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Preview this Book. Woods, Florence E. Add to Wish List. Close Preview. Toggle navigation Additional Book Information. Description Table of Contents Editor s Bio. Summary As a review of the status of biogeography in the West Indies in the s, the first edition of Biogeography of the West Indies: Past, Present, and Future provided a synthesis of our current knowledge of the systematics and distribution of major plant and animal groups in the Caribbean basin. The totally new and revised Second Edition, Biogeography of the West Indies: Patterns and Perspectives, emphasizes recent ideas and hypotheses in the field and includes many new chapters and contributions.

The authors use the broadest possible interpretations of the concepts of biogeography, consider anthropological and geological factors, and discuss the conservation of endemic species.


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Drawing together contributions from the leading experts in biogeography and biodiversity, this book introduces new patterns and developments that add to our understanding of how plants and animals are dispersed throughout the region. Many contributions use new techniques such as molecular systematics to test older studies based strictly on morphological data.

Unique in its inclusion of a wide variety of organisms and in its coordination of scientific data and conservation strategies, Biogeography of the West Indies: Patterns and Perspectives, Second Edition provides the only encyclopedic discussion available on the biogeography of the Antilles. Table of Contents Introduction, C. Curtis, M. Brenner, and D. McNab Phylogeny and Biogeorgraphy of Lyonia sect.

Lyonia Ericaceae , W. Genaro and A. Miller and L. Hass, L. Certain morphological similarities have even led some authors to propose African affinities with, for instance, the subtribe Ypthimina Satyrini [ 33 ] and the satyrine tribe Dirini [ 34 ],[ 35 ].

Regardless of the phylogenetic position of Calisto , a continental origin of the genus is the most plausible explanation, as no other extant satyrine butterflies with the potential of being a closely related group are found in the Greater Antilles; thus, its ancestors would have necessarily arrived to the Caribbean from the nearby American continent [ 31 ],[ 33 ],[ 36 ]. Once Calisto colonized the Greater Antilles, further differentiation by vicariance [ 31 ],[ 37 ], within-island diversification [ 28 ],[ 36 ] or adaptive radiation [ 27 ] might have shaped the evolution of these butterflies.

In this study, we aim to elucidate the phylogenetic affinities and to identify the main drivers of the diversification and distribution of Calisto by using a secondarily calibrated molecular phylogeny. We also aim to reconstruct the historical biogeography of Calisto and to evaluate possible changes in diversification rates throughout the evolution of the genus.

Intra-island differentiation appears to be an important factor for the radiation of these butterflies, a phenomenon observed in other Caribbean animal lineages [ 2 ],[ 4 ],[ 9 ],[ 38 ]-[ 40 ]. However, even though rapid diversification driven by ecological evolution is plausible explanation considering the diversity of Caribbean habitats, niche saturation and island size may have imposed diversification limits [ 38 ] which could have restricted the diversity and geographical distribution of Calisto.

Our phylogenetic inferences using single gene datasets are congruent with the combined analyses, recovering the main clades within Calisto Additional file 1. Moreover, the combined analyses were consistent regardless of the method used and the partitioning strategy Figure 1. A summary of the dataset properties is presented in Table 1.

BI consensus phylogeny using the combined dataset partitioned by gene. Support values are represented by symbols on the left of each node, where the upper symbol is the bootstrap BS support value from the ML analysis, and the left bottom symbol is the posterior probability PP of the Bayesian Inference BI from the gene partition analysis and from the partition-by-bins analysis on the right. Filled stars are strong support values of 0.

Calisto nubila split early in the evolution of the genus, becoming an old and separate entity. The lineage did not apparently diversify further within Puerto Rico, although C. Three main monophyletic groups from Hispaniola are identified: the lyceius -, the confusa - hysius and the chrysaoros clades.

The monophyletic group consisting of Cuban and Bahamian Calisto is closely related to Hispaniolan lineages such as C. A revised checklist of the genus Calisto is presented in Table 2. Dated phylogram and a consensus biogeographical history. The ultrametric tree is scaled in Ma.


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Colours on each symbol represent the level of support. The phylogeny in the bottom left is the Satyrini tree, with the Calisto clade showing in red. Extant distributions of Calisto , following the subdivision of the Greater Antilles, are represented by coloured squares. The main geological events through time are depicted on top of the figure following the time scale in Ma.

