Adding Biotic Interactions into Paleodistribution Models: A Host-Cleptoparasite Complex of Neotropical Orchid Bees

Orchid bees compose an exclusive Neotropical pollinators group, with bright body coloration. Several of those species build their own nests, while others are reported as nest cleptoparasites. Here, the objective was to evaluate whether the inclusion of a strong biotic interaction, such as the presence of a host species, improved the ability of species distribution models (SDMs) to predict the geographic range of the cleptoparasite species. The target species were Aglae caerulea and its host species Eulaema nigrita. Additionally, since A. caerulea is more frequently found in the Amazon rather than the Cerrado areas, a secondary objective was to evaluate whether this species is increasing or decreasing its distribution given South American past and current climatic conditions. SDMs methods (Maxent and Bioclim), in addition with current and past South American climatic conditions, as well as the occurrences for A. caerulea and E. nigrita were used to generate the distribution models. The distribution of A. caerulea was generated with and without the inclusion of the distribution of E. nigrita as a predictor variable. The results indicate A. caerulea was barely affected by past climatic conditions and the populations from the Cerrado savanna could be at least 21,000 years old (the last glacial maximum), as well as the Amazonian ones. On the other hand, in this study, the inclusion of the host-cleptoparasite interaction complex did not statistically improve the quality of the produced models, which means that the geographic range of this cleptoparasite species is mainly constrained by climate and not by the presence of the host species. Nonetheless, this could also be caused by unknown complexes of other Euglossini hosts with A. caerulea, which still are still needed to be described by science.     

Conservation Value and Permeability of Neotropical Oil Palm Landscapes for Orchid Bees

The proliferation of oil palm plantations has led to dramatic changes in tropical landscapes across the globe. However, relatively little is known about the effects of oil palm expansion on biodiversity, especially in key ecosystem-service providing organisms like pollinators. Rapid land use change is exacerbated by limited knowledge of the mechanisms causing biodiversity decline in the tropics, particularly those involving landscape features. We examined these mechanisms by undertaking a survey of orchid bees, a well-known group of Neotropical pollinators, across forest and oil palm plantations in Costa Rica. We used chemical baits to survey the community in four regions: continuous forest sites, oil palm sites immediately adjacent to forest, oil palm sites 2km from forest, and oil palm sites greater than 5km from forest. We found that although orchid bees are present in all environments, orchid bee communities diverged across the gradient, and community richness, abundance, and similarity to forest declined as distance from forest increased. In addition, mean phylogenetic distance of the orchid bee community declined and was more clustered in oil palm. Community traits also differed with individuals in oil palm having shorter average tongue length and larger average geographic range size than those in the forest. Our results indicate two key features about Neotropical landscapes that contain oil palm: 1) oil palm is selectively permeable to orchid bees and 2) orchid bee communities in oil palm have distinct phylogenetic and trait structure compared to communities in forest. These results suggest that conservation and management efforts in oil palm-cultivating regions should focus on landscape features.     

The Ecological Basis for Biogeographic Classification: an Example in Orchid Bees (Apidae: Euglossini)

Biogeography has been difficult to apply as a methodological approach because organismic biology is incomplete at levels where the process of formulating comparisons and analogies is complex. The study of insect biogeography became necessary because insects possess numerous evolutionary traits and play an important role as pollinators. Among insects, the euglossine bees, or orchid bees, attract interest because the study of their biology allows us to explain important steps in the evolution of social behavior and many other adaptive tradeoffs. We analyzed the distribution of morphological characteristics in Colombian orchid bees from an ecological perspective. The aim of this study was to observe the distribution of these attributes on a regional basis. Data corresponding to Colombian euglossine species were ordered with a correspondence analysis and with subsequent hierarchical clustering. Later, and based on community proprieties, we compared the resulting hierarchical model with the collection localities to seek to identify a biogeographic classification pattern. From this analysis, we derived a model that classifies the territory of Colombia into 11 biogeographic units or natural clusters. Ecological assumptions in concordance with the derived classification levels suggest that species characteristics associated with flight performance, nectar uptake, and social behavior are the factors that served to produce the current geographical structure.     

