .
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 ) .
several findings add further support to recent suggested taxonomic changes . in agreement with scriber et al . ( 1991 ) and caterino and sperling ( 1999 ) , we found pterourus sensu hancock ( 1983 ) to be paraphyletic with respect to pyrrhosticta . monophyly of heraclides , and relationships within it , were strongly supported in all analyses . there is some evidence against division of heraclides into four subgenera , heraclides , calaides , troilides and priamides , as suggested in tyler et al . ( 1994 ) . at least for troilides , we found relatively strong support for paraphyly . however , additional samples are needed . our results support the previously weak placement of papilio alexanor near pterourus , as found by caterino and sperling ( 1999 ) and reed and sperling ( 1999 ) . previous suggestions that this species belongs near the p . machaon species group ( munroe , 1961 ; hancock , 1983 ) can now be considered refuted , and the resemblance of p . alexanor and p . machaon is considered the result of convergence due to similar host plants utilized by larvae .
other uncertainties with reconstruction of the ancestral states are related to incomplete information about resting larval behavior and proleg structures . optimization of larval feeding habits may be tentative in the absence of a well - supported phylogeny for papilio outgroups . the tribe troidini was believed to be the sister group for papilionini , i . e . , papilio sensu miller ( 1987 ) , and the tribe leptocircini was considered their sister group ( hancock , 1983 ; caterino et al . , 2001 ) . however , some studies have suggested a basal position of the genus meandrusa within papilionini ( aubert et al . , 1999 ) . although we were not able to sample this genus , we performed alternative character optimizations placing meandrusa as a sister taxon of papilio to test different hypothesis . differences in ancestral state reconstructions due to choice of sister taxa are outlined in the discussion .
the phylogenetic relationships within papilio obtained here ( fig . 4 ) suggest that ancestral feeding on rutaceae is at least equally plausible based on mp character optimization when meandrusa was included as the sister taxon of papilio . reconstruction of the ancestral larval feeding habits under the ml model supported rutaceae feeding as a primitive trait even when meandrusa was included in character optimization . we have estimated the first split within papilio at approximately 55–65 mya , fitting the proposed age of the initial diversification within rutales at about 67 mya ( magallón et al . , 1999 ) . under this hypothesis , there would be only one major shift to lauraceae , in the stem lineage of chilasa and pterourus ( after the divergence of p . alexanor ) . another shift to lauraceae in the african p . hesperus group ( ackery et al . , 1995 ) is not associated with major diversification and can be compared with the shift from rutaceae to fabaceae feeding in the australian subspecies of p . demoleus , p . d . sthenelus ( braby , 2000 ) . another observation supporting rutaceae as the ancestral host is that even those species whose larvae normally develop on members of other plant families possess the ability to feed on rutaceae , both in the wild and in captivity ( tyler et al . , 1994 ) .
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
we added sequences for 31 species of papilio sensu lato to the 23 available previously ( caterino and sperling , 1999 ; reed and sperling , 1999 ; caterino et al . , 2001 ) . overall , we include sequences for 54 specimens from 51 papilio species , which represent almost 25 % of all species in this large genus . species were selected to give more complete representation of major species groups and subgenera within papilio . representatives of 26 of 42 species groups ( sensu hancock , 1983 ) were included . the sampled species represent all genera recognized by munroe ( 1961 ) , hancock ( 1983 ) , and haüser et al . , ( 2002 ) and all except two genera ( agehana and euchenor ) recognized by igarashi ( 1984 ) . outgroups were 16 species of papilionidae from tribes other than papilionini , and pieris napi from the putative sister family , pieridae . newly sampled taxa , sources of material , and genbank accession numbers for new materials are given in table 2 .
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 .
