Insects have developed various adaptations to survive harsh winter conditions. Among freeze-intol... more Insects have developed various adaptations to survive harsh winter conditions. Among freeze-intolerant species, some produce “antifreeze proteins” (AFPs) that bind to nascent ice crystals and inhibit further ice growth. Such is the case of the spruce budworm, Choristoneura fumiferana (Lepidoptera: Tortricidae), a destructive North American conifer pest that can withstand temperatures below −30°C. Despite the potential importance of AFPs in the adaptive diversification of Choristoneura, genomic tools to explore their origins have until now been limited. Here, we present a chromosome-scale genome assembly for C. fumiferana, which we used to conduct comparative genomic analyses aimed at reconstructing the evolutionary history of tortricid AFPs. The budworm genome features 16 genes homologous to previously reported C. fumiferana AFPs (CfAFPs), 15 of which map to a single region on chromosome 18. Fourteen of these were also detected in five congeneric species, indicating Choristoneura AF...
Insects have developed various adaptations to survive harsh winter conditions. Among freeze-intol... more Insects have developed various adaptations to survive harsh winter conditions. Among freeze-intolerant species, some produce “antifreeze proteins” (AFPs) that bind to nascent ice crystals and inhibit further ice growth. Such is the case of the spruce budworm, Choristoneura fumiferana (Lepidoptera: Tortricidae), a destructive North American conifer pest that can withstand temperatures below −30°C. Despite the potential importance of AFPs in the adaptive diversification of Choristoneura, genomic tools to explore their origins have until now been limited. Here, we present a chromosome-scale genome assembly for C. fumiferana, which we used to conduct comparative genomic analyses aimed at reconstructing the evolutionary history of tortricid AFPs. The budworm genome features 16 genes homologous to previously reported C. fumiferana AFPs (CfAFPs), 15 of which map to a single region on chromosome 18. Fourteen of these were also detected in five congeneric species, indicating Choristoneura AF...
The mountain pine beetle (Dendroctonus ponderosae Hopkins) (Coleoptera: Curculionidae) is a signi... more The mountain pine beetle (Dendroctonus ponderosae Hopkins) (Coleoptera: Curculionidae) is a significant destructive force in the pine forests of western Canada and has the capacity to spread east into a novel host tree species, jack pine (Pinaceae). New populations have been documented in central Alberta, Canada, but the source populations for these outbreaks have yet to be identified. In this study, we use genetic data to identify parent populations for recent outbreak sites near Slave Lake, Lac La Biche, and Hinton, Alberta. We found the northern population cluster that entered Alberta near Grande Prairie was the source of the most eastern established population near Lac La Biche, and the range expansion to this leading-edge population has been too rapid to establish evidence of population structure. However, some dispersal from a population in the Jasper and Hinton area has been detected as far north and east as Slave Lake, Alberta. We also identified two potential source populat...
We investigated the population structure of Grosmannia clavigera (Gc), a fungal symbiont of the m... more We investigated the population structure of Grosmannia clavigera (Gc), a fungal symbiont of the mountain pine beetle (MPB) that plays a crucial role in the establishment and reproductive success of this pathogen. This insect–fungal complex has destroyed over 16 million ha of lodgepole pine forests in Canada, the largest MPB epidemic in recorded history. During this current epidemic, MPB has expanded its range beyond historically recorded boundaries, both northward and eastward, and has now reached the jack pine of Alberta, potentially threatening the Canadian boreal forest. To better understand the dynamics between the beetle and its fungal symbiont, we sampled 19 populations in western North America and genotyped individuals from these populations with eight microsatellite markers. The fungus displayed high haplotype diversity, with over 250 unique haplotypes observed in 335 single spore isolates. Linkage equilibria in 13 of the 19 populations suggested that the fungus reproduces s...
