JOURNAL OF

LEPIDOPTERISTS' SOCIETY

Volume 57 2003 Number 1 j ournal of the Lepidopterists' Society 57(1), 2003, 1-16

AN'J'-ASSOCIATION AMONG SOUTHERN AFRICAN

ALAN HEATH! Department ofZoolo!,'Y, University of Cape Town, Rondebosch 7700, South

AND

ANDRE J, M. CLAASSENS 2m, High Level Road, Sea Point 8005,

ABSTRACT. Known life histOIY data, -associations and larval feeding hahits for southern African lycaenids are summarized and dis­ cussed with a focus on recently acquired knowledge. Of the 392 lycaenid represented in southern Africa, over three quarters are ant­ associated, two thirds of which are obligate. The is represented hy two tribes of algae/lichen feeders that are not ant-associated. Of lhe , three qWllters are obligately ant-associated, and all are believed to prey on Homoptera or their secretions during their larval stage. Within the , the tribe Theelini are all believed to he facultative. The trihe Aphnaeini constitutes one third of aillycaenids, almost all ohligately ant-associated. The tribe accounts for a third of aillycaenids and is 95% ant-associated, containing similar proportions of facultative and obligate associations. The presence, absence and function of myrmecophilous organs at various larval stages is discussed. Ovipositing below the soil or on sand su rface is recorded for the first time. The various trophit: strategies oflarvae are discussed. () and () together account for almost 80% of obligate relationships. It is suggested that many synonymies exist among obligately ant -associated taxa. Additional key words: evolution, myrmccopbily, trophallaxis, aphytophagy, entomophagy, detritus, acoustics, South Africa.

Of 3607 Afrotropical species, 42% are in head protected beneath a tough carapace. Many such the family Lycaenidae (Ackery et al. 1995), a similar larvae have organs that serve speCific functions in their figure (47%) is given for southern Africa2 by Pringle et association with ants. A dorsal nectary organ (DNO), al. (1994). Of the 397 Australian , 36% are in present on the seventh abdominal segment of some the family Lycaenidae (Braby 2000). These figures im­ larvae provides honeydew for ants to imbibe. A pair of ply that a higher level of diversification has occurred in tentacle organs (TOs) located on the eighth abdominal the Lycaenidae than in other families, and Pierce segment and several minute perforated cupola organs (1984) suggested that this may have been caused by (peOs) are present on the larva's cuticle which all se­ ant-association (myrmecophily). Many species of ly­ crete substances that can influence ant behavior (Cot­ caenids are ant-associated in the early stages (Fiedler treIl1984). Note that ant-organs found in ant-associated 1991), ants being among the leading predators of in­ Riodininae differ from. those in other lycaenids; see sects (Holldobler & Wilson 1990). Lycaenid larvae are DeVries (1991, 1997) for a comparison. By using these vulnerable to predatory ant species, hence various organs, free-living mutualistic caterpillars proVide strategies have evolved to prevent or reduce attack. nutritious secretions and emit chemical signals (Hen­ Dense and long hair serves as one effective strategy, ning 1983b) that can manipulate ant behavior to re­ endophagy (feeding inside plant material) is another. duce aggression and obtain protection from predators Other lycaenids possess an extra-thick cuticle with the and parasitoids (Lenz 1917, Hinton 1951, Pierce & Mead 1981, Pierce 1984, Pierce et al. 1987, DeVries I Address for correspondence: 1 The Close, Limekiln Lane Bal­ dock, Herts, SG7 6PJ U.K. (Email: alan.heath:[email protected]). 1988, 1991, Fiedler 1991). In some cases this manipu­ 2 Southern Africa: Countries south of , Zambia and the lation is extended to inducing regurgitations from Zambezi River in Mozambique. The region includes Namibia, Botswana, Zi m bahwe, South Africa, Swaziland, and south­ adult ants (trophallaxis) or enabling the larva to prey ern Mozambique. directly on the ant brood (parasitism) (Henning 1983b, 2 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

Pierce 1995, Heath 1998, Elmes et aI. 1991, Thomas Our classification here follows Scott (1985), Eliot (in 1983, Thomas & Wardlaw 1992). Corbet et al. 1992), Ackery et al. (1999) and Pierce ct The DNO was first described by Guenee (1867), al. (2002), who treat Riodininae3 as a subfamily within subsequently more knowledge was gathered about the Lycaenidae, along with Poritiinae (including Pentilini myrmecophilous organs (Newcomer 1912, Ehrhardt and Liptenini), Miletinae (Miletini, Liphyrini and 1914, Hinton 1951, Clark & Dickson 1956). Their Spalgini) and Lycaeninae (Theclini, Aphnaeini, Ly­ structure and function was studied in greater detail by caenini and Polyommatini). Examples of adult Iy­ Malicky (1969, 1970). More recently, secretions from caenids from the region are illustrated (Figs. 1-20). these organs were chemically analyzed (Maschwitz et al. 1975, Pierce 1983, DeVries & Baker 1989). Many, METHODS USED FOR STUDYING LYCAENIDS IN THE but not all ant-associated Iycaenids possess these or­ FIELD AND LABORATORY gans, whose presence and pOSSibly function can vary Inducing oviposition. Inducing oviposition facili­ throughout the larval stage (Cottrell 1984). Some Iy­ tates life history studies in the laboratory. The method caenid tribes and genera are known to be more for inducing oviposition among Chrysoritis Butler is to strongly ant-associated than others, but even within a place not more than three known or 'suspected' host Single , the type of association can be quite var­ ants together with some stems preferably of a known ied (Pierce 1989). larval foodplant, inside an open plastiC container about Ant-associated Iycaenid larvae are known to pro­ 130 x 100 mm and 70 mm deep covered tightly with duce a substrate borne call to recruit ants (DeVries fine netting (e.g., ladies' stockings) and kept warm, but 1988, 1mJO, 1991, Travassos & Pierce 2000). Southern not hot, or in the sun. A female Chrysoritis is added African aphnaeine larvae also produce sounds (Heath once the ants have settled down. More than three ants 1998), as do pupae (Schlosz 1991, Claassens 1991). often results in an escape hole chewed in the netting. Numerous field and laboratory studies have con­ The female butterfly is fed daily with a small lump of tributed to our knowledge of African ant-Iycaenid as­ tissue paper soaked in weak sugar water placed on the sociations. One of the earliest accounts detailing ant­ netting. Eggs are often laid on the netting. Attempts to association in African Iycaenids was Lamborn (1914) induce oviposition among Hubner in this who recorded observations on early stages of 27 manner have been unsuccessful, but oviposition species (14 genera) hom southern Nigeria. Nine of among captive Thestor Hubner occurred on almost these genera are also represented in southern Africa. any surface without either ants or vegetation present Observations in and Uganda by Jackson (1937) (Heath & Claasscns 2000). described obligate ant-associations in seven species Obtaining early stages in the field. Methods for and discussed 25 facultative associations. The first ma­ finding early stages in the field vary with life history jor study on South African Iycaenid early stages was by characteristics. Many myrmecophilous species spend Clark and Dickson (1971), who described and illus­ part or all of their juvenile phases in or close to subter­ trated 125 species at different stages of their develop­ ranean ant nests which makes them difficult to dis­ ment, including myrmecophilous organs. Behavioral cover and study. Others are found under rocks and as­ studies under laboratory conditions were done subse­ sociated with a particular plant or ant nest. The quently by Claassens (1972, 1976) and Claassens and method used to find Chrysoritis larvae is to search the Dickson (1977). On a global scale, lycaenid-ant associ­ debris beneath potential food plants where ants are ations have been reviewed by Cottrell (1984), Fiedler present. On occasions, the larvae are found in small fi­ (1991) and Pierce et al. (2002). brous shelters built on the plant stems by the ants, pre­ This paper summarizes and discusses our under­ sumably intended for Homoptera. For Aloeides we standing of myrmecophilous lycaenids in southern search in the soil beneath potential food plants or be­ Africa during the past ten years. Recent accounts in­ neath nearby rocks. In Trimenia Tite & Dickson, the clude: Schlosz and Brinkman (1991), Williams and early stages are sought by digging in the gravel or by Joannou (1996), Heath and Brinkman (1995a, b ), prizing open cracks in bedrock where nests of the host Heath (1997a, b, 1998), Claassens and Heath (1997), ants occur (Heath & Brinkman 1995b). For Phasis Edge and Pringle (1996), Heath and Claassens (2000), Hubner, hollowed out stems of the foodplants are ex­ Lu and Samways (2001 ). The number of species for amined where ants are present. For Thestor and T.1f­ each genus follows Pringle et al. (1994), Ackery et al. lopaedia Tite & Dickson, we search beneath large (199,5), Williams (1999) and Heath (2001), A compre­ boulders covering ant nests in areas where the adults hensive list of food plants and ant-associates for south­ ern African is found in Kroon (1999). 3 Riodininae is not represented in southern Africa. VOLUME .57, NUMBER] 3

early stages can be studied in the natural environment. It is nevertheless possible to rear these in captivity with or without the host ant.