when a different outgroup was chosen , no substantial differences were observed for reconstruction of ancestral states of any characters except larval host plant . when meandrusa was placed as the closest papilio outgroup , an uncertain mp reconstruction of the ancestral host plant was obtained at the root of papilio ( node 88 ) , indicating equal probabilities for larval feeding on rutaceae and lauraceae . in ml reconstructions , with meandrusa as a sister taxon for papilio , all changes except larval hosts at nodes 88 and 101 ( fig . 4 ) had the same transitions but with slightly higher or equal log - likelihood support ( data not shown ) . inclusion of meandrusa decreased the likelihood ( from 97 . 3 % to 87 . 1 % out of total likelihood ) that rutaceae feeding was the primitive trait for papilio larvae . as the ancestral larval host , lauraceae had less than 0 . 6 % of total likelihood . inclusion of meandrusa increased this estimate up to 11 . 6 % . uncertainty remains for the ancestral state of node 101 , where none of the reconstructions gave a strong indication for larvae feeding on any one of rutaceae , apiaceae , or lauraceae as the primitive trait . incomplete data did not allow ml character optimizations for presence or absence of female secretion on eggs , shape of larval body hairs , number of crochet rows on larval prolegs , and presence or absence of a silk pad on larval resting sites .
partly because of the limitations of traditional approaches and with the development of new systematic techniques , the classification of papilio has received significant attention in recent years . relationships among species within the p . machaon and p . glaucus - troilus species groups were studied based on allozyme variation ( sperling , 1987 ; hagen and scriber , 1991 ) . restriction fragment length polymorphism of mitochondrial dna ( mtdna ) was used to compare taxa within the same species groups in later studies ( sperling , 1991 , 1993a , 1993b ; sperling and harrison , 1994 , tyler et al . , 1994 ) . phylogenetic relationships within papilio have also been analyzed using dna sequences of a variety of genes ( 28s , cyt b , ef - 1 α , nd1 , nd5 , coi , coii ) , but these studies have been confined to single species groups ( vane - wright et al . , 1999 ) or local geographic areas ( yagi et al . , 1999 ) or have included limited sampling across papilio subdivisions ( aubert et al . , 1999 ; caterino and sperling , 1999 ; reed and sperling , 1999 ) .
papilio can be considered to fall into two major clades : one ( with few exceptions ) , restricted to the new world and the other comprising mostly old world species . this fact , along with the suggested center of the origin for papilio in the north america–europe landmass ( hancock , 1983 ) , gives reason to assume that initial diversification was related to disconnection between the present nearctic and palearctic . this event was almost completed by the end of the cretaceous about 65 mya ( dietz and holden , 1970 ) . the latest direct land connection between europe and eastern north america , known as the thulean route , was apparently available into the miocene and persisted up to as recently as approximately 20 mya ( tiffney , 1985 ) . however , because climatic cooling after the eocene / oligocene boundary apparently rendered the thulean route unsuitable for many warm temperature–adapted groups , the vicariance event may have been approximately 35 mya ( noonan , 1988 ) or even earlier ( chapco et al . , 2001 ) . based on this geological time frame , we constrained the maximum and minimum time of the initial split within papilio to 65 and 35 mya , respectively .
some morphological traits of papilio larvae ( such as the morphology of prolegs ) are tightly related to their behavior . the number of crochet rows in abdominal prolegs is related to the ability of the larva to attach to the host plant ( hagen , 1999 ) . larvae with clasping prolegs , with only a single row of crochets , can cling more firmly to thin twigs , whereas larvae with gripping prolegs and two rows of crochets usually produce a silk pad on the leaf surface and hold on to this pad . twig - clasping prolegs seem to be plesiomorphic in papilio . although data on proleg morphology are incomplete , there are several examples of apparently independent gains or losses of the two - row gripping prolegs and the ability to spin a silk pad ( fig . 4 ) .
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 .
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 ) .
