(Lepidoptera). Part 5

Zootaxa 3724 (1): 001–085 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Monograph ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3724.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:7D05BB2E-4373-4AFB-8DD3-ABE203D3BEC1 ZOOTAXA

3724

Observations on the Biology of Afrotropical Hesperiidae (Lepidoptera). Part 5. Hesperiinae incertae sedis: Dicotyledon Feeders

MATTHEW J. W. COCK1 & T. COLIN E. CONGDON2 1 C/o CABI Europe – UK, Bakeham Lane, Egham, TW20 9TY, UK (e–mail: [email protected]; [email protected]) 2 African Butterfly Research Institute (ABRI), P.O. Box 14308, Nairobi, Kenya (e-mail: [email protected])

Magnolia Press Auckland, New Zealand

Accepted by C. Prieto: 29 Aug. 2013; published: 25 Oct. 2013 MATTHEW J. W. COCK & T. COLIN E. CONGDON Observations on the Biology of Afrotropical Hesperiidae (Lepidoptera). Part 5. Hesperiinae incertae sedis: Dicotyledon Feeders (Zootaxa 3724) 85 pp.; 30 cm. 25 Oct. 2013 ISBN 978-1-77557-278-7 (paperback) ISBN 978-1-77557-279-4 (Online edition)

FIRST PUBLISHED IN 2013 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

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ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition)

Abstract ...... 4 INTRODUCTION ...... 4 HESPERIINAE Latreille, 1809...... 5 From Ampittia group ...... 5 Gorgyra Holland, 1896 ...... 5 Gorgyra bibulus Riley, 1929 (in Eltringham et al. 1929) ...... 7 Gorgyra johnstoni Butler, 1894...... 12 Gorgyra mocquerysii Holland, 1896...... 15 Gorgyra subflavidus Holland, 1896 ...... 17 Gorgyra discussion ...... 18 From Ceratrichia group...... 18 Teniorhinus Holland, 1892 ...... 18 Acada Evans, 1937 ...... 19 Acada biseriata Mabille, 1893 ...... 19 Acada discussion...... 20 Parosmodes Holland, 1896 ...... 20 Parosmodes morantii morantii Trimen, 1873 ...... 20 Parosmodes discussion ...... 25 From Acleros group ...... 25 Acleros Mabille, [1885] (in Mabille, 1885–[1887]) ...... 26 Acleros mackenii instabilis Mabille, 1890 ...... 26 Acleros ploetzi Mabille, 1890 ...... 30 Acleros discussion ...... 31 Meza Hemming, 1939 ...... 31 Meza larea Neave, 1910 ...... 32 Paronymus Aurivillius, 1925...... 32 Paronymus ligora Hewitson, 1876 ...... 32 Andronymus Holland, 1896...... 33 Andronymus caesar Fabricius, 1793 ...... 34 Andronymus caesar caesar Fabricius, 1793 ...... 36 Andronymus caesar philander Hopffer, 1855 ...... 42 Andronymus hero Evans, 1937 ...... 42 Andronymus helles Evans, 1937 ...... 45 Andronymus marina Evans, 1937 ...... 49 Andronymus neander neander Plötz, 1884 ...... 49 Andronymus discussion ...... 49 From Gegenes group ...... 49 Melphinyet Larsen, 2012...... 49 Melphinyet flavina Evans, 1937 ...... 49 Melphinyet tarace Mabille, 1891 ...... 53 Melphinyet discussion ...... 55 Fresna Evans, 1937...... 55 Fresna nyassae Hewitson, 1878 ...... 56 Fresna discussion ...... 57 Platylesches Holland, 1896...... 58 Platylesches ayresii Trimen, 1889...... 59 Platylesches galesa Hewitson, 1877 ...... 60 Platylesches langa Evans, 1937 ...... 61 Platylesches sp. moritili group ...... 63 Platylesches moritili Wallengren, 1857...... 63 Platylesches neba Hewitson, 1877 ...... 70 Platylesches picanini Holland, 1894...... 71 Platylesches robustus robustus Neave, 1910 ...... 72 Platylesches shona Evans, 1937 ...... 73 Platylesches tina Evans, 1937...... 74 Platylesches discussion ...... 77 DISCUSSION ...... 78 ACKNOWLEDGEMENTS...... 80 REFERENCES ...... 80

