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The taxonomy, biogeography and conservation of the myrmecophilous Chrysoritis (: ) in

R.F. TERBLANCHE and H. VAN HAMBURG

Terblanche, R.F. and H. Van Hamburg. 2003. The taxonomy, biogeography and conser- vation of the myrmecophilous Chrysoritis butterflies (Lepidoptera: Lycaenidae) in South Africa. Koedoe 46(2): 65–81. Pretoria. ISSN 0075-6458. The relevance and integration of scientific knowledge to conservation management of the locally popular and highly endemic Chrysoritis are investigated within the research fields of taxonomy and biogeography. The butterfly genus Chrysori- tis contains at least 41 species endemic to South Africa. The taxonomy of Chrysoritis has reached a state where revisions could easily result in a plethora of names between “lumping and splitting”. In practice, the state of the taxonomy of these butterflies on species level may alter their conservation priority. The two most species rich species groups in Chrysoritis have different centres of endemism, however, a butterfly atlas becomes a necessity to reveal more about their biogeography. There is an absence of butterfly species lists in many of our National Parks and Nature Reserves. Legislation should facilitate rather than limit the valuable role of the amateur lepidopterist to add distribution records. In turn, the amateur lepidopterists should adapt and make an effort to explore unknown localities, apart from monitoring butterflies at their well-known localities. The red listing of localised butterflies in South Africa, including a number of Chrysoritis species, is in need of an urgent review in the light of the most recent IUCN categories. A species such as Chrysoritis dicksoni should be protected by law – but at its known localities. The scenario that real conservation action is only needed if the last known locality of a butterfly is threatened, should be abolished. A paradigm shift to con- serve the metapopulations of the highly endemic Chrysoritis genus and not merely a few of its species as items that appear on lists, seems necessary. Key words: Chrysoritis, myrmecophilous, endemic, conservation, research priorities, environmental management, South Africa, taxonomy, biogeography, red list

R.F. Terblanche and H. Van Hamburg, School of Environmental Sciences and Develop- ment, Private Bag X6001, Potchefstroom University for CHE, Potchefstroom, 2520 South Africa.

Introduction Chrysoritis seem to be myrmecophilous (Heath 1997a; 2001). Myrmecophilous but- The purpose of this paper is to consider the terflies pose special challenges for conserva- conservation of the genus Chrysoritis as a tion management due to their intricate habi- real example of conservation in South tat requirements and populations are often Africa. Interactions between disciplines, localised. such as taxonomy and legislation, as well as the role of various interested parties such as A summary of the distribution of the the amateur Lepidopterists and professionals, Chrysoritis species in Lesotho, Swaziland have been poorly investigated to date. The and the provinces of South Africa is present- main objective of this overview is to identify ed for the first time (Table 1). Biogeographic research needs and to create a framework notes with regard to Chrysoritis is given along which research priorities could be from a regional perspective on conservation directed towards sustainable environmental priorities and will hopefully stimulate further management with a view to butterfly conser- research on the distribution of these butter- vation. All the members of the genus flies.

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Table 1 The distribution of Chrysoritis species in the provinces of South Africa as well as the Kingdom of Lesotho and Swaziland. A number 1 indicates that the Chrysoritis species occurs in that area. The major regional literature sources used were Clark & Dickson (1971), Terblanche (1991), Keller, Amodio & Stänz (1997), Kloppers & Van Son (1978), Pringle, Henning & Ball (1994), Duke, Saunders & Saunders (1999), and Claassens (2000). Chrysoritis Species Lesotho, Swaziland and Provinces of South Africa EC FS GP KZ LE LI MP NC NW WC SW Chrysaor Species Group Chrysoritis aethon (Trimen & Bowker, 1887) 1 1 1 Chrysoritis aureus (Van Son, 1966) 1 1 Chrysoritis chrysaor (Trimen, 1864) 1 1 1 1 1 1 1 Chrysoritis lycegenes (Trimen, 1874) 1 1 1 Chrysoritis lyncurium (Trimen, 1868) 1 1 Chrysoritis midas (Pennington, 1962) 1 Chrysoritis natalensis (Van Son, 1966) 1 1 Chrysoritis phosphor (Trimen, 1866) 1 1 1 Chrysantas Species Group Chrysoritis chrysantas (Trimen, 1868) 1 1 Oreas Species Group Chrysoritis dicksoni (Gabriel, 1946) 1 Chrysoritis oreas (Trimen, 1891) 1 Zeuxo Species Group Chrysoritis zeuxo (Linnaeus, 1764) 1 1 Chrysoritis zonarius (Riley, 1938) 1 1 Pyroeis Species Group Chrysoritis felthami (Trimen, 1904) 1 1 Chrysoritis pyroeis (Trimen, 1864) 1 1 1 Thysbe Species Group Chrysoritis adonis (Pennington, 1962) 1 Chrysoritis aridus (Pennington, 1953) 1 1 Chrysoritis azurius (Swanepoel, 1975) 1 Chrysoritis beaufortius (Dickson, 1966) 1 Chrysoritis beulah (Quickelberge, 1966) 1 Chrysoritis blencathrae (Heath and Ball, 1992) 1 Chrysoritis braueri (Pennington, 1967) 1 Chrysoritis brooksi (Riley, 1938) 1 Chrysoritis daphne (Dickson, 1975) 1 Chrysoritis endymion (Pennington, 1962) 1 Chrysoritis irene (Pennington, 1968) 1 Chrysoritis nigricans (Aurivillius, 1924) 1 Chrysoritis orientalis (Swanepoel, 1976) 1 Chrysoritis palmus subsp. palmus (Stoll, 1781) 1 1 Chrysoritis pan (Pennington, 1962) 1 1 1 Chrysoritis pelion (Pennington, 1953) 1 1 1 1 Chrysoritis penningtoni (Riley, 1938) 1 Chrysoritis perseus (Henning, 1977) 1 Chrysoritis plutus (Pennington, 1976) 1 1 Chrysoritis pyramus (Pennington, 1953) 1 Chrysoritis rileyi (Dickson, 1966) 1 Chrysoritis swanepoeli (Dickson, 1965) 1 Chrysoritis thysbe (Linnaeus, 1764) 1 1 Chrysoritis trimeni (Riley, 1938) 1 Chrysoritis turneri (Riley, 1938) 1 1 Chrysoritis uranus (Pennington, 1962) 1 Chrysoritis violescens (Dickson, 1971) 1 EC FS GP KZ LE LI MP NC NW WC SW TOTAL 13 3 1 9 2 0 5 14 0 25 1

