Discordance Between Spatial Distributions of Y-Chromosomal and Mitochondrial Haplotypes in African Green Monkeys (Chlorocebus spp.): A Result of Introgressive Hybridization or Cryptic Diversity?

Introgressive hybridization may cause substantial discordances among phylogenies based on different genetic markers. Such discordances have been found in diverse mammal species including primates. A recent study of mitochondrial DNA (mtDNA) revealed several poly- and paraphyletic relationships in African green monkeys (Chlorocebus), suggesting contemporary and/or ancient introgressive hybridization among almost all parapatric species of the genus. However, mtDNA analyses alone do not allow us to draw conclusions concerning introgression events. In this study we analyzed two Y chromosomal (Y-chr) markers for 30 African green monkey samples and compared the resulting genetic relationships to those based on published mtDNA data. In line with the results for mtDNA, we found no Y-chr evidence of hypothesized hybridization among Chlorocebus sabaeus and C. tantalus in the northern part of the contact zone in West Africa, and we found two distinct and distantly related Y-chr haplotypes within the range of C. tantalus, suggesting possible cryptic genetic diversity rather than ancient introgressive hybridization in this species. In contrast, Y-chr data revealed monophyletic relationships within Chlorocebus pygerythrus from East Africa, suggesting that mtDNA paraphylies found in this species are most likely to be the result of ancient introgressive hybridization and subsequent cytonuclear extinction of an earlier taxon. Our results accentuate the importance of analyzing sex chromosomal data in addition to mtDNA to obtain more information on the potential outcomes of hybridization with respect to genetic and species diversity. Analysis of more diverse nuclear marker sets is needed to obtain a more complete picture of the African green monkey evolution.     

Coalescent analyses support multiple mainland-to-island dispersals in the evolution of Malagasy Triaenops bats (Chiroptera: Hipposideridae)

Aim We investigate the directionality of mainland‐to‐island dispersals, focusing on a case study of an African‐Malagasy bat genus, Triaenops (Hipposideridae). Taxa include T. persicus from east Africa and three Triaenops species from Madagascar (T. auritus, T. furculus, and T. rufus). The evolution of this bat family considerably post‐dated the tectonic division of Madagascar from Africa, excluding vicariance as a viable hypothesis. Therefore, we consider three biogeographical scenarios to explain these species' current ranges: (A) a single dispersal from Africa to Madagascar with subsequent speciation of the Malagasy species; (B) multiple, unidirectional dispersals from Africa to Madagascar resulting in multiple, independent Malagasy lineages; or (C) early dispersal of a proto‐species from Africa to Madagascar, with later back‐dispersal of a descendant Malagasy taxon to Africa. Location East Africa, Madagascar, and the Mozambique Channel. Methods We compare the utility of phylogenetic and coalescent methodologies to address the question of directionality in a mainland‐to‐island dispersal event for recently diverged taxa. We also emphasize the application of biologically explicit demographic systems, such as the non‐equilibrium isolation‐with‐migration model. Here, these methods are applied to a four‐species haploid genetic data set, with simulation analyses being applied to validate this approach. Results Coalescent simulations favour scenario B: multiple, unidirectional dispersals from Africa to Madagascar resulting in multiple, independent Malagasy bat lineages. From coalescent dating, we estimate that the genus Triaenops was still a single taxon approximately 2.25 Ma. The most recent Africa to Madagascar dispersal occurred much more recently (c. 660 ka), and led to the formation of the extant Malagasy species, T. rufus. Main conclusions Haploid genetic data from four species of Triaenops are statistically most consistent with multiple, unidirectional dispersals from mainland Africa to Madagascar during the late Pleistocene.     

