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Naomi E Pierce - One of the best experts on this subject based on the ideXlab platform.
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recent diversification of chrysoritis butterflies in the south african cape lepidoptera Lycaenidae
Molecular Phylogenetics and Evolution, 2020Co-Authors: Gerard Talavera, Alan Heath, Zofia A Kaliszewska, Naomi E PierceAbstract:Abstract Although best known for its extraordinary radiations of endemic plant species, the South African fynbos is home to a great diversity of phytophagous insects, including butterflies in the genus Chrysoritis (Lepidoptera: Lycaenidae). These butterflies are remarkably uniform morphologically; nevertheless, they comprise 43 currently accepted species and 68 currently valid taxonomic names. While many species have highly restricted, dot-like distributions, others are widespread. Here, we investigate the phylogenetic and biogeographic history underlying their diversification by analyzing molecular markers from 406 representatives of all described species throughout their respective ranges. We recover monophyletic clades for both C. chrysaor and C. thysbe species-groups, and identify a set of lineages that fall between them. The estimated age of divergence for the genus is 32 Mya, and we document significantly rapid diversification of the thysbe species-group in the Pleistocene (~2 Mya). Using ancestral geographic range reconstruction, we show that West Fynbos is the most likely region of origin for the radiation of the thysbe species-group. The colonization of this region occurred 9 Mya and appears to have been followed by a long period of relative stasis before a recent increase in diversification. Thus, the thysbe radiation does not appear to have resulted from the colonization of new biogeographic areas. Rather, the impact of species interactions (with ants and plants), the appearance of key innovations, and/or the opening of new ecological niche space in the region might explain the sudden burst of speciation that occurred in this group 2 Mya. The biogeographic model suggests two different diversification processes with few historical cross-colonisations, one in eastern South Africa for the C. chrysaor group and the other in western South Africa for the remaining taxa. Distributional range assessments and ecological niche models for each species show important niche overlap, and in a few cases, complete overlap. However, these shared traits are not explained by phylogenetic history. Chrysoritis taxa frequently fly in sympatry and gene tree reticulation appears to be widespread at the species level, suggesting that several episodes of range shifts might have led to secondary sympatries, allowing limited gene flow that challenges species delimitation efforts. In addition, the unusually high diversification rate for the thysbe clade of 1.35 [0.91–1.81] lineages per million years also suggests the possibility of taxonomic oversplitting. The phylogeny presented here provides a framework for a taxonomic revision of the genus. We highlight cases of potential synonymy both in allopatry and sympatry, and stress the importance of dedicated studies to assess potential pre- and post-zygotic barriers giving rise to species delimitations of the thysbe group.
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reinforcement of pre zygotic isolation and karyotype evolution in agrodiaetus butterflies
Nature, 2005Co-Authors: Vladimir A Lukhtanov, Nikolay P Kandul, Joshua B Plotkin, A Dantchenko, David Haig, Naomi E PierceAbstract:theory has shown that reinforcement is a possible mechanism that can lead to speciation 6‐8 , empirical evidence has been sufficiently scarce to raise doubts about the importance of reinforcement in nature 6,9,10 . Agrodiaetus butterflies (Lepidoptera: Lycaenidae) exhibit unusual variability in chromosome number. Whereas their genitalia and other morphological characteristics are largely uniform, different species vary considerably in male wing colour, and provide a model system to study the role of reinforcement in speciation. Using comparative phylogenetic methods, we show that the sympatric distribution of 15 relatively young sister taxa of Agrodiaetus strongly correlates with differences in male wing colour, and that this pattern is most likely the result of reinforcement. We find little evidence supporting sympatric speciation: rather, in Agrodiaetus, karyotypic changes accumulate gradually in allopatry, prompting reinforcement when karyotypically divergent races come into contact.
