Contrasting temporal dynamics and spatial patterns of population genetic structure correlate with differences in demography and habitat between two closely-related African freshwater snails

The relationship between habitat stability, demography, and population genetic structure was explored by comparing temporal microsatellite variability spanning a decade in two closely-related hermaphroditic freshwater snails from Cameroon, Bulinus forskalii and Bulinus camerunensis . Although both species show similar levels of preferential selfing, microsatellite analysis revealed significantly greater allelic richness and gene diversity in populations of the highly endemic B. camerunensis compared to those of the geographically-widespread B. forskalii . Additionally, B. camerunensis populations showed significantly lower spatial genetic differentiation, higher dispersal rates, and greater temporal stability compared to B. forskalii populations over a similar spatial scale. This suggests that a more stable demography and greater gene flow account for the elevated genetic diversity observed in this geographically-restricted snail. This contrasts sharply with a metapopulation model (which includes extinction/contraction, recolonization/expansion, and passive dispersal) invoked to account for population structuring in B. forskalii . As intermediate hosts for medically important schistosome parasites, these findings have ramifications for determining the scale at which local adaptation may occur in the coevolution of these snails and their parasites.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 747–760.     

Relationship between microspatial population genetic structure and habitat heterogeneity in Pomatias elegans (O.F. Müller 1774) (Caenogastropoda, Pomatiasidae)

In the present study the population genetic structure of the terrestrial snail Pomatias elegans was related to habitat structure on a microspatial scale. The genetic variability of 1607 individuals from 51 sampling sites in five different populations in Provence, France, was studied with an allozyme marker using population genetic methods, Mantel tests and spatial autocorrelation techniques were applied to different connectivity networks accounting for the structural features of the landscape. It is suggested that the population structure is, to a large extent, a function of the habitat quality, quantified as population density, and of the spatial arrangement of the habitat in the landscape and not of the geographical distance per se . In fragmented habitats, random genetic drift was the prevailing force for sampling sites separated by a few hundred meters.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 565–575.     

Selection and disequilibrium in Cepaea nemoralis

The snail Cepaea nemoralis (L.) is usually polymorphic for linked loci conferring pink or yellow shell ground colour and banding or lack of bands. An unlinked locus, mid‐banded, modifies the appearance of banded shells. These characters have a very wide range of frequencies across the species range and even within limited areas. In Britain, frequencies differ between populations from shaded woodland habitats and those from closely adjacent open habitats. Using data from the Evolution MegaLab Project, it is shown that the greater frequency of pink unbanded in shaded habitats is associated with a greater excess of the favoured allele combinations, measured as linkage disequilibrium. Examination of the relationship of disequilibrium to allele frequency in samples from the full geographical range and all habitats and climates shows that, in general, the allele combinations at high frequencies are in gametic excess. This suggests that selection tends to diversify rather than move frequencies towards stable equilibria. The result is important in relation to the as yet unresolved question how the polymorphism is maintained. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Dispersal history of a spider (Stegodyphus lineatus) across contiguous deserts: vicariance and range expansion

Israel marks a crossroads between three continents encompassing several phytogeographical and zoogeographical zones. In this complex area, the flow of species from different biogeographical regions creates opportunities to study how geographical division and colonization routes affect current distribution and structure of resident populations of organisms associated with desert and arid environments, habitats that may have persisted throughout Pleistocene glacial periods. The present paper analyses the population history of the spider Stegodyphus lineatus in the contiguous Negev and Judean deserts in Israel using allozyme and mtDNA variation. The distinct patterns of variation indicate that Judean and Negev populations are vicariant lineages. The residence time was longer in Judea, where populations were more polymorphic for mtDNA, showed isolation by distance and were less structured than in the Negev. The Negev population, possibly linked to other Mediterranean populations of S. lineatus , consisted of two subdivisions derived from a recent eastward expansion across the central Negev watershed. Despite differences in age and level of structure, all lineages show similar dispersal processes dominated by restricted gene flow. The distribution patterns of allozyme and mtDNA markers are unrelated to geographical patterns of precipitation and vegetation. Rather, they follow large-scale topographic features, namely the water divide between Mediterranean and Afro-Syrian rift drainages and between eastern and western Negev drainages.   © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 739–754.     