Migration, isolation and hybridization in island crop populations: the case of Madagascar rice

Understanding how crop species spread and are introduced to new areas provides insights into the nature of species range expansions. The domesticated species Oryza sativa or Asian rice is one of the key domesticated crop species in the world. The island of Madagascar off the coast of East Africa was one of the last major Old World areas of introduction of rice after the domestication of this crop species and before extensive historical global trade in this crop. Asian rice was introduced in Madagascar from India, the Malay Peninsula and Indonesia approximately 800-1400 years ago. Studies of domestication traits characteristic of the two independently domesticated Asian rice subspecies, indica and tropical japonica, suggest two major waves of migrations into Madagascar. A population genetic analysis of rice in Madagascar using sequence data from 53 gene fragments provided insights into the dynamics of island founder events during the expansion of a crop species' geographic range and introduction to novel agro-ecological environments. We observed a significant decrease in genetic diversity in rice from Madagascar when compared to those in Asia, likely the result of a bottleneck on the island. We also found a high frequency of a unique indica type in Madagascar that shows clear population differentiation from most of the sampled Asian landraces, as well as differential exchange of alleles between Asia and Madagascar populations of the tropical japonica subspecies. Finally, despite partial reproductive isolation between japonica and indica, there was evidence of indica/japonica recombination resulting from their hybridization on the island.     

The antiquity of Madagascar’s grasslands and the rise of C4 grassy biomes

Aim Grasslands and savannas, which make up > 75% of Madagascar’s land area, have long been viewed as anthropogenically derived after people settled on the island c. 2 ka. We investigated this hypothesis and an alternative – that the grasslands are an insular example of the post‐Miocene spread of C₄ grassy biomes world‐wide. Location Madagascar, southern Africa, East Africa. Methods We compared the number of C₄ grass genera in Madagascar with that in southern and south‐central African floras. If the grasslands are recent we would expect to find fewer species and genera in Madagascar relative to Africa and for these species and genera to have very wide distribution ranges in Madagascar. Secondly, we searched Madagascan floras for the presence of endemic plant species or genera restricted to grasslands. We also searched for evidence of a grassland specialist fauna with species endemic to Madagascar. Plant and animal species endemic to C₄ grassy biomes would not be expected if these are of recent origin. Results Madagascar has c. 88 C₄ grass genera, including six endemic genera. Excluding African genera with only one or two species, Madagascar has 86.6% of southern Africa’s and 89.4% of south‐central Africa’s grass genera. C₄ grass species make up c. 4% of the flora of both Madagascar and southern Africa and species : genus ratios are similar (4.3 and 5.1, respectively). Turnover of grasses along geographical gradients follows similar patterns to those in South Africa, with Andropogoneae dominating in mesic biomes and Chlorideae in semi‐arid grassy biomes. At least 16 monocot genera have grassland members, many of which are endemic to Madagascar. Woody species in frequently burnt savannas include both Madagascan endemics and African species. A different woody flora, mostly endemic, occurs in less frequently burnt grasslands in the central highlands, filling a similar successional niche to montane C₄ grasslands in Africa. Diverse vertebrate and invertebrate lineages have grassland specialists, including many endemic to Madagascar (e.g. termites, ants, lizards, snakes, birds and mammals). Grassland use of the extinct fauna is poorly known but carbon isotope analysis indicates that a hippo, two giant tortoises and one extinct lemur ate C₄ or CAM (crassulacean acid metabolism) plants. Main conclusions The diversity of C₄ grass lineages in Madagascar relative to that in Africa, and the presence of plant and animal species endemic to Madagascan grassy biomes, does not fit the view that these grasslands are anthropogenically derived. We suggest that grasslands invaded Madagascar after the late Miocene, part of the world‐wide expansion of C₄ grassy biomes. Madagascar provides an interesting test case for biogeographical analysis of how these novel biomes assembled, and the sources of the flora and fauna that now occupy them. A necessary part of such an analysis would be to establish the pre‐settlement extent of the C₄ grassy biomes. Carbon isotope analysis of soil organic matter would be a feasible method for doing this.     

