Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails

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.

The latitudinal diversity gradient in New World swallowtail butterflies is caused by contrasting patterns of out‐of‐ and into‐the‐tropics dispersal

“…The sole species of the temperate subgenus Alexanoria is sister to the clade composed of Agehana + Chilasa + Pterourus . Both subgenus and species group‐level relationships are consistent with results of previous studies (Condamine et al, ; Lewis et al, ; Shiraiwa et al, ; Zakharov et al, ). Within the New World Papilio , temperate and tropical taxa do not constitute monophyletic assemblages (Figure ; see also Supporting Information Appendix S2 Figure S2.2).…”

“…Our biogeographical reconstructions indicate that the most recent common ancestor of New World swallowtails is likely to have occupied both northern temperate and tropical latitudes and climatic tropics and temperate zones. Corroborating previous work (Lewis et al, ), we found that after a putative vicariance event c . 20 Ma, Heraclides diversified in the climatic and latitudinal tropics, with some lineages then dispersing to temperate latitudes and cooler climates in the late Miocene.…”

“…We sequenced two mitochondrial genes, cytochrome oxidase I ( COI , 1,495 nucleotides) and cytochrome oxidase II ( COII , 680 nucleotides), and two nuclear genes, elongation factor 1‐alpha ( EF‐1α , 1,240 nucleotides) and wingless ( Wg , 403 nucleotides). We also included 133 morphological characters for Heraclides (Lewis et al, ) to improve phylogenetic resolution among Papilio species. Further details on sequencing procedures and sequence cleaning are provided in Supporting Information Appendix S4.…”

“…The clade is distributed primarily in the New World, from northern Alaska and Canada south to Argentina, although subgenus Chilasa is an Old‐World Southeast Asian lineage, and Alexanoria is spread throughout Mediterranean Europe and the Middle East (Zakharov, Caterino, & Sperling, ). These species have an extensive history of research by amateur naturalists and career researchers, resulting in a wealth of available biological specimens, molecular sequences and occurrence data (e.g., Condamine et al, ; Lewis et al, ; Shiraiwa, Cong, & Grishin, ; Tyler, Brown, & Wilson, ; Wu et al, ). Still, no study has yet assessed whether geographical shifts in lineages of Papilio are correlated with shifts in diversification rates, or how abiotic niche evolution may have related to geographical shifts.…”

Species delimitation and evolutionary relationships among Phoebis New World sulphur butterflies (Lepidoptera, Pieridae, Coliadinae)

“…As suggested by present‐day behaviour of some species (Urquhart & Urquhart, ; Oliveira et al ., ; Srygley & de Oliveira, ), some ancestral populations were probably strongly vagile and colonized the continent across the open ocean. Reverse colonization from the Greater Antilles to the continent has been inferred for other groups, including Urocoptidae land snails (Uit de Weerd et al ., ), Heraclides swallowtail butterflies (Lewis et al ., ), Exophthalmus weevils (Zhang et al ., ), Anolis lizards (Nicholson et al ., ), Eleutherodactylus frogs (Heinicke et al ., ) and Amazona parrots (Russello & Amato, ).…”