bioRxiv Science⌕ Search

Biology subjects

Germain, R. M.

Publications and source records attributed to Germain, R. M..

12 recordsLinked to original sources

Niche differentiation confers coexistence prior to the species boundary in an aquatic plant

Despite the prevailing view that ecological divergence drives speciation, we know little about when or how nascent lineages evolve the ecological differences needed to coexist upon secondary contact. Here, we apply ecological coexistence theory to quantify the potential for coexistence among 126 allopatric lineages of the globally distributed duckweed Spirodela polyrhiza. Using competition experiments simulating secondary contact, we found that rapid accumulation of niche differences stabilized coexistence to permit sympatry among potentially interbreeding lineages. Competition against sister-species Spirodela intermedia further showed that niche differences accumulate more slowly post-speciation, revealing that niche differences enabling coexistence evolve well before timescales at which speciation is complete. Our findings suggest that rapid coexistence may thus contribute to time-lags in speciation, shaping both the origin and maintenance of biodiversity.

ecology↗

Sink invaders can increase resident metapopulation size by modifying local conditions and regional connectivity

Populations are regulated by processes operating at both local and regional scales. As a result, species can interact not only within local communities but also through regional-scale processes. However, the feedback between local and regional processes of species interactions in shaping metapopulation dynamics have rarely been tested experimentally. Using nectar microbes as a model system, we examine how a bacterial species dispersing to a sink habitat, sustained by external supply, can interact with a resident yeast species directly by modifying local nectar chemistry and indirectly by altering metapopulation connectivity. We show that the external supply of bacteria increases resident yeast densities, potentially through increased local productivity, and that this benefit is enhanced when the dispersal rate of yeast is reduced in patches supplied with bacteria. Analysis of a metapopulation model indicates that the benefit of dispersal modification on metapopulation size occurs only when local productivity differs across patches, consistent with our interpretation that bacterial supply increases local productivity. Taken together, our results underscore the role of dispersal in shaping how metapopulations experience density-dependence across heterogeneous landscapes, highlight the relevance of material flows across ecosystem boundaries, and suggest a plausible mechanism in which transient species may leave lasting impacts on ecological communities.

ecology↗

Dispersal kernels influence the magnitude of environmental, biotic, and stochastic effects on the maintenance of metacommunity diversity

Dispersal plays a central role in shaping patterns of diversity in metacommunities. However, a primary focus on emigration rates may mischaracterize dispersal effects that actually arise from dispersal kernels. Kernels describe probabilistic movements between donor and recipient patches, but the influence of kernel shape on metacommunity diversity remains unclear. We used simulations to measure how kernels affect diversity across metacommunity scales and ecological contexts. We disentangled causes of these patterns using a novel approach quantifying the effects of environmental filtering, competition, stochasticity, and dispersal on fitness. Although metacommunities with shallow kernels followed expectations where emigration increased alpha-but decreased beta- and gamma-diversity, metacommunities with steeper kernels did not. Steeper kernels maintained regional diversity by reducing interspecific competition and stochastic extinctions, with dispersal conferring weaker benefits but less homogenization. Our work suggests dispersal kernels and emigration rates jointly regulate exposure to environmental variation and the balance of assembly mechanisms in metacommunities.

ecology↗

Spatial fingerprint of consumer body size and habitat preference on resource distribution

Consumers shape spatial patterns on landscapes by amplifying or dampening environmental heterogeneity through feeding, excretion, and movement of resources. The degree to which the environment is modified by consumers depends on species traits, including body mass, movement and foraging behavior, and habitat specialization. Global change is altering the size and traits of consumer populations, but our understanding of how this may impact resource heterogeneity is limited. Here, we developed an individual-based model of habitat specialists and generalists movement and activity in a patchy landscape and investigated the impact of changes in population and mean body sizes on landscape-scale resource heterogeneity. We found that consumers specializing on low-resource habitats (a common risk avoidance strategy) increased spatial resource heterogeneity regardless of their population and body size. By contrast, generalists eroded differences among habitats, and we further found that resource heterogeneity decreased with the average body size of generalist consumers, even while controlling for total consumer biomass. Larger perceptual ranges increased the area over which generalist consumers could select foraging habitat, and reduced the extent to which they eroded landscape structure. These nuanced spatial outcomes of consumer-resource interactions emerge from how metabolic demands, which scale nonlinearly with body size, play out among habitat types which attract different consumers, as well as the scale at which those consumers make habitat selection decisions. Since global change disproportionately impacts larger species and specialists, indirect consequences on ecosystems may arise via biotic processes, affecting spatial heterogeneity of future landscapes.

ecology↗

Macroecological variation in movement profiles: body size does not explain it all