Deciphering patterns of transoceanic dispersal: the evolutionary origin and biogeography of coastal lizards (Cryptoblepharus) in the Western Indian Ocean region

Aim Cryptoblepharus is a genus of small arboreal or rock‐dwelling scincid lizards, widespread through the Indo‐Pacific and Australian regions, with a disjunct outlier in the Malagasy region. The taxonomy within this genus is controversial, with different authors ranking the different forms (now some 36) at various levels, from different species to subspecies of a single species, Cryptoblepharus boutonii. We investigated the biogeography and genetic differentiation of the Cryptoblepharus from the Western Indian Ocean region, in order to understand their origin and history. Location Western Indian Ocean region. Methods We analysed sequences of mitochondrial DNA (partial 12s and 16s rRNA genes, 766 bp) from 48 specimens collected in Madagascar, Mauritius, the four Comoros islands and East Africa, and also in New Caledonia, representing the Australo‐Pacific unit of the distribution. Results Pairwise sequence divergences of c. 3.1% were found between the New Caledonian forms and the ones from the Western Indian Ocean. Two clades were identified in Madagascar, probably corresponding to the recognized forms cognatus and voeltzkowi, and two clades were identified in the Comoro islands, where each island population formed a distinct haplotype clade. The East African samples form a monophyletic unit, with some variation existing between Pemba, Zanzibar and continental Tanzania populations. Individuals from Mauritius form a divergent group, more related to populations from Moheli and Grand Comore (Comoros islands) than to the others. Main conclusions The level of divergence between the populations from the Western Indian Ocean and Australian regions and the geographic coherence of the variation within the Western Indian Ocean group are concordant with the hypothesis of a colonization of this region by a natural transoceanic dispersal (from Australia or Indonesia). The group then may have diversified in Madagascar, from where it separately colonized the East African coast, the Comoros islands (twice), and Mauritius. The genetic divergence found is congruent with the known morphological variation, but its degree is much lower than typically seen between distinct species of reptiles.     

The taxonomy, reproductive biology and economic potential of Parkia (Leguminosae: Mimosoideae) in Africa and Madagascar

The taxonomy, reproductive biology and economic potential of Parkia (Leguminosae: Mimosoideae) in Africa and Madagascar. Three species of the pantropical woody genus Parkia are recognized from continental Africa (including São Tomé), P. biglobosa, P. bicolor and P.filicoidea. A fourth species, P. madagascariensis , is confined to Madagascar. All have capitula of similar structure although details of shape vary. Relatives sharing this capitulum structure occur in the Far East and the Neotropics. Patterns of infraspecific variation are described for the mainland species. Parkia biglobosa shows clinal variation in several leaf characters and P. clappertoniana must be regarded as synonymous. Parkia bicolor shows geographical variation especially in capitulum shape and pod width, and P. filicoidea varies geographically in its pod morphology. The red, balllike capitula are pollinated by bats; other visitors include birds, insects and non-flying mammals. In P. biglobosa a proportion of the capitula is functionally male. The pods of the African species contain mealy pulp. Dispersal involves mammals, especially primates, and birds. Information on uses, chemical composition of the seeds and insect pests is summarized.     

A new citrus-feeding psyllid from the Comoro Islands, with a review of the Diaphorina amoena species group (Homoptera)

ABSTRACT. Analysis of host plant data for the seventy-one species of Diaphorina shows the genus to be unusually polyphagous, the species developing on eighteen families of dicotyledonous angiosperms. Apart from the common Asian citrus pest, Diaphorina citri , a second citrus-feeding species is described and discussed. This species belongs to the amoena species-group which is defined and a key to nine of the ten included species is given. Seven new species are described: flavipennis sp.n. and nigripennis sp.n. from East Africa on Strychnos usambarensis; huila sp.n. from Angola on Ptaeroxylon obliquum; turned sp.n., from South Africa (host unknown); orantimina sp.n. from Madagascar (host unknown); auberti sp.n. from Comoro Islands on citrus trees; and bispinulata sp.n. from Madagascar (host unknown).     

