Pronophilina

Pronophilina
Montagna Mountain Satyr (Pedaliodes montagna), Arvi Park, Colombia
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Lepidoptera
Superfamily:	Papilionoidea
Family:	Nymphalidae
Subfamily:	Satyrinae
Tribe:	Satyrini
Subtribe:	Pronophilina Reuter, 1896[1]

Pronophilina is a Neotropical subtribe of butterflies of the subfamily Satyrinae. They are a species-rich group with highest diversity in the tropical and subtropical mountains, especially the Andes. Before 1970, they were poorly studied, but recent interest has resulted in high rates of species description from previously unexplored mountain ranges. However, there is still a lack of knowledge on their biology and ecology. Their relationship to other groups of Satyrine butterflies and their complex patterns of speciation within and among mountain ranges have led to several biogeographic discussions.

Systematics and Taxonomy

Traditionally the name Pronophilini (or Pronophilidi) was used to describe a tribe of Neotropical Satyrines,^[1][2] but modern arrangement place them as a subtribe within the Satyrini tribe of the Satyrinae.^[3] The number of genera included in Pronophilina is disputed, since some genera were formally transferred to the Erebiina and Hypocystina (now part of Coenonymphina) subtribes,^{[4] [5]} but some authors reject this arrangement.^[6]

Morphological analysis indicates there is a distinct core group of Pronophilina sensu stricto, and one or two additional groups (Neotropical Erebiina and Hypocistina in their original designation),^[4] but molecular analysis suggest they are each other's sister taxa and form a monophyletic group.^[7]

Illustrations of new species of exotic butterflies Pronophila VIII

By the time Reuter proposed Pronophilidi as a formal tribe, there were some 230 described species.^[4] That number rose to 300 species by 1907, and 370 by 1970, primarily due to the work on museum and collection specimens by (in chronological order) William Chapman Hewitson, Cajetan Freiherr von Felder & Rudolf Felder, Arthur Gardiner Butler, Otto Staudinger, Theodor Otto Thieme and Gustav Weymer.^[3] More detailed field studies in the northern Andes by Adams and Bernard during the 1970s and 1980s resulted in many new taxa descriptions and a better understanding of their distribution and ecology and lead to an increased interest in this group after the 1990s. More than 100 species have been described since 1970, mostly due to contribution from A.L. Viloria, T.W. Pyrcz and G. Lamas,^[3] and it is estimated that the number of known taxa (including several yet unpublished species and subspecies descriptions) has nearly doubled in that period. ^[8][9]

Description

The subtribe Pronophilina can be separated from other American satyrines by the following three external morphological synapomorphies: eyes always densely hairy; HW cross vein m1-m2 always curved or angled basally into the discal cell; maximum length of HW discal cell equal to or longer than half the total maximum length of HW (excluding tails).^[4] These characters separate the Pronophilina sensu stricto from other Neotropical montane satyrids previously included in the group. This arrangement has been adopted by Lamas,^[3] but phylogenetic analysis based on molecular data suggests a larger, more inclusive delimitation of Pronophilina is needed.

The background color of most species is dominated by brown, dark grey or black, with few and slight distinctive features in the wings, but some species show colorful variations between white, yellow, orange, red and iridescent blue.^[4]

Genera

Pedaliodes hewitsoni

Argyrophorus argenteus is distributed in Argentina and Chile

Corades enyo

Steroma superba

Pronophilina sensu stricto[4] Altopedaliodes Forster, 1964 Antopedaliodes Forster, 1964 Apexacuta Pyrcz, 2004 Arhuaco Adams & Bernard, 1977 Calisto (Hübner, 1823) Cheimas Thieme, 1907 Corades Doubleday, [1849] Corderopedaliodes Forster, 1964 Daedalma Hewitson, 1858 Dangond Adams & Bernard, 1979 Dioriste Thieme, 1907 Drucina Butler, 1872 Eretris Thieme, 1905 Eteona Doubleday, 1848 Foetterleia Viloria, 2004 Junea Hemming, 1964 Lasiophila C. & R.Felder, 1859 Lymanopoda Westwood, 1851 Mygona Thieme, 1907 Neopedaliodes Viloria, Miller & Miller, 2004 Oxeoschistus Butler, 1867 Panyapedaliodes Forster, 1964 Paramo Adams & Bernard, 1977 Parapedaliodes Forster, 1964 Pedaliodes Butler, 1867 Pherepedaliodes Forster, 1964 Physcopedaliodes Forster, 1964 Praepedaliodes Forster, 1964 Praepronophila Forster, 1964 Pronophila Doubleday, [1849] Protopedaliodes Viloria & Pyrcz, 1994 Pseudomaniola Röber, 1889 Punapedaliodes Forster, 1964 Redonda Adams & Bernard, 1981 Sierrasteroma Adams & Bernard, 1977 Steremnia Thieme, 1905 Steroma Westwood, [1850] Steromapedaliodes Forster, 1964 Thiemeia Weymer, 1912

