Volume 24: 57–68 METAMORPHOSIS ISSN 1018–6490 (PRINT) LEPIDOPTERISTS’ SOCIETY OF ISSN 2307–5031 (ONLINE)

Publications on Afrotropical during 2013 Published online: 31 December 2013

Mark C. Williams

University of Pretoria, Pretoria, South Africa. E-mail : [email protected]

Copyright © Lepidopterists’ Society of Africa

Abstract: The articles published during 2013 which deal with scientific research into Afrotropical Lepidoptera are listed alphabetically by author and abstracts are provided if they were published. Articles dealing with control of Lepidoptera as pests are excluded.

Citation: Williams, M.C. 2013. Publications on Afrotropical Lepidoptera during 2013. Metamorphosis 24: 57–68.

AFROTROPICAL LEPIDOPTERA in the western Congo Basin and southern Tanzania/Mozambique. Main conclusions. Sampling effort is highly biased and controlling for it in large-scale BALLESTEROS-MEJIA, L., KITCHING, I.J., compilations of presence-only data is critical for drawing JETZ, W., NAGEL, P. & BECK J. 2013. inferences from our still limited knowledge of Mapping the biodiversity of tropical : species invertebrate distributions. Our study shows that a baseline richness and inventory completeness of African of estimate of broad-scale diversity patterns in sphingid . Global Ecology and Biogeography understudied taxa can be derived from combining 22(5): 586–595. numerical estimators of richness, models of main Abstract: Aim. Many taxa, especially invertebrates, environmental effects and spatial interpolation. Inventory remain biogeographically highly understudied and even completeness can be partly predicted from human baseline assessments are missing, with too limited and geographic features and such models may offer fruitful heterogeneous sampling being key reasons. Here we set guidance for prioritization of future sampling to further out to assess the human geographic and associated refine and validate estimated patterns of species richness. environmental factors behind inventory completeness for the hawkmoths of Africa. We aim to separate the causes BARTSCH, D. 2013. Revisionary checklist of the of differential sampling from those affecting gradients of Southern African Sesiini (Lepidoptera: Sesiidae) species richness to illustrate a potential general avenue with description of new species. Zootaxa 3741(1): for advancing knowledge about diversity in understudied 1–54. groups. Location. Sub-Saharan Africa. Methods. Using a Abstract: A revisionary checklist of the Sesiini of database of distributional records of hawkmoths, we southern Africa is presented. All known genera of the computed rarefaction curves and estimated total species region and their type species are redescribed. The richness across 200km x 200km grid cells. We fitted following genera are placed in Sesiini for the first time: multivariate models to identify environmental predictors Alonina Walker, 1856; Anaudia Wallengren, 1863; of species richness and used environmental co-kriging to Austrosetia Felder & Felder, 1874; Megalosphecia Le map region-wide diversity patterns. We estimated cell- Cerf, 1916 and Vespanthedon Le Cerf, 1917. The genera wide inventory completeness from observed and Ci-cinnoscelis Holland, 1893 rev. stat. (from synonymy estimated data, and related these to human geographic with Alonina Walker, 1856) and Felderiola Naumann, factors. Results. Observed patterns of hawkmoths species 1971 rev. stat. (from synonymy with Monopetalotaxis richness are strongly determined by the number of Wallengren, 1859) are resurrected from synonymy and available records in grid cells. Both show spatially included in Sesiini. One new , Hagnogyna new structured distributions. Variables describing vegetation gen., and seven new species, Cicinnoscelis flavipes new type, emerge as important predictors of estimated total sp., C. krooni new sp., Alonina rufa new sp., A. fusca richness, and variables capturing heat, energy availability new sp., Anaudia thyranthrena new sp., Felderiola and topographic heterogeneity all show a strong positive karooensis new sp. and F. xanthogaster new sp., are relationship. Patterns of interpolated richness identify described. The following new combinations are three centres of diversity: Cameroon coastal mountains, introduced: Cicinnoscelis longipes Holland, 1893 rev. and the northern and southern East African montane comb., Alonina pyrethra (Hampson, 1919) new comb., areas. Inventory completeness is positively influenced by A. pyrocraspis (Hampson, 1910) new comb., population density, accessibility, protected areas and A. luteopunctata (Freina, 2011) new comb., Felderiola colonial history. Species richness is still under-recorded candescens (Felder & Felder, 1874) rev. comb., Received: 1 December 2013 Hagnogyna sanguicosta (Hampson, 1919) new comb., Accepted: 31 December 2013 H. bartschi (Freina, 2011) new comb. and Vespanthedon chalciphora (Hampson, 1919) new comb. Alonina Copyright: This work is licensed under the Creative difformis Hampson, 1919 new syn. is considered a Commons Attribution-NonCommercial-NoDerivs 3.0 subjective junior synonym of A. rygchiiformis Walker, Unported License. To view a copy of this license, send a 1856. A lecto-type is designated for Austrosetia semirufa letter to Creative Commons, Second Street, Suite 300, Felder & Felder, 1874 and the previously unknown males San Francisco, California, 94105, USA, or visit: of this species and of Anaudia felderi Wallengren, 1863 http://creative commons.org/licenses/by-nc-nd/3.0/

Williams/ Metamorphosis 24: 57–68 58 are described. Bionomical data for some of the species Genetic diversity was dramatically different among discussed are given for the first time. species and not easily explained by either species-specific traits or effects of fragmentation. galene, the BASQUIN, P. 2013. Contribution à la species most impacted by fragmentation, exhibited very connaissance du complexe de Maltagorea rostaingi high diversity, whereas G. betsimena, a broadly (Griveaud, 1961) à Madagascar (Lepidoptera, distributed, very common species, with relaxed habitat Saturniidae). Saturnafrica 15: 19–25, 27–33. fidelity, was genetically depauperate. There was limited evidence of genetic erosion from the sacred groves despite these small forest patches accounting for less than BASQUIN, P. 2013. Nouveaux Eupterotidae 1-10 % of the total area of the forest reserves, which africains (Lepidoptera). Entomologia Africana indicates these forest relics have high conservation value. 18(1): 23–26. Abstract: Two new Eupterotidae species are described: BOUYER, T. 2013. Description d’une nouvelle Stenoglene bouyeri n. sp. from Gabon and Hoplojana espèce d’Iridana Aurivillius, 1920 du Congo (R. politzari n. sp. from Somalia. D.) (Lepidoptera, Lycaenidae). Entomologia Africana 18(1): 4–6. [Iridana ducarmei Bouyer, BASSI, G. 2013. Notes on some Old World 2013] Prionapterygini Landry, 1995 (Lepidoptera: Pyraloidea, , ), with BOUYER, T. & DUCARME, R. 2013. descriptions of new species. Revue Suisse de Description de la femelle de Cymothoe bouyeri Van Zoologie 120(1): 131–160. de Weghe, 2011 (Lepidoptera, ). Abstract: Notes on some Old World Prionapterygini Entomologia Africana 18(1): 27–28. Landry, 1995 (Lepidoptera: Pyraloidea, Crambidae, Crambinae), with descriptions of new species. – Zovax COACHE, A., RAINON, B. & ZANNOU, E. venus Bassi sp. n. along with two new species of Mesolia 2013. Enfin un Cymothoe Hübner, 1919 en Ragonot, 1889 (M. meyi Bassi sp. n. and M. alborzella République du Bénin (Lepidoptera, Nymphalidae). Bassi sp. n.) and four new species of Stephens, 1834 (P. banaadirensis Bassi sp. n., P. eberti L’Entomologiste 69(3): 133–135. [Cymothoe druryi Bassi & Mey sp. n., P. helena Bassi sp. n. and P. somala van Velzen & Larsen, 2009 recorded from Bénin] Bassi sp. n.) are described. A lectotype is designated for Z. whiteheadii (Wollaston), 1879. Loxophantis Meyrick, COACHE, A., RAINON, B. & ZANNOU, E. 1935 is newly considered as a synonym of Prionapteryx, 2013. Découverte du mâle de Tumerepedes flava which causes the following changes: Prionapteryx Bethune-Baker, 1913, en République du Bénin triplecta (Meyrick, 1935), comb. n., and Prionapteryx (Lepidoptera Lycaenidae). L’Entomologiste 69(1): albimaculalis (Hampson), 1919 = Loxophantis 57–58. pretoriella Bleszynski, 1970 syn. n. P. triplecta is compared to P. diaplecta (Meyrick), P. albimaculalis is COCK, M.J.W. & CONGDON, T.C.E. 2013. compared to P. plumbealis (Hampson) and M. meyi is Observations on the biology of Afrotropical compared to M. uniformella Janse 1922. All species are Hesperiidae (Lepidoptera). Part 5. Hesperiinae fully illustrated. The habitus of P. amathusia Bassi & incertae sedis: Dicotyledon feeders. Zootaxa Mey, 2011 is illustrated and corrections to Bassi & Mey (2011) are given. A list of species presently placed in 3724(1): 1–85. Prionapteryx and which should be transferred to Walker, 1863 is given. COCKBURN, K.N.A. 2013. Observations on the larval host plant and life history of the Dwarf Blue, BOSSART, J.L. & ANTWI, J.B. 2013. Species- Oraidium barberae Trimen, 1868 (Lycaenidae: specific traits predict genetic structure but not Polyommatinae). Metamorphosis 24: 12–14. genetic diversity of three fragmented Afrotropical forest species. Conservation Genetics COETZER, A.J. & GARDINER, A.J. 2013. The 14(2): 511–528. behaviour and life history of Anthene crawshayi Abstract: The Upper Guinean forests of Ghana, West juanitae Henning & Henning, 1993 (Lepidoptera: Africa, are considered among the most threatened and Lycaenidae). Metamorphosis 24: 35–37. fragmented in the world. Little is known about the genetic consequences of fragmentation on Ghana’s COLLINS, S.C., CONGDON, T.C.E., forest-associated species, but this genetic signature is HENNING, G.A., LARSEN, T.B. & generally expected to differ across species. We compared WILLIAMS, M.C. 2013. A review of d’Abrera’s patterns of mtDNA cytochrome oxidase I (COI) variation of the Afrotropical Region – Part III of three Nymphalid forest butterfly species that differ (second edition), 2009 – Part 1. Metamorphosis 24: with respect to their relative dispersibilities (Aterica galene: high habitat fidelity, low dispersal ability; 25–34. Euphaedra medon: high habitat fidelity, strong dispersal Abstract: Part III of the Butterflies of the Afrotropical ability; betsimena: relaxed habitat fidelity, Region, dealing with the Lycaenidae and Riodinidae, was low dispersal ability). Individuals were collected from published by d’Abrera in 2009. The text has been two large forest reserves and five small sacred forest reviewed in detail and errors, omissions and groves. Patterns of differentiation across species were misattributions are listed in this Part 1. The taxonomic broadly coincident with our predicted hierarchy of changes that have been made in these butterfly families relative species dispersibility and suggested that genetic by other authors since d’Abrera (2009) are also listed and connectivity is most compromised by strict fidelity to annotated if necessary. forest habitat rather than by raw capacity for sustained flight. Connectivity was uncorrelated with geographic COLLINS, S.C., CONGDON, T.C.E., distance, but instead seemed best explained by HENNING, G.A., LARSEN, T.B. & urbanization and the sequential pattern of forest loss. WILLIAMS, M.C. 2013. A review of d’Abrera’s Williams/ Metamorphosis 24: 57–68 59

