River network structure shapes interannual feedbacks between adult sea lamprey migration and larval habitation

Adult sea lampreys (Petromyzon marinus) migrating upstream to spawn follow a pheromone released by instream larvae. The size (i.e. flow) of a tributary dilutes the concentration of this pheromone, such that the downstream propagation pattern of larval pheromone must be influenced by patterns in the relative sizes and numbers of confluent tributaries. We developed an individual-based model to explicitly test the resulting hypothesis that river network structure influences the migration decisions of adult lampreys following the larval pheromone, and in turn the distribution of larvae. First, we initialized the model using randomly generated river networks, and found a strong positive relationship between network diameter and larval aggregation. Larvae aggregated over time, and the degree and rate of this aggregation depended on network diameter. Second, we initialized the model using a river network based on the Muskegon River, Michigan, and compared model-generated larval distribution to available field survey data. We found a significant correlation between model-generated larval abundance and field-measured larval densities (r₂ = 0.54; p < 0.0001). We also found an inverse relationship between subwatershed area and the degree to which path-dependent effects influenced larval abundance in that subwatershed. Our results overall suggest that larval distribution across a watershed results from a system of context-dependent interannual feedbacks shaped by network structure and the past migratory and spawning behavior of adults.     

Shape and efficiency of wood ant foraging networks

We measured the shape of the foraging trail networks of 11 colonies of the wood ant Formica aquilonia (Formica rufa group). We characterized these networks in terms of their degree of branching and the angles between branches, as well as in terms of their efficiency. The measured networks were compared with idealized model networks built to optimize one of two components of efficiency, total length (i.e., total amount of trail) and route factor (i.e., average distance between nest and foraging site). The analysis shows that the networks built by the ants obtain a compromise between the two modes of efficiency. These results are largely independent of the size of the network or colony size. The ants’ efficiency is comparable to that of networks built by humans but achieved without the benefit of centralized control.     

Linking local to global properties in branching modular networks: gorgonian coral colonies

Branching growth is present both in plants and animals, either marine or terrestrial. Although cellular and other modular levels of organization in plants and animals have evolved through different molecular and physiological mechanisms, several aspects of their branching modular system and morphology are similar. We studied vessel organization and colony integration, in order to comprehend underlying relationships between different structural components in a gorgonian coral network. The theoretical formalism was validated in the gorgonian coral Eunicea mammosa (Plexauridae, Octocorallia) in Belize. As in vascular plants, these colonial animals create a complex network of connections among modular branches integrated in stem canals downstream toward the base. A new formalism is proposed for describing gorgonian branching. A global property of a colony is for instance the size of its base or its weight whereas a local property is the size of branch in a particular place of the colony. However, a global property is not the simple addition of local modular properties, as the case of stem canals in the colony base. Theoretically, the process of branching is tightly intertwined with the internal network organization. The colony network centralization is driven by a linear relationship between the total number of branches and the stem canals at the base of the colony. If stem canals play important roles in the transport of nutrients throughout the colony and the biomechanical support from the base up to the tips, we can assume that there is an underlying association between the number of stem canals at the base and the number of for example, terminal branches. These associations may provide new findings that extend our understanding of the functional organization of tree-like networks in octocorals and their vascular systems. The idea that the external components of a tree-like plant network are directly correlated and connected down to the main trunk seems to be analogous in an animal system.     

Genetic structure of local populations and divergence between growth forms in a clonal invertebrate,the Caribbean octocoral Briareum asbestinum