Sex versus parasitism versus density

The Red Queen hypothesis predicts that sexual reproduction should be favoured when the risk of infection by parasites is high. However, this prediction may also be achieved independently by means of the reproductive assurance and the density-dependent transmission hypotheses because higher densities increase the chances of fertilization (i.e. mates are easier to find) and increase the risk of infection. Additionally, the Red Queen hypothesis makes a long-term prediction that infection should be genotype-specific, with time-lagged dynamics. In the present study, we attempt to consolidate these two predictions by investigating whether sexuality and parasitism alternate in a time-lagged fashion on an ecological time-scale. In a 3-year study, we examined the time-lagged dynamics between male frequencies, infection levels (by Centrocestus sp.), and snail densities of four natural populations of sexual and asexual Melanoides tuberculata snails. The results obtained provide no evidence of either prediction of the Red Queen hypothesis. We found partial support for reproductive assurance in a single population, and could not discriminate between the three hypotheses in another population. The remaining two populations did not support any of the hypotheses in question.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 537–544.     

Microgeographical diversification of threespine stickleback: body shape–habitat correlations in a small, ecologically diverse Alaskan drainage

Adaptive radiations are a major source of evolutionary diversity in nature, and understanding how they originate and how organisms diversify during the early stages of adaptive radiation is a major problem in evolutionary biology. The relationship between habitat type and body shape variation was investigated in a postglacial radiation of threespine stickleback in the upper Fish Creek drainage of Cook Inlet, Alaska. Although small, the upper Fish Creek drainage includes ecologically diverse lakes and streams in close proximity to one another that harbour abundant stickleback. Specimens from ancestral anadromous and derived resident freshwater populations differed substantially and could be distinguished by body shape alone, suggesting that the initial stages of adaptation contribute disproportionately to evolutionary divergence. Body shape divergence among resident freshwater populations was also considerable, and phenotypic distances among samples from freshwater populations were associated with habitat type but not geographical distance. As expected, stream stickleback from slow-moving, structurally complex environments tended to have the deepest bodies, stickleback from lakes with a mostly benthic habitat were similar but less extreme, and stickleback from lakes with a mostly limnetic habitat were the most shallow-bodied, elongate fish. Beyond adapting rapidly to conditions in freshwater environments, stickleback can diversify rapidly over small geographical scales in freshwater systems despite opportunities for gene flow. This study highlights the importance of ecological heterogeneity over small geographical scales for evolutionary diversification during the early stages of adaptive radiation, and lays the foundation for future research on this ecologically diverse, postglacial system.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 139–151.     

Do current hypotheses explain continental and seasonal variation in female morph frequencies of the damselfly,Nehalennia irene?

Female-limited colour polymorphism occurs in many damselfly species, where one morph resembles the male (andromorph) and the other is dissimilar (gynomorph). Explanations for this phenomenon vary, but most assume that andromorphism has arisen in odonates, as a response to excessive male harassment. Here, we quantify the extent of continental and seasonal variation in female morph frequencies in a widely-distributed damselfly and ask whether the spatiotemporal patterns in andromorph frequency can be understood on the basis of sexual harassment theory. We sampled the damselfly, Nehalennia irene (Hagen) among regions across Canada, and at several sites, over the reproductive season, within Central Canada. Andromorph frequencies ranged from 0 to > 90% across Canada. In particular, sites in Western Canada had consistently high andromorph frequencies, whereas andromorph frequencies among Central sites were lower and variable and, among Eastern sites, were lower still (except one site) and relatively invariant. For populations in Central Canada, both andromorph frequencies and population densities varied significantly over time, reaching a peak mid-season. As expected, morph frequency covaried significantly with estimates of male harassment in some cases, but estimates of male harassment did not consistently account for variation in morph frequencies within all regions. Additional factors such as genetic drift may influence morph frequency at the edge of a species' range. Future work also should test, and attempt to explain causation, for seasonal variation in morph frequency.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 501–508.     