Integrating Life Stages into Ecological Niche Models: A Case Study on Tiger Beetles

Detailed understanding of a species’ natural history and environmental needs across spatial scales is a primary requisite for effective conservation planning, particularly for species with complex life cycles in which different life stages occupy different niches and respond to the environment at different scales. However, niche models applied to conservation often neglect early life stages and are mostly performed at broad spatial scales. Using the endangered heath tiger beetle (Cicindela sylvatica) as a model species, we relate presence/absence and abundance data of locally dispersing adults and sedentary larvae to abiotic and biotic variables measured in a multiscale approach within the geographic extent relevant to active conservation management. At the scale of hundreds of meters, fine-grained abiotic conditions (i.e., vegetation structure) are fundamental determinants of the occurrence of both life stages, whereas the effect of biotic factors is mostly contained in the abiotic signature. The combination of dense heath vegetation and bare ground areas is thus the first requirement for the species’ preservation, provided that accessibility to the suitable habitat is ensured. At a smaller scale (centimetres), the influence of abiotic factors on larval occurrence becomes negligible, suggesting the existence of important additional variables acting within larval proximity. Sustained significant correlations between neighbouring larvae in the models provide an indication of the potential impact of neighbourhood crowding on the larval niche within a few centimetres. Since the species spends the majority of its life cycle in the larval stage, it is essential to consider the hierarchical abiotic and biotic processes affecting the larvae when designing practical conservation guidelines for the species. This underlines the necessity for a more critical evaluation of the consequences of disregarding niche variation between life stages when estimating niches and addressing effective conservation measures for species with complex life cycles.     

Honey Pollen: Using Melissopalynology to Understand Foraging Preferences of Bees in Tropical South India

The aim of the study was to use melissopalynology to delineate the foraging preferences of bees in tropical environs. This was done by comparing pollen spectra obtained from the same hives every three months for three years at four sampling locations (in two sites) within a confined landscape mosaic. If melissopalynology is highly replicable, the spatial variation of the pollen spectrum from the honey samples would be much more than the temporal (inter-annual) variations. In other words, given the three factors, Month, Year and Location, honey pollen from different Locations, in a given Year and Month, would be much less similar than samples from different Years, in a given Location and Month. We then determined how the factors, Month, Year and Location, influenced the pollen influx of honey. The pollen analyses of the 42 honey samples collected during the three years yielded 80 pollen taxa/types: 72 dicotyledonous and 8 monocotyledonous, encompassing 41 botanical families spread into seven life forms namely, trees, shrubs, epiphytes, herbs, climbers, grasses, and sedges. Our results showed that pollen spectra were equally comparable between Locations and between Months and Years; the importance of this result is that it helped to demonstrate the complexity of ecological/environmental phenomena involved in the process of foraging by bees in a heterogeneous and complex landscape.     

Cuticular Hydrocarbons of Orchid Bees Males: Interspecific and Chemotaxonomy Variation

Recent studies have investigated the composition of compounds that cover the cuticle in social insects, but few studies have focused on solitary bees. Cuticular hydrocarbons may provide a tool for chemotaxonomy, and perhaps they can be used as a complement to morphology and genetic characters in phylogenetic studies. Orchid bees (Tribe Euglossini) are a highly diverse group of Neotropical bees with more than 200 species. Here, the cuticular hydrocarbons of 17 species were identified and statistical analysis revealed 108 compounds, which allowed for the taxonomic classification according to the genera. The most significant compounds discriminating the four genera were (Z)-9-pentacosene, (Z,Z)-pentatriacontene-3, (Z)-9-tricosene, and (Z)-9-heptacosene. The analyses demonstrated the potential use of CHCs to identify different species.     

