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Volume 26: 102–108 METAMORPHOSIS www.metamorphosis.org.za ISSN 1018–6490 (PRINT) LEPIDOPTERISTS’ SOCIETY OF AFRICA ISSN 2307–5031 (ONLINE) Classification of the Afrotropical butterflies to generic level Published online: 25 December 2015 Mark C. Williams 183 van der Merwe Street, Rietondale, Pretoria, South Africa. E-mail: [email protected] Copyright © Lepidopterists’ Society of Africa Abstract: This paper applies the findings of phylogenetic studies on butterflies (Papilionoidea) in order to present an up to date classification of the Afrotropical butterflies to genus level. The classification for Afrotropical butterflies is placed within a worldwide context to subtribal level. Taxa that still require interrogation are highlighted. Hopefully this classification will provide a stable context for researchers working on Afrotropical butterflies. Key words: Lepidoptera, Papilionoidea, Afrotropical butterflies, classification. Citation: Williams, M.C. (2015). Classification of the Afrotropical butterflies to generic level. Metamorphosis 26: 102–108. INTRODUCTION Suborder Glossata Fabricius, 1775 (6 infraorders) Infraorder Heteroneura Tillyard, 1918 (34 Natural classifications of biological organisms, based superfamilies) on robust phylogenetic hypotheses, are needed before Clade Obtectomera Minet, 1986 (12 superfamilies) meaningful studies can be conducted in regard to their Superfamily Papilionoidea Latreille, 1802 (7 evolution, biogeography, ecology and conservation. families) Classifications, dating from the time of Linnaeus in the Family Papilionidae Latreille, 1802 (32 genera, 570 mid seventeen hundreds, were based on morphology species) for nearly two hundred and fifty years. Classifications Family Hedylidae Guenée, 1858 (1 genus, 36 species) based on phylogenies derived from an interrogation of Family Hesperiidae Latreille, 1809 (570 genera, 4113 the genome of individual organisms began in the late species) 20th century. In less than 50 years an increasing Family Nymphalidae Rafinesque, 1815 (559 genera, number of progressively more sophisticated studies, 6152 species some incorporating both morphology and molecules, Family Pieridae Swainson, 1820 (91 genera, 1164 have produced robust phylogenies for many taxa in the species) order Lepidoptera. Perhaps the most exciting recent Family Riodinidae Grote, 1895 (146 genera, 1532 development is the placement of the “butterfly” species) superfamilies Papilionoidea, Hesperioidea and Family Lycaenidae Leach, 1815 (416 genera, 5201 Hedylidae within the Lepidoptera (Heikkila et al., species) 2012; Kawahara & Breinholt, 2014; Mutanen et al., 2010; Regier et al., 2013; Wahlberg et al., 2013). The order of the families above is modified from that given by Van Nieukerken et al., (2011) to reflect the RESULTS findings of Heikkila et al., 2012. In their study the latter authors found that, within Papilionoidea, Superfamilies and families Papilionidae is the sister-group to Hedyloidea, Hesperioidea, and all other butterflies. Hence, the The position of the seven families considered to reside families in the current three superfamilies should be in the Superfamily Papilionoidea Latreille, [1802] in placed in a single family Papilionoidea. They also the Order Lepidoptera Linnaeus, 1758 were given in found that Hedylidae is sister to Hesperiidae. Their the consensus of Van Nieukerken et al., 2011, as parsimony analysis showed Papilionidae as basal, follows: followed by Hedylidae + Hesperiidae, Nymphalidae, Pieridae and Riodinidae + Lycaenidae. They used 71 Order Lepidoptera Linnaeus, 1758 (4 suborders) butterfly species and included all seven families, the (15,578 genera, 157,424 species) only recent study to have done so. Received: 15 December 2015 The Superfamily Papilionoidea has a total of about Accepted: 25 December 2015 19000 species in about 1820 genera. In the Copyright: This work is licensed under the Creative Commons Afrotropical Region there are 4325 species (about 23% Attribution-NonCommercial-NoDerivs 3.0 Unported License. of the world total) in 318 genera (about 17% of the To view a copy of this license, send a letter to Creative world total). Commons, Second Street, Suite 300, San Francisco, California, 94105, USA, or visit: http://creative commons.org/licenses/by- nc-nd/3.0/ Willliams / Metamorphosis 26: 102–108 103 FAMILY PAPILIONIDAE Latreille, [1802] & Larsen, 2014; Apallaga Strand, 1911; Eretis Mabille, 1891; Sarangesa Moore, [1881]; Alenia