Genetic differentiation on multiple spatial scales in an ecotype-forming marine snail with limited dispersal: Littorina saxatilis

The population genetic structure of marine species lacking free-swimming larvae is expected to be strongly affected by random genetic drift among populations, resulting in genetic isolation by geographical distance. At the same time, ecological separation over microhabitats followed by direct selection on those parts of the genome that affect adaptation might also be strong. Here, we address the question of how the relative importance of stochastic vs. selective structuring forces varies at different geographical scales. We use microsatellite DNA and allozyme data from samples of the marine rocky shore snail Littorina saxatilis over distance scales ranging from metres to 1000 km, and we show that genetic drift is the most important structuring evolutionary force at distances > 1 km. On smaller geographical scales (< 1 km), divergent selection between contrasting habitats affects population genetic structure by impeding gene flow over microhabitat borders (microsatellite structure), or by directly favouring specific alleles of selected loci (allozyme structure). The results suggest that evolutionary drivers of population genetic structure cannot a priori be assumed to be equally important at different geographical scales. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 31–40.     

Heterosis in Cepaea

A test has been made of the association of heterozygosity with shell breadth in the polymorphic snail Cepaea nemoralis. The material was collected by C. B. Goodhart from a series of paired sites at which individuals reached different adult breadths. Dominant phenotypes, in which a large fraction was heterozygous, had a greater breadth and lower variance than recessive phenotypes regardless of whether the measurement was of shell ground colour, banding or the double recessive vs. the rest. Most of the difference was contributed by samples from the habitat where animals reached the largest size. The result is consistent with existence of heterotic sections of chromosome that include the colour and banding loci, and may help to explain the persistence of the polymorphism.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 49–53.     

Colonization,stability, and adaptation in a transplant experiment of the polymorphic land snail Cepaea nemoralis (Gastropoda: Pulmonata) at the edge of its geographical range

At the eastern margins of the geographical distribution in Europe, populations of Cepaea nemoralis are sparse and limited to urban environments to which they are possibly confined by relatively warmer climates. In 1999 we introduced 1101 C. nemoralis individuals originating from nine urban populations to a rural location in the area. The snails established a viable population, which suggests that confinement to urban settings is dispersal‐ rather than climate‐limited. The snails filled available habitats at a rate of approximately 400–600 m² year^?1. On the whole, morph frequencies remained remarkably stable; changes that occurred are attributable to segregation of alleles or chromosomes. However, snails responded to habitat heterogeneity: consistent and predictable divergence occurred between habitat types, such that light‐shelled snails were repeatedly more frequent in the open than in adjoining shaded habitats. This suggests the operation of climatic and/or visual selection. As the whole area encompassing seven distinct habitat patches was only 0.3 ha, and the maximum duration of population divergence was only 11 years (fewer than four snail generations), these results indicate extremely small temporal and spatial scales of adaptation during initial phases of population establishment and spread. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 462–470.     

The importance of energetic versus pelvic constraints on reproductive allocation by the eastern fence lizard (Sceloporus undulatus)

In ectothermic species, females often produce larger eggs in colder environments. Models based on energetic constraints suggest that this pattern is an adaptation to compensate for the slower growth of offspring in the cold. Yet, females in cold environments also tend to be larger than females in warm environments. Consequently, thermal clines in egg size could be caused by pelvic constraints, which stem from the inability of large eggs to pass through a small pelvic aperture. Models based on energetic constraints and models based on pelvic constraints predict similar relationships between maternal size and egg size. However, pelvic constraints should produce these relationships both within and among populations, whereas energetic constraints would not necessarily do so. If pelvic constraints are important, we might also expect small females to compensate by producing eggs that are relatively rich in lipids (i.e. high energy density). The present study aimed to assess whether energetic or pelvic constraints generate geographical variation in egg size of the lizard Sceloporus undulatus . Pelvic width is very highly correlated with body length in S. undulatus , making maternal size a suitable measure of pelvic constraint. Although maternal size and egg mass (dry and wet) covaried among populations, these variables were generally not related within populations. Energetic density of eggs tended to increase with decreasing egg mass (dry and wet), but this relationship was strongest in populations where no relationship between maternal size and egg mass was observed. Our results do not support the pelvic constraint model and thus indicate energetic constraints play a greater role in generating geographical variation in egg size.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 513–521.     