Using Networks To Understand Medical Data: The Case of Class III Malocclusions

A system of elements that interact or regulate each other can be represented by a mathematical object called a network. While network analysis has been successfully applied to high-throughput biological systems, less has been done regarding their application in more applied fields of medicine; here we show an application based on standard medical diagnostic data. We apply network analysis to Class III malocclusion, one of the most difficult to understand and treat orofacial anomaly. We hypothesize that different interactions of the skeletal components can contribute to pathological disequilibrium; in order to test this hypothesis, we apply network analysis to 532 Class III young female patients. The topology of the Class III malocclusion obtained by network analysis shows a strong co-occurrence of abnormal skeletal features. The pattern of these occurrences influences the vertical and horizontal balance of disharmony in skeletal form and position. Patients with more unbalanced orthodontic phenotypes show preponderance of the pathological skeletal nodes and minor relevance of adaptive dentoalveolar equilibrating nodes. Furthermore, by applying Power Graphs analysis we identify some functional modules among orthodontic nodes. These modules correspond to groups of tightly inter-related features and presumably constitute the key regulators of plasticity and the sites of unbalance of the growing dentofacial Class III system. The data of the present study show that, in their most basic abstraction level, the orofacial characteristics can be represented as graphs using nodes to represent orthodontic characteristics, and edges to represent their various types of interactions. The applications of this mathematical model could improve the interpretation of the quantitative, patient-specific information, and help to better targeting therapy. Last but not least, the methodology we have applied in analyzing orthodontic features can be applied easily to other fields of the medical science.     

Niche Divergence in a Brown Lemur (<Emphasis Type="Italic">Eulemur</Emphasis> spp.) Hybrid Zone: Using Ecological Niche Models to Test Models of Stability

Endogenous selection is often implicated in the maintenance of stability of natural hybrid zones. Environmental conditions often vary across these zones, suggesting that local adaptation to ecological conditions could also play a role in this process. We used niche modeling to investigate these alternatives in a hybrid zone between two species of brown lemur (Eulemur rufifrons and E. cinereiceps) in southeastern Madagascar. We produced ecological niche models (ENMs) for parental and hybrid populations and compared values of niche overlap to null expectations using identity and background tests. All three taxonomic groups had nonequivalent ENMs with limited spatial overlap, supporting a role for niche divergence and local adaptation in the maintenance of this zone. However, values of niche overlap between ENMs were not greater than null expectations controlling for background environmental differences. These results could suggest that taxa in this hybrid zone inhabit portions of their environments that are more similar to their backgrounds, i.e., niche conservatism. Nevertheless, we did find evidence of niche divergence when using background tests that examined environmental variables separately. Although we could not rule out models indicating selection against hybrids, most lines of evidence were consistent with predictions for the bounded superiority model of hybrid zone stability. This study thus provides support that exogenous, environmental selection may be responsible for maintaining the hybrid zone, and may be implicated in the evolutionary divergence of these taxa.     

The Geography of Speciation: Case Studies from Birds

Our understanding of the origin of species, or speciation, is sometimes viewed as incomplete, a ??mystery of mysteries.?? We in fact know a lot about speciation, especially when we consider its two basic components, the geography of speciation and the biology of speciation (changes in phenotype and genotype that occur during the process). Our understanding of the geography of speciation is quite clear. The process involves the separation of a once-continuous range into two or more geographically isolated, or allopatric parts, which over time accrue genetic changes that result in new daughter species. Current distributions show that recently evolved species are currently allopatric, supporting the role of allopatry in speciation. However, many species originated in the early- to mid-Pleistocene, meaning that they persisted through the environmental perturbations of multiple glacial cycles. It has been assumed that species maintained allopatric distributions during these episodes of glacial advance and retreat. I used Grinnelian niche models to estimate species distributions at the Last Glacial Maximum and the Last Interglacial. For a pair of recently isolated warbler species, and a pair of relatively old sister species of gnatcatchers, allopatry was observed at all time periods. Thus, there is no mystery about the geography of speciation: at least in birds, allopatry predominates. The sentiment that speciation is mysterious comes, I argue, from the biological species concept, which requires populations to be reproductively isolated before recognizing them as species. Reproductive isolation is a complicated process that rarely occurs the same way twice, and I argue that this lack of generality has been misinterpreted as a mystery.     