This is the second smallest family (after the Hedylidae) Evans, 1935; Triskelionia Larsen & Congdon, in the Papilionoidea. The family is cosmopolitan, with 2011) about 570 species in 32 genera. There are 101 species Tribe Carcharodini Verity, 1940 (Afrotropical in 3 genera in the Afrotropical Region. A robust genera: Gomalia Moore, 1879; Carcharodus phylogeny for the family, to subtribal level, has been Hübner, [1819]; Spialia Swinhoe, [1912]) hypothesized and is given below. The following Tribe Erynnini Brues & Carpenter, 1932 publications were consulted: Braby et al., 2005 Tribe Achlyodidini Burmeister, 1878 (Troidini); Condamine et al., 2012 (Papilionidae); Tribe Pyrgini Burmeister, 1878 Nazari et al., 2007 (Parnassinae); Simonsen et al., Subfamily Heteropterinae Aurivillius, [1925] 2011 (Papilioninae). Afrotropical taxa are highlighted (Afrotropical genera: Metisella Hemming, 1934; in yellow. Hovala Evans, 1937) Subfamily Trapezitinae Waterhouse & Lyell, 1914 Subfamily Baroniinae Bryk, 1913 Subfamily Hesperiinae Latreille, 1809 Subfamily Parnassiinae Duponchel, [1835] Tribe Aeromachini Tutt, 1906 (Afrotropical Tribe Parnassiiini Duponchel, [1835] genus: Ampittia Moore, [1881]) Tribe Luehdorfiini Chapman, 1895 Incertae sedis (Afrotropical genera: Tsitana Tribe Zerynthiini Grote, 1899 Evans, 1937; Lepella Evans, 1937; Astictopterus Subfamily Papilioninae Latreiile, [1802] Felder & Felder, 1860; Arnetta Watson, 1893; Tribe Leptocircini Kirby, 1896 (Afrotropical Prosopalpus Holland, 1896; Kedestes Watson, genus: Graphium Scopoli, 1777) 1893; Fulda Evans, 1937; Gorgyra Holland, Tribe Teinopalpini Grote, 1899 1896; Gyrogra Lindsey & Miller, 1965; Tribe Troidini Talbot, 1939 Teniorhinus Holland, 1892; Heriila Larsen & Subtribe Battina Munroe & Ehrlich, 1960 Collins, 2012; Ceratrichia Butler, 1870; Subtribe Troidina Talbot, 1939 Ceratricula Larsen, 2013; Flandria Larsen, 2013; (Afrotropical genus: Pharmacophagus Pardaleodes Butler, 1870; Hollandus Larsen & Haase, 1891) Collins, 2015; Ankola Evans, 1937; Xanthodisca Tribe Papilionini Latreille, [1802] (Afrotropical Aurivillius, [1925]; Acada Evans, 1937; genus: Papilio Linnaeus, 1758) Rhabdomantis Holland, 1896; Osmodes Holland, FAMILY HESPERIIDAE Latreille, 1809 1892; Parosmodes Holland, 1896; Osphantes Holland, 1896; Acleros Mabille, 1885; Paracleros This family is cosmopolitan, with about 4, 150 species Berger, 1978; Semalea Holland, 1896; Hypoleucis in 570 genera. There are 612 species in 84 genera in Mabille, 1891; Meza Hemming, 1939; Paronymus the Afrotropical Region. A phylogeny for the family as Aurivillius, [1925]; Andronymus Holland, 1896; a whole was recently hypothesized by Warren and Malaza Mabille, [1904]; Perrotia Oberthür, 1916; coworkers (Warren et al., 2008, 2009). Within the Ploetzia Saalmüller, 1884; Moltena Evans, 1937; subfamily Hesperiinae, which contains the majority of Chondrolepis Mabille, [1904]; Zophopetes species in the family, the tribal placement of a large Mabille, [1904]; Gamia Holland, 1896; Artitropa number of genera is ambiguous. These genera are Holland, 1896; Mopala Evans, 1937; Gretna therefore currently designated as incertae sedis. Evans, 1937; Pteroteinon Watson, 1893; Leona Evans, 1937; Caenides Holland, 1896; Monza Subfamily Coeliadinae Evans, 1937 (Afrotropical Evans, 1937; Melphina Evans, 1937; Melphinyet genera: Coeliades Hübner, [1818]; Pyrrhiades Larsen, 2012; Noctulana Larsen, 2012; Fresna Lindsey & Miller, 1965; Pyrrhochalcia Mabille, 1904) Evans, 1937; Platylesches Holland, 1896; Subfamily Euschemoninae Kirby, 1897 Erionota Mabille, 1878) Subfamily Eudaminae Mabille, 1877 Tribe Baorini Doherty, 1886 (Afrotropical Subfamily Pyrginae Burmeister, 1878 genera: Brusa Evans, 1937; Zenonia Evans, 1935; Tribe Pyrrhopygini Mabille, 1877 Gegenes Hübner, [1819]; Parnara Moore, [1881]; Subtribe Pyrrhopygina Mabille, 1877 Borbo Evans, 1949; Pelopidas Walker, 1870) Subtribe Oxynetrina Mielke, 2001 Tribe Taractrocerini Voss, 1952 Subtribe Passovina Mielke, 2001 Tribe Thymelicini Tutt, 1905 Subtribe Zoniina Mielke, 2001 Tribe Calpodini Clark, 1948 Tribe Tagiadini Mabille, 1878 (Afrotropical Tribe Anthoptini Warren, 2009 genera: Eagris Guénée, 1862; Procampta Tribe Moncini Warren, 2008 Holland, 1892; Calleagris Aurivillius, [1925]; Tribe Hesperiini Latreille, 1809 Kobelana Larsen & Collins, 2013; Tagiades Hübner, [1819]; Netrobalane Mabille, [1903]; FAMILY NYMPHALIDAE Rafinesque, 1815 Caprona Wallengren, 1857; Leucochitonea Wallengren, 1857; Abantis Hopffer, 1855) This family is cosmopolitan, with about 6, 160 species Tribe Celaenorrhinini Swinhoe, 1912 in 560 genera. There are 1, 580 species in 86 genera in (Afrotropical genera: Ortholexis Karsch, 1895; the Afrotropical Region. A classification of the family, Katreus Watson, 1893; Celaenorrhinus Hübner, shown below, was derived after consulting the [1819]; Scopulifera Libert, 2014; Bettonula Libert following publications: Brower et al., 2009 (Danaini); Willliams / Metamorphosis 26: 102–108 104 Henning & Williams, 2010 (Acraeini); Lang, 2010 Subtribe Ragadiina Herrich-Schaeffer, (Chalangini);