Population genetic structure of New Zealand's endemic corophiid amphipods: evidence for allopatric speciation

Allozyme electrophoresis was used to examine population genetic structure at inter- and intraspecific levels for the New Zealand endemic corophiid amphipods, Paracorophium lucasi and P. excavatum. Individuals were collected from estuarine and freshwater habitats from North, South and Chatham Islands. Analyses of genetic structure among interspecific populations indicated clear allelic differentiation between the two Paracorophium species (Nei's genetic distance, D  = 1.62), as well as considerable intraspecific substructuring ( D  = 0.15–0.65). These levels of divergence are similar to interspecific levels for other amphipods and it is proposed that at least two groups from the P. lucasi complex and three from the P. excavatum complex correspond to sibling species. In most cases allopatry can account for the differentiation among the putative sibling species. For populations that share a common coastline we found low levels of differentiation and little or no correlation with geographical distance, suggesting that gene flow is adequate to maintain homogeneous population genetic structure. By contrast, populations on separate coastlines (i.e. isolated by land) showed moderate levels of geographical differentiation indicating restricted gene flow. The juxtaposition of population genetic and biogeographical data for Paracorophium in conjunction with the geological record infers past histories of glacial extirpation, and possible isolating effects of sea-level and landmass changes that have occurred throughout the Plio-Pleistocene.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 119–133.     

Osteology and ecology of Megantereon cultridens SE311 (Mammalia; Felidae; Machairodontinae), a sabrecat from the Late Pliocene – Early Pleistocene of Senéze, France

Megantereon cultridens was a derived, Late Pliocene to Early Pleistocene sabrecat, and although fossils of animals referred to the genus and species have been found at several locations throughout Eurasia, most are fragmentary. However, the specimen SE311 from Senéze in France represents an almost complete and well-preserved skeleton, and this is only known from very few other sabrecats, thus providing a rare glimpse into the full anatomy of an unusual and derived sabrecat. In this monograph, we provide a complete overview of the anatomy of Megantereon cultridens SE311, and compare it with extant large felids, and the few other derived sabrecats from which ample fossil material is known, although most frequently representing several specimens. SE311 was a large specimen of M. cultridens and would have had a body mass of 100–110 kg and a head–body length or around 160 cm, which is similar to a small lioness or large male jaguar. Megantereon sp. were sexually dimorphic, and the size of SE311 suggests that it was a male. As with several other derived sabrecats, it was powerfully built, and had particularly robust forequarters with very well-developed muscle attachment sites, indicating a powerful forelimb and shoulder musculature. The neck was proportionally much longer than in extant felids, and the thoracic and particularly lumbar region was proportionally shorter, mimicking the condition in other derived sabrecats from which large parts of the vertebral column is known. Megantereon probably lived in open-forest environments and preyed on cervids, which were dispatched with a shearing bite from the hypertrophied and blade-like upper canines to the throat of the prey, while the prey was held immobile with the massive forelimbs, thus minimizing the risk of damage to the fangs.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 833–884.     

