Building modern coexistence theory from the ground up: The role of community assembly

Modern coexistence theory (MCT) is one of the leading methods to understand species coexistence. It uses invasion growth rates—the average, per-capita growth rate of a rare species—to identify when and why species coexist. Despite significant advances in dissecting coexistence mechanisms when coexistence occurs, MCT relies on a ‘mutual invasibility’ condition designed for two-species communities but poorly defined for species-rich communities. Here, we review well-known issues with this component of MCT and propose a solution based on recent mathematical advances. We propose a clear framework for expanding MCT to species-rich communities and for understanding invasion resistance as well as coexistence, especially for communities that could not be analysed with MCT so far. Using two data-driven community models from the literature, we illustrate the utility of our framework and highlight the opportunities for bridging the fields of community assembly and species coexistence.     

Shifts of community composition and population density substantially affect ecosystem function despite invariant richness

There has been considerable focus on the impacts of environmental change on ecosystem function arising from changes in species richness. However, environmental change may affect ecosystem function without affecting richness, most notably by affecting population densities and community composition. Using a theoretical model, we find that, despite invariant richness, (1) small environmental effects may already lead to a collapse of function; (2) competitive strength may be a less important determinant of ecosystem function change than the selectivity of the environmental change driver and (3) effects on ecosystem function increase when effects on composition are larger. We also present a complementary statistical analysis of 13 data sets of phytoplankton and periphyton communities exposed to chemical stressors and show that effects on primary production under invariant richness ranged from ?75% to +10%. We conclude that environmental protection goals relying on measures of richness could underestimate ecological impacts of environmental change.     

Effects of Juvenile Host Density and Food Availability on Adult Immune Response,Parasite Resistance and Virulence in a Daphnia-Parasite System

Host density can increase infection rates and reduce host fitness as increasing population density enhances the risk of becoming infected either through increased encounter rate or because host condition may decline. Conceivably, potential hosts could take high host density as a cue to up-regulate their defence systems. However, as host density usually covaries with food availability, it is difficult to examine the importance of host density in isolation. Thus, we performed two full-factorial experiments that varied juvenile densities of Daphnia magna (a freshwater crustacean) and food availability independently. We also included a simulated high-density treatment, where juvenile experimental animals were kept in filtered media that previously maintained Daphnia at high-density. Upon reaching adulthood, we exposed the Daphnia to their sterilizing bacterial parasite, Pasteuria ramosa, and examined how the juvenile treatments influenced the likelihood and severity of infection (Experiment I) and host immune investment (Experiment II). Neither juvenile density nor food treatments affected the likelihood of infection; however, well-fed hosts that were well-fed as juveniles produced more offspring prior to sterilization than their less well-fed counterparts. By contrast, parasite growth was independent of host juvenile resources or host density. Parasite-exposed hosts had a greater number of circulating haemocytes than controls (i.e., there was a cellular immune response), but the magnitude of immune response was not mediated by food availability or host density. These results suggest that density dependent effects on disease arise primarily through correlated changes in food availability: low food could limit parasitism and potentially curtail epidemics by reducing both the host’s and parasite’s reproduction as both depend on the same food.     

Intuitive and broadly applicable definitions of niche and fitness differences

Explaining nature’s biodiversity is a key challenge for science. To persist, populations must be able to grow faster when rare, a feature called negative frequency dependence and quantified as ‘niche differences’ () in modern coexistence theory. Here, we first show that available definitions of differ in how link to species interactions, are difficult to interpret and often apply to specific community types only. We then present a new definition of that is intuitive and applicable to a broader set of (modelled and empirical) communities than is currently the case, filling a main gap in the literature. Given , we also redefine fitness differences () and illustrate how and determine coexistence. Finally, we demonstrate how to apply our definitions to theoretical models and experimental data, and provide ideas on how they can facilitate comparison and synthesis in community ecology.     