Butterflies of the Afrotropical Region – Part III with known functions in metabolism, pigment (second edition), 2009 – Part 2 (Miletinae and production, development and heat shock responses. These Poritiinae). Metamorphosis 24: 44–56. include genes for which we expect a priori a role in Abstract: Part III of the Butterflies of the Afrotropical thermal adaptation and, thus, varying selection pressures Region, dealing with the Lycaenidae and Riodinidae, was along a latitudinal cline, and genes we do not expect to published by d’Abrera in 2009. The text of this work has vary clinally and can be used as controls. We identified been reviewed in detail and in this part of the review the amino acid substitution polymorphisms in 13 genes and taxonomic changes implemented or suggested by the tested these for clinal variation by correlation analysis of author in the subfamilies Miletinae and Poritiinae are allele frequencies with latitude. In addition, we used two assessed, and whilst many are accepted, others are FST-based outlier methods to identify loci with higher reversed or revised. population differentiation than expected under neutral evolution, while accounting for potentially confounding COLLINS, S.C. & LARSEN, T.B. 2013. A new effects of population structure and demographic history. species in the Afrotropical skipper genus Artitropa Two metabolic enzymes of the glycolytic pathway, UGP and Treh, showed clinal variation. The same loci showed from São Tomé and Principe (Lepidoptera: elevated population differentiation and were identified as Hesperiidae: Hesperiinae (incertae sedis)). significant outliers. We found no evidence of clines in the Metamorphosis 24: 20–24. pigmentation genes, heat shock proteins and Abstract: A distinct new species in the genus Artitropa developmental genes. However, we identified outlier loci (Lepidoptera: Hesperiidae: Hesperiinae (incertae sedis)) in more localized parts of the range in the pigmentation inhabiting the islands of Principe and São Tomé is genes yellow and black. We discuss that the observed described as Artitropa principetome. It was first collected clinal variation and elevated population divergence in in 2008 and almost certainly has been overlooked before UGP and Treh may reflect adaptation to a geographic that date. Colour patterns and the male genitalia indicate thermal gradient. that it is most closely related to Artitropa comus, the only member of the genus to be found in West Africa. The DUBATOLOV, V.V. 2013. A new genus of food plant was found to be Dracaena arborea. Most African tiger moths, with a review of the Amsacta endemic species on these islands are found on only one of melanogastra Holland species group (Lepidoptera, these islands. Since members of the genus do not ). Zootaxa 3682(4): 579–583. generally disperse it is likely that it evolved from A. comus over an extended time period on one island, EDGE, D.A. 2013. Book review: Lepidoptera and reaching the other recently as there are no morphological differences between their populations. Conservation – Tim R. New. Metamorphosis 24: 75. CONGDON, T.C.E. & BAYLISS, J.L. 2013. Butterflies of Mt Mecula and Mt Yao, Niassa EDGE, D.A. 2013. Obituary – Johan (Scottie) Province, Northern Mozambique. Metamorphosis Buys (15.06.1949 – 21.12.2013). Metamorphosis 23: 26–34. 24: v–vi. Abstract: A multidisciplinary expedition visited Mt Mecula and Mt Yao in northern Mozambique from 6th to EL-HAWAGRY, M.S., KHALIL, M.W., 16th May, 2012. Butterflies found on the two mountains SHARAF, M.R., FADL, H.H. & ALDAWOOD, are scheduled, and those of scientific interest are A.S. 2013. A preliminary study on the fauna discussed. A preliminary report on the flora of Mt Yao is of Al-Baha Province, Saudi Arabia, with provided. Biogeographical inferences are drawn. descriptions of two new species. ZooKeys 274: 1– 88. D’ABRERA B. Undated. Saturniidae Mundi: Abstract: A preliminary study was carried out on the Saturniid Moths of the World. Part II. Hill House insect fauna of Al-Baha Province, south-western part of Publishers, Malvern. Saudi Arabia. A total number of 582 species and subspecies (few identified only to the genus level) DARGE, P. 2013. Nouvelles données sur la faune belonging to 129 families and representing 17 orders entomologique de Zambie, avec description de were recorded. Two of these species are described as plusieurs espèces des genres Adafroptilum et new, namely: Monomorium sarawatensis Sharaf & Aurivillius (Lepidoptera, Saturniidae). Saturnafrica Aldawood, sp. n. (Formicidae, Hymenoptera) and 15: 3–17. Anthrax alruqibi El-Hawagry sp. n. (Bombyliidae, Diptera). Another eight species are recorded for the first DARGE, P. 2013. Nouvelles espèces de time in Saudi Arabia, namely: Xiphoceriana arabica (Uvarov, 1922) (Pamphagidae, Orthoptera), Rhopopteriana Bouyer, 2012. (Lepidoptera, Pyrgomorpha conica (Olivier, 1791) (Pyrgomorphidae, Eupterotidae). Saturnafrica 15: 35–42, 43–47. Orthoptera), Catopsilia florella (Fabricius, 1775) (Pieridae, Lepidoptera), Anthrax chionanthrax (Bezzi, DE JONG, M.A., COLLINS, S., BELDADE, P., 1926) (Bombyliidae, Diptera), Spogostylum near BRAKEFIELD, P.M. & ZWAAN, B.J. 2013. tripunctatum Pallas in Wiedemann, 1818 (Bombyliidae, Footprints of selection in wild populations of Diptera), Cononedys dichromatopa (Bezzi, 1925) Bicyclus anynana along a latitudinal cline. (Bombyliidae, Diptera), Mydas sp. (Mydidae, Diptera), Molecular Ecology 22(2): 341–353. and Hippobosca equina Linnaeus, 1758 (Hippoboscidae, Abstract: One of the major questions in ecology and Diptera). Al-Baha Province is divided by huge and steep evolutionary biology is how variation in the genome Rocky Mountains into two main sectors, a lowland enables species to adapt to divergent environments. Here, coastal plain at the west, known as “Tihama”, and a we study footprints of thermal selection in candidate mountainous area with an elevation of 1500 to 2450 m genes in six wild populations of the afrotropical butterfly above sea level at the east, known as “Al-Sarat or Al- Bicyclus anynana sampled along a c. 3000 km latitudinal Sarah” which form a part of Al-Sarawat Mountains cline. We sequenced coding regions of 31 selected genes range. Insect species richness in the two sectors (Tihama Williams/ Metamorphosis 24: 57–68 60 and Al-Sarah) was compared, and the results showed that bacterium Spiroplasma. Hypothesis: The different colour each of the two sectors of Al-Baha Province has a unique forms of D. chrysippus represent incipient species that insect community. The study generally concluded that the have undergone hybridization as a result of Spiroplasma insect faunal composition in Al-Baha Province has an invasion, because many females of the frequently Afrotropical flavor, with the Afrotropical elements infected forms are forced to mate with the males of the predominant, and a closer affiliation to the Afrotropical less infected forms. Prediction: Some forms are more region than to the Palearctic region or the Eremic zone. susceptible to Spiroplasma infection than others. Times Consequently, we tend to agree with those and places: and Sudan during 2005, 2006, and biogeographers who consider that parts of the Arabian 2007. Analytical method: We collected D. chrysippus Peninsula, including Al-Baha Province, should be butterflies in the wild. We recorded their colour pattern included in the Afrotropical region rather than in the and sex. We then used polymerase chain reaction (PCR) Palaearctic region or the Eremic zone. to determine whether they were infected with Spiroplasma. We estimated the morph ratio, the sex ratio, FRANKE, K. & FISCHER, K. 2013. Effects of and the prevalence of Spiroplasma in different inbreeding and temperature stress on life history populations and regions. The association between colour and immune function in a butterfly. Journal of pattern and Spiroplasma infection was subjected to Evolutionary Biology 26(3): 517–528. statistical analysis. Conclusion: We found no significant Abstract: Theory predicts that inbreeding depression difference in the sex ratio or the prevalence of should be more pronounced under environmental stress Spiroplasma between different forms. Colour forms do due to an increase in the expression of recessive not vary in their susceptibility to Spiroplasma infection. deleterious alleles. If so, inbred populations may be especially vulnerable to environmental change. Against HASSAN, S.S.M., IDRIS, E. & MAJERUS, this background, we here investigate effects of M.E.N. 2013. Male-Killer Dynamics in the inbreeding, temperature stress and its interactions with Tropical Butterfly Acraea encedon (Lepidoptera: inbreeding in the tropical butterfly Bicyclus anynana. Nymphalidae). African Entomology 21(2): 209-214. We use a full-factorial design with three levels of Abstract: Sex ratio distortion in the tropical butterfly inbreeding (F=0/0.25/0.38) and three temperature Acraea encedon is caused by infection with a male- treatments (2h exposure to 1, 27 or 39°C). Despite using killing bacterium of the genus Wolbachia. Severely relatively low levels of inbreeding significant inbreeding female-biased populations with extremely high bacterial depression was found in pupal mass, pupal time, thorax prevalences were previously reported in A. encedon. In mass, abdomen fat content, egg hatching success and this paper, we describe a study in which the spatial fecundity. However, stress resistance traits (heat variations and the temporal changes in the population sex tolerance, immune function) were not affected by ratio and the prevalence of the male-killing Wolbachia inbreeding and interactions with temperature treatments were thoroughly investigated in the wild populations of were virtually absent. We thus found no support for an Acraea encedon in Uganda between 2005 and 2007. The increased sensitivity of inbred individuals to two parameters were found to vary extensively over environmental stress, and suspect that such patterns are space and time, indicating that the male-killer is highly restricted to harsher conditions. Our temperature dynamic. Both the female-bias and the Wolbachia treatments evidently imposed stress, significantly prevalence were lower than that recorded in the literature. reducing longevity, fecundity, egg hatching success and haemocyte numbers, while fat content, protein content HEATH, A. & KALISZEWSKA, Z.A. 2013. and lysozyme activity remained unaffected. Males and Further life history observations in the females differed in all traits measured except pupal time, myrmecophilous genus Chrysoritis Butler, plus protein content and phenoloxidase (PO) activity. notes on dwarfism in reared specimens. Correlation analyses revealed, among others, a trade-off Metamorphosis 23: 16–21. between PO and lysozyme activity, and negative Abstract: Associated host-ant species and larval host- correlations between fat content and several other traits. plants are recorded for Chrysoritis pelion (Pennington, We stress that more data are needed on the effects of 1953), Chrysoritis irene (Pennington, 1968) and inbreeding, temperature variation and sexual differences Chrysoritis natalensis (Van Son, 1966). Instances of on insect immune function before more general undersize adults obtained when reared in captivity from conclusions can be drawn. eggs, without ant attendance, are recorded and discussed.