The swallowtail genus Papilio (Lepidoptera: Papilionidae) is species rich, distributed worldwide,... more The swallowtail genus Papilio (Lepidoptera: Papilionidae) is species rich, distributed worldwide, and has broad morphological habits and ecological niches. Because of its elevated species richness, it has been historically difficult to reconstruct a densely sampled phylogeny for this clade. Here we provide a taxonomic working list for the genus, resulting in 235 Papilio species, and assemble a molecular dataset of seven gene fragments representing ca. 80% of the currently described diversity. Phylogenetic analyses reconstructed a robust tree with highly supported relationships within subgenera, although a few nodes in the early history of the Old World Papilio remain unresolved. Contrasting with previous results, we found that Papilio alexanor is sister to all Old World Papilio and that the subgenus Eleppone is no longer monotypic. The latter includes the recently described Fijian Papilio natewa with the Australian Papilio anactus and is sister to subgenus Araminta (formerly included in subgenus Menelaides) occurring in Southeast Asia. Our phylogeny also includes rarely studied (P. antimachus, P. benguetana) or endangered species (P. buddha, P. chikae). Taxonomic changes resulting from this study are elucidated. Molecular dating and biogeographic analyses indicate that Papilio originated ca. 30 million years ago (Oligocene), in a northern region centered on Beringia. A rapid early Miocene radiation in the Paleotropics is revealed within Old World Papilio, potentially explaining their low early branch support. Most subgenera originated in the early to middle Miocene followed by synchronous southward biogeographic dispersals and repeated local extirpations in northern latitudes. This study provides a comprehensive phylogenetic framework for Papilio with clarification of subgeneric systematics and species taxonomic changes enumerated, which will facilitate further studies to address questions on their ecology and evolutionary biology using this model clade.
The swallowtail genus Papilio (Lepidoptera: Papilionidae) is species rich, distributed worldwide,... more The swallowtail genus Papilio (Lepidoptera: Papilionidae) is species rich, distributed worldwide, and has broad morphological habits and ecological niches. Because of its elevated species richness, it has been historically difficult to reconstruct a densely sampled phylogeny for this clade. Here we provide a taxonomic working list for the genus, resulting in 235 Papilio species, and assemble a molecular dataset of seven gene fragments representing ca. 80% of the currently described diversity. Phylogenetic analyses reconstructed a robust tree with highly supported relationships within subgenera, although a few nodes in the early history of the Old World Papilio remain unresolved. Contrasting with previous results, we found that Papilio alexanor is sister to all Old World Papilio and that the subgenus Eleppone is no longer monotypic. The latter includes the recently described Fijian Papilio natewa with the Australian Papilio anactus and is sister to subgenus Araminta (formerly included in subgenus Menelaides) occurring in Southeast Asia. Our phylogeny also includes rarely studied (P. antimachus, P. benguetana) or endangered species (P. buddha, P. chikae). Taxonomic changes resulting from this study are elucidated. Molecular dating and biogeographic analyses indicate that Papilio originated ca. 30 million years ago (Oligocene), in a northern region centered on Beringia. A rapid early Miocene radiation in the Paleotropics is revealed within Old World Papilio, potentially explaining their low early branch support. Most subgenera originated in the early to middle Miocene followed by synchronous southward biogeographic dispersals and repeated local extirpations in northern latitudes. This study provides a comprehensive phylogenetic framework for Papilio with clarification of subgeneric systematics and species taxonomic changes enumerated, which will facilitate further studies to address questions on their ecology and evolutionary biology using this model clade.
The exuberant proliferation of herbivorous insects is attributed to their associations with plant... more The exuberant proliferation of herbivorous insects is attributed to their associations with plants. Despite abundant studies on insect-plant interactions, we do not know whether host-plant shifts have impacted both genomic adaptation and species diversification over geological times. We show that the antagonistic insect-plant interaction between swallowtail butterflies and the highly toxic birthworts began 55 million years ago in Beringia, followed by several major ancient host-plant shifts. This evolutionary framework provides a unique opportunity for repeated tests of genomic signatures of macroevolutionary changes and estimation of diversification rates across their phylogeny. We find that host-plant shifts in butterflies are associated with both genome-wide adaptive molecular evolution (more genes under positive selection) and repeated bursts of speciation rates, contributing to an increase in global diversification through time. Our study links ecological changes, genome-wide a...
