Diet specialization and brood parasitism in cuckoo species

1. Brood parasitism is a breeding strategy adopted by many species of cuckoos across the world. This breeding strategy influences the evolution of life histories of brood parasite species.
2. In this study, we tested whether the degree on diet specialization is related to the breeding strategy in cuckoo species, by comparing brood parasite and nonparasite species. We measured the gradient of diet specialization of cuckoos, by calculating the Gini coefficient, an index of inequality, on the multiple traits describing the diet of species. The Gini coefficient is a measure of statistical dispersion on a scale between 0 and 1, reflecting a gradient from low to high specialization, respectively. First, we tested the strength of the phylogenetic signal of diet specialization index among cuckoo species worldwide. Then, we ran phylogenetic generalized least square (PGLS) models to compare diet specialization, distribution range, and body mass of parasitic and nonparasitic cuckoo species, considering the phylogenetic signal of data.
3. After adjusting for the phylogenetic signal of the data and considering both, species distribution range and species body mass, brood parasitic cuckoos were characterized by higher diet specialization than nonbrood parasitic species. Brood parasitic species were also characterized by a larger breeding distribution range than nonparasitic species.
4. The findings of this study provide an additional understanding of the cuckoos’ ecology, relating diet and breeding strategies, information that could be important in conservation ecology.

     

Resource predictability and specialization in avian malaria parasites

We tested the hypothesis that avian haemosporidian (malaria) parasites specialize on hosts that can be characterized as predictable resources at a site in Amazonian Ecuador. We incorporated host phylogenetic relationship and relative abundance in assessing parasite specialization, and we examined associations between parasite specialization and three host characteristics – abundance, mass and longevity – using quantile regression, phylogenetic logistic regression and t‐tests. Hosts of specialist malaria parasite lineages were on average more abundant than hosts of generalist parasite lineages, but the relationship between host abundance and parasite specialization was not consistent across analyses. We also found support for a positive association between parasite specialization and host longevity, but this also was not consistent across analyses. Nonetheless, our findings suggest that the predictability of a host resource may play a role in the evolution of specialization. However, we also discuss two alternative explanations to the resource predictability hypothesis for specialization: (i) that interspecific interactions among the parasites themselves might constrain some parasites to a specialist strategy, and (ii) that frequent encounters with multiple host species, mediated by blood‐sucking insects, might promote generalization within this system.     

Host and habitat specialization of avian malaria in Africa

Studies of both vertebrates and invertebrates have suggested that specialists, as compared to generalists, are likely to suffer more serious declines in response to environmental change. Less is known about the effects of environmental conditions on specialist versus generalist parasites. Here, we study the evolutionary strategies of malaria parasites (Plasmodium spp.) among different bird host communities. We determined the parasite diversity and prevalence of avian malaria in three bird communities in the lowland forests in Cameroon, highland forests in East Africa and fynbos in South Africa. We calculated the host specificity index of parasites to examine the range of hosts parasitized as a function of the habitat and investigated the phylogenetic relationships of parasites. First, using phylogenetic and ancestral reconstruction analyses, we found an evolutionary tendency for generalist malaria parasites to become specialists. The transition rate at which generalists become specialists was nearly four times as great as the rate at which specialists become generalists. We also found more specialist parasites and greater parasite diversity in African lowland rainforests as compared to the more climatically variable habitats of the fynbos and the highland forests. Thus, with environmental changes, we anticipate a change in the distribution of both specialist and generalist parasites with potential impacts on bird communities.     

Seasonality,species richness and poor dispersion mediate intraspecific trait variability in stonefly community responses along an elevational gradient

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Shifts in species richness, herbivore specialization, and plant resistance along elevation gradients

Environmental gradients have been postulated to generate patterns of diversity and diet specialization, in which more stable environments, such as tropical regions, should promote higher diversity and specialization. Using field sampling and phylogenetic analyses of butterfly fauna over an entire alpine region, we show that butterfly specialization (measured as the mean phylogenetic distance between utilized host plants) decreases at higher elevations, alongside a decreasing gradient of plant diversity. Consistent with current hypotheses on the relationship between biodiversity and the strength of species interactions, we experimentally show that a higher level of generalization at high elevations is associated with lower levels of plant resistance: across 16 pairs of plant species, low-elevation plants were more resistant vis-à-vis their congeneric alpine relatives. Thus, the links between diversity, herbivore diet specialization, and plant resistance along an elevation gradient suggest a causal relationship analogous to that hypothesized along latitudinal gradients.     

