Experimentally induced helper dispersal in colonially breeding cooperative cichlids

The ‘benefits of philopatry’ hypothesis states that helpers in cooperatively breeding species derive higher benefits from remaining home, instead of dispersing and attempting to breed independently. We tested experimentally whether dispersal options influence dispersal propensity in the cooperatively breeding Lake Tanganyika cichlids Neolamprologus pulcher and N. savoryi. Cooperative groups of these fishes breed in densely packed colonies, surrounded by unoccupied, but apparently suitable breeding habitat. Breeding inside colonies and living in groups seems to benefit individuals, for example by early detection and deterrence of predators. We show that despite a slight preference of both species for habitat with a higher stone cover, 40% of the preferred habitat remained unoccupied. On average, the colonies contained a higher number of (1) predators of adults, juveniles and eggs, (2) shelter competitors, and (3) other species including potential food competitors, compared to the outside colony habitat. Apparently, habitat differences cannot explain why these cichlids breed in colonies. Accordingly, dispersal may not be limited by a lack of suitable breeding shelters, but by the relatively higher risk of establishing an outside- compared to a within-colony breeding territory. To test whether cichlids prefer within- to outside-colony breeding territories, we provided breeding shelters inside the colony and at the colony edge and studied helper dispersal. As expected, significantly more shelters were occupied within the colony compared to the edge. New breeding pairs with several helpers occupied these shelters. We conclude that although breeding habitat is plentiful outside the colonies, helpers delay dispersal to obtain a higher quality breeding position within the group or colony eventually, or they disperse in groups. Our results suggest that (1) group augmentation and Allee effects are generally important for dispersal decisions in cooperatively breeding cichlids, consistent with the ‘benefits of philopatry hypothesis’, and (2) habitat saturation cannot fully explain delayed dispersal in these species.     

Morphological and behavioral antipredatory adaptations of decapod zoeae

Summary Zoeae of some species of estuarine decapods are retained in the estuary throughout development while others are exported into nearshore coastal waters. The horizontal migrations of decapod zoeae to coastal waters may have evolved to reduce the probability of encountering planktivorous fishes which are most abundant in the estuary. If so, then the morphological vulnerability of zoeae to fish predation should be inversely related to the number of predators occurring where they develop. Six species of estuarine decapod zoeae were offered to Menidia menidia and Fundulus heteroclitus. The behavioral interactions were observed to determine the prey's vulnerability to predation, and the mode of operation and relative effectiveness of their defenses. Feeding trials and behavioral observations both demonstrated that M. menidia 6–16 mm long preferred Uca minax and Callinectes sapidus zoeae, which are exported from the estuary, to Rhithropanopeus harrisii, Sesarma reticulatum and Palaemonetes pugio, which are retained within estuaries. Pinnotheres ostreum zoeae develop in the lower estuary and fish demonstrated an intermediate preference for the zoeae. Menidia menidia 20–40 mm long showed similar preferences for R. harrisii S. reticulatum, P. ostreum and U. minax as did small silversides. Large-mouthed demersal fish, Fundulus heteroclitus 6–10 mm long, also preferred U. minax to R. harrisii, but more readily preyed on zoeae than did M. menidia. The exported species of zoeae have shorter spines and smaller bodies than do retained zoeae, except P. ostreum which is small, spineless and passively sinks when attacked by fish. Other retained species of zoeae also have postcontact behavioral defenses which enhance the effectiveness of their morphological defenses. Zoeae do not evade attacks by fishes, but fishes quickly learned to avoid zoeae, which increases the effectiveness of the zoeae's antipredatory adaptations.     

Trait plasticity alters the range of possible coexistence conditions in a competition–colonisation trade‐off

Most of the classical theory on species coexistence has been based on species‐level competitive trade‐offs. However, it is becoming apparent that plant species display high levels of trait plasticity. The implications of this plasticity are almost completely unknown for most coexistence theory. Here, we model a competition–colonisation trade‐off and incorporate trait plasticity to evaluate its effects on coexistence. Our simulations show that the classic competition–colonisation trade‐off is highly sensitive to environmental circumstances, and coexistence only occurs in narrow ranges of conditions. The inclusion of plasticity, which allows shifts in competitive hierarchies across the landscape, leads to coexistence across a much broader range of competitive and environmental conditions including disturbance levels, the magnitude of competitive differences between species, and landscape spatial patterning. Plasticity also increases the number of species that persist in simulations of multispecies assemblages. Plasticity may generally increase the robustness of coexistence mechanisms and be an important component of scaling coexistence theory to higher diversity communities.     