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Discussion high. Although the latest synonymies pre- sented by Heath (2001) have been followed in this paper and in the most recent checklist Taxonomy of the genus Chrysoritis of Southern African butterflies (Vári et al. Most of the present Chrysoritis species were 2002), there is currently much debate among previously known as the Poecilmitis species. Lepidopterists of Africa whether all the syn- Based on wing pattern, genitalia features and onymies suggested by Heath (2001) should larval morphology, Heath (1997a) treated the be accepted. A few factors that could have genera Chrysoritis, Poecilmitis, and Oxychaeta as synonyms of Chrysoritis. His treatment with regard to Chrysoritis, Poecilmitis and Oxychaeta was supported by the phylogeny inferred from mitochondrial cytochrome oxidase I sequences (Rand et al. 2000). Rand et al. (2000) could not ascertain the phylogenetic position of Bowkeria due to a lack of fresh specimens, being a rare canopy dweller and difficult to obtain. The inclusion of Bowkeria in Chrysoritis needs further study, especially since its life history remains unknown. The studies of Heath (1997a) that led to the inclusion of Chrysori- tis, Oxychaeta and Poecilmitis being syn- onyms of Chrysoritis, as well as Rand et al. (2000) expanded the understanding of the genus Chrysoritis considerably, including the ecological functional similarities among the species in this group. For instance, apart from Chrysoritis dicksoni (previously known as Oxychaeta dicksoni) that might have a derived aphytophagy, the rest of the Chrysoritis species studied to date are phy- tophagous with a mutualism that includes myrmicine ants of the genera Myrmicaria and Crematogaster. Figure 1 illustrates some members of the species groups of Chrysoritis. Heath (2001) is followed for the species groups within the Chrysoritis genus, which in turn is largely based on Heath (1997a) and Rand et al. (2000) (see Table 1 for the species and species groups). Heath (2001) instated a number of synonymies within the genus Chrysoritis, which reduced the total number of species from 59 to 42. It was suggested that there may be a number of synonymies among the remaining 42 species as well. Even if the number of species was to be low- ered further, the number of endemic species Fig. 1. Examples of Chrysoritis species. Top: of this genus could still be regarded as very C. aethon; Middle: C. pelion; Bottom: C. zeuxo.

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led to the confusion in Lepidopterists’ circles reader could not confirm or repeat the judge- are briefly discussed below. ments. One example, Chrysoritis lycia, has Heath (2001) stated that his conclusions rely been synonimised with Chrysoritis chrysaor. in part on the paper by Rand et al. (2000) and Based on breeding experiments, of which no a revision of Aphnaeini (Heath 1997a) but further details apart from the host plants and did not specify to what extent for each of the localities are presented, the conclusion is synonymies. Rand et al. (2000) investigated drawn that Chrysoritis lycia could be consid- mitochondrial DNA sequences of only 19 ered an extreme and more yellowish form of species of the former 59 species of Chrysori- Chrysoritis chrysaor. However, important tis species. Although this investigation was other morphological characteristics used to to a great extent sufficient to clarify and con- distinguish Chrysoritis lycia from Chrysori- firm the species groups and other phyloge- tis chrysaor, such as the series of markings netic relationships within the genus on the hind wing underside margins are not Chrysoritis, not enough information was discussed. The Chrysoritis lycia material has gained to base synonymies of species on, also been bred from material at with the possible exception of the Chrysori- Leipoldtville, which is a few hundred kilo- tis zeuxo species group which is the only metres from the type locality near Matjes- group of Chrysoritis of which some DNA fontein. Even if the synonymies and Heaths’ information exists for all the species in the judgements based on his reference collection group. It seems that the synonymies present- were correct in these cases, the above lack of ed by Heath (2001) of most of the Chrysori- details, illustrations as well as more specific tis species are still largely based on morpho- details about the material and methods could logical and life history information and not have led to the present confusion. It is to be the DNA study by Rand et al. (2000). questioned if sufficient information is avail- Heath (1997a, 1997b) showed that the mor- able to synonimise many of the Chrysoritis phology of Chrysoritis genitalia, even species at present. A similar analogy exists among the majority of the species in the in the plant genus Haworthia that also con- genus, is very uniform in structure, and that tains variations between and within popula- this might be related to their myrmecophily. tions and which is also of considerable ama- Therefore, the separation of Chrysoritis teur and collectors’ interest (see Terblanche species relies on other morphological char- et al. 1993; Bayer 1999). acteristics. Unfortunately, other morphologi- Heath (2001) suggested that other syn- cal details and life history information that onymies might be extant among more were presented to sink species in the Heath Chrysoritis species, especially within the paper (2001), were not presented or illustrat- ed in any specific detail for any of the syn- Chrysoritis chrysaor species group. This fol- onymies. Reference to characterisics speci- lows the suggestion by Owen-Johnston fied in the original descriptions as well as the (1991) that Chrysoritis lycegenes and type series of all the entities that were syn- Chrysoritis lyncurium and an unidentified onomised, were lacking at large. Heath Chrysoritis entity that occurs at Morgenzon (2001) states that rearing Chrysoritis over in southern , might be a cline several years from egg to adult from cater- belonging to the same species (Owen-John- pillars collected in the field from various ston 1991). In addtion Heath (2001) suggest- localities, has demonstrated the extent to ed that the relation between Chrysoritis which the environment could, during the lar- aethon and Chrysoritis aureus should be val stage, affect adult wing shape, size and investigated. The habitats of Chrysoritis lyn- the markings of both wing surfaces. No ref- curium seem to be in great danger, especial- erence to controlled experiments in a climat- ly from alien invasion (Woodhall 1996). This ic breeding room, were provided. Therefore, may butterfly escape as a conservation prior- in the comparisons of the life histories, the ity due to taxonomic uncertainties.