LTT plots follow the time scale of the phylogeny in Ma. Confidence intervals for LTT are displayed as coloured ranges. The genus Calisto was not recovered within any valid Satyrini subtribes. Instead, our BEAST reconstructions place it sister to all sampled subtribes except Euptychiina with low support values posterior probability around 0.

The exclusion of the genus Euptychia which apparently caused long branch attraction in a different dataset [ 32 ] only increases the support for such a placement to moderate values around 0. Height posterior distributions displayed normally whereas summarizing the trees as means or medians height showed no significant difference.

There was no statistical difference in the global likelihood between the non-time-stratified analyses NS0 and NS1 Table 3. The time-stratified analysis TS1 favoured vicariance over dispersal in all cases and TS2 inferred a PR-sH origin of Calisto and vicariance for the origin of Cuban diversity. However, TS2 analysis did not improve the global likelihood of the inference over TS1. Root optimizations significantly favoured a PR-sH distribution and vicariance as the cause of the Cuban clade split from its sister Hispaniolan lineages. Geological history of the Greater Antilles and Bahamas and the evolution of Calisto.

Maps were modified from [ 6 ]. Area connectivity and dispersal rates used in our biogeographical analyses are shown below each time period a: 31—20 Ma, b: 10—20 Ma, c: 5—10 Ma, and d: 5 Ma to present. Dispersal probability, as used in TS analyses, are displayed below on each table. LD is long-distance dispersal including one extra area.

Values of 0. The estimation of the parameter j founder-event speciation significantly improved the DEC models. NS1- j preferred dispersal in critical nodes, i. From all rate-variable models in DDD , only those with a whole Calisto shift under diversity-dependent process are preferred with Akaike weights higher than 0. The decoupling of parameters for the Cuban taxa alone from the main Calisto tree was not enough to explain the radiation of the genus. Cuban and Hispaniolan taxa analyzed separately did not have constant diversification rates; rates changed possibly due to increased speciation, diversity-dependence processes, or a combination of both Akaike Weights were unable to discern among models.

Including the number of missing taxa into the models when possible did not affect the recovered estimations Table 6. Previous attempts to date the diversification of Calisto were done based only on a pairwise substitution rate for mitochondrial evolution [ 27 ]. This latter study deduced younger ages 4—8 Ma but did not actually carry out a timing of the divergence analysis, rather they only calculated pairwise genetic distances with Kimura 2-parameter without an adequate model testing. It is not the first time that GAARlandia is invoked to explain butterfly geographic range expansion.

It is the case for the nymphaline subtribe Phyciodina [ 52 ], the satyrine subtribe Pronophilina [ 32 ], and certain lineages within the papilionid tribe Troidini [ 53 ]. Adult butterflies respond to incoming bad weather by taking refuge [ 36 ] whereas a high mortality of eggs, larvae and pupae is observed when they are exposed to marine water [ 54 ]. Calisto , when compared to most other butterflies, are rather sedentary, and hence the direct and indirect dispersal capabilities of Calisto make a dispersalist model less likely.

Therefore, the evolution of Calisto is better explained by the main predictions of the Caribbean paleogeographical model of colonization rather than the stochastic dispersalist scenario. The inclusion of Jamaica into the vicariance model is less supported by the paleogeographical reconstructions, although a remote connection between the Blue Mountains block with GAARlandia has not been discarded [ 4 ],[ 6 ]. At that time, large portions of Jamaica began to uplift and the entire island remained above water afterwards, and hence the colonization of Jamaica by rare long-distance dispersal events is the most likely explanation for the origin of the endemic sole species found there, Calisto zangis.

Biogeographical reconstructions were significantly improved when we constrained dispersal probability and area-connectivity following the paleogeographic history of the Caribbean. Moreover, we found for the first time, statistical support for long-distance dispersal in the colonization of the Bahamas and Jamaica by estimating a founder-event parameter using a more general DEC model.

This could be due primarily to, first, the assumptions made by the models and, second, the different approaches to node reconstruction. In the first case, vicariance is favoured when incorporating area connectivity through time TS but dispersal is recovered by adding the parameter j founder-event or long-distance dispersal speciation. This difference is evidenced in NS1 analysis replicated using both software programs where vicariance is only reconstructed under the global most-likely inference by BioGeoBEARS. We believe, given the paleogeographic scenario and our dating estimations which correlate with the former, that the most plausible explanation for the colonization of Cuba is vicariance.

