Current geographical ranges of Malagasy dung beetles are not delimited by large rivers

Aim We investigated whether the largest river (Mangoro) on the east coast of Madagascar acts as a barrier to dispersal in dung beetles by comparing species composition and genetic differentiation of the most common species on the two banks of the river. Moreover, by analysing the current geographical ranges of all wet forest dung beetle species, possible long‐term effects of the largest rivers on the distribution of species were assessed. Location Madagascar. Methods Dung beetles were sampled with baited pitfall traps at a downstream and an upstream locality on the two banks of the Mangoro River. The most common species, Nanos binotatus (Canthonini), was sequenced for cytochrome c oxidase subunit I (COI; 804 bp) to characterize within‐population diversity and between‐population genetic differentiation. For the analysis of species geographical range boundaries in relation to the position of the largest rivers on the east coast, a database including all the records for 158 wet forest species was used. The congruence of species range boundaries with the positions of the rivers was tested with a randomization test. Results All common species were found on both sides of the Mangoro River. In Nanos binotatus, haplotype and nucleotide diversities ranged from 0.25 to 0.85 and 0.001 to 0.01, respectively. Population differentiation was high and significant in all comparisons (P < 0.01; average F_ST = 0.61). The differentiation was not significantly higher across than along the river, as would be expected by the riverine barrier hypothesis. There was no indication that the range boundaries of wet forest dung beetle species would generally coincide with the largest rivers in eastern Madagascar. Main conclusions The results provide little support for the riverine barrier hypothesis as an explanation for the current range boundaries of dung beetles in eastern Madagascar. However, extensive deforestation of the coastal regions in eastern Madagascar may have caused a great shrinkage of the ranges of many forest‐dwelling species. Thus the present‐day distributions may not reflect accurately the patterns of the past geographical ranges of the species.     

How did an annual plant react to Pleistocene glaciations? Postglacial history of Rhinanthus angustifolius in Europe

The impact of climate fluctuations during the Pleistocene on the geographic structure of genetic variation in plant populations is well documented, but there is a lack of studies of annual species at the European scale. The present study aimed to infer the history of the widespread European annual Rhinanthus angustifolius C. C. Gmelin (Orobanchaceae). We explored variation in chloroplast DNA (cpDNA) sequences and amplified fragment length polymorphism (AFLP) in twenty-nine populations covering the entire distribution area of the species. Five AFLP groups were identified, suggesting at least two glacial refugial areas: one area in southwestern Europe and one large eastern area in the Balkan/Caucasus. Recolonization of previously glaciated areas mainly took place from the east of Europe. Despite the difference in life-history traits, the patterns found for the annual R. angustifolius show similarities with those of perennial species in terms of genetic diversity and geographic organization of genetic variation. Although organelle markers have typically been preferred in phylogeographic studies, the cpDNA variation in R. angustifolius did not show any clear geographic structure. The absence of geographic structure in the cpDNA variation may reflect persistence of ancestral polymorphisms or hybridization and introgression with closely-related species.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 1–13.     

Speciation mirrors geomorphology and palaeoclimatic history in African laminate-toothed rats (Muridae: Otomyini) of the Otomys denti and Otomys lacustris species-complexes in the 'Montane Circle' of East Africa

We adopted an integrated systematic approach to delimit evolutionary species and describe phylogeographic, morphometric and ecological relationships in Otomys denti (from the Albertine Rift, Southern Rift in Malawi and the northern Eastern Arc Mountains) and Otomys lacustris (from the Southern Rift in Tanzania and Zambia, and the southern Eastern Arc Mountains). Molecular [cytochrome (cyt) b sequences, 1143 bp, N  = 18], craniometric (classical, N  = 100 and geometric, N  = 60) and ecological (Partial Least Squares regression of shape and ecogeographic variables) approaches show a profound, parallel disjunction between two groups: (1) Eastern Arc and Southern Rift (including the Malawi Rift) ( O. lacustris and Otomys denti sungae ) and (2) Albertine Rift ( Otomys denti denti and Otomys denti kempi ) taxa. Within both groups, cyt b sequences or craniometric analysis provided evidence for the differentiation of both southern and northern Eastern Arc from Southern Rift lineages (across the so-called Makambako Gap). Within the Albertine Rift ( denti – kempi ) lineage, populations from individual mountain ranges differed significantly in skull shape (but not size), but were similar genetically. Over-reliance in the past on very few morphological characters (e.g. number of molar laminae) and a polytypic species concept has obscured phylogenetic relationships and species discrimination in this group. We recognize at least three species in this group, and distinct lineages within two of these species. Each species or lineage was endemic to one of three regions: the Albertine Rift, the Malawi Rift or the Eastern Arc. Our result echo conclusions of recent studies of other mammalian and bird taxa and reflect the geomorphology and palaeoclimatic history of the region.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 913–941.     