Although the genetic structure of many populations of marine organisms show little deviation from panmixia, in those marine species with limited larval dispersal, patterns of microgeographic genetic differentiation may be common. The octocoral Briareum asbestinum should show local population differentiation because colonies reproduce asexually by fragmentation, most matings occur between colonies in very close proximity, and the sexually produced larvae and sperm appear to disperse only short distances. Variability in secondary chemistry of individual B. asbestinum colonies from different populations in close proximity also suggests local population differentiation. We determined the genetic composition of local populations by surveying allozyme variation of three shallow and two deep populations within a 300 m² area at San Salvador Island, Bahamas and at a site 161 km away on Little San Salvador, Bahamas in July 1990. As B. asbestinum occurs as either an erect branching form or an encrusting mat often at the same sites, we sampled both morphs to examine the extent of genetic exchange between them. Five of 21 loci were polymorphic and most populations showed a deficit of heterozygotes. Allele frequencies differed significantly between morphs at each site where they occurred together. The mean genetic distance (D=0.065) between morphs is consistent with the interpretation that the two morphs are genetically isolated. Despite the close spatial proximity of the San Salvador populations, both the branching and encrusting morphs showed significant genetic heterogeneity among neighboring populations. Similarly, pooled allelic frequencies for samples collected from the islands of San Salvador and Little San Salvador differed significantly at 1 locus for the branching morph and at 3 out of 5 loci for the encrusting morph.     

Exploring Population Dynamics Patterns in a Rare Fish, Zingel asper, through Capture-Mark-Recapture Methods

Abstract:  We performed a capture-mark-recapture study on one of the last populations of Zingel asper , an endemic percid species of the Rhône River basin in France. The distribution of Z. asper has decreased dramatically during the last century. We sampled three sites in suitable habitats in the Beaume River. No impact of individual tagging on survival was found. The demography of the population was analyzed using capture-recapture methods that allow the estimation of survival, recruitment, and demographic growth rates. Annual survival rates were low (0.35–0.50). The level of transience was high (5% to 25%), suggesting that a significant number of individuals were highly mobile or shifted to suboptimal habitats. Seniority rates suggested random highly variable recruitment between years. The three sites had similar variation patterns in all demographic parameters, indicating broad spatial covariation in population dynamics. We found some local differences in demographic parameters, which could be linked to local habitat quality. Individual tagging allowed for the estimation of demographic parameters that improved our understanding of Z. asper population dynamics and revealed mechanisms that may affect population persistence, such as stochastic recruitment, low survival, and frequent dispersal. The fragmentation of habitat through river damming inhibits dispersal and represents a threat to the persistence of Z. asper in the Rhône basin. Our results offer evidence of the importance of dispersal in nonmigratory fishes and confirm the usefulness of individual tagging methods in rare fish demography.     

Trends in Sex Ratios of Turtles in the United States: Implications of Road Mortality

Abstract:  Road mortality has been implicated as a significant demographic force in turtles, particularly for females, which are killed disproportionately on overland nesting movements. Moreover, the United States' road network has expanded dramatically over the last century. We therefore predicted that historical trends in sex ratios of turtle populations would be male biased. To test this prediction, we synthesized published estimates of population-level sex ratios in freshwater and terrestrial turtles in the United States (165 estimates for 36 species, published 1928–2003). Our analysis suggests that the proportion of males in populations has increased linearly ( p = 0.001); the trend in male bias is synchronized with the expansion of the surfaced portion of the road network since 1930; sex ratios became more male biased in states with higher densities of roads; and populations have become more male biased in aquatic species, in which movement differentials between males and females are greatest, and are least biased in semiaquatic and terrestrial species, in which overland movements are more comparable between sexes. Our results suggest an ongoing depletion of breeding females from wild turtle populations over the last century because of many factors, including, and perhaps chiefly, road mortality.     

Modelling dispersal with diffusion and habitat selection: Analytical results for highly fragmented landscapes

Quantifying dispersal is crucial both for understanding ecological population dynamics, and for gaining insight into factors that affect the genetic structure of populations. The role of dispersal becomes pronounced in highly fragmented landscapes inhabited by spatially structured populations. We consider a landscape consisting of a set of habitat patches surrounded by unsuitable matrix, and model dispersal by assuming that the individuals follow a random walk with parameters that may be specific to the habitat type. We allow for spatial variation in patch quality, and account for edge-mediated behavior, the latter meaning that the individuals bias their movement towards the patches when close to an edge between a patch and the matrix. We employ a diffusion approximation of the random walk model to derive analytical expressions for various characteristics of the dispersal process. For example, we derive formulae for the time that an individual is expected to spend in its current patch i, and for the time that it will spend in the matrix, both conditional on the individual hitting next a given patch j before hitting any of the other patches or dying. The analytical formulae are based on the assumptions that the landscape is infinitely large, that the patches are circularly shaped, and that the patches are small compared to interpatch distances. We evaluate the effect of these assumptions by comparing the analytical results to numerical results in a real patch network that violates all of the three assumptions. We then consider a landscape that fulfills the assumptions, and show that in this case the analytical results are in a very good agreement with the numerical results. The results obtained here allow the construction of computationally efficient dispersal models that can be used as components of metapopulation models.     