In macroevolution, the Red Queen (RQ) model posits that biodiversity dynamics depend mainly on sp... more In macroevolution, the Red Queen (RQ) model posits that biodiversity dynamics depend mainly on species-intrinsic biotic factors such as interactions among species or life-history traits, while the Court Jester (CJ) model states that extrinsic environmental abiotic factors have a stronger role. Until recently, a lack of relevant methodological approaches has prevented the unraveling of contributions from these two types of factors to the evolutionary history of a lineage. Here we take advantage of the rapid development of new macroevolution models that tie diversification rates to changes in paleoenvironmental (extrinsic) and/or biotic (intrinsic) factors. We inferred a robust and fully-sampled species-level phylogeny, as well as divergence times and ancestral geographic ranges, and related these to the radiation of Apollo butterflies (Parnassiinae) using both extant (molecular) and extinct (fossil/morphological) evidence. We tested whether their diversification dynamics are better e...
Publisher Summary This chapter discusses various entomology-teaching resources. Those who teach a... more Publisher Summary This chapter discusses various entomology-teaching resources. Those who teach about insects in the entomological sense, the best resources have always been a good entomological library, a selection of live insects in culture, a well-maintained collection of preserved specimens, and somewhere to take students to see live insects behaving in a natural fashion. These are still the basics, along with contagious enthusiasm and genuine personal knowledge of the subject. In recent years, two changes in social attitudes have strongly affected entomological teaching. Websites are the most important recent development in entomological teaching, and take various forms, such as clearinghouses to other sites, museum databases, exercises for instruction at various levels, and taxon-based compilations and/or species accounts. Internet is a wonderful tool, but the library is still the rightful place for scholarly information in entomology. Books and journals are for the most part much more carefully written, reviewed, and designed than their web-based counterparts, and therefore they are the guardians of mainstream entomological knowledge. Additionally, a number of excellent television programs, as well as regular series, have featured insects as their subject matter. Some of these are available on videotape or DVD. In general, video narration is not peer-reviewed, and is notorious for inaccuracies. Nonetheless, the value of video in education is difficult to overestimate.
The bacterial microbiome of ticks is notoriously diverse, but the factors leading to this diversi... more The bacterial microbiome of ticks is notoriously diverse, but the factors leading to this diversity are poorly understood. We sequenced bacterial 16S rRNA amplicons from individual winter ticks, Dermacentor albipictus, to assess whether their one-host life cycle is associated with reduced bacterial diversity. On average, about 100 bacterial genera were found for individual ticks. Francisella-like endosymbiont (FLE) dominated bacterial communities, particularly in female ticks and in ticks that had fed. The remainder of the winter tick microbiome was highly variable. In addition to FLE, the main bacterial genera associated with winter ticks on elk were Pseudomonas, Ehrlichia, Asinibacterium, Acinetobacter and Streptococcus, although sequences associated with hundreds of other minor bacterial genera were detected. A complex interaction between richness and evenness was revealed in comparisons among tick life stages, using the Hill number series to show trends in diversity with decreasing emphasis on rare members of the assemblage. Male ticks had a significantly greater number of bacterial genera than females or nymphs, while males had greater evenness than females and similar evenness to nymphs. We intentionally sampled ticks from a single host species, North American elk, from a single location in Alberta, Canada, to constrain the ecological and blood meal variation that individuals experience through their life cycle. In spite of this, we found that the number of bacterial genera detected in this one-host tick system was remarkably diverse. The high taxonomic variability of the minor components of the winter tick microbiome suggests that this part of their microbiome diversity should be examined for functional significance.