CATEGORIES OF ANT-ASSOCIATION AND FEEDING STRATEGIES There have been many attempts to categorize feed­ ing strategies and relationships between lycaenids and ants; e.g., Fiedler (1991), DeVries (1991), Pierce (1995), Eastwood and Fraser (1999), Heath and Claassens (2000) and Pierce et a1. (2002). The follow­ ing three broad categories of ant-association are used here: (1) 'Not ant-associated' (direct and close interac­ tion with ants is absent or rare, even if ants are pres­ ent). (2) 'Facultative' (intermittently attended by ants but not wholly dependent on them for survival under field conditions). (3) 'Obligate' (dependent on one species of ant for survival under field conditions). Three categories of larval feeding strategy used here are 'algae/lichen-feeding', 'herbivorous' and 'ento­ FIGS. 1-20. Lycaenidae from southern Africa. 1, lolaus (10- mophagous'. The latter term being re-defined by laphilous) trimeni Wallengren. 2, Pilasis thero thero (Linnaeus) fe­ Pierce et a1. (2002) to mean dependant upon any in­ male. 3, Myrina .Ii/anus ficedula Trimen male. 4, Thestor protUnt.nl1s protl1mm.ls (Linnaeus) female. 5, (Wallengren) sect-derived resource, and may include homopteran male. 6, Aloeides pallida grandis Tite & Dickson male. 7, Erikssonia secretions, ant regurgitations ancl!or the them­ acraeina Trimen male. S, tropicalis tropicalis (De Boisduval) selves (carnivory). More than one of these categories mal e. 9, Tylopaedia sardonyx sardonyx (Trimen) female. 10, Cigari­ tis natalensis (Westwood) mal e. 11, LepicZochrysops trimeni may be employed during the larval phase. (Bethnne-Baker) male. 12, Lepidochrysops oreas areas Tite female. 13, Chrysoritis hrook.\·j brooksi (Riley) female. 14, Chrysuritis GENERA, LIFE HISTORY AND ANT-ASSOCIATIONS thysbe thysbe (Linnaeus) female. 15, Chrysuritis nigricans nigricans (AUlivillius) male. 16, ChnjSoritis palmus palmus (Stoll) female. 17, For completeness, all 641ycaenid genera occurring Lycaena orus Stoll male. IS, Lepiduchrysops robertsoni Cottrell in southern Africa are included here, whatever their male. 19, Lachrwcnema bihull1S (Fabricius) female. 20, Axiocerces ant-association. More speCific details are shown in amanga (Westwood) male. Table 1 and summarized in Table 2. are known to fly. Rocks under which juvenile Thestor PORITIINAE: Larvae of Pentilini and Liptenini species are found, are often turned over by baboons (together including 13 genera) are clothed with long looking for Thestor larvae and pupae, and scorpions. hairs and not directly ant-associated; they possess no Study of behavior in captivity. Due to the sub­ DNO or TOs. Larval food is mostly algae or lichen terranean habits of many juveniles, dose study of the (Bampton 1995), but were recorded taking interactions between ants and larvae in nature is not honeydew from Homoptera in the company of ants feasible. Hence, for taxa like Aloeides, Thestor, Lepi­ (Pringle et a1. 1994), although ants ignore the larvae dochrysops, Orachrysops and Trimenia, where a (Jackson 1937). formicarium similar to that described by Claassens Pentilini (three genera); De Boisdllval, (1972,1974) is required, a nest of host ants with brood 1847; Pentila Westwood, 1852; and queen is installed within the formicarium. Before Bethune-Baker, 1914. being introduced into the nest section of the formicar­ Liptenini (ten genera): Trimen, 1862; ium, larvae are left among the ant brood for about two Durbaniella van Son, 1959; Durbaniopsis van Son, hours. This is to acquire the scent of the ant colony 1959; H. H. Druce, 1905; and avoid subsequent attack by ants, but also to see Butler, 1872; Euthecta Bennett, 1954; what interaction takes place between larvae and Kirby, 1887; Stempffer & Bennett, 1953; brood. Pieces of suspected food plant are kept in water Cnodontes Stempffer & Bennett, 1953; Deloneura within the arena, as described by Claassens and Dick­ Trimen, 1868. son (1977). Chrysoritis species do not generally enter MILETINAE: Larval food can be regurgitations ant nests but are constantly attended while feeding on from ants but is mostly Homoptera or their secretions. the foodplant and resting in the debris beneath it; their Homoptera are almost always attended by ants but di- 4 JOURNAL OF THE LEPIDOPTERISTS' S OCIElY

TA.BLE l. Life history details for all known Southern African lycaenids, showing presence of D NO or TOs, lmval feeding category' and ant-as- sociation. Abbreviations: * = probable but unconfirmed. DNO, TOs: + = yes; - = no; Feeding category: A = Algae or Lichen feeder; H = Her- bivorous; E = Entomophagous. Ant-association: N = none; F = facultative; 0 = obligate; ? = unknown. Principal sources: 1 = Clark and Dickson (1971); 2 = Pringle et al. (1994); 3 = Jackson (1937); 4 = Heath and Claassens (2000); 5 = Heath (1997); 6 = Bampton and Congdon (pers. com.); 7 = Stempffer (1967); 8 = Henning (1983a, b); 9 = Claassens (1976); 10 = Edge and Pringle (1996); 11 = Schlosz and Brinkmann (1991 ); 12 = Heath (pers. obs.); 13 = Kroon (1999); 14 = Lamborn (1914); 15 = Heath and Blinkmann (1995b); 16 = Williams and Joannou (1996); 17 = Lu and Samways (2001 ); 18 = Larsen (1991 ); 19 = Pennington (1956); 20 = Atsatt (1981); 21 = Braby and Woodger (1994); 22 = Henning and Hen- ning (1982); 23 = Henning (1984a); 24 = Acke,y and Rajan (1990); 25 = van SOll1 eren (1974); 26 = Claassens and Dickson (1977); 27 = Claassens and Dickson (1980); 28 = Fiedler and Hagemann (1992); 29 Clark and Dickson (1956); 30 = Congdon and Bampton (1995); 31 = Edge (1990).

