The search for pattern in the ecology and evolutionary biology of insect-plant associations has fascinated biologists for centuries. High levels of tropical （tow-latitude） plant and insect diversity relative to pole...The search for pattern in the ecology and evolutionary biology of insect-plant associations has fascinated biologists for centuries. High levels of tropical （tow-latitude） plant and insect diversity relative to poleward latitudes and the disproportionate abundance of host-specialized insect herbivores have been noted. This review addresses several aspects of local insect specialization, host use abilities （and loss of these abilities with specialization）, host-associated evolutionary divergence, and ecological （including ＂hybrid＂） speciation, with special reference to the generation of biodiversity and the geographic and taxonomic identification of ＂species borders＂ for swallowtail butterflies （Papilionidae）. From ancient phytochemically defined angiosperm affiliations that trace back millions of years to recent and very local specialized populations, the Papilionidae （swallowtail butterflies） have provided a model for enhanced tmderstanding of localized ecological patterns and genetically based evolutionary processes. They have served as a useful group for evaluating the feeding specialization/physiological efficiency hypothesis. They have shown how the abiotic （thermal） environment interacts with host nutritional suitability to generate ＂voltinisrn/suitability＂ gradients in specialization or preference latitudinally, and geographical mosaics locally. Several studies reviewed here suggest strongly that the oscillation hypothesis for speciation does have considerable merit, but at the same time, some species-level host specializations may lead to evolutionary dead-ends, especially with rapid environmental/habitat changes involving their host plants. Latitudinal gradients in species richness and degree of herbivore feeding specialization have been impacted by recent developments in ecological genetics and evolutionary ecology. Localized insect- plant associations that span the biospectrum from polyphenisms, polymorphisms, biotypes, demes, host races, to cryptic species, remain academically contentious, with simple definitions still debated. However, molecular analyses combined with ecological, ethological and physiological studies, have already begun to unveil some answers for many important ecological/evolutionary questions.展开更多
The ecology and evolutionary biology of insect-plant associations has real- ized extensive attention, especially during the past 60 years. The classifications （categorical designations） of continuous variation in bi...The ecology and evolutionary biology of insect-plant associations has real- ized extensive attention, especially during the past 60 years. The classifications （categorical designations） of continuous variation in biodiversity, ranging from global patterns （e.g., lat- itudinal gradients in species richness/diversity and degree of herbivore feeding specialization） to localized insect-plant associations that span the biospectrum from polyphenisms, polymorphisms, biotypes, demes, host races, to cryptic species, remain academically contentious. Semantic and biosystematic （taxonomical） disagreements sometimes detract from more important ecological and evolutionary processes that drive diversification, the dynamics of gene flow and local extinctions. This review addresses several aspects of insect specialization, host-associated divergence and ecological （including ＂hybrid＂） speciation, with special reference to the climate warming impacts on species borders of hybridiz- ing swallowtail butterflies （Papilionidae）. Interspecific hybrid introgression may result in collapse of multi-species communities or increase species numbers via homoploid hybrid speciation. We may see diverging, merging, or emerging genotypes across hybrid zones, all part of the ongoing processes of evolution. Molecular analyses of genetic mosaics and genomic dynamics with ＂divergence hitchhiking＂, combined with ecological, ethological and physiological studies of＂species porosity＂, have already begun to unveil some answers for some important ecological/evolutionary questions. （i） How rapidly can host-associated divergence lead to new species （and why doesn＇t it always do so, e.g., resulting in ＂incom- plete＂ speciation）？ （ii） How might ＂speciation genes＂ function, and how/where would we find them？ （iii） Can oscillations from specialists to generalists and back to specialists help explain global diversity in herbivorous insects？ （iv） How could recombinant interspecific hybridization lead to divergence and speciation？ From ancient phytochemically defined angiosperm affiliations to recent and very local geographical mosaics, the Papilionidae （swallowtail butterflies） have provided a model for enhanced understanding of ecological patterns and evolutionary processes, including host-associated genetic divergence, ge- nomic mosaics, genetic hitchhiking and sex-linked speciation genes. Apparent homoploid hybrid speciation in Papilio appears to have been catalyzed by climate warming-induced interspecific introgression of some, but not all, species diagnostic traits, reflecting strong divergent selection （discordant）, especially on the Z （= X） chromosome. Reproductive isolation of these novel recombinant hybrid genotypes appears to be accomplished via a delayed post-diapause emergence or temporal isolation, and is perhaps aided by the thermal landscape. Changing thermal landscapes appear to have created （and may destroy） novel recombinant hybrid genotypes and hybrid species.展开更多
基金Acknowledgments This review reflects decades of personally treasured friendships and assistance from colleagues in the insectplant interactions field of study. The Papilionidae research from our lab was supported in part by the National Science Foundation （DEB-9201122 DEB-9510044+1 种基金 DEB 0716683 DEB 0918879） and the Michigan Agricultural Experiment Station （Project # 01644）. I wish to thank all lab members and friends for their enthusiasm and assistance over the years, and most recently, Matthew Aardema, Rodrigo Mereader and Gabe Ording.