Patterns in the genetic variation of species can be used to infer their specific demographic and ... more Patterns in the genetic variation of species can be used to infer their specific demographic and evolutionary history and provide insight into the general mechanisms underlying population divergence and speciation. The Macoun’s Arctic (Oeneis macounii (W.H. Edwards, 1885); MA) butterfly occurs across Canada and parts of the northern United States in association with jack pine (Pinus banksiana Lamb.) and lodgepole pine (Pinus contorta Douglas ex Loudon). MA’s current distribution is highly fragmented, and the extent of reproductive isolation among allopatric populations is unknown. Furthermore, although MA is biennial, adults emerge every year in some populations. These populations presumably consist of two alternate-year cohorts, providing the opportunity for sympatric divergence via allochronic isolation. Using mitochondrial DNA (mtDNA) and amplified fragment length polymorphism (AFLP) markers, we analyzed MA’s genetic structure to determine the current and historical role of allop...
ABSTRACT Numerous hypotheses on the evolution of Neotropical biodiversity have stimulated researc... more ABSTRACT Numerous hypotheses on the evolution of Neotropical biodiversity have stimulated research to provide a better understanding of diversity dynamics and distribution patterns of the region. However, few studies integrate molecular and morphological data with complete sampling of a Neotropical group, and so there has been little synthesis of the multiple processes governing biodiversity through space and time. Here, a total-evidence phylogenetic approach is used to reconstruct the evolutionary history of the butterfly subgenus Heraclides. We used DNA sequences for two mitochondrial genes and one nuclear gene and coded 133 morphological characters of larvae and adults. A robust and well-resolved phylogeny was obtained using several analytical approaches, while molecular dating and biogeographical analyses indicated an early Miocene origin (22 Mya) in the Caribbean Islands. We inferred six independent dispersal events from the Caribbean to the mainland, and three from the mainland to the Caribbean, and we suggest that cooling climates with decreasing sea levels may have contributed to these events. The time-calibrated tree is best explained by a museum model of diversity in which both speciation and extinction rates remained constant through time. By assessing both continental and fine-scale biodiversity patterns, this study provides new findings, for instance that islands may act as source of diversity rather than as a sink, to explain spatio-temporal macroevolutionary processes within the Neotropical region.
Insects have developed various adaptations to survive harsh winter conditions. Among freeze-intol... more Insects have developed various adaptations to survive harsh winter conditions. Among freeze-intolerant species, some produce “antifreeze proteins” (AFPs) that bind to nascent ice crystals and inhibit further ice growth. Such is the case of the spruce budworm, Choristoneura fumiferana (Lepidoptera: Tortricidae), a destructive North American conifer pest that can withstand temperatures below −30°C. Despite the potential importance of AFPs in the adaptive diversification of Choristoneura, genomic tools to explore their origins have until now been limited. Here, we present a chromosome-scale genome assembly for C. fumiferana, which we used to conduct comparative genomic analyses aimed at reconstructing the evolutionary history of tortricid AFPs. The budworm genome features 16 genes homologous to previously reported C. fumiferana AFPs (CfAFPs), 15 of which map to a single region on chromosome 18. Fourteen of these were also detected in five congeneric species, indicating Choristoneura AF...
Insects have developed various adaptations to survive harsh winter conditions. Among freeze-intol... more Insects have developed various adaptations to survive harsh winter conditions. Among freeze-intolerant species, some produce “antifreeze proteins” (AFPs) that bind to nascent ice crystals and inhibit further ice growth. Such is the case of the spruce budworm, Choristoneura fumiferana (Lepidoptera: Tortricidae), a destructive North American conifer pest that can withstand temperatures below −30°C. Despite the potential importance of AFPs in the adaptive diversification of Choristoneura, genomic tools to explore their origins have until now been limited. Here, we present a chromosome-scale genome assembly for C. fumiferana, which we used to conduct comparative genomic analyses aimed at reconstructing the evolutionary history of tortricid AFPs. The budworm genome features 16 genes homologous to previously reported C. fumiferana AFPs (CfAFPs), 15 of which map to a single region on chromosome 18. Fourteen of these were also detected in five congeneric species, indicating Choristoneura AF...
