Editing Telegonus fulgerator

{{Short description|Species of butterfly}}
{{Speciesbox
| name = Two-barred flasher
| image = Two-barred flasher (Astraptes fulgerator).jpg
| taxon = Telegonus fulgerator
| authority = ([[Johann Ernst Immanuel Walch|Walch]], 1775)
| synonyms = * ''Astraptes fulgerator'' <small>(Walch, 1775)</small>
}}

'''''Telegonus fulgerator''''', the '''two-barred flasher''', is a species of [[Spread-winged skipper|spread-wing]] [[skipper (butterfly)|skipper]] [[butterfly]] in the family [[Hesperiidae]] which may constitute a possible [[cryptic species complex]]. It ranges all over the [[Americas]], from the southern United States to northern Argentina.

==Description==
{{Lepidopteran glossary hatnote}}

''Telegonus fulgerator'' is a mid-sized skipper butterfly with the wing shape typical of this group. The upperside is black, with basal to postbasal blue corners which are more extensive on the forewings. There is one discal-tomal{{clarify|date=June 2020|reason=both missing in glossary}} and one apical band on the forewing; these are usually off white to light blue but the former may be quite white towards the costal margin. The [[Thorax (insect anatomy)|thorax]] has bluish hair on the back, the underside is yellow to orange.<ref name = hebertetal2004>Hebert ''et al.'' (2004)</ref>

The [[caterpillar]]s and [[pupae]] show a wide range of colors and patterns, and the caterpillars also vary in food preference. Last-[[instar]] caterpillars are black with a pattern consisting of light to bright yellow dots along the sides, or rings of varying thickness, sometimes interrupted on the back, in a range of colors varying from white to orange red.<ref name="hebertetal2004"/>

===Larval food plants===
This species is highly [[phagy|polyphagous]], with most food plants belonging to the [[Fabaceae]] (legume [[family (biology)|family]]):

'''Selected primary food plants'''<ref name = nwcr>[[Area de Conservación Guanacaste World Heritage Site]], NW [[Costa Rica]]: Hébert ''et al.'' (2004)</ref>
* [[Fabaceae]]
** ''[[Inga]]'': ''[[Inga exalata|I. exalata]]'', ''[[Inga oerstediana|I. oerstediana]]'', ''[[Inga punctata|I. punctata]]'', ''[[Inga sapindoides|I. sapindoides]]'', ''[[Inga vera|I. vera]]'' and probably others
** ''[[Lonchocarpus]]'' (see also [[#Cryptic variation|below]])
** ''[[Senna (plant)|Senna]]'': [[candle bush]] (''S. alata''), ''[[Senna cobanensis|S. cobanensis]]'', ''[[Senna hayesiana|S. hayesiana]]'', ''[[Senna pallida|S. pallida]]'', ''[[Senna papillosa|S. papillosa]]'', ''[[Senna undulata|S. undulata]]'' and probably others<ref>Hébert ''et al.'' (2004) refer to ''"Cassia emarginata"'', which today is either of ''[[Chamaecrista pilosa]]'', [[Rambling Senna]] (''S. bicapsularis'') or ''[[Senna candolleana]]''. The last does not occur in their study area; given the general importance of ''Senna'' species and the lack of records for ''Chamaecrista'' species as food plants in the study area, ''S. bicapsularis'' seems to be the plant in question.</ref>
* [[Malvaceae]]
** ''[[Hampea appendiculata]]'' (but see [[#Cryptic variation|below]])
* [[Sapindaceae]]
** ''[[Cupania]]'': ''[[Cupania glabra|C. glabra]]'', ''[[Cupania guatemalensis|C. guatemalensis]]'' and probably others

'''Selected secondary and accidental food plants'''<ref name = nwcr />
* [[Capparaceae]]
** ''[[Capparis frondosa]]''
* [[Fabaceae]]
** ''[[Canavalia brasiliensis]]''
** ''[[Cassia (genus)|Cassia]]'': [[golden shower tree]] (''C. fistula''), ''[[Cassia grandis|C. grandis]]'' and possibly others
** ''[[Centrosema]]'' (butterfly peas): ''[[Centrosema macrocarpum|C. macrocarpum]]'', ''[[Centrosema plumieri|C. plumieri]]'' and possibly others
** ''[[Desmodium glabrum]]''
** ''[[Dioclea (plant)|Dioclea]]'': ''[[Dioclea malacocarpa|D. malacocarpa]]'', ''[[Dioclea violacea|D. violacea]]'' and possibly others
** ''[[Mucuna holtonii]]''
** ''[[Piscidia carthagenensis]]''
* [[Malvaceae]]
** ''[[Byttneria catalpaefolia]]''
* [[Rhamnaceae]]
** ''[[Karwinskia calderoni]]''
* [[Salicaceae]]
** ''[[Casearia sylvestris]]''
* [[Styracaceae]]
** ''[[Styrax argenteus]]''