Feeding Ant- Principal Genus Species DNO TOs categOly" association Ant sources PORITIINAE Pentilini (8) Alaena amazoula A N 1,29 brainei A N 2,6 nyassa A N 2,6 Inar[f0dtacea A N 2, 6 Pentila pat! 'I A N 12 swynnertoni A N 2,6 tropicalis A N 2, 6 Ornipholidotos peucetia A N 2, 12, 18 Liptenini (22) A N 1, 12,29 limbata A N 2, 12 Durbaniella clarki A N 2, 12 Dllrhaniopsis saga A N 2, 12 Cooksonia neavei A "l 2 Mimacraea marshalli A N 2 neokoton A N 2 Eutheeta cooksoni ? ? A* N" 2 Teriomima pllellaris ? ? A N 2 puella ? ? A N 2 zulllana A N 2 Balioehil" aslanga ') ') A N 2 harnesi ? ? A 1\ 2 neavei ? ? A N 2 lipara ? ? A N 2 singlaris ? ? A N 2 Cnodontes 3 species ? ? A* N* 19 Deloneura shel,pardi A N 1 'I'm an A N 2, ,3 , 19 subfusea A 1\ 2,.3,6,19 MILETINAE Spalgini (1 ) Shalgis lenwlea ? ? E N 3,8,14 Lac nocnemini (32) Lachnoenema bibulus E N 1, Il, 14, 29 durbani E N 1,2, 29 brimo - * - * E 1\* 2 Thestor bastltus E 0 A custodiens 1, 4, 16, 29 brachycerus E 0 A custodiens 12 pictus E 0 A eustodiens 12 protumntls E 0 A C11stodiens 1, 12 riley; E 0 A cllstodiens 12 -rOSSOl1tvi E 0 A eustodiens 12 strutti E 0 A custodiens 4 yilclizae E 0 A ellstodieHs 4 dicksoni E 0 A Gllstodiens 1,2, 29 +20 species E* 0* A cust()diens * 2 Liphyrini (3) AsZau ga 3 species +* E* N* 3, 6,7, 14 J YCAEN1NAE Aphnaeini (131) Aphnaeus eriksson; +* +* H* 0 Crematogaster sp. 2 hutchinsonii + + H 0 Crc1Jwtogaster sp. 1, 3,29, .31 marshalli +* +* H* 0* Crematogaster sp. * 2 + + H 0 C. castanea 1,2,6,29 ella + + H 0 C. castanea 1,2,6, 18 namaqua + + H 0 Crematogaster sp. 8,6 phanes + + H 0 C. castanea 8,6 apelles +* +* II 0 Crerrwtogaster sp. 2,6 +5 species +* +* H* O' Crematogaster sp. * 2,6 L~ aphnaells aclerna +* +* H 0* Crematogaster sp.' 6,30 C lorosela, pselldozeritis + + H 0 C. gerstaeekeri 2, 6,3 argentea +* +* H 0 Crematogaster sp. 2, 6 1nazoensis +* +* H 0 Crematogaster sp. 2,6,19 VOLUME 57, NUMBER 1 5

TABLE 1. Continued.

Feeding Ant- Principal Genus Species DNO TOs categOlY' association Ant sources Zeritis sorhagenii ? H ? 2 Axiocerses tjoane + + H F 1, 6, 29b amanga + + H F' 3,6 ~unlce~ +* +* H F 2, 6 Phasis raue" +* + H 0 C. peringueyi 1,4 clavum +* +* H 0* Crematogaster sp. * 2 pringlei + + H 0 C. peringueyi 12, 13 thero +d + H 0 C. peringueyi 1,4,12,29 Tylopaedia sardonyx + + H 0 C. melanogaster 11, 29 Argyraspodes argyraspis ? ? II ? ? 5 Trimenia arg~roplaga ? + II 0 A Gustodiens 12 ma agrida ? + ? 0 A. GUstodiens 15 +3 species ? + ? 0* A. GUstodiens* 2 Aloeides apicalis +' + H 0 M on01Jwrium fridae 4 aranda +f + II 0 capensis 1,4,29 clarki + + H 0* Unidentified sp' 1,2 damarensis + + H 0* Unidentified sp' 1,2 dentatis ?g + H 0 L. cileensis 2, 4, 8 depicta + + H 0* Uni entified sp. * 1,2 gowani + + II 0* Unidentified sp. * 1 pallida + + H, Eh 0 L. capensis 4 pierns + + II 0 L. capensis 4,29 thyra + + H 0 L. capensis 2,4,26 trimeni + + II 0 L. capensis 1, 2 rossouwi +* +* II 0 sp. 22 taikosama + +* H 0* Unidentified sp. * 2,29 almeida + +* H 0* Unidentified sp' 2,29 +39 species +* +* II 0* Unidentified sp. * 2 Erikssonia acraeina + + II 0 Lepisiota sp. 2,23 Chrysoritis adonis + + II 0 C.liengmei 5 aethon + + H 0 C.liengmei 5 aridus + + H 0 Crematogaster sp. 5 aureus + + H 0 C.liengmei 5 azurius + + II 0 C.liengmei 5 beaufortia + + II 0 C. peringueyi 5 beulah + + II 0 Crematogaster sp. 5 blencathra + + II 0 Crernatogaster sp. 5 braueri + + II 0 C.liengmei 5 brooksi + + II 0 C. peringueyi 5 chrysantas + + H 0 C. melanogaster 5 chrysaor + + II 0 C. liengmei 5,29 daphne + + II 0 C.liengmei 5 dieksard + + E 0 C. peringuey 5 endk,mian + + II 0 C. peringuey 5 felt ami + + H 0 C. peringuey 5,29 irene + + H 0 Crematogaster sp. 5 lyeegenes + + H 0 C.liengmei 5 lynGUrium + + II 0 C. liengmei 5 midas + + H 0 C. peringueyi 5 natalensis + + II o· Crematogaster sp. * 5 nigricans + + H 0 Crematogaster sp. 5 oreas + + H 0 Myrmicaria nigra 5 orientalis + + II 0 C. liengmei 5 palmus + + II 0 C. liengmei 5, 29 pan + + H 0 C.liengmei 5 pelion + + II 0* Crematogaster sp. * 5 penni.ngtoni + + H 0 Crernatogaster sp. 5 perseus + + H 0 C. melanogaster 5 phosphor +* +* II* 0* Crmnatogaster sp. * 5 plutus + + II 0 C. peringueyi 5 pyramus + + H 0 C. peringueyi 5 pr°eiS + + H 0 Myrmicaria nigra 5, 29 rt ey' + + H 0 C. reringueyi 5 swanep()eli + + H 0 C . .iengmei 5 thysbe + + H 0 C. peringueyi 5, 29 trinwni + + H 0 C. ~ eringueYi 5 turneri + + H 0 C. iengmei 5 uranus + + H 0 C.liengmei 5 violescens + + H 0 C. peringueyi ,5 zeuxo + + H 0 C liengmei ,5 zonarillS + + H 0 C. perinfteyi 5 Crndaria leroma + + II 0 A. Gusto iens 1,5, 12,29 wykehami + + H 0 A. eustocliens 12 capensis +* +* H* 0* A. GUstodiens* 12 Lycaenini (2) Lycaena clarki H N 1, 12 onts H N 1, 12 6 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

TABLE 1. Continued.

Feeding Ant- Principal Genus Species DNO TOs category" association Ant sources Theclini (47) Amblypodiiti [2] + + H F 1,3,14,29 dermaptera + + H F 1, 29 lolaiti [21] lolaus aemu.!"s + + H F* 1,2,29 alienus + + H F 1,2,29 ap hnaeoides +* +* H F* 2 australis +* +* H F* 2 bakeri +* +* H F* 2 bowkeri + + H F 1, 2,12,29 diarnetra +* +* H F* 2 lalos +* +* H F* 2 lulua +* +* H F* 2 lnimosae + + II F 1,2,12,29 nasisi-i +* +* II F* 2, 12 obscurus +* +* II F* 2 pallene +* +* II F* 2, 12 penningtoni +* +* II F* 2 poultoni +* +* H F* 2 sidus + + H F* 1,2, 12,29 silarus + + II F 2, 12 silas + + II F 1, 2, 12,29 suhinfuscata +* +* H F 2, 12 tri:meni +* +* II F* 2,8,12 violacea +* +* H F* 2, 12 Hypolycaenili (9) Hypolycaena philippus + II F 1,3, 14, 29 lochmophila +* * II F* 2 caeculus ? ? H F 2,6 +2 species +* II* F* 2 Leptomyrina hirunclo + II F 1,2 lara + II F 1, 2,27,29 ~orgias + II F 1, 2 eni'ingi +* * II F* 2 Oeudorigiti [15] 15 + H F 1,2,29 dariaves +* II F* 2 dinochares + H F 1,2,24 dinomenes +* II F* 2 dioc/es + II F 1,2,29 lorisona +* II F* 2 magda +* H F* 2 penningtoni +* H F* 2 vansoni +* II F* 2 caerulea +* H* F* 2 zeloides +* - * II* F* 2 Capys alphaeus + II F 1,27,29 penningtoni + H F 2, 12 disjunctus + H F 1,2,12 connexivus + H F 2, 12 Poiyommatini (146) Lycaenesthiti [26] + + II F 1,3, 29 butleri + + II F* 1,29 contrastata +* +* H F* 2 crawsho,yi + + II F* .3 definita + + II F 1,27,29 kersteni + + H F l ' lernnos + + H F* 1,29 liodes + + II F* 2,14 111nlliata + + II F 3 wilsoni ? 0 Unidentihed sp. 3 atacili" + + II F 1, 6,25,29 talboti + + H F* 1 nigeriae + + H 0 Unidentified sp 3 +13 species ? ? H* F* 2 Polyommatiti [120] Cllpido"sis cissu,,;' + + tJ y * 1, .3 johates + + H y * 1 sichela ? ? H ? 2 Lampides boeticus + + H F 1,3, 27, 29 UrarwthaunUl antinorii * * H N* 2 poggei H N 2,6 VOLUME 57, N UMBER 1 7

TABLE l. Continued.