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the ecology and evolution of ant association in the Lycaenidae lepidoptera
Annual Review of Entomology, 2002Co-Authors: Naomi E Pierce, Michael F Braby, Alan Heath, David J Lohman, John Mathew, Douglas B Rand, Mark A TravassosAbstract:The estimated 6000 species of Lycaenidae account for about one third of all Papilionoidea. The majority of lycaenids have associations with ants that can be facultative or obligate and range from mutualism to parasitism. Lycaenid larvae and pupae employ complex chemical and acoustical signals to manipulate ants. Cost/benefit analyses have demonstrated multiple trade-offs involved in myrmecophily. Both demographic and phylogenetic evidence indicate that ant association has shaped the evolution of obligately associated groups. Parasitism typically arises from mutualism with ants, and entomophagous species are disproportionately common in the Lycaenidae compared with other Lepidoptera. Obligate associations are more common in the Southern Hemisphere, in part because highly ant-associated lineages make up a larger proportion of the fauna in these regions. Further research on phylogeny and natural history, particularly of the Neotropical fauna, will be necessary to understand the role ant association has played in the evolution of the Lycaenidae.
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phylogeny and life history evolution of the genus chrysoritis within the aphnaeini lepidoptera Lycaenidae inferred from mitochondrial cytochrome oxidase i sequences
Molecular Phylogenetics and Evolution, 2000Co-Authors: Douglas B Rand, Alan Heath, Tamara Suderman, Naomi E PierceAbstract:Phylogenetic relationships among 26 South African species in the tribe Aphnaeini (Lepidoptera: Lycaenidae) were inferred from DNA characters of the mitochondrial gene cytochrome oxidase I (COI), using maximum-parsimony methods. The resulting phylogenetic estimate supports the systematic hypothesis made by Heath (1997, Metamorphosis, supplement 2), based on morphological characters, that at least three preexisting genera (Chrysoritis, Poecilmitis, and Oxychaeta) should be collapsed into the single monophyletic genus Chrysoritis. Two of the species groups described by Heath within Chrysoritis are also monophyletic, while one is paraphyletic and thus unsupported by the molecular data. Strong node support and skewed transition/transversion ratios suggest that two Chrysoritis clades contain synonymous species. Aphytophagy appears as a derived feeding strategy. Evolutionary patterns of ant association indicate lability at the level of ant genus, while association with different ant subfamilies may have played an ancestral and chemically mediated role in the diversification of South African aphnaeines.
Gerard Talavera - One of the best experts on this subject based on the ideXlab platform.
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recent diversification of chrysoritis butterflies in the south african cape lepidoptera Lycaenidae
Molecular Phylogenetics and Evolution, 2020Co-Authors: Gerard Talavera, Alan Heath, Zofia A Kaliszewska, Naomi E PierceAbstract:Abstract Although best known for its extraordinary radiations of endemic plant species, the South African fynbos is home to a great diversity of phytophagous insects, including butterflies in the genus Chrysoritis (Lepidoptera: Lycaenidae). These butterflies are remarkably uniform morphologically; nevertheless, they comprise 43 currently accepted species and 68 currently valid taxonomic names. While many species have highly restricted, dot-like distributions, others are widespread. Here, we investigate the phylogenetic and biogeographic history underlying their diversification by analyzing molecular markers from 406 representatives of all described species throughout their respective ranges. We recover monophyletic clades for both C. chrysaor and C. thysbe species-groups, and identify a set of lineages that fall between them. The estimated age of divergence for the genus is 32 Mya, and we document significantly rapid diversification of the thysbe species-group in the Pleistocene (~2 Mya). Using ancestral geographic range reconstruction, we show that West Fynbos is the most likely region of origin for the radiation of the thysbe species-group. The colonization of this region occurred 9 Mya and appears to have been followed by a long period of relative stasis before a recent increase in diversification. Thus, the thysbe radiation does not appear to have resulted from the colonization of new biogeographic areas. Rather, the impact of species interactions (with ants and plants), the appearance of key innovations, and/or the opening of new ecological niche space in the region might explain the sudden burst of speciation that occurred in this group 2 Mya. The biogeographic model suggests two different diversification processes with few historical cross-colonisations, one in eastern South Africa for the C. chrysaor group and the other in western South Africa for the remaining taxa. Distributional range assessments and ecological niche models for each species show important niche overlap, and in a few cases, complete overlap. However, these shared traits are not explained by phylogenetic history. Chrysoritis taxa frequently fly in sympatry and gene tree reticulation appears to be widespread at the species level, suggesting that several episodes of range shifts might have led to secondary sympatries, allowing limited gene flow that challenges species delimitation efforts. In addition, the unusually high diversification rate for the thysbe clade of 1.35 [0.91–1.81] lineages per million years also suggests the possibility of taxonomic oversplitting. The phylogeny presented here provides a framework for a taxonomic revision of the genus. We highlight cases of potential synonymy both in allopatry and sympatry, and stress the importance of dedicated studies to assess potential pre- and post-zygotic barriers giving rise to species delimitations of the thysbe group.