Evidence from morphological and genetic data confirms that Colossendeis tenera Hilton, 1943 (Arthropoda: Pycnogonida), does not belong to the Colossendeis megalonyx Hoek, 1881 complex

Within the Pycnogonida, genetic studies have revealed that Colossendeis megalonyx Hoek (Challenger Report, Zoology, 3(X), 1–167, 1881), consists of a complex of several cryptic or overlooked species. Colossendeis megalonyx is a typical Southern Hemisphere species complex distributed primarily on the continental shelves in the Antarctic and Subantarctic. However, a different Colossendeis species with a completely different geographic distribution range, Colossendeis tenera Hilton (Journal of Entomology and Zoology, Pomona College, Claremont, 35(1), 2–4, 1943), was considered a subspecies of Colossendeis megalonyx by Turpaeva (Trudy Instituta Okeanology "P. P. Shirshova", Akademy Nauk SSSR, 103, 230–246, 1975). Colossendeis tenera occurs predominantly along the Pacific Coast of North America from the Bering Sea to central California. Prominent differences between these two currently distinct species are found in body proportions and other characters that were interpreted by Turpaeva as a possible case of pedomorphosis induced by deep-sea conditions. In this study, we tested the hypothesis that Colossendeis tenera belongs to the Colossendeis megalonyx complex by analyzing available and novel sequence data (CO1 and H3) of both Colossendeis megalonyx and Colossendeis tenera as well as a similar, apparently closely related species, Colossendeis angusta Sars (Archiv for Mathematik og Naturvidenskab, 2, 237–271, 1877). We compared morphometric data and SEM of the ovigera of these species. Our results clearly indicate that Colossendeis tenera and Colossendeis angusta are not a part of the Colossendeis megalonyx complex. A sister-group relationship of Colossendeis tenera and Colossendeis angusta is strongly supported, but Colossendeis tenera is not clearly resolved as monophyletic with respect to Colossendeis angusta. This work highlights the need for further examination of the variation found in the tenera-angusta clade. It also gives a first hint of the phylogenetic affinities of species within Colossendeis.     

Parasite survey of a Daphnia hybrid complex: host-specificity and environment determine infection

1. Hybridization between species is a common phenomenon in plants and animals. If parasite prevalence differs for hybrids and parental species (i.e. taxa) there may be considerable consequences for relative hybrid fitness. Some studies have investigated hybrid complexes for infection, and complex-specific differences in parasite prevalence have been detected. 2. Based on the results of a field study on a hybridizing Daphnia population from a single lake, it has been hypothesized that permanently over- or under-infected hybrids do not exist. The observed field-patterns can only be temporal because taxa, in addition to single genotypes, might be the subject of parasite driven host frequency-dependent selection. Thus, parasites will track any common taxon within a hybrid complex. 3. In the present study, hybridizing Daphnia populations from 43 lakes were screened for parasite infections to obtain indirect evidence for coevolutionary cycles. It was hypothesized that, due to time lags between the evolution of resistance in host populations and the evolution of the parasite towards tracking of a common host taxon, the same Daphnia taxon will be over-infected in some lakes, while being under-infected in others. 4. Two of the four parasite species were specialists: their prevalence differed among coexisting Daphnia taxa. The varying infection patterns detected across spatially segregated hybridizing Daphnia populations are consistent with theoretical predictions for coevolutionary cycles. Thus the infection patterns, as observed under natural conditions, are temporal and unstable. 5. Additionally, the spatial distribution of the four parasite species was analysed with respect to habitat differences. The results show that the presence of a particular parasite on a host taxon was determined not only by the host-specificity of the parasite, but also by host-habitat relations.     

Epidemiology of a <Emphasis Type="Italic">Daphnia</Emphasis> brood parasite and its implications on host life-history traits

Parasites influence host life-history traits and therefore might crucially shape host populations in natural systems. In a series of laboratory experiments, we studied the impact of an oomycete brood parasite on its Daphnia (waterflea) host. We asked whether Daphnia dump the infected brood and subsequently are able to reproduce again as was occasionally observed in a preliminary study. No viable offspring developed from infected clutches, but 78% of the infected females produced healthy offspring after releasing the infected brood while molting. Neither those offsprings’ development success nor their mothers’ reproductive potential was affected by the brood parasite. However, infected Daphnia had a reduced life-span and suffered an increased susceptibility to another parasite, an unidentified bacterium. Additionally, we studied the prevalence of this brood parasite and the unidentified bacterium in a natural Daphnia assemblage in a pre-alpine lake, across changing demographic and environmental conditions. The brood parasite epidemic seemed to be host-density dependent. Our results show that the brood parasite’s impact on the host population is enhanced when combined with the unidentified bacterium.     