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Table 2 A summary of the red data book status of Chrysoritis species in the past. Literature sources: S.F. & G.A. Henning (1989) and G.A & S.F. Henning (1995). The taxonomic status and taxonomic research recommen- dations according to Heath (2001) are given in the last column. Species Group and Species Red Data Status Proposed Taxonomic Status 1989 1995 Taxonomic status (Heath, 2001) Chrysaor species group Chrysoritis aureus (Trimen, 1864) R R Relation with C. aethon needs research Chrysoritis lyncurium (Trimen, 1868) R R Relation with C. lycegenes needs research Chrysoritis phosphor subsp. borealis R R Status quo (Quickelberge, 1972) Oreas Species Group Chrysoritis dicksoni (Gabriel, 1946) V E Status quo Chrysoritis oreas (Trimen, 1891) R R Status quo Zeuxo Species Group Chrysoritis cottrelli (Dickson, 1975) E R Regarded as a synonym of C. zeuxo Pyroeis Species Group Chrysoritis pyroeis subsp. hersaleki R R Status quo (Dickson, 1970) Thysbe Species Group Chrysoritis adonis (Pennington, 1962) R I Status quo Chrysoritis azurius (Swanepoel, 1975) R I Status quo Chrysoritis balli R I Synomym (Dickson & G.A. Henning, 1980) Chrysoritis brooksi subsp. taerei R I Status quo (Dickson, 1966) Chrysoritis daphne (Dickson, 1975) R I Relation with C. swanepoeli needs research Chrysoritis endymion (Pennington, 1962) R I Status quo Chrysoritis irene (Pennington, 1968) R R Status quo Chrysoritis henningi (Bampton, 1981) R R Regarded as a synonym of C. pan Chrysoritis hyperion(Dickson, 1975) R I Regarded as a synonym of C. swanepoeli Chrysoritis kaplani (S.F. Henning, 1980) R R Regarded as a synonym of C. beaufortius Chrysoritis lyndseyae (S.F. Henning, 1979) I I Regarded as synonym of C. thysbe subsp. bamptoni Chrysoritis mithras (Pringle, 1995) – R Regarded a subsp. of C. thysbe by Heath (2001) Chrysoritis nigricans subsp. nigricans I I Status quo (Aurivillius, 1924) Chrysoritis nigricans subsp. zwartbergae I I Status quo (Dickson, 1982) Chrysoritis orientalis (Swanepoel, 1976) I I Relation with C. pelion needs research Chrysoritis pan (Pennington, 1962) I I Contains several new synonymies Chrysoritis penningtoni (Riley, 1938) R I Status quo Chrysoritis pyramus (Pennington, 1953) R I Contains C. balli as a new synonymy. Relation with C. thysbe needs research Chrysoritis rileyi (Dickson, 1966) R I Status quo Chrysoritis stepheni (Dickson, 1978 R I Regarded as synonym of C. beaufortius Chrysoritis swanepoeli (Dickson, 1965) R I Contains C. hyperion as a new synonymy. Relation with C. daphne needs research Chrysoritis trimeni (Riley, 1938) I I Relation with C. pan needs research Chrysoritis wykehami (Dickson, 1980) R I Regarded as a synonym of C. turneri by Heath (2001) R =Rare ; V = Vulnerable; E = Endangered; I = Indeterminate

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The taxonomic uncertainties regarding the are encountered. Such localities include Chrysoritis species are of much more than Lambert’s Bay, Hondeklip Bay, and Port academic importance, since the conservation Nolloth along the western coast in the of unique entities are at stake. For instance, Northern Cape. These localised populations most often a species has a higher priority in are indicative of unique sets of habitat practice as a conservation priority than a requirements. A small colony of a wide- subspecies or a form that is part of a “cline”. spread Chrysoritis that deserves conserva- Furthermore, species and subspecies appear tion is the colony of Chrysoritis thysbe at on the protected or red lists and not unique Hout Bay in the Cape Peninsula (Claassens habitats. More specifically, it means that the 1994). For insect conservation, the impact of conservation priority for entities such as exotic and invasive flora is of particular con- Chrysoritis hyperion, Chrysoritis lyndseyae, cern in such rare and restricted habitat types Chrysoritis kaplani and Chrysoritis stepheni, (McGeoch 2002). The butterflies of which were all regarded as rare or intermedi- Chrysoritis with their either metallic orange ate in the red data book of Henning & Hen- or metallic orange and blue upper sides of ning (1989), could change considerably (see wings could all be regarded as an aesthetic Table 2). To name one more example: asset and much could be lost before science Chrysoritis cottrelli was previously regarded unravels the full significance of their phy- as endangered (1989) but may dissapear logeny. from the revised list (Table 2). These species are now all regarded as synonyms of other species according to Heath (2001). A taxo- Distribution of Chrysoritis species nomic impediment could lead directly to a Obviously, a better knowledge of the distrib- conservation impediment (New 1997). ution of Chrysoritis species will enhance The above illustrates the importance of tax- research (taxonomy and ecology) and the onomy and its implications for conservation applications of environmental impact assess- of these localised butterfly species. Howev- ments and management. Despite many short- er, it is perhaps the perception of conserva- comings in the study of the distribution of tion of rare butterflies that must change. The butterflies, proportionally far more informa- genus Chrysoritis in its entirety is highly tion is available compared to many other endemic and contains a large number of pop- groups of in Africa. At least one ulations and subpopulations of which most knows to some extent that a butterfly is rare are very localised (even for some of the more or not—this cannot apply to many other widespread species). One is also struck by of which the information on distribu- the variability within and among the local tion and the manpower to obtain these are (sub) populations of Chrysoritis species. very poor. Furthermore, relatively few but- With regard to wing pattern and wing mor- terflies await discovery in South Africa, phology, some of them are seemingly on which is not true for many other insect their way to become species, and other groups. Despite the possibilities that such a forms/entities perhaps becoming less plenti- well-known group presents in terms of dis- ful – these life processes should be con- tribution, it was is not yet been applied suffi- served. It is especially the mixture of ciently in the African context. A butterfly Chrysoritis thysbe forms along the western atlas is becoming an increasing need to inter- and southern coast of South Africa as well as pret the distribution of South African butter- the adjacent inland areas, that reflects enti- flies (Terblanche & Taylor 2000). ties in a stage of speciation difficult to esti- mate. Sympatry and allopatry have been Table 1 gives a summary of the distribution poorly resolved in Chrysoritis, especially for of all the Chrysoritis species in all the new example along the western coast and the provinces (since 1994) of South Africa as Swartberg. It is often difficult to know well as other African countries where these whether different populations or variations butterflies have been recorded. Political

