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.     

Notes on African species of the genus Anisopappus Hook. & Arn. (Asteraceae)

Two new species of the genus Anisopappus Hook. & Arn. (Asteraceae) are described A. athanasioides Paiva & Ortiz, from Angola, and A. pseudopinnatifidus Ortiz & Paiva, from Namibia. The morphological variability of the A. chinensia-A. africanus complex is studied and two new combinations are proposed: A. chinensis subsp. africanus (Hook. fil.) Ortiz & Paiva and A. chinensis subsp. lobatus (Wild) Ortiz & Paiva.     

Notes on African species of the genus Anisopappus Hook. & Arn. (Asteraceae)

Two new species of the genus Anisopappus Hook. & Arn. (Asteraceae) are described: A. athanasioides Paiva & Ortiz, from Angola, and A. pseudopinnaididus Ortiz & Paiva, from Namibia. The morphological variability of the A. chinensis-A. africanus complex is studied and two new combinations are proposed: A. chinensis subsp. africanus (Hook, fil.) Ortiz & Paiva and A. chinensis subsp. lobatus (Wild) Ortiz & Paiva.     

An Inordinate Fondness? The Number,Distributions, and Origins of Diatom Species

The number of extant species of diatoms is estimated here to be at least 30,000 and probably ca. 100,000, by extrapolation from an eclectic sample of genera and species complexes. Available data, although few, indicate that the pseudocryptic species being discovered in many genera are not functionally equivalent. Molecular sequence data show that some diatom species are ubiquitously dispersed. A good case can be made that at least some diatom species and even a few genera are endemics, but many such claims are still weak. The combination of very large species numbers and relatively rapid dispersal in diatoms is inconsistent with some versions of the “ubiquity hypothesis” of protist biogeography, and appears paradoxical. However, population genetic data indicate geographical structure in all the (few) marine and freshwater species that have been examined in detail, sometimes over distances of a few tens of kilometres. The mode of speciation may often be parapatric, in the context of a constantly shifting mosaic of temporarily isolated (meta) populations, but if our “intermediate dispersal hypothesis” is true (that long‐distance dispersal is rare, but not extremely rare), allopatric speciation could also be maximized.     

Ecological and morphological differentiation among cryptic evolutionary lineages in freshwater limpets of the nominal form-group Ancylus fluviatilis (O.F. Müller, 1774)

The phylogeny and potential mode of speciation of the river limpet Ancylus fluviatilis (Basommatophora) was examined using mitochondrial DNA sequences from 16S ribosomal RNA, cytochrome c oxidase subunit I (COI) and nuclear DNA from internal transcribed spacer (ITS-1) regions from 103 populations across Europe. Four highly divergent lineages were observed within Ancylus. Clade 1, representing the nominal taxon Ancylus fluviatilis (O.F. Müller, 1774), is mainly found in central and northern Europe, Clade 2 is present in a single Portuguese population, Clade 3 is distributed on the Canary islands, North Africa and the eastern Mediterranean region, whereas Clade 4 inhabits the Northern Mediterranean coasts. Phylogenetic analyses revealed an overall consistent topology of nuclear and mitochondrial gene trees. Based on a molecular clock, we estimated that the basic radiation occurred in the late Pliocene. Although clades differ significantly in size independent shell shape, morphological differentiation of lineages is not feasible without genetic data. Environmental data related to climate (precipitation, temperature, etc.) showed a significant differentiation of clades. Clade 1 dwells in relatively colder and more stable habitats than Clades 3 and 4, whose habitats in turn differ in a low or high amount of precipitation during spring and autumn, respectively. Based on the combined data sets on mitochondrial DNA, nuclear DNA, morphological and ecological differentiation, we conclude that Ancylus represents a cryptic species complex of reproductively and genetically isolated lineages. In addition, the joint analysis suggests that ecological speciation is probable to explain current patterns.     

Morphological stasis in the Eurytemora affinis species complex (Copepoda: Temoridae)