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ecological specialization is associated with genetic structure in the ant associated butterfly family Lycaenidae
Proceedings of The Royal Society B: Biological Sciences, 2018Co-Authors: Sami Schar, Rodney Eastwood, Kimberly G Arnaldi, Gerard Talavera, Zofia A Kaliszewska, John H Boyle, Marianne EspelandAbstract:The role of specialization in diversification can be explored along two geological axes in the butterfly family Lycaenidae. In addition to variation in host-plant specialization normally exhibited by butterflies, the caterpillars of most Lycaenidae have symbioses with ants ranging from no interactions through to obligate and specific associations, increasing niche dimensionality in ant-associated taxa. Based on mitochondrial sequences from 8282 specimens from 967 species and 249 genera, we show that the degree of ecological specialization of lycaenid species is positively correlated with genetic divergence, haplotype diversity and an increase in isolation by distance. Nucleotide substitution rate is higher in carnivorous than phytophagous lycaenids. The effects documented here for both micro- and macroevolutionary processes could result from increased spatial segregation as a consequence of reduced connectivity in specialists, niche-based divergence or a combination of both. They could also provide an explanation for the extraordinary diversity of the Lycaenidae and, more generally, for diversity in groups of organisms with similar multi-dimensional ecological specialization.
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when caterpillars attack biogeography and life history evolution of the miletinae lepidoptera Lycaenidae
Evolution, 2015Co-Authors: Zofia A Kaliszewska, David J Lohman, John Mathew, Douglas B Rand, Kathrin Sommer, Glenn Adelson, Gerard TalaveraAbstract:Of the four most diverse insect orders, Lepidoptera contains remarkably few predatory and parasitic species. Although species with these habits have evolved multiple times in moths and butterflies, they have rarely been associated with diversification. The wholly aphytophagous subfamily Miletinae (Lycaenidae) is an exception, consisting of nearly 190 species distributed primarily throughout the Old World tropics and subtropics. Most miletines eat Hemiptera, although some consume ant brood or are fed by ant trophallaxis. A well-resolved phylogeny inferred using 4915 bp from seven markers sampled from representatives of all genera and nearly one-third the described species was used to examine the biogeography and evolution of biotic associations in this group. Biogeographic analyses indicate that Miletinae likely diverged from an African ancestor near the start of the Eocene, and four lineages dispersed between Africa and Asia. Phylogenetic constraint in prey selection is apparent at two levels: related miletine species are more likely to feed on related Hemiptera, and related miletines are more likely to associate with related ants, either directly by eating the ants, or indirectly by eating hemipteran prey that are attended by those ants. These results suggest that adaptations for host ant location by ovipositing female miletines may have been retained from phytophagous ancestors that associated with ants mutualistically.
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biogeography and systematics of aricia butterflies lepidoptera Lycaenidae
Molecular Phylogenetics and Evolution, 2013Co-Authors: Claudia P Sanudorestrepo, Gerard Talavera, Vlad Dincă, Roger VilaAbstract:Butterflies of the Aricia species group represent a paradigm of unresolved taxonomy, both at the genus and species levels. We studied phylogenetic relationships, biogeography, and systematics based on genetic – nuclear and mitochondrial – and morphometric – external (wings) and internal (genitalia) – data. We show that Aricia is a monophyletic genus comprising the taxa Pseudoaricia, Ultraaricia and Umpria, which are here considered junior synonyms of Aricia. The taxa allous, inhonora, issekutzi, mandzhuriana, myrmecias and transalaica, which have often been raised to species rank, are shown to probably represent subspecies or synonyms. We show that montensis is likely a good species that is sister to all A. artaxerxes populations across the Palearctic region. The species A. anteros and A. morronensis are shown to display deep intraspecific divergences and they may harbor cryptic species. We also discovered that A. cramera and A. agestis exhibit a pattern of mutual exclusion on islands, and a parapatric distribution in mainland with a narrow contact zone where potential hybrids were detected. The lack of a prezygotic barrier that prevents their coexistence could explain this phenomenon. This study will hopefully contribute to the stability of the systematics of Aricia, a group with potential for the study of the link between speciation and biogeography.