==Systematics==
Due to the diversity of caterpillar colors and food plants it was suspected that the butterflies referred to as ''Telegonus fulgerator'' might be more than a single species. A controversial 2004 [[DNA barcode|DNA barcoding]] study of a 648 [[base pair]] sequence from the [[Cytochrome c oxidase]] (COI) [[DNA sequence]] by [[Paul D. N. Hebert|Paul D. N. Hébert]] ''et al''. led the authors to claim that at least ten [[sympatric]] populations in the [[Area de Conservación Guanacaste World Heritage Site]] in north-western Costa Rica were in various stages of [[reproductive isolation]].<ref name = hebertetal2004 /> However, a later reanalysis of the same DNA sequence data using [[neighbor joining]] [[Bootstrapping (statistics)|bootstrap]], population aggregation analysis and [[cladistic]] [[haplotype]] analysis found that: "[a]t least three, but not more than seven [[mtDNA]] clades that may correspond to cryptic species are supported by the evidence". Also, where a specific DNA sequence did not correspond to its purported host plant, it was "[[Fabrication (science)|simply dismissed]] with an [[ad hoc hypothesis]] and manifestly incorrect explanation" by Hébert and his coworkers. The improper use of [[Taxonomy (biology)|taxonomic]] vocabulary was also criticized; Hébert ''et al''. apply the terms "[[species]]" and "[[taxa]]" as if they were synonymous, but never actually validly described their proposed "species".<ref name = brower2006>Brower (2006)</ref>

{{gallery |mode=packed |height=130
|File:Two-barred flasher (Astraptes fulgerator azul).jpg|''A. f. azul''<br>Costa Rica
|File:Two-barred flasher (Astraptes fulgerator azul) underside.jpg|''A. f. azul''<br>Costa Rica
}}

===Cryptic variation===
The exact number of taxa involved is disputed, most of the "species" detected by the [[DNA barcoding]] study seem to be nothing more than [[Morph (zoology)|morph]]s or incipient [[subspecies]], coupled with a serious underestimation of variation. Still, two lineages appear to be well distinct and separable at least as [[subspecies]]:<ref name = brower2006 /> "CELT" has larvae with bold orange bands in the last instar, which were recorded only on ''[[Celtis iguanaea]]'' ([[Ulmaceae]]); "TRIGO" last-instar larvae have bold yellow bands and were found on the [[Malpighiales]] ''[[Trigonia (plant)|Trigonia]]'' (''[[Trigonia arborea|T. arborea]]'', ''[[Trigonia laevis|T. laevis]]'' and ''[[Trigonia rugosa|T. rugosa]]'') and, apparently accidentally, on ''[[Licania arborea]]''.<ref name = hebertetal2004 />

Three more lineages are in need of further study. One, "NUMT", was initially dismissed as a [[numt]] [[pseudogene]] combined with sequencing error<ref name = hebertetal2004 /> but may represent a hitherto unrecognized taxon.<ref name = brower2006 /> Two other lineages, "LOHAM" and "LONCHO", were considered highly distinct in the barcoding study<ref name = hebertetal2004 /> but the reanalysis showed that this might be an error.<ref name = brower2006 /> The latter two are peculiar in some aspects, such as apparently never having bands or orange colors in the last instar stage and showing a preference for ''[[Lonchocarpus costaricensis]]'', ''[[Lonchocarpus oliganthus]]'', and ''[[Hampea appendiculata]]'' as larval food but are not [[phagy|monophagous]]. They appear to be an intermediate stage in [[lineage sorting]] and might be considered one or two subspecies if the two most distinct lineages are split off as species.<ref name = hebertetal2004 /><ref name = brower2006 />

The other lineages show a marked lack of agreement between morphological, ecological and genetical variation in the reanalysis of the supposed clusters. The entire range of caterpillar colors and patterns is found across one huge ill-structured cluster of genetic diversity. They are [[phagy|polyphagous]], feeding preferentially on ''[[Inga]]'' and ''[[Senna (plant)|Senna]]'' as well as a variety of other plants, but apparently not on those preferred by the more distinct lineages except ''[[Hampea appendiculata]]''.<ref name = brower2006 />

The proposed divergence times for the lineages are derived from a standard [[molecular clock]] model, which is today known to be incorrect.<ref name = brower2006 />

==Footnotes==
{{Reflist}}

==References==
* {{aut|Brower, Andrew V.Z.}} (2006): Problems with DNA barcodes for species delimitation: 'ten species' of ''Astraptes fulgerator'' reassessed (Lepidoptera: Hesperiidae). ''Systematics and Biodiversity'' '''4'''(2): 127–132. <small>{{doi|10.1017/S147720000500191X}}</small> [https://web.archive.org/web/20081217031406/http://www.bolinfonet.org/pdf/Brower_2006SYSTBIODIVERSITY%5B1%5D.pdf PDF fulltext]
* {{aut|[[Paul D. N. Hebert|Hébert, Paul D.N.]]; Penton, Erin H.; Burns, John M.; Janzen, Daniel H. & Hallwachs, Winnie}} (2004): Ten species in one: DNA barcoding reveals cryptic species in the semitropical skipper butterfly ''Astraptes fulgerator''. ''[[PNAS]]'' '''101'''(41): 14812-14817. <small>{{doi|10.1073/pnas.0406166101}}</small> [http://www.pnas.org/cgi/reprint/101/41/14812.pdf PDF fulltext] [http://www.pnas.org/cgi/content/full/0406166101/DC1 Supporting Appendices]

==External links==
* [https://web.archive.org/web/20110612145256/http://www.barcodinglife.com/views/imagecomparison.php?& Gallery of specimens] as per Hébert ''et al.'' (2004). Retrieved 2007-DEC-17.

{{Taxonbar|from1=Q97672923|from2=Q3503407}}

[[Category:Astraptes]]
[[Category:Butterflies of North America]]
[[Category:Butterflies described in 1775]]
[[Category:Hesperiidae of South America]]
[[Category:Taxa named by Johann Ernst Immanuel Walch]]