Disputed, Pronophilina or Hypocystina(=Coenonymphina)[5] Argyrophorus Blanchard, 1852 Auca Hayward, 1953 Chillanella Herrera, 1966 Cosmosatyrus C. & R.Felder, 1867 Elina Blanchard, 1852 Etcheverrius Herrera, 1965 Faunula C. & R.Felder, 1867 Haywardella Herrera, 1966 Homoeonympha C. & R.Felder, 1867 (including Erebina and Stygnolepis) Nelia Hayward, 1953 Neomaenas Wallengren, 1858 (sometimes including Spinantenna) Neosatyrus Wallengren, 1858 Palmaris Herrera, 1965 Pampasatyrus Hayward, 1953 (including Pseudocercyonis) Pamperis Heimlich, 1959 Punargentus Heimlich, 1963 Quilaphoetosus Herrera, 1966 Spinantenna Hayward, 1953 (sometimes included in NeoMaenas) Tetraphlebia C. & R.Felder, 1867 Disputed, Pronophilina or Erebiina[5] Diaphanos Adams & Bernard, 1981 Ianussiusa Pyrcz & Viloria 2004 Idioneurula Strand, 1932 (including Tamania Pyrcz, 1995) Manerebia Staudinger, 1897 Neomaniola Hayward, 1949 Sabatoga Staudinger, 1897 Stuardosatyrus Herrera & Etcheverry, 1965 Previously in Pronophilina but of uncertain position Amphidecta Butler, 1867 Gyrocheilus Butler, 1867 Catargynnis Röber, 1892 Druphila Pyrcz, 2004 Proboscis Thieme, 1907

Biology

The life cycle of pronophiline butterflies has been scarcely documented. Schultze^[10] described incomplete life histories for Pedaliodes phoenissa (Hewitson), Lymanopoda samius Westwood, and Junea doraete (Hewitson). Other authors have observed oviposition on Chusquea (Poaceae) or other woody bamboos,^[11][12] or loosely over grass dominated vegetation.^[13] Early stages of several species found in Costa Rica were published by DeVries.^[14] Recently, life cycle description have been documented for Parapedaliodes parepa (Hewitson) in Ecuador,^[15] Pedaliodes zingara Viloria & Heredia in Colombia,^[16] Pedaliodes poesia (Hewitson) and Corades medeba Doubleday in Ecuador,^[17][18] and Daedalma dinias emma Pyrcz & Greeney and Daedalma rubroreducta Pyrcz & Willmott.^[8]

Biotic associations

Host plants

Chusquea cumingii Nees by Pato Novoa - 002

All reported host plants are in the Poaceae family, with the genus Chusquea featuring prominently, and a few records in Cynodon, Saccharum, Bambusa, Guada, Rhipidocladum, Merostachys and Zea, among others.^[4][19][20]

Parasitism

Parasitoids in the early stages of pronophiline butterflies have not been properly documented, although they might be locally important.^[4] Incidence of ectoparasitic Diptera (probably Ceratopogonidae) have been documented for seven species of the genera Corades, Lasiophila, Lymanopoda, Mygona and Pedaliodes.^[21]

Mimicry

There are 18 documented examples of convergent coloration patterns between coexisting pairs of pronophiline species from different genera (three examples), between pronophiline species and other satyrines (eight examples), and between pronophiline species and other butterflies or skippers (seven examples).^[4] Most examples involve species of Lymanopoda or Eretris. Some of these observations have been described as mimetic relationships, but the degree of resemblance is not so accurate as in other mimetic butterfly groups, there is no direct evidence of unpalatability of Pronophiline butterflies, and no clear understanding of the ecological consequences of such resemblance.^[4]

Diversity

Depending on the classification adopted, the pronophilini include between 592 and 711 species.^[3][4] This represents approximately 50-60% of the Neotropical,^[3] and 23-27% of the worldwide Satyrinae.^[22]

Most species have a geographically and altitudinally restricted distribution in the tropical and subtropical Andes, and other mountain ranges in Brasil, the Guayana Shield, Central America, and México. The genus Calisto has a Caribbean distribution, and some other genera are distributed in Patagonia.^[4]