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boundaries were used due to the lack of data by the Chrysoritis species. There is also a in an atlas format. These political boundaries high number of local varieties of subpopula- are used for practical reasons since the distri- tions of Chrysoritis species in the sandy bution in standard reference works on butter- (dune) areas of the Western Cape. The extent flies are commonly described according to to which the entities are sympatric and provinces (Cotrell 1985). These political allopatric has not been clarified sufficiently. boundaries are still useful, since it is known More distribution records will add to the tax- for example that much of the Fynbos biome onomic and ecological insights. Some gaps is confined to the Western Cape as well as in the distribution of species might be real smaller parts of the Eastern and Northern rather than artificial, warranting subspecific Cape and that the most of the con- status of a number of “clines”. sists of grassland (see Rutherford & Westfall 1994). Furthermore, the list is now available No Chrysoritis species have yet been report- to conservation authorities in the different ed from the Highlands of Zimbabwe or fur- regions. Much of the knowledge on the dis- ther northwards in Africa. This contrasts tribution of Chrysoritis species has been with species of other ant-associated lycaenid accumulated by many amateur lepidopter- genera with high concentrations of endemic ists. species in South Africa, especially the West- ern Cape (Fynbos). Such genera include The genus Chrysoritis is almost entirely , Lepidochrysops and even Thestor. endemic to South Africa, Lesotho and The latter has one recorded species from Swaziland (Table 1). The only Chrysoritis parts of Zimbabwe. A few Aloeides species species collected outside the borders of and many more Lepidochrysops species are South Africa, Lesotho or Swaziland, is also found in Equatorial Africa. These distri- Chrysoritis chrysantas. This was found in butions of ant-associated genera are in con- the southern parts of Namibia, by Plowes on trast with the genus Chrysoritis and highlight the Tiras Mountains (Pringle et al. 1994). the high level of endemism of the genus. The genera Chrysoritis and Poecilmitis, The distribution of the two most species rich which are today only known as one genus, groups, the Chrysoritis chrysaor species namely Chrysoritis, had been described as group and the Chrysoritis thysbe species Cape Province centred (an area now known group is discussed in more detail to identify as the Eastern Cape, Western Cape and further research priorities of the biogeogra- Northern Cape) by Cottrell (1985). It falls phy of the genus. into the category of species mainly confined to the old Cape Province (although some All the species within the Chrysoritis have been to a limited extent recorded out- chrysaor species group (which include side this area. The work of Cottrell (1985) Chrysoritis aureus) lack the metallic blue preceeded the synonymies suggested by colour on the upper side of the wings. Heath (2001) and the new provinces in South Instead, they only have an orange upper side Africa so that the levels of endemism are with black markings and a metallic sheen on briefly reviewed here. From Table 1, con- the orange background on the upper side of taining the species list according to the revi- the wings—from which the vernacular/triv- sion by Heath (2001), it can be seen that the ial name “copper” originated (Fig. 1). The highest number of Chrysoritis species occurs metallic blue (opal) colouring with orange in the Western Cape, namely 25 (59.5% of and black markings are most prominent in the total), followed by the 14 species from most members of the Chrysoritis thysbe the Northern Cape (33%) and 13 species of group (Fig. 1). This group is largely confined the Eastern Cape (31%). This echoes Cottrell to the Fynbos biome, with a few exceptions. (1985) that Chrysoritis is Cape-centred. The Simplified, the distribution of Chrysoritis richest area of rare endemic Lycaenidae is species follows a pattern from the southern found in the Cape Fold Mountains (Western and Eastern Cape) part of South (Samways 1993), a trend strongly followed Africa with northerly extensions to the west-

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ern escarpment to the coast and eastern lands National Park and the Sentinel, drop escarpment to the coast. Chrysoritis is poor- considerably in height above see level (down ly represented in the central grassland to a maximum height of 2350 m). Chrysori- plateau. No Chrysoritis species occur in the tis pelion represents the northernmost outlier savanna biome, hence their absence in the of the Chrysoritis thysbe group on the east- North West and Limpopo provinces of South ern side of Southern Africa, where the but- Africa (Table 1; Fig. 2). terfly is found on the Alti Mountain grass- land (Granger & Bredenkamp 1996). Patch- The most northern locality record for a es of Afromontane vegetation, affiliated to species belonging to the Chrysoritis thysbe Fynbos, are scattered throughout this grass- species group, on the eastern (moister) side land (Granger & Bredenkamp 1996). This of South Africa, is that of Chrysoritis pelion. distribution of Chrysoritis pelion represents The northernmost locality records for a faunal reflection of extensions of such Chrysoritis pelion is at the Golden Gate Fynbos-affiliated vegetation. Highlands National Park. M.C. Williams dis- covered the locality (Terblanche 1991). In contrast, Chrysoritis aethon of the Chrysoritis pelion was also collected by Chrysoritis chrysaor species group, has been Terblanche, in January 1991, near Sentinel found much further north on Mariepskop Mountain, where the Drakensberg and Malu- (Fig. 2) (Pringle et al. 1994) and represents ti mountains meet. The butterfly is more at present the most northerly distributed widespread to the south, in the highlands of Chrysoritis species. Mariepskop is 510 km Lesotho (Fig. 2). Both the above localities of NNE of the Golden Gate Highlands Nation- Chrysoritis pelion are found at an altitude al Park. The Chrysoritis chrysaor species above 2700 m, in areas with rocks and group is significant in the sense that it con- shrubs, among the grasslands. The high- tains the most north-easterly recorded lands, northwards of the Golden Gate High- species of the Chrysoritis genus, namely

Fig. 2. Map of South Africa showing the nine provinces and three localities of importance.