文摘The search for pattern in the ecology and evolutionary biology of insect-plant associations has fascinated biologists for centuries. High levels of tropical （tow-latitude） plant and insect diversity relative to poleward latitudes and the disproportionate abundance of host-specialized insect herbivores have been noted. This review addresses several aspects of local insect specialization, host use abilities （and loss of these abilities with specialization）, host-associated evolutionary divergence, and ecological （including ＂hybrid＂） speciation, with special reference to the generation of biodiversity and the geographic and taxonomic identification of ＂species borders＂ for swallowtail butterflies （Papilionidae）. From ancient phytochemically defined angiosperm affiliations that trace back millions of years to recent and very local specialized populations, the Papilionidae （swallowtail butterflies） have provided a model for enhanced tmderstanding of localized ecological patterns and genetically based evolutionary processes. They have served as a useful group for evaluating the feeding specialization/physiological efficiency hypothesis. They have shown how the abiotic （thermal） environment interacts with host nutritional suitability to generate ＂voltinisrn/suitability＂ gradients in specialization or preference latitudinally, and geographical mosaics locally. Several studies reviewed here suggest strongly that the oscillation hypothesis for speciation does have considerable merit, but at the same time, some species-level host specializations may lead to evolutionary dead-ends, especially with rapid environmental/habitat changes involving their host plants. Latitudinal gradients in species richness and degree of herbivore feeding specialization have been impacted by recent developments in ecological genetics and evolutionary ecology. Localized insect- plant associations that span the biospectrum from polyphenisms, polymorphisms, biotypes, demes, host races, to cryptic species, remain academically contentious, with simple definitions still debated. However, molecular analyses combined with ecological, ethological and physiological studies, have already begun to unveil some answers for many important ecological/evolutionary questions.
基金This review reflects decades of personally treasured friendships and assistance from colleagues in the insectplant interactions field of study. The Papilionidae research from our laboratory was supported by such academic synergism and also in part by the National Science Foundation （DEB-9201122 DEB-9510044+1 种基金 DEB 0716683 DEB 0918879） and the Michigan Agricultural Experiment Station （Project # 01644）. I wish to thank all laboratory members and friends for their enthusiasm and assistance over the years, and most recently, Matthew Aardema, Rodrigo Mercader and Gabe Ording. Special thanks are extended to one particularly helpful reviewer.
文摘The ecology and evolutionary biology of insect-plant associations has real- ized extensive attention, especially during the past 60 years. The classifications （categorical designations） of continuous variation in biodiversity, ranging from global patterns （e.g., lat- itudinal gradients in species richness/diversity and degree of herbivore feeding specialization） to localized insect-plant associations that span the biospectrum from polyphenisms, polymorphisms, biotypes, demes, host races, to cryptic species, remain academically contentious. Semantic and biosystematic （taxonomical） disagreements sometimes detract from more important ecological and evolutionary processes that drive diversification, the dynamics of gene flow and local extinctions. This review addresses several aspects of insect specialization, host-associated divergence and ecological （including ＂hybrid＂） speciation, with special reference to the climate warming impacts on species borders of hybridiz- ing swallowtail butterflies （Papilionidae）. Interspecific hybrid introgression may result in collapse of multi-species communities or increase species numbers via homoploid hybrid speciation. We may see diverging, merging, or emerging genotypes across hybrid zones, all part of the ongoing processes of evolution. Molecular analyses of genetic mosaics and genomic dynamics with ＂divergence hitchhiking＂, combined with ecological, ethological and physiological studies of＂species porosity＂, have already begun to unveil some answers for some important ecological/evolutionary questions. （i） How rapidly can host-associated divergence lead to new species （and why doesn＇t it always do so, e.g., resulting in ＂incom- plete＂ speciation）？ （ii） How might ＂speciation genes＂ function, and how/where would we find them？ （iii） Can oscillations from specialists to generalists and back to specialists help explain global diversity in herbivorous insects？ （iv） How could recombinant interspecific hybridization lead to divergence and speciation？ From ancient phytochemically defined angiosperm affiliations to recent and very local geographical mosaics, the Papilionidae （swallowtail butterflies） have provided a model for enhanced understanding of ecological patterns and evolutionary processes, including host-associated genetic divergence, ge- nomic mosaics, genetic hitchhiking and sex-linked speciation genes. Apparent homoploid hybrid speciation in Papilio appears to have been catalyzed by climate warming-induced interspecific introgression of some, but not all, species diagnostic traits, reflecting strong divergent selection （discordant）, especially on the Z （= X） chromosome. Reproductive isolation of these novel recombinant hybrid genotypes appears to be accomplished via a delayed post-diapause emergence or temporal isolation, and is perhaps aided by the thermal landscape. Changing thermal landscapes appear to have created （and may destroy） novel recombinant hybrid genotypes and hybrid species.