Biogeography

The Pronophilines have been involved in diverse biogeographical discussions that aim to explain their current distribution, diversity and endemism.^[23] The high diversity within the Pronophilina, and the distinctive speciation patterns in mountain and caribbean taxa, has also motivated discussion about parapatric and sympatric speciation.^[24][25]

Origins

The affinities of the Pronophilina to other species-groups within the specious Satyrinae suggest different interpretations about its origin. Viloria^[4] reviewed morphological characters and separated three groups that might have different phylogenetic and biogeographic affinities: a major group of 39 true Neotropical Pronophilina, including the genus Calisto with a Caribbean distribution, eight genera related to the Holarctic Erebiina, and 19 genera which he placed as Neotropical representatives of the otherwise Australasian Hypocystina. This grouping would suggest an amphipacific distribution of the Hypocystina, and imply a Gondwanan origin according to a panbiogegraphic interpretations.^[23] This hypothesis was rebutted after a large phylogenetic analysis of the Satyrinae recovered a representative sample of Pronophiline genera as a monophyletic group, and suggested a completely Neotropical, or at least, a common origin for the group.^[7] However, the relationships of the group remain unresolved, as subsequent work has found that at least two genera -- Eretris, and the Caribbean Calisto—might be closer related to Holarctic Satyrines,^[26] but larger taxonomic sampling and a better integration of molecular and morphological data is needed to rule out analytical artifacts.^[6]

Speciation

It has been observed that most mountain species, especially in the species-rich genera close to Pedaliodes, have very narrow altitudinal ranges, and are restricted to one or few mountain ranges. This leads to high turn-over of species along altitudinal gradients within a mountain range, and high turn-over between ranges.^[24] Most species appear to be more closely related to those occupying a similar altitudinal range in neighboring mountains, than to the species below or above.^[27] A possible mechanism was proposed by Adams^[24] and extensively discussed by Viloria.^[27] This involves a series of alternative events of colonization and isolation that would be linked to repeated cycles of cold-wet and warm-dry climate (glaciations and interglaciation periods).

The genus Calisto also shows a distinctive radiation in the Caribbean, where it is the only representative of the Satyrinae.^[25] At least 54 named taxa have been recognized, most of them restricted to particular habitats in the island of Hispaniola, with fewer species in Cuba, Jamaica and Puerto Rico.^[3] The diversification of Calisto is referred to as an example of adaptive radiation into contrasting habitat types.^[25]

