Exploitative competition between invasive herbivores benefits a native host plant

Although biological invasions are of considerable concern to ecologists, relatively little attention has been paid to the potential for and consequences of indirect interactions between invasive species. Such interactions are generally thought to enhance invasives' spread and impact (i.e., the "invasional meltdown" hypothesis); however, exotic species might also act indirectly to slow the spread or blunt the impact of other invasives. On the east coast of the United States, the invasive hemlock woolly adelgid (Adelges tsugae, HWA) and elongate hemlock scale (Fiorinia externa, EHS) both feed on eastern hemlock (Tsuga canadensis). Of the two insects, HWA is considered far more damaging and disproportionately responsible for hemlock mortality. We describe research assessing the interaction between HWA and EHS, and the consequences of this interaction for eastern hemlock. We conducted an experiment in which uninfested hemlock branches were experimentally infested with herbivores in a 2 x 2 factorial design (either, both, or neither herbivore species). Over the 2.5-year course of the experiment, each herbivore's density was approximately 30% lower in mixed- vs. single-species treatments. Intriguingly, however, interspecific competition weakened rather than enhanced plant damage: growth was lower in the HWA-only treatment than in the HWA + EHS, EHS-only, or control treatments. Our results suggest that, for HWA-infested hemlocks, the benefit of co-occurring EHS infestations (reduced HWA density) may outweigh the cost (increased resource depletion).     

Can rare positive interactions become common when large carnivores consume livestock?

Livestock populations in protected areas are viewed negatively because of their interaction with native ungulates through direct competition for food resources. However, livestock and native prey can also interact indirectly through their shared predator. Indirect interactions between two prey species occur when one prey modifies either the functional or numerical responses of a shared predator. This interaction is often manifested as negative effects (apparent competition) on one or both prey species through increased predation risk. But indirect interactions can also yield positive effects on a focal prey if the shared predator modifies its functional response toward increased consumption of an abundant and higher-quality alternative prey. Such a phenomenon between two prey species is underappreciated and overlooked in nature. Positive indirect effects can be expected to occur in livestock-dominated wildlife reserves containing large carnivores. We searched for such positive effects in Acacia-Zizhypus forests of India's Gir sanctuary where livestock (Bubalus bubalis and Bos indicus) and a coexisting native prey (chital deer, Axis axis) are consumed by Asiatic lions (Panthera leo persica). Chital vigilance was higher in areas with low livestock density than in areas with high livestock density. This positive indirect effect occurred because lion predation rates on livestock were twice as great where livestock were abundant than where livestock density was low. Positive indirect interactions mediated by shared predators may be more common than generally thought with rather major consequences for ecological understanding and conservation. We encourage further studies to understand outcomes of indirect interactions on long-term predator-prey dynamics in livestock-dominated protected areas.     

From interference to predation: type and effects of direct interspecific interactions of small mammals

Indirect exploitative competition, direct interference and predation are important interactions affecting species coexistence. These interaction types may overlap and vary with the season and life-history state of individuals. We studied effects of competition and potential nest predation by common shrews (Sorex araneus) on lactating bank voles (Myodes glareolus) in two seasons. The species coexist and may interact aggressively. Additionally, shrews can prey on nestling voles. We studied bank vole mothers’ spatial and temporal adaptations to shrew presence during summer and autumn. Further, we focused on fitness costs, e.g. decreased offspring survival, which bank voles may experience in the presence of shrews. In summer, interference with shrews decreased the voles’ home ranges and they spent more time outside the nest, but there were no effects on offspring survival. In autumn, we found decreased offspring survival in enclosures with shrews, potentially due to nest predation by shrews or by increased competition between species. Our results indicate a shift between interaction types depending on seasonal constraints. In summer, voles and shrews seem to interact mainly by interference, whereas resource competition and/or nest predation by shrews gain importance in autumn. Different food availability, changing environmental conditions and the energetic constraints in voles and shrews later in the year may be the reasons for the varying combinations of interaction types and their increasing effects on the inclusive fitness of bank voles. Our study provides evidence for the need of studies combining life history with behavioural measurements and seasonal constraints.     

