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How species richness scales spatially is a foundational concept of community ecology, but how biotic interactions scale spatially is poorly known. Previous studies have proposed interactions-area relationships (IARs) based on two competing relationships for how the number of interactions scale with the number of species, the ‘link-species scaling law’ and the ‘constant connectance hypothesis.’ The link-species scaling law posits that the number of interactions per species remains constant as the size of the network increases. The constant connectance hypothesis says that the proportion of realized interactions remains constant with network size. While few tests of these IARs exist, evidence for the original interactions-species relationships are mixed. We propose a novel IAR and test it against the two existing IARs. We first present a general theory for how interactions scale spatially and the mathematical relationship between the IAR and the species richness-area curve. We then provide a new mathematical formulation of the IAR, accounting for connectance varying with area. Employing data from three mutualistic networks (i.e. a network which specifies interconnected and mutually-beneficial interactions between two groups of species), we evaluate three competing models of how interactions scale spatially: two previously published IAR models and our proposed IAR. We find the new IAR described by our theory-based equation fits the empirical datasets equally as well as the previously proposed IAR based on the link-species scaling law in one out of three cases and better than the previously-proposed models in two out of three cases. Our novel IAR improves upon previous models and quantifies mutualist interactions across space, which is paramount to understanding biodiversity and preventing its loss.

 

Phenological shifts are a widely studied consequence of climate change. Little is known, however, about certain critical phenological events, nor about mechanistic links between shifts in different life-history stages of the same organism. Among angiosperms, flowering times have been observed to advance with climate change, but, whether fruiting times shift as a direct consequence of shifting flowering times, or respond differently or not at all to climate change, is poorly understood. Yet, shifts in fruiting could alter species interactions, including by disrupting seed dispersal mutualisms. In the absence of long-term data on fruiting phenology, but given extensive data on flowering, we argue that an understanding of whether flowering and fruiting are tightly linked or respond independently to environmental change can significantly advance our understanding of how fruiting phenologies will respond to warming climates. Through a case study of biotically and abiotically dispersed plants, we present evidence for a potential functional link between the timing of flowering and fruiting. We then propose general mechanisms for how flowering and fruiting life history stages could be functionally linked or independently driven by external factors, and we use our case study species and phenological responses to distinguish among proposed mechanisms in a real-world framework. Finally, we identify research directions that could elucidate which of these mechanisms drive the timing between subsequent life stages. Understanding how fruiting phenology is altered by climate change is essential for all plant species but is particularly critical to sustaining the large numbers of plant species that rely on animal-mediated dispersal, as well as the animals that rely on fruit for sustenance. 

Habitat loss disrupts species interactions through local extinctions, potentially orphaning species that depend on interacting partners, via mutualisms or commensalisms, and increasing secondary extinction risk. Orphaned species may become functionally or secondarily extinct, increasing the severity of the current biodiversity crisis. While habitat destruction is a major cause of biodiversity loss, the number of secondary extinctions is largely unknown. We investigate the relationship between habitat loss, orphaned species, and bipartite network properties. Using a real seed dispersal network, we simulate habitat loss to estimate the rate at which species are orphaned. To be able to draw general conclusions, we also simulate habitat loss in synthetic networks to quantify how changes in network properties affect orphan rates across broader parameter space. Both real and synthetic network simulations show that even small amounts of habitat loss can cause up to 10% of species to be orphaned. More area loss, less connected networks, and a greater disparity in the species richness of the network's trophic levels generally result in more orphaned species. As habitat is lost to land‐use conversion and climate change, more orphaned species increase the loss of community‐level and ecosystem functions. However, the potential severity of repercussions ranges from minimal (no species orphaned) to catastrophic (up to 60% of species within a network orphaned). Severity of repercussions also depends on how much the interaction richness and intactness of the community affects the degree of redundancy within networks. Orphaned species could add substantially to the loss of ecosystem function and secondary extinction worldwide.

 

Abstract There is growing realization that intraspecific variation in seed dispersal can have important ecological and evolutionary consequences. However, we do not have a good understanding of the drivers or causes of intraspecific variation in dispersal, how strong an effect these drivers have, and how widespread they are across dispersal modes. As a first step to developing a better understanding, we present a broad, but not exhaustive, review of what is known about the drivers of intraspecific variation in seed dispersal, and what remains uncertain. We start by decomposing ‘drivers of intraspecific variation in seed dispersal’ into intrinsic drivers (i.e. variation in traits of individual plants) and extrinsic drivers (i.e. variation in ecological context). For intrinsic traits, we further decompose intraspecific variation into variation among individuals and variation of trait values within individuals. We then review our understanding of the major intrinsic and extrinsic drivers of intraspecific variation in seed dispersal, with an emphasis on variation among individuals. Crop size is the best-supported and best-understood intrinsic driver of variation across dispersal modes; overall, more seeds are dispersed as more seeds are produced, even in cases where per seed dispersal rates decline. Fruit/seed size is the second most widely studied intrinsic driver, and is also relevant to a broad range of seed dispersal modes. Remaining intrinsic drivers are poorly understood, and range from effects that are probably widespread, such as plant height, to drivers that are most likely sporadic, such as fruit or seed colour polymorphism. Primary extrinsic drivers of variation in seed dispersal include local environmental conditions and habitat structure. Finally, we present a selection of outstanding questions as a starting point to advance our understanding of individual variation in seed dispersal. 