it is a species that eluded me until 2013 . i visited known sites in mid - june but in this late year , when a poor spring left most species about two or three weeks late , i was clearly too early . as i was due to be in the region of another known site in mid - july , i took the opportunity to go there and was not disappointed . there were about six alexanor flying , roughly half males and half females . i observed them for about four hours in total over three days but i have to say that during that entire period not one stopped for any reason , even when the sky became slightly overcast .
however , estimation of divergence times remains a difficult task that involves substantial uncertainties due to a lack of reliable fossil records or well - dated vicariance events . in this study , we relied on several calibration points that were applied in different combinations . the inferred estimate for the divergence of papilio from the troidini ( up to 100 mya , when all other calibration points were enforced with no constraint age at the tree root ) and the age of subdivisions within papilio ( around 30–50 mya ) fit well with the results of one other insect molecular clock calibration ( gaunt and miles , 2002 ) . these estimates agree with dating of the diversification of the major group of host plants in swallowtails , the rutaceae ( magallón et al . , 1999 ) . with few exceptions , the dates conform with likely vicariant dispersal patterns in the family ( see scriber et al . , 1995 ) and are consistent with known fossil records ( see emmel et al . , 1992 ) .
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 .
papilio remains a taxonomic enigma . few if any papilio subdivisions can be delineated precisely using adult or larval color pattern or other morphological characters ( munroe , 1961 ; hancock , 1983 ; igarashi , 1984 ; miller , 1987 ) . no consistent conclusions on taxonomic rank and species relationships can be derived from the comparison with hybrid incompatibility index ( ae , 1979 ) . although distantly related taxa with up to 15 % nucleotide sequence divergence usually have high incompatibility scores and rare cases of successful matings between species from different subgenera may produce sterile hybrids ( generally in the heterogametic sex ; sperling , 1990 , 2003 ) , crosses between species within the same species group may either completely fail or result in viable and fertile hybrids ( fig . 6 ) . although there is much overlap between ae ' s differentiation index and groupings based on comparisons within species groups , between species groups , and between subgenera , there is much better correspondence between mtdna divergence and these taxonomic groupings . an uncorrected mtdna sequence divergence of about 5 % seems to provide a clear boundary between comparisons within versus between species groups .
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 .
another significant finding is the support for paraphyly of princeps with respect to papilio sensu stricto and eleppone . hancock ( 1983 ) divided princeps into four subgeneric lineages . only one , menelaides , is supported here as a monophyletic taxonomic unit ; a second group , sinoprinceps , is represented here by a single species . hancock ' s other two subgenera , druryia and princeps sensu stricto , were found to be paraphyletic . some clades within princeps , however , are strongly supported : ( 1 ) menelaides ( sensu hancock , 1983 ) ; ( 2 ) the p . demoleus species group ; ( 3 ) achillides ( sensu shimogori , 1997 ) ; ( 4 ) p . nireus species group ; and ( 5 ) the ( ( rex , phorcas ) delalandei ) species groups from subdivisions druryia and princeps . in congruence with another study ( vane - wright et al . , 1999 ) , p . nobilis is strongly associated with this latter group . the present results offer the strongest support yet for relationships among the three species of the phorcas group ; our finding of ( ( dardanus , constantinus ) phorcas ) contrasts with the ( ( dardanus , phorcas ) constantinus ) resolution suggested by vane - wright et al . ( 1999 ) .