Behavioural interference between ungulate species: roe are not on velvet with fallow deer

Interference is expected to occur at feeding areas between species with a similar diet, but few studies have tested this idea for wild ungulates. We analysed interactions between fallow deer, European roe deer and wild boar, in three sites, in a Mediterranean area. We expected that interference should be greater between deer than between them and wild boar. We documented the negative effects of behavioural interference by fallow on foraging behaviour of roe deer, under field conditions. Deer species built up 90% interference interactions, with fallow always dominant on roe, also through direct aggression. Although roe deer decreased feeding and increased vigilance levels in proximity (<50 m) of either fallow deer or wild boar, they were displaced significantly more often by the former than by the latter. Fallow deer were neither displaced nor alarmed by roe and rarely by wild boar. No deer species displaced wild boar. Interference was significantly greater on solitary roe deer, especially females, in spring and roe left the feeding ground most often in the smallest site (13 ha). Roe deer avoided areas where the local density of fallow deer was the highest. During our 4-year-study, roe deer density decreased whereas fallow deer numbers increased. Behavioural interference may explain how fallow deer outcompete roe deer through spatial exclusion from feeding sites and avoidance of areas with high densities of the former. Fallow deer evolved in semi-arid, relatively poor habitats of Asia Minor: interspecific defence of crucial resources could have developed as a beneficial tactic for its survival.     

Biophysical feedback mediates effects of invasive grasses on coastal dune shape

Vegetation at the aquatic-terrestrial interface can alter landscape features through its growth and interactions with sediment and fluids. Even similar species may impart different effects due to variation in their interactions and feedbacks with the environment. Consequently, replacement of one engineering species by another can cause significant change in the physical environment. Here we investigate the species-specific ecological mechanisms influencing the geomorphology of U.S. Pacific Northwest coastal dunes. Over the last century, this system changed from open, shifting sand dunes with sparse vegetation (including native beach grass, Elymus mollis), to densely vegetated continuous foredune ridges resulting from the introduction and subsequent invasions of two nonnative grass species (Ammophila arenaria and Ammophila breviligulata), each of which is associated with different dune shapes and sediment supply rates along the coast. Here we propose a biophysical feedback responsible for differences in dune shape, and we investigate two, non-mutually exclusive ecological mechanisms for these differences: (1) species differ in their ability to capture sand and (2) species differ in their growth habit in response to sand deposition. To investigate sand capture, we used a moveable bed wind tunnel experiment and found that increasing tiller density increased sand capture efficiency and that, under different experimental densities, the native grass had higher sand capture efficiency compared to the Ammophila congeners. However, the greater densities of nonnative grasses under field conditions suggest that they have greater potential to capture more sand overall. We used a mesocosm experiment to look at plant growth responses to sand deposition and found that, in response to increasing sand supply rates, A. arenaria produced higher-density vertical tillers (characteristic of higher sand capture efficiency), while A. breviligulata and E. mollis responded with lower-density lateral tiller growth (characteristic of lower sand capture efficiency). Combined, these experiments provide evidence for a species-specific effect on coastal dune shape. Understanding how dominant ecosystem engineers, especially nonnative ones, differ in their interactions with abiotic factors is necessary to better parameterize coastal vulnerability models and inform management practices related to both coastal protection ecosystem services and ecosystem restoration.     

An Assessment of Invasion Risk from Assisted Migration

Abstract:  To reduce the risk of extinction due to climate change, some ecologists have suggested human-aided translocation of species, or assisted migration (AM), to areas where climate is projected to become suitable. Such intentional movement, however, may create new invasive species if successful introductions grow out of control and cause ecologic or economic damage. We assessed this risk by surveying invasive species in the United States and categorizing invaders based on origin. Because AM will involve moving species on a regional scale within continents (i.e., range shifts), we used invasive species with an intracontinental origin as a proxy for species that would be moved through AM. We then determined whether intracontinental invasions were more prevalent or harmful than intercontinental invasions. Intracontinental invasions occurred far less frequently than invasions from other continents, but they were just as likely to have had severe effects. Fish and crustaceans pose a particularly high threat of intracontinental invasion. We conclude that the risk of AM to create novel invasive species is small, but assisted species that do become invasive could have large effects. Past experience with species reintroductions may help inform policy regarding AM.     