The mountain pine beetle (Dendroctonus ponderosae Hopkins) (Coleoptera: Curculionidae) is a signi... more The mountain pine beetle (Dendroctonus ponderosae Hopkins) (Coleoptera: Curculionidae) is a significant destructive force in the pine forests of western Canada and has the capacity to spread east into a novel host tree species, jack pine (Pinaceae). New populations have been documented in central Alberta, Canada, but the source populations for these outbreaks have yet to be identified. In this study, we use genetic data to identify parent populations for recent outbreak sites near Slave Lake, Lac La Biche, and Hinton, Alberta. We found the northern population cluster that entered Alberta near Grande Prairie was the source of the most eastern established population near Lac La Biche, and the range expansion to this leading-edge population has been too rapid to establish evidence of population structure. However, some dispersal from a population in the Jasper and Hinton area has been detected as far north and east as Slave Lake, Alberta. We also identified two potential source populat...
We investigated the population structure of Grosmannia clavigera (Gc), a fungal symbiont of the m... more We investigated the population structure of Grosmannia clavigera (Gc), a fungal symbiont of the mountain pine beetle (MPB) that plays a crucial role in the establishment and reproductive success of this pathogen. This insect–fungal complex has destroyed over 16 million ha of lodgepole pine forests in Canada, the largest MPB epidemic in recorded history. During this current epidemic, MPB has expanded its range beyond historically recorded boundaries, both northward and eastward, and has now reached the jack pine of Alberta, potentially threatening the Canadian boreal forest. To better understand the dynamics between the beetle and its fungal symbiont, we sampled 19 populations in western North America and genotyped individuals from these populations with eight microsatellite markers. The fungus displayed high haplotype diversity, with over 250 unique haplotypes observed in 335 single spore isolates. Linkage equilibria in 13 of the 19 populations suggested that the fungus reproduces s...
The swallowtail genus Papilio (Lepidoptera: Papilionidae) is species rich, distributed worldwide,... more The swallowtail genus Papilio (Lepidoptera: Papilionidae) is species rich, distributed worldwide, and has broad morphological habits and ecological niches. Because of its elevated species richness, it has been historically difficult to reconstruct a densely sampled phylogeny for this clade. Here we provide a taxonomic working list for the genus, resulting in 235 Papilio species, and assemble a molecular dataset of seven gene fragments representing ca. 80% of the currently described diversity. Phylogenetic analyses reconstructed a robust tree with highly supported relationships within subgenera, although a few nodes in the early history of the Old World Papilio remain unresolved. Contrasting with previous results, we found that Papilio alexanor is sister to all Old World Papilio and that the subgenus Eleppone is no longer monotypic. The latter includes the recently described Fijian Papilio natewa with the Australian Papilio anactus and is sister to subgenus Araminta (formerly included in subgenus Menelaides) occurring in Southeast Asia. Our phylogeny also includes rarely studied (P. antimachus, P. benguetana) or endangered species (P. buddha, P. chikae). Taxonomic changes resulting from this study are elucidated. Molecular dating and biogeographic analyses indicate that Papilio originated ca. 30 million years ago (Oligocene), in a northern region centered on Beringia. A rapid early Miocene radiation in the Paleotropics is revealed within Old World Papilio, potentially explaining their low early branch support. Most subgenera originated in the early to middle Miocene followed by synchronous southward biogeographic dispersals and repeated local extirpations in northern latitudes. This study provides a comprehensive phylogenetic framework for Papilio with clarification of subgeneric systematics and species taxonomic changes enumerated, which will facilitate further studies to address questions on their ecology and evolutionary biology using this model clade.