our proposed dating suggests slow rates of evolution in mtdna in the genus papilio . substitution rates for the mtdna genes coi and coii and the nuclear gene ef - 1 α are 7 . 8–10 . 2 × 10 −9 and 1 . 32–2 . 0 × 10 −9 per site per year , respectively . these rates are 2–4 ( for mtdna ) and up to 30 ( for ef - 1 α ) times slower than the “standard” substitution rates for coi in drosophila ( 20 . 0–29 . 0 × 10 −9 substitutions per site per year ; beckenbach et al . , 1993 ) and fig - pollinating wasps ( 19 . 0 × 10 −9 ; machado et al . , 2001 ) . however , there is now abundant evidence for a diversity of mtdna rates in insects . low transformation rates have been reported for carabid beetles ( 2 . 8 × 10 −9 substitutions per site per year ; su et al . , 1998 ) , halobates sea skaters ( 4–7 × 10 −9 per year ; andersen et al . , 2000 ) , bacillus phasmids ( 3–4 . 7 × 10 − 9 per year ; mantovani et al . , 2001 ) , and limnoporus water - striders ( 4–4 . 5 × 10 −9 per year ; sperling et al . , 1997 ) . accelerated substitution rates have been reported for the honeybee apis mellifera ( approximately 34 . 0 - 49 . 0 × 10 − 9 ; crozier et al . , 1989 ) and parasitic lice of the genus dennyus ( up to 90 × 10 − 9 ; page et al . , 1998 ) .
southern europe , balkans , asia minor to middle east , middle asia to iran and afghanistan .
in armenia the species distributed in a patchy manner with fairly big population on the slopes of gegham mountains and other smaller populations in central and southern armenia . mostly it inhabits stony semi - deserts , and xerophyte mountain steppes . the elevation range occupied by the species is from 1000 to 2100 m a . s . l .
the species shows steep declining trend ( p < 0 . 01 ) trend in most of the areas of its range in armenia . the host plant is not influenced directly by livestock grazing , although the general degradation of habitat and subsequent erosion might affect it . nevertheless
species in armenia are widely used as an eatable herb , which provokes extensive collection of the plants by local inhabitants . also it was notified that the species abundance shows dependence on average humidity of particular season and therefore can be affected by unsustainable watershed management practices . the trade of the species exists in armenia , however its volume and the level of harm for populations is not well studied .
conservation measures : the species is not evaluated yet for iucn red list , nevertheless it has a status of least concern in european red list . it is included in red book
of animals of armenia as vulnerable and included in appendix ii of bern convention . at current it is protected in zangezur biosphere reserve , in khosrov nature reserve and in gnisheek community protection area . however still some important populations of the species in central armenia are not covered by the network of protected areas . re -
evelopment of the network of prime butterfly area are the first steps of species and habitat conservation , since they can lead to change of the land status into emerald site and therefore to make them internationally protected under bern convention . in addition to that , it is important to study the trade of the species
also it is important to develop sensitive mechanisms of eatable herb collection , as well as sustainable pastureland management in the areas of its distribution .
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van swaay , c . , wynhoff , i . , verovnik , r . , wiemers , m . , lópez munguira , m . , maes , d . , sasic , m . , verstrael , t . , warren , m . & settele , j .
van swaay , c . & cuttelod , a . ( iucn red list unit )
justification : this species is listed as least concern , as it has been evaluated against all the iucn criteria and does not meet , nor is it close to meeting ( e . g . populations have not been declining by more than 25 % in the past 10 years ) , the thresholds for any category of threat .
this species occurs in the southeast of france and just in the adjacent alps of northwestern italy , extreme south of italy ( aspromonte ) and east of sicily , the balkans , croatia ( coastal area ) , albania and large parts of greece including several greek islands . mostly between 300 - 1 , 500 m . its range extends to turkey and the middle east , middle asia to iran and afghanistan . the global distribution area of the species is situated both within and outside europe .
this is a local species , restricted to ( semi - ) natural areas . strong decline in distribution or population size of more than 30 % have been reported from fyr of macedonia . decline in distribution or population size of 6 - 30 % have been reported from croatia and france ( data provided by the national partners of butterfly conservation europe ) .
