Preferences for different nitrogen forms by coexisting plant species and soil microbes

The growing awareness that plants might use a variety of nitrogen (N) forms, both organic and inorganic, has raised questions about the role of resource partitioning in plant communities. It has been proposed that coexisting plant species might be able to partition a limited N pool, thereby avoiding competition for resources, through the uptake of different chemical forms of N. In this study, we used in situ stable isotope labeling techniques to assess whether coexisting plant species of a temperate grassland (England, UK) display preferences for different chemical forms of N, including inorganic N and a range of amino acids of varying complexity. We also tested whether plants and soil microbes differ in their preference for different N forms, thereby relaxing competition for this limiting resource. We examined preferential uptake of a range of 13C15N-labeled amino acids (glycine, serine, and phenylalanine) and 15N-labeled inorganic N by coexisting grass species and soil microbes in the field. Our data show that while coexisting plant species simultaneously take up a variety of N forms, including inorganic N and amino acids, they all showed a preference for inorganic N over organic N and for simple over the more complex amino acids. Soil microbes outcompeted plants for added N after 50 hours, but in the long-term (33 days) the proportion of added 15N contained in the plant pool increased for all N forms except for phenylalanine, while the proportion in the microbial biomass declined relative to the first harvest. These findings suggest that in the longer-term plants become more effective competitors for added 15N. This might be due to microbial turnover releasing 15N back into the plant-soil system or to the mineralization and subsequent plant uptake of 15N transferred initially to the organic matter pool. We found no evidence that soil microbes preferentially utilize any of the N forms added, despite previous studies showing that microbial preferences for N forms vary over time. Our data suggest that coexisting plants can outcompete microbes for a variety of N forms, but that such plant species show similar preferences for inorganic over organic N.     

Niche complementarity due to plasticity in resource use: plant partitioning of chemical N forms

Niche complementarity, in which coexisting species use different forms of a resource, has been widely invoked to explain some of the most debated patterns in ecology, including maintenance of diversity and relationships between diversity and ecosystem function. However, classical models assume resource specialization in the form of distinct niches, which does not obviously apply to the broadly overlapping resource use in plant communities. Here we utilize an experimental framework based on competition theory to test whether plants partition resources via classical niche differentiation or via plasticity in resource use. We explore two alternatives: niche preemption, in which individuals respond to a superior competitor by switching to an alternative, less-used resource, and dominant plasticity, in which superior competitors exhibit high resource use plasticity and shift resource use depending on the competitive environment. We determined competitive ability by measuring growth responses with and without neighbors over a growing season and then used 15N tracer techniques to measure uptake of different nitrogen (N) forms in a field setting. We show that four alpine plant species of differing competitive abilities have statistically indistinguishable uptake patterns (nitrate > ammonium > glycine) in their fundamental niche (without competitors) but differ in whether they shift these uptake patterns in their realized niche (with competitors). Competitively superior species increased their uptake of the most available N form, ammonium, when in competition with the rarer, competitively inferior species. In contrast, the competitively inferior species did not alter its N uptake pattern in competition. The existence of plasticity in resource use among the dominant species provides a mechanism that helps to explain the manner by which plant species with broadly overlapping resource use might coexist.     

Nitrogen niches revealed through species and functional group removal in a boreal shrub community

Most theories attempting to explain the coexistence of species in local communities make fundamental assumptions regarding whether neighbors exhibit competitive, neutral, or positive resource-use interactions; however, few long-term data from naturally assembled plant communities exist to test these assumptions. We utilized a 13-year experiment consisting of factorial removal of three shrub species (Vaccinium myrtillus, V. vitis-idaea, and Empetrum hermaphroditum) and factorial removal of two functional groups (tree roots and feather mosses) to assess how neighbors affect N acquisition and growth of each of the three shrub species. The removal plots were established on each of 30 lake islands in northern Sweden that form a natural gradient of resource availability. We tested the hypotheses that: (1) the presence of functionally similar neighbors would reduce shrub N acquisition through competition for a shared N resource; (2) the removal of functional groups would affect shrub N acquisition by altering the breadth of their niches; and (3) soil fertility would influence the effects of neighbor removals. We found that the removal of functionally similar neighbors (i.e., other shrub species) usually resulted in higher biomass and biomass N, with the strength of these effects varying strongly with site fertility. Shrub species removals never resulted in altered stable N isotope ratios (delta(15)N), suggesting that the niche breadth of the three shrubs was unaffected by the presence of neighboring shrub species. In the functional group removal experiment, we found positive effects of feather moss removal on V. myrtillus biomass and biomass N, and negative effects on E. hermaphrotium N concentration and V. vitis-idaea biomass and biomass N. Tree root removal also caused a significant shift in foliar delta(15)N of V. myrtillus and altered the delta(15)N, biomass, and biomass N of E. hermaphroditum. Collectively, these results show that the resource acquisition and niche breadth of the three shrub species are often affected by neighbors, and further that both the identity of neighbors and site fertility strongly determine whether these interactions are positive, negative, or neutral. These findings have implications for understanding species coexistence and the reciprocal relationships between productivity and species diversity in this ecosystem.     

