This dissertation focuses on the systematics of the Paussinae, a lineage of approximately 775 beetle species all of which produce explosive defensive chemicals and many which are obligate associates of ants (myrmecophiles). This dissertation includes six discrete studies, which taken together span the spectrum of systematic research. It includes taxonomic projects on both larval and adult paussines. It also includes phylogenetic investigations, based on DNA sequence data and morphological data, that address the evolutionary relationships among the major lineages within Paussinae as well as the evolutionary position of the Paussinae within the beetle suborder Adephaga. The inferred trees are used to study character evolution of physiological and behavioral traits of these organisms in a phylogenetic framework. This work sets the stage for a lifetime of work on this fascinating group.

The Brenton Blue butterfly, Orachrysops niobe (Trimen) is currently only known to be extant at one locality in the southern Cape (the Brenton Blue Butterfly Reserve = BBBR), and currently globally Red Listed as "Critically Endangered". This research investigates the life history and ecological interactions of o. niobe and recommends management techniques for the BBBR. Adult nectar sources and female oviposition behaviour are described. The first two larval instars feed on the leaflets of the host plant Indigofera erecta Thunb., and the 3rd and 4th instar larvae feed on this plant's woody rootstock, attended by ants Camponotus baynei Arnold. Cannibalism takes place in the early larval stages. Adults emerge from late October - early December, from late January to early March, and occasionally there is a third brood in April. Morphological and behavioural features of the larvae are described, and larval growth patterns are compared to other polyommatines. The nature of the myrmecophilous behaviour is assessed and the phylogenetic relationships between Orachrysops, Euchrysops, Lepidochrysops, and other polyommatine genera are discussed. Ant assemblages at the BBBR are described from pitfall trap sampling and field observations of interactions between O. niobe larvae and ants. The ant assemblages at the BBBR, Nature's Valley Fynbos Reserve (NVFR) and Uitzicht 216/ 40 are compared. The potential impact of the Argentine ant at the BBBR is discussed. Adult and egg counts are used to study the population dynamics of O. niobe, leading to an estimate of the adult butterfly population and its fluctuations. A population dynamics model is constructed, and several factors impacting fecundity and mortality are assessed. Population studies on other polyommatines are compared and discussed. The morphology, reproductive biology and autoecology of the papilionoid legume I. erecta are described. An explanation is offered why the larvae of O. niobe are monophagous on this plant. Microhabitat variations cause changes in its vegetative morphology and this further explains the restricted range of O. niobe. The population dynamics of I. erecta is investigated and the effects of biotic and abiotic factors (including fire) assessed. Braun Blanquet methodology is used to sample and classify the vegetation communities at the BBBR. Ordination techniques are used to confirm the classification and to diagnose for environmental gradients. Hypotheses are generated about the ecological processes functioning at the site, and the environmental niche occupied by I. erecta. Correlations between the occurrence xxii of vegetation types and other plant species and I. erecta are sought and a strong association with Pterocelastrus tricuspidatus is demonstrated. Three study sites are described and the ecological history of the region is reconstructed. Ecosystems are no longer fully functional because natural fire regimes and megaherbivores are absent. It is suggested that O. niobe currently inhabits an interglacial refugium under stress from the current global warming trend. The remaining suitable habitat is threatened by coastal property development and environmental degradation. An increased population of o. niobe at the BBBR, and establishment of new populations is essential to avoid extinction. Management techniques are evaluated and a management strategy for the BBBR is proposed. / Thesis (Ph.D. (Environmental Science))--North-West University, Potchefstroom Campus, 2006.