GARDINER, A.J. 2013. Description of a new HENNING, G.A. & BALL, J.B. 2013. A new species of Erikssonia Trimen, 1891 (Lepidoptera: species of Lepidochrysops Hedicke (Lepidoptera: Lycaenidae) from the Democratic Republic of Lycaenidae) from the Northern Cape, South Africa. Congo. Metamorphosis 23: 22–25. Metamorphosis 23: 1–7. Abstract: A fourth species of Erikssonia Trimen, 1891 Abstract: The genus Lepidochrysops Hedicke from the Democratic Republic of Congo, E. bouyeri sp. (Lycaenidae) is briefly discussed and a new species in the nov., is added to the three species previously recognised L. methymna group, Lepidochrysops frederikiae sp. nov. in the genus: E. acraeina Trimen, 1891, E. cooksoni from the Northern Cape, is described and figured. The Druce, 1905, and E. edgei Gardiner & Terblanche, 2010. habitat, a small inselberg (at an altitude of over 1200m) A key to the species is provided. of the Fynbos vegetation unit Roggeveld Shale Renosterveld surrounded by succulent Karoo vegetation HASSAN, S.S.M., IDRIS, E. & MAJERUS, units, is discussed in relation to other apparently new M.E.N. 2013. Is it just a coincidence that and plant taxa found there. aposematic polymorphism and sex ratio distortion co-occur in a tropical butterfly? Evolutionary INTERNATIONAL COMMISSION ON Ecology Research 15(1): 93–102. ZOOLOGICAL NOMENCLATURE 2013. Abstract: Background: The cosmopolitan butterfly Opinion 2319. Case 3503). Papilio hesperus Danaus chrysippus is polymorphic only within the Westwood, 1843 (Insecta, Lepidoptera, geographic zone where it is infected by the male-killing PAPILIONIDAE): conserved by suppression of Williams/ Metamorphosis 24: 57–68 61