Feeding Ant- Principal Genus Species DNO TOs category" association Ant sources nubifer H N 2,3 vansomereni * H N* 2 Cacyreus dicksoni H N 1, 2, 27 lingeus + H N 1, .3, 27, 29 marshalli H N 1, 2,27, 29 tespis H N 1,2,27, 29) viri/is + H N 1,2, 27 H arpendyretts notoba + H F* 1 tsomo +* * H F* 2 l10qttaSa +* * H F* 2 Leptates 'birithotts + + H F* 1, 3,27,29' revidentatus + + H F* 1 jeanneli + + H F* 2,29 babattlti +* +* H F* 2 pulcher +* +* H F* 2 calice + + H F* 1, 2 melaena + + H F* 1, 2,29' hesperis +* +* H F* 2 Tamcus SKharis + + H F* 1 t esp's + + H F 2, 27, 29 bowker·j + + H F* 1, 29 Z.intha hintza + + H F 1, 2,3 Zizina antanossa + + H F* 1 Zizeeria knysna + + H F 1,12, 28 , 29 + + H F* 1,29 stellata + + H F* 1, 29 Zizula hylax + + H F* 1, 21 Brephidium m etophis + + H F* 1,20 Oraidium barberae ? ? H F* 1, 2 Az.antts ubaldus + + H F* 1, 29 jesous + + H F* 1,3,29 natalensis + + H F 1,3 moriqua + + H F* 1,29 1nirza +* +* H F* 2 Eicochrysops rnessapus + + H F* 1, 29 eicatrachi/us +* +* H F* 2 hippocrates + + H F* 1 + + H F 1,3 barkeri + + II F 1 malathana + + H F 1,3,14 dolorosa + + H F 1, 2,29 subpallida + * +* H F* 2 Lepidochrysops asteri" + H , E*'" 0* Camponotus sp. * 1,2, 29" bacchus + H , E* 0* Campanottts sp. * 1, 2,29 ignota + H, E 0 C. niveosettts 2, 8 ketsi + H.E* 0* Camponotus sp. * 1,2 methymna + H,E 0 C. rnacuiatlls 1, 2,29 areas + H , E 0 C. rnacu/atlls 1, 2 patricia + H , E 0 C. maculatlls l , 2, 29 puncticilia + H , E* 0* Camponotlls sp. * 1, 2 trirneni + H,E 0 C. niveosetus 1,2 oariablis + H , E 0 C. niveosetlts 1, 2 +49 species +* * H ,E* 0* Campanotus sp. * 2 Orachrysops iacrirnosa + + H* F* 1, 2,29 niobe + + H* F*o 10, 2 ariadne + + H* 0 C. nataiensis 17,2 + 7 species +* +* H* F* 2 OberU'ftiu bueronica ? ? H F 2 traC!~lu s + + H F* 1 Themwniphas mlcYlts ? ? H N 2,6 'For Feeding category, and Ant-association, see section "Categories of ant-association and feeding strategies" above. b As A. bambana (m isidentified ). ' A. aman~a is attended only by Pheidole species but often found unattended (Bampton pers. com. ). d P thera las DNO but not in final instar. "Has DNO but absent in final instar (also A. depicta, A. pallida and A. thyra ). ' DNO present in final instar (also A. piems ).

TABLE 2. Summary of ant-association: Conf. = based on published and confirmed observations. Predict. = confirmed + predicted but un­ confirm ed associations.

Obligate Facultative None Unknown Total species Conf. Predic. Conf. Predic. Conf. Predic. Conf. Predic. PORITIINAE 30 Pentilini 8 0 0 0 0 8 8 0 0 Liptenini 22 0 0 0 0 18 22 4 0 MILETINAE 36 Spalgini 1 0 0 0 0 1 1 0 0 Lachnocnemini 32 9 29 0 0 2 3 21 0 Liphyrini 3 0 0 0 0 0 3 3 0 LYCAENINAE 326 Aphnaeini 131 66 126 3 3 0 0 62 2 Lycaenini 2 0 0 0 0 2 2 0 0 Theclini 47 0 0 20 47 0 0 27 0 Polyommatini 146 9 62 15 73 8 10 114 1 TOTALS 392 84 217 38 123 39 49 231 4 rect interaction between ant and larva may not neces­ South Africa. Early ins tar larvae of LachnocnenUt bibu­ sarily occur. lus feed on young psyllids (Homoptera) and their Liphyrini (one genus): Aslauga Kirby, 1890 (14 droppings. Mature larvae creep up behind adult psyl­ Afrotropical species, three in southern Africa). Scarce, lids, seize their wings, and devour them. Larvae do not arboreal butterflies with a distinctive wingshape. Vir­ associate with ants directly but ants are always present tually nothing is known of the early stages of southern tending the Homoptera. African Aslauga species but suspected to be the same Thestor Hubner, 1819 (29 species, all endemic to as those observed by Jackson (1937) and Lamborn southern Africa). Medium-size, moth-like, with stout (1914). Larval skin is leathery, the carapace being ex­ bodies, and either blackish or yellow and brown; they traordinarily heavy; head small and carried on an ex­ always settle on the ground or rocks. Adults possess a tendable neck which can be retracted under a cara­ vestigial probOSCiS and do not nectar. Most Thestor pace; TOs present, but DNO absent. Although the species are univoltine. All species appear to associate presence of TOs is unique in this subfamily, they are with Anoplolepis (Formicinae) ants (Claassens & small and do not evert. Larvae feed on Coccidae (Ho­ Dickson 1980, Cottrell 1984, Claassens & Heath moptera) tended by CrenUltogaster ants that do not in­ 1997). During the first three ins tars of T proturnnus teract directly with the larvae. aridus Van Son and T basutus (Wallengren), larvae Spalgini (one genus): Spalgis Moore, 1879 (four feed on Homoptera (Clark & Dickson 1960, 1971, Afrotropical species, one in southern Africa). Larvae Williams & Joannou 1996). In T yildizae Kor;:ak, T pic­ feed on Coccidae (Homoptera), do not associate with tus Van Son and T basutus food of the final two instars the ants directly, but ants are always present tending is regurgitated food from ants (Fig. 21 ). We suspect Homoptera. Larvae cover themselves with a waxy se­ that all Thestor are entomophagous but only in T ba­ cretion produced by the Homoptera and lack both sutus is the life history fully known. The remarkable DNO and TOs (see Lamborn 1914, Jackson 1937). Thestor larva (Fig. 22) lacks both DNO and TOs, and Lachnocnemini (two genera): In these genera, lar­ possesses an extremely small head with an extendable vae lack DNO and TOs. Lachnocnerna Trimen & fleshy "neck". Larval antennae are elongate and pro­ Bowker, 1887 (38 Afrotropical species, 3 in southern ject forward; each with a long terminal seta. In Africa). Feeding habits of L. bibulus Fab. have been T yildizae these antennae remain in contact with the described by South African authors, but by a different ant's mandibles during exchange of food. It is possible account of LachnocnenUt hom Kenya (Cripps & Jack­ these antennae simulate the ant's mandibles, facilitat­ son 1940), larvae were carried by ants down to their ing the larva's acceptance as a nestmate anclJor to stim­ nest, and trophallaxis occured. The genus was revised ulate the ant to regurgitate. by Libert (1996a-c) who described 25 new species, Williams and Joannou (1996) observed females of hence the life history accounts from Kenya most prob­ Thestor basutus capeneri (Dickson) ovipositing on ably apply to what are now considered different blades of grass infested by grass-feeding coccids Pulvi­ species. Accounts are given by Clark and Dickson naria iceryi (Signoret) (Homoptera: Coccidae) which (1971) of L. bibulus and L. durbani Trimen from in turn were tended by Anoplolepis custodiens Smith VOLliME 57, NUMBEH 1 9