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Chrysoritis phosphor subsp. phosphor, Bankenveld (now Rocky Highveld Grass- Chrysoritis aethon, Chrysoritis aureus and land) as a fire-maintained grassland that the widespread Chrysoritis chrysaor. Signif- would develop into savanna if fire were icantly, the northern extensions of the genus excluded. Fire, as variable, plays at least an on the western side of South Africa contain important role in the maintenance of grass- mostly members of the Chrysoritis thysbe land (Bredenkamp & Van Rooyen 1996; group. On the northern extension of their O’Connor & Bredenkamp 1997). In the light range and on the eastern side of the country, of the above distribution of Chrysoritis the Chrysoritis chrysaor group is better rep- species, it is significant that Chrysoritis resented. All the Chrysoritis species that aureus extends the distribution of Chrysori- occur in the moister (eastern) side of South tis to inland grassland areas to the west of the Africa belong to the Chrysoritis chrysaor escarpment at altitudes between 1600 m and species group and the centre of endemism of 1800 m. These distributions show that even the Chrysoritis chrysaor species group among the species groups of Chrysoritis dif- seems to be outside the Fynbos biome. Only ferent ecological pressures may have shaped two of the eight species (that is 25 %) that the species. belongs to the Chrysoritis chrysaor species group has been recorded in the Western A study of the vegetation of three localities Cape, of which one is also widespread in the of Chrysoritis aureus points to the possible eastern side of South Africa. This is in con- importance of fire in maintaining a viable trast with the tendency of the rest of the open grassland habitat for the butterfly genus (Table 1). It seems, therefore, that the (Terblanche et al. 2003). Since many centre of endemism of the Chrysoritis colonies of species of the Chrysoritis chrysaor group is in the highlands of chrysaor group is often found in grassy areas KwaZulu/ Natal (6 species), Mpumalanga (5 not far from forests or well-wooded kloofs, species) and the Eastern Cape (4) species. fire and climate may have been important in The exception is Chrysoritis natalensis that shaping these species and their ant associa- occurs in the moist, subtropical, eastern tions. coastal areas. Nevertheless, the centre of After decades of exploration by a number of endemism of the Chrysoritis chrysaor group Lepidopterists only five confirmed localities is not in the Fynbos biome but in the eastern of Chrysoritis aureus were reported parts of the Grassland biome. Furthermore, (Terblanche et al. 2003). A fortunate contri- when the northerly extensions of the bution towards the conservation of Chrysori- Chrysoritis genus is considered, the tis aureus was the discovery of the locality at Chrysoritis chrysaor group has its centre of the Alice Glockner Nature Reserve (near endemism within the summer rainfall area, Heidelberg, ) in 1998 by P.S. Roos whereas the Chrysoritis thysbe group has its and G.A. Henning (Roos & Henning 2000). centre of endemism in the winter rainfall This discovery has increased the conserva- area. The only Chrysoritis associated with tion priority of the, at that stage, undevel- afromontane forest, namely Chrysoritis oped Alice Glockner Nature Reserve consid- phosphor, is also a member of the Chrysori- erably, since this locality would probably tis chrysaor species group, although have been destroyed by urbanisation. The Chrysoritis phosphor, that superficially future of Chrysorities aureus has been looks atypical to the genus might not be con- secured by the co-operation between devel- generic. Interestingly, Chrysoritis phosphor opers, government officials and concerned is often recorded at forest margins bordering NGO’s such as the Lepidopterists’ Society of grassland. The only Chrysoritis species, Africa. apart from the ubiquitous Chrysoritis chrysaor that is encountered sparsely in the Chrysoritis aureus is very localised in all the Rocky Highveld Grassland, is Chrysoritis areas where it occurs. Most of the individu- aureus. Acocks (1975, 1988) interpreted the als in such a colony will for most of the time

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Table 3 The five confirmed and apparantly viable localities of Chrysoritis aureus (Heidelberg Copper butterfly) Locality Province Grid reference 1 Altitude (m) Alice Glockner Nature Reserve Gauteng 26º34'S, 28º22E(2628CB) 1690 8 km S Heidelberg Hill N of Greylingstad Mpumalanga 26º44'S, 28º44'E(2628DA) 1810 Malanskraal locality14 km W Balfour Mpumalanga 26º39'S, 28º26'E(2628CB) 1780 Suikerbosrand Nature Reserve Gauteng 26º29'S, 28º18'E(2628AD) 1720 Type locality at the South African Gauteng 26º29'S, 28º21'E(2628AD) 1600 National Force premises 1 The grid reference reading is from the north-western corner of each minute by minute square.

