Alternative models of conspecific attraction in flies and crabs

Animals are often attracted to one another when selecting habitats,but little is known about the rules governing conspecific attraction.We use Akaike Information Criterion to evaluate alternativemodels of the effects of conspecifics on individual choice inthe context of habitat selection. One set of models was testedusing data collected on virgin female flies, Drosophila melanogaster,selecting habitats in the laboratory; a second set of modelswas tested using data collected on crabs, Ocypode rotundata,selecting foraging patches in the field. Patterns of space usein the flies were most consistent with models indicating thatindividuals were attracted to other females that selected trapsduring the same hour, as well as to cues left by females thathad entered traps during the previous hour. Results for thecrabs were most consistent with a model which assumes that individualspreferred to join the patch with the most crabs but that theirability to assess the number of crabs in alternative patcheswas constrained by basic psychophysical principles (Weber'slaw). These results provide support for hypotheses about thefunctional significance of conspecific attraction in the fliesand the crabs and illustrate the richness of information aboutconspecific attraction that can be obtained when the same dataset is confronted with alternative models of the ways that animalsrespond to one another when selecting habitats.     

The role of ecological knowledge in explaining biogeography and biodiversity in Amazonia

Biogeographical studies in Amazonia have commonly taken a historical, rather than an ecological approach. General patterns have been sought in the distribution maps of different species, and these have been explained in terms of past or present distribution barriers, especially past climates and large rivers. Implicitly, and often also explicitly, it is assumed that Amazonia is ecologically so uniform that present-day ecological conditions are rather insignificant in determining species distribution patterns and speciation. However, this assumption is more based on the lack of relevant data than on actual observations of environmental uniformity or ecological unspecialization of the species. Recent studies have indeed documented ecological heterogeneity and floristic differences among sites that were previously thought similar. In the absence of direct knowledge of the past, more complete ecological and environmental understanding of the present-day Amazonia are needed for evaluating the relative roles of historical and ecological factors in Amazonian biogeography and biodiversity.     

The phlebotomine sandflies of Venezuelan Amazonia

Phlebotomine sandflies were surveyed in two ecologically contrasted areas, the northern moist and southern wet tropical forests, of the Territorio Federal Amazonas, Venezuela. Three new taxa and twenty-one new records were added to the previously known species list for Venezuelan sandflies, which now totals eighty species. Both sexes of Lutzomyia (Psychodopygus) killicki sp.n., L. (Trichophoryomyia) bettinii sp.n., L. (Nyssomyia) olmeca reducta subsp.n. and and the females of L. bernalei Osorno et al., Brumptomyia pintoi Costa Lima and L. begonae (Ortiz & Torres) are described and illustrated.     

Soil carbon dynamics in regrowing forest of eastern Amazonia

The future flora of Amazonia will include significant areas of secondary forest as degraded pastures are abandoned and secondary succession proceeds. The rate at which secondary forests regain carbon (C) stocks and re-establish biogeochemical cycles that resemble those of primary forests will influence the biogeochemistry of the region. Most studies have focused on the effects of deforestation on biogeochemical cycles. In this study, we present data on the recuperation of carbon stocks and carbon fluxes within a secondary forest of the eastern Amazon, and we compare these measurements to those for primary forest, degraded pasture, and productive pasture. Along a transect from a 23-y-old degraded pasture, through a 7-y-old secondary forest, through a 16-year-old secondary forest, and to a primary forest, the δ¹³C values of soil organic matter (SOM) in the top 10 cm of soil were – 21.0, – 26.5, – 27.4, and – 27.9‰, respectively, indicating that the isotopic signature of SOM from C3 forest plants was rapidly re-established. The degraded pasture also had significant inputs of C from C3 plants. Radiocarbon data indicated that most of the C in the top 10 cm of soil had been fixed by plants during the last 30 years. Differences in soil C inventory among land use types were small compared to uncertainties in their measurement. Root inputs were nearly identical in primary and secondary forests, and litterfall in the secondary forest was 88% of the litterfall rate of the primary forest. In contrast, the secondary forest had only 17% of the above ground biomass. Because of rapid cycling rates of soil C and rapid recovery of C fluxes to and from the soil, the below ground C cycle in this secondary forest was nearly identical with those of the unaltered primary forest.     