Subdivided population structure and phylogeography of an endangered freshwater snail, Notopala sublineata (Conrad, 1850) (Gastropoda: Viviparidae), in Western Queensland, Australia

Populations of the Australian freshwater snail Notopala sublineata (Conrad, 1850) have declined rapidly over the last decade, but are still abundant in most river systems of Western Queensland. These rivers are characterized by the unpredictable and highly variable nature of their climatic and hydrological regimes, with episodic periods of very large flow and many periods of little or no flow. We used mitochondrial sequences and allozymes to investigate the genetic structure and infer patterns of dispersal of N. sublineata within this unique environment. We sampled 24 waterholes throughout the four major catchments of the Lake Eyre Basin. Based on a 457-bp fragment of the mitochondrial cytochrome oxidase subunit I gene, we identified 55 haplotypes in a sample of 256 individuals. Both nuclear and mitochondrial genetic datasets indicated high levels of genetic subdivision and restricted gene flow among populations within and among catchments. The mitochondrial haplotypes clustered into two main geographical clades, corresponding with two groups of adjacent catchments: Cooper–Bulloo and Diamantina–Georgina, which appear to have diverged 300 000 years ago. Populations of N. sublineata within these adjacent catchments seem to have diverged relatively recently, roughly 130 000 years ago. Contemporary dispersal seems to be absent between catchments but we suggest that climate fluctuations during the Pleistocene resulted in extensive floods that promoted historical movement of aquatic organisms across catchment boundaries.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 1–16.     

The influence of recent geography, palaeogeography and climate on the composition of the fauna of the central Aegean Islands

Partial Mantel tests and structural equation models were used to investigate the influence of recent geography, palaeogeography and climate on the composition of the fauna of the central Aegean Islands. The composition of land snail and isopod island faunas was significantly influenced by recent and by Pliocene geography. Only Pleistocene palaeogeography had a significant influence on the composition of tenebrionid beetle island faunas. The composition of butterfly island faunas was influenced by recent and by Miocene geographical distances. The composition of reptile island faunas was correlated with recent and Pliocene geography as well as with Pleistocene and/or Miocene geographical distances. Island area influenced only the composition of the island faunas of the volant butterflies, and not that of the less mobile land snails, land isopods, tenebrionid beetles and reptiles. This might indicate that butterflies are able to colonize large islands with suitable habitats even if such islands are comparatively far from source areas more frequently than can the nonvolant groups. Influence of a climatic parameter, namely annual precipitation, on faunal composition was found only for reptiles.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 785–795.     

Evolution of jumping capacity in Tropidurinae lizards: does habitat complexity influence obstacle‐crossing ability?

Jumping performance is relevant for lizards in many ecological contexts and might be favoured during the colonization of structurally complex habitats. Although ground-dwelling lizards use jumps to overcome small obstacles in their natural environments, jumping capacity has been mostly studied in arboreal species. Here, we analysed the evolution of jumping behaviour and performance in lizards from eight ground-dwelling species of Tropidurinae attempting to cross obstacles of different heights in a jumping track, both when undisturbed and under continuous stimulation. To establish ecological correlates with habitat complexity, individuals from two contrasting Brazilian habitats, the arid Caatingas (sand species) and the savannah-like Cerrados (rock species), were compared. Rock species jumped more often and crossed higher obstacles than sand ones in both tests, and performed more vertical than horizontal jumps. Although sand species performed less jumps, they were more successful at crossing the obstacles presented in comparison with rock species. Phylogenetic analyses confirmed these findings and demonstrated a large divergence in jumping capacity between sister-species from different habitats. Therefore, the differences in propensity and endurance for jumping activity appear to be independent of phylogenetic relationships in Tropidurinae and likely reflect an adaptation to the contrasting environments inhabited. The ecological implications of these findings are discussed.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 393–402.     

Terrestrial versus aquatic phenotypes correlate with hydrological predictability of habitats in a semiterrestrial salamander (Urodela, Plethodontidae)