Molecular taxonomy and phylogeography of Miniopterus schreibersii (Kuhl, 1817) (Chiroptera: Vespertilionidae), in the Eurasian transition

Miniopterus schreibersii is a polytypic bat species, with one of the widest distribution ranges among the mammals. We studied the genetic differentiation and taxonomy of this species in the transition zone between south-eastern Europe and Anatolia (in Asia), where two subspecies have been described. The results indicated a sharp genetic break between the samples from western Anatolia and south-eastern Europe and those of eastern Anatolia. In addition, the samples from western Anatolia and south-eastern Europe were seen to be reciprocally monophyletic, although the differentiation was less drastic. These patterns of genetic differentiation suggest the presence of two distinct groups within the M. schreibersii complex in the region, concordant with previous subspecific recognition. The cause of this genetic break is most likely differentiation in separate glacial refugia followed by secondary contact. However, more samples are needed to assess whether these represent different species, as well as to understand more clearly the causes of this differentiation.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 577–582.     

Natural colonization or introduction? Phylogeographical relationships and morphological differentiation of house geckos (Hemidactylus) from Madagascar

Several house gecko species of the genus Hemidactylus are almost cosmopolitan lizards, with distributions that have probably been shaped by natural transoceanic dispersal as well as by more recent human introductions. Here we revise the Hemidactylus populations of Madagascar and compare them genetically with populations from other sites in the Indian Ocean region. Morphological data strongly confirm the occurrence of three Hemidactylus species on Madagascar: Hemidactylus frenatus , distributed along the western coast of Madagascar; H. platycephalus , restricted to the north-west and the widespread H. mercatorius that occurs throughout the island, including coastal areas at sea level as well as big cities (Antananarivo, Fianarantsoa) at altitudes of 1200–1300 m above sea level. Analyses of partial sequences of the 16S rRNA gene in 46 Hemidactylus specimens from Madagascar, East Africa, South Asia, and the Comoro and Mascarene archipelagos demonstrated the presence of a fourth species, H. brooki, on the Mascarenes (Réunion, Rodrigues, and Mauritius) and Comoros (Moheli). The Malagasy populations of H. platycephalus were genetically uniform and differentiated from the African and Comoroan specimens studied. H. frenatus had a relatively low genetic differentiation over the whole region with no recognizable phylogeographical structure, indicating more recent colonizations or introductions. In contrast, H. mercatorius showed a strong phylogeographical structure of haplotypes, with two distinctly different lineages in Madagascar. Moreover, all Malagasy specimens differed strongly from the single African specimen included. This indicates that populations of H. mercatorius in Madagascar have a long history that predates human settlement.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 115–130.     

Genetic diversity and phylogeography of the European roe deer: the refuge area theory revisited

The extant taxa of central and northern Europe are commonly believed to derive from Pleistocene ancestors, who moved to the north from three separate glacial refugia: the Iberian and Italian peninsulae, as well as the southern Balkans. The issue of postglacial dispersal patterns was addressed through the investigation of population structure and phylogeography of the European roe deer, Capreolus capreolus . The genetic diversity in 376 individuals representing 14 allegedly native populations across their European range was assessed, using ten autosomal microsatellite loci and restriction fragment length polymorphisms of the mitochondrial D-loop and NADH dehydrogenase 1 gene segments. Our results suggest the existence of three major genetic lineages of roe deer in Europe. One comprises populations in the south-western limit of the species' distribution (i.e. Iberia), where an internal substructure splits a northern from a southern sublineage. A second lineage includes populations of southern and eastern Europe, as well as a separate sublineage sampled in central-southern Italy, where the existence of the subspecies Capreolus c. italicus was supported. In central-northern Europe, a third lineage is present, which appeared genetically rather homogeneous, although admixed, and equally divergent from both the eastern and western lineages. Current patterns of intraspecific genetic variation suggest that postglacial recolonization routes of this cervid to northern Europe could be due to range expansion from one or more refugia in central-eastern Europe, rather than proceeding from the Mediterranean areas.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 85–100.     