Interference competition between introduced black rats and endemic Galápagos rice rats

Replicated field experiments were used to quantify and to describe the mechanism of competition between the introduced black rat Rattus rattus and the endemic Santiago rice rat Nesoryzomys swarthi on Santiago Island, Galápagos Islands, Ecuador. The removal of R. rattus significantly slowed the rate of seasonal population decline in N. swarthi. This effect was particularly evident for female, relative to male, N. swarthi and appeared to be driven solely by enhanced immigration; no other fitness or space use parameters were affected. The candidate hypotheses to explain the mechanism of competition were exploitation competition, interference by resource defense, and interference by aggressive encounter. To distinguish between hypotheses, we conducted a replicated resource supplementation experiment with patchy food, scattered food, and no food (control) treatments. The opportunistic R. rattus responded to the extra resources with increased adult immigration and juvenile recruitment, resulting in a significant abundance boost of sevenfold on patchy grids and fourfold on scattered grids. Females increased in body mass, and the breeding season was lengthened. In contrast, there was no change in the abundance of N. swarthi and no obvious benefit to reproduction. Instead, the costs of interference apparently outweighed the benefits of extra food: female N. swarthi increased in mass with supplementary food, but female (relative to male) immigration and residency were repressed on all supplemented areas. This response supported the hypothesis of interference by aggressive encounter, and we were able to rule out the alternative hypotheses. Although periodic population crashes of R. rattus on the arid north coast of Santiago may ameliorate its competitive impact, climate change may tip the balance. Control or eradication of R. rattus should improve future survival prospects for N. swarthi, but wildlife managers must be prepared for the potential eruption of the introduced house mouse Mus musculus, because this species experienced a release from interference competition and immigrated to removal areas.     

Neighborhood phylodiversity affects plant performance

Facilitation and competition are ecological interactions that are crucial for the organization of plant communities. Facilitative interactions tend to occur among distantly related species, while the strength of competition tends to decrease with phylogenetic distance. The balance between both types of interactions will ultimately determine the specific composition of multispecies associations. Although multispecies patches are the arena in which coexistence develops among different phylogenetic groups within communities, the specific processes that occur across life stages have not been explored. Here we study how different species, in composing discrete patches in central Mexico, exert competitive or facilitative effects on seeds and seedlings. We relate these interactions to phylogenetic relationships among nurse species and beneficiary species, and among members of the patches. Survivorship and growth rates of the columnar cactus Neobuxbaumia mezcalaensis were highly positively related to increasing phylogenetic distance to different nurse species, to the presence of related species in patches, and to mean phylogenetic distances to the rest of the species in the patch. Each of these three elements influenced N. mezcalaensis differently, with different nurse species varying substantially in their early effects on emergence, and the nearest relatives and species composition of patches varying in their late effects on survival and growth. Our results emphasize that evolutionary relationships among co-occurring species in vegetation clumps exert direct and indirect effects on plants, affecting individual performance and species coexistence.     

Importance of greater interdisciplinarity and geographic scope when tackling the driving forces behind biological invasions

Invasive non-native species are important drivers of ecosystem change, yet the driving forces of biological invasions themselves are poorly understood. Such information is essential to ensure policies focus on the most relevant drivers, and that future scenarios capture the full range of potential outcomes for invasive non-native species. I carried out a bibliometric analysis of articles published from 2000 to 2020 that address either invasive non-native species or biodiversity and ecosystem services and that also mention 1 or more drivers of ecosystem change. I examined 5 indirect drivers (demographic, economic, governance, sociocultural, and technological) and 6 direct drivers (climate change, invasive non-native species, land-use or sea-use change, natural hazards, pollution, and resource extraction). Using the Web of Science core collection of citation indexes, I undertook searches of article titles and keywords and retrieved 27,462 articles addressing invasive non-native species and 110,087 articles dealing with biodiversity or ecosystem services. Most research to date on biological invasions as well as on biodiversity and ecosystem services has focused on anthropogenic direct drivers of ecosystem change rather than indirect drivers. Yet currently, less than 18% of articles addressing biological invasions examined drivers of ecosystem change, a similar level to that found over 20 years ago for biodiversity or ecosystem services. Knowledge of the drivers of biological invasions is limited, emphasizes tractable drivers over those that require an interdisciplinary approach, and is biased toward developed economies. Drivers generally deemed important for biological invasions, such as governance and resource extraction, accounted for less than 2% of research effort. The absence of a systematic understanding of the forces that drive invasive non-native species and how they interact means that attempts to mitigate or forecast biological invasions are likely to fail. To address biological invasions requires a much better orientation of national and international research on drivers in relation to both their actual importance as well as their policy relevance.     