BEMOVI,software for extracting behavior and morphology from videos,illustrated with analyses of microbes

Microbes are critical components of ecosystems and provide vital services (e.g., photosynthesis, decomposition, nutrient recycling). From the diverse roles microbes play in natural ecosystems, high levels of functional diversity result. Quantifying this diversity is challenging, because it is weakly associated with morphological differentiation. In addition, the small size of microbes hinders morphological and behavioral measurements at the individual level, as well as interactions between individuals. Advances in microbial community genetics and genomics, flow cytometry and digital analysis of still images are promising approaches. They miss out, however, on a very important aspect of populations and communities: the behavior of individuals. Video analysis complements these methods by providing in addition to abundance and trait measurements, detailed behavioral information, capturing dynamic processes such as movement, and hence has the potential to describe the interactions between individuals. We introduce BEMOVI, a package using the R and ImageJ software, to extract abundance, morphology, and movement data for tens to thousands of individuals in a video. Through a set of functions BEMOVI identifies individuals present in a video, reconstructs their movement trajectories through space and time, and merges this information into a single database. BEMOVI is a modular set of functions, which can be customized to allow for peculiarities of the videos to be analyzed, in terms of organisms features (e.g., morphology or movement) and how they can be distinguished from the background. We illustrate the validity and accuracy of the method with an example on experimental multispecies communities of aquatic protists. We show high correspondence between manual and automatic counts and illustrate how simultaneous time series of abundance, morphology, and behavior are obtained from BEMOVI. We further demonstrate how the trait data can be used with machine learning to automatically classify individuals into species and that information on movement behavior improves the predictive ability.     

Individual specialization in a shorebird population with narrow foraging niche

Individual specialization in resource use is a widespread driver for intra-population trait variation, playing a crucial evolutionary role in free-living animals. We investigated the individual foraging specialization of Black-tailed Godwits (Limosa limosa islandica) during the wintering period. Godwits displayed distinct degrees of individual specialization in diet and microhabitat use, indicating the presence of both generalist and specialist birds. Females were overall more specialist than males, primarily consuming polychaetes. Specialist males consumed mainly bivalves, but some individuals also specialized on gastropods or polychaetes. Sexual dimorphism in bill length is probably important in determining the differences in specialization, as longer-billed individuals have access to deep-buried polychaetes inaccessible to most males. Different levels of specialization within the same sex, unrelated to bill length, were also found, suggesting that mechanisms other traits are involved in explaining individual specialization. Godwits specialized on bivalves achieved higher intake rates than non-specialist birds, supporting the idea that individual foraging choices or skills result in different short-term payoffs within the same population. Understanding whether short-term payoffs are good indicators of long-term fitness and how selection operates to favour the prevalence of specialist or generalist godwits is a major future challenge.     

The ecology of differences: assessing community assembly with trait and evolutionary distances

Species enter and persist in local communities because of their ecological fit to local conditions, and recently, ecologists have moved from measuring diversity as species richness and evenness, to using measures that reflect species ecological differences. There are two principal approaches for quantifying species ecological differences: functional (trait‐based) and phylogenetic pairwise distances between species. Both approaches have produced new ecological insights, yet at the same time methodological issues and assumptions limit them. Traits and phylogeny may provide different, and perhaps complementary, information about species' differences. To adequately test assembly hypotheses, a framework integrating the information provided by traits and phylogenies is required. We propose an intuitive measure for combining functional and phylogenetic pairwise distances, which provides a useful way to assess how functional and phylogenetic distances contribute to understanding patterns of community assembly. Here, we show that both traits and phylogeny inform community assembly patterns in alpine plant communities across an elevation gradient, because they represent complementary information. Differences in historical selection pressures have produced variation in the strength of the trait‐phylogeny correlation, and as such, integrating traits and phylogeny can enhance the ability to detect assembly patterns across habitats or environmental gradients.     