Terrestrial adaptations by small, desmognathine salamanders in Appalachian stream communities have been viewed as evolved responses to large, aquatic, predatory congeners. We tested the role of moisture in the development of adaptive phenotypes in the small, semiterrestrial Desmognathus ocoee . We measured indicators of relative terrestrial vs. aquatic phenotypes (i.e. tolerance of desiccation, anatomical depression of body/tail) using individuals from stream habitats having different flow regimes within each of three geographical locations. Salamanders from intermittent, first-order streams exhibited greater desiccation tolerance, took longer to reach tolerance limits, and had more anatomically depressed bodies/tails than those from second-order, perennial streams, regardless of location. Phenotypes of salamanders from second-order streams showed aquatic tendencies. Phenotypic variation was consistent with relative habitat stability and not with relative abundance of large congeners. MtDNA analysis revealed that individuals grouped together based on geographical location and not habitat type, implying that the observed phenotypic differences arose in situ at each geographical location. Recent phylogenetic reconstructions of the Desmognathus indicate that large, aquatic forms are derived, and their appearance corresponds with the late Cenozoic development of modern Appalachian topography. We propose that radiation of desmognathine lifestyles and associated phenotypes tracked the formation of moisture-stability gradients associated with the creation of cool, wet, montane habitats.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 227–238.     

Risk of predation as a promoting factor of species divergence in threespine sticklebacks (Gasterosteus aculeatus L.)

Icelandic freshwater systems are geologically young and contain only six species of freshwater fish. As these species colonized Icelandic fresh waters they were presented with a diversity of unique, uncontested habitats and food resources, promoting the evolution of new behaviour strategies crucial to the formation of new morphs and speciation. To determine the likelihood that predation threat could affect the antipredator behaviour and possibly the sympatric divergence of prey populations, we analysed antipredator behaviour of seven groups of Icelandic threespine sticklebacks ( Gasterosteus aculeatus ): two marine groups, one group from a lake without piscine predators, and two polymorphic lake populations, each with two groups occupying unique habitats. Shoaling cohesion, school formation and duration, and vigilance in predator inspection/avoidance behaviour varied greatly among groups. The differences appeared to be related to the risk of predation as well as to opportunities and constraints set by the different habitats. Antipredator behaviour was especially pronounced and differed extensively in two polymorphic forms from the lake Thingvallavatn, where predation risk is very high. By keeping the two morphs separate in their respective habitats, high predation risk may be a contributing factor in promoting the habitat-specific divergence of G. aculeatus seen in the lake. This suggests that in situations where refuge habitats are spatially separated, the risk of predation may contribute to the evolution of separate sympatric forms of small fish such as G. aculeatus .  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 189–203.     

Diversity patterns of the terrestrial snail fauna of Nyungwe Forest National Park (Rwanda), a Pleistocene refugium in the heart of Africa

We investigated the land snail fauna of Nyungwe Forest National Park in south‐western Rwanda. Fifty plots at altitudes between 1718 and 2573 m were studied. In total, 3461 specimens were collected and were assigned to 102 land snail species. With respect to land snail species, Nyungwe Forest is the richest forest known in Africa. A comparison with other forests in the northern Albertine Rift indicates that land snail species richness in this region is significantly correlated with distance from Pleistocene forest refugia. The high beta diversity in Nyungwe is the result of a high species turnover between sites, which has biogeographical and ecological origins. Nyungwe Forest is situated on the Congo–Nile divide where species of different geographical origin may meet. Moreover, Nyungwe Forest offers a high diversity of habitats because it extends across a wide range of altitudinal zones. Species richness decreased with increasing altitude. It was also correlated with the presence of bare rocks that offer additional microhabitats and shelter. Although the occurrences of different land snail species in Nyungwe Forest were significantly clustered, only a minority of the species could be assigned to a group of species with similar occurrences. The majority of the species respond individualistically to environmental variables. The significant nestedness of the occurrences of the land snail species in Nyungwe was mainly correlated with altitude. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 363–375.     

A new species of Ilex (Aquifoliaceae) from Espinhaço Range, Bahia, Brazil

A new species, Ilex mucugensis Groppo sp. nov. (Aquifoliaceae), is described and illustrated. The species is known only from 'campos rupestres' (vegetation of rock outcrops at high altitude) in the vicinity of Mucugê, Bahia State, Brazil, in the mountains of the Espinhaço Range. Comments on affinities, geographical distribution and phenology of the new species are also provided.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 153–156.