the southern swallowtail is mostly found on warm , dry calcareous slopes with a flower - rich vegetation and low - growing bushes . they prefer slopes that are steep and rocky . they are especially active during the hottests hours of the day . different foodplants are known , all of them umbellifers . ptychotis saxifraga is the most important one in the western part of its range , but eggs are also laid on opopanax chironium , seseli montanum and trinia glauca . in the eastern part , the caterpillars feed mostly on various fennels ( ferula spp . ) and also on opopanax hispidus , burnet saxifrage ( pimpinella saxifraga ) , scaligeria cretica and wild parsnip ( pastinaca sativa ) . the caterpillars eat the flowers and ripening seeds . they seem to prefer plants growing in very sparse vegetation near bare patches . the southern swallowtail has one generation a year and passes the winter in the pupal stage . habitats : phrygana ( 40 % ) , sclerophyllous scrub ( 20 % ) , heath and scrub ( 20 % ) , dry calcareous grasslands and steppes ( 10 % ) , dry siliceous grasslands ( 10 % ) .
although this species shows a decline in a part of its european range , it is not believed to face major threats at the european scale .
the species is listed on the habitats directive annex 4 and bern convention annex 2 . this species occurs in a number of protected areas across its range . in bosnia and herzegovina , the species only occurs in protected areas . no specific action is needed at a european level , but in countries where the species is declining , important habitats should be protected and managed . the effects should be monitored by butterfly monitoring schemes .
van swaay , c . , wynhoff , i . , verovnik , r . , wiemers , m . , lópez munguira , m . , maes , d . , sasic , m . , verstrael , t . , warren , m . & settele , j . 2010 .
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a very large species , found across the south and southeast mediterranean but it is local and not frequently encountered . it can however be found
casually
with a good dose of luck .
shares its range in europe . the tiger stripes are distinctive and are reminiscent only of the scarce swallowtail ,
, but that butterfly has a ) a white or pale cream ground colour rather than yellow and b ) the tiger stripes are paired .
se france ( plus neighbouring italy ) , greece and its larger islands . low levels , rising to 1000m or more in hills . it flies from april in hotter places , more usually june ( eg samos gr ) into early july ( se france ) . northwest sicily , i ' ve heard there are no recent records .
hilly country , with hot dry slopes . often with open woodland . larvae take various umbelliferae ( carrots ) . frequently takes nectar from thistles and red valarian .
france , is found exceedingly rarely in just north of toulon . it is larger and appears earlier than elsewhere in france . it is very highly threatened , but every few years or so there is a report of an adult or larva found in this very restricted region . over collecting , particularly recently when already very highly threatened , may have played ( and continue to play ) a role in the demise of this subspecies .
near toulon , a region where the ssp . destelensis just may still fly .
2017 photographs highlighted in yellow . click on any photograph to go to an enlarged picture , or simply scroll down the page .
a very localised species in france , with the main localities being in the alpes - de - haute - provence . there are some historic sites in var , which i have visited on numerous occasions , but it may well be that the subspecies
it is principally a species of south - eastern europe , where it is often common , with isolated populations further west .
. the female is usually larger and of a paler yellow . males , borne out by my very limited experience of this species , are more yellow and can be quite small compared with the females . it is univoltine , the flight period being april to july , depending on altitude , although in provence the period seems to be mid - june to mid - july .
but they did this on the wing without stopping . my only chance to get any sort of photograph was to set the camera up to focus on the plant they had been laying on , and wait , and even then i had to use the rapid shoot feature ( 5 frames per second ) and even that did not freeze the subject . so , the photograph 33902 below represented my best effort at this site in 2013 .
was indeed still flying there , still flying non - stop . by pure chance , i had received information as to where
was currently flying in the alpes - maritimes and this spot was on my planned route for the next day . i was fortunate to get the exact location ( thank you , james w ) and was not disappointed . there were several