Papilio hesperus Fabricius, 1793 of male behavior in determining mating success may (NYMPHALIDAE). Bulletin of Zoological result in sexual conflict. Nomenclature 70(2): 127–128. Abstract: The Commission has conserved the name KASANGAKI, P., AKOL, A.M. & BASUTA, Papilio hesperus Westwood, 1843 by suppression of G.I. 2013 (2012). Butterfly species list for selected Papilio hesperus Fabricius, 1793. Coincidentally this also west Albertine rift forests. Dataset Papers in conserves the name Harma chalcis C. & R. Felder, 1860, Biology 2013 1–4 Article ID 451461. which is in widespread use in much of Africa in the Abstract: This dataset gives details on a study which combination Euryphura chalcis. was conducted in seventeen forests in western Uganda (Albertine region) between 1993 and 1995. Different KARL, I. & FISCHER, K. 2013. Old male mating methods were used to sample butterflies from these advantage results from sexual conflict in a butterfly. forests. A total of 630 butterfly species were collected in Animal Behaviour 85(1): 143–149. 5 main families. Key characteristics of the forests were Abstract: Accumulating evidence suggests that sexual recorded, and these were found to influence the butterfly conflict is all pervasive, although cooperation is evidently species richness of the forests. needed in sexually reproducing organisms. While male mating success is typically positively associated with KAWAHARA, A.Y. 2013. Systematic revision resource-holding potential, old male mating success and review of the extant and fossil snout butterflies prevails in some organisms despite old males being in (Lepidoptera: Nymphalidae: ). Zootaxa greatly reduced condition. We tested whether this 3631(1): 1, 3–74. counterintuitive pattern is the outcome of (1) sexual Abstract: Extant and fossil genera and species in the conflict or (2) female preference for older males in the Libytheinae (Nymphalidae) are revised and reviewed. tropical butterfly Bicyclus anynana. Our results support The Libytheinae includes two genera: Fabricius the first hypothesis as females that mated with older and Libytheana Michener. Fifteen species and an compared to younger males showed reduced hatching additional 24 subspecies are recognized and 41 names are success of eggs and higher remating propensity, and synonymized. Species recognized are: Libythea tended to show shorter refractory periods. Thus, females (Laicharting), L. collenettei Riley, L. cinyras Trimen apparently try to compensate for reduced fitness arising revised status, L. geoffroyi Godart, L. labdaca Westwood, from mating with older (low-quality) males by being L. laius Trimen, L. lepita Moore, L. myrrha Godart, L. more willing to remate, especially with young and/or narina Godart, Libytheana carinenta (Cramer), L. virgin (high-quality) males. As mating with older males florissanti (Scudder), L. fulvescens (Lathy), L. motya negatively affects female fitness, old males may acquire a (Hübner), L. terena (Godart), and L. vagabunda mating advantage by being more aggressive and (Scudder). New synonymies that are established for persistent during courtship, in line with the residual (Laicharting (1782)) include: Libythea reproductive value hypothesis. Thus, males increase their (Libythea) celtis f. obscura Milliere 1879 syn. nov.; own reproductive success at the expense of their female Libythea celtis f. denudata Dannehl 1925 syn. nov.; partners, such that the ‘cooperation-conflict balance’ Libythea celtis f. separata Dannehl 1925 syn. nov.; between the sexes seems to be shifted towards conflict in Libythea celtis livida Saggara 1926 syn. nov.; Libythea B. anynana. (Libythea) celtis f. albonervulata Verity 1950 syn. nov.; Libythea (Libythea) celtis f. latefulva Verity 1950 syn. KARL, I., HEUSKIN, S. & FISCHER, K. 2013. nov.; Libythea (Libythea) celtis f. obscurenervulata Dissecting the mechanisms underlying old male Verity 1950 syn. nov.; Libythea (Libythea) celtis f. mating advantage in a butterfly. Behavioral pallida Verity 1950 syn. nov.; Libythea (Libythea) celtis Ecology and Sociobiology 67(5): 837–849. f. pygmaea Verity 1950 syn. nov.; Libythea (Libythea) Abstract: Selection is expected to maximize an celtis f. rubescens Verity 1950 syn. nov.; Libythea individual's own genetic reward regardless of the (Libythea) celtis f. subochracea Verity 1950 syn. nov.; potential fitness consequences for its sexual partners, Libythea (Libythea) celtis f. violacea Verity 1950 syn. which may cause sexual conflict. Although performance nov.; and Libythea celtis platooni Korb 2005 syn. nov. in holometabolous insects typically diminishes with age, Synonyms of Libythea geoffroyi geoffroyi Godart 1822 old male mating advantage has been documented in a few include: Libythea antipoda Boisduval, 1859 syn. nov.; species. Whether this pattern arises from female Libythea orientalis Godman and Salvin, 1888 syn. nov.; preference for older males based on, e.g., pheromone Libythea hauxwelli Moore, 1901 syn. nov.; Libythea blends (intersexual selection), or from increased (Libythea) geoffroy var. sumbensis Pagenstecher, 1901 eagerness to mate in older compared to younger males is syn. nov.; Libythea geoffroy deminuta Fruhstorfer, 1909 currently debated. We explore the mechanistic basis of syn. nov.; and Libythea geoffroy maenia Frühstorfer, old male mating advantage, using a series of experiments 1909 syn. nov. Libythea batchiana Wallace 1869 syn. including behavioral as well as manipulative approaches, nov. is a synonym of Libythea geoffroyi ceramensis in the tropical butterfly Bicyclus anynana. Consistent Wallace 1869. Synonyms of Libythea geoffroyi with the residual reproductive value hypothesis, old male philippina Staudinger 1889 include: Libythea geoffroyi mating advantage was associated with a greater eagerness var. celebensis Staudinger 1889 syn. nov.; and Libythea to mate, evidenced by a two times higher flying and geoffroyi bardas Frühstorfer 1914 syn. nov. Libythea courting activity in older than in younger males. In labdaca subintegra Aurivillius 1921 syn. nov. is a contrast, we found only limited support for a contribution synonym of ancoata Grose-Smith of female preference for older males based on pheromone 1891. Synonyms of lepita Moore (1858) composition, although male sex pheromones clearly do include: Libythea celtis celtoides Frühstorfer 1909 syn. play a role in mating success. Our results suggest that nov.; Libythea celtis chinensis Frühstorfer 1909 syn. male behavior may play a primary role in old male nov.; and Libythea celtis sophene Frühstorfer 1914 syn. mating advantage, and that pheromones are likely of nov. Libythea celtis yayeyamana Fujioka, 1975 syn. nov. secondary importance only. Male mating success was is a synonym of Libythea lepita amamiana Shirozu 1956. related to higher overall pheromone titers rather than myrrhina Frühstorfer 1909 syn. nov. variation in a single component. A dominant importance and Libythea myrrha yawa Frühstorfer 1914 syn. nov. are synonyms of Libythea myrrha myrrha Godart 1819. Williams/ Metamorphosis 24: 57–68 62

Libythea myrrha borneensis Frühstorfer 1914 syn. nov. Abstract: A revision of the genus Acrasia Rogenhofer, and Libythea myrrha iwanagai Hayashi 1976 syn. nov. 1875 is presented as part of an effort to revise a large are synonyms of Libythea myrrha hecura Frühstorfer assemblage of ennomine geometrids in the Afrotropical 1914. Libythea myrrha carma Frühstorfer 1914 syn. nov. Region putatively placed in the southern hemisphere tribe is a synonym of Libythea myrrha rama Moore 1872. . All accessible primary types were Libythea myrrha thira Frühstorfer 1914 syn. nov. is a examined. Acrasia is redefined, and a total of 13 species synonym of Libythea myrrha sanguinalis Frühstorfer placed in three informally named species groups are 1898. Subspecies of L. narina Godart 1819 have been described as new: Acrasia dukei spec. nov., A. monticola synonymized with the nominal species as there are no spec. nov., A. robusta spec. nov.; A. accepta spec. nov.; diagnostic characters that can be used to separate A. amoena spec. nov.; A. lechriogramma spec. nov. subspecies. Synonyms of Libythea narina include: (crinita group); A. grandis spec. nov. (grandis group); A. Libythea rohini Marshall 1880 syn. nov. Libythea similis spec. nov.; A nyangica spec. nov.; A. avellanata luzonica Moore 1901 syn. nov. Libythea narina canuleia spec. nov.; A. lenzi spec. nov.; A. sympatrica spec. nov.; Frühstorfer, 1909 syn. nov.; Libythea narina neratia and A. chimalensis spec. nov. (ava group). Three new Frühstorfer 1909 syn. nov.; Libythea narina sangha combinations are proposed: Acrasia drakensbergensis Frühstorfer 1914 syn. nov.; Libythea narina sumbawana (Herbulot, 1992) comb. nov., A. ava (Prout, 1938) comb. Frühstorfer 1914 syn. nov. The status of the North nov., and A. punctillata (Prout, 1938) comb. nov., stat. American taxon has been confused in the literature, and it nov. (all from Mauna Walker, 1865); the latter taxon is is here treated as Libytheana carinenta bachmanii also raised to specific status. A lectotype is designated for (Kirtland 1851) revised status. The fossil Oligodonta Mauna ava Prout, 1938. A monotypic new genus, florissantensis Brown, 1976 (Pieridae) is a new synonym Panhyperochia gen. nov., is erected to accommodate P. of Libytheana florissanti (Scudder 1892). Keys to genera, ingens spec. nov., a taxon close to Acrasia Rogenhofer species, and subspecies are included along with a but distinguished from it by the development of the juxta discussion of the taxonomic history of the subfamily. in the male genitalia. Keys to the genera, as well as to Diagnostic characters are also presented along with brief species groups and species within Acrasia are provided, comments on the distribution and biology of each as are illustrations of adults and genitalia and distribution species. maps to the southern African species. The revision is complemented by a gazetteer. KEHL, T., KARL, I. & FISCHER, K. 2013. Old- male paternity advantage is a function of LARSEN, T.B. 2013. Ceratricula and Flandria – accumulating sperm and last-male precedence in a two new genera of Afrotropical Hesperiidae butterfly. Molecular Ecology 22(16): 4289–4297. (Hesperiinae (incertae sedis)) for species currently Abstract: Old-male mating advantage has been placed in the genus Ceratrichia Butler. Zootaxa convincingly demonstrated in Bicyclus anynana 3666(4): 476–488. butterflies. This intriguing pattern may be explained by Abstract: Two genera of Afrotropical Hesperiidae are two alternative hypotheses: (i) an increased described to receive species currently placed in aggressiveness and persistence of older males during Ceratrichia Butler, where they do not belong. courtship, being caused by the older males’ low residual Ceratricula gen. nov. comprises a single species reproductive value; and (ii) an active preference of spanning most of the main rainforest zone from Sierra females towards older males what reflects a good genes Leone to Uganda and south to in three hypothesis. Against this background, we here investigate subspecies, ssp. congdoni ssp. nov. being new and ssp. postcopulatory sexual selection by double-mating indeterminabilis Strand raised from synonymy. Flandria Bicyclus anynana females to older and younger males, gen. nov. consists of three rare or very rare species that thus allowing for sperm competition and cryptic mate are limited to eastern Cameroun and the western choice, and by genotyping the resulting offspring. Virgin Democratic Republic of Congo, which gives this genus a females were mated with a younger virgin (2-3days old) more restricted range than any other among the African and afterwards an older virgin male (12-13days old) or forest skippers; the three Flandria had previously been vice versa. Older males had a higher paternity success placed in Ceratrichia, Pardaleodes, or Andronymus. than younger ones, but only when being the second (=last) mating partner, while paternity success was equal LARSEN, T.B. & COLLINS, S.C. 2013. among older and younger males when older males were Kobelana: a new genus for two South African the first mating partner. Older males produced larger skippers currently placed in Calleagris Aurivillius spermatophores with much higher numbers of fertile (as C. kobela Aurivillius, 1925 and C. krooni Vári, sperm than younger males. Thus, we found no evidence 1974) (Lepidoptera: Hesperiidae: Pyrginae: for cryptic female mate choice. Rather, the findings reported here seem to result from a combination of last- Tagiadini). Metamorphosis 23: 8–15. male precedence and the number of sperm transferred Abstract: Two species, currently placed as Calleagris upon mating, both increasing paternity success. kobela (Trimen, 1864) and C. krooni Vári, 1974, are removed to the new genus Kobelana (Lepidoptera; KELLY-MAARTENS, K. & STAUDE, H.S. Hesperiidae; Pyrginae; Tagiadini) on the basis of 2013. Obituary – John G. Joannou (22.01.1949- differences in external features and especially in the structurally different male genitalia. Evans (1937) 25.02.2013). Metamorphosis 23: iv–vi. included five species in the genus Calleagris – C. krooni

being described at a later date. The four species KRÜGER, M. 2013. Revision of Acrasia remaining in Calleagris have very different colour Rogenhofer, 1875 and description of patterns but are united by distinctive, homologous Panhyperochia gen. nov., a new genus of putative genitalia. Kobelana is endemic to South Africa and does nacophorine geometrids from South Africa not overlap with any of the four Calleagris, one or more (Lepidoptera: Geometridae: Ennominae). Annals of of which are found throughout tropical Africa except in the Ditsong National Museum of Natural History 3: the driest habitats. 99–152. Williams/ Metamorphosis 24: 57–68 63