Asymmetric competition via induced resistance: specialist herbivores indirectly suppress generalist preference and populations

Species may compete indirectly by altering the traits of a shared resource. For example, herbivore-induced responses in plants may make plants more resistant or susceptible to additional herbivorous insect species. Herbivore-induced plant responses can significantly affect interspecific competition and herbivore population dynamics. These herbivore-herbivore indirect interactions have been overlooked in aquatic ecosystems where previous studies used the same herbivore species to induce changes and to assess the effects of these changes. We asked whether seaweed grazing by one of two herbivorous, congeneric snail species (Littorina obtusata or Littorina littorea) with different feeding strategies and preferences would affect subsequent feeding preferences of three herbivore species (both snails and the isopod Idotea baltica) and population densities of three herbivore species (both snails and a third periwinkle snail, Lacuna vincta). In addition, we measured phlorotannin concentrations to test the hypothesis that these metabolites function as induced defenses in the Phaeophyceae. Snail herbivory induced cue-specific responses in apical tissues of the seaweed Fucus vesiculosus that affected the three herbivore species similarly. When compared to ungrazed controls, direct grazing by Littorina obtusata reduced seaweed palatability by at least 52% for both snail species and the isopod species. In contrast, direct grazing by L. littorea did not decrease seaweed palatability for any herbivore, indicating herbivore-specific responses. Previous grazing by L. obtusata reduced populations of L. littorea on outplanted seaweeds by 46% but had no effect on L. obtusata populations. Phlorotannins, a potential class of inducible chemicals in brown algae, were not more concentrated in grazed seaweed tissues, suggesting that some other trait was responsible for the induced resistance. Our results indicate that marine herbivores may compete via inducible responses in shared seaweeds. These plant-mediated interactions were asymmetric with a specialist (L. obtusata) competitively superior to a generalist (L. littorea).     

Potential Interference Between a Threatened Endemic Thistle and an Invasive Nonnative Plant

Although the establishment of nonnative plants is recognized as a threat to native ecosystems, there are few documented examples of an invasive species directly influencing a rare native plant. The Eurasian biennial Dipsacus sylvestris (teasel) is invading the central New Mexico habitat of Cirsium vinaceum , an endemic thistle that is federally listed as threatened. We documented changes in teasel distribution and abundance between 1989 and 1993 that suggest the potential for direct interactions with the native thistle. We then compared habitat characteristics, germination behavior, and performance in greenhouse and field competition trials to evaluate the potential outcome of interference between these two species. There were no significant differences in measured habitat characteristics between sites supporting C. vinaceum and those with D. sylvestris. Dipsacus was better able to germinate in low light than the thistle, suggesting that D. sylvestris might invade C. vinaceum populations but that thistle recruitment would be unlikely in dense stands of the nonnative plants. In the greenhouse growth of C. vinaceum rosettes was significantly reduced by the presence of Dipsacus , but the invader was unaffected by the thistle; results of a short-term field experiment were equivocal but suggestive of interference between the two. We suggest criteria for managers to use in determining whether invading species pose problems for specific rare native taxa, and we discuss the constraints on experimental work where protected taxa are involved.     