Felipe Gilt - One of the best experts on this subject based on the ideXlab platform.
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estudio sobre la influencia de parasitoides hymenoptera ichneumonoidea en poblaciones del raro lepidoptero iolana iolas ochsenheimer 1816 lepidoptera Lycaenidae
2001Co-Authors: Felipe GiltAbstract:C/ Corr. Veleta, 1, 2-A, E-18008 Granada. Resumen: En este trabajo se analizan los datos obtenidos en el muestreo realizado durante tres anos sobre la existencia de varias especies de parasitoides Ichneumonoidea en larvas y pupas del muy localizado y escaso lepidoptero Lycaenidae Iolana iolas Ochsenheimer, 1816 y la influencia del parasitismo en las poblaciones de imagos adultos en posteriores generaciones. De uno de los parasitoides estudiados, Anisobas cephalotes (Kriechbaumer, 1882), citado por primera vez para Espana en SELFA & BORDERA (1995: 473), especie muy interesante dada la escasez de especimenes conocidos (4) hasta ahora, se da la segunda cita en este trabajo, ademas de informacion sobre nuevas localidades, total de nuevos especimenes obtenidos y descripcion de su biologia, siendo las provincias de Granada y Madrid los unicos lugares conocidos con registros de este parasitoide en la peninsula Iberica.
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estudio sobre la influencia de parasitoides hymenoptera ichneumonoidea en poblaciones del raro lepidoptero iolana iolas ochsenheimer 1816 lepidoptera Lycaenidae
2001Co-Authors: Felipe GiltAbstract:C/ Corr. Veleta, 1, 2-A, E-18008 Granada. Resumen: En este trabajo se analizan los datos obtenidos en el muestreo realizado durante tres anos sobre la existencia de varias especies de parasitoides Ichneumonoidea en larvas y pupas del muy localizado y escaso lepidoptero Lycaenidae Iolana iolas Ochsenheimer, 1816 y la influencia del parasitismo en las poblaciones de imagos adultos en posteriores generaciones. De uno de los parasitoides estudiados, Anisobas cephalotes (Kriechbaumer, 1882), citado por primera vez para Espana en SELFA & BORDERA (1995: 473), especie muy interesante dada la escasez de especimenes conocidos (4) hasta ahora, se da la segunda cita en este trabajo, ademas de informacion sobre nuevas localidades, total de nuevos especimenes obtenidos y descripcion de su biologia, siendo las provincias de Granada y Madrid los unicos lugares conocidos con registros de este parasitoide en la peninsula Iberica.
Freitas, André Victor Lucci - One of the best experts on this subject based on the ideXlab platform.
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Butterflies collected using malaise traps as useful bycatches for ecology and conservation
'Wildlife Information Liaison Development Society', 2020Co-Authors: Rosa, Augusto Henrique Batista, Perillo, Lucas Neves, Neves, Frederico Siqueira, Ribeiro, Danilo Bandini, Freitas, André Victor LucciAbstract:From 2013 to 2016, a study using malaise was carried out throughout the Espinhaço Mountain range in the Brazilian states of Minas Gerais and Bahia. A total of 1,000 butterflies belonging to six families, namely Pieridae (n=353), Nymphalidae (n=274), Hesperiidae (n=205), Lycaenidae (n=139), Riodinidae (n=17), and Papilionidae (n=12), were captured during this period. Because of the collecting method, most butterflies became mangled and resulted in faded colours, making them difficult to be identified. Nevertheless, three interesting species were identified in the collected material as they were either threatened, endemic, or undescribed. These new records highlight the importance of storing and making available all collected material, including bycatches, since this can provide important information for studies on ecology, taxonomy, and conservation11