Echolocation, olfaction and fruit display: how bats find fruit of flagellichorous cucurbits

1. The relationship between plant morphology and the senses used by dispersal agents to find fruit was examined. 'Flagellichory' (fruit borne on pendulous structures), a costly morphology associated with dispersal by bats, is focused on.
2. Using Gurania spinulosa , a flagellichorous vine, and its major dispersal agent, Phyllostomus hastatus , the hypothesis was tested that flagellichory increases the conspicuousness of fruit to bats that use echolocation to find fruit.
3. The responses of wild-caught P. hastatus to various fruiting branch morphologies and fruit odour were recorded. Phyllostomus hastatus used echolocation rather than olfaction to detect fruit, and consistently chose fruit displayed on pendulous leafless branches, ignoring fruit held among leaves on horizontal branches.
4. By comparing echolocation signals with the distance between fruiting branches of G. spinulosa and surrounding vegetation, it was shown that pendulous fruiting branches present clear, clutter-free targets that can be detected by echolocating bats. This is the first demonstration of neotropical frugivorous bats using echolocation to find fruit.     

ON THE EVOLUTION OF MIGRATION IN HETEROGENEOUS ENVIRONMENTS

Populations often experience variable conditions, both in time and space. Here we develop a novel theoretical framework to study the evolution of migration under the influence of spatially and temporally variable selection and genetic drift. First, we examine when polymorphism is maintained at a locus under heterogeneous selection, as a function of the pattern of spatial heterogeneity and the migration rate. In a second step, we study how levels of migration evolve under the joint action of kin competition and local adaptation at a polymorphic locus. This analysis reveals the existence of evolutionary bistability, whereby a low or a high migration rate may evolve depending on the initial conditions. Last, we relax several assumptions regarding selection heterogeneity commonly made in previous studies and explore the consequences of more complex spatial and temporal patterns of variability in selection on the evolution of migration. We found that small modifications in the pattern of environmental heterogeneity may have dramatic effects on the evolution of migration. This work highlights the importance of considering more general scenarios of environmental heterogeneity when studying the evolution of life‐history traits in ecologically complex settings.     

Linking dispersal, immigration and scale in the neutral theory of biodiversity

In the classic spatially implicit formulation of Hubbell's neutral theory of biodiversity a local community receives immigrants from a metacommunity operating on a relatively slow timescale, and dispersal into the local community is governed by an immigration parameter m . A current problem with neutral theory is that m lacks a clear biological interpretation. Here, we derive analytical expressions that relate the immigration parameter m to the geometry of the plot defining the local community and the parameters of a dispersal kernel. Our results facilitate more rigorous and extensive tests of the neutral theory: we conduct a test of neutral theory by comparing estimates of m derived from fits to empirical species abundance distributions to those derived from dispersal kernels and find acceptable correspondence; and we generate a new prediction of neutral theory by investigating how the shapes of species abundance distributions change theoretically as the spatial scale of observation changes. We also discuss how our main analytical results can be used to assess the error in the mean-field approximations associated with spatially implicit formulations of neutral theory.     

Comparison of three seed trap types in a chalk grassland: toward a standardised protocol

In a chalk grassland in north-western France, vegetation and seed rain were studied along two transects. We compared the efficiency of three types of seed traps (funnel traps, sticky traps, pots exposed in the field) to estimate the annual seed rain and to reveal various processes involved in plant community regeneration. Data were analysed to compare seed density, species richness and composition across seed traps and vegetation. Geostatistical analyses (semivariograms. were used to detect possible autocorrelation and to examine patterns of spatial variation in seed rain and vegetation. The results show that (1) funnel traps are most efficient (56 species, 8079 seeds.m^–2). (2) Each type of seed traps brings different information about the processes involved in seed dispersal and seedling establishment. Sticky traps help to understand import and export of seeds in a community through anemochory. Funnel traps may provide a measure of the auto-regeneration capacity of the vegetation through its local seed production. Pot traps integrate various processes involved between seed dispersal and seedling emergence. (3.) The exposition to environmental conditions generates marked spatial patterns in species richness measured in pots and in vegetation, selecting species able to survive. Pot traps indicate that numerous species producing seeds in standing vegetation are unable to establish in the field through seedling recruitment. Our study emphasizes the need of using different types of seed traps and spatial sampling configurations to assess the regenerative potential of standing vegetation and the various mechanisms involved in seed dispersal. A standardised protocol for seed rain sampling was finally developed.     

Vegetation and seed bank in a calcareous grassland restored from a Pinus forest

Question: What is the importance of the seed bank in the maintenance of the restoration potential of a 60‐year‐old abandoned calcareous grassland overgrown by Pinus trees? Location: ‘Les Pairées’, province of Luxembourg, Belgium. Methods: The seed bank and the above‐ground vegetation were surveyed in three adjacent stands, previously forming a unique calcareous grassland: a 60‐year‐old Pinus forest, a four‐year‐old clear‐cutting and a typical calcareous grassland. Floristic diversity was compared among stands and between vegetation and seed bank. Results: The species richness of the vegetation and the seed bank was significantly lower in the Pinus forest. More floristic similarities were found between the clear‐cutting and the calcareous grassland. Seed bank was essentially transient, dominated by annual species. Its correspondence with the above‐ground vegetation was weak. Conclusion: Very few calcareous grassland species have persisted in the Pinus stand. Four years after clear‐cutting, the stand was nearing restoration towards a calcareous grassland. Seed longevity in the soil was not the most explicative factor. Dispersal of propagules from adjacent sources was also important.