The significance of metal hyperaccumulation for biotic interactions

Metal hyperaccumulating plants contain very high metal contents. Because of the general toxicity of metals, chemically-mediated biotic interactions involving hyperaccumulating plants may differ greatly from those of non-hyperaccumulators. Recent research has demonstrated a defensive function for hyperaccumulated metals against herbivores and pathogens. We predict that some herbivore/pathogen species have evolved metal tolerance, and suggest that resulting high metal levels in herbivores/pathogens may defend them against their own predators. Little is known regarding interference and commensal interactions involving hyperaccumulating plants. Decreased competition may occur through an interference interaction similar to allelopathy, in which enrichment of metal in the soil under a hyperaccumulator plant's canopy may inhibit another plant species, thus resulting in “elemental allelopathy”. Metal enrichment of soil under hyperaccumulators also may result in commensalism if another plant species (possibly another hyperaccumulator) derives a benefit from growing in the metal-enriched soil under the canopy of a hyperaccumulating overstory plant. It seems likely that high-metal plant litter will host a specialized microflora of decomposers and may affect nutrient cycling rates. Mutualist biotic interactions also may be affected by the elevated metal contents of hyperaccumulating species. Mycorrhizal fungi may form mutualisms with hyperaccumulators, but the phenomenon is poorly-explored. The few cases investigated to date have not detected mycorrhizae. Pollination and seed dispersal mechanisms may require biotic vectors that might be affected by plant metal content. Hyperaccumulating plants may have solved this dilemma in three ways. First, some may rely on abiotic vectors for pollen or seed dispersal. Second, biotic vectors used by these species may have varied diets and thus dilute metal intake to non-toxic levels. Finally, biotic vectors may have evolved tolerance of elevated dietary levels of metals, and perhaps have become specialists on hyperaccumulator species. Received 7 November 1997; accepted 28 December 1997.     

Spatial and temporal variation in distribution and protein ubiquitination for <Emphasis Type="Italic">Mytilus</Emphasis> congeners in the California hybrid zone

Along the west coast of North America, the invasive mussel Mytilus galloprovincialis and a native congener M. trossulus overlap in range and compete for habitat in an extensive hybrid zone along central California. The two species have been shown to exhibit differential abiotic tolerances in laboratory studies, yet little is known about how such tolerances affect spatial and temporal patterns of geographic distribution, particularly in areas of competition. We examined distributions of the two congeners and their hybrids in neighboring intertidal and subtidal habitats in Bodega Bay, CA over 2 years, and compared shell length and seasonal ubiquitin (Ub) conjugates to estimate protein turnover and physiological stress for the species at each site. The two species were spatially segregated, with M. galloprovincialis dominating the subtidal habitat, and M. trossulus constituting a majority of the intertidal mussel population. Hybrid individuals appeared in low numbers at both sites. For each habitat, there was no statistical difference between shell lengths of M. galloprovincialis and hybrids but M. trossulus mussels were statistically smaller than the other two. In regards to physiological performance, ubiquitin conjugate values showed different seasonal cycles for the two species, suggesting different periods of peak environmental stress. The highest levels of Ub-conjugated proteins were observed in winter for M. galloprovincialis and in summer for M. trossulus, consistent with the respective range edges for their distributions since Bodega Bay is near the northern range edge of the invader and the southern edge of the native species. These findings suggest that future assessments of Mytilus populations along the California coast may need to consider vertical distributions and seasonal cycles as part of monitoring and research activities.     

Causes of habitat divergence in two species of agamid lizards in arid central Australia

