Caffeine as a novel toxicant for slugs and snails

In this study, caffeine is shown to act as both a repellent and toxicant against slugs and snails. This research is the first to document the potential of caffeine as a molluscicide. A drench treatment using a 1% or 2% solution of caffeine caused 100% of slugs (Veronicella cubensis) to exit treated soil, and the majority of these slugs subsequently died from caffeine poisoning. A 2% solution of caffeine applied to the growing medium of orchids killed 95% of orchid snails, (Zonitoides arboreus), and gave better control than a liquid metaldehyde product representing the standard commercial control for this pest. Using leaf‐dip bioassays, we discovered that slugs tended to avoid feeding on plant material treated with caffeine solutions 0.1%, and caffeine solutions as low as 0.01% significantly reduced overall feeding by slugs. Due to concerns about chemical residues, available molluscicides generally cannot be applied directly to food crops for control of slug and snail pests. Caffeine is a natural product which is approved as a food additive. Therefore, caffeine may prove useful for protecting food crops from slugs and snails.     

Microsatellites for the slugs Deroceras reticulatum and Arion intermedius

Microsatellites were developed for two pest slug species: self‐fertilizing Arion intermedius and outcrossing Deroceras reticulatum. Six microsatellites from each species were tested on individuals collected from within and between arable fields. The 39 individuals analysed per species had 3–9 and 7–26 alleles per microsatellite for A. intermedius and D. reticulatum, respectively. Gene diversity, [1 ? Σp_i²] , ranged from 0.19 to 0.75 and 0.74 to 0.95, respectively.     

Behavioural avoidance by slugs and snails of the parasitic nematode Phasmarhabditis hermaphrodita

The nematode Phasmarhabditis hermaphrodita has been developed as a biological control agent for slugs and snails. Slugs avoid areas where P. hermaphrodita is present. We investigated whether behavioural avoidance of P. hermaphrodita is a common feature of slugs and snails by exposing eight species to P. hermaphrodita. We showed that slugs generally avoided P. hermaphrodita, whereas snails did not. We also showed that slugs specifically avoided the commercial strain and a natural isolate of P. hermaphrodita and were not deterred by other nematodes such as Steinernema kraussei or Turbatrix aceti. We also showed that slugs avoided the dauer stage of P. hermaphrodita and not mixed-stage cultures. Furthermore, slugs do not avoid dead P. hermaphrodita or exudates from live nematodes. Taken together, we have unravelled further factors that are essential for slugs to avoid P. hermaphrodita in soil, which could have important implications for the biological control of slugs and snails.     

Invasive slugs as under-appreciated obstacles to rare plant restoration: evidence from the Hawaiian Islands

Introduced slugs have invaded many parts of the world where they were recognized as important pests of gardens and agriculture, but we know little about the effects of introduced slugs on rare plants in natural areas. The Hawaiian Islands have no native slugs, but over a dozen introduced slug species are now established. We reviewed Rare Plant Recovery Plans produced by the U.S. Fish and Wildlife Service for Hawaii and found that introduced slugs were specifically mentioned as threats or potential threats to 59 rare plant species (22% of all endangered and threatened plants), based mainly on anecdotal observations by field biologists. We then initiated an experimental field study to assess the impact of slug herbivory on the growth and survival of two endangered plant species (Cyanea superba, and Schidea obovata), one non-endangered native species (Nestegis sandwicensis) and two co-occurring invasive plant species (Psidium cattleianum and Clidemia hirta). In mesic forest on the Island of Oahu, we tracked the fate of outplanted seedlings in replicated 1 m² plots, with and without slug control. Slugs decreased seedling survival of the endangered species by 51%, on average. Slugs did not significantly affect survival of the non-endangered or invasive plant species. Introduced slugs seem to be under-appreciated as a direct cause of plant endangerment. Invasive slugs may also facilitate the success of some invasive plant species by reducing competition with more palatable, native plant competitors. Slug control measures are relatively inexpensive and could facilitate rare plant establishment and population recovery.     