Ecological Patterns and Biological Invasions: Using Regional Species Inventories in Macroecology

Macroecology depends heavily on a comparative methodology in order to identify large-scale patterns and to test alternative hypotheses that might generate such patterns. With the advent and accessibility of large electronic databases of species and their life history and ecological attributes, ecologists have begun seeking generalities, and examining large-scale ecological hypotheses involving core themes of range, abundance and diversity. For example, combinations of ecological, life history and phylogenetic data have been analysed using large species sets to test hypotheses in invasion biology. Analysis of regional species inventories can contribute cogently to our understanding of invasions. Here we examine several ways in which database analysis is effective. We review 19 studies of comparative invasions biology, each using >100 species of plants in their analyses, and show that invader success is linked to seven correlates: short life cycle, abiotic (mostly wind) dispersal, large native range size, non-random taxonomic patterns (emphasizing certain families or orders), presence of clonal organs, occupying disturbed habitats, and earlier time of introduction. These phylogenetically influenced, comparative analyses using regional species inventories are only just beginning and have much potential.     

Comparative Development of Anurans: Using Phylogeny to Understand Ontogeny

Hypotheses of relationships are critical to describing and understandingpatterns of evolution within groups of organisms. But rarelyhas a comparative, historical approach been employed to studydevelopmental change, particularly among anurans. A recent resurgenceof interest in collecting basic ontogenetic information providesus with the opportunity to compare ontogenetic trajectoriesin a phylogenetic framework. Larval skeletons and osteologicaldevelopment were examined for 22 taxa and compared to two hypothesesof relationships—that of Cannatella, and one proposedherein based on 41 morphological characters from larvae and62 from adults. Larval characters were mapped on the alternatecladograms using the ACCTRAN optimization criterion. Severallarval features are highly conserved among some anurans, suggestingthat there is some level of canalization of morphology earlyin ontogeny. In contrast, a number of morphologies vary amonggroups, supporting the fact that there have been major evolutionarymodifications to anuran larval morphologies early in ontogenyand in the early evolutionary history of anurans.     

Climate Change and Risk of Leishmaniasis in North America: Predictions from Ecological Niche Models of Vector and Reservoir Species

Background

Climate change is increasingly being implicated in species'' range shifts throughout the world, including those of important vector and reservoir species for infectious diseases. In North America (México, United States, and Canada), leishmaniasis is a vector-borne disease that is autochthonous in México and Texas and has begun to expand its range northward. Further expansion to the north may be facilitated by climate change as more habitat becomes suitable for vector and reservoir species for leishmaniasis.

Methods and Findings

The analysis began with the construction of ecological niche models using a maximum entropy algorithm for the distribution of two sand fly vector species (Lutzomyia anthophora and L. diabolica), three confirmed rodent reservoir species (Neotoma albigula, N. floridana, and N. micropus), and one potential rodent reservoir species (N. mexicana) for leishmaniasis in northern México and the United States. As input, these models used species'' occurrence records with topographic and climatic parameters as explanatory variables. Models were tested for their ability to predict correctly both a specified fraction of occurrence points set aside for this purpose and occurrence points from an independently derived data set. These models were refined to obtain predicted species'' geographical distributions under increasingly strict assumptions about the ability of a species to disperse to suitable habitat and to persist in it, as modulated by its ecological suitability. Models successful at predictions were fitted to the extreme A2 and relatively conservative B2 projected climate scenarios for 2020, 2050, and 2080 using publicly available interpolated climate data from the Third Intergovernmental Panel on Climate Change Assessment Report. Further analyses included estimation of the projected human population that could potentially be exposed to leishmaniasis in 2020, 2050, and 2080 under the A2 and B2 scenarios. All confirmed vector and reservoir species will see an expansion of their potential range towards the north. Thus, leishmaniasis has the potential to expand northwards from México and the southern United States. In the eastern United States its spread is predicted to be limited by the range of L. diabolica; further west, L. anthophora may play the same role. In the east it may even reach the southern boundary of Canada. The risk of spread is greater for the A2 scenario than for the B2 scenario. Even in the latter case, with restrictive (contiguous) models for dispersal of vector and reservoir species, and limiting vector and reservoir species occupancy to only the top 10% of their potential suitable habitat, the expected number of human individuals exposed to leishmaniasis by 2080 will at least double its present value.