The deserts of central Australia contain richer communities of lizards than any other arid regions, with the highest diversity occurring in sand dune habitats dominated by hummock-forming spinifex grasses. To investigate the mechanisms that permit coexistence, we studied two species of coexisting agamid lizards that exhibit striking divergence in their use of habitat in the Simpson Desert of central Australia. Here, the military dragon Ctenophorus isolepis is restricted primarily to sites providing > 30% cover of hard spinifex Triodia basedowii, whereas the central netted dragon C. nuchalis occurs in areas with much sparser (< 10%) cover. We constructed four mechanistic models to explain this pattern and then derived hypotheses to test them. One hypothesis, that competition restricts each species to its preferred habitat, was rejected after dyad encounters in field enclosures failed to elicit any habitat shift or any overt interactions between the species. Our next hypotheses were that each species exhibits preferences for different thermal environments or different prey types and that each selects the habitats that maximize access to them. Both were supported. C. isolepis preferred lower temperatures when active and specialized in eating ants < 5 mm long and selected spinifex-dominated areas where these requirements were met. In contrast, C. nuchalis preferred higher temperatures and a diversity of prey, both of which were available mostly in open areas. Finally, we used plasticine models to test the hypothesis that each species faced lower risk of predation in its selected habitat. This was partly supported, as models of both species were attacked more often in the open than under spinifex cover. The results show that habitat divergence occurs along several, probably covarying, niche axes. We suggest that different levels of spinifex cover provide the template for a broad range of ecological interactions, allowing lizard species to partition biotic and abiotic resources and achieve the extraordinarily high levels of local diversity that are observed.     

Invasional meltdown: invader-invader mutualism facilitates a secondary invasion

In multiply invaded ecosystems, introduced species should interact with each other as well as with native species. Invader-invader interactions may affect the success of further invaders by altering attributes of recipient communities and propagule pressure. The invasional meltdown hypothesis (IMH) posits that positive interactions among invaders initiate positive population-level feedback that intensifies impacts and promotes secondary invasions. IMH remains controversial: few studies show feedback between invaders that amplifies their effects, and none yet demonstrate facilitation of entry and spread of secondary invaders. Our results show that supercolonies of an alien ant, promoted by mutualism with introduced honeydew-secreting scale insects, permitted invasion by an exotic land snail on Christmas Island, Indian Ocean. Modeling of land snail spread over 750 sites across 135 km2 over seven years showed that the probability of land snail invasion was facilitated 253-fold in ant supercolonies but impeded in intact forest where predaceous native land crabs remained abundant. Land snail occurrence at neighboring sites, a measure of propagule pressure, also promoted land snail spread. Site comparisons and experiments revealed that ant supercolonies, by killing land crabs but not land snails, disrupted biotic resistance and provided enemy-free space. Predation pressure on land snails was lower (28.6%), survival 115 times longer, and abundance 20-fold greater in supercolonies than in intact forest. Whole-ecosystem suppression of supercolonies reversed the probability of land snail invasion by allowing recolonization of land crabs; land snails were much less likely (0.79%) to invade sites where supercolonies were suppressed than where they remained intact. Our results provide strong empirical evidence for IMH by demonstrating that mutualism between invaders reconfigures key interactions in the recipient community. This facilitates entry of secondary invaders and elevates propagule pressure, propagating their spread at the whole-ecosystem level. We show that identification and management of key facilitative interactions in invaded ecosystems can be used to reverse impacts and restore resistance to further invasions.     

Effects of Invasive Plants on Arthropods

Non‐native plants have invaded nearly all ecosystems and represent a major component of global ecological change. Plant invasions frequently change the composition and structure of vegetation communities, which can alter animal communities and ecosystem processes. We reviewed 87 articles published in the peer‐reviewed literature to evaluate responses of arthropod communities and functional groups to non‐native invasive plants. Total abundance of arthropods decreased in 62% of studies and increased in 15%. Taxonomic richness decreased in 48% of studies and increased in 13%. Herbivorous arthropods decreased in response to plant invasions in 48% of studies and increased in 17%, likely due to direct effects of decreased plant diversity. Predaceous arthropods decreased in response to invasive plants in 44% of studies, which may reflect indirect effects due to reductions in prey. Twenty‐two percent of studies documented increases in predators, which may reflect changes in vegetation structure that improved mobility, survival, or web‐building for these species. Detritivores increased in 67% of studies, likely in response to increased litter and decaying vegetation; no studies documented decreased abundance in this functional group. Although many researchers have examined effects of plant invasions on arthropods, sizeable information gaps remain, specifically regarding how invasive plants influence habitat and dietary requirements. Beyond this, the ability to predict changes in arthropod populations and communities associated with plant invasions could be improved by adopting a more functional and mechanistic approach. Understanding responses of arthropods to invasive plants will critically inform conservation of virtually all biodiversity and ecological processes because so many organisms depend on arthropods as prey or for their functional roles, including pollination, seed dispersal, and decomposition. Given their short generation times and ability to respond rapidly to ecological change, arthropods may be ideal targets for restoration and conservation activities. Efectos de las Plantas Invasoras sobre los Artrópodos     