Plant growth forms: an ecological and evolutionary perspective

Trees, shrubs, lianas and herbs have widely different mechanical architectures, which can also vary phenotypically with the environment. This review investigates how environmental effects, particularly mechanical perturbation, can influence biomechanical development in self-supporting and climbing growth forms. The bifacial vascular cambium is discussed in terms of its significance to growth form variation, ecology and evolution among extant plants, and during its appearance and early evolution. A key aspect of this developmental innovation concerned its potential for architectural and mechanical variation in response to environmental effects as well as optimizing hydraulic supply before the appearance of laminate leaves. Growth form diversity and its importance to past and present ecosystems are discussed in relation to both evolutionary constraints and ecological factors such as climatic change and atmospheric CO2 concentrations. We discuss how widely ranging growth forms such as climbers show a large range of developmental and phenotypic variation that has much to offer in understanding how the environment can modify plant development, particularly in terms of the bifacial vascular cambium. The broad approach we propose would benefit a wide range of studies from research into wood development to long-term ecological censuses of today's potentially changing ecosystems.     

Plastid-bearing sea slugs fix CO2 in the light but do not require photosynthesis to survive

Several sacoglossan sea slugs (Plakobranchoidea) feed upon plastids of large unicellular algae. Four species—called long-term retention (LtR) species—are known to sequester ingested plastids within specialized cells of the digestive gland. There, the stolen plastids (kleptoplasts) remain photosynthetically active for several months, during which time LtR species can survive without additional food uptake. Kleptoplast longevity has long been puzzling, because the slugs do not sequester algal nuclei that could support photosystem maintenance. It is widely assumed that the slugs survive starvation by means of kleptoplast photosynthesis, yet direct evidence to support that view is lacking. We show that two LtR plakobranchids, Elysia timida and Plakobranchus ocellatus, incorporate ¹⁴CO₂ into acid-stable products 60- and 64-fold more rapidly in the light than in the dark, respectively. Despite this light-dependent CO₂ fixation ability, light is, surprisingly, not essential for the slugs to survive starvation. LtR animals survived several months of starvation (i) in complete darkness and (ii) in the light in the presence of the photosynthesis inhibitor monolinuron, all while not losing weight faster than the control animals. Contrary to current views, sacoglossan kleptoplasts seem to be slowly digested food reserves, not a source of solar power.     

Denitrification: ecological niches,competition and survival

Organisms with the denitrification capacity are widely distributed and in high density in nature. It is not well understood why they are so successful. A survey of denitrifying enzyme content of various habitats is presented which indicates a role of carbon and oxygen, but not nitrate, in affecting denitrifier populations. It is suggested that organic carbon is more important than oxygen status in determining denitrifying enzyme content of habitats. In low oxygen environments, denitrifiers compete with organisms that dissimilate nitrate to ammonium, a process which conserves nitrogen. The energetic and kinetic parameters that affect this competition are evaluated. The latter is examined using Michaelis-Menten theoretical models by varying V_max, K_m, and S_o (substrate concentration) for the two competing populations. The outcome predicted by these models is presented and discussed in relation to previous data on population densities and K_m values for representatives of these competing groups. These models suggest the conditions required to achieve changes in partitioning between the two fates of nitrate. These considerations are important if one is to be able to evaluate and successfully “manage” the fate of nitrate in any habitat.     

The diversification of Halenia (Gentianaceae): ecological opportunity versus key innovation

The plant genus Halenia (Gentianaceae) consists of herbs growing in temperate and tropical alpine habitats and most species possess flowers in which nectar is produced in spurs. This probably helps reward only specialized long-tongued pollinators, and a narrow pollinator/flower relationship is thought to accelerate diversification rates (a key innovation). To test the pattern of diversification of Halenia against the unspurred sister group we reconstructed phylogenetic relationships among 22 species plus outgroups using nuclear ITS and chloroplast rpl16 intron sequence data. We show that Halenia originated in East Asia and migrated via North America into Central America. From there, it colonized South America three times independently, probably within the last million years. Significant changes in diversification rates were found during the evolution of Halenia using a sister group method, a likelihood method, and a diversity-through-time plot. In contrast to other studies, we could not observe a direct speciation rate effect of the evolution of nectar spurs in comparison with the unspurred sister group of Halenia. Rather, increases in diversification occurred following the colonization of Central and South America by spurred progenitor taxa. This later switch in diversification may have resulted from the availability of new geographical and ecological opportunities, or from the availability of more and different pollinators in these regions. Following the latter hypothesis, the nectar spurs were a preadaption and functioned as a key innovation only in this new biotic environment. After an initial rapid increase, a reduction in diversification rate was observed in Central America, probably illustrating density dependence of speciation rates. Finally, we found preliminary evidence for the key innovation hypothesis in geologically young spurred and unspurred lineages of Halenia in South America.     