Animals couple habitats by three types of movement: dispersal, migration, and foraging, which dynamically link populations, communities, and ecosystems. Spatial distances of movement tend to correlate with each other, reflecting shared allometric scaling with body size, but may diverge due to biomechanical, phylogenetic, and ecological constraints. While these constraints have been investigated within specific taxa, the macroecological and macroevolutionary constraints on movement distances, and causes of those constraints, are still unknown. Here, we synthesized distances of all three movement types across 300+ vertebrate species, and investigated how the relationships between movement types and body size were modified by movement medium, taxonomy, and trophic guild (carnivore, herbivore, etc.). We found that the strength of relationships between movement types and body size varied among environments, taxa, and trophic guilds. Movement profiles interacted with physiological, taxonomic, and ecological traits to depart from expected body mass scaling. Overall, we find that there are systematic patterns to movement distances, and that movement types with very distinct ecological consequences (foraging, migration) can be correlated and subject to similar constraints. This implies that the scales of population dynamics in ecological communities are not entirely determined by the environment and likely reflect general biomechanical, evolutionary and metabolic constraints.

ecology↗

Linking intrinsic scales of ecological processes to characteristic scales of biodiversity and functioning patterns

Ecology is a science of scale, which guides our description of both ecological processes and patterns, but we lack a systematic understanding of how process scale and pattern scale are connected. Recent calls for a synthesis between population ecology, community ecology, and ecosystem ecology motivate the integration of phenomena at multiple levels of organization. Furthermore, many studies leave out the scaling of a critical process: species interactions, which may be non-local through movement or foraging and must be distinguished from dispersal scales. Here, we use simulations to explore the consequences of three different process scales (species interactions, dispersal, and the environment) on emergent patterns of biodiversity, ecosystem functioning, and their relationship, in a spatially-explicit landscape and stable equilibrium setting. A major result of our study is that the spatial scales of dispersal and species interactions have opposite effects: a larger dispersal scale homogenizes spatial biomass patterns, while a larger interaction scale amplifies their heterogeneity. Interestingly, the specific scale at which dispersal and interaction scales begin to influence landscape patterns depends on the scale of environmental heterogeneity - in other words, the scale of one process allows important scales to emerge in other processes. This interplay between process scales, i.e., a situation where no single process dominates, can only occur when the environment is heterogeneous and the scale of dispersal small. Finally, contrary to our expectations, we observe that the spatial scale of ecological processes is more clearly reflected in landscape patterns (i.e., distribution of local outcomes) than in global patterns such as Species-Area Relationships or large-scale biodiversity-functioning relationships. Overall we conclude that long-range interactions often act differently and even in opposite ways to dispersal, and that the landscape patterns that emerge from the interplay of long-ranged interactions, dispersal and environmental heterogeneity are not well captured by often-used metrics like the Species-Area Relationship.

ecology↗

Dispersal across habitat boundaries: uncovering the demographic fates of populations in unsuitable habitat

Patchy landscapes are characterized by abrupt transitions among distinct habitat types, forcing species to cross habitat boundaries in order to spread. Since seed dispersal is a probabilistic process, with a kernel that decays with distance, most individuals will fail to reach new, suitable habitat. Although failed dispersers are presumed dead in population models, their demographic fates may not be so simple. If transient survival is possible within unsuitable habitat, then through time, individuals may be able to reach distant, suitable habitat, forming new populations and buffering species from extinction. In a fragmented Californian grassland, we explored the fates of individuals that crossed habitat boundaries, and if those fates differed among specialists dispersing from two habitat types: serpentine habitat patches and the invaded non-serpentine matrix. We surveyed the diversity of seedbank and adult life stages along transects that crossed boundaries between patches and the matrix. First, we considered how patch specialists might transiently survive in the matrix via seed dormancy or stepping-stone populations. Second, we investigated the dispersal of an invasive matrix specialist (Avena fatua) into patches, to assess if sink populations existed across the habitat boundary. We found that dormancy maintained populations of patch specialists deep into the matrix, as abundances of seedbanks and of adult plant communities differed with distance into the matrix. We found evidence that these dormant seeds disperse secondarily with vectors of material flows in the landscape, suggesting that they could eventually reach suitable patches even if they first land in the matrix. We found that A. fatua were largely absent deep in patches, where reproductive outputs plummeted and there was no evidence of a dormant seedbank. Our results not only reveal the demographic fates of individuals that land in unsuitable habitat, but that their ecological consequences differ depending on the direction by which the boundary is crossed (patch [->] matrix = matrix [->] patch). Dormancy is often understood as a mechanism for persisting in face of temporal variability, but it may serve as a means of traversing unsuitable habitat in patchy systems, warranting its consideration in estimates of habitat connectivity.

ecology↗

Climate warming weakens local adaptation

Anthropogenic climate change is generating mismatches between the environmental conditions that populations historically experienced and those in which they reside. Understanding how climate change affects population performance is a critical scientific challenge. We combine a quantitative synthesis of field transplant experiments with a novel statistical approach based in evolutionary theory to quantify the effects of temperature and precipitation variability on population performance. We find that species average performance is affected by both temperature and precipitation, but populations show signs of local adaptation to temperature only. Contemporary responses to temperature are strongly shaped by the local climates under which populations evolved, resulting in performance declines when temperatures deviate from historic conditions. Adaptation to other local environmental factors is strong, but temperature deviations as small as 2{degrees}C erode the advantage that these non-climatic adaptations historically gave populations in their home sites. One sentence summaryClimate change is pulling the thermal rug out from under populations, reducing average performance and eroding their historical home-site advantage.