The swallowtail genus Papilio (Lepidoptera: Papilionidae) is species rich, distributed worldwide,... more The swallowtail genus Papilio (Lepidoptera: Papilionidae) is species rich, distributed worldwide, and has broad morphological habits and ecological niches. Because of its elevated species richness, it has been historically difficult to reconstruct a densely sampled phylogeny for this clade. Here we provide a taxonomic working list for the genus, resulting in 235 Papilio species, and assemble a molecular dataset of seven gene fragments representing ca. 80% of the currently described diversity. Phylogenetic analyses reconstructed a robust tree with highly supported relationships within subgenera, although a few nodes in the early history of the Old World Papilio remain unresolved. Contrasting with previous results, we found that Papilio alexanor is sister to all Old World Papilio and that the subgenus Eleppone is no longer monotypic. The latter includes the recently described Fijian Papilio natewa with the Australian Papilio anactus and is sister to subgenus Araminta (formerly included in subgenus Menelaides) occurring in Southeast Asia. Our phylogeny also includes rarely studied (P. antimachus, P. benguetana) or endangered species (P. buddha, P. chikae). Taxonomic changes resulting from this study are elucidated. Molecular dating and biogeographic analyses indicate that Papilio originated ca. 30 million years ago (Oligocene), in a northern region centered on Beringia. A rapid early Miocene radiation in the Paleotropics is revealed within Old World Papilio, potentially explaining their low early branch support. Most subgenera originated in the early to middle Miocene followed by synchronous southward biogeographic dispersals and repeated local extirpations in northern latitudes. This study provides a comprehensive phylogenetic framework for Papilio with clarification of subgeneric systematics and species taxonomic changes enumerated, which will facilitate further studies to address questions on their ecology and evolutionary biology using this model clade.
The exuberant proliferation of herbivorous insects is attributed to their associations with plant... more The exuberant proliferation of herbivorous insects is attributed to their associations with plants. Despite abundant studies on insect-plant interactions, we do not know whether host-plant shifts have impacted both genomic adaptation and species diversification over geological times. We show that the antagonistic insect-plant interaction between swallowtail butterflies and the highly toxic birthworts began 55 million years ago in Beringia, followed by several major ancient host-plant shifts. This evolutionary framework provides a unique opportunity for repeated tests of genomic signatures of macroevolutionary changes and estimation of diversification rates across their phylogeny. We find that host-plant shifts in butterflies are associated with both genome-wide adaptive molecular evolution (more genes under positive selection) and repeated bursts of speciation rates, contributing to an increase in global diversification through time. Our study links ecological changes, genome-wide a...
In macroevolution, the Red Queen (RQ) model posits that biodiversity dynamics depend mainly on sp... more In macroevolution, the Red Queen (RQ) model posits that biodiversity dynamics depend mainly on species-intrinsic biotic factors such as interactions among species or life-history traits, while the Court Jester (CJ) model states that extrinsic environmental abiotic factors have a stronger role. Until recently, a lack of relevant methodological approaches has prevented the unraveling of contributions from these two types of factors to the evolutionary history of a lineage. Here we take advantage of the rapid development of new macroevolution models that tie diversification rates to changes in paleoenvironmental (extrinsic) and/or biotic (intrinsic) factors. We inferred a robust and fully-sampled species-level phylogeny, as well as divergence times and ancestral geographic ranges, and related these to the radiation of Apollo butterflies (Parnassiinae) using both extant (molecular) and extinct (fossil/morphological) evidence. We tested whether their diversification dynamics are better e...