Effects of forest disturbance on the structure of ground-foraging ant communities in central Amazonia

This study evaluates biotic responses, using ants as bio-indicators, to relatively recent anthropogenic disturbances to mature forest in central Amazonia. The structure of the ground-foraging ant community was compared in four habitats that represented a gradient of disturbance associated with differences in land use. Ants were collected in undisturbed, mature forest, in an abandoned pasture, in a young regrowth forest (situated in a former pasture area), and in an old regrowth forest (established where mature forest was just cleared and abandoned). More ant species were found in mature and old regrowth forest than in the abandoned pasture. By contrast, ant abundance tended to decrease with forest maturity. Both pasture and young regrowth forest exhibited a distinct ant species composition compared to mature forest, whereas species composition in the old regrowth forest showed greater similarity to that of mature forest. In spite of differences in fallow time between former pasture areas and non-pasture areas, there is evidence that different land-management practices do result in different rates of recovery of the ant forest fauna after land abandonment. In any case, recuperation of the ground-foraging ant fauna appears to be faster than regeneration of the woody-plant community. In this sense, regrowth forests may be valuable for the conservation of ground-foraging ants and perhaps for other components of mature-forest leaf-litter fauna within the context of a fragmented landscape.     

Alternative adaptations, sympatric speciation and the evolution of parasitic, inquiline ants

Inquiline ant species are workerless social parasites whose queens reproduce in colonies of other species alongside the host queens. Inquilines arise either when one non-parasitic species evolves into an inquiline parasite of another non-parasitic species (the interspecific hypothesis), or by the speciation of intraspecific inquilines from their host stock (the intraspecific hypothesis): it is unlikely that inquilines evolve from other forms of social parasite. This paper reviews the evidence for and against the inter-and intraspecific hypotheses. All inquilines are close phylogenetic relatives of their host species (loose ‘Emery's rule’), and some are their host's closest relative (strict ‘Emery's rule’). A problem for the interspecific hypothesis is how to explain the strict Emery's rule, because phylogenetic constraints on host choice are probably quite weak. By contrast, the intraspecific hypothesis has difficulty accounting for the parasites' sympatric reproductive isolation. Facultative polygyny, in which queens may found colonies alone or by adoption into an existing multi-queen colony, should promote the evolution of small intraspecific inquilines. This is because small colony-founding queens should preferentially seek adoption, which provides the opportunity to produce a sexual-only brood. We suggest that microgynes, i.e. miniature queens found in some polygynous ants, represent such parasites. We review the evidence that inquiline species have evolved intraspecifically from microgynes in Myrmica ants. The coexistence within a species of a monogynous (singly-queened) and a polygynous form is probably a phenomenon usually unconnected with inquiline evolution. The reproductive isolation of intraspecific inquilines plausibly arises from divergent breeding behaviour associated with the parasites' small size. Such divergence could involve either a temporal separation in mating episodes, with small parasites maturing early, or a spatial separation, with small males being sexually-selected to mate near the nest with small queens seeking adoption, instead of in mating aggregations. We conclude that inquiline species strictly following Emery's rule could have evolved by the intraspecific route. If so, such species provide evidence for West-Eberhard's “alternative adaptation” hypothesis that between-species diversity frequently stems from diversity within species. They also represent likely cases of sympatric speciation. We suggest work on the parasites' phytogeny, genetics, behaviour and mating biology to test these conclusions further.     

Assessment of lead speciation by organic ligands using speciation models

Abstract

The biogeochemical behaviour of lead (Pb) in ecosystems greatly depends on its chemical species. Organic ligands strongly influence Pb species and mobility in soil solution. In the present study, two metal speciation models, Windermere Humic Aqueous Model (WHAM) and Visual Minteq are used to compute Pb speciation in nutrient solution in the presence and absence of organic ligands. The three organic ligands used include ethylenediamine tetraacetic acid (EDTA), citric acid (CA) and fulvic acid (FA). The results show that in the absence of organic ligands, Pb²⁺ is the dominant form under acidic conditions and Pb–OH under alkaline conditions. The presence of organic ligands strongly influences Pb speciation. EDTA is more effective than are CA and FA concerning its influence on Pb speciation due to high Pb binding capacity. The results also indicate that Pb binding capacity of organic ligands varies with solution pH.     