evolutionary biology↗

Environmental variability and the evolution of bet hedging strategies

Bet-hedging strategies, such as dispersal and dormancy, are predicted to evolve in varying and uncertain environments and are critical to ecological models of biodiversity maintenance. Theories of the specific ecological scenarios that favor the evolution of dispersal, dormancy, or their covariance are rarely tested, particularly for naturally-evolved populations that experience complex patterns of spatiotemporal environmental variation. We grew 23 populations of Vulpia microstachys, an annual grass native to California, in a greenhouse, and on the offspring generation measured seed dispersal ability and dormancy rates. We hypothesized that seed dormancy rates and dispersal abilities would be highest in populations from more productive, temporally variable sites, causing them to covary positively. Contrary to our hypothesis, our data suggest that both dispersal and dormancy evolve to combat different axes and scales of spatial heterogeneity, and are underlain by different seed traits, allowing them to evolve independently. Dormancy appears to have evolved as a strategy for overcoming microgeographic heterogeneity rather than temporal climate fluctuations, an outcome that to our knowledge has not been considered by theory. In sum, we provide much needed empirical data on the evolution of bet hedging, as well as a new perspective on the ecological function dormancy provides in heterogeneous landscapes.

ecology↗

An experimental manipulation of species’ phenologies overturns competitive hierarchies

Ecological theory produces opposing predictions about whether differences in the timing of life history transitions, or phenology, promote or limit coexistence. Phenological separation is predicted to create temporal niche differences, increasing coexistence, yet phenological separation may competitively favour one species, increasing fitness differences and hindering coexistence. We experimentally manipulated relative germination timing, a critical phenological event, of two annual grass species, Vulpia microstachys and V. octoflora, to test these contrasting predictions. We parameterized a competition model to estimate within-season niche differences, fitness differences, and coexistence, and to estimate coexistence when among-year fluctuations of germination timing occur. Increasing germination separation caused parallel changes in niche and fitness differences, with the net effect of weakening within-year coexistence. Both species experienced a competitive advantage by germinating earlier, strongly enough to allow the generally inferior competitor to exclude the other with at least a four day head start. The overall consequence of germination separation was to limit coexistence within a given year, although among-year variation in relative timing of germination was sufficient to support long-term coexistence. Our results clarify how phenological differences structure competitive interactions, and highlight the need to quantify among-year variation in these differences to better understand species coexistence.

ecology↗

Cryptic Dispersal Networks Connect Habitat Patches in ways not Captured by Distance

Species interact with the physical world in complex ways, and life history strategies might cause species to differ in how they experience connectedness of the same landscape. As a consequence, dispersal limitation might be present but not captured by distance-based measures of connectivity. To test these ideas, we surveyed plant communities that associate with serpentine soils but differ in dispersal mode (gravity, animal, or wind), and used satellite imagery to quantify forms of landscape connectivity associated with each dispersal mode. Our data yielded two key insights: First, dispersal limitation appeared to be absent using a conventional distance-based measure of connectivity, but emerged after considering forms of landscape connectivity relevant to each dispersal mode. Second, the landscape variables that emerged as important to each dispersal mode were generally consistent with our predictions based on putative dispersal vectors, and included interactive effects that allude to the altered efficacy of animal dispersal in invaded landscapes. Our results have broad implications for understanding how ecological communities reorganize as landscapes are fragmented, patches are lost, and the function of dispersal life histories is altered.

ecology↗

Frequency-dependent Maternal Effects across Species and Environments

O_LIMaternal provisioning of offspring in response to environmental conditions (\"maternal environmental effects\") has been argued as the missing link in plant life histories. Although empirical evidence suggests that maternal responses to abiotic conditions are common, there is little understanding of the prevalence of maternal provisioning in competitive environments.\nC_LIO_LIWe tested how competition in two soil moisture environments affects maternal provisioning of offspring seed mass. Specifically, we varied conspecific frequency from 90% (intraspecific competition) to 10% (interspecific competition) for 15 pairs of annual plant species that occur in California.\nC_LIO_LIWe found that conspecific frequency affected maternal provisioning (seed mass) in 48% of species, and that these responses included both increased (20%) and decreased (24%) seed mass. In contrast, 68% of species responded to competition through changes in per capita fecundity (seed number), which generally decreased as conspecific frequency increased. The direction and magnitude of frequency-dependent seed mass depended on the identity of the competitor, even among species in which fecundity was not affected by competitor identity.\nC_LIO_LISynthesis. Our research demonstrates how species responses to different competitive environments manifest through maternal provisioning, and that these responses alter previous estimates of environmental maternal effects and reproductive output; future study is needed to understand their combined effects on population and community dynamics.\nC_LI

ecology↗