Publisher Summary This chapter discusses various entomology-teaching resources. Those who teach a... more Publisher Summary This chapter discusses various entomology-teaching resources. Those who teach about insects in the entomological sense, the best resources have always been a good entomological library, a selection of live insects in culture, a well-maintained collection of preserved specimens, and somewhere to take students to see live insects behaving in a natural fashion. These are still the basics, along with contagious enthusiasm and genuine personal knowledge of the subject. In recent years, two changes in social attitudes have strongly affected entomological teaching. Websites are the most important recent development in entomological teaching, and take various forms, such as clearinghouses to other sites, museum databases, exercises for instruction at various levels, and taxon-based compilations and/or species accounts. Internet is a wonderful tool, but the library is still the rightful place for scholarly information in entomology. Books and journals are for the most part much more carefully written, reviewed, and designed than their web-based counterparts, and therefore they are the guardians of mainstream entomological knowledge. Additionally, a number of excellent television programs, as well as regular series, have featured insects as their subject matter. Some of these are available on videotape or DVD. In general, video narration is not peer-reviewed, and is notorious for inaccuracies. Nonetheless, the value of video in education is difficult to overestimate.
The bacterial microbiome of ticks is notoriously diverse, but the factors leading to this diversi... more The bacterial microbiome of ticks is notoriously diverse, but the factors leading to this diversity are poorly understood. We sequenced bacterial 16S rRNA amplicons from individual winter ticks, Dermacentor albipictus, to assess whether their one-host life cycle is associated with reduced bacterial diversity. On average, about 100 bacterial genera were found for individual ticks. Francisella-like endosymbiont (FLE) dominated bacterial communities, particularly in female ticks and in ticks that had fed. The remainder of the winter tick microbiome was highly variable. In addition to FLE, the main bacterial genera associated with winter ticks on elk were Pseudomonas, Ehrlichia, Asinibacterium, Acinetobacter and Streptococcus, although sequences associated with hundreds of other minor bacterial genera were detected. A complex interaction between richness and evenness was revealed in comparisons among tick life stages, using the Hill number series to show trends in diversity with decreasing emphasis on rare members of the assemblage. Male ticks had a significantly greater number of bacterial genera than females or nymphs, while males had greater evenness than females and similar evenness to nymphs. We intentionally sampled ticks from a single host species, North American elk, from a single location in Alberta, Canada, to constrain the ecological and blood meal variation that individuals experience through their life cycle. In spite of this, we found that the number of bacterial genera detected in this one-host tick system was remarkably diverse. The high taxonomic variability of the minor components of the winter tick microbiome suggests that this part of their microbiome diversity should be examined for functional significance.
Patterns in the genetic variation of species can be used to infer their specific demographic and ... more Patterns in the genetic variation of species can be used to infer their specific demographic and evolutionary history and provide insight into the general mechanisms underlying population divergence and speciation. The Macoun’s Arctic (Oeneis macounii (W.H. Edwards, 1885); MA) butterfly occurs across Canada and parts of the northern United States in association with jack pine (Pinus banksiana Lamb.) and lodgepole pine (Pinus contorta Douglas ex Loudon). MA’s current distribution is highly fragmented, and the extent of reproductive isolation among allopatric populations is unknown. Furthermore, although MA is biennial, adults emerge every year in some populations. These populations presumably consist of two alternate-year cohorts, providing the opportunity for sympatric divergence via allochronic isolation. Using mitochondrial DNA (mtDNA) and amplified fragment length polymorphism (AFLP) markers, we analyzed MA’s genetic structure to determine the current and historical role of allop...