fly within an area of 200 m2 to 400 m2. The females were recorded on separate dates, on size of these localities is being measured at the northern aspect below the summit of the present. Migration behaviour is not associat- hill South of Diepkloof (26º29'S; 28º12'E). ed with any of the Chrysoritis species and All these specimens were recorded at an alti- the populations of every species tend to live tude of 1860–1900 m, which is higher than in relatively small confined areas. The the normal elevation at which Chrysoritis Chrysoritis populations and sub populations aureus occurs (see Table 3). The markings of are therefore referred to as colonies in litera- the undersides of these specimens left one in ture. Males of the Chrysoritis aureus seem to no doubt about the identification. This is the choose an outpost (usually some branch of a first time that such “isolated” Chrysoritis shrub) from where it controls its small terri- aureus individuals were reported, which is tory, whereas females often wander around especially significant in the light of the within the borders of such a colony. Despite the fact that confinement of Chrysoritis above-mentioned surveys that the Lepi- species to its habitats is appreciated by all dopterists Society of Southern Africa had the lepidopterists experienced with the done in Suikerbosrand Nature Reserve Chrysoritis butterflies, scientific information (Henning 1986a, 1986b, 1987). The full sig- on the dispersal of Chrysoritis species is nificance of these findings is being further absent. Two single males and two single researched but may point to dispersal behav- females of Chrysoritis aureus were recorded iour. by R.F. Terblanche on 1 October 2002 and The above outline of the knowledge on the 23 January 2002 at Suikerbosrand Nature distribution of Chrysoritis aureus highlights Reserve 10.5 km W and 6.5 km WSW of the known locality in this reserve. No host plant a few important aspects of research and con- (Clutia pulchella) occurs near these spots, servation management. The importance of and only a single specimen that was clearly metapopulation studies and the lack of staying in a small area of not more than knowledge on the dispersal of Chrysoritis 25 m2 was recorded each time. The two species become clear. Corridors and linkages males were observed, each in an isolated are important for the conservation of insect spot more than 100 m apart, at Perdekop diversity (Samways 1993; 1994; Pryke & (26º30'S; 28º15'E). They were near the sum- Samways 2001) and the possible dispersal mit, defending their small area in the manner behaviour of Chrysoritis should be investi- they would do in a normal colony. The gated in more detail.

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Conservation history (NGOs, politics, law, the only known locality of the butterfly for a education and environmental management) long time (De Wet 1987). Unfortunately, all the members of the genus Charaxes were Science is vital, but alone cannot curb the also put on the protected wild list at decline of the earth’s biodiversity (Kohm et the same time. Many of the Charaxes species al. 2000). Kohm et al. (2000) further state are very widespread in the savanna and had that new partnerships and improved commu- not been under any threat at any stage. The nication should be forged among scientists, placement of all the Charaxes species on the managers and policy makers so that conser- protected list therefore compromised the vation science could be available and under- validity of the above Ordinance and did not standable to all parties. Whitten et al. (2001) help to highlight the importance of protect- that ask if conservation biology is merely ing Chrysoritis aureus. This showed that it is another scientific discipline, safely nestled essential that effective consultation with within the confines of academia, is very stakeholders should take place before laws appropriate for butterfly conservation (and are instated. An NGO that has played a very not only the forests of Indonesia that are dis- important role in Lepidoptera conservation cussed in the relevant paper). The question is to date is the Lepidopterists’ Society of further asked whether conservation is not a Southern Africa that was founded in 1983. mission-concerned with judicial reform, One of the aims of the Lepidopterists’ Soci- political economy, other people’s spatial ety of Africa (previously of Southern Africa) planning, community participation, poverty is to promote the conservation of Lepi- alleviation, human and institutional capacity, doptera in the Afrotropical region. Although consumption, population growth and agri- this society consists largely of amateur lepi- cultural planning (Whitten et al. 2001). The dopterists, major contributions towards but- great challenge of conserving butterfly habi- terfly (insect) conservation in Africa was ini- tats in South Africa thus not only depends on tiated by its members. The proclamation of the level of biological scientific information, the Ruimsig Entomological Reserve for the but on how that information is presented and threatened subsp. dentatis linked with management that takes into by the Roodepoort City Council in 1985 – account socio-economic factors. the first of its kind in Africa – was a land- mark for conservation of localised butterflies Conservation history in Africa. This event draw attention to the urgency and significance of conserving the The conservation of butterflies in South vulnerable habitats of localised myrme- Africa has a proud history, but has not been cophilous butterflies. No issue on insect con- without difficulties. Henning (1997) gave an servation in Africa entered the realm of gov- outline of major events that enhanced the ernment policies and public perceptions to conservation of butterflies in South Africa. such an extent as the saga that led to the pro- Four of sixteen butterflies that were added to tection of the last known locality of Schedule 2 of the list of protected wild ani- niobe (the Brenton Blue butter- mals in the Cape Province in 1976, due to fly). The “Brenton Blue Saga” has been motivation by C.G.C Dickson, were described in detail by Steenkamp & Stein Chrysoritis species. These were Chrysoritis (1999). The Minister of Environmental endymion, Chrysoritis lyncurium, Chrysori- Affairs and Tourism invoked Section 31A of tis nigricans and Chrysoritis rileyi. In the the Environment Conservation Act, 73 of northern areas (the old Transvaal), Chrysori- 1989, whereby a landowner was deprived of tis aureus was added to the protected wild developing his property in order to conserve animal list of Transvaal (now known as four the habitat of the Brenton Blue butterfly provinces: Gauteng, Mpumalanga, Northern (Orachrysops niobe) (Steenkamp & Stein Province and North West) under Ordinance 1999). Any similar conflict to follow will be 12 of 1983, since the type locality had been able to deal with the matter at hand with far