Conclusions

These models predict that climate change will exacerbate the ecological risk of human exposure to leishmaniasis in areas outside its present range in the United States and, possibly, in parts of southern Canada. This prediction suggests the adoption of measures such as surveillance for leishmaniasis north of Texas as disease cases spread northwards. Potential vector and reservoir control strategies—besides direct intervention in disease cases—should also be further investigated.     

Niche Divergence versus Neutral Processes: Combined Environmental and Genetic Analyses Identify Contrasting Patterns of Differentiation in Recently Diverged Pine Species

Background and Aims

Solving relationships of recently diverged taxa, poses a challenge due to shared polymorphism and weak reproductive barriers. Multiple lines of evidence are needed to identify independently evolving lineages. This is especially true of long-lived species with large effective population sizes, and slow rates of lineage sorting. North American pines are an interesting group to test this multiple approach. Our aim is to combine cytoplasmic genetic markers with environmental information to clarify species boundaries and relationships of the species complex of Pinus flexilis, Pinus ayacahuite, and Pinus strobiformis.

Methods

Mitochondrial and chloroplast sequences were combined with previously obtained microsatellite data and contrasted with environmental information to reconstruct phylogenetic relationships of the species complex. Ecological niche models were compared to test if ecological divergence is significant among species.

Key Results and Conclusion

Separately, both genetic and ecological evidence support a clear differentiation of all three species but with different topology, but also reveal an ancestral contact zone between P. strobiformis and P. ayacahuite. The marked ecological differentiation of P. flexilis suggests that ecological speciation has occurred in this lineage, but this is not reflected in neutral markers. The inclusion of environmental traits in phylogenetic reconstruction improved the resolution of internal branches. We suggest that combining environmental and genetic information would be useful for species delimitation and phylogenetic studies in other recently diverged species complexes.     

Using Uncertainty and Sensitivity Analyses in Socioecological Agent-Based Models to Improve Their Analytical Performance and Policy Relevance

Agent-based models (ABMs) have been widely used to study socioecological systems. They are useful for studying such systems because of their ability to incorporate micro-level behaviors among interacting agents, and to understand emergent phenomena due to these interactions. However, ABMs are inherently stochastic and require proper handling of uncertainty. We propose a simulation framework based on quantitative uncertainty and sensitivity analyses to build parsimonious ABMs that serve two purposes: exploration of the outcome space to simulate low-probability but high-consequence events that may have significant policy implications, and explanation of model behavior to describe the system with higher accuracy. The proposed framework is applied to the problem of modeling farmland conservation resulting in land use change. We employ output variance decomposition based on quasi-random sampling of the input space and perform three computational experiments. First, we perform uncertainty analysis to improve model legitimacy, where the distribution of results informs us about the expected value that can be validated against independent data, and provides information on the variance around this mean as well as the extreme results. In our last two computational experiments, we employ sensitivity analysis to produce two simpler versions of the ABM. First, input space is reduced only to inputs that produced the variance of the initial ABM, resulting in a model with output distribution similar to the initial model. Second, we refine the value of the most influential input, producing a model that maintains the mean of the output of initial ABM but with less spread. These simplifications can be used to 1) efficiently explore model outcomes, including outliers that may be important considerations in the design of robust policies, and 2) conduct explanatory analysis that exposes the smallest number of inputs influencing the steady state of the modeled system.     

Diversity and Distributional Patterns of Neotropical Freshwater Copepods (Calanoida: Diaptomidae)

The distributional patterns and diversity of the diaptomid calanoid copepods were analysed to assess the faunistic affinity of North and South America with respect to Mexico and Central America. In the Neotropical region, the most speciose genera of Diaptomidae are Leptodiaptomus and Mastigodiaptomus. The former genus is a Nearctic form, and Mastigodiaptomus is Neotropical. Based on the current distribution of their diversity, it is probable that these genera radiated into Mexico and Central America from North America and the insular Caribbean, respectively. Arctodiaptomus dorsalis is a primarily Palaearctic taxon, it is widely distributed between North and Central America. This species probably radiated in the Americas as a Tethyan derivate. Prionodiaptomus is the only member of the highly diverse South American diaptomid fauna that has expanded beyond the subcontinent. Despite the high diversity present in South America, its influence in Mexico and Central America appears to be weak; this is probably a consequence of the geologically recent union of the two main subcontinental landmasses. Mexico shares 33% of its species with NA, and no species are shared between NA and SA. For the Diaptomidae, the Nearctic influence is strongest in Mexico. (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Mapping the Potential Risk of Mycetoma Infection in Sudan and South Sudan Using Ecological Niche Modeling