The Brenton Blue butterfly, Orachrysops niobe (Trimen) is currently only known to be extant at one locality in the southern Cape (the Brenton Blue Butterfly Reserve = BBBR), and currently globally Red Listed as "Critically Endangered". This research investigates the life history and ecological interactions of o. niobe and recommends management techniques for the BBBR. Adult nectar sources and female oviposition behaviour are described. The first two larval instars feed on the leaflets of the host plant Indigofera erecta Thunb., and the 3rd and 4th instar larvae feed on this plant's woody rootstock, attended by ants Camponotus baynei Arnold. Cannibalism takes place in the early larval stages. Adults emerge from late October - early December, from late January to early March, and occasionally there is a third brood in April. Morphological and behavioural features of the larvae are described, and larval growth patterns are compared to other polyommatines. The nature of the myrmecophilous behaviour is assessed and the phylogenetic relationships between Orachrysops, Euchrysops, Lepidochrysops, and other polyommatine genera are discussed. Ant assemblages at the BBBR are described from pitfall trap sampling and field observations of interactions between O. niobe larvae and ants. The ant assemblages at the BBBR, Nature's Valley Fynbos Reserve (NVFR) and Uitzicht 216/ 40 are compared. The potential impact of the Argentine ant at the BBBR is discussed. Adult and egg counts are used to study the population dynamics of O. niobe, leading to an estimate of the adult butterfly population and its fluctuations. A population dynamics model is constructed, and several factors impacting fecundity and mortality are assessed. Population studies on other polyommatines are compared and discussed. The morphology, reproductive biology and autoecology of the papilionoid legume I. erecta are described. An explanation is offered why the larvae of O. niobe are monophagous on this plant. Microhabitat variations cause changes in its vegetative morphology and this further explains the restricted range of O. niobe. The population dynamics of I. erecta is investigated and the effects of biotic and abiotic factors (including fire) assessed. Braun Blanquet methodology is used to sample and classify the vegetation communities at the BBBR. Ordination techniques are used to confirm the classification and to diagnose for environmental gradients. Hypotheses are generated about the ecological processes functioning at the site, and the environmental niche occupied by I. erecta. Correlations between the occurrence xxii of vegetation types and other plant species and I. erecta are sought and a strong association with Pterocelastrus tricuspidatus is demonstrated. Three study sites are described and the ecological history of the region is reconstructed. Ecosystems are no longer fully functional because natural fire regimes and megaherbivores are absent. It is suggested that O. niobe currently inhabits an interglacial refugium under stress from the current global warming trend. The remaining suitable habitat is threatened by coastal property development and environmental degradation. An increased population of o. niobe at the BBBR, and establishment of new populations is essential to avoid extinction. Management techniques are evaluated and a management strategy for the BBBR is proposed. / Thesis (Ph.D. (Environmental Science))--North-West University, Potchefstroom Campus, 2006.

The presented thesis deals with the evolution of the social parasitism of Phengaris (Maculinea) butterflies. It consists of three related problems: I) The phylogeny of Glaucopsyche section (Lepidoptera: Lycaenidae: Polyommatinae) {--} the social parasitism occured only once in the Glaucopsyche section. II) The conditions in which the social parasitism of Phengaris evolved {--} the process was probably facilitated by unique species composition of the myrmecofauna of the biotopes inhabited by the ancestor of Phengaris. III) The host specificity of Phengaris {--} today{\crq}s view on the host specificity of Phengaris is poorly supported.

Although the research on myrmecophilous insects is dated back to the 19th century, certain aspects of fascinating relationship between hosts and their social parasites are not satisfactorily understood. The latest studies focus mostly on the systematics of myrmecophilous taxa, and bionomy remains neglected. Most of the myrmecophilous insect can be found among beetles (Coleoptera). These myrmecophiles reveal a broad spectrum of adaptations that allow them to live in close association with their host ants. These adaptations significantly differ among the well-integrated species on one hand and poorly integrated or non-integrated species of myrmecophilous beetles on the other. This study focuses at the comparison of behaviour in two species of myrmecophilous beetles, Claviger testaceus (Staphylinidae: Pselaphinae) as the member of integrated myrmecophiles, and Pella spp. (Staphylinidae: Aleocharinae) as poorly integrated or non-integrated beetle. My work shows significant differences in behaviour taking place after encounter of a beetle and an ant. Ants are more aggressive towards the non-integrated species, and at the same time, these beetles are well-adapted for aggressive behaviour from the ants. On the other hand, the integrated beetles do not possess many behavioural adaptations for the...

No description available.