Hidden diversity in the freshwater planktonic diatom Asterionella formosa

Many freshwater and marine algal species are described as having cosmopolitan distributions. Whether these widely distributed morphologically similar algae also share a similar gene pool remains often unclear. In the context of island biogeography theory, stronger spatial isolation deemed typical of freshwater lakes should restrict gene flow and lead to higher genetic differentiation among lakes. Using nine microsatellite loci, we investigate the genetic diversity of a widely distributed freshwater planktonic diatom, Asterionella formosa, across different lakes in Switzerland and the Netherlands. We applied a hierarchical spatial sampling design to determine the geographical scale at which populations are structured. A subset of the isolates was additionally analysed using amplified fragment length polymorphism (AFLP) markers. Our results revealed complex and unexpected population structure in A. formosa with evidence for both restricted and moderate to high gene flow at the same time. Different genetic markers (microsatellites and AFLPs) analysed with a variety of multivariate methods consistently revealed that genetic differentiation within lakes was much stronger than among lakes, indicating the presence of cryptic species within A. formosa. We conclude that the hidden diversity found in this study is expected to have implications for the further use of A. formosa in biogeographical, conservation and ecological studies. Further research using species‐level phylogenetic markers is necessary to place the observed differentiation in an evolutionary context of speciation.     

Hybridization in the Daphnia galeata complex: are hybrids locally produced?

Within the species complex of Daphnia galeata,D. cucullata and D. hyalina variouscombinations of hybrids and parental taxa occur inlakes throughout Europe. Since daphnids are cyclicparthenogens and mostly reproduce asexually, hybridpopulations can be maintained by asexual reproductionand without recurrent hybridization events. Therefore,it is possible that hybridization events have beenrare, with range expansion occurring by dispersal ofhybrids.Allozyme data from seven European populations wereused to compare genetic variation within and betweenhybrid and parental taxa. An UPGMA cluster analysis ofgenetic distances showed that D. cucullata × galeatahybrids from different lakes grouped indifferent clusters according to the lake from whichthey were isolated, suggesting multiple hybridizationevents. Clonal diversity within hybrid taxa wascomparable to parental taxa. Furthermore, evidence wasfound for introgression of the Pgi-S allele fromD. cucullata to D. galeata in three lakes.These results indicate that multiple hybridizationevents within this species complex are likely, andthat hybrid taxa can reproduce sexually.     

Sexual reproduction in Daphnia: interspecific differences in a hybrid species complex

To investigate whether reproductive variation in the Daphnia galeata-cucullata-hyalina hybrid species complex can explain the occurrence of hybrids and backcrosses, a total of 43 clones from three north German lakes were tested for allocation to sexual reproduction under equal stress conditions. Six replicates per clone were followed until the seventh adult instar. I used the following cues to promote sexual reproduction: short photoperiod, water from a crowded Daphnia culture, fish-conditioned water and low food concentration. For each animal, clutch size and clutch sex were recorded. Ephippia, which were empty since the animals were cultured individually, were only produced by D. cucullata and D. cucullata x hyalina (26% and 6% of the broods, respectively), whereas almost all taxa produced males (the range was 2–15%). Intraspecific variation for male and ephippial production was also found. The fact that the taxa tested show different responses to the stimuli of-fered may indicate that there is reproductive isolation between them. This is supported by field data from the Tjeukemeer (The Netherlands) which shows that D. galeata mostly produces ephippia in spring and D. cucullata in autumn. The presence of hybrids with ephippia in both seasons, however, shows that backcrossing is also possible.