LEHMANN, I. Undated. Description of two new flavirubralis (Hampson, 1917), n. comb., Hypsopygia genera and ten new species of Metarbelidae aurocilialis (Hampson, 1891), n. comb., Arippara (Lepidoptera: ) from western, north- vibicalis (Lederer, 1863), n. comb., Euryzonella central and eastern Africa with notes on habitats sichimensis (Hampson, 1917), n. comb., Lantzodes alcardi (Leraut, 2007), n. comb., Lantzodes bernardii and biogeography. Published by the author, (Leraut, 2007), n. comb., Lantzodes semicircularis University of Bonn, Zoological Research Museum (Hampson, 1917), n. comb., Nussodes africalis (Leraut, Alexander Koenig, Adenauerallee 160, 53113 2007), n. comb. The genus Episindris Ragonot, 1891, n. Bonn, Germany. E-mail: stat., is restored, distinct from Sindris Boisduval, 1833. [email protected]. LI, H., ZHEN, H. & MEY, W. 2013. Notes on LEHMANN, I. & RAJAEI SH, H. 2013. Dichomeris Huebner, 1818 from Southern Africa Description of a new genus and three new species and , with description of seven new species of Metarbelidae (Lepidoptera: Cossoidea) from (Lepidoptera, Gelechiidae, Dichomeridinae). East and Central Africa, with notes on Zootaxa 3608(7): 561–574. biogeography. Bonn Zoological Bulletin 62(1): Abstract: This paper deals with 12 species of the genus 100–110. Dichomeris Hübner, 1818 from Southern Africa and Abstract: Shimonia gen. nov., a new genus of Kenya. Seven species are described as new: D. arquata Metarbelidae (Lepidoptera: Cossoidea), is described from sp. nov., D. brevicornuta sp. nov., D. dentata sp. nov., D. East and Central Africa (Afrotropical Region). It ligulacea sp. nov., D. paulidigitata sp. nov., D. currently comprises four species, three of which are tenextrema sp. nov. and D. ventriprojecta sp. nov.; and described as new: S. timberlakei sp. n., S. oyiekeae sp. n. other five known species are listed. Images of adults and and S. fischeri sp. n. The fourth species, S. splendida illustrations of genitalia are provided. (Fletcher, 1968) is here transferred to the new genus from Metarbela Holland, 1893. Illustrations of adult LIBERT, M. 2013. Analyse de la distribution morphology and notes on ecology and biogeography of muséologique des types des taxons africains these species are presented. d’Anthene Doubleday et des genres apparentés (Lepidoptera, Lycaenidae). Bulletin de la Société LERAUT, P. 2013. Espèces et genres nouveaux de entomologique de France, 118(3): 417–419. (Lepidoptera, Pyraloidea, ). Abstract: Analysis of the museological distribution of Bulletin de la Societe Entomologique de France the types of African taxa of Anthene Doubleday and 118(1): 41–72. allied genera (Lepidoptera, Lycaenidae). Some 280 types Abstract: New species and genera of Pyralinae have been examined by the author when he revised the (Lepidoptera, Pyraloidea, Pyralidae). Eighteen genera of African species of Anthene Doubleday and closely allied Pyralidae (Pyralinae) are studied, of which six are genera, Triclema Karsch, 1893, Cupidesthes Aurivillius, described as new: Mabokia n. gen., Cassouodes n. gen., 1895, Neurellipes Bethune-Baker, 1910 and Monile Lantzina n. gen., Barbutodes n. gen, Lantzodes n. gen. Ungemach, 1932. In the present paper, the distribution of and Nussodes n. gen. Fifty-one species are studied, these types in the various museums is examined and including the description of twenty-one species new for briefly commented on. science: Maradana modestalis n. sp., Maradana yapoensis n. sp., Mabokia delicatalis n. sp., Mabokia LIBERT, M. & COLLINS, S.C. 2013. boudinotalis n. sp, Nussodes nimbaensis n. sp., Nussodes Description of two new species of African abidjanalis n. sp., Epacternis bicoloralis n. sp. Goateria Lycaenidae in the collection of the African adelinae n. sp., Goateria bourgoinalis n. sp., Goateria Butterfly Research Institute. Metamorphosis 24: 3– ambralis n. sp., Zitha ouvrardi n. sp., Zitha andoboalis n. 6. sp., Zitha gracilalis n. sp., Sindris rogueti n. sp., Abstract: Two new species of African Lycaenidae Haplosindris gabonensis n. sp., Lantzina hodeberti n. sp., recently discovered in East Africa are described, Aslauga Lantzina rufalis n. sp., Barbutodes lantzi n. sp., gallmannae Collins & Libert sp. nov. from Kenya, and Lantzodes lantoalis n. sp., Lantzodes subpurpuralis n. Pseudaletis barnetti Libert & Collins sp. nov. from sp., Lantzodes nienokonealis n. sp. Six new synonymies Uganda. are established: Sindris catalalis Viette, 1953, n. syn. for

Sindris sganzini Boisduval, 1833, Sindris sganzini comorensis Viette, 1960, n. syn. for Sindris sganzini LISEKI, S.D. & VANE-WRIGHT, R.I. 2013. Boisduval, 1833, Sybrida Walker, 1865, n. syn. for Butterflies (Lepidoptera: Papilionoidea) of Mount Sacada Walker, 1862, Marionana Viette, 1953, n. syn. Kilimanjaro: family Pieridae, subfamily Coliadinae. for Sacada Walker, 1862, Sacada vinolentalis Viette, Journal of Natural History 47(19–20): 1309–1323. 1960, n. syn. for Sacada paulianalis (Viette, 1953), Abstract: This paper forms part of a series on the Arippara roborealis Swinhoe, 1885, n. syn. for Arippara butterfly fauna of Mount Kilimanjaro. Three genera vibicalis (Lederer, 1863). Twenty-two new combinations (Catopsilia, Colias and Eurema) with eight species of are proposed: Maradana claveriei (Rougeot, 1977), n. Coliadinae are believed to occur within the main forest comb., Goateria madegassalis (Viette, 1960), n. comb., zone. However, of these, one or two may prove to be no Goateria fotakalis (Viette, 1960), n. comb., Episindris more than variants of a third species, Eurema minutalis (Viette, 1960), n. comb., Episindris desjardinsii. A fourth conventionally recognized member boisduvalalis (Viette, 1953), n. comb., Cassouodes of the complex may occur on the lower slopes below camptoceralis (Hampson, 1906), n. comb., Lantzina 1800m. The widespread species Eurema hecabe occurs prasinalis (Hampson, 1906), n. comb., Barbutodes on the lower slopes, but records are sparse, and no albizziae (Tams, 1941), n. comb., Lantzodes vinacealis records of its close relative Eurema floricola, with which (Leraut, 2007), n. comb., Lantzodes dargei (Leraut, it has often been confused, have been found. The need for 2007), n. comb., Lantzodes obovalis (Hampson, 1917), n. original field and laboratory research on the of comb., Loryma centrafricalis (Leraut, 2007), n. comb., African Eurema species is stressed. Keys to adult Zitha seyrigalis (Marion & Viette, 1956), n. comb., Zitha Coliadinae found in Tanzania, with colour illustrations, sakarahalis (Marion & Viette, 1956), n. comb., Zitha are included as online supplementary material. Williams/ Metamorphosis 24: 57–68 64