杠杆状雄蕊及其进化生态学意义

对被子植物一类特化雄蕊——杠杆状雄蕊的结构多样性及其进化生态学意义进行了归纳总结。植物的花在进化过程中, 常会发生雄蕊群的改变, 包括雄蕊数目及其形态结构的变化, 同时雄蕊功能也会发生相应的适应性转变。杠杆状雄蕊是指结构特化为杠杆状或距状, 在传粉过程中具有类似杠杆功能的一类特化的雄蕊类型。目前, 已在唇形科不同亚科以及姜科6个属中发现杠杆状雄蕊, 根据其结构和形态发生方式总体上可分为2大类: 一类是唇形科中以鼠尾草属(Salvia)为代表的, 由2个可育雄蕊平行发育、药隔组织增长所形成的杠杆状雄蕊; 另一类是姜科植物中由一个可育雄蕊特化形成的带有距状附属体的雄蕊类型。在生态功能上, 两类雄蕊均能通过传粉者推动其距状下臂做杠杆运动进行传粉, 被认为是一种促进异交的传粉机制, 可通过精确传粉和花粉分发等途径影响植物的繁殖成功。杠杆状雄蕊在不同的类群中是独立起源与进化的; 仅在唇形科鼠尾草属中, 杠杆状雄蕊发生了3次独立进化, 而且它可能是触发该属物种适应性辐射的关键性状。将来需在宏观进化和微观进化两个水平深入探讨杠杆状雄蕊的进化生态学意义。     

Interference competition between coyotes and raccoons: a test of the mesopredator release hypothesis

Some predator species appear to conform to the mesopredatorrelease hypothesis (MRH), in which larger predators help limitpopulations of smaller predators. This hypothesis has been usedto explain the possible relationship between coyotes, mesopredators,and resultant cascades involving nonpredators. However, relationshipsbetween coyotes and noncanid mesopredators are poorly understood,and predictions from the MRH have rarely been rigorously tested.We monitored sympatric raccoon and coyote populations to assess2 predictions derived from the MRH: coyote predation is an importantcause of mortality in raccoon populations or raccoons avoidareas used by coyotes. Between March 2000 and September 2001,we recorded 3553 locations for 27 radio-collared raccoons and1393 locations for 13 coyotes captured on the Max McGraw WildlifeFoundation in Illinois, USA. No raccoon mortality from coyotepredation was observed during the study, and raccoon survivalwas >0.7 each season. All raccoon 95% home ranges exhibitedoverlap with 95% coyote home ranges in each season. The meanproportion of raccoon locations within 95% coyote home rangesdid not vary by sex but did vary by season. Raccoon overlapof coyote core areas varied considerably among individuals withinseasons, ranging from 0% to 83%. However, 45% of raccoons had<10% overlap with coyote core areas, whereas only 14% ofraccoons exhibited >50% overlap. Mean overlap with core areasdid not vary by season or sex. For those raccoons with homeranges overlapping coyote core areas, mean proportion of observedraccoon locations within coyote core areas was generally greaterthan the mean proportion of random locations. Scent-stationexperiments failed to document raccoon avoidance of specificsites that had been marked with coyote urine. We did not findsupport for a mortality prediction or avoidance prediction tosupport MRH with regard to raccoons and coyotes. These resultssuggest that relationships among mammalian predators may notbe simply dictated by body size, particularly for species outsidethe Canidae.     