Parents influence asymmetric sibling competition: experimental evidence with partially dependent young

Asymmetric sibling competition, which occurs when some siblings hatch as stronger competitors than others, is an important component of avian reproductive strategies. Here, we report two experiments on the burying beetle Nicrophorus vespilloides investigating how parents might influence the outcome of asymmetric sibling competition. In this species, as in altricial birds, different-aged offspring compete for resources provided by the parents. However, unlike altricial birds, offspring depend only partially on their parents for resources, and parents adjust the brood size directly through filial cannibalism. In the first experiment, we compared the growth and survivorship of different-aged offspring when parents could and could not influence asymmetric sibling competition. In the second experiment, we recorded behavioral interactions between different-aged offspring and parents. We found that senior offspring (early-hatched) grew faster than juniors (late-hatched) when parents were present and could influence the outcome of sibling competition, whereas seniors and juniors grew at similar rates when parents were removed. Thus, seniors benefited more than did juniors when the offspring could obtain resources by begging from the female parent. There was no difference in the survivorship of seniors and juniors. We also found that seniors and juniors spent a similar amount of time feeding from female parents, but juniors spent more time begging and were less effective at begging than seniors. Interestingly, juniors spent more time begging only as long as seniors also begged, suggesting that juniors adjusted their begging effort in response to direct competition against seniors for resources provided by parents. Our study provides novel insights into the ecological significance of asymmetric sibling competition by showing that asymmetric sibling competition took place when parents were present and offspring could obtain resources by begging. In contrast, we found no evidence of asymmetric sibling competition when parents were absent and offspring obtained resources solely by self-feeding.     

Importance of competition mechanisms in successive invasions by polyphagous tephritids in La Reunion

Understanding the strength and modes of interspecific interactions between introduced and resident species (native or previously introduced) is necessary to predict invasion success. We evaluated different mechanisms of interspecific competition among four species of polyphagous fruit flies (Diptera: Tephritidae) from the island of La Reunion: one endemic species, Ceratitis catoirii, and three exotic species, C. capitata, C. rosa, and Bactrocera zonata, that have successively invaded the island. Larval competition experiments, i.e., co-infestations of the same fruit, and behavioral interference experiments measuring the ability of one female to displace another from a fruit, were performed among all pairs of the four species. We observed asymmetric and hierarchical interactions among species in both larval and adult interference competition. In agreement with the hypothesis that invasion is competition-limited, the competitive hierarchy coincided with the temporal sequence of establishment on the island, i.e., each newly established species tended to be competitively dominant over previously established ones.     

The social and mating systems of two labrid fishes,Halichoeres maculipinna and H. garnoti,off the Caribbean coast of Panama

Large males of both species are territorial, particularly when spawning. Small Halichoeres maculipinna (but not H. garnoti) are territorial towards equal sized conspecifics, but not towards dissimilar sized conspecifics or any individuals of an abundant congener. Territoriality of small H. maculipinna may be related to the process of sex-change rather than defense of food. Both sexes of both species, but particularly H. maculipinna, migrate from feeding areas to spawning sites at the edges of reefs at specific times of day. In the two species a large male spawns both with females that live in his feeding area and with many others. Differences in the mating system of H. maculipinna in Panama and Florida seem related to habitat differences at those sites. The relative spatial distributions of feeding and spawning sites may affect (a) whether a female spawns with a large male with whom she feeds, and (b) the ability of large males to monopolize mates at the expense of small males. The spawning success of small males seems to depend on whether large males can penalize their participation in spawning rather than on the ability of small males to predict the occurrence of (and then interfere in) spawnings by large males. In both species, the male-specific color pattern is most intensely exhibited by large males during sexual interactions and tends to be lost when they are being aggressive towards conspecifics. The evolutionary development of such a color pattern in labroid fishes seems to be a response to intersexual rather than intrasexual interactions.