Magadi tilapia ecological specialization: filling the early gap in the speciation continuum

Cichlid fish are well known for their high speciation rates, which are usually accompanied by spectacular and rapid diversification in eco‐morphological and secondary sexual traits. This is best illustrated by the famous repeated explosive radiations in the African Great Lakes Tanganyika, Malawi and Victoria, each lake harbouring several hundreds of mostly endemic species. Correspondingly, cichlids diversified very rapidly in many other lakes across their range. Although the larger radiations, unparalleled in vertebrates, are certainly the most intriguing, they are also the most intricate and difficult to address because of their complex nature. This is where smaller, simpler systems may prove to be the most useful. In this issue of Molecular Ecology, Kavembe et al. ( 2016 ) report very recent genetic diversification accompanied by ecological specialization in cichlids of the small and ecologically extreme Lake Magadi, in Kenya. Combining geometric morphometrics, stable isotope analysis, population genomics using RADSeq data and coalescent‐based modelling techniques, the authors characterize the eco‐morphological differences between genetically distinct populations of Magadi tilapia (Alcolapia grahami), which are consistent with the different environmental conditions they experience, and infer their history of divergence. The simplicity of the focal system and the use of a multidisciplinary approach make this work particularly important for our understanding of the early stages of speciation, in both cichlids and other organisms.     

Phylogeny, specialization, and brood parasite--host coevolution: some possible pitfalls of parsimony

Coevolutionary hypotheses (COEV) predict that parasitic birdsbecome more specialized in host selection over time as morehost species evolve defenses. A contrasting model, PHYLO, suggeststhat brood parasites exhibit a phylogenetic trajectory towardincreasing generalization because there is a positive correlationbetween present-day numbers of host species and the branchingorder of parasitic cowbird species in a DNA-based phylogeny. However, this apparent phylogenetic pattern does not conflictwith COEV, as some have concluded. Assuming allopatric speciation,which is supported by an area cladogram, COEV predicts a correlationbetween branching order and host number because the potentialhosts of the earliest cowbirds to branch off have had the greatestamount of time to evolve defenses. Although PHYLO is more parsimoniousthan COEV, the difference is trivial, with the latter requiring only one more evolutionary change in the entire cowbird cladeto produce the pattern that exists today. Support for COEVover PHYLO comes from brood parasitic cuckoos, which are muchmore specialized than parasitic cowbirds and represent an olderclade, as shown by new DNA data. Cuckoos also have lower interspecificvariance in host numbers than do cowbirds, which conflicts with PHYLO. Unlike COEV, which assumes that the number of hosts aparasite uses is related at least as much to present ecologicalconditions as to phylogenetic history, PHYLO assumes that currenthost numbers reflect historical character states. However,host number is labile, with as much variation within as betweenspecies. Nor are published host numbers reliable measures ofparasite host selectivity, as they are due in part to researchereffort and range size. Although the comparative approach canprovide insights into evolutionary history, some coevolvedfeatures may be too dynamic to retain a phylogenetic signature,and, in the case of parasitic birds, neither PHYLO nor COEVcan be invalidated, although the latter is more consistentwith available evidence. Strict adherence to parsimony mayoften be inappropriate when assessing coevolved characters.     

A global meta‐analysis of the relative extent of intraspecific trait variation in plant communities

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta‐analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole‐plant (e.g. plant height) vs. organ‐level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait‐based community and ecosystem studies.     

Eco-evolutionary drivers of avian migratory connectivity

Migratory connectivity, reflecting the extent by which migrants tend to maintain their reciprocal positions in seasonal ranges, can assist in the conservation and management of mobile species, yet relevant drivers remain unclear. Taking advantage of an exceptionally large (~150,000 individuals, 83 species) and more-than-a-century-long dataset of bird ringing encounters, we investigated eco-evolutionary drivers of migratory connectivity in both short- and long-distance Afro-Palearctic migratory birds. Connectivity was strongly associated with geographical proxies of migration costs and was weakly influenced by biological traits and phylogeny, suggesting the evolutionary lability of migratory behaviour. The large intraspecific variability in avian migration strategies, through which most species geographically split into distinct migratory populations, explained why most of them were significantly connected. By unravelling key determinants of migratory connectivity, our study improves knowledge about the resilience of avian migrants to ecological perturbations, providing a critical tool to inform transboundary conservation and management strategies at the population level.     

Gender specialization and distyly in hoary puccoon,LITHOSPERMUM CROCEUM (Boraginaceae)

This 2-yr study of gender and its relation to the sizes of pin and thrum plants in Lithospermum croceum was performed in central Michigan. The population was composed of 51-52% thrums, 42-43% pins, and ≈6% homostyles in 1988 and 1989. Pins produced more flowers and fruits than thrums, and a greater proportion of their ovaries produced more than one nutlet. However, the number of fruits produced by pins and thrums in 1988 did not differ when plant size is a covariate in ANCOVA. The number of seeds produced by plants in 1989 is also unaffected by floral morph when size is a covariate. Instead, plant size is the most important predictor of fecundity in both years. The average percent fruit sets (fruits/ovaries) of the morphs did not differ in either year. Herbivory was more common in 1988 than in 1989, but there was no difference in the frequency with which pins and thrums were attacked. Furthermore, the effect of herbivory on fecundity did not differ for the morphs. The fecundity of neither morph was limited by pollen in 1989. Apparent gender specialization was a consequence of subtle differences in the vegetative sizes of the morphs.     