Flc. 21. Final instar larva of Thestor yildizae K09ak being fcd re­ gurgttatlOns lrom a "Pugnacious ant" Anoplolepis custodiens F. Smith. ants. They were observed to oviposit on a wide variety of vegetation, often without ants or Homoptera being present. In the wild and in captivity the first three in­ stars were predaceous on coccids, the ants taking no interest in the larvae. After moulting, fourth instar lar­ vae in captivity refused to feed on coccids and were 1 em left in the formicary where they entered the artificial FTC. 22. Final instar lmva of Thestor yildiwe, lateral view (up­ ant nest. While in the ant nest, they were ignored by per), dorsal view with head extended (lower). Scale har = 1 cm. the ants but subsequently died, presumably of starva­ tion (Williams & Joannou 1996). Similar observations The behavior of T basutus differed from the ob­ were recorded hy Clark and Dickson (1960, 1972) for served behavior of T yildizae and T pictus larvae T hasutus basutus. which solicited regurgitations from individual passing In November 2002 four final instar larva of T hasu­ ants. The ant species associated with Thestor have so tus basutus (Wallengren) were taken from an far been recorded as A. cllstodiens but this is currently Anoplolepis custodiens ant nest in KwaZulu-Natal and believed to constitute a species complex. A prelimi­ studied in captivity together with Anoplolepis ants nary molecular study of A. custodiens at Harvard Uni­ from a nearby locality (AH, AJMC, S. P. Quek). A larva versity suggcsts that at least three distinct exist was often seen to approach two ants cngaged in (S.P. Quek unpublished). trophallaxis and insert its head between theirs. It pro­ LYCAENINAE: Some ant-association occurs in all ceeded to imbibe regurgitations passed between them tribes except Lycaenini. Almost all species have a and continued to accept regurgitations from the donor DNO and most have TOs. ant after the other ant had departed. The larva was oc­ Theclini (six genera): The number oflarval instal'S caSionally seen to 'pull' on ant eggs and larvae and drag is normally four and the D NO first appears in the sec­ them beneath its carapace; it is assumed these were ond or third ins tar, the TOs appearing in the second eaten. The larva also appeared to scavenge for detritus instal' except in endophytic feeders, where they are ab­ on the nest substrate. Although T basutus appears to sent in all instars. utilize four food sour(;es in its larval stage, during the Myrina Fahricius, 1807 (five Afrotropical species, week it was studied (prior to pupation) the major food two in southern Africa); lolaus Hubner, [1819] (116 was obtained by trophallaxis. An earlier study in cap­ Afrotropical species, 21 in southern Africa); Hypoly­ tivity of T basutus larvae from KwaZulu-Natal with an caena Felder, 1862 (extralimitaI4 : 28 Afrotropical ant species from Cape Town resulted in the larva feed­ species, five in southern Africa); Leptomyrina Butler, ing on detritus only and then dying after almost four 1898 (nine Afrotropical specie s, four in southern weeks without any trophallaxis having been observed Africa); Capys Hewitson, 1965 (14 Afrotropical (Heath & Claassens 2000). This would indicate larvae species, four in southern Africa); Deudorix Hewitson, are highly specific to certain Anoplolepis ants. 1863 (extralimital: 89 Afrotropical species, nine in The larva occasionally 'groomed' an ant, the latter southern Africa). All six genera are regarded as having remaining motionless while the former may have de­ a facultative association with ants. Ants (often Cre­ rived some form of detritus from the process (AI-I, matogaster species) may be present but do not attend AJMC unpublished obs. ). This grooming behavior was the larvae permanently, and larvae do not depend on also recorded in the case of Lepidochrysops larvae any species of ant for survival. Larvae of the first three (Polyommatini) and its ant-associate Camponotus genera possess DNO and TOs but those of Leptomy- maculatus Fab. (Formicidae) (Claassens 1976). 4 Extralimital: Cenus also represented outside Africa. 10 JOURNAL OF THE L EPIDOPTERISTS' SOCIETY