Partial life histories of 24 Hesperiinae incertae sedis that feed on Dicotyledons are described and illustrated. The genera dealt with are: Acada, Acleros, Andronymus, Fresna, Gorgyra, Melphinyet, Meza, Paronymus, Parosmodes, Platylesches, and Teniorhinus. The food plants of Gorgyra (mainly Connaraceae), Melphinyet (Euphorbiaceae) and Platylesches (Chrysobalanaceae) stand out as clearly distinct from those of the other six genera, which share one or two food plant fam- ilies with each other. Thus four genera include Fabaceae in their food plants, two include Sapindaceae, two include Com- bretaceae, and Andronymus includes all three. Ova are only reported for five genera, but they do point to some significant differences: Gorgyra, Parosmodes and Platylesches are different from each other, and different from Acleros and Andro- nymus. The features of the caterpillars do not indicate clear patterns. The pupal shelters point to Parosmodes and Mel- phinyet showing some affinity and certainly differ from all others. Acleros and Andronymus have similar pupal shelters, and in common with Meza (dicotyledon-feeding species) and Fresna, the pupal chamber is not closed and the pupa rests on the leaf upper surface. The pupae of Gorgyra differ from all others. The pupae of Acleros and Andronymus have their colouring in common, probably reflecting their very similar pupal shelters. Similarly the pupae and shelters of Meza (dicotyledon-feeding species) and Fresna seem analogous. The other pupae, hidden in their shelters are less differentiated, although the completely wax-free shelter of Acada is superficially different. We conclude that the likeliest scenario is that in the Afrotropical Region members of the Hesperiinae incertae sedis have made the switch from monocotyledons to dicotyledons several times, and then radiated.

Key words: Acada, Acleros, Andronymus, Fresna, Gorgyra, Melphinyet, Meza, Paronymus, Parosmodes, Platylesches, Teniorhinus, food plant, life history, leaf shelter, parasitoid

INTRODUCTION

This is the fifth in a series of papers on the biology of Afrotropical Hesperiidae, based on the first author’s observations primarily in Kenya, and the second author’s observations primarily in Tanzania. Earlier parts covered Coeliadinae (Cock 2010), Pyrginae: Tagiadini (Cock & Congdon 2011a) and Celaenorrhinini (Cock & Congdon 2011b), and Hesperiinae: Aeromachini and Baorini (Cock & Congdon 2012). The remaining genera of Hesperiinae are currently incertae sedis and not placed in tribes (Warren et al. 2009). It seems likely that characters of the early stages will be useful in clarifying the classification of many Hesperiinae genera incertae sedis, and we hope our observations will provide relevant and useful information for the revision of African Hesperiidae currently being undertaken by T.B. Larsen (see e.g. Larsen & Congdon 2012). A detailed introduction is provided in Cock (2010) and supplemented in Cock & Congdon (2011a), so no further introduction is provided here. In this paper, unacknowledged information is the work of the first author (MJWC) who undertook the bulk of the writing, while the second author (TCEC) provided additional material which is attributed, in particular the important documentation of Platylesches spp. life histories. Although, the title of the series refers to observations principally from Kenya, we have dropped this for this part, as only three of the many species reared were from Kenya. We have included photographs of living adult skippers when available, but in many cases there are good photographs available on the internet (e.g. SABCA 2012 for South African species); we have not referred to these, unless directly relevant to the points we make, and more particularly if pictures of the early stages are available. Material referred to as TCEC in this paper reflects the outputs of the Butterfly/Host Plant Research Programme which has run in Tanzania and neighbouring countries for over 20 years under the auspices of the Tanzania Commission for Science and Technology (COSTECH). The key researchers have been T.C.E. Congdon, I. Bampton (deceased May 2010) and S.C. Collins. TCEC did all of his rearing in collaboration with the late Ivan Bampton, and it is not possible to separate the efforts of these co-workers. However, TCEC did all the photography and writing up of what they reared together. Hence, reference to TCEC collections in this paper should automatically be considered to include Bampton. Both Congdon and Bampton have been Research Associates of the African Butterfly Research Institute (ABRI) since its inception in 1996, and all material collected and reared under the COSTECH programme has been incorporated in the ABRI collection in Nairobi.