Genetic and morphological divergence reveals local subdivision of perch (Perca fluviatilis L.)

The level of gene flow is an important factor influencing genetic differentiation between populations. Typically, geographic distance is considered to be the major factor limiting dispersal and should thus only influence the degree of genetic divergence at larger spatial scales. However, recent studies have revealed the possibility for small-scale genetic differentiation, suggesting that the spatial scale considered is pivotal for finding patterns of isolation by distance. To address this question, genetic and morphological differentiation were studied at two spatial scales (range 2–13 km and range 300 m to 2 km) in the perch ( Perca fluviatilis L.) from the east coast archipelago of Sweden, using seven microsatellite loci and geometric morphometrics. We found highly significant genetic differentiation between sampled locations at both scales. At the larger spatial scale, the distance per se was not affecting the level of divergence. At the small scale, however, we found subtle patterns of isolation by distance. In addition, we also found morphological divergence between locations, congruent with a spatial separation at a microgeographic scale, most likely due to phenotypic plasticity. The present study highlights the importance of geographical scale and indicates that there might be a disparity between the dispersal capacity of a species and the actual movement of genes. Thus, how we view the environment and possible barriers to dispersal might have great implications for our ability to fully understand the evolution of genetic differentiation, local adaptation, and, in the end, speciation.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 746–758.     

A new Mantalania species (Rubiaceae) from Madagascar

Mantalania longipedunculata De Block & A.P. Davis, a new species of Rubiaceae (Gardenieae) from Île Sainte Marie, Madagascar, is described, illustrated, and compared with the three other species in the genus. An extinction risk assessment is given, using the categories and criteria of the World Conservation Union (IUCN).  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 421–424.     

High genetic variability in Neotropical myophilous orchids

The patterns of genetic variability of seven Bulbophyllum Thouars species were investigated using 14 enzymatic loci, and compared with those of other Neotropical myophilous orchid species. The genetic variability estimated was very high ( H _e = 0.39–0.61, P  = 86–100%, A  = 2.6–3.8), despite the small population sizes, which may be explained by the vegetative reproduction. Of the three species with multiple populations, only B. epiphytum showed moderate values of genetic structuring. Environmental characteristics and seed dispersal mechanisms are important in understanding the differences in the population structure observed. Almost all pairs of species showed low genetic similarity, indicating a long period of divergence. Our results suggest that B. adiamantinum and B. insectiferum , species currently classified in different sections of the genus, are more closely related than previously thought.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 33–40.     

Different phylogeographic patterns in two Japanese Silpha species (Coleoptera: Silphidae) affected by climatic gradients and topography

To reveal differences in phylogeographic patterns of flightless insect species occurring in different regions of Japan, we studied the phylogeography and demographic history of Silpha beetles occurring in cool-temperate habitats of two major islands, Honshu and Hokkaido, using sequences of the mitochondrial cytochrome oxidase subunit I (COI) gene. Honshu has a more mountainous topography, and cool-temperate habitats occur discontinuously, whereas Hokkaido, located to the north of Honshu, has more continuous cool-temperate habitats. A species endemic to Honshu, S. longicornis occurs on Honshu, whereas S. perforata occurs on Hokkaido and the East Asian continent. Our results indicate that the ancestors of S. longicornis colonized Honshu via a south-west route c . 0.7 Mya and the species has highly divergent populations in isolated mountainous areas of Honshu, whereas S. perforata colonized Hokkaido via a northern route less than 90 000 years ago and has less divergent geographic populations. During the last glacial period, S. perforata was probably restricted to refugia in southern Hokkaido and later expanded into northern Hokkaido, whereas S. longicornis populations existed in many isolated refugia, probably because of the complex topography of Honshu. Thus, our study demonstrates that, even between closely related species, interactions among biology, latitudinal climatic gradients and topography can produce different phylogeographic patterns.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 452–467.     