Deep phylogenetic divergence between Scolytoplatypus and Remansus,a new genus of Scolytoplatypodini from Madagascar (Coleoptera,Curculionidae, Scolytinae)

Scolytoplatypodini Blandford is a monotypic tribe of ambrosia beetles found in Asia, Madagascar and Africa. Only three species are currently known from Madagascar and four additional species are here described as new to science. Phylogenetic analyses of morphological and molecular data revealed that four of the seven endemic species are deeply separated from all other species by genetic and distinct morphological characters and therefore placed in a new genus Remansus Jordal. The split between this ancient lineage and Scolytoplatypus Schaufuss was estimated to approximate Palaeocene age (63 Ma), extending the minimum age of ambrosia feeding for this tribe to the beginning of the Palaeocene‒Eocene thermal maximum (PETM). In addition to the ancient origin of Remansus in Madagascar during the Palaeocene, a second origin occurred in Scolytoplatypus no more than 13 Ma. A geographical origin of the latter in South-Eastern Africa was unequivocally inferred from the phylogenies.     

A RAPD study of the Sarcostemma group of Cynanchum (Apocynaceae-Asclepiadoideae-Asclepiadeae)

One hundred and thirty-seven accessions of Cynanchum viminale and its relatives, formerly known as Sarcostemma, were studied using randomly amplified polymorphic DNA (RAPD). A fingerprinting technique was used because sequencing had failed to differentiate between morphologically separable groups. Chromosome counts were conducted to establish the ploidy level of the accessions. The banding patterns resulting from RAPD analysis were evaluated with Canonical Analysis of Principal Coordinates, Permanova and neighbour-joining. A strong geographic component was found in the structure of the group. Taxa considered species or subspecies based on morphology often formed coherent groups. The data are interpreted to reflect at least two cycles of diversification: the first one from Madagascar and the second one most likely from the East African–Arabian region, reaching Madagascar again. In mainland Africa, polyploidisation has occurred several times.     

Two new species of Acutogordius (Nematomorpha), with a brief review of literature data of this genus

We describe two new species of the nematomorph genus Acutogordius, A. sulawensis from Sulawesi and A. protectus from Madagascar. For comparative reasons, the other known species are briefly described and discussed. The genus Acutogordius now comprises nine species regarded as valid. They are distributed on the entire southern hemisphere, from South and Central America over Africa and South‐east Asia to Australia. The three male specimens of Acutogordius protectus differ in the shape of a cuticular structure, the postcloacal crescent, which has implications for the potential of this character as a taxonomic tool.     

Systematic revision of the genus Aglyptodactylus Boulenger, 1919 (Amphibia: Ranidae), and analysis of its phylogenetic relationships to other Madagascan ranid genera (Tomopterna, Boophis, Mantidactylus, and Mantella)

Recent field studies revealed two new species of the genus Aglyptodactylus (Amphibia: Anura: Ranidae), which was hitherto considered as monotypic and confined to humid eastern Madagascar. Both new species, Aglyptodactylus laticeps sp. n. and Aglyptodactylus securifer sp. n. , occur syntopically in the deciduous dry forest of Kirindy in western Madagascar. In comparison to Aglyptodactylus madagascariensis from eastern rainforests, the new species A. laticeps shows a remarkable morphological divergence, which may be partly due to adaptations to burrowing habits in dry environments. Despite of the morphological differentiation, advertisement calls and osteology indicate that all three species of Aglyptodactylus are closely related. A phylogenetic analysis of the Madagascan ranid genera Aglyptodactylus, Mantella, Mantidactylus, Boophis , and Tomopterna (the latter including species from Madagascar, Africa, and Asia) strongly supports a sister group relationship of Aglyptodactylus with the ranine genus Tomopterna . We therefore transfer Aglyptodactylus from the Rhacophorinae to the Raninae and discuss implications of this rearrangement for ranoid systematics. The existence of the endemic genus Aglyptodactylus in Madagascar as well as its close phylogenetic relationships to Tomopterna confirm that the Raninae were already present on the Madagascan plate before its separation from Africa. The Madagascan Tomopterna labrosa shows several important differences both to Asian and to African species of the genus, and is therefore transferred from the subgenus Sphaerotheca (now restricted to Asia) to a new subgenus Laliostoma subgen. n .     

Why Africa matters: evolution of Old World Salvia (Lamiaceae) in Africa

Background and Aims

Salvia is the largest genus in Lamiaceae and it has recently been found to be non-monophyletic. Molecular data on Old World Salvia are largely lacking. In this study, we present data concerning Salvia in Africa. The focus is on the colonization of the continent, character evolution and the switch of pollination systems in the genus.