A panmictic fiddler crab from the coast of Brazil? Impact of divergent ocean currents and larval dispersal potential on genetic and morphological variation in Uca maracoani

Marine species tend to exhibit relatively less population structuring than terrestrial species owing to fewer barriers to gene flow and increased connectivity resulting from greater dispersal abilities. Thus, in many cases, life history plays a more important role in phylogeography of marine taxa than do oceanographic features. Littoral species are of particular phylogeographic interest because they possess life histories that have both marine and terrestrial characteristics. This study evaluates the synergistic impact of divergent ocean currents and a high larval dispersal potential on the phylogeography of the fiddler crab, Uca maracoani, distributed along the coast of Brazil. Patterns of genetic variation were assessed with sequence data for a portion of the mitochondrial COI gene and AFLPs. Geometric morphometric techniques were used to evaluate morphological variation. Results revealed a lack of discernible genetic subdivision. However, geometric morphometrics showed statistically significant morphological differentiation. The absence of a clear phylogeographic pattern appears to be determined primarily by life history characteristics permitting a high level of connectivity. One, or a combination of several factors, may explain the incongruity between genetic and morphologic signatures, including phenotypic plasticity, incomplete lineage sorting, or recent and ongoing genetic divergence.     

Estimating Plecoglossus altivelis altivelis migration using a mass balance model expressed by hydrological distribution parameters in a major limpid river basin in Japan

We examined the influence of several hydrological and meteorological parameters on the migratory movements of ayu Plecoglossus altivelis altivelis in central Japan. When comprehensively evaluating rivers and ayu behaviour on a catchment scale, the subjects of analysis typically include human activities and hydrological and meteorological phenomena. However, limiting analyses to such factors may be too restrictive when human activities are being conducted. Accordingly, we incorporated a biological viewpoint into the evaluation method, analysing hydrological data (river discharge, river water temperature, sea water temperature) to determine watershed characteristics and examining the relationship between these characteristics and the habitat conditions of ayu. Then we constructed a numerical model for ayu migratory runs that incorporated ayu ecology and watershed characteristics. Analyses of ayu movements from a lower estuarine dam demonstrated that downstream displacements were associated with high water flows of more than 200 m³ s⁻¹ at the beginning of summer. We conclude that it is important to consider the effects of environmental parameters on the movements of different fish species to understand the causes of spatial variation in fish distribution in lowland rivers.     

Dispersal behaviour and colony structure in a colonial spider

Group living in spiders is characterised by two principle modes, the cooperative social mode and the colonial non-cooperative mode. Kin-relationships due to reduced dispersal determine population genetic structure in social spiders, but the dispersal mechanisms underlying group structure remain poorly understood in colonial spiders. Assuming similar ecological benefits of group living, we address the question whether reduced dispersal shapes population structure in a colonial spider, Cyrtophora citricola (Araneidae). We analysed dispersal by studying settling decisions under semi-natural conditions in experimental trees with and without colonies, and in natural populations, we estimated dispersal and colony structure using population genetic analyses. The propensity to disperse decreased with increasing age in experimental colonies. Adult females did not disperse in the experiment. Sub-adult female spiders preferred trees with a colony to trees without a colony. Dispersal in third instar juveniles was influenced significantly by wind but not by the presence of a colony. Thus, we showed that being in a colony did not inhibit juvenile dispersal, but pre-mating females were philopatric. Genetic differentiation among colonies in natural populations was heterogeneous, colonies being either little or highly differentiated. The heterogeneous structure is likely caused by colony founding by one or a few females followed by dispersal among perennial colonies. Gene flow, however, was slightly male-biased. The experimental and indirect, genetic approaches combined showed that dispersal and the breeding system of C. citricola resemble that of solitary spiders, with juvenile dispersal occurring in both sexes, while the colonial distribution is maintained by female philopatry.     