Markedly discordant mitochondrial DNA and allozyme phylogenies of tube-nosed fruit bats, Nyctimene, at the Australian–Oriental biogeographical interface

Australo-Papuan tube-nosed fruit bats of the genus Nyctimene reach their most westerly distribution on the island archipelagos of eastern Indonesia. A recent morphological examination indicates three species occur on Moluccan islands. Both allozyme electrophoresis and mitochondrial DNA (mtDNA) sequencing reveal there are only two species present, Nyctimene cephalotes and Nyctimene albiventer , but there is considerable disparity between the results obtained from the two genetic data sets. Allozyme data indicate N. albiventer occurs on Wokam, which sits on the Australian continental shelf and was joined to Australo-Papua during the last glacial maximum, and N. cephalotes on the other Moluccan islands, all of which are off the continental shelf. Divergence of these two species is dated at approximately 1.5 Mya. By contrast, the mtDNA gene tree shows two deep clades, one containing all specimens from Wokam and Yamdena, and the second all the specimens from the other islands. This especially marked incongruence between the two genetic data sets is ascribed to either a single introgression event of N. albiventer mtDNA into N. cephalotes on Yamdena at around 100 000 ya or lineage sorting of very old mtDNA lineages that coalesce a rather long time before the speciation event. These results highlight that caution should be exercised when relying on mtDNA as barcodes in species taxonomy. The continental-associated N. albiventer on Wokam has higher levels of allozyme heterozygosity and mtDNA nucleotide diversity than the N. cephalotes populations occurring on the more remote islands, indicating that this colonizing species has experienced bottlenecks and/or a low effective population size since speciation.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 589–602.     

Low genetic diversity and allozymic evidence for autopolyploidy in the tetraploid Pyrenean endemic larkspur Delphinium montanum (Ranunculaceae)

Allozyme electrophoresis was conducted to survey the levels and distribution of genetic diversity in the tetraploid perennial larkspur Delphinium montanum (Ranunculaceae), which is endemic to the eastern Pyrenees of Spain and France and considered to be endangered. Seven populations were sampled, resolving 14 putative enzymatic loci belonging to eight enzymes. Banding patterns stained in gels revealed several enzymatic duplications attributable to autotetraploidy, such as the presence of both balanced and unbalanced heterozygotes and the lack of fixed heterozygosity. However, variability in D. montanum ( P  = 23.8%, A  = 1.48, and H _e = 0.082) was lower than that expected for an autotetraploid species. This, in addition to the scarcity of loci showing three or four different alleles, could indicate that this species has suffered genetic erosion through population bottlenecks, or, alternatively, that it is undergoing diploidization.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 211–222.     

Glacial history of high alpine Ranunculus glacialis (Ranunculaceae) in the European Alps in a comparative phylogeographical context

There is considerable controversy concerning the fate of Alpine plants during Pleistocene glaciations. While some studies have found evidence for nunatak survival, others have explained the present genetic patterns by survival only in peripheral refugia. We investigated 75 populations of high alpine Ranunculus glacialis from its entire Alpine distribution. Phylogeographical analyses of AFLP data revealed four groups of populations. Two of them, located in the western Alps, were genetically isolated from each other and from the eastern groups, whereas the two eastern Alpine groups were genetically more similar to each other. This suggests longer isolation and/or lower levels of gene flow in the two western groups. As all groups are close to, or overlap with, presumed glacial refugia, invoking glacial survival on nunataks is unnecessary to explain the present genetic pattern. Similar to the phylogeographical patterns of R. glacialis , the previously investigated alpine Phyteuma globulariifolium and Androsace alpina , which are also confined to siliceous bedrock, showed strong geographical affinities to peripheral refugial areas and there were large-scale congruencies in the location of these refugia for all three species.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 183–195.     

Sound production in four damselfish (Dascyllus) species: phyletic relationships?