Size patterns among competitors: ecological character displacement and character release in mammals, with special reference to island populations

Morphological relationship among sympatric animal species have often been seen as indirect evidence for competition. Many early ecomorphological studies revealed patterns that were taken as indicating character displacement and character release, driven by competition or lack thereof. These patterns may result from a coevolutionary morphological response or from species sorting according to size. Thus, the relationship between morphology and competition may be crucial for understanding both the morphological evolution of animals and the role of competition in structuring communities. Some earlier research perceived as indicating morphological relationships conditioned by interaction of species was conducted on mammals, particularly carnivores. Subsequent criticism in the ecological literature demonstrated that many of the perceived patterns could not be statistically confirmed, thus calling into question this line of evidence for competition. More recent ecological literature relies on strong statistical analyses and careful consideration both of guild composition and of which morphological traits should be examined. This literature, resting largely on mammals, includes several cases that suggest a coevolutionary morphological response to interspecific competition. These studies have focused on the thropic apparatus directly related to food procurement by mammals — the teeth. Island mammals often show striking morphological patterns, some of which have been interpreted as resulting from release from competition with mainland species that have not reached islands. However, few of these patterns were critically evaluated to demonstrate their support for the hypothesis of character release. Despite several decades of interest and research, many questions regarding competitively induced morphological patterns remain unresolved and require further research. Mammals are especially promising subjects for such researh.     

Positive feedback between ecological and reproductive character displacement in a young avian hybrid zone

Character displacement can reduce costly interspecific interactions between young species. We investigated the mechanisms behind divergence in three key traits-breeding habitat choice, timing of breeding, and plumage coloration-in Ficedula flycatchers. We found that male pied flycatchers became expelled from the preferred deciduous habitat into mixed forest as the superior competitor, collared flycatchers, increased in numbers. The peak in food abundance differs between habitats, and the spatial segregation was paralleled by an increased divergence in timing of breeding between the two species. Male pied flycatchers vary from brown to black with brown coloration being more frequent in sympatry with collared flycatchers, a pattern often proposed to result from selection against hybridization, that is, reinforcement. In contrast to this view, we show that brown male pied flycatchers more often hybridize than black males. Male pied flycatcher plumage coloration influenced the territory obtained in areas of co-occurrence with collared flycatchers, and brown male pied flycatchers experienced higher relative fitness than black males when faced with heterospecific competition. We suggest that allopatric divergence in resource defense ability causes a feedback loop at secondary contact where male pied flycatchers with the most divergent strategy compared to collared flycatchers are favored by selection.     

Character displacement as the "best of a bad situation": fitness trade-offs resulting from selection to minimize resource and mate competition

Abstract Character displacement has long been considered a major cause of adaptive diversification. When species compete for resources or mates, character displacement minimizes competition by promoting divergence in phenotypes associated with resource use (ecological character displacement) or mate attraction (reproductive character displacement). In this study, we investigated whether character displacement can also have pleiotropic effects that lead to fitness trade-offs between the benefits of avoiding competition and costs accrued in other fitness components. We show that both reproductive and ecological character displacement have caused spadefoot toads to evolve smaller body size in the presence of a heterospecific competitor. Although this shift in size likely arose as a by-product of character displacement acting to promote divergence between species in mating behavior and larval development, it concomitantly reduces offspring survival, female fecundity, and sexual selection on males. Thus, character displacement may represent the "best of a bad situation" in that it lessens competition, but at a cost. Individuals in sympatry with the displaced phenotype will have higher fitness than those without the displaced trait because they experience reduced competition, but they may have reduced fitness relative to individuals in allopatry. Such a fitness trade-off can limit the conditions under which character displacement evolves and may even increase the risk of "Darwinian extinction" in sympatric populations. Consequently, character displacement may not always promote diversification in the manner that is often expected.     

Consequences of competition on the reproduction and mortality of three species of terrestrial slugs

The interaction of three species of terrestrial slugs was studied in six field cages containing monocultures and 2-way combinations. A. columbianus and A. ater were non-aggressive species and L. maximus was highly aggressive during the summer. All three species were similar in body design and resource utilization. Reproduction (egg size, eggs/batch, number of batches, wet-weight productivity) and mortality were monitored through the 1977 season. L. maximus drastically reduced reproductive success of the non-aggressive slugs and increased their mortality. There were also significant interactions between the non-aggressive species. Compared to other invertebrates, the life history characteristics of A. columbianus are strongly K-selected and probably evolved through intraspecific exploitative competition in predictable, stable forest habitats. A. ater is best classified as a refuging r-selected species adapted to less predictable, unstable field environments. L. maximus appears to be α-selected and its overall design is suited to density-dependent regulation in fields.     