Maximum feasible distance of windborne cross-pollination in Brassica napus: A ‘mass budget’ model

A theory of gene dispersal by wind pollination can make an important contribution to understanding the viability and evolution of important plant groups in the Earth's changing landscape and it can be applied to evaluate concerns about the spread of engineered genes from genetically modified (GM) crops into conventional varieties via windborne pollen. Here, we present a model of cross-pollination between plant populations due to the wind. We perform a ‘mass budget’ of pollen by accounting for the number of pollen grains from release in the source population, dispersal from the source to the sink population by the wind, and deposition on receptive surfaces in the sink population. Our model can be parameterised for any wind-pollinated species, but we apply it to Brassica napus (oilseed rape or canola) to investigate the threat posed by wind pollination to GM confinement in agriculture. Specifically, we calculate the maximum feasible distance at which a particular level of windborne gene dispersal could be attained. This is equivalent to the separation distance between populations or fields required to achieve a given threshold of gene dispersal or adventitious GM presence. As required, model predictions of the upper bounds on levels of wind-mediated gene dispersal exceed observations from a wide range of published studies. For a level of gene dispersal below 0.9%, which is the EU threshold for GM adventitious presence, we predict that the maximum feasible distance for agricultural fields of B. napus is 1000 m, regardless of field shape and direction of prevailing winds. For fields closer than 1000 m, our model results do not necessarily imply that the 0.9% threshold is likely to be breached, because in this instance we have conservatively set the values of parameters where current knowledge is limited. We also predict that gene dispersal is reduced by 50% when the lag in peak flowering between the source and sink populations is 13 days, and reduced by 90% when the lag is 24 days. We identify further measurements necessary to improve the accuracy of the model predictions.     

The role of environmental and spatial processes in structuring lake communities from bacteria to fish

We assessed the relative roles of local environmental conditions and dispersal on community structure in a landscape of lakes for the major trophic groups. We use taxonomic presence-absence and abundance data for bacteria, phytoplankton, zooplankton, and fish from 18 lakes in southern Quebec, Canada. The question of interest was whether communities composed of organisms with more limited dispersal abilities, because of size and life history (zooplankton and fish) would show a different effect of lake distribution than communities composed of good dispersers (bacteria and phytoplankton). We examine the variation in structure attributable to local environmental (i.e., lake chemical and physical variables) vs. dispersal predictors (i.e., overland and watercourse distances between lakes) using variation partitioning techniques. Overall, we show that less motile species (crustacean zooplankton and fish) are better predicted by spatial factors than by local environmental ones. Furthermore, we show that for zooplankton abundances, both overland and watercourse dispersal pathways are equally strong, though they may select for different components of the community, while for fish, only watercourses are relevant dispersal pathways. These results suggest that crustacean zooplankton and fish are more constrained by dispersal and therefore more likely to operate as a metacommunity than are bacteria and phytoplankton within this studied landscape.     

Unveiling human-assisted dispersal mechanisms in invasive alien insects: Integration of spatial stochastic simulation and phenology models

Capturing the spread of biological invasions in heterogeneous landscapes is a complex modelling task where information on both dispersal and population dynamics needs to be integrated. Spatial stochastic simulation and phenology models have rarely been combined to assist in the study of human-assisted long-distance dispersal events.Here we develop a process-based spatially explicit landscape-extent simulation model that considers the spread and detection of invasive insects. Natural and human-assisted dispersal mechanisms are modelled with an individual-based approach using negative exponential and negative power law dispersal kernels and gravity models. The model incorporates a phenology sub-model that uses daily temperature grids for the prediction and timing of the population dynamics in each habitat patch. The model was applied to the study of the invasion by the important maize pest western corn rootworm (WCR) Diabrotica virgifera ssp. virgifera in Europe. We parameterized and validated the model using maximum likelihood and simulation methods from the historical invasion of WCR in Austria.WCR was found to follow stratified dispersal where international transport networks in the Danube basin played a key role in the occurrence of long-distance dispersal events. Detection measures were found to be effective and altitude had a significant effect on limiting the spread of WCR. Spatial stochastic simulation combined with phenology models, maximum likelihood methods and predicted versus observed regression showed a high degree of flexibility that captured the salient features of WCR spread in Austria. This modelling approach is useful because it allows to fully exploit and the often limited and heterogeneous information available regarding the population dynamics and dispersal of alien invasive insects.     