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firmer legislative and procedural foundations Monitoring of these localised butterflies at their disposal (Steenkamp & Stein 1999). should be in place for such species that could become threatened in a relatively short time The Red Data Book of butterflies (Henning by alien vegetation and perhaps also global & Henning 1989; Henning & Henning 1995) climate changes. According to the most was yet another major achievement for recent IUCN categories, the red listing is insect conservation in Africa. Again, without being addressed at present. Blue lists of the endeavours of many professional ento- threatened species may later be introduced to mologists but many more of the amateur fra- motivate people in their conservation efforts ternity over the years, the Red Data Book (Gigon et al. 2000). Such lists could be of would have remained only a dream. Table 2 special value to the conservation of gives a summary of the Chrysoritis species Chrysoritis species. that were regarded as Red data species. A number of synonymies are present for for- merly Red Data categorised butterflies Conservation, collecting and the law (Table 2), which confirms that the taxonomy of the Chrysoritis genus, should be more sat- The legal regime for the conservation and isfactory resolved, especially if the aim was sustainable use of biodiversity is centred on to compile species lists for red listing. the 1992 Convention on Biological Diversi- ty. The primary focus of the 1992 Conven- The red listing of some of the Chrysoritis tion is the conservation of biodiversity and species needs to be reviewed to an extent equitable distribution of its benefits that one can follow conservation progress (Glazewski 2000). The inclusion of an envi- clearly. The latest categories by the IUCN ronmental clause, 108 of 1996, in the Bill of (2001)—adopted in 1994—will help to stan- Rights chapter of the South African Consti- dardise and establish a red listing, especially tution was a major step in the development in the case of Chrysoritis, for those species of substantial environmental law in South that were previously in the rare and interme- Africa (Glazewski 2000). Not only does the diate categories of the Red Data Book. A environmental clause constitute statements review falls beyond the scope of this paper that clearly promotes conservation, these but a few examples are mentioned to illus- have also been further concretised in the set trate some practical aspects of such a review. of environmental management principles In the case of Chrysoritis dicksoni one is underpinning the National Environmental sure at present that the butterfly is in the Management Act, 107 of 1998 (NEMA) process of vanishing if proper conservation (Glazewski 2000). In effect, these inclusions strategies were not implemented in the near in law had and will have a substantial con- future (this butterfly should be entered in the structive effect on conservation of sensitive critically endangered category). As men- habitats and endangered species in South tioned earlier in the paper, the numbers of Africa in future. Some species, such as the Chrysoritis aureus are also dwindling (it is red listed species, are protected in various almost absent at its type locality) but it ways in the schedules of ordinances of the seems that conservation efforts may be in provinces in South Africa. Such a system is time to save the butterfly, and especially thought to be regionally adaptable to local much of its metapopulation structure, from needs and ecological circumstances the impact of urbanisation. Localised (Glazewski 2000). The four nature conserva- Chrysoritis species in remote farm areas tion ordinances that applied to the former such as those in the Sutherland area four provinces in South Africa, still apply in (Roggeveld escarpment) in the Northern many cases to the nine provinces of South Cape should probably be transferred from Africa (see Glazewski 2000). In practice the the old rare to the new least concern catego- above necessitates a constant vigilance by ry or be abandoned from the red list if more the scientific community to monitor the sta- localities become known (see IUCN 2001). tus of species in each province and thus

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demands a sophisticated administrative and valuable in the past. This is relevant for the technical infrastructure that many of the Chrysoritis species, e.g., the new localities under-resourced provinces lack (Glazewksi that have been found for Chrysoritis dick- 2000). soni. In South Africa, there is no evidence of any butterfly populations being destroyed by The particular challenges of the monitoring collecting. It is recommended that collection of insect species in the vast South African take place strictly for scientific purposes, countryside should be appreciated here. In such as for SEM studies, at any of the known the case of Chrysoritis (which already localities of Chrysoritis dicksoni. Collection belongs to a well-known insect group, the of more specimens at these localities, for butterflies) neither the scientific community purposes of expanding collections, will be of nor the conservation authorities has the man- no value. The same applies to other localised power and funds to monitor the status of the butterflies from the genus Orachrysops rare species regularly enough to follow (Orachrysops niobe and Orachrysops ari- progress or decline sufficiently, not even in adne). It is not certain whether the conserva- nature reserves or national parks. For per- tion legislation would be better served by a spective, note that for many of the nature national body than legislation by the differ- reserves and national parks in South Africa a ent provinces or at least a streamlined coor- species list of butterflies (the most baseline dination from a national level. The issuing of information) does not exist. To complicate permits should be streamlined and a more matters, some of the Chrysoritis species, pragmatic approach should be adapted for such as Chrysoritis endymion, are not seen facilitating collectors who wants to make a every year despite visits by lepidopterists at contribution towards the knowledge on Lep- the right time of the year. To identify the idoptera. As stated in Glazewski (2000): more rare Chrysoritis species in the field and “much remains to be done particularly as even in the laboratory, takes a specialist – a regards the co-ordination of the administra- skill that would be nearly impossible in most tion of a diffuse set of laws”. cases for the already overburdened conserva- tion officials to acquire. One possible solu- There remains, as pointed out by Steenkamp tion is to involve the insect collectors (in this & Stein (1999), a number of strategic issues case the butterfly collector) in the monitor- with which the conservation establishment ing as well as the location of the insect (but- will have to deal if they want to ensure ongo- terfly) habitats. ing civil society participation in environmen- tal decision-making. The conservation An important issue, inevitably touched on in efforts in connection with Aloeides dentatis many of the above discussions regarding leg- and Orachrysops niobe have led to an aware- islation that seriously needs to be addressed, ness that will possibly enhance the conserva- is the role of the butterfly collector. There- tion of all the localised lycaenid butterflies in fore, this is an area that needs urgent atten- South Africa. Personal experience is that tion. New (1997) stated that it is vital to sus- since the application of the law in the case of tain the symbiosis and co-operation between Orachrysops niobe, people, including professional scientists and amateurs, as effi- landowners and developers, have been much cient assessment of many butterfly popula- more aware of possible red listed butterfly tions can only be enhanced in this way. Arm- species and possible unique habitats. To strong (1998) gives a valuable perspective what extent, remains to be seen. Biodiversi- on the importance of the contribution of the ty and the quality of such data in environ- amateur lepidopterist but also the vital role mental impact assessments deserve attention of conservation authorities. The information beyond the scope of this overview. Another that could be made available by bona fide key problem in saving threatened butterfly amateur butterfly collectors for properly species is the notion that the habitat must be conserving butterflies and ecosystems by the last known habitat before the protection conservation authorities have been very really becomes urgent. Conservation of but-