rina, Capys and Deudorix are endophytic and lack by the host ant. The DNO was present in the second TOs (Clark & Dickson 1971). Leptomyrina larvae feed instar, but secreted no honeydew, however, when inside the fleshy leaves of Crassuleaceae (Jaekson, everted it attracted, but stupified nearby ants. When 1937). Capys larvae feed inside flower buds offered a red Homopteran juvenile, the second instar (Proteaceae) (Murray 1935, Clark & Dickson 1971); larva ate it and half a second one, but died the follow­ the hollowed out buds are later often occupied by ing day. The final instar also fed on ant regurgitations Crematogaster ants (AH pers. obs.). Deudorix larvae living and pupating within the carton nest of the host feed inside pods and immature fruit of a variety of ant Crematogaster peringueyi Emery. It is assumed trees (Pinhey 1965, Clark & Dickson 1971). that the intervening three instars had the same feeding Aphnaeini (14 genera); All known life histories of behavior as first and last instars. In areas where C. the Aphnaeini indicate an obligate ant-association. The dicksoni flies , armored scale insects (Homoptel'a) are number of larval ins tars is normally six, the DNO first always nearby and may somehow be necessmy for the appearing in second or third instars, but TOs occur in survival of the larva (Heath 1998). The DNO was still all ins tars . present in the sixth (final) instal', although it was infre­ Chrysoritis Butler, 1898 (42 species, all endemic to quently visited by ants. The final instar also had spe­ southern Africa). A revision and molecular phylogeny cialized setae on each segment that frequently at­ of Chrysoritis can be found in Heath (2001) and Rand tracted the host ant who seemed to nibble at them. et al. (2000). The genus comprises small to medium­ These setae appeared as a curved structure resembling sized robust butterflies, associated with open veld, a bottle-brush, and are illustrated by Heath (1998). coastal and montane '. Most Chrysoritis are Heath and Brinkman (1997b) inferred that in the wild, multivoltine, except in montane areas, but C. dicksoni females are only stimulated to oviposit by ants attend­ (Gabriel) which flies in non-montane localities, is en­ ing this unidentified species of scale . Colonies of tomophagous and univoltine. With this one exception, C. dicksani adults occupy small areas, seldom larger Chrysoritis are all herbivorous feeding on a wide vari­ than a tennis court, and where females oviposit on a ety of plants (see Heath 1997a) and can be reared in wide variety of plants near the host ant nest. captivity without ants. They are nevertheless consid­ Aloeides Butler, 1819 (58 Afrotropical species, 53 in ered obligate, as larvae are continuously attended by southern Africa). A large genus of small to medium­ ants and oviposition only occurs in the presence of the size brown or orange butterflies obligately associated correct ant species (Heath 1997a). Crematogaster ants with ants. All species are found in open or grassland are the most common associates for Chrysoritis, but habitat. The genus is in need of taxonomic revision, as one species of Myrmicaria (Myrmicinae) is known to we believe it contains many taxa of dubious status. Un­ associate in two species of this genus (Heath 1997a). til recently, the only ant genus known to be associated Excepting C. dicksoni, larvae in nature rest in a corral with Aloeides larvae was Lepisiota (=Acantholepis) (or byre) beneath the food plant and are tended by (Formicinae). However, two additional ant hosts have three or more ants. Larvae were often found singly or recently been discovered (Heath & Claassens 2000)­ in pairs but as many as five or six of varying sizes could Monomorium fridae Forel (Myrmicinae) and PheicZole be found together within a corral (Heath 1997a). capensis Mayr (Myrmicinae). Larval food plants are In Chrysoritis the DNO appears in the second in­ species of (Fabaceae), Hermannia (Stercu­ star, a feature shared only with Crudaria Wallengren. liaceae), SicZa (Malvaceae) (Kroon 1999), and Gnidia The larva's DNO is frequently visited and stimulated (Thyrnelaeaceae) (A. Gardiner pers. corn.). Aloeides by the ant's antennae and the secretion is eagerly taken larvae have TOs in all instars but the DNO first ap­ by the ant. The TOs are active, everted qUickly and as pears in the third. In some species, e.g., A. apicalis quickly withrawn; this happens whenever the larva is Tite & Dickson, A. pallida grandis 'fite & Dickson, disturbed. When ants are over-eager to access honey­ A. thyra (Linnaeus) and A. dentatis (Swierstra), the dew, the larva will evert its TOs causing them to jump DNO is absent in the final instar (Heath & Claassens in alarm. Among this large and otherwise herbivorous 2000, Henning 1983a, b). However, in at least four genus, C. dicksani larvae are entomophagous and were species, A. pierus (Cramer) A. gowani Tite & Dickson, observed in the field and laboratory by Heath (1998). A. trimeni southeyae Tite & Dickson and A. aranda The first two instars fed on ant regurgitations but (Wallengren), larvae retain their DNO until pupation stayed on the plant close to scale insects, also tended (Clark & Dickson 1971, Heath & Claassens 2000). Early instar larvae of A. pallicZa grancZis are assumed ., Fynbos: Characteristic. treeless shrublancl vegetation of the to feed on species of Aspalathus always found close to southern and south-western Cape of South Africa. where the larvae are found. Heath and Claassens (2000) VOLUME 57, NUMBER 1 11 observed that, in captivity final ins tar larvae remained south-western region of South Africa; they are univol­ inside the nest of Mayr for four tine and all are presumed to be aphytophagous (Heath months, and grew without foraging outside. Despite 1997a). A final instar larva and pupa of Trimenia mala­ ample ant brood of all stages present in the nest, the grida maryae (Dickson) were found in small fissures larvae fed only on ant eggs. about 5 cm deep inside the bedrock tended by Despite the DNO of A. thyra being absent in its final Anoplolepis custodiens ants (Heath & Brinkman ins tar, it is herbivorous on Aspalathus species and rests 1995b). In captivity it could not be determined what in an L. capensis ant nest (Claassens & Dickson 1977) the larval food was since the larva shunned any light. but it is not known if trophallaxis or ant eggs form a When disturbed, it was tended by many ants, with a supplementary PaIt of its diet. In contrast, A. apicalis concentration around the head of the larva (AH pers. and A. aranda which associate with Monorrwrium and obs.). There was no vegetation within a meter of the Pheidole ants respectively, were not inside the ant nest site where the larva was found, but after collection it but were generally tended by four or five ants in a cor­ survived among ants without vegetation for two weeks ral just below the soil surface close to the food plant, before pupating, supporting the notion that it is aphy­ often two or three meters from the ant nest (AH, tophagous, at least in the final instar. We suspect it was AJMC unpublished). Ovipositing females of A. maZomo feeding on ant eggs or ant regurgitations. Despite the coalescens THe & Dickson were observed inserting presence of TOs, the final instal' larva had no DNO their abdomens deep into the sand beneath a species of and was very similar to that of Trimenia argymplaga Gnidia (Thymelaeaceae) (A. Gardiner pers. com.). (Pringle in Pringle et al. 1994). A T argymplaga larva Eggs of A. aranda (Wallengren) were found buried ca. in its penultimate instar (presumed) had an active 1 cm in the sand beneath its food plant Aspalathus sp. DNO, but in the final ins tar it was absent. The larva (c. Penz, P. De Vries, AH pers. obs.). was seen to accept ant regurgitations, presumably its Erikssonia Trimen, 1891 (three Afrotropical species, sole diet as it did not feed on ant brood or eggs. The one in southern Africa). E. acraeina Trimen is scarce larva often sought a dark place between the nest and and local, its larval food plant is Gnidia kraussiana arena to rest, and sometimes at night, it would go into Meisner and it has an obligate ant association with the arena but the ants would manoever it back to the Lepisiota sp. (Henning 1984a). The final instar have nest entrance again (Heath & Claassens 2(00). both DNO and TOs. This orange-red butterfly appar­ Argyraspodes Tite & Dickson, 1973 (one species, ently mimics unpalatable species of Fabricius endemic to southern Africa); Zeritis De Boisduval, (Nymphalidae). Erikssonia is closely related to Aloe­ 1836 (extralimital; six Afrotropical species, one in ides and could be synonymized with Aloe'ides based southern Africa). Life histories of these two genera are upon genitalia (Heath 1997a), but other small differ­ unknown, although the former is closely related to Tri­ ences occur in the adult (Henning & Henning 200l). menia and may also be aphytophagous. Eggs are laid among soil particles beneath the food­ Tylopaedia Tite & Dickson, 1973 (one species, en­ plant (Pringle et al. 1994). demic to southern Africa). This large, robust lycaenid Phasis Hubner, 1819 (four species, all endemic to is orange and black and univoltine. T.1f1opaedia sar­ southern Africa). Large brown Iycaenids restricted to donyx peringueyi (Aurivillius) is recorded as using a the southern and south-western Cape; their larval food species of Aspalathus for its lalval food plant (Schlosz plants include Rhus (Anacardiaceae) and Melianthus & Brinkman 1991). The same authors observed the (Melianthaceae) and they have an obligate ant associa­ ant-associate to be Crematogaster melanogaster tion with Crematogaster peringueyi. The DNO on (Emery) and noted that the female would not oviposit Phasis them (Linnnaeus) larvae appears on third and without the presence of ants. subsequent instars (Clark & Dickson 1971) but is ab­ Lipaphnaeus Aurivillius, 1916 (four Afrotropical sent in the final instar (Heath & Claassens 2(00). TOs species, one in southern Africa); Chlomselas Butler, are present in all instars (Clark & Dickson 1971). Lar­ 1885 (=Desmolycaena, Trimen) (13 Afrotropical vae and pupae are found inside hollow stems and al­ species, three in southern Africa); Crudaria Wallen­ though associated with their host ants, interaction has gren, 1875 (three Afrotropical species, all in southern not been studied closely. Africa); Aphnaeus Hubner, 1819 (20 Afrotropical Trimenia Tite & Dickson, 1973 (five species, all en­ species, three in southern Africa); Axiocerses Hubner, demic to southern Africa). Medium-to-large orange 1819 (25 Afrotropical species, five in southern Africa); and brown butterflies with silvery spots on the ventral Cigaritis Donzel, 1847 (= Spindasis Wallengren; surface of all wings. The genitalia are all very similar. Apharitis Riley) (extralimital; 37 Afrotropical species, They are restricted to arid habitats in the southern and ten in southern Africa). The life history of these six 12 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