MARTINS, D.J., COLLINS, S.C., CONGDON, showed a strong positive relationship with nectar volume, C. & PIERCE, N.E. 2013. Association between even after phylogenetic correction, which implies that the African lycaenid, Anthene usamba, and an plants with long corolla tubes are under selection to offer obligate acacia ant, Crematogaster mimosae. relatively large amounts of nectar to entice visits by polyphagous long-tongued hawkmoths. Our study shows Biological Journal of the Linnean Society 109(2): that diffusely co-evolved pollination systems involving 302–312. long-tongued hawkmoths are clearly asymmetrical, with Abstract: The African lycaenid butterfly, Anthene plants exhibiting a high degree of floral specialization, usamba, is an obligate myrmecophile of the acacia ant, while hawkmoths exhibit polyphagous behaviour. Crematogaster mimosae. Female butterflies use the presence of C. mimosae as an oviposition cue. The eggs MECENERO, S., BALL, J.B., EDGE, D.A., are laid on the foliage and young branches of the host HAMER, M.L., HENNING, G.A., PRINGLE, plant, Acacia drepanolobium. Larvae shelter in the swollen thorns (domatia) of the host tree, where they live E.L., TERBLANCHE, R.F. & WILLIAMS, in close association with the acacia ants, and each larva M.C. (eds). 2013. Conservation assessment of occupies a domatium singly. Anthene usamba are tended butterflies of South Africa, Lesotho and Swaziland: by ants that feed from the dorsal nectary organ at regular Red List and Atlas. Saftronics (Pty) Ltd., intervals. Larvae also possess tentacle organs flanking the Johannesburg & Animal Demography Unit, Cape dorsal nectary organ and appear to signal to ants by Town. everting these structures. Larvae were observed to spend most of their time within the domatia. Stable isotope MONTEIRO, A., CHEN, B., RAMOS, D.M., analysis of matched host plant/ant/butterfly samples OLIVER, J.C., TONG, X., GUO, M., WANG, revealed that Anthene usamba are 15N enriched relative W-K., FAZZINO, L. & KAMAL, F. 2013. to the ants with which they associate. These data, based Distal-less regulates eyespot patterns and on the increase in 15N through trophic levels, indicate melanization in Bicyclus butterflies. Journal of that the caterpillars of these butterflies are aphytophagous and either exploit the ant brood of C. mimosae within the Experimental Zoology Part B Molecular and domatia, or are fed mouth to mouth by adult workers via Developmental Evolution 320B(5): 321–331. trophallaxis. This is the first documented case of Abstract: Butterfly eyespots represent novel complex aphytophagy in African Anthene. Pupation occurs inside traits that display substantial diversity in number and size the domatium and the imago emerges and departs via the within and across species. Correlative gene expression hole chewed by the larva. The adult females remain studies have implicated a large suite of transcription closely associated with their natal patch of trees, whereas factors, including Distal-less (Dll), Engrailed (En), and males disperse more widely across the acacia savannah. Spalt (Sal), in eyespot development in butterflies, but Females prefer to oviposit on trees with the specific host direct evidence testing the function of any of these ant, C. mimosae, an aggressive obligate mutualist, and proteins is still missing. Here we show that the avoid neighbouring trees with other ant species. Adult characteristic two-eyespot pattern of wildtype Bicyclus butterflies are active during most months of the year, and anynana forewings is correlated with dynamic there are at least two to three generations each year. progression of Dll, En, and Sal expression in larval wings Observations made over a 5-year period indicate that a from four spots to two spots, whereas no such decline in number of different lycaenid species utilize ant-acacias in gene expression ensues in a four-eyespot mutant. We East Africa, and these observations are summarized, then conduct transgenic experiments testing whether together with comparisons from the literature. over-expression of any of these genes in a wild-type genetic background is sufficient to induce eyespot MARTINS, D. & JOHNSON, S.D. 2013. differentiation in these pre-patterned wing compartments. Interactions between hawkmoths and flowering We also produce a Dll-RNAi transgenic line to test how plants in East Africa: polyphagy and evolutionary Dll down-regulation affects eyespot development. Finally we test how ectopic expression of these genes during the specialization in an ecological context. Biological pupal stages of development alters adults color patters. Journal of the Linnean Society 110(1): 199–213. We show that over-expressing Dll in larvae is sufficient Abstract: Hawkmoths (Lepidoptera, Sphingidae) are to induce the differentiation of additional eyespots and considered important pollinators in tropical regions, but increase the size of eyespots, whereas down-regulating the frequency and degree of reciprocal specialization of Dll leads to a decrease in eyespot size. Furthermore, interactions between hawkmoths and flowers remain ectopic expression of Dll in the early pupal wing led to poorly understood. Detailed observations at two sites in the appearance of ectopic patches of black scales. We Kenya over a two-year period indicate that adult conclude that Dll is a positive regulator of focal hawkmoths are routinely polyphagous and opportunistic, differentiation and eyespot signaling and that this gene is regardless of their proboscis length. About 700 also a possible selector gene for scale melanization in individuals of 13 hawkmoth species were observed butterflies. visiting a wide range of plant species at the study sites, including 25 taxa that appear to be specifically adapted MOOLMAN, H.J., VAN DEN BERG, J., for pollination by hawkmoths. We estimate that 277 plant CONLONG, D., CUGALA, D., SIEBERT, S.J. species in Kenya (c. 4.61% of the total angiosperm flora) & LE RU, B.P. 2013. Diversity of stem borer are adapted for pollination by hawkmoths. Floral tube lengths of these plants have a bimodal distribution, parasitoids and their associated wild host plants in reflecting the existence of two hawkmoth guilds differing South Africa and Mozambique. Phytoparasitica in tongue length. Hawkmoths exhibited strongly 41(1): 89–104. crepuscular foraging patterns with activity confined to a Abstract: The diversity of lepidopterous stem borers, 20-min period at dusk and, in some cases, a similar their parasitoids and their associated wild host plants was period just before dawn. Corolla tube length appears to studied in South Africa between 2006 and 2009 and in act as a mechanical filter as the longest-tubed plants were Mozambique between 2005 and 2010. In South Africa, visited by the fewest hawkmoth species and these were 20 species of parasitoids were recovered from 17 stem exclusively from the long-tongued guild. Tube length borer species collected on 16 wild host plant species. Williams/ Metamorphosis 24: 57–68 65