Multiscale patterns of genetic structure in a marine snail (Solenosteira macrospira) without pelagic dispersal

The northern Gulf of California (NGC) is one of the most dynamic and productive marine ecosystems in the world, yet knowledge about population connectivity and dispersal patterns is lacking for many of its resident species. Using nuclear and mitochondrial markers, we investigated the effects of open water, geographical distance and suitable habitat on patterns of genetic structure of Solenosteira macrospira, a benthic buccinid whelk with direct development. We collected samples in April 2004, 2005 and May 2007 from the upper NGC (31°34.39″N, 114°44.45″W). Phylogenetic analyses, hierarchical analyses of variance and Bayesian assignment tests substantiated a break between the east and west coasts. Genetic distance between population pairs increased with geographical distance, but only when assuming a U-shaped dispersal pathway over the open water of the NGC. Given S. macrospira’s association with rocky intertidal habitats, and its limited dispersal potential, we assumed that the geographical distribution of rocky habitat would play a significant role in genetic differentiation of S. macrospira. Nevertheless, populations separated by sand were more similar than populations separated by rocks. The influence of open water, geographical distance and suitable habitat (rocks vs. sand) also varied significantly across different genetic markers that presumably evolve at different rates. Specifically, the more rapidly evolving nuclear microsatellites suggested that physical transport processes strongly influence genetic differentiation on contemporary time scales, even in a species with direct benthic development. This underscores the strong, and potentially homogenizing, effect of present-day ocean circulation patterns in the NGC.     

Importance of Habitat Quality and Landscape Connectivity for the Persistence of Endangered Natterjack Toads

Abstract: The natterjack toad (Bufo calamita) is endangered in several parts of its distribution, including Belgium, where it occurs mainly in artificial habitats. We parameterized a general model for natterjack population viability analysis (PVA) and tested its sensitivity to changes in the values of basic parameters. Then we assessed the relative efficiency of various conservation measures in 2 situations: a small isolated population and a system of 4 populations connected by rare dispersal movements. We based the population viability analysis on a stage‐structured model of natterjack population dynamics. We parameterized the model in the RAMAS GIS platform with vital rates obtained from our own field experience and from published studies. Simulated natterjack populations were highly sensitive to habitat quality (particularly pond drying), to dispersal from surrounding local populations, and to a lesser extent to values of fecundity and survival of terrestrial stages. Population trajectories were nearly insensitive to initial abundances, carrying capacities, and the frequency of extreme climatic conditions. The simulations showed that in habitats with highly ephemeral ponds, where premetamorphosis mortality was high, natterjack populations nearly always had a very high extinction risk. We also illustrated how low dispersal rates (<1 dispersing individual/generation) efficiently rescued declining local populations. Such source‐sink dynamics demonstrate that the identification and management of source populations should be a high priority.     

Assessing the current state of ecological connectivity in a large marine protected area system

The establishment of marine protected areas (MPAs) is a critical step in ensuring the continued persistence of marine biodiversity. Although the area protected in MPAs is growing, the movement of individuals (or larvae) among MPAs, termed connectivity, has only recently been included as an objective of many MPAs. As such, assessing connectivity is often neglected or oversimplified in the planning process. For promoting population persistence, it is important to ensure that protected areas in a system are functionally connected through dispersal or adult movement. We devised a multi-species model of larval dispersal for the Australian marine environment to evaluate how much local scale connectivity is protected in MPAs and determine whether the extensive system of MPAs truly functions as a network. We focused on non-migratory species with simplified larval behaviors (i.e., passive larval dispersal) (e.g., no explicit vertical migration) as an illustration. Of all the MPAs analyzed (approximately 2.7 million km²), outside the Great Barrier Reef and Ningaloo Reef, <50% of MPAs (46-80% of total MPA area depending on the species considered) were functionally connected. Our results suggest that Australia's MPA system cannot be referred to as a single network, but rather a collection of numerous smaller networks delineated by natural breaks in the connectivity of reef habitat. Depending on the dispersal capacity of the taxa of interest, there may be between 25 and 47 individual ecological networks distributed across the Australian marine environment. The need to first assess the underlying natural connectivity of a study system prior to implementing new MPAs represents a key research priority for strategically enlarging MPA networks. Our findings highlight the benefits of integrating multi-species connectivity into conservation planning to identify opportunities to better incorporate connectivity into the design of MPA systems and thus to increase their capacity to support long-term, sustainable biodiversity outcomes.     