In 2013, the World Health Organization (WHO) recognized mycetoma as one of the neglected tropical conditions due to the efforts of the mycetoma consortium. This same consortium formulated knowledge gaps that require further research. One of these gaps was that very few data are available on the epidemiology and transmission cycle of the causative agents. Previous work suggested a soil-borne or Acacia thorn-prick-mediated origin of mycetoma infections, but no studies have investigated effects of soil type and Acacia geographic distribution on mycetoma case distributions. Here, we map risk of mycetoma infection across Sudan and South Sudan using ecological niche modeling (ENM). For this study, records of mycetoma cases were obtained from the scientific literature and GIDEON; Acacia records were obtained from the Global Biodiversity Information Facility. We developed ENMs based on digital GIS data layers summarizing soil characteristics, land-surface temperature, and greenness indices to provide a rich picture of environmental variation across Sudan and South Sudan. ENMs were calibrated in known endemic districts and transferred countrywide; model results suggested that risk is greatest in an east-west belt across central Sudan. Visualizing ENMs in environmental dimensions, mycetoma occurs under diverse environmental conditions. We compared niches of mycetoma and Acacia trees, and could not reject the null hypothesis of niche similarity. This study revealed contributions of different environmental factors to mycetoma infection risk, identified suitable environments and regions for transmission, signaled a potential mycetoma-Acacia association, and provided steps towards a robust risk map for the disease.     

Using Plant Functional Traits and Phylogenies to Understand Patterns of Plant Community Assembly in a Seasonal Tropical Forest in Lao PDR

Plant functional traits reflect different evolutionary responses to environmental variation, and among extant species determine the outcomes of interactions between plants and their environment, including other plant species. Thus, combining phylogenetic and trait-based information can be a powerful approach for understanding community assembly processes across a range of spatial scales. We used this approach to investigate tree community composition at Phou Khao Khouay National Park (18°14’-18°32’N; 102°38’- 102°59’E), Laos, where several distinct forest types occur in close proximity. The aim of our study was to examine patterns of plant community assembly across the strong environmental gradients evident at our site. We hypothesized that differences in tree community composition were being driven by an underlying gradient in soil conditions. Thus, we predicted that environmental filtering would predominate at the site and that the filtering would be strongest on sandier soil with low pH, as these are the conditions least favorable to plant growth. We surveyed eleven 0.25 ha (50x50 m) plots for all trees above 10 cm dbh (1221 individual trees, including 47 families, 70 genera and 123 species) and sampled soils in each plot. For each species in the community, we measured 11 commonly studied plant functional traits covering both the leaf and wood economic spectrum traits and we reconstructed a phylogenetic tree for 115 of the species in the community using rbcL and matK sequences downloaded from Genebank (other species were not available). Finally we compared the distribution of trait values and species at two scales (among plots and 10x10m subplots) to examine trait and phylogenetic community structures. Although there was strong evidence that an underlying soil gradient was determining patterns of species composition at the site, our results did not support the hypothesis that the environmental filtering dominated community assembly processes. For the measured plant functional traits there was no consistent pattern of trait dispersion across the site, either when traits were considered individually or when combined in a multivariate analysis. However, there was a significant correlation between the degree of phylogenetic dispersion and the first principle component axis (PCA1) for the soil parameters. Moreover, the more phylogenetically clustered plots were on sandier soils with lower pH. Hence, we suggest that the community assembly processes across our site may reflect the influence of more conserved traits that we did not measure. Nevertheless, our results are equivocal and other interpretations are possible. Our study illustrates some difficulties in combining trait and phylogenetic approaches that may result from the complexities of integrating spatial and evolutionary processes that vary at different scales.