Morphological stasis has long been regarded as one of the most challenging problems in evolutionary biology. This study focused on the copepod species complex, Eurytemora affinis, as a model system to determine pattern and degree of morphological stasis. This study revealed discordant rates of morphological differentiation, molecular evolution, and reproductive isolation, where speciation was accompanied by lack of morphological differentiation in secondary sex characters. Comparisons were made among phylogenies based on morphometrics, nuclear (allozyme) loci, and mitochondrial DNA (mtDNA) sequences from cytochrome oxidase I, for a total of 43 populations within the complex. These systematic relationships were also compared to patterns of reproductive isolation. In addition, genetic subdivision of nuclear molecular (allozyme) markers (G _ST) and quantitative (morphological) characters (Q _ST) were determined to infer evolutionary forces driving morphological differentiation. The morphometric phylogeny revealed that all clades, excluding the European clade, were morphologically undifferentiated and formed a polytomy (multifurcation). Morphometric distances were not correlated with mtDNA distances, or with patterns of reproductive isolation. In contrast, nuclear and mtDNA phylogenies were mostly congruent. Reproductive isolation proved to be the most sensitive indicator of speciation, given that two genetically and morphologically proximate populations showed evidence of hybrid breakdown. Quantitative genetic (morphological) subdivision (Q _ST = 0.162) was lower than nuclear genetic subdivision (G _ST = 0.617) for four laboratory-reared North American populations, indicating retarded evolution of morphological characters. This result contrasts with most other species, where Q _ST typically exceeds G _ST as a result of directional selection. Thus, in all but the European populations, evolution of the secondary sex characters was marked by morphological stasis, even between reproductively-isolated populations.     

Geographical patterns of old and young species in African forest biota: the significance of specific montane areas as evolutionary centres

A widely accepted paradigm for speciation in tropical forests, the refuge theory, requires periodic habitat fragmentation driven by global climatic fluctuations to provide conditions for allopatric speciation. This implies that comparative species richness in refugia is due to loss of diverse communities in areas affected by climatic cycles. In this study we compare distribution patterns of bird and plant taxa which we consider to be of either deep phylogenetic lineages or recent radiations. It is demonstrated that lowland areas which have been postulated as Pleistocene refugia are dominated by species which represent lineages of pre-Pleistocene age. Since variations in species richness within these forest tracts reflect currently apparent environmental variables which might be considered to determine carrying capacity, we do not need to postulate that richness is the result of changes in forest cover in the past. Recently diversified taxa of plants and birds are found mainly at the periphery of the main rain forest blocks and in habitat islands outside them. Here, peak concentrations of young restricted-range species are often congruent with clusters of old and biogeographically relictual species. It is suggested that this reflects special intrinsic environmental properties of these areas, in the form of long-term environmental stability caused mainly by persistent orographic rain or mist. In this case, richness is not necessarily due to extinction outside these areas. Stability not only enables survival of relictual taxa, but also promotes morphological differentiation of radiating taxa, leading to aggregates of taxa of restricted distribution.     

Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies

Paracoccidioides brasiliensis is the etiologic agent of paracoccidioidomycosis, a disease confined to Latin America and of marked importance in the endemic areas due to its frequency and severity. This species is considered to be clonal according to mycological criteria and has been shown to vary in virulence. To characterize natural genetic variation and reproductive mode in this fungus, we analyzed P. brasiliensis phylogenetically in search of cryptic species and possible recombination using concordance and nondiscordance of gene genealogies with respect to phylogenies of eight regions in five nuclear loci. Our data indicate that this fungus consists of at least three distinct, previously unrecognized species: S1 (species 1 with 38 isolates), PS2 (phylogenetic species 2 with six isolates), and PS3 (phylogenetic species 3 with 21 isolates). Genealogies of four of the regions studied strongly supported the PS2 clade, composed of five Brazilian and one Venezuelan isolate. The second clade, PS3, composed solely of 21 Colombian isolates, was strongly supported by the alpha-tubulin genealogy. The remaining 38 individuals formed S1. Two of the three lineages of P. brasiliensis, S1 and PS2, are sympatric across their range, suggesting barriers to gene flow other than geographic isolation. Our study provides the first evidence for possible sexual reproduction in P. brasiliensis S1, but does not rule it out in the other two species.     

Endemism, diversity, and the threat of tropical moist forest extinctions

Extinction rates have risen to perhaps 10⁴ the background rate. Much of this increase is due to projected influences of habitat loss on regions of the world with tropical moist forest. This ecosystem, home to a disproportionate amount of global biodiversity and a major regulator of regional and global climate, also faces disproportionately severe threats. In this study, we collect diversity and endemism data for tropical forested countries of the world, along with areal and socioeconomic data. While a correlation between overall numbers of species and endemic species per country is expected, we demonstrate that endemism patterns among birds and mammals remain very strongly convergent even after statistically rendering all countries equal in size and overall species richness and after adjusting for spatial autocorrelation. On a per country basis, mammals are generally more threatened than birds in these tropical moist forested countries. Human population growth rates and rising debt among these nations should be viewed as priorities for amelioration by the developed countries. Reserve network extent is not related to numbers of endemic mammals or birds at this large spatial scale.