Competition among eucalyptus trees depends on genetic variation and resource supply

Genetic variation and environmental heterogeneity fundamentally shape the interactions between plants of the same species. According to the resource partitioning hypothesis, competition between neighbors intensifies as their similarity increases. Such competition may change in response to increasing supplies of limiting resources. We tested the resource partitioning hypothesis in stands of genetically identical (clone-origin) and genetically diverse (seed-origin) Eucalyptus trees with different water and nutrient supplies, using individual-based tree growth models. We found that genetic variation greatly reduced competitive interactions between neighboring trees, supporting the resource partitioning hypothesis. The importance of genetic variation for Eucalyptus growth patterns depended strongly on local stand structure and focal tree size. This suggests that spatial and temporal variation in the strength of species interactions leads to reversals in the growth rank of seed-origin and clone-origin trees. This study is one of the first to experimentally test the resource partitioning hypothesis for intergenotypic vs. intragenotypic interactions in trees. We provide evidence that variation at the level of genes, and not just species, is functionally important for driving individual and community-level processes in forested ecosystems.     

Neighborhood analyses of canopy tree competition along environmental gradients in New England forests.

We use permanent-plot data from the USDA Forest Service's Forest Inventory and Analysis (FIA) program for an analysis of the effects of competition on tree growth along environmental gradients for the 14 most abundant tree species in forests of northern New England, USA. Our analysis estimates actual growth for each individual tree of a given species as a function of average potential diameter growth modified by three sets of scalars that quantify the effects on growth of (1) initial target tree size (dbh), (2) local environmental conditions, and (3) crowding by neighboring trees. Potential growth of seven of the 14 species varied along at least one of the two environmental axes identified by an ordination of relative abundance of species in plots. The relative abundances of a number of species were significantly displaced from sites where they showed maximum potential growth. In all of these cases, abundance was displaced to the more resource-poor end of the environmental gradient (either low fertility or low moisture). The pattern was most pronounced among early successional species, whereas late-successional species reached their greatest abundance on sites where they also showed the highest growth in the absence of competition. The analysis also provides empirical estimates of the strength of intraspecific and interspecific competitive effects of neighbors. For all but one of the species, our results led us to reject the hypothesis that all species of competitors have equivalent effects on a target species. Most of the individual pairwise interactions were strongly asymmetric. There was a clear competitive hierarchy among the four most shade-tolerant species, and a separate competitive hierarchy among the shade-intolerant species. Our results suggest that timber yield following selective logging will vary dramatically depending on the configuration of the residual canopy, because of interspecific variation in the magnitude of both the competitive effects of different species of neighbors and the competitive responses of different species of target trees to neighbors. The matrix of competition coefficients suggests that there may be clear benefits in managing for specific mixtures of species within local neighborhoods within stands.     

Mechanistic analysis of interspecific competition using foraging trade-offs: implications for duck assemblages

The role of interspecific competition and resource partitioning in determining the composition of species assemblages is often controversial. In many cases data on species co-occurrence or resource use (prey or habitat) have been interpreted without a clear understanding of how, or even whether, phenotypic differences constrain performance to allow resource partitioning or how these constraints and the density of resources and competitors should shape resource selection by each species. Instead, predictions have been based on assumed constraints, possibly leading to conflicting results. One such controversy involves the role of bill morphology in mediating resource partitioning among dabbling ducks (Anas spp.). To determine whether incorrect assumptions may have contributed to this controversy, I constructed mechanistic models that predict filter-feeding performance for seven species of ducks directly from bill morphology and kinetics and compared these predictions to those of earlier studies that tested the bill morphology hypothesis. The models predicted that species should share a preference for their most profitable (primary) prey while partitioning their less profitable (secondary) prey by size. Consequently, ducks should forage in the same habitats and exhibit high overlap in prey size when competitor/resource ratios are either high or low. In contrast, earlier studies expected that resource partitioning should always be evident, which implicitly assumes that species partition their primary resources. The models also predicted that the ecological similarity of species in assemblages should increase as prey abundance and size variability declines, contrary to the expectations of an earlier study. A more consistent understanding of the mechanisms regulating assemblages of dabbling ducks, and other species, might emerge if patterns of resource use and species co-occurrence were predicted directly from a mechanistic understanding of how performance trade-offs affect resource selection in the context of varying resource and competitor densities.     