The 1883 discovery, research into generic life histories and taxonomic investigations into the aphytophagous butterfly Thestor brachycerus brachycerus (the Knysna Skolly) from Knysna, South Africa are described. Its slide towards extinction through loss and degradation of habitat in the late 20th century left only one known colony by 2009 leading to it being assessed as Critically Endangered in the 2013 South African Red List and Atlas. During searches for more populations in 2011, a second T. b. brachycerus colony was found which differed markedly with respect to aspect, altitude and vegetation. Study areas were defined around the original colony on the Pezula Golf Estate (Pezula) and the newly discovered coastal colony east of Coney Glen (Coastal site). An assessment is made of the ecological differences and similarities between the two sites. The extent and size of the two known populations was measured; the Extent of Occupancy was found to be 7.5 km2 and the Area of Occupancy as 1.05 x 10-3 km2. During observations over 5 seasons in December and January 2009 – 2014 the average number of butterflies observed at the original Pezula colony varied from 1 – 8 males and 0 – 3 females per survey, compared with 1 – 6 males and 1 – 3 females observed over 3 seasons from 2011 to 2014 at the Coastal colony. The mating, territoriality, oviposition and predation avoidance behaviour of the adult T. b. brachycerus butterflies was studied by direct observation. Males were found to exhibit territorial behaviour, spending long periods of time perched in one spot, particularly in areas where the vegetation was open or low. Oviposition was rapid and no preference was shown by females for any particular plant. Between the two colonies, oviposition was observed on a total of 16 different plants. The vegetation composition at the two study sites was analysed using Braun–Blanquet methodology. Only one plant, Tarchonanthus littoralis, was found to be common to both. There appeared to be no correlation between floristic composition and the occurrence of T. b. brachycerus. All the Thestor species so far studied by other workers have been found to be myrmecophilous, having an association with the pugnacious ant Anoplolepis custodiens. Ant communities were sampled by employing pitfall traps and direct sampling by hand. At both the Pezula and Coastal sites A. custodiens was the most ubiquitous ant species. The insights gained have enabled implementation of a habitat management plan. Experimental management methods are recommended at the Pezula site which include small scale burning and selective cutting of vegetation. Search for more colonies is ongoing, using the improved ecological knowledge of the butterfly. / MSc (Environmental Sciences), North-West University, Potchefstroom Campus, 2015

The 1883 discovery, research into generic life histories and taxonomic investigations into the aphytophagous butterfly Thestor brachycerus brachycerus (the Knysna Skolly) from Knysna, South Africa are described. Its slide towards extinction through loss and degradation of habitat in the late 20th century left only one known colony by 2009 leading to it being assessed as Critically Endangered in the 2013 South African Red List and Atlas. During searches for more populations in 2011, a second T. b. brachycerus colony was found which differed markedly with respect to aspect, altitude and vegetation. Study areas were defined around the original colony on the Pezula Golf Estate (Pezula) and the newly discovered coastal colony east of Coney Glen (Coastal site). An assessment is made of the ecological differences and similarities between the two sites. The extent and size of the two known populations was measured; the Extent of Occupancy was found to be 7.5 km2 and the Area of Occupancy as 1.05 x 10-3 km2. During observations over 5 seasons in December and January 2009 – 2014 the average number of butterflies observed at the original Pezula colony varied from 1 – 8 males and 0 – 3 females per survey, compared with 1 – 6 males and 1 – 3 females observed over 3 seasons from 2011 to 2014 at the Coastal colony. The mating, territoriality, oviposition and predation avoidance behaviour of the adult T. b. brachycerus butterflies was studied by direct observation. Males were found to exhibit territorial behaviour, spending long periods of time perched in one spot, particularly in areas where the vegetation was open or low. Oviposition was rapid and no preference was shown by females for any particular plant. Between the two colonies, oviposition was observed on a total of 16 different plants. The vegetation composition at the two study sites was analysed using Braun–Blanquet methodology. Only one plant, Tarchonanthus littoralis, was found to be common to both. There appeared to be no correlation between floristic composition and the occurrence of T. b. brachycerus. All the Thestor species so far studied by other workers have been found to be myrmecophilous, having an association with the pugnacious ant Anoplolepis custodiens. Ant communities were sampled by employing pitfall traps and direct sampling by hand. At both the Pezula and Coastal sites A. custodiens was the most ubiquitous ant species. The insights gained have enabled implementation of a habitat management plan. Experimental management methods are recommended at the Pezula site which include small scale burning and selective cutting of vegetation. Search for more colonies is ongoing, using the improved ecological knowledge of the butterfly. / MSc (Environmental Sciences), North-West University, Potchefstroom Campus, 2015

The state of the populations of many insects is critical in the Czech Republic, other species have already extinct. Among the species balancing at the limit of survival is myrmecophilus Alcon Blue (Phengaris alcon), especially its ecological form (P. alcon f. alcon), which is associated with the Marsh Gentian (Gentiana pneumonanthe) during the early larval stages. The general aim of my work was to evaluate the current distribution of the Alcon Blue butterfly (P. alcon f. alcon) in the Czech Republic, to estimate the size of the remaining populations and to propose a practical action plan for conservation of the species in the Czech Republic. The analysis of the databases showed that the species survives theoretically in 11 localities. The field survey in 2016 revealed, however, that at five other sites a species in the recent past was extinct. The remaining 6 localities are concentrated in Southwestern Bohemia on a total area of only 20 hectares. Two population is however close to extinction. The cause of local extinction of Alcon Blues was the extinction of host plants. The main factors influencing the size of butterfly populations in the last localities were the abundace and vitality of host plants (number of fertile shoots, height). Part of my work was an attempt to reintroduce Alcon Blue to a...