genera have not been studied in recent years. Their studied the interaction benveen larvae and ants in the larvae all possess DNO and TOs (Clark & Dickson laboratory, and observed that larvae of L. trimeni 1971, Edge 1990), and are all believed to be herbivo­ (Bethune-Baker) and L. rnethymna (Trimen) did not rous and obligately ant-associated, excepting Axio­ possess TOs at any stage, but the DNO appeared in cerses, which are facultative. TOs occur in all instars, the second instar and remained until pupation. the DNO first appears in the third instar, excepting Claassens also observed that trophallaxis supple­ Crudaria where it appears in the second. Clark and mented the diet of ant brood, also that the larva some­ Dickson (1971) described additional saucer-like times groomed an ant. Henning (1983b) studied L. ig­ glands referred to as "dewpatches" on the dorsum of nota (Trimen & Bowker), and made similar feeding the A2-A4 segments of Cigaritis and Crudaria final observations. Henning (1983b) demonstrated that lar­ instar larvae. These glands secrete a fluid the ants vae of L. ignota and chemically consume. These six genera are mostly associated with mimic the brood of their attending ants Carnponotus Crematogaster ants, except for Crudaria which asso­ niveosetosus and Lepisiota capensis. Corn grits were ciate with Anoplolepis custodiens (Heath 1997a). Lar­ soaked in epidermal extracts of ant brood or larvae and vae and pupae of Crudaria wykehami Dickson taken then offered to the appropriate species of ant. Treated from an ant nest beneath a large flat stone were heav­ grits were carried by ants to their brood chamber, un­ ily parasitized (ca. 80%) by wasps and a tachinid fly treated grits were ignored. Gas chromatograms of epi­ (AH unpublished obs. ) dermal extracts confirmed that chemicals found OIl the Lycaenini (one genus): Lycaena Fabricius, 1807 larva were similar to those on the ant brood. In captiv­ (extralimital; three Afrotropical species, two in south­ ity, a fourth ins tar larva of L. p. plebeia (Butler) was ern Africa). Small coppery-red butterflies. Larvae observed feeding on ant eggs as soon as they were laid, naked, lacking DNO and TOs, and not ant-associated. when ant brood supply was exhausted (Williams 1990). Polyommatini (2.5 genera): Normally having four Orachrysops Vari, 1986 (11 species, all endemic to larval instars, the DNO appears in the second or third southern Africa). Formerly included in Lepi­ instar, the presence of TOs variable. dochrysops, these medium-sized dull blue and gray Euchrysops Butler, 1900 (five in southern Africa). lycaenids are superficially similar to Lepidochrysops, Medium-sized blue or brown lycaenids. The phy­ but their genitalia differ. Little is known about ant­ tophagous larvae possess DNO and TOs, the former association in their early stages. Formerly thought to appearing in the second instar, the latter in the third be phytopredacious, as in Lepidochrysops (Henning and have a facultative ant-association (Clark & Dick­ & Henning 1994) but recent work suggests otherwise son 1971). The genitalia are of the same type as Lepi­ (Edge & Pringle 1996). The larvae of O. niobe (Tri­ dochrysops from which Euchrysops are not easily sep­ men) feed on Indigofera (Fabaceae), and in captivity, arated on morphological grounds (see Gardiner 1998). were bred to adult without ant presence. However it Lepidochrysops Hedicke, 1923 (127 Afrotropical is unknown if ant-association is obligate or facultative species, 59 in southern Africa). Medium to large, blue under natural conditions and whether other sources or brown with spotted undersides frequenting open supplement its diet. The DNO and TOs appear in the and sparsely wooded grassland. Morphologically simi­ second and subsequent instars, the TOs are not well lar to Euchrysops with remarkably uniform genitalia. developed. The ant suspected of being an associate is All are assumed to be phytopredacious (herbivorous in Camponotus niveosetosus (Mayr) (Edge & Pringle early ins tars, later becoming predacious on other in­ 1996), recorded among the roots of the food plant sects), associated with Campunotus ants, and in many where the larvae shelter. Recently, Lu and Samways respects, similar in appearance and behavior to the (2001) showed that O. ariadne (Butler) has an oblig­ palaearctic Maculinea (see Frohawk 1916, Elmes et al. ate ant-association. Larvae were found in soil beneath 1991, Thomas 1983, 1995). The larva feeds on flower the foodplant at depths up to 10 cm, always attended buds for the first two instars, then it is carried by a by C. natalensis (Smith). species of Carnponotus ant to its brood chamber in the Anthene Doubleday (25 species in southern Africa), third instar where it is predacious on ant brood. De­ Eliot (one species in southern Africa), Tuxentius spite the large size of this genus, few life histories have Larsen (three species in southern Africa), Scud­ been studied in any depth. Our current knowledge is der (five species in southern Africa), Larnpid.es Hubner based upon the life history of eleven species by Clark (one species in southern Africa), Tarucus Moore (three and Dickson (1971), Claassens (1972, 1974, 1976), species in southem Africa), Harpendyreus Heron (three Henning (1983a, b) and Williams (1990), some being species in southern Africa), Pseudonacaduba Stempffer incomplete. However, Claassens (1972, 1974, 1976) (one species in southem Africa), Eicochrysops Bethune- VOLUME 57, NUMBEH 1 13

Baker (three species in southern Africa), Cupidopsis TABLE 3. Lycaenids obligately associated with ant genera. Karsch (two species in southern Africa), Thermoniphas Karsch (one species in southern Africa), Oboronia Number of obligately ant-associated Iycaenids (%) Karsch (one species in southern Africa), Actizera Chap­ man (two species in southern Africa), Chapman Confirmed Confirmed Zizeeria Ant genus only and predicted (one species in southern Africa), Zizina Chapman (one Myrmicinae species in southern Africa), Brephidium Scudder (one Pheidole 1 (1 ) 1 (0 ..5) species in southern Africa), Oraidium Bethune-Baker lvlonoJrUJrium 1 (1) 1 (0.5) (one species endemic to southern Africa), Azanus Moore C rematogaster .51 (61 ) 62 (28 ) Myrmicaria 2 (2) 2 (1) (five species in southern Africa), Chilades Moore (one Formicinae species in southern Africa), Zizula Chapman (one Anoplolepis 13 (15) 37 (17) species in southern Africa). All 20 of these genera (71 Lepisiota 7 (9) 7 (3) Camponotus 7 (9) 60 (28) species) are facultatively ant-associated, most having Unidentified 2 (2) 47 (22) DNO and TOs but often without attendant ants (see Totals 84 217 Clark & Dickson 1971). Uranothauma Butler (four species in southern It can be seen from Table 1 that algae or lichen Africa), Cacyreus Butler (five species in southern feeders are unique to the two tribes of Poritiinae. The Africa). Not ant-associated, TOs and DNO are absent, Miletinae feed mostly on Homoptera but also on ho­ except for two species of Cacyreus which have a DNO. mopteran secretions, ant regurgitations and possibly Vibrational communication. Using a cardboard detritus and ant early stages at times. Among the Ly­ poster tube with a paper membrane at one end, final caeninae, almost all the entomophagous records are instars of Chrysoritis thysbe (Linnaeus), C. dicksoni, from the Polyommatiti, with only two confirmed cases Aloeides pierus and A. pallida grandis were heard to from the Afrotropical Aphnaeini, although there may produce ticking or drumming sounds, but C. thysbe well be more still undiscovered7. also made an intermittent high-pitched buzzing sound Symbiont ant genera. The two main ant genera (Heath 1998). Sounds have also been noted in the pu­ confirmed in obligate association with lycaenids are pae of Chrysoritis brooksi Riley and P irene Penning­ Crematogaster (Myrmicinae) and Anoploplepis ton (Schlosz 1991). These sounds likely play an addi­ (Formicinae), together 76%, whereas Camponotus tional role in communication with ants (De Vries 1990, and Lepisiota (Formicinae), represent 9% each (Table 1991). 3). Three accounts of ant distribution (Samways 1983, Ant-associations and feeding strategies. More Donnelly & Giliomee 1985, H. Robertson pers. com.) than three quarters of the 392 southern African indicate the two dominant ant genera in Southern lycaenids are ant-associated, over two thirds of these Africa are Pheidole (Myrmicinae) ~nd Anoplolepis, the have obligate relationships. The Poritiinae constituting latter more so in open habitats. Crematogaster species 8% of lycaenids (30 species) have no direct ant­ are also numerous in denser vegetation while Lepi­ association° and the Miletinae (36 species) represent­ siota and Camponotus although not always numerous, ing 9%, three quarters of which are obligately ant­ are to be found in most habitat types in southern associated although the remainder feed in the Africa (H. Robertson pers. com.). Despite the com­ presence of ants. The subfamily Lycaeninae (326 parative dominance of Pheidole ants, only one species species) accounts for 83% of species, of which the has been identified as an obligate symbiont (to a wide­ Aphnaeini (131 species) contains a third of all southern spread species of Aloeides).