bucculatrix sexnotata is a moth in the bucculatricidae family .

it is found in north america , where it has been recorded from california , kentucky , maine , new brunswick , north carolina , nova scotia , ohio , ontario , pennsylvania and quebec .

the wingspan is about 7-7.5 mm .

the forewings are dark , almost black , faintly shining .

there are six silvery white spots .

the hindwings are brown .

adults have been recorded on wing in april and from june to september .

the larvae feed on aster species .

they mine the leaves of their host plant .

the mines are very long and thread-like .

older larvae feed freely on the underside of the leaf .

pupation takes place in a pale greyish brown cocoon .

bucculatrix sexnotata

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powell , j . a . & p . a . opler , moths of western north america , pl . 4 . 6m ; p . 57 . book review and ordering
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hodges , r . w . et al . , eds . 1983 . check list of the lepidoptera of america north of mexico . e . w . classey limited and the wedge entomological research foundation , london . 284 pp .
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ridgely , r . s . , t . f . allnutt , t . brooks , d . k . mcnicol , d . w . mehlman , b . e . young , and j . r . zook . 2003 . digital distribution maps of the birds of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
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the green bush squirrel ( paraxerus poensis ) is a species of rodents in the sciuridae family found in benin , cameroon , the republic of the congo , the democratic republic of the congo , ivory coast , equatorial guinea , gabon , ghana , guinea , liberia , nigeria , and sierra leone .

its natural habitats are subtropical or tropical moist lowland forests and plantations .

green bush squirrel

a young / baby of a green bush squirrel is called a ' pup , kit or kitten ' . the females are called ' doe ' and males ' buck ' . a green bush squirrel group is called a ' dray or scurry ' .
the green bush squirrel is listed as least concern . does not qualify for a more at risk category . widespread and abundant taxa are included in this category , on the iucn red list of threatened species
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justification : listed as least concern in view of its wide distribution , presumed large population , it occurs in a number of protected areas , has a tolerance of a degree of habitat modification , and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category .
this species is widely distributed in subsaharan africa , ranging from sierra leone to ghana ( west of the volta basin ) ; then from southeastern nigeria ( east of the niger river ) to eastern congo ; also recorded from three localities in northeastern democratic republic of the congo ( bata , medje , niapa ) . this species is present on the island of bioko ( equatorial guinea ) . it has been recorded up to 1 , 600 m asl on mount bintamane ( sierra leone ) .
rosevear ( 1969 ) noted that this species was the commonest and most widespread of all squirrels in west africa , that it was well represented in museum collections , and not infrequently found as a pet .
this species is found in tropical moist forest , forest fringes , secondary growth around villages , and deserted farmland ( rosevear 1969 ) . it is apparently a pest species in cocoa plantations . despite the fact they seem to be relatively common and widespread , not a great deal is known about this species .
there appear to be no major threats to this widespread and somewhat adaptable species .
this species has been recorded from many protected areas . additional studies are needed into the distribution , abundance , general ecology and threats to this poorly - known species .
this errata assessment has been created because the map was accidentally left out of the version published previously .
( errata version published in 2017 ) . the iucn red list of threatened species 2016 : e . t16211a115132527 .
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a curated database of candidate human ageing - related genes and genes associated with longevity and / or ageing in model organisms .
a curated database of genes associated with dietary restriction in model organisms either from genetic manipulation experiments or gene expression profiling .
projects focused on gene expression profiling of ageing and of dietary manipulations of ageing , such as caloric restriction .
software for ageing research , including the ageing research computational tools ( arct ) perl toolkit .
a curated database of ageing and life history information in animals , including extensive longevity records .
the benchmark genome assembly and annotation of the long - lived , cancer - resistant naked mole - rat ( heterocephalus glaber ) .
a high - coverage genome of the bowhead whale ( balaena mysticetus ) , the longest - lived mammal .
analyses using the anage database to study the evolution of longevity and ageing in vertebrate lineages .
a portal of ageing changes covering different biological levels , integrating molecular , physiological and pathological age - related data .
the whosage database contains people and biotech companies that are contributing to increase our understanding of ageing and life - extension .
little is known about the longevity of these endangered animals , but one specimen lived 7 . 3 years in captivity [ 0671 ] .
[ 0978 ] jones et al . ( 2009 ) , pantheria : a species - level database of life history , ecology , and geography of extant and recently extinct mammals
[ 0455 ] virginia hayssen et al . ( 1993 ) , asdell ' s patterns of mammalian reproduction : a compendium of species - specific data
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thorington , richard w . , jr . , and robert s . hoffman / wilson , don e . , and deeann m . reeder , eds .
mammal species of the world : a taxonomic and geographic reference , 3rd ed . , vol . 2
thorington , richard w . , jr . , john l . koprowski , michael a . steele , and james f . whatton
mammal species of the world : a taxonomic and geographic reference , 2nd ed . , 3rd printing
with contributions by bernadette n . graham , adam p . potter , and mariana m . upmeyer
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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 .
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