Borboletas neotropicais, como os licenídeos, são conhecidas pela notável diversidade, entretanto aspectos ecológicos fundamentais desse grupo taxonômico, particularmente dos imaturos, como informações sobre suas plantas hospedeiras e interações mirmecófilas são praticamente desconhecidos para a maioria das comunidades neotropicais. Além do que, o conhecimento limitado da história natural desses insetos, impede uma melhor compreensão do seu papel ecológico-evolutivo. Com isso, a presente tese teve como objetivo inicial caracterizar a comunidade de imaturos de licenídeos florívoros em uma área de cerrado descrevendo o padrão de sazonalidade, identificando as plantas hospedeiras, a amplitude de dieta e interações com formigas e inimigos naturais. A partir dessa base de informações, o objetivo seguinte foi avaliar a ocorrência de licenídeos florívoros no cerrado de acordo com fatores ligados à disponibilidade de alimento nas plantas e à presença de formigas. Adicionalmente, foram estudados diferentes sistemas de interação licenídeos-plantas-formigas com o intuito de inferir o papel da formiga sobre a seleção de plantas hospedeiras. Em suma, nosso estudo mostrou uma comunidade composta por 31 espécies de Lycaenidae registradas em 41 espécies (17 famílias) de plantas hospedeiras e caracterizada pela marcante sazonalidade, cuja ocorrência de imaturos foi maior durante a estação seca. De forma geral, a comunidade de licenídeos apresentou variação de acordo com a oferta temporal de alimento, acompanhando a fenologia de suas plantas hospedeiras disponíveis no campo. Contudo, a comunidade estudada aqui, formada por licenídeos oligófagos e polífagos teve uma tendência em utilizar plantas com menor quantidade de botões e também selecionar plantas que possuem nectários extraflorais e plantas ocupadas por formigas. Particularmente, nosso estudo revelou que a seleção de plantas hospedeiras mediadas por formigas pode variar de acordo com o sistema estudo. Licenídeos como Allosmaitia strophius e Rekoa marius utilizaram a xi presença de formigas como pista durante a seleção de plantas Heteropterys byrsonimifolia, porém os ovos presentes em ramos com formigas sofreram maior ação de parasitóides. Enquanto que Leptotes cassius mostrou selecionar inflorescências de Bionia coriacea independente da presença de Camponotus, formigas as quais mantém associações facultativas. / The Lycaenidae butterflies are known for their remarkable diversity in the Neotropics; however basic ecological features of this group, especially of the immature stages, such as information about host plants and ant-interactions (myrmecophily) are barely known for most species. Therefore the limited knowledge of lycaenids natural history restrains a better understanding of important ecological-evolutionary aspects. In this thesis we initially characterized the community of florivorous lycaenids in a Brazilian Savanna area, describing the seasonal occurrence of immatures, identifying host plants, the diet breadth and interactions with ants and natural enemies. Then, we evaluated the occurrence of the lycaenids according to factors related to resource availability (host plants) and the presence/absence of ants. In summary, we sampled 31 species of Lycaenidae in 41 plant species (17 families). Immatures were more abundant in the dry season indicating a seasonal pattern of occurrence; in addition lycaenids were related to the temporal availability of food resource, following the phenology of the host plants in the field. The oligophagous and polyphagous species were found predominately in plants which produce few flowers buds. The presence of immatures in extrafloral nectaried plants was commonplace, which was expected given that larvae can benefit from living with ants. A further detailed experimental study revealed that ant presence is a cue for female oviposition in two facultative myrmecophilous lycaenid species Allosmaitia strophius and Rekoa marius, but contrary to our predictions, ants failed to protect lycaenid eggs from parasitoids. In other study system, Leptotes cassius did not discriminate between plants with or without ants, despite the fact that larvae had stable association with ants. Our study indicates that there can be striking differences among lycaenid species with regard to association with ants and the benefits granted from these associations. So generalizations about lycaenid ant associations should be avoided and every study system must be examined with detail.