Stability of displacement clines arising in allospecies competition

Summary A stability analysis for geographic displacement clines between competing allospecies is presented. The competition model incorporates the effects of annual dispersal and of selective recruitment determined by geographically varying conditions at the breeding sites. It is assumed that a species gains a local competitive advantage wherever it attains sufficient numerical predominance. This assumption is valid when the species crosspair but the hybrids produced are not recruited into the adult population, because the minority species loses proportionately more of its reproductive potential. It is shown that no stable equilibria occur when the competitive balance is independent of location, but that even a slight geographic variation in the competitive balance allows stable equilibria. The greater the length of the combined breeding ranges of the two species, the smaller the geographic shift in competitive balance needed to produce stability.     

Parallel evolution and ecological selection: replicated character displacement in spadefoot toads

Ecological character displacement—trait evolution stemming from selection to lessen resource competition between species—is most often inferred from a pattern in which species differ in resource-use traits in sympatry but not in allopatry, and in which sympatric populations within each species differ from conspecific allopatric populations. Yet, without information on population history, the presence of a divergent phenotype in multiple sympatric populations does not necessarily imply that there has been repeated evolution of character displacement. Instead, such a pattern may arise if there has been character displacement in a single ancestral population, followed by gene flow carrying the divergent phenotype into multiple, derived, sympatric populations. Here, we evaluate the likelihood of such historical events versus ongoing ecological selection in generating divergence in trophic morphology between multiple populations of spadefoot toad (Spea multiplicata) tadpoles that are in sympatry with a heterospecific and those that are in allopatry. We present both phylogenetic and population genetic evidence indicating that the same divergent trait, which minimizes resource competition with the heterospecific, has arisen independently in multiple sympatric populations. These data, therefore, provide strong indirect support for competition''s role in divergent trait evolution.     

Scaling of Size and Dimorphism in Primates II: Macroevolution

Previous researchers found positive scaling of body size and sexual size dimorphism (SSD) in primates, known as Rensch's rule. The pattern is present in Haplorhini, but absent in Strepsirhini. I found that positive evolutionary correlations between size and SSD drive positive scaling relationships within Haplorhini as a whole and Platyrrhini, Cercopithecinae, Colobinae, and Hominoidea individually at the generic level and higher, but that evolutionary correlations within genera in these clades are often nonsignificant or negative. I suggest that positive evolutionary correlations result from greater change in male than in female size, usually because of sexual selection acting on polygynous populations. I suggest that negative evolutionary correlations result from greater change in female size, owing to either natural selection or, in Callitrichidae, sexual selection acting on polyandrous populations. The high incidence of negative evolutionary correlations within Haplorhini suggests a relatively large influence of natural selection on SSD, at least with regard to differences in SSD between congeners. I propose two possible explanations for the difference in intrageneric and supergeneric evolutionary patterns: 1) natural selection is a relatively weak force for modifying SSD and has a noticeable effect only when one compares related species experiencing similar levels of sexual selection, and 2) natural selection is a relatively strong force for modifying SSD but is less likely than sexual selection to affect higher level taxonomic comparisons noticeably because of the cumulative effect over time of marginal differences in mortality rates of these two types of selection. I discuss types of data required to test these explanations and implications for reconstructing fossil behavior.     

Evidence for large-scale effects of competition: niche displacement in Canada lynx and bobcat

Determining the patterns, causes and consequences of character displacement is central to our understanding of competition in ecological communities. However, the majority of competition research has occurred over small spatial extents or focused on fine-scale differences in morphology or behaviour. The effects of competition on broad-scale distribution and niche characteristics of species remain poorly understood but critically important. Using range-wide species distribution models, we evaluated whether Canada lynx (Lynx canadensis) or bobcat (Lynx rufus) were displaced in regions of sympatry. Consistent with our prediction, we found that lynx niches were less similar to those of bobcat in areas of sympatry versus allopatry, with a stronger reliance on snow cover driving lynx niche divergence in the sympatric zone. By contrast, bobcat increased niche breadth in zones of sympatry, and bobcat niches were equally similar to those of lynx in zones of sympatry and allopatry. These findings suggest that competitively disadvantaged species avoid competition at large scales by restricting their niche to highly suitable conditions, while superior competitors expand the diversity of environments used. Our results indicate that competition can manifest within climatic niche space across species’ ranges, highlighting the importance of biotic interactions occurring at large spatial scales on niche dynamics.