From Mozambique, 14 parasitoid species were recorded within the Miridae. The intromission of the male organ is from 16 stem borer species collected on 14 wild host not performed by the usual genital route, i.e. at the base plant species. The highest diversity of parasitoids was of the ovipositor via the vulva and the vestibulum. The recorded on stem borers that attacked the host plants phallus is introduced into the ovipositor itself, at its distal Phragmites australis (7 spp.) and Panicum maximum (6 extremity, and when inserted it extends up to the genital spp.), in South Africa and Mozambique, respectively. chamber. Bracon sp. (Hymenoptera: Braconidae) and Procerochasmias nigromaculatus (Hymenoptera: PRINGLE, E.L. 2013. A new species of Ichneumonidae) were the most recorded parasitoid Cassionympha Van Son (Nymphalidae: ) species in South Africa while Cotesia sesamiae from the southern coast of the Western Cape, with a (Hymenoptera: Braconidae) parasitized most stem borer discussion of its possible evolutionary origins. species in Mozambique. The most common tachinid Metamorphosis 24: 38–43. parasitoid recorded in this study was Sturmiopsis Abstract: The genus Cassionympha Van Son is parasitica (Diptera: Tachinidae). Parasitism of stem discussed and reviewed, and a Neotype of the type borers during the off season was previously thought to species of the genus Cassionympha cassius (Godart, occur mainly in natural habitats but this study shows that 1824) designated. A new species from the southern coast although natural habitats provided refuges for some region of the Western Cape is described, and a discussion parasitoid species, stem borer parasitism was generally about the evolutionary origins of the group provided. low in wild host plants, irrespective of whether collections were done during the cropping or off-season. PRZYBYLOWICZ, L. 2013. Review of subgenus Parasitoid beta diversity did not depend on habitat or host plant species, but was determined by stem borer diversity. Compsochromia Kiriakoff 1953 (Lepidoptera: Erebidae: , genus Balacra) with NEW, T.R. 2013. Lepidoptera and conservation. identification keys and description of a new species Wiley Blackwell. from Cameroon. Annales de la Societe Entomologique de France 49(1): 53–60. NÓBREGA, R.P. 2013. Diurnal butterflies of Abstract: Subgenus Compsochromia Kiriakoff 1953 is . CIBIO, Porto & IICT, Lisbon. 288pp. briefly reviewed. An overview of the taxonomic history of the group is provided along with a checklist covering OTTO, H.H.H., GARDINER, A.J. & SHARP, all synonyms. Balacra flava n. sp. is described on the I.C. 2013. New larval host plant records for basis of eight specimens of both sexes collected in butterflies of the Kruger National Park and its Cameroon. Three separate keys, based on male and surrounds, Limpopo & Mpumalanga, South Africa. female habitus as well as on the male genitalia, are presented for the determination of Compsochromia Metamorphosis 24: 69–74. specimens. The vesica of B. compsa is described and Abstract: New butterfly larval host plants are illustrated for the first time. recorded for the Kruger National Park area, by means of oviposition observations, raising larvae to adults and/ or PRZYBYLOWICZ, L. 2013. Thyretini of Africa. larval feeding observations. Some oviposition An Illustrated Catalogue of the Thyretini observations are provided and discussed. (Lepidoptera: Arctiidae: Syntominae) of the PLUOT-SIGWALT, D. & CHEROT, F. 2013. Afrotropical Region. Apollo Books. Donnees biologiques et anatomiques, regime PYRCZ, T.W., WARREN-GASH, H., alimentaire et taxonomie d’un nouveau Fulvius LORENC-BRUDECKA, J., KNOOP, D., afrotropical (Insecta, Heteroptera, Miridae, OREMANS, P. & SAFIAN, S. 2013. Taxonomy Cylapinae, Fulviini). Zoosystema 35(1): 45–68. Abstract: Biological and anatomical data, feeding habits and distribution pattern of the African rain forest and taxonomy of a new species of an afrotropical Fulvius butterfly genus Euphaedra Hübner sensu stricto (Insecta, Heteroptera, Miridae, Cylapinae, Fulviini). with the description of three new subspecies of Fulvius carayoni n. sp. is described on the basis of Euphaedra cyparissa (Cramer) and one of E. abundant material mainly from Cameroon and Ghana. sarcoptera (Butler) (Lepidoptera, Nymphalidae, Adults can be easily distinguished among the , Adoliadini). ZooKeys 298: 1–37. Afrotropical Fulvius Sail, 1862 by the longitudinal strips Abstract: Updated data on the distribution, ecology and arising from the base of the hemelytra, by the colouration taxonomy of Euphaedra cyparissa (Cramer) and of the second antennal segment and of the scutellum, and Euphaedra sarcoptera (Butler) are presented. Three new by the male genital structures, particularly by the subspecies of E. cyparissa and one of E. sarcoptera are concomitant presence on the phallus of a hair-like apical described and their geographic distribution is presented. filamentous structure and a pair of spiculi with bases The monophyly of the genus Euphaedra sensu Hecq is posterior to the secondary gonopore. The egg and assessed based on morphological, in particular male and nymphal instars are briefly described. The presence of a female genitalia, and behavioural traits. Possible pair of ventral cephalic trichobothria is documented for evolutionary reasons for the convergence of colour the immature stages as well as for the imago. Information pattern between the sympatric subspecies of E. cyparissa about the internal anatomy of the male reproductive and and E. sarcoptera are discussed. digestive systems is supplied, as well as data concerning feeding habits, breeding and mating. Laboratory breeding RAZAFIMANANTSOA, T.M., RAMINOSOA, of F. carayoni n. sp. shows that the species is predatory: N., RAKOTONDRASOA, O.L., RAJOELISON, for four years, it fed exclusively on eggs and neonate G.L., BOGAERT, J., RABEARISOA, M.R., larvae of the pyralid Lepidoptera, Anagasta kuehniella RAMAMONJISOA, B.S., PONCELET, M., (Zeller, 1879). The anatomy, clearly of the mind type, does not reveal any peculiarity except the fact that testes HAUBRUGE, E. & VERHEGGEN, F.J. 2013. are each composed of a single follicle. In contrast, the Silkworm moths inventory in their natural tapia modalities of the mating are peculiar, never observed forest habitat (Madagascar): diversity, population Williams/ Metamorphosis 24: 57–68 66 dynamics and host plants. African Entomology Meyrick, 1920; Spilonota sinuosa Meyrick, 1917, and 21(1): 137–150. Laspeyresia ichthyura Meyrick, 1926. Abstract: Endemic silk moths (Lepidoptera: ) in Madagascar have been collected and RICHARDSON, I.D. 2013. A study of the exploited for centuries by local populations either for genitalia of actia (Distant), 1880 and Precis food or as a source of silk cocoons from which textiles pelarga (Fabricius), 1775 (Lepidoptera: are made. natural forest habitat has also been Nymphalidae). Metamorphosis 24: 7–11. degraded, leading to a drastic decrease in silk moth Abstract: The diagnostic character based on the position populations. However, very few scientific reports of a dot in a light coloured patch on the forewing is highlighted these observations well known by the local quantified for each male specimen in short series of people. We have inventoried silk moths species in tapia Precis actia and Precis pelarga from Zambia. A study of (Uapaca bojeri Baill.) forests located in the central the male genitalia reveals an additional diagnostic highlands of Madagascar. Inventories have been character, which correlates well with the quantified dot- conducted during one year from August 2009 to July position character. This additional character is proposed 2010 by sampling transects in Imamo forests. Three as a more definitive means of separating the two very species of Lasiocampidae belonging to two genera were similar species. found: Borocera cajani Vinson, Borocera marginepunctata Guerin-Meneville and Europtera SÁFIÁN, S., LIBERT, M. & COLLINS, S.C. punctillata Guenee. These three silk moth species are 2013. Two new Aphnaeus (Lepidoptera: endemic to Madagascar but only one (B. cajani) is Lycaenidae: Theclinae) species from Liberia. commercially exploited in the silk industry. The habitat, Zootaxa 3718(2): host plants, abundance, life cycle and feeding behaviour Abstract: Two new butterfly species belonging to the of these species in their natural habitat are described. genus Aphnaeus (Lycaenidae: Theclinae), A. mirabilis sp. n., A. nimbaensis sp. n. were found in the Nimba RAZOWSKI, J. & KRÜGER, M. 2013. An Mountains, Western Range, Liberia in February 2012. illustrated catalogue of the type specimens of Both are illustrated and described in comparison with in the Iziko South African Museum, closely related species: A. flavescens Stempffer, 1954 and Cape Town (Lepidoptera: Tortricidae). SHILAP A. liberti Bouyer, 1996, respectively. Genitalia of the Revista de Lepidopterologia 41(162): 213–240. male holotype of A. nimbaensis are also illustrated. Abstract: The type specimens of 36 species of Tortricidae in the collection of the Iziko South African SHARP, I.C. & SHARP, T. 2013. Observations Museum, Cape Town, South Africa are discussed and on the life history of the Southern Pied Woolly illustrated in colour, and illustrations of their genitalia are Legs, Lachnocnema laches Fabricius, 1793 provided where possible. A total of 18 new combinations (Lycaenidae: Miletinae) in the Mpumalanga is proposed: Clepsis crispata (Meyrick, 1912), comb. n. lowveld. Metamorphosis 24: 18–19. (Tortrix); Doridostoma symplecta (Meyrick, 1910), comb. n. (Tortrix); Epichoristodes adustana SIRCOULOMB, G. 2013. Les Geometridae du (Walsingham, 1881), comb. n. (Cacoecia); Parc national de Dzanga Ndoki (R.C.A): inventaire Epichoristodes cinerata (Meyrick, 1920), comb. n. et description d’une nouvelle espèce du genre (Epichorista); Epichoristodes dorsiplagana Microlyces Herbulot, 1981 (Lepidoptera) (1ere (Walsingham, 1881), comb. n. (Cacoecia); Epichoristodes exanimata (Meyrick, 1920), comb. n. contribution). Entomologia Africana 18(1): 29–36. (Epichorista); Epichoristodes niphosema (Meyrick, 1917), comb. n. (Epichorista); Epichoristodes phalaraea SHI, Q-H., ZHAO, F., HAO, J-s & YANG, Q. (Meyrick, 1920), comb. n. (Epichorista); Hectaphelia 2013. Complete mitochondrial genome of the mensaria (Meyrick, 1912), comb. n. (Tortrix); Common Evening Brown, Melanitis leda Linnaeus Hectaphelia sporadias (Meyrick, 1920), comb. n. (Lepidoptera: Nymphalidae: Satyrinae). (Tortrix); Hectaphelia vestigialis (Meyrick, 1914), comb. Mitochondrial DNA 24(5): 492–494. n. (Epichorista); Clepsis biformis (Meyrick, 1920, comb. Abstract: The complete mitochondrial genome n. (Tortrix); Nkandla macrostoma (Meyrick, 1920), (mitogenome) of Melanitis leda (Lepidoptera: comb. n. (Cnephasia); Eccopsis orichlora (Meyrick, Nymphalidae: Satyrinae) is a circular molecule of 15,122 1920), comb. n. (Argyroploce); Eudemis globigera bp in length, containing 37 typical animal mitochondrial (Meyrick, 1914), comb. n. (Argyroploce); Strepsicrates genes and 1 control region, known in insects as the AT- sinuosa (Meyrick, 1917), comb. n. (Spilonota); Fulcrifera rich region. Its gene content and order are identical to all ocnogramma (Meyrick, 1910), comb. n. (Laspeyresia); other available butterfly mitogenomes. All protein-coding and Fulcrifera ichthyura (Meyrick, 1926), comb. n. genes (PCGs) start with a typical ATN initiation codon, (Laspeyresia). Lectotypes were designated for the except for COI, which is initiated by the CGA codon as following 20 taxa in an effort to stabilize the observed in other butterfly species. A total of 97 bp of nomenclature: Pharmacis stigmatica Meyrick, 1909; intergenic spacers are interspersed in 11 regions, ranging Tortrix crispata Meyrick, 1912; Cacoecia heliaspis in size from 1 to 45 bp. The 314-bp-long AT-rich region Meyrick, 1909; Proselena ionephela Meyrick, 1909; is the smallest of all the butterfly corresponding regions Cacoecia adustana Walsingham, 1881; Cacoecia available and contains some conserved structures similar dorsiplagana Walsingham, 1881; Tortrix biformis to those found in other butterfly mitogenomes, including Meyrick, 1920; Lozotaenia elegans Walsingham, 1881; the motif ATAGA followed by a 19-bp poly-T stretch Cnephasia macrostoma Meyrick, 1920; Cnephasia and a microsatellite-like (AT)6 element preceded by the chlorocrossa Meyrick, 1926; Acharneodes atrinodis ATTTA motif. Meyrick, 1926; Argyroploce orichlora Meyrick, 1920; Eccopsis fluctuatana Walsingham, 1881; Argyroploce SMITH, D.A.S. 2013. On refining hypotheses for globigera Meyrick, 1914; Doliochastis homograpta biased sex ratios and aposematic polymorphism in Meyrick, 1920; Eucosma galactitis Meyrick, 1912; African butterflies: a commentary. Ideas in Ecology Eucosma insolens Meyrick, 1912; Eucosma tenax and Evolution 6: 17–19. Williams/ Metamorphosis 24: 57–68 67