References

1. E.R. Reuter 1896 Über die Palpen der Rhopaloceren: ein Beitrag zur Erkenntnis der verwandtschaftlichen Beziehungen unter den Tagfaltern Acta Societas Scientiarum Fennica 22: 1-577.
2. L.D. Miller 1968 The higher classification, phylogeny, and zoogeography of the Satyridae (Lepidoptera) Memories of the American Entomological Society 24: 1-174.
3. Lamas, G. 2004 Atlas of Neotropical Lepidoptera. CheckList: Part 4A Hesperioidea-Papilionoidea Scientific Publishers.
4. A.L Viloria 2007 The Pronophilini: Synopsis of their Biology and Systematics Tropical Lepidoptera 2004(15): 1–17.
5. A.L. Viloria 2007 Some Gondwanan and Laurasian elements in the satyrine fauna of South America (Lepidoptera: Nymphalidae: Satyrinae) Tropical Lepidoptera (2004)15: 53-55.
6. MA Marín , C. Peña , AVL Freitas , N. Wahlberg , and SI Uribe 2011 From the Phylogeny of the Satyrinae Butterflies to the Systematics of Euptychiina (Lepidoptera: Nymphalidae): History, Progress and Prospects Neotropical Entomology 40: 1--13.
7. C. Peña , N. Wahlberg , E. Weingartner , U. Kodandaramaiah , S. Nylin , A.V.L. Freitas , and A.V.Z. Brower 2006 Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data Molecular Phylogenetics and Evolution 40: 29–49.
8. T.W. Pyrcz , H.F. Greeney , K.R. Willmott , and J. Wojtusiak 2011 A taxonomic revision of the genus Daedalma Hewitson with the descriptions of twenty new taxa and the immature stages of two species (Lepidoptera: Nymphalidae: Satyrinae) Zootaxa 2898: 1–68.
9. Viloria, Ángel L.; Ferrer-Paris, José R.; Camacho, Jesús; Costa, Mauro (2013) [2015]. "New satyrine butterflies from the Venezuelan Andes (Lepidoptera: Nymphalidae)" (PDF). Anartia. 25: 95–160. doi:10.5281/zenodo.18332.
10. A. Schultze 1929 Die ersten Stände von drei kolumbianischen hochandinen Satyriden Deutsche Entomologische Zeitschrift 'Iris' 43: 157–165, 3 plates.
11. M.J. Adams 1986 Pronophiline butterflies (Satyridae) of the three Andean Cordilleras of Colombia Zoological Journal of the Linnean Society 87: 235–320.
12. T.W. Pyrcz , K.R. Willmott , and J.P. Hall 1999 Contributions to the knowledge of Ecuadorian Pronophilina, Part 3, Three new species and five new subspecies of Lymanopoda (Lepidoptera: Nymphalidae: Satyrinae) Genus 10: 497–522.
13. Viloria, A. L.; Pyrcz, T. W.; Wojtusiak, J.; Ferrer-Paris, J. R.; Beccaloni, G. W.; Sattler, K.; Lees, D. C. (7 August 2003). "A brachypterous butterfly?". Proceedings of the Royal Society B: Biological Sciences. 270 (Suppl_1): S21–S24. doi:10.1098/rsbl.2003.0015.
14. P. J DeVries 1987 The butterflies of {Costa Rica} and their natural history. Papilionidae, {Pieridae}, Nymphalidae Princeton University PressPrinceton.
15. V. Pelz 1997 Life history of Pedaliodes parepa from Ecuador (Lepidoptera: Nymphalidae: Satyrinae) Tropical Lepidoptera 8: 41–45.
16. M.D. Heredia and A.L. Viloria 2004 Description and life history of Pedaliodes zingara, a new satyrine species from Colombia Journal of the Lepidopterist’s Society 58: 80–87.
17. H. Greeney , T.W. Pyrcz , L.A. Dyer , and M.Z. Sanches 2010 The early stages and natural history of Corades medeba Hewitson, 1850 in eastern Ecuador (Lepidoptera, Nymphalidae, Satyrinae, Pronophilina) Tropical Lepidoptera Research 20: 8–13.
18. H. Greeney , T.W. Pyrcz , P.J. DeVries , and L.A. Dyer 2009 The early stages of Pedaliodes poesia (Hewitson, 1862) in eastern Ecuador (Lepidoptera: Satyrinae: Pronophilina) Journal of Insect Science 9: 1–9.
19. George W. Beccaloni, Angel Luis Viloria, Stephen R. Hall, and Gaden S. Robinson 2008 Catalogue of the hostplants of the Neotropical butterflies. Catálogo de las plantas huésped de las mariposas neotropicales RIBES-CYTED / Sociedad Entomológica AragonesaZaragoza.
20. Ferrer-Paris, José R.; Sánchez-Mercado, Ada; Viloria, Ángel L.; Donaldson, John (2013). "Congruence and Diversity of Butterfly-Host Plant Associations at Higher Taxonomic Levels". PLoS ONE. 8 (5): e63570. doi:10.1371/journal.pone.0063570. Retrieved 29 June 2016.
21. R. Lichy 1946 Documents pour servir a l'etude des Lépidoptérès du Vénézuéla (3e Note). Un cas de parasitisme sur les ailes des Lépidoptérès Bol. Ent. Venez. 5: 1-4.
22. G. Lamas 2008 La sistemática sobre mariposas (Lepidoptera: Hesperioidea y Papilionoidea) en el mundo: estado actual y perspectivas futuras ': 57 -- 70.
23. A.L. Viloria 2003 Historical biogeography and the origins of the satyrine butterflies of the tropical Andes (Lepidoptera: Rhopalocera), In: Llorente J. Morrone, J.J. (Eds.) Una perspectiva latinoamericana de la biogeografía Universidad Autónoma de México, México.
24. Michael J Adams 1985 Speciation in the pronophiline butterflies (Template:Satyridae) of the northern {Andes} Journal of Research on the Lepidoptera Supl. 1: 33-49..
25. Andrei Sourakov and Evgeny Zakharov 2011 “Darwin’s butterflies”? DNA barcoding and the radiation of the endemic Caribbean butterfly genus Calisto (Lepidoptera, Nymphalidae, Satyrinae) Comparative Cytogenetics 5: 191--210.
26. N. Wahlberg , J. Leneveu , U. Kodandaramaiah , C. Peña , S. Nylin , A. V. L. Freitas , and A. V. Z. Brower 2009 Nymphalid butterflies diversify following near demise at the Cretaceous/Tertiary boundary Proceedings of the Royal Society of London Series B Biological Sciences 276: 4295--4302.
27. A.L. Viloria 1999 Studies on the systematics and biogeography of some montane satyrid butterflies (Lepidoptera). Imperial College, London. PhD Thesis