Population genetics of Manihot esculenta ssp. flabellifolia gives insight into past distribution of xeric vegetation in a postulated forest refugium area in northern Amazonia

The Guianas have often been proposed as a forest refugium; however, this view has received little testing. Studies of population genetics of forest taxa suggest that the central part of French Guiana remained forested, while the southern part (currently forested) may have harboured more open vegetation. Insights into the population structure of species restricted to non-forested habitats can help test this hypothesis. Using six microsatellite loci, we investigated the population genetics of French Guianan accessions of Manihot esculenta ssp. flabellifolia , a taxon restricted to coastal savannas and to rocky outcrops in the densely forested inland. Coastal populations were highly differentiated from one another, and our data suggest a recent colonization of these savannas by M. esculenta ssp. flabellifolia in a west-to-east process. Coastal populations were strongly differentiated from inselberg populations, consistent with an ancient separation of these two groups, with no or low subsequent gene flow. This supports the hypothesis that the central part of the region may have remained forested since the Last Glacial Maximum, impeding the establishment of Manihot . Contrary to coastal populations, inselberg Manihot populations were strikingly homogeneous at a broad spatial scale. This suggests they were connected until recently, either by a large continuous savanna area or by smaller, temporary disturbed areas shifting in space.     

Metapopulations: an arena for local speciation

Speciation in many groups of plants and animals is thought to occur in single, isolated populations. Recent investigations of metapopulation longevity and genetic structure indicate that metapopulations have demographic and genetic properties which may be more conducive for local speciation than properties of single populations. It is argued that metapopulations may persist longer than single populations, that the former may retain genetic variation more readily, and that the former offers more opportunity for genomic reorganization.     

Geographical range size heritability: what do neutral models with different modes of speciation predict?

Aim Phylogenetic conservatism or heritability of the geographical range sizes of species (i.e. the tendency for closely related species to share similar range sizes) has been predicted to occur because of the strong phylogenetic conservatism of niche traits. However, the extent of such heritability in range size is disputed and the role of biology in shaping this attribute remains unclear. Here, we investigate the level of heritability of geographical range sizes that is generated from neutral models assuming no biological differences between species. Methods We used three different neutral models, which differ in their speciation mode, to simulate the life‐history of 250,000 individuals in a square lattice of 50 × 50 cells. These individuals can speciate, reproduce, migrate and die in the metacommunity according to stochastic events. We ran each model for 3000 steps and recorded the range size of each species at each step. The heritability of geographical range size was assessed using an asymmetry coefficient between range sizes of sister species and using the coefficient of correlation between the range sizes of ancestors and their descendants. Results Our results demonstrated the ability of neutral models to mimic some important observed patterns in the heritability of geographical range size. Consistently, sister species exhibited higher asymmetry in range sizes than expected by chance, and correlations between the range sizes of ancestor–descendant species pairs, although often weak, were almost invariably positive. Main conclusions Our findings suggest that, even without any biological trait differences, statistically significant heritability in the geographical range sizes of species can be found. This heritability is weaker than that observed in some empirical studies, but suggests that even here a substantial component of heritability may not necessarily be associated with niche conservatism. We also conclude that both present‐day and fossil data sets may provide similar information on the heritability of the geographical range sizes of species, while the omission of rare species will tend to overestimate this heritability.     

Systematics, biogeography and host plant associations of the Pseudomyrmex viduus group (Hymenoptera: Formicidae), Triplaris- and Tac/Mga/t-inhabiting ants

The Pseudomyrmex viduus group is a Neotropical clade of arboreal ants containing 13 species, of which three are newly described here: P. insuavis, P. ultrix, and P. vitabilis. Most species inhabit the domatia of specialized ant-plants. The ants keep brood and scale insects (Coccoidea) in the plant cavities, and defend their nest sites much more aggressively than do generalist species of Pseudomyrmex nesting in dead twigs. Five species are obligate associates of trees in the genus Triplaris (Polygonaceae) and five taxa are restricted to Tachigali (Fabaceae: Caesalpinioideae). One species, P. viduus (F. Smith), is much less host-specific, inhabiting Cordia, Coussapoa, Ocotea, Pseudobombax, Pterocarpus, Sapium, Triplaris and other myrmecophytes. Nothing is known about the biology of P. vitabilis, a close relative of/?viduus. The last member of the species group, P. kuenckeli (Emery), is a non-specialist but aggressive species that nests in dead branches of various plants. A cladistic analysis indicates that this aggressive behaviour evolved before obligate associations with specialized ant-plants, and that the Triplaris and Tachigali inhabitants each form their own clade. P. viduus, which is nested within the Triplaris-associated clade, suggests a possible model for host plant evolution in these ants wherein shifts from one ant-plant to another involve an intermediate phase of expanded host plant use. At least nine other Pseudomyrmex species, from two different species groups (not closely related to the P. viduus group), have evolved specialized associations with Triplaris or Tachigali including five new species: P. crudelis, P. deminutus, P. eculeus. P. ferox and P. hospitalis. Although the P. viduus group is centred in the Amazon basin, the geographic ranges of most species do not coincide with the Pleistocene forest refugia proposed by Haffer and others. A consideration of the phylogenetic relationships, distribution patterns, and host plant specificity of the ants indicates that much of the diversification of the P. viduus group occurred before the Pleistocene, and that the interactions with Triplaris and Tachigali plants are also of Tertiary origin.     