SMITH, D.A.S. 2013. African Queens and their a major causal factor of mate-choice decisions and sexual Kin: a Neo-Darwinian Odyssey. Brambleby Books, selection. Harpenden, U.K. VAN DEN HEUVEL, J., SAASTAMOINEN, M., TAMBARA, E., MURWIRA, A. & KATIVU, S. BRAKEFIELD, P.M., KIRKWOOD, T.B.L., 2013. From natural woodlands to cultivated land: ZWAAN, B.J. & SHANLEY, D.P. 2013. The diversity of fruit-feeding butterflies and beetles in predictive adaptive response: Modeling the life- the mid-Zambezi valley, northern Zimbabwe. history evolution of the butterfly Bicyclus anynana African Journal of Ecology 51(2): 263–269. in seasonal environments. American Naturalist Abstract: We tested the effect of cultivation on butterfly 181(2): E28–E42. (Nymphalidae: Charaxes) and beetle (Coleoptera: Abstract: A predictive adaptive response (PAR) is a type Scarabaeidae: Cetoniinae) species richness and of developmental plasticity where the response to an abundance along a cultivation intensification gradient. environmental cue is not immediately advantageous but Results showed significant differences in species richness instead is later in life. The PAR is a way for organisms to and abundance between natural woodlands and cultivated maximize fitness in varying environments. Insects living landscapes with larger differences in areas of high in seasonal environments are valuable model systems for cultivation intensity. The results indicate that natural testing the existence and form of PAR. Previous woodland clearing for cultivation purposes has negative manipulations of the larval and the adult environments of impacts on diversity, a situation more severe in the butterfly Bicyclus anynana have shown that highly intensified cultivated areas. Our results imply that individuals that were food restricted during the larval mosaics of different land-use units, each in a different stage coped better with forced flight during the adult phase of clearance-cultivation-abandonment-recovery- stage compared to those with optimal conditions in the clearance cycle could counter the negative effects of larval stage. Here, we describe a state-dependent energy cultivation intensity on arthropod diversity. allocation model, which we use to test whether such a Cetoniinae, Charaxes: Farming and agriculture, response to food restriction could be adaptive in nature Agricultural intensification, Richness, abundance, where this butterfly exhibits seasonal cycles. The results Conservation, Species diversity, Richness, abundance, from the model confirm the responses obtained in our Zimbabwe, mid-Zambezi valley, Mbire Rural District, previous experimental work and show how such an Dande Communal Area. outcome was facilitated by resource allocation patterns to the thorax during the pupal stage. We conclude that for B. TROPEK, R., JANSTA, P. & LESTINA, D. anynana, early-stage cues can direct development toward 2013. Acraea wigginsi occidentalis (Bethune- a better adapted phenotype later in life and, therefore, that Baker, 1926), a new butterfly for , with a PAR has evolved in this species. remarks on its habitat and known distribution VAN VELZEN, R., WAHLBERG, N., SOSEF, (Lepidoptera: Nymphalidae). SHILAP Revista de M.S.M. & BAKKER, F.T. 2013. Effects of Lepidopterologia 41(161): 163–165. changing climate on species diversification in Abstract: Acraea wigginsi occidentalis (Bethune-Baker, tropical forest butterflies of the genus Cymothoe 1926) [now Telchinia wigginsi occidentalis] is recorded in Nigeria for the first time, in the Gotel Mountains, close (Lepidoptera: Nymphalidae). Biological Journal of to the Cameroonian border. All of its known distribution the Linnean Society 108(3): 546–564. is summarized, including four new records from Abstract: Extant clades may differ greatly in their Cameroon. Its occurrence in the degraded submontane species richness, suggesting differential rates of species grasslands is also discussed as the reason why it has been diversification. Based on phylogenetic trees, it is possible rarely observed. to identify potential correlates of such differences. Here, we examine species diversification in a clade of 82 VAN BERGEN, E., BRAKEFIELD, P.M., tropical African forest butterfly species (Cymothoe), HEUSKIN, S., ZWAAN, B.J. & NIEBERDING, together with its monotypic sister genus Harma. Our aim C.M. 2013. The scent of inbreeding: a male sex was to test whether the diversification of the Harma/Cymothoe clade correlates with end-Miocene pheromone betrays inbred males. Proceedings of global cooling and desiccation, or with Pleistocene the Royal Society Biological Sciences Series B habitat range oscillations, both postulated to have led to 280(1758): 1–8. habitat fragmentation. We first generated a species-level Abstract: Inbreeding depression results from mating phylogenetic tree for Harma and Cymothoe, calibrated among genetically related individuals and impairs within an absolute time scale, and then identified reproductive success. The decrease in male mating temporal and phylogenetic shifts in species success is usually attributed to an impact on multiple diversification. Finally, we assessed correlations between fitness-related traits that reduce the general condition of species diversification and reconstructed global inbred males. Here, we find that the production of the temperatures. Results show that, after the divergence of male sex pheromone is reduced significantly by Harma and Cymothoe in the Miocene (15Mya), net inbreeding in the butterfly Bicyclus anynana. Other traits species diversification was low during the first 7Myr. indicative of the general condition, including flight Coinciding with the onset of diversification of Cymothoe performance, are also negatively affected in male around 7.5Mya, there was a sharp and significant increase butterflies by inbreeding. Yet, we unambiguously show in diversification rate, suggesting a rapid radiation, and that only the production of male pheromones affects correlating with a reconstructed period of global cooling mating success. Thus, this pheromone signal informs and desiccation in the late Miocene, rather than with females about the inbreeding status of their mating Pleistocene oscillations. Our estimated age of 4Myr for a partners. We also identify the specific chemical clade of montane species corresponds well with the uplift component (hexadecanal) probably responsible for the of the Eastern Arc Mountains where they occur. We decrease in male mating success. Our results advocate conclude that forest fragmentation caused by changing giving increased attention to olfactory communication as climate in the late Miocene as well as the Eastern Arc Mountain uplift are both likely to have promoted species Williams/ Metamorphosis 24: 57–68 68 diversification in the Harma/Cymothoe clade. Cymothoe colonized Madagascar much later than most other insect lineages and, consequently, had less time available for diversification on the island. We consider the diversification of Cymothoe to be a special case compared with other butterfly clades studied so far, both in terms of its abrupt diversification rate increase and its recent occurrence (7Myr). It is clear that larval host plant shift(s) cannot explain the difference in diversification between Cymothoe and Harma; however, such a shift(s) may have triggered differential diversification rates within Cymothoe.

WILLIAMS, M.C. 2013. Publications on Afrotropical Lepidoptera during 2013. Metamorphosis 24: 57–68. Abstract: The articles published during 2013 which deal with scientific research into Afrotropical Lepidoptera are listed alphabetically by author and abstracts are provided if they were published. Articles dealing with control of Lepidoptera as pests are excluded.

WILLIS, C.K. 2013. Book review: Conservation assessment of butterflies of South Africa, Lesotho and Swaziland: Red List and Atlas – Mecenero et al. (eds.). Metamorphosis 24: 15–17.

WOODHALL, S.[E.] 2013. Observations on the life history of the Blotched Leopard, Lachnoptera ayresii Trimen, 1879 (Nymphalidae: Heliconiinae). Metamorphosis 24: 1–2.

YAKOVLEV, R., IVINSKIS, P., RIMSAITE, J. & SALDAITIS, A. 2013. Description of two new species of Meharia Chretien, 1915 (Lepidoptera: ) from East Africa. Zootaxa 3635(5): 587– 590.

YAKOVLEV, R. & MURPHY, R.J. 2013. The Cossidae (Lepidoptera) of Malawi with descriptions of two new species. Zootaxa 370(4): 371–393. Abstract: An annotated list of Cossidae of Malawi including 30 species from 13 genera and 3 subfamilies is presented for the first time. Two new species are described: Macrocossus grebennikovi Yakovlev, sp. nov. and Strigocossus elephas Yakovlev, sp. nov. Three new synonyms are established: Strigocossus moderatus (Walker, 1856) = Strigocossus leucopteris Houlbert, 1916, syn. nov. = Xyleutes sjoestedti vosseleri Gaede, 1930, syn. nov. and Strigocossus capensis (Walker, 1856) = Azygophleps kilimandjarae Le Cerf, 1914, syn. nov.