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  • Intraspecific Body Size Frequency Distributions of Insects

    Intraspecific Body Size Frequency Distributions of Insects

    Intraspecific Body Size Frequency Distributions of Insects E. Jeanne Gouws1¤, Kevin J. Gaston2, Steven L. Chown1* 1 Department of Botany and Zoology, Centre for Invasion Biology, Stellenbosch University, Stellenbosch, South Africa, 2 Biodiversity and Macroecology Group, Department of Animal and Plant Sciences, University of Sheffield, Sheffield, United Kingdom Abstract Although interspecific body size frequency distributions are well documented for many taxa, including the insects, intraspecific body size frequency distributions (IaBSFDs) are more poorly known, and their variation among mass-based and linear estimates of size has not been widely explored. Here we provide IaBSFDs for 16 species of insects based on both mass and linear estimates and large sample sizes (n$100). In addition, we review the published IaBSFDs for insects, though doing so is complicated by their under-emphasis in the literature. The form of IaBSFDs can differ substantially between mass- based and linear measures. Nonetheless, in non-social insects they tend to be normally distributed (18 of 27 species) or in fewer instances positively skewed. Negatively skewed distributions are infrequently reported and log transformation readily removes the positive skew. Sexual size dimorphism does not generally cause bimodality in IaBSFDs. The available information on IaBSFDs in the social insects suggests that these distributions are usually positively skewed or bimodal (24 of 30 species). However, only c. 15% of ant genera are polymorphic, suggesting that normal distributions are probably more common, but less frequently investigated. Although only 57 species, representing seven of the 29 orders of insects, have been considered here, it appears that whilst IaBSFDs are usually normal, other distribution shapes can be found in several species, though most notably among the social insects.
  • Biology of the Queen of Spain Fritillary, Issoria Lathonia (Lepidoptera: Nymphalidae)

    Biology of the Queen of Spain Fritillary, Issoria Lathonia (Lepidoptera: Nymphalidae)

    Zobar & Genc: Biology of Queen of Spain Fritillary 237 BIOLOGY OF THE QUEEN OF SPAIN FRITILLARY, ISSORIA LATHONIA (LEPIDOPTERA: NYMPHALIDAE) DAMLA ZOBAR1 AND HANIFE GENC1 1Department of Plant Protection, Agricultural Faculty, Canakkale Onsekiz Mart University, 17100 Canakkale, Turkey ABSTRACT The biology and the life cycle of Issoria lathonia (Nymphalidae) (Linnaeus 1758) on its host plant, Viola tricolor L. (Violaceae), are described from laboratory studies. In the laboratory eggs are laid singly on the host plant leaves as well as on the surfaces of plastic screen cages. Newly hatched larvae aggregate and feed on the host plant leaves. Later instars disperse on the plant and continue to feed on leaves and flowers. Head capsule widths, and weight and size measurements show that larvae develop through 5 instars. The larvae crawl off the host plant and pupate off the host. The life cycle from egg to adult requires 23-31 d at 26°C, and 16:8 (L:D) photoperiod in the laboratory. The butterfly has been reared continuously in the laboratory for about 2 years. Key Words: Issoria lathonia, Nymphalidae, Argynnini, Viola tricolor RESUMEN Se describe la biología y el ciclo de vida de Issoria lathonia (Linnaeus, 1758) sobre su planta hospedera, Viola tricolor L. (Violaceae) basado sobre estudios de laboratorio. En el laborato- rio los huevos están puestos individualmente sobre las hojas de la planta hospedera igual como sobre la superficie de la tela plástica de las jaulas. Las larvas recién nacidas se agregan y se alimentan sobre las hojas de la planta hospedero. Los instares posteriores se dispersan sobre la planta y continúan su alimentación sobre las hojas y flores.