Methods

Maximum likelihood and Bayesian inference were used for phylogenetic reconstruction. Analyses were based on two nuclear markers [internal transcribed spacer (ITS) and external transcribed spacer (ETS)] and one plastid marker (rpl32-trnL). Sequence data were generated for 41 of the 62 African taxa (66 %). Mesquite was used to reconstruct ancestral character states for distribution, life form, calyx shape, stamen type and pollination syndrome.

Key Results

Salvia in Africa is non-monophyletic. Each of the five major regions in Africa, except Madagascar, was colonized at least twice, and floristic links between North African, south-west Asian and European species are strongly supported. The large radiation in Sub-Saharan Africa (23 species) can be traced back to dispersal from North Africa via East Africa to the Cape Region. Adaptation to bird pollination in southern Africa and Madagascar reflects parallel evolution.

Conclusions

The phenotypic diversity in African Salvia is associated with repeated introductions to the continent. Many important evolutionary processes, such as colonization, adaptation, parallelism and character transformation, are reflected in this comparatively small group. The data presented in this study can help to understand the evolution of Salvia sensu lato and other large genera.     

A revision of the brassoline genus Selenophanes (Lepidoptera: Rhopalocera)

The Brassoline genus Selenophanes is revised: three new subspecies are described and three lectotypes designated. The nine known holotypes and two of the three lectotypes are figured. Along the Andes there is a fairly close agreement in the distribution of some of the species and subspecies of Selenophanes with the recently recognized endemic centres. East of the Andes there is no close correspondence between subspecies and endemic centres; each subspecies has, in general, a distribution across several centres. A similar distribution has been noted previously in the Brassoline genus Catoblepia.     

Biogeography and diversification of hermit spiders on Indian Ocean islands (Nephilidae: Nephilengys)

The origin of the terrestrial biota of Madagascar and, especially, the smaller island chains of the western Indian Ocean is relatively poorly understood. Madagascar represents a mixture of Gondwanan vicariant lineages and more recent colonizers arriving via Cenozoic dispersal, mostly from Africa. Dispersal must explain the biota of the smaller islands such as the Comoros and the chain of Mascarene islands, but relatively few studies have pinpointed the source of colonizers, which may include mainland Africa, Asia, Australasia, and Madagascar. The pantropical hermit spiders (genus Nephilengys) seem to have colonized the Indian Ocean island arc stretching from Comoros through Madagascar and onto Mascarenes, and thus offer one opportunity to reveal biogeographical patterns in the Indian Ocean. We test alternative hypotheses on the colonization route of Nephilengys spiders in the Indian Ocean and simultaneously test the current taxonomical hypothesis using genetic and morphological data. We used mitochondrial (COI) and nuclear (ITS2) markers to examine Nephilengys phylogenetic structure with samples from Africa, southeast Asia, and the Indian Ocean islands of Madagascar, Mayotte, Réunion and Mauritius. We used Bayesian and parsimony methods to reconstruct phylogenies and haplotype networks, and calculated genetic distances and fixation indices. Our results suggest an African origin of Madagascar Nephilengys via Cenozoic dispersal, and subsequent colonization of the Mascarene islands from Madagascar. We find strong evidence of gene flow across Madagascar and through the neighboring islands north of it, while phylogenetic trees, haplotype networks, and fixation indices all reveal genetically isolated and divergent lineages on Mauritius and Réunion, consistent with female color morphs. These results, and the discovery of the first males from Réunion and Mauritius, in turn falsify the existing taxonomic hypothesis of a single widespread species, Nephilengys borbonica, throughout the archipelago. Instead, we diagnose three Nephilengys species: Nephilengys livida (Vinson, 1863) from Madagascar and Comoros, N. borbonica (Vinson, 1863) from Réunion, and Nephilengys dodo new species from Mauritius. Nephilengys followed a colonization route to Madagascar from Africa, and on through to the Mascarenes, where it speciated on isolated islands. The related golden orb-weaving spiders, genus Nephila, have followed the same colonization route, but Nephila shows shallower divergencies, implying recent colonization, or a moderate level of gene flow across the archipelago preventing speciation. Unlike their synanthropic congeners, N. borbonica and N. dodo are confined to pristine island forests and their discovery calls for evaluation of their conservation status.     

