Allopatric origin of cryptic butterfly species that were discovered feeding on distinct host plants in sympatry

Surveys of tropical insects are increasingly uncovering cryptic species – morphologically similar yet reproductively isolated taxa once thought to comprise a single interbreeding entity. The vast majority of such species are described from a single location. This leaves us with little information on geographic range and intraspecific variation and limits our ability to infer the forces responsible for generating such diversity. For example, in herbivorous and parasitic insects, multiple specialists are often discovered within what were thought to be single more generalized species. Host shifts are likely to have contributed to speciation in these cases. But when and where did those shifts occur, and were they facilitated by geographic isolation? We attempted to answer these questions for two cryptic species within the butterfly Cymothoe egesta that were recently discovered on different host plants in central Cameroon. We first used mtDNA markers to separate individuals collected on the two hosts within Cameroon and then extended our analysis to incorporate individuals collected across the entire pan‐Afrotropical range of the original taxon. To our surprise, we found that the species are almost entirely allopatric, dividing the original range and overlapping only in the narrow zone of West‐Central Africa where they were first discovered in sympatry. This finding, combined with analyses of genetic variation within each butterfly species, strongly suggests that speciation occurred in allopatry, probably during the Pleistocene. We discuss the implications of our results for understanding speciation among other cryptic species recently discovered in the tropics and argue that more work is needed on geographic patterns and host usage in such taxa.     

Sympatric speciation in a genus of marine reef fishes

Sympatric speciation has been contentious since its inception, yet is increasingly recognized as important based on accumulating theoretical and empirical support. Here, we present a compelling case of sympatric speciation in a taxon of marine reef fishes using a comparative and mechanistic approach. Hexagrammos otakii and H. agrammus occur in sympatry throughout their ranges. Molecular sequence data from six loci, with complete sampling of the genus, support monophyly of these sister species. Although hybridization occurrs frequently with an allopatric congener in an area of slight distributional overlap, we found no F₁ hybrids between the focal sympatric taxa throughout their coextensive ranges. We present genetic evidence for complete reproductive isolation based on SNP analysis of 382 individuals indicating fixed polymorphisms, with no shared haplotypes or genotypes, between sympatric species. To address questions of speciation, we take a mechanistic approach and directly compare aspects of reproductive isolation between allopatric and sympatric taxa both in nature and in the laboratory. We conclude that the buildup of reproductive isolation is strikingly different in sympatric vs. allopatric taxa, consistent with theoretical predictions. Lab reared hybrids from allopatric species crosses exhibit severe fitness effects in the F₁ or backcross generation. No intrinsic fitness effects are observed in F₁ hybrids from sympatric species pairs, however these treatments exhibited reduced fertilization success and complete pre‐mating isolation is implied in nature because F₁ hybrid adults do not occur. Our study addresses limitations of previous studies and supports new criteria for inferring sympatric speciation.     

Ribosomal RNA Sequencing of Members of the Crypthecodinium cohnii (Dinophyceae) Species Complex; Comparison with Soluble Enzyme Studies

ABSTRACT. Sixty-five members of the Ctypthecodinium cohnii species complex were analyzed for sequence differences within the D2 region of the 23S ribosomal RNA molecule. On the basis of 46 sequence differences the strains fell into 19 distinct ribosets (strains of identical sequence), some with many members. Members of four of the seven major sibling species (widespread breeding groups) were each found within single ribosets. Members of three other major sibling species were each, however, divided into two ribosets by a single sequence difference correlated with geographic separation and with previously reported electrophoretic polymorphisms of soluble enzymes within the sibling species. In addition to members of major sibling species, some ribosets include many minor sibling species (each represented by only one strain). Of 38 minor sibling species, 22 shared sequence with a major sibling species. Of these 22, 14 were identical in soluble enzymes to their related major sibling species or differed by only one of three enzymes. Other minor sibling species appear to have diverged extensively from any others in both rRNA sequence and electrophoretic profile. As a group, major sibling species differ markedly in the number of minor sibling species associated with them, suggesting differences in frequency of sexually isolating events in their past histories. These findings are discussed in the context of the previously proposed model of sympatric speciation.     