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terflies (and other insects) should be seen in of butterflies—or even new scientists—than a much wider integrated sense of conserving no involvement caused by too strict legisla- metapopulations. tion. In the end such involvement will enhance the conservation of butterflies (or other insects) rather than decrease the Education and awareness chances of their survival. A fear that the lay- Without education and public awareness the man collector might collect butterflies such loss of touch with the environment, in which as Chrysoritis aureus and destroy colonies of most modern and developing societies seem the butterfly is probably overrated for the to find themselves, would continue. Conser- following three reasons. The butterfly is easy vation in South Africa has shifted in recent to recognise by anybody with a good refer- years from a protectionist approach towards ence guide and reasonable experience of one that involves the community in conser- Lepidoptera, but unlikely to be found by a vation (Wynberg 2002). Butterflies such as novice. Furthermore, the confirmed locali- Chrysoritis should be taken to the communi- ties are clearly protected in nature reserves, ty. Education, and use of a more indigenous especially the established Alice Glockner curriculum in schools is needed to appreciate Nature Reserve. Any lepidopterist that the existence of biological intricacies such as would want to survey butterflies in such a the ecology of Chrysoritis species in Africa. nature reserve would have to submit a Nature reserves such as the Alice Glockner research proposal (which always foreruns Nature Reserve and Suikerbosrand Nature the issuing of any permit), stating that they Reserve provide excellent opportunities for know about the butterfly and will avoid to such educational programmes and should be collect any of these. managed as such. In the case of Chrysoritis aureus, the efforts of Gauteng Nature Con- Integrating science,environmental manage- servation to enhance co-operation in ment, policies, law and education research and conservation have been most valuable. Hopefully, these initiatives will The integration of the efforts of scientists, prove valuable in a country that has many the public sector and the government can socio-economic issues that have to be only be successful if proper communication addressed urgently. Encouraging is that the takes place and therefore proper forums are above conservation efforts are directed at the available for such communication. The jour- ‘small things’ in natural veld, a possible nal Metamorphosis (Journal of the Lepi- source for ecotourism and education – very dopterists’ Society of Southern Africa) has poorly exploited up to now. Synecological acted as a forum for debate in the past (e.g. studies, which are conducted to conserve the De Wet 1990; Vlok 1993; Woodhall 1994) butterfly, could also benefit the development between the lepidopterists and conservation of the Alice Glockner Nature Reserve as a authorities, with regard to the issue of col- whole, addressing other aspects of biodiver- lecting butterflies and approaches in conser- sity apart from butterfly conservation. At vation. The discussions between conserva- present the management of urbanisation tion authorities, researchers and other stake- close to the Alice Glockner Nature Reserve holders on the issuing of permits for collect- deserves further attention, especially with ing insects as well as the law governing the regard to community involvement and public protection of material collected in South awareness. Africa, was initiated at the 13th Congress of the Entomological Society of Southern The involvement of the younger generation Africa in Pietermaritzburg were encourag- (school learners and students), if only to start ing. Another important event that coincided collecting butterflies in a back yard, may with the 13th Congress was a symposium on mean much more to science in the end, if it diversity and conservation in results, for instance, in new locality records Southern Africa, so that emphasis was also

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on insect conservation and not only those takes responsibility— aided by codes of con- insects of economic agricultural importance duct—to maintain optimal co-operation (not that it is not all part of environmental between the Government and the NGO management). should be in place. Species such as Chrysori- tis dicksoni should be protected by law (also from collecting) at their known and con- Conclusions served localities. Since the genus contains a number of Distribution records are not in any atlas for- localised populations and subpopulations of mat, a fact which limits our understanding of significant biological and aesthetic value, especially a genus such as Chrysoritis that Chrysoritis genus should be seen as endemic contains a number of highly localised and as a whole. It is clear that the genus contains endemic butterflies. A Southern African but- bioindicators that should be verified accord- terfly atlas would be appropriate and in fact ing to the categories set out by McGeoch a necessity, since it would enhance the (1998). Since the butterfly populations of understanding of the biogeography of a Chrysoritis tend to be localised, it is espe- group of butterflies that might be good bio- cially their role as ecological indicators and indicators on a large geographical scale. It is biodiversity indicators that are worth consid- to be hoped that butterflies, that seems to be ering. The scales within which the Chrysori- ideal for atlas work, would be included as tis species could be used are important. On a part of the Biodiversity and Conservation for larger geographical scale their dwindling South African Invertebrates initiative number of populations is certainly indicative (Hamer & Underhill 2001) or any other sim- of anthropogenic pressures on sensitive habi- ilar initiatives. tats. The red listing of localised (myrme- Despite the fact that butterflies are such a cophilous) butterflies such as the Chrysoritis relatively well studied group of insects in is in need of urgent review in the light of the South Africa, limitations in our knowledge most recent IUCN categories (IUCN 2001) still exist even on the taxonomic and distrib- that were adopted in 1994. The environmen- utional level. Being a relatively well-known tal management priorities are linked to invertebrate group, the taxonomy of butter- applied ecology, but the infrastructure to flies such as Chrysoritis has reached a state build the necessary bridges in practice, is where revisions could easily result in a num- often lacking. A sound ecosystem approach ber of unnecessary names. It is recommend- in conservation biology lies in the intersec- ed that the relationships between the entities tion between ecological, socio-economic and are carefully analysed in such a manner that institutional perspectives (Meffe & Carol the work can be traced and repeated. More 1997). This cannot be more true for the con- specific distribution records will be a valu- servation of Chrysoritis species. It is there- able aid for resolving the taxonomy and fore clear that the prosperity of Chrysoritis responses of such habitat-sensitive butter- species not only depends on our understand- flies to environmental changes. Legislation ing of its interactions in nature in the context should facilitate rather than limit the valu- of communities, but also of our human inter- able role that the amateur may play to con- actions as scientists, NGO members, policy- tribute in the gathering of distribution makers, law makers, educators, environmen- records. In turn, the amateur lepidopterists talists and citizens in whatever capacity— should adapt and make an effort to include especially when it comes to the integration unknown localities, apart from monitoring of our efforts. The above that the organisa- butterflies at their well-known localities. tions/institutions involved with conservation This information should also be readily strategies, the monitoring ideals and the leg- available to conservation authorities. A sys- islation, should appreciate the challenges tem in which the amateur fraternity also particular to insect conservation.

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