A craniometric study of the black and white Colobus Illiger 1811 (Primates: Ceropithecoidea).

This study examines the craniometry of Black and White Colobus monkeys using 1072 specimens representing all the recognized subspecies (after Rahm, '70) of the genus. Seventy-six measurements were taken on each individual, and examined using canonical variates analysis and clustering by Ward's Error Sum method. The assumptions of the analytical techniques are shown to be met, and the results demonstrated to be stable. Examination of the populations for statistical difference and taxonomic distinctiveness using a multivariate extension of Mayr's Rule indicates that the taxonomy presented by Rahm ('70) is essentially correct, except that the subspecies of guereza across the northern part of Central Africa should be lumped into a single group--C. g. occidentalis--and the subspecies of montane angolan colobus in Eastern Zaire should all be lumped into C. a. ruwenzorii. The systematic patterns of the genus illustrate the whole range of the process of speciation, from barely distinct subspecies, to subspecies almost as distinct as allopatric species of the genus, and on the fully sympatric species Three major zoogeographic areas may be delineated: an East African area dominated by the effects of the Rift Valley, with a large number of subspecies isolated in forest islands; a Central African area with little subspeciation and sympatric overlap of the major species of Colobus; and a West African area with a clinal pattern of distribution of subspecies, and secondary intergradation. The arbitrary nature of Mary's Rule is also apparent. Lastly, the CVA indicated major differences across the genus to be located in the teeth and jaws, suggesting diet might be an important distinguishing feature in Colobus.     

A chloroplast DNA phylogeny of eastern Phlox (Polemoniaceae): implications of congruence and incongruence with the ITS phylogeny

The comparison of independent phylogenies is a valuable approach to the study of evolutionary pattern and process. Available data on eastern North American Phlox, including our recent ITS phylogeny, suggest that relationships are complicated in the group and that hybridization may have been a contributing factor. We used restriction site data from the chloroplast genome to develop a second phylogeny for eastern Phlox. Sampling was the same as that for the ITS study and consisted of 79 samples (including all 22 eastern Phlox species and most eastern subspecies, as well as multiple populations of many taxa). The resulting cpDNA phylogeny agrees with the ITS phylogeny in many respects, strengthening earlier conclusions. Nevertheless, incongruence between the trees is noteworthy: many samples, particularly of members of the P. pilosa and P. glaberrima complexes, are placed in different clades. A variety of tests were carried out to assess congruence in terms of topological patterns, character congruence, and homogeneity of data sets. Significant conflict between the phylogenies is discussed in light of the hypothesis that hybridization has affected relationships in this genus.     

High species diversity in one of the dominant groups of spiders in East African montane forests (Araneae: Pholcidae: Buitinga n. gen., Spermophora Hentz)

The six-eyed pholcid spiders of East Africa are revised. A new genus is recognized, Buitinga n. gen ., with 17 new species and three species transferred from Spermophora Hentz. The new genus is characterized by the presence of a scape on the epigynum. This scape may be straight or tightly curled up at rest, and is usually highly expandable. Seven additional African and Comoran species are newly described and tentatively assigned to Spermophora . A data matrix with 60 characters and 77 taxa (including 20 East African species and 25 additional Spermophora and ' Spermophora -like' species) is analysed cladistically. Buitinga is closer to the genera Paramicromerys Millot (endemic in Madagascar) and Spermophorides Wunderlich (Mediterranean and Canary Islands) as well as to several African and Comoran species tentatively assigned to Spermophora than to the type species of Spermophora . It is argued that current estimates of species numbers may be inaccurate and that pholcids may turn out to be one of the most diverse spider families.  © 2003 The Linnean Society of London. Zoological Journal of the Linnean Society, 2003, 137, 555−619