Case studies and mathematical models of ecological speciation. 2. Palms on an oceanic island

A recent study of a pair of sympatric species of palms on the Lord Howe Island is viewed as providing probably one of the most convincing examples of sympatric speciation to date. Here we describe and study a stochastic, individual-based, explicit genetic model tailored for this palms system. Overall, our results show that relatively rapid (<50,000 generations) colonization of a new ecological niche, and sympatric or parapatric speciation via local adaptation and divergence in flowering periods are theoretically plausible if (i) the number of loci controlling the ecological and flowering period traits is small; (ii) the strength of selection for local adaptation is intermediate; and (iii) an acceleration of flowering by a direct environmental effect associated with the new ecological niche is present. We discuss patterns and time-scales of ecological speciation identified by our model, and we highlight important parameters and features that need to be studied empirically in order to provide information that can be used to improve the biological realism and power of mathematical models of ecological speciation.     

The impact of global climate change on tropical forest biodiversity in Amazonia

Aim To model long‐term trends in plant species distributions in response to predicted changes in global climate. Location Amazonia. Methods The impacts of expected global climate change on the potential and realized distributions of a representative sample of 69 individual Angiosperm species in Amazonia were simulated from 1990 to 2095. The climate trend followed the HADCM2GSa1 scenario, which assumes an annual 1% increase of atmospheric CO₂ content with effects mitigated by sulphate forcing. Potential distributions of species in one‐degree grid cells were modelled using a suitability index and rectilinear envelope based on bioclimate variables. Realized distributions were additionally limited by spatial contiguity with, and proximity to, known record sites. A size‐structured population model was simulated for each cell in the realized distributions to allow for lags in response to climate change, but dispersal was not included. Results In the resulting simulations, 43% of all species became non‐viable by 2095 because their potential distributions had changed drastically, but there was little change in the realized distributions of most species, owing to delays in population responses. Widely distributed species with high tolerance to environmental variation exhibited the least response to climate change, and species with narrow ranges and short generation times the greatest. Climate changed most in north‐east Amazonia while the best remaining conditions for lowland moist forest species were in western Amazonia. Main conclusions To maintain the greatest resilience of Amazonian biodiversity to climate change as modelled by HADCM2GSa1, highest priority should be given to strengthening and extending protected areas in western Amazonia that encompass lowland and montane forests.     

The geographical pattern of speciation and floral diversification in the neotropics: the tribe sinningieae (gesneriaceae) as a case study

The geographical pattern of speciation and the relationship between floral variation and species ranges were investigated in the tribe Sinningieae (Gesneriaceae), which is found mainly in the Atlantic forests of Brazil. Geographical distribution data recorded on a grid system of 0.5 x 0.5 degree intervals and a near-complete species-level phylogenetic tree of Sinningieae inferred from a simultaneous analysis of seven DNA regions were used to address the role of geographical isolation in speciation. Geographical range overlaps between sister lineages were measured across all nodes in the phylogenetic tree and analyzed in relation to relative ages estimated from branch lengths. Although there are several cases of species sympatry in Sinningieae, patterns of sympatry between sister taxa support the predominance of allopatric speciation. The pattern of sympatry between sister taxa is consistent with range shifts following allopatric speciation, except in one clade, in which the overlapping distribution of recent sister species indicates speciation within a restricted geographical area and involving changes in pollinators and habitats. The relationship between floral divergence and regional sympatry was also examined by analyzing floral contrasts, phenological overlap, and the degree of sympatry between sister clades. Morphological contrast between flowers is not increased in sympatry and phenological divergence is more apparent between allopatric clades than between sympatric clades. Therefore, our results failed to indicate a tendency for sympatric taxa to minimize morphological and phenological overlap (geographic exclusion and/or character displacement hypotheses). Instead, they point toward adaptation in phenology to local conditions and buildup of sympatries at random with respect to flower morphology. Additional studies at a lower geographical scale are needed to identify truely coexisting species and the components of their reproductive isolation.     