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As borboletas (Lepidoptera: Papilionoidea e Hesperioidea) do Parque Ecológico João Vasconcelos Sobrinho, Caruaru, Pernambuco, Brasil
Instituto Virtual da Biodiversidade | BIOTA - FAPESP, 2015Co-Authors: Paluch Márlon, Mielke, Olaf Hermann Hendrik, Nobre, Carlos Eduardo Beserra, Casagrande, Mirna Martins, Melo, Douglas Henrique Alves, Freitas, André Victor LucciAbstract:Comprising a natural reserve with 359 ha of montane forest inserted on the Brazilian semi-arid, the Parque Ecológico João Vasconcelos Sobrinho (PEJVS), locally known as Brejo dos Cavalos is currently under high anthropogenic pressure. A list of 197 species of butterflies belonging to six families is presented, being 59 species of Hesperiidae, 4 of Papilionidae, 18 of Pieridae, 17 of Lycaenidae, 12 of Riodinidae and 87 of Nymphalidae. The butterfly community was composed mainly by widespread species commonly found in open habitats. There were also many species typical of forested areas such as Scada karschina delicata Talbot, 1932 (Danainae: Ithomiini), which is an endangered butterfly.O Parque Ecológico João Vasconcelos Sobrinho (PEJVS), conhecido localmente como Brejo dos Cavalos, é um fragmento de mata serrana de 359 ha inserido no semi-árido brasileiro, e que atualmente encontra-se sobre alta pressão antrópica. Uma lista com 197 espécies de borboletas pertencentes a seis famílias é apresentada, sendo 59 espécies de Hesperiidae, 4 de Papilionidae, 18 de Pieridae, 17 de Lycaenidae, 12 de Riodinidae e 87 de Nymphalidae. A fauna é composta principalmente de espécies de áreas abertas e com ampla distribuição geográfica. Entretanto, diversas espécies típicas de áreas florestadas também estão presentes, uma delas, Scada karschina delicata Talbot, 1932 (Danainae: Ithomiini), é uma espécie de borboleta criticamente ameaçada de extinção
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As borboletas (Lepidoptera: Papilionoidea e Hesperioidea) do Parque Ecológico João Vasconcelos Sobrinho, Caruaru, Pernambuco, Brasil
'FapUNIFESP (SciELO)', 2015Co-Authors: Paluch Márlon, Mielke, Olaf Hermann Hendrik, Nobre, Carlos Eduardo Beserra, Casagrande, Mirna Martins, Melo, Douglas Henrique Alves, Freitas, André Victor LucciAbstract:Comprising a natural reserve with 359 ha of montane forest inserted on the Brazilian semi-arid, the Parque Ecológico João Vasconcelos Sobrinho (PEJVS), locally known as Brejo dos Cavalos is currently under high anthropogenic pressure. A list of 197 species of butterflies belonging to six families is presented, being 59 species of Hesperiidae, 4 of Papilionidae, 18 of Pieridae, 17 of Lycaenidae, 12 of Riodinidae and 87 of Nymphalidae. The butterfly community was composed mainly by widespread species commonly found in open habitats. There were also many species typical of forested areas such as Scada karschina delicata Talbot, 1932 (Danainae: Ithomiini), which is an endangered butterfly.O Parque Ecológico João Vasconcelos Sobrinho (PEJVS), conhecido localmente como Brejo dos Cavalos, é um fragmento de mata serrana de 359 ha inserido no semi-árido brasileiro, e que atualmente encontra-se sobre alta pressão antrópica. Uma lista com 197 espécies de borboletas pertencentes a seis famílias é apresentada, sendo 59 espécies de Hesperiidae, 4 de Papilionidae, 18 de Pieridae, 17 de Lycaenidae, 12 de Riodinidae e 87 de Nymphalidae. A fauna é composta principalmente de espécies de áreas abertas e com ampla distribuição geográfica. Entretanto, diversas espécies típicas de áreas florestadas também estão presentes, uma delas, Scada karschina delicata Talbot, 1932 (Danainae: Ithomiini), é uma espécie de borboleta criticamente ameaçada de extinção.229238Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq
Alan Heath - One of the best experts on this subject based on the ideXlab platform.