Furthermore, the whole of extant Cuban diversity is monophyletic and sister to a Hispaniola lineage, as predicted by the vicariant model. Vicariance driving speciation within islands is significantly recovered for Hispaniolan fauna during two instances, at 10—13 Ma and 4—6 Ma. The first vicariant instance is independently evidenced in two lineages with simultaneous shifts in ancestral ranges, the lyceius and the confusa-hysius clades.

The dating estimates are congruent with the major uplift of the Cordillera Central which might have provided new ecological opportunities and created isolated populations [ 39 ],[ 40 ]. For instance, sister species-pairs within both major clades feed, as larvae, exclusively on distinct bunch grasses, and have morphologically adapted to specific altitudinal ranges. Species inhabiting lower altitude and warmer areas are smaller than their sister montane species [ 27 ],[ 46 ], suggesting an adaptation for thermoregulatory efficiency [ 57 ].

The second instance of vicariant process within Hispaniola occurred during the Pliocene as evidenced in the lyceius and chrysaoros clades. Ecological niche shifts might be another plausible explanation for the lyceus clade members having differentiated during the Pliocene. As larvae, they feed on the bunchgrass Uniola virgata , which provides a unique niche and would have required significant adaptations [ 56 ].

Acknowledgments

Its sister taxa are Hispaniolan lineages that occur in the northwestern Cordillera Central Massif du Nord in Haiti [ 22 ],[ 46 ], which is the closest region to eastern Cuba. The two Bahamian lineages have distinct ancestral areas, while C. The species richness of Calisto across islands is largely unequal. Such a pattern has been previously reported as the consequence of island size and age, ecological limits and habitat diversity [ 8 ],[ 38 ],[ 58 ]. Munroe [ 59 ],[ 60 ] pointed out that extant Calisto diversity is distributed unequally among islands more likely due to speciation rather than to differential immigration, and that extinction was extremely low, especially in Hispaniola.

The calculation of diversification rates and ancestral states in this study suggested that the extant geographical distribution of Calisto reflects the rapid diversification within Hispaniola and Cuba during two instances, at 25 and 14 Ma, while inter-island flow was negligible for the entire genus. Calisto is the most species-rich butterfly genus in the West Indies because it was able to expand its ecological niche e.

The recovered date of this shift at 14 Ma is congruent with an increase in ecological opportunity in Hispaniola and Cuba and a time at which new environments were being created as a result of geological processes e. Nonetheless, the arrival of Calisto to an unoccupied island of Cuba did certainly provide for new heretofore empty niches to be colonized. The most likely scenario for such a decoupling was at 14 Ma, as recovered in DR1 analysis.

However, because such a date is confounded with the availability of new niches in Hispaniola, a model including one single shift in K for the whole genus was preferred. Adaptive radiation and the origin of island endemism of West Indies insects remain statistically untested. Under a phylogenetic framework, indirect evidence of adaptive radiation could be inferred based on diversification rate shifts: i. It is unlikely that the extinction rate rose, as it was near zero in all of our estimations.

The second major radiation took place at 14 Ma discussed above , but it is more plausible that the diversification rate increased due to a shift in K rather than by a sole increase in speciation rate. The Cuban clade, when analyzed independently, better fits a 2-yule-rate, with 5 times larger speciation rate at 10 Ma than when the lineage branched off at 21 Ma.

An intriguing question is why the observed diversification dynamics of Calisto on Cuba and Hispaniola were not replicated on Jamaica and Puerto Rico, the third and fourth largest islands of the West Indies, respectively. Whereas Calisto are usually locally adapted to particular habitats within Cuba and Hispaniola, the single species on each of the other two islands are widespread.

While some diverse Hispaniolan lineages feed as larvae on bunch grasses, the Puerto Rican C. According to Turner, similar, relatively adaptable oviposition behaviour is exhibited by C. Perhaps in this indiscriminate behaviour lies the explanation for the fact that these two species were able to colonize their entire respective islands instead of forming separate disjunctive populations as did their Hispaniolan congeners. Such wide distribution and relatively good dispersal abilities of these relatively larger Calisto species Sourakov, pers.

Further research on the natural history, dietary preferences and behaviour of Calisto is necessary to corroborate our speculations. The phylogenetic and biogeographical evidence presented in this study agrees with the Caribbean paleogeographical model of colonization Figures 2 and 3. Vicariant models explaining the diversification of Calisto have already been proposed based on their extant geographical distribution [ 31 ],[ 33 ],[ 36 ],[ 60 ], although some authors had favoured the alternative dispersalist explanation [ 27 ],[ 63 ]. Kwet, A.

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