The chromosomes of the hystricomorphous family Ctenodactylidae (Rodentia: ?Sciuromorpha) and their bearing on the relationships of the four living genera

Karyotype studies support the view that modern genera of the family Ctenodactylidae originated in Africa. Karyotype differences between the genera are less obvious than morphological differences but coincide in relating Massoutiera to Felovia and deriving this line from the Pectinator -like ancestor which, in turn, was closely related to a Ctenodactylus ancestor. 43% of the chromosomes are standard throughout the family; 25% seem to be very susceptible to fragmentation, translocation and inversion. These changeable chromosomes are the only ones that show differences in their G-band patterns. The ctenodactylid karyotype resembles caviomorph karyotypes in its NF, predominantly metacentric chromosomes and in its nucleolar organiser, or marker, chromosomes.     

Description of a new species from Laguna Chichancanab,Yucatan, Mexico: <Emphasis Type="Italic">Cyprinodon suavium</Emphasis> (Pisces: Cyprinodontidae)

Cyprinodon suavium is a new species that belongs to the endemic species flock from Laguna Chichancanab, Yucatan, Mexico, which is proposed to have evolved by sympatric speciation in the lake during the last 8000years. C. suaviumis distinguished from all other known Cyprinodon species by a flattened and concave inter- and postorbital skull roof and a terminal mouth with distinctive thickened lips. The short gut length and dietary items found in the gut indicate that C. suavium is one of the carnivorous members of the flock.     

Lake Tana large barbs: phenetics, growth and diversification

Among the Lake Tana large barbs of about 10 cm SL only representatives of the' acute'morphotype can be distinguished, in the size range 10–20 cm SL' bigmouth big eye' can be identified also. As for the rest, very few individuals can be confidently affiliated with a particular morphotype most of them looking like' intermedius'. Even within the range of 20–30 cm SL some individuals are still difficult to identify. Principal component analysis of cranial characters revealed discrete groups of morphotypes. Differences in both external and cranial characters of the morphotypes result from divergence which is most pronounced when fish are 4–5 years old and 20–25 cm SL. This divergence cannot be related exclusively with differences in the growth rates of individuals representing different morphotypes. Differences in food composition between the morphotypes probably increase in parallel with their morphological divergence. Differences between the morphotypes in the lateral line (11) and the gill rakers (Sb) counts were revealed using ANOVA. Comparison of the Lake Tana Barbus complex of forms with that previously known from Lake Lanao (Philippines) suggests that in both lakes the different forms arose sympatrically but that sympatric speciation in Lake Lanao has advanced further than in Lake Tana.     

A two-pollinator model for the evolution of floral complexity

For a new, more complex floral form to become established in a population it must overcome the problem of frequency-dependent constancy to successfully attract pollinators. This may be achieved by complex floral forms offering absolute greater rewards than the simpler forms, or by complex flowers offering a higher probability of being rewarding because fewer pollinators are able to visit them. In this paper we examine the effect of three pollinator foraging strategies on the ratio of flights within and between floral morphs and hence on the probability of a new morph establishing in a population without offering a greater reward. We incorporate pollinator behaviour based around observations of two pollinator species systems into three models of competition for pollinators. In the first model the constancy of the pollinator of the new floral morph is a function only of the foraging strategy of the existing pollinator of the original floral morph. In the next model the constancy of the second pollinator is determined by the number of rewarding flowers of each floral morph left by the original pollinator and in the third model it is determined by the ratio of rewarding flowers of each morph left by the original pollinator. The results demonstrate that under conditions of intense competition for pollinators, new, more complex floral forms are indeed able to attract high levels of constant pollinators without offering intrinsically higher rewards. However, for this to occur constancy in one of the pollinators must be a function of the ratio of rewarding to non-rewarding flowers of both floral forms. One prediction from our results is that sympatric speciation of floral complexity based on a higher probability of reward is more likely to occur in flowers offering rewards of pollen rather than nectar. This is because the cost of visiting non-rewarding flowers is usually higher where the reward is pollen rather than nectar. We also predict that complex flowers occurring at low frequency, which offer rewards of nectar, may need intrinsically greater rewards if they are to successfully attract pollinators.