Transition rates between specialization and generalization in phytophagous insects

Although most species of animals exhibit specialized patterns of resource use, it is unclear whether specialization evolves at a faster rate than generalization. To test this hypothesis, transition rates toward specialization and toward generalization were estimated using phylogenies from 15 groups of phytophagous insects. Among the groups studied, maximum-likelihood analyses showed that the forward transition rate from generalization to specialization was significantly higher than the reverse transition rate from specialization to generalization (mean ratio of forward to reverse transition rate = 1.47 using uniform branch lengths and 1.76 using Grafen branch lengths). Although phylogenetic conservatism of host-plant use is common, the results suggest that the evolution of specialization is a highly dynamic process. For example, higher transitions rates both toward and away from specialization as well as equal transition rates were inferred. Collectively, the results reveal a tendency for directional evolution toward increased specialization but also indicate that specialization does not always represent an evolutionary dead-end that strongly limits further evolution.     

Scavenging affects persistence of avian carcasses resulting from window collisions in an urban landscape

ABSTRACT Collisions with windows remain an important human‐related threat to bird survival in urban landscapes. Accurately estimating the magnitude of avian mortality at windows is difficult and may be influenced by many sources of error, such as scavenging of carcasses. Failure to account for removal of carcasses by scavengers can bias estimates of window mortality. We tested the hypothesis that carcass survival depends on local habitat factors known to influence scavenger behavior. Scavenger activity on bird carcasses was documented at 20 buildings in an urban landscape in northwestern Illinois for 1 week during each season of a year. Known‐fate models were used to relate carcass survival to local habitat composition and to evaluate temporal variation in survival. We also documented species of scavengers and the timing of scavenging using motion‐triggered cameras. Daily carcass survival was greater in winter than during spring, summer, and fall. Survival was related negatively to canopy cover (trees and shrubs within a 50‐m buffer) and window area, and positively to pavement cover. Using an exponential model of survival time, estimated mean time of survival of carcasses (t± SE) was 82.9 ± 11.7 d for winter and 11.8 ± 7.2 d for other seasons. Raccoons (Procyon lotor) scavenged more carcasses than other species. Our results suggest that (1) carcass survival times may be short at locations with preferred habitats of known scavengers and predictable sources of food, and (2) knowledge of scavenger distribution and activity can inform predictive models of persistence. In studies of bird‐window collisions, the influence of scavenger bias can be minimized by maintaining short time intervals between carcass searches. Search intervals can be inferred by estimating the number of days that a carcass should persist at a site, which can be calculated using predicted daily survival probabilities of carcasses at study buildings.     

Effect of environmental and spatial factors on the phylogenetic and functional diversity of the Mediterranean tree communities of Europe

• The tree flora of the Mediterranean Basin contains an outstanding taxonomic richness and a high proportion of endemic taxa. Contrary to other regions of the Mediterranean biome, a comprehensive phylogenetic analysis of the relationship between phylogenetic diversity, trait diversity and environmental factors in a spatial ecological context is lacking.
• We inferred the first calibrated phylogeny of 203 native tree species occurring in the European Mediterranean Basin based on 12 DNA regions. Using a set of four functional traits, we computed phylogenetic diversity for all 10,042 grid cells of 10 × 10 km spatial resolution to completely cover Mediterranean Europe. Then, we tested the spatial influence of environmental factors on tree diversity.
• Our results suggest that the nature of the relationship between traits and phylogeny varies among the different studied traits and according to the evolutionary distance considered. Phylogenetic diversity and functional diversity of European Mediterranean trees correlated strongly with species richness. High values of these diversity indices were located in the north of the study area, at high altitude, and minimum temperature of the coldest month. In contrast, the two phylogenetic indices that were not correlated with species richness (Mean Phylogenetic Distance, Phylogenetic Species Variability) were located in the south of the study area and were positively correlated with high altitude, soil organic carbon stock and sand soil texture.
• Our study provides support for the use of phylogenies in conservation biology to assess ecosystem functioning, and provides insights for the implementation of sustainable forest ecosystem management.