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recent diversification of chrysoritis butterflies in the south african cape lepidoptera Lycaenidae
Molecular Phylogenetics and Evolution, 2020Co-Authors: Gerard Talavera, Alan Heath, Zofia A Kaliszewska, Naomi E PierceAbstract:Abstract Although best known for its extraordinary radiations of endemic plant species, the South African fynbos is home to a great diversity of phytophagous insects, including butterflies in the genus Chrysoritis (Lepidoptera: Lycaenidae). These butterflies are remarkably uniform morphologically; nevertheless, they comprise 43 currently accepted species and 68 currently valid taxonomic names. While many species have highly restricted, dot-like distributions, others are widespread. Here, we investigate the phylogenetic and biogeographic history underlying their diversification by analyzing molecular markers from 406 representatives of all described species throughout their respective ranges. We recover monophyletic clades for both C. chrysaor and C. thysbe species-groups, and identify a set of lineages that fall between them. The estimated age of divergence for the genus is 32 Mya, and we document significantly rapid diversification of the thysbe species-group in the Pleistocene (~2 Mya). Using ancestral geographic range reconstruction, we show that West Fynbos is the most likely region of origin for the radiation of the thysbe species-group. The colonization of this region occurred 9 Mya and appears to have been followed by a long period of relative stasis before a recent increase in diversification. Thus, the thysbe radiation does not appear to have resulted from the colonization of new biogeographic areas. Rather, the impact of species interactions (with ants and plants), the appearance of key innovations, and/or the opening of new ecological niche space in the region might explain the sudden burst of speciation that occurred in this group 2 Mya. The biogeographic model suggests two different diversification processes with few historical cross-colonisations, one in eastern South Africa for the C. chrysaor group and the other in western South Africa for the remaining taxa. Distributional range assessments and ecological niche models for each species show important niche overlap, and in a few cases, complete overlap. However, these shared traits are not explained by phylogenetic history. Chrysoritis taxa frequently fly in sympatry and gene tree reticulation appears to be widespread at the species level, suggesting that several episodes of range shifts might have led to secondary sympatries, allowing limited gene flow that challenges species delimitation efforts. In addition, the unusually high diversification rate for the thysbe clade of 1.35 [0.91–1.81] lineages per million years also suggests the possibility of taxonomic oversplitting. The phylogeny presented here provides a framework for a taxonomic revision of the genus. We highlight cases of potential synonymy both in allopatry and sympatry, and stress the importance of dedicated studies to assess potential pre- and post-zygotic barriers giving rise to species delimitations of the thysbe group.
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the ecology and evolution of ant association in the Lycaenidae lepidoptera
Annual Review of Entomology, 2002Co-Authors: Naomi E Pierce, Michael F Braby, Alan Heath, David J Lohman, John Mathew, Douglas B Rand, Mark A TravassosAbstract:The estimated 6000 species of Lycaenidae account for about one third of all Papilionoidea. The majority of lycaenids have associations with ants that can be facultative or obligate and range from mutualism to parasitism. Lycaenid larvae and pupae employ complex chemical and acoustical signals to manipulate ants. Cost/benefit analyses have demonstrated multiple trade-offs involved in myrmecophily. Both demographic and phylogenetic evidence indicate that ant association has shaped the evolution of obligately associated groups. Parasitism typically arises from mutualism with ants, and entomophagous species are disproportionately common in the Lycaenidae compared with other Lepidoptera. Obligate associations are more common in the Southern Hemisphere, in part because highly ant-associated lineages make up a larger proportion of the fauna in these regions. Further research on phylogeny and natural history, particularly of the Neotropical fauna, will be necessary to understand the role ant association has played in the evolution of the Lycaenidae.
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phylogeny and life history evolution of the genus chrysoritis within the aphnaeini lepidoptera Lycaenidae inferred from mitochondrial cytochrome oxidase i sequences
Molecular Phylogenetics and Evolution, 2000Co-Authors: Douglas B Rand, Alan Heath, Tamara Suderman, Naomi E PierceAbstract:Phylogenetic relationships among 26 South African species in the tribe Aphnaeini (Lepidoptera: Lycaenidae) were inferred from DNA characters of the mitochondrial gene cytochrome oxidase I (COI), using maximum-parsimony methods. The resulting phylogenetic estimate supports the systematic hypothesis made by Heath (1997, Metamorphosis, supplement 2), based on morphological characters, that at least three preexisting genera (Chrysoritis, Poecilmitis, and Oxychaeta) should be collapsed into the single monophyletic genus Chrysoritis. Two of the species groups described by Heath within Chrysoritis are also monophyletic, while one is paraphyletic and thus unsupported by the molecular data. Strong node support and skewed transition/transversion ratios suggest that two Chrysoritis clades contain synonymous species. Aphytophagy appears as a derived feeding strategy. Evolutionary patterns of ant association indicate lability at the level of ant genus, while association with different ant subfamilies may have played an ancestral and chemically mediated role in the diversification of South African aphnaeines.