Abstract The distribution and abundance of plants across the world depends in part on their ability to move, which is commonly characterized by a dispersal kernel. For seeds, the total dispersal kernel (TDK) describes the combined influence of all primary, secondary and higher-order dispersal vectors on the overall dispersal kernel for a plant individual, population, species or community. Understanding the role of each vector within the TDK, and their combined influence on the TDK, is critically important for being able to predict plant responses to a changing biotic or abiotic environment. In addition, fully characterizing the TDK by including all vectors may affect predictions of population spread. Here, we review existing research on the TDK and discuss advances in empirical, conceptual modelling and statistical approaches that will facilitate broader application. The concept is simple, but few examples of well-characterized TDKs exist. We find that significant empirical challenges exist, as many studies do not account for all dispersal vectors (e.g. gravity, higher-order dispersal vectors), inadequately measure or estimate long-distance dispersal resulting from multiple vectors and/or neglect spatial heterogeneity and context dependence. Existing mathematical and conceptual modelling approaches and statistical methods allow fitting individual dispersal kernels and combining them to form a TDK; these will perform best if robust prior information is available. We recommend a modelling cycle to parameterize TDKs, where empirical data inform models, which in turn inform additional data collection. Finally, we recommend that the TDK concept be extended to account for not only where seeds land, but also how that location affects the likelihood of establishing and producing a reproductive adult, i.e. the total effective dispersal kernel. 

Understanding patterns of diversity is central to ecology and conservation, yet estimates of diversity are often biased by imperfect detection. In recent years, multi‐species occupancy models (MSOM) have been developed as a statistical tool to account for species‐specific heterogeneity in detection while estimating true measures of diversity. Although the power of these models has been tested in various ways, their ability to estimate gamma diversity—or true community size,Nis a largely unrecognized feature that needs rigorous evaluation.

We use both simulations and an empirical dataset to evaluate the bias, precision, accuracy and coverage of estimates ofNfrom MSOM compared to the widely applied iChao2 non‐parametric estimator. We simulated 5,600 datasets across seven scenarios of varying average occupancy and detectability covariates, as well as varying numbers of sites, replicates and true community size. Additionally, we use a real dataset of surveys over 9 years (where species accumulation reached an asymptote, indicating trueN), to estimateNfrom each annual survey.

Simulations showed that both MSOM and iChao2 estimators are generally accurate (i.e. unbiased and precise) except under unideal scenarios where mean species occupancy is low. In such scenarios, MSOM frequently overestimatedN. Across all scenarios, MSOM estimates were less certain than iChao2, but this led to over‐confident iChao2 estimates that showed poor coverage. Results from the real dataset largely confirmed the simulation findings, with MSOM estimates showing greater accuracy and coverage than iChao2.

Community ecologists have a wide choice of analytical methods, and both iChao2 and MSOM estimates ofNare substantially preferable to raw species counts. The simplicity of non‐parametric estimators has obvious advantages, but our results show that in many cases, MSOM may provide superior estimates that also account more accurately for uncertainty. Both methods can show strong bias when average occupancy is very low, and practitioners should show caution when using estimates derived from either method under such conditions.

 

Abstract Although dispersal is generally viewed as a crucial determinant for the fitness of any organism, our understanding of its role in the persistence and spread of plant populations remains incomplete. Generalizing and predicting dispersal processes are challenging due to context dependence of seed dispersal, environmental heterogeneity and interdependent processes occurring over multiple spatial and temporal scales. Current population models often use simple phenomenological descriptions of dispersal processes, limiting their ability to examine the role of population persistence and spread, especially under global change. To move seed dispersal ecology forward, we need to evaluate the impact of any single seed dispersal event within the full spatial and temporal context of a plant’s life history and environmental variability that ultimately influences a population’s ability to persist and spread. In this perspective, we provide guidance on integrating empirical and theoretical approaches that account for the context dependency of seed dispersal to improve our ability to generalize and predict the consequences of dispersal, and its anthropogenic alteration, across systems. We synthesize suitable theoretical frameworks for this work and discuss concepts, approaches and available data from diverse subdisciplines to help operationalize concepts, highlight recent breakthroughs across research areas and discuss ongoing challenges and open questions. We address knowledge gaps in the movement ecology of seeds and the integration of dispersal and demography that could benefit from such a synthesis. With an interdisciplinary perspective, we will be able to better understand how global change will impact seed dispersal processes, and potential cascading effects on plant population persistence, spread and biodiversity.