Inverting the null-hypothesis of speciation: a marine snail perspective

Speciation is currently an intensely debated topic, much more so than 20–30 years ago when most biologists held the view that new species (at least of animals) were formed through the split of evolutionary lineages by the appearance of physical barriers to gene flow. Recent advances have, however, lent both theoretical and empirical support to speciation in the presence of gene flow. Nevertheless, the allopatric hypothesis of speciation is still the default model. The consequence of this is that to support sympatric and parapatric modes of speciation all allopatric alternatives must be rejected, while an allopatric explanation is usually accepted without rejecting possible non-allopatric alternatives. However, classical cases of allopatric speciation can be challenged by alternative non-allopatric explanations, and this begs for a more respectful view of how to deal with all models of speciation. An appealing approach is studying parallel evolution of reproductive barriers, which allows for comparative approaches to distinguish between allopatric and non-allopatric events, and explicit tests of a suitable null-hypothesis. Parallel evolution of reproductive isolation in a strongly polymorphic marine snail species serves as an illustrative example of such an approach. In conclusion, a more balanced debate on allopatric and non-allopatric speciation is needed and an urgent issue is to treat both allopatric and nonallopatric hypotheses critically, rather than using allopatry as the default model of speciation.     

Multi-variate models are essential for understanding vertebrate diversification in deep time

Statistical models are helping palaeontologists to elucidate the history of biodiversity. Sampling standardization has been extensively applied to remedy the effects of uneven sampling in large datasets of fossil invertebrates. However, many vertebrate datasets are smaller, and the issue of uneven sampling has commonly been ignored, or approached using pairwise comparisons with a numerical proxy for sampling effort. Although most authors find a strong correlation between palaeodiversity and sampling proxies, weak correlation is recorded in some datasets. This has led several authors to conclude that uneven sampling does not influence our view of vertebrate macroevolution. We demonstrate that multi-variate regression models incorporating a model of underlying biological diversification, as well as a sampling proxy, fit observed sauropodomorph dinosaur palaeodiversity best. This bivariate model is a better fit than separate univariate models, and illustrates that observed palaeodiversity is a composite pattern, representing a biological signal overprinted by variation in sampling effort. Multi-variate models and other approaches that consider sampling as an essential component of palaeodiversity are central to gaining a more complete understanding of deep time vertebrate diversification.     

Marine vertebrate zoonoses: an overview of the DAO special issue

The role of marine birds, mammals, turtles and fish as vectors of infectious agents of potential risk to humans can be examined from a variety of perspectives. The studies in this DAO Special include a broad survey of multiple agents and species, a sequencing study of Giardia intestinalis haplotypes known to be pathogenic to humans, an assessment of risks to humans working with marine mammals, a source tracking study using E. coli ribotypes, studies of regional Salmonella and Brucella epizootiology, a serology survey and a case report of a herpes simplex infection in a dolphin. Additionally, a recently published study (Venn-Watson et al. 2008; Dis Aquat Org 79:87-93) classifying pure cultures of bacteria from a captive dolphin colony also pertains to this theme. These studies raise the following questions: whether the presence of zoonotic agents in marine vertebrates represents a risk to other marine vertebrates, humans, or both; what are the routes by which these marine vertebrate zoonotic infections are acquired and circulated in the marine ecosystem; to what degree are such agents subclinical versus causes of overt disease in marine vertebrates; what are the subsets of the human population most likely to be affected by such infections; and which human health preventive measures would seem reasonable?     

Evidence of extreme habitat stability in a Southeast Asian biodiversity hotspot based on the evolutionary analysis of neotenic net-winged beetles

The diversification of neotenic beetle lineages has not been studied, despite the potential for defining biodiversity hotspots and elucidating the history of regional faunas. Additionally, neotenics may provide insight into the process of speciation in small populations with extremely low dispersal ability and a limited range. Here, we used two rDNA and three mtDNA markers to investigate the phylogeny of Scarelus, a neotenic lineage endemic to Southeast Asian rainforests. Most genetic differentiation was associated with Palaeogene geographical divisions, which remain distinct despite temporary connections. Dispersal events were rare, with only two inferred for Scarelus: from Borneo to the Philippines 28.3 million years ago (Ma) and from Sumatra to Java 13.9 Ma. We suggest that speciation resulted from allopatric range fragmentation, and Scarelus diversified readily when conditions were favourable; in this case, at different times in the eastern (19.3-39.1 Ma) and western (3.5-13.9 Ma) parts of Sundaland. The observed strong phenotypic similarity was preserved under speciation through complete allopatry. Neotenic Lycidae have survived for a long time in very stable habitats, and extremely low dispersal activity has not limited their persistence; however, the long-term diversification rate of neotenics is low and diversification is nonexistent under stable conditions. The modern ranges of neotenic lineages are indicative of ancient rainforest refugia and may be used in biodiversity conservation management. Most neotenics are at risk of extinction because of their small ranges and a low dispersal potential.