African lycaenids, almost all being obligate. Theclini DISCUSSION (47 species) and Polyomrnatini (146 species) account for 12% and 37% respectively; the former being fac­ Reliability of data. Considerable disparity exists in ultative, and latter split between facultative and oblig­ the Polyommatini between confirmed and predicted ate. The Lycaenini has only two spccics, neither ant­ ant ~ association (Table 2) mainly because Lepi­ associated. do chrysops (Polyommatiti) life histories are widely be­ lieved to be uniform, but few have been confirmed.

01n Deloneurrt and certain other Liptenini further north in Africa e.g., , larvae are /(mnd in the company of ants but no interac­ 7 Cottrell (1984), Elmes et aJ. (2001) list Cigaritis takanonis Mat­ tion has been observed. It is possible that ant-derived detritus sup­ sumura from Japan as accepting ant regurgitations. Jackson (1937) plements the algae on which the larvae feed, or it may enrich the al­ infers insect feces as additional f,)od for ChZoroselfls pseudozeritis gae, making it more attractive, but these hypotheses have yet to be (Trimen). Fiedler (1991) infers possible ant regurgitations for several confirmed. Aphnaeini. 14 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

Obligately ant-associated Afi-otropicallycaenid species ACKNOWLEDGMENTS show little morphological differences among closely We thank Adrian Armstrong and the Natal Parks Board (South related species (Heath 1997b), and recently Heath Africa) and the Chief Executive Officer of the Western Cape Nature (2001) synonymized many Chrysoritis taxa, reducing Conservation Board for permission to study and collect material in areas under their control. Hamish Robertson, South African Mu­ the species by 28%, inferring that similar syn­ seum, Cape Town is acknowledged for identifying ant species pro­ onymies probably exist in Aloeicles, and we believe vided by the authors and for giving related input on this paper. Ivan Bampton and Colin Congdon are acknowledged for some recent life this is the case for Lepiclochrysops. We suspect that history data. Simon Joubert and Swee Peck Quek are thanked for southern Africa has an oversplit among the helping to locate the T hasutus larvae and ant associates in ant-associated lycaenids, particularly those with an ob­ KwaZulu-Natal. Phil DeVries is espeCially thanked for his many ligate relationship. helpful comments on the manuscript. Changes in feeding strategy. Although most lep­ LITERATURE CITED idopterous larvae are herbivores, some lycaenid ACKERY, P. R., R. DE JONG & R. I. VANE-WRIGHT. 1999. The but­ species are known to switch from one trophic strategy terflies: Hedyloiclea, Hesperioiclea and . In Kris­ to another midway through their larval phase; e.g., the tensen, N.r. (ed.), Handbook of Zoology 4 (35):263-300. Berlin, de Gruyter. Maculinea (Thomas 1983, 1995). In southern Africa ACKERY, P. R&D. RAJAN. 1990. A manuscript host-list of the this occurs in Lepiclochrysops, Thestor, and at least Afrotropical butterflies. Unpub!. list. Nat. Hist. Mus. London. one Aloeicles species (Cottrell 1984, Heath & ACKERY, P. R , C. R. SMITH & RI. VANE-WRIGHT. 1995. Carcasson's African Butterflies: an annotated catalogue of the Papilionoidea Claassens 2000). More than one food source can also and Hesperioidea of the Afrotropical region. CS IRO, Australia. be explOited at the same time e.g., trophallaxis and car­ ATSATT, P. R 1981. Lycaenid butterflies and ants: selection for en­ nivory in Cigaritis (Sanetra & Fiedler 1996), and in cmy-free space. Amer. Nal. 118(5):638-654. BAMPTON, I. 1995. A discussion on the larval food of the suhfamily some Maculinea (Thomas & Wardlaw 1992), and in Lipteninae (Lepidoptera: Lycaenidae). MetamorphoSiS Lepidochrysops (Claassens 1976, Henning 1983) and 6:162- 166. Thestor (AH, AJMC) in southern Africa. BRABY, M. F. 2000. Butterflies of Australia: their identification, bi­ ology and distribution. Vols. 1 & 2. CSIRO Publishing, Aus­ Dorsal nectary organ and tentacle organs. The tralia. 976 pp. DNO and TOs are completely absent in the Poritiinae BRABY, M. F. & T. A. WOODGER. 1994. The life history ofZizula hy­ and Milctinae except in Aslauga (Liphyrini) but one or lax attenuata (T. P. Lucus) (Lepidoptera: Lycaeniclae). Aus­ tralian Entomologist 21:39-42. both are usually present among the Lycaeninae. In CLAASSENS, A. J. M. 1972. Lepidochrysops trimeni (Bethune­ some Aphnaeini (Phasis, some Aloeicles) the DNO is Baker) (Lepidoptera: Lycaenidae) reared from larvae found in present in earlier ins tars but absent in the final instar. the nest of Camponatus maculatus Fab. (: Formi­ cidae). J. ent. Soc. sth. Afr. 35:359-360. Second instar Chrysoritis dicksoni had a DNO that did ---. 1974. Methods employed in the successful rearing, nnder not secrete honeydew but rather it gave off a artificial conditions, of two Cape species of Lepidachrysaps pheromone, causing the ants to remain close and (Lepidoptera: Lycaenidae). J. ent. Soc. sth. Afr. 37:387-392. ---. 1976. Observations on the myrmecophilous relationships stupified (Heath & Brinkmann 1995a). Although the and the parasites of Lepidochrysaps methymna methymna (Tri­ TOs are absent in all instars of Lepiclochrysops, the men) and L. trirneni (Bethune-Baker) (Lepidoptera: Ly­ DNO appears in the second instar (Clark & Dickson caenidae). J. ent. Soc. sth. Afr. 39:279-289. - - - . 1991. The mystery of the squeaking pupae of myrme­ 1971) and is retained until pupation, as in Palaearctic cophilous Lycaenidae. Metamorphosis 2:19-20. Maculinea. Fiedlcr (1998) suggested DNO secretions CLAASSENS, A. J. M. & c. G. c. DICKSON. 1977. A study of the supplement the adoption procedure. However, Hen­ myrmeeophilous behaviour of the immature stages of Aloeides thyra (L.) (Lep.: Lycaenidae) with speCial reference to the func­ ning's (1983b) experiments using corn cob grits do not tion of the retractile tubercles and with additional notes on the support this for Lepidochrysops. biology of the species. Entomologist's Ree. J. Var. 89:225-231. Fiedler (1998) attributed the lack of TOs in Macu­ - --. 1980. Butterflies of the Table Mountain range. Struik, linea to the endophytic life-habit of the early larval in­ Cape Town. 160 pp. CLAASS ENS , A. J. M. & A. HEATH. 1997. Notes on the myrme­ stars rather than its life-habit within the ant nest, but cophilous early stages of two species of Thestor Hubner (Lepi­ in Afrotropical Polyommatini (including Euchrysops) doptera: Lycaenidae) from South Africa. Metamorphosis 8:56-61. the TOs generally appear in the third or fourth instar CLARK, G. C. & C. G. C. DICKSON . 1956. The honey gland and tu­ bercles of the Lycaenidae. Lepid. News 10:225-2.31. (Clark & Dickson 1971). Lepidochrysops larvae cease ---. 1960. The life histories of two species of Thestor Hbn. to be herhivorous after the second instar, and hence (Lepidoptera: Lycaenidae). J. ent. Soc. sth. Afr. 2:3:278-281. the TOs would not have been present during any early ---. 1971. Life histories of the South African lycaenid butter­ flies. Purnell , Cape Town. 272 pp. instar. An al ternative hypothesis for Lepiclochrysops is CONGDON. T. C. E. & 1. BAMPTON. 1995. Ants and lycaenid butter­ that within the nest, the TOs became redundant as a flies- some observations. Lambilliollea XCV 4:572-574. means of recmiting ants and may also have been a hin­ CORHET, A. S. , H. M. PE!\IDLEIlURY & J. N. ELIOT. 1992. The but­ terflies of the Malay Peninsula. Malayan Nature SOCiety, Kuala derance within the confined space. Lampur. VOLUME 57, NUMBER 1 15

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