Most studies of fish sounds show that the sounds are species-specific, with unique spectral and timing characteristics. This raises the question as to whether these sounds can be used to understand phyletic relationships between species and which acoustic parameters are subject to variation between species. In the present study, 597 sounds (and 2540 pulses) related to signal jumps of four Dascyllus species ( Dascyllus aruanus , Dascyllus trimaculatus , Dascyllus albisella , and Dascyllus flavicaudus ) from different geographic regions (Madagascar, Moorea, Rangiroa, and Hawaii) were analysed. It was possible to discern species-specific sounds, but also variation in sounds between populations. Large variations in sound length were found between Dascyllus species, whereas differences in interpulse duration were found to be variable between populations. In the regions where species live in sympatry, it appears that they restrict the variability in their sounds. This could comprise evidence of adaptation with character displacement of sonic characteristics where different species co-occur. However, sonic characteristics still overlapped substantially between species, suggesting that females would need to sample more than one sound and potentially use other cues to discriminate between species.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 928–940.     

A taxonomic revision of the genus Flagenium Baill. (Rubiaceae–Octotropideae)

A taxonomic revision of the genus Flagenium Baill. is presented. Flagenium is endemic to Madagascar and comprises six species; three new species are described here ( F. farafanganensis , F. petrikensis , and F. pedunculatum ) and one species ( F. arboreum ) is transferred into synonymy. Each species is fully described, and summaries of distribution, habitat and ecology, and phenology are given; conservation assessments are provided for each species. Flagenium is characterized by having ovaries with at least two erect and two pendulous ovules per locule, which, depending on the species, can display an additional two to four horizontal ovules per locule arranged between the uppermost and lowermost ovules. In addition, Flagenium has a longitudinally ten-ribbed fruit. These features are potentially unique within the Octotropideae.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 557–570.     

The importance of looking at small-scale patterns when inferring Gondwanan biogeography: a case study of the centipede Paralamyctes (Chilopoda, Lithobiomorpha, Henicopidae)

Gondwanan biogeography has fascinated zoologists and botanists for over a century, but most biogeographical work has used continent-scale areas as analytical units. More finely resolved patterns, as can be obtained from small invertebrates with limited dispersal abilities, will be obscured in those studies. A common case is treating Australia as a single biogeographical region. In the present study, the necessity of splitting Australia into multiple microareas is demonstrated using centipedes as an example. The lithobiomorph centipede Paralamyctes is distributed on fragments of Gondwana, with species in southern Africa, Madagascar, southern India, Patagonia, eastern Australia, and New Zealand. A cladogram for Paralamyctes is based on morphology and sequences for four molecular markers for 30 terminals that sample 20 of 26 known ingroup species and four outgroups. Analysis with direct optimization across a range of indel costs and transversion : transition cost ratios identifies two main clades: Paralamyctes ( Paralamyctes ) unites species from southern Africa, Madagascar, tropical and warm temperate Australia, and New Zealand. The other group includes the temperate Australian/New Zealand Paralamyctes ( Haasiella ) and Paralamyctes ( Thingathinga ) and a Chilean clade. Subtree analysis finds that different parts of Australia have closest affinities to other Gondwanan fragments, and some of these relationships (such as that between north Queensland and New Zealand) are based on taxonomically stable clades. Area delimitation for large continental fragments should use sufficiently fine resolution to test the 'monophyly' of those fragments and attempt to eliminate spurious geographical paralogy.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 65–78.     

Population structure in an isolated Arctic fox, Vulpes lagopus, population: the impact of geographical barriers

The genetic composition of a population reflects several aspects of the organism and its environment. The Icelandic Arctic fox population exceeds 8000 individuals and is comprised of both coastal and inland foxes. Several factors may affect within-population movement and subsequent genetic population structure. A narrow isthmus and sheep-proof fences may prevent movement between the north-western and central part and glacial rivers may reduce movement between the eastern and central part of Iceland. Moreover, population density and habitat characteristics can influence movement behaviour further. Here, we investigate the genetic structure in the Icelandic Arctic fox population ( n  = 108) using 10 microsatellite loci. Despite large glacial rivers, we found low divergence between the central and eastern part, suggesting extensive movement between these areas. However, both model- and frequency-based analyses suggest that the north-western part is genetically differentiated from the rest of Iceland (F_ST = 0.04, D_S = 0.094), corresponding to 100–200 generations of complete isolation. This suggests that the fences cannot be the sole cause of divergence. Rather, the isthmus causes limited movement between the regions, implying that protection in the Hornstrandir Nature Reserve has a minimal impact on Arctic fox population size in the rest of Iceland.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 18–26.