Duke of Burgundy Hamearis lucina, copyright Gilles San Martin.

Belongs within: Obtectomera.
Contains: Nymphidiini.

The Riodinidae, metalmarks, are a group of butterflies that are most diverse in the Neotropical region. Authors have differed whether this family should be included as a subgroup of the related Lycaenidae.

Characters (from Braby 2000, as Riodininae): Male fore leg non-functional, less than half length of mid and hind legs, tarsus fused into single segment, rarely with claws; hind wing usually with basal vein along costal margin, tornus may be produced, lobed or dentate but rarely tailed; males rarely with sex-scales. Larva with prothorax bearing pair of vibratory papillae, metathorax bearing pair of dorsolateral eversible organs, pair of eversible nectary organs present on segment 8.

<==Riodinidae [Nemeobiidae]
    |--Hamearinae [Nemeobiinae]J03
    |    |--AbisaraJ03
    |    |--ZemerosJ03
    |    `--Hamearis lucina [=Nemeobius lucina]RD77
         |  i. s.: Praetaxila Fruhstorfer 1914B00
         |           `--P. segecia (Hewitson 1861)B00
         |                |--P. s. segeciaB00
         |                |--P. s. punctaria (Fruhstorfer 1904)B00
         |                `--P. s. yaniya Fruhsorfer 1914B00
         |         Styx [Styginae]B00
         |           `--S. infernalis Staudinger 1875B00
         |         Semomesia capaneaKP19
         |         ApodemiaS72
         |           |--A. mormoHH02
         |           |    |--A. m. mormoUSDI77
         |           |    |--A. m. langeiUSDI77
         |           |    `--A. m. tuolumnensisS72
         |           `--A. naisS72
         |         EmesisS72
         |           |--E. guttata [=Audre guttata]HH02
         |           |--E. mandana (Cramer 1780)S55
         |           |--E. xanthosaHH02
         |           `--E. zelaS72
         |         LepheliscaS72
         |           |--L. borealisS72
         |           `--L. virginiensisS72
Riodinidae incertae sedis:
  Petrocerus catienaHH02
  Calydna caietaHH02
    |--P. emyliusS55
    |    |--P. e. emyliusS55
    |    `--P. e. crispinella Stichel 1911S55
    `--P. lasthenesFS54
         |--P. l. lasthenesFS54
         `--P. l. zeurippa Boisduval 1836FS54
  Calephelis virginiensisFS54
    |--C. v. virginiensisFS54
    `--C. v. laverna (Godman & Salvin 1886)FS54
    |--L. cephise (Ménétriés 1855)FS54
    `--L. pixe (Boisduval 1910)FS54
  Esthemopsis caeruleata (Godman & Salvin 1878)FS54
    |--M. jeziellaS55
    |    |--M. j. jeziellaS55
    |    `--M. j. thyestes Druce 1878S55
    `--M. judicialis Butler 1874S55
  Lyropteryx apollonia Westwood 1851S55
  Cyrenia martia Westwood 1851S55
    |--A. aulestesS55
    |    |--A. a. aulestesS55
    |    `--A. a. eryxo (Saunders 1858)S55
    `--A. incaGE05
    |--R. arciusGE05
    `--R. periander (Cramer 1777)S55
    |--C. mantinea (Felder 1861)S55
    `--C. trochilusS55
         |--C. t. trochilusS55
         `--C. t. arete (Felder 1861)S55
  Baeotis nesaeaS55
    |--B. n. nesaeaS55
    `--B. n. bacaenita Schaus 1902S55
  Amarynthis meneria [incl. A. meneria f. maecenas (Fabricius 1793), A. meneria f. micalia (Cramer 1776)]S55
  Peplia lamis [incl. P. lamis f. molpe (Hübner 1816)]S55
    |--H. acisR13
    `--H. cupidoGE05
  Riodinella nymphaGE05
    |--V. danforthiV-W04
    |--V. drambaGE05
    `--V. umbraV-W04
  Mesenopsis melanochloraGE05
    |--Alesa amesisHH02
         |--E. nicaea (Fabricius 1775)S55
         `--E. patronaGE05

*Type species of generic name indicated


[B00] Braby, M. F. 2000. Butterflies of Australia: their identification, biology and distribution vol. 2. CSIRO Publishing: Collingwood (Victoria).

[FS54] Franz, E., & H. Schröder. 1954. Tagfalter (Lep. Rhopalocera) aus El Salvador. Senckenbergiana Biologica 35: 75–87.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[HH02] Hall, J. P. W., & D. J. Harvey. 2002. Basal subtribes of the Nymphidiini (Lepidoptera: Riodinidae): phylogeny and myrmecophily. Cladistics 18: 539–569.

[J03] Jong, R. de. 2003. Are there butterflies with Gondwanan ancestry in the Australian region? Invertebrate Systematics 17: 143–156.

[KP19] Kawahara, A. Y., D. Plotkin, M. Espeland, K. Meusemann, E. F. A. Toussaint, A. Donath, F. Gimnich, P. B. Frandsen, A. Zwick, M. dos Reis, J. R. Barber, R. S. Peters, S. Liu, X. Zhou, C. Mayer, L. Podsiadlowski, C. Storer, J. E. Yack, B. Misof & J. W. Breinholt. 2019. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the USA 116 (45): 22657–22663.

[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms’ General Textbook of Entomology 10th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[S55] Schröder, H. 1955. Eine Falter-Ausbeute aus dem westlichen Bolivien. (Ins. Lepid. Rhopal.) Senckenbergiana Biologica 36: 329–338.

[S72] Shields, O. 1972. Flower visitation records for butterflies (Lepidoptera). Pan-Pacific Entomologist 48 (3): 189–203.

[USDI77] United States Department of the Interior. 1977. Endangered and threatened wildlife and plants—republication of list of species. Federal Register 42: 36420–36431.

[V-W04] Vane-Wright, D. 2004. Butterflies at that awkward age. Nature 428: 477–480.

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