ABSTRACT Numerous hypotheses on the evolution of Neotropical biodiversity have stimulated researc... more ABSTRACT Numerous hypotheses on the evolution of Neotropical biodiversity have stimulated research to provide a better understanding of diversity dynamics and distribution patterns of the region. However, few studies integrate molecular and morphological data with complete sampling of a Neotropical group, and so there has been little synthesis of the multiple processes governing biodiversity through space and time. Here, a total-evidence phylogenetic approach is used to reconstruct the evolutionary history of the butterfly subgenus Heraclides. We used DNA sequences for two mitochondrial genes and one nuclear gene and coded 133 morphological characters of larvae and adults. A robust and well-resolved phylogeny was obtained using several analytical approaches, while molecular dating and biogeographical analyses indicated an early Miocene origin (22 Mya) in the Caribbean Islands. We inferred six independent dispersal events from the Caribbean to the mainland, and three from the mainland to the Caribbean, and we suggest that cooling climates with decreasing sea levels may have contributed to these events. The time-calibrated tree is best explained by a museum model of diversity in which both speciation and extinction rates remained constant through time. By assessing both continental and fine-scale biodiversity patterns, this study provides new findings, for instance that islands may act as source of diversity rather than as a sink, to explain spatio-temporal macroevolutionary processes within the Neotropical region.
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Papers by Felix Sperling
morphological habits and ecological niches. Because of its elevated species richness, it has been historically
difficult to reconstruct a densely sampled phylogeny for this clade. Here we provide a taxonomic working list for
the genus, resulting in 235 Papilio species, and assemble a molecular dataset of seven gene fragments representing
ca. 80% of the currently described diversity. Phylogenetic analyses reconstructed a robust tree with
highly supported relationships within subgenera, although a few nodes in the early history of the Old World
Papilio remain unresolved. Contrasting with previous results, we found that Papilio alexanor is sister to all Old
World Papilio and that the subgenus Eleppone is no longer monotypic. The latter includes the recently described
Fijian Papilio natewa with the Australian Papilio anactus and is sister to subgenus Araminta (formerly included in
subgenus Menelaides) occurring in Southeast Asia. Our phylogeny also includes rarely studied (P. antimachus,
P. benguetana) or endangered species (P. buddha, P. chikae). Taxonomic changes resulting from this study are
elucidated. Molecular dating and biogeographic analyses indicate that Papilio originated ca. 30 million years ago
(Oligocene), in a northern region centered on Beringia. A rapid early Miocene radiation in the Paleotropics is
revealed within Old World Papilio, potentially explaining their low early branch support. Most subgenera originated
in the early to middle Miocene followed by synchronous southward biogeographic dispersals and repeated
local extirpations in northern latitudes. This study provides a comprehensive phylogenetic framework for Papilio
with clarification of subgeneric systematics and species taxonomic changes enumerated, which will facilitate
further studies to address questions on their ecology and evolutionary biology using this model clade.
morphological habits and ecological niches. Because of its elevated species richness, it has been historically
difficult to reconstruct a densely sampled phylogeny for this clade. Here we provide a taxonomic working list for
the genus, resulting in 235 Papilio species, and assemble a molecular dataset of seven gene fragments representing
ca. 80% of the currently described diversity. Phylogenetic analyses reconstructed a robust tree with
highly supported relationships within subgenera, although a few nodes in the early history of the Old World
Papilio remain unresolved. Contrasting with previous results, we found that Papilio alexanor is sister to all Old
World Papilio and that the subgenus Eleppone is no longer monotypic. The latter includes the recently described
Fijian Papilio natewa with the Australian Papilio anactus and is sister to subgenus Araminta (formerly included in
subgenus Menelaides) occurring in Southeast Asia. Our phylogeny also includes rarely studied (P. antimachus,
P. benguetana) or endangered species (P. buddha, P. chikae). Taxonomic changes resulting from this study are
elucidated. Molecular dating and biogeographic analyses indicate that Papilio originated ca. 30 million years ago
(Oligocene), in a northern region centered on Beringia. A rapid early Miocene radiation in the Paleotropics is
revealed within Old World Papilio, potentially explaining their low early branch support. Most subgenera originated
in the early to middle Miocene followed by synchronous southward biogeographic dispersals and repeated
local extirpations in northern latitudes. This study provides a comprehensive phylogenetic framework for Papilio
with clarification of subgeneric systematics and species taxonomic changes enumerated, which will facilitate
further studies to address questions on their ecology and evolutionary biology using this model clade.