Assessing biodegradation benefits from dispersal networks

The performance of biodegradation of organic pollutants in soil often depends on abiotic conditions and the bioavailability of these pollutants to degrading bacteria. In this context, bacterial dispersal is an essential aspect. Recent studies on the potential promotion of bacterial dispersal by fungal hyphae raised the idea of specifically applying fungal networks to accelerate bacterial degradation processes in situ. Our objective is to investigate these processes and their performance via simulation modelling and address the following questions: (1) Under what abiotic conditions can dispersal networks significantly improve bacterial degradation? and (2) To what extent does the spatial configuration of the networks influence the degradation performance? To answer these questions, we developed a spatially explicit bacterial colony model, which is applied to controlled laboratory experiments with Pseudomonas putida G7 organisms as a case study. Using this model, we analyzed degradation performance in response to different environmental scenarios and showed that conditions of limited bacterial dispersal also limit degradation performance. Under such conditions, dispersal networks have the highest potential for improving the bioavailability of pollutants to bacteria. We also found that degradation performance significantly varies with the spatial configuration of the dispersal networks applied and the time horizon over which performance is assessed. Regarding future practical applications, our results suggest that (1) fungal networks may dramatically improve initially adverse conditions for biodegradation of pollutants in soil, and (2) the network's spatial structure and accessibility are decisive for the success of such tasks.     

Genetic differentiation in Littorina saxatilis (Gastropoda)

Littorina saxatilis, an ovoviviparous gastropod, displays considerable morphological variability, perhaps due to limited dispersal abilities. Gene and genotype frequencies of 2 polymorphic enzymatic loci were determined electrophoretically for 7 populations of L. saxatilis, to estimate degree of population differentiation. Results show that significant population differentiation may occur over distances of as little as 2 km, while widely separated populations may be nearly identical. An index of probability of genotypic identity calculated for populations of L. saxatilis shows variable and heterogeneous values. Comparison of this index for L. saxatilis with an index for 12 populations of Nassarius obsoletus —which possesses strong dispersal abilities due to long-term veliger larva and high, homogeneous indices of probability of genotypic identity between populations — indicates a strong correlation between population differentiation and dispersal ability of the organism. Possible factors influencing population differentiation in L. saxatilis are discussed.     

Proposed Conservation Landscape for Giant Pandas in the Minshan Mountains,China

Abstract: The giant panda (Ailuropoda melanoleuca), is one of the world's most endangered species. Habitat loss and fragmentation have reduced its numbers, shrunk its distribution, and separated the population into isolated subpopulations. Such isolated, small populations are in danger of extinction due to random demographic factors and inbreeding. We used least‐cost modeling as a systematic approach to incorporate satellite imagery and data on ecological and behavioral parameters of the giant panda collected during more than 10 years of field research to design a conservation landscape for giant pandas in the Minshan Mountains. We identified 8 core habitats and 4 potential linkages that would link core habitats CH3, CH4, and CH5 with core habitats CH6, CH7, and CH8. Establishing and integrating the identified habitats with existing reserves would create an efficient reserve network for giant panda conservation. The core habitats had an average density of 4.9 pandas/100 km² and contained approximately 76.6% of the giant panda population. About 45% of the core habitat (3245.4 km²) existed outside the current nature reserves network. Total estimated core habitat decreased between 30.4 and 44.5% with the addition of residential areas and road networks factored into the model. A conservation area for giant panda in the Minshan Mountains should aim to ensure habitat retention and connectivity, improve dispersal potential of corridors, and maintain the evolutionary potential of giant pandas in the face of future environmental changes.