Sympatric sexual signal divergence among North American Calopteryx damselflies is correlated with increased intra- and interspecific male–male aggression

Divergence of sexual signals in sympatry can arise as a consequence of (1) interspecific competition for resources, (2) selection against maladaptive hybridization, or (3) as a result of selection to reduce the cost of interspecific aggression; termed agonistic character displacement (ACD). Calopterygid damselflies have emerged as a model system for studying the evolution of divergent sexual signals due to the repeated evolution of sympatric species pairs with fully and partially melanized wings. Damselfly wing patterns function during both courtship and territory defense. However, the relative contributions of natural and sexual selection to phenotypic divergence and enhanced isolation in sympatry remain unclear in many cases. Here, we investigated the hypothesis that interference competition, in the form of increased interspecific male–male aggression, drives the evolution of character displacement in sympatry between two species of North American damselflies, Calopteryx aequabilis and Calopteryx maculata, that show no evidence of ecological divergence or ongoing hybridization. In paired behavioral trials, we found that interspecific male aggression related to territory defense varied between site, species, and as a function of the relative abundance of con- vs. hetero-specific males. Specifically, we found that large-spotted C. aequabilis males received increased intra- and interspecific aggression but that aggression against large-spotted males declined during the middle of the flight season when both species were equally abundant. Based on these results, we suggest that ACD leads to enhanced species recognition, and may be a common outcome of the antagonism between interspecific male–male competition and the countervailing force of intraspecific sexual selection favoring increased wing melanization among territorial damselfly species.     

巨枝人工林灌木层优势种的种间联结性分析

种间联结的研究对于人工林群落的生物多样性保育和可持续经营与管理具有重要的意义,目前对巨桉人工林灌木层物种之间的作用关系知之甚少。根据野外60个样方的调查数据,运用x^2检验、Jaccard指数和Spearman秩相关系数研究了巨桉（Eucalyptusgrandis）人工林灌木层18个优势种的种间关联和相关关系。结果表明,灌木层优势种具有总体上的正联结性,但在153个种对中,具有显著正联结性的种对数约占总种对数的10％,绝大数种对表现为弱联结性或无联结性,种间关系较为松散,表明巨桉林下灌木种群具有相对独立分布的特性。研究种间关系时,种间关联结合种间相关得出的结果较好。根据分析结果将18个优势种划分为4个生态种组,生态种组内的种其资源利用方式和生态要求相似,而组间表现出明显的差异。生态种组的划分能为巨桉人工林林下灌木层的管理提供理论依据。     

Testing hypotheses on shape and distribution of ecological response curves

Niche theory with hypotheses on shape and distribution of ecological response curves is used in the studies of resource sharing of competing plant species. Predictions based on theory should be applicable when, e.g., effects of competing species on the ecological tolerances are assessed or species’ diversity along a resource gradient is evaluated. We studied the ecological response curves of competing plant species along a resource gradient in boreal forests. The study was based on nation-wide soil and vegetation data collected from 455 sample plots on boreal forests in Finland. Species response curves along a soil fertility gradient (in terms of C/N ratio) were estimated using generalized additive models. Distribution of species optima and the relationship of niche width and skewness to the location of the optimum were analyzed with new bootstrap tests. The developed tests can account for the effects of truncation observed in the response curves of several species and for the uneven distribution of observations on the gradient.The estimated response curves of the major field layer species of boreal forests were not evenly distributed along soil C/N gradient. The density of optima peaked with relatively high nitrogen availability. Species with optima at low nitrogen availability had relatively broad realized niches. Niche width was negatively correlated with the density of optima. Species optima were packed and niches were narrow at high resource levels. This result suggests that a greater number of more specialized species can occur and interspecific competition decreases niche widths at high resource levels. Species were packed in the gradient where the C/N ratio was lower than 25, i.e., in conditions where nitrification can take place. This indicates that the majority of the vascular plants of boreal forests are favoured by the availability of NO₃⁻. Those few species thriving at high C/N ratios have broader realized niches.     

Niche complementarity for nitrogen: an explanation for the biodiversity and ecosystem functioning relationship?

The relationship between plant diversity and productivity has largely been attributed to niche complementarity, assuming that plant species are complementary in their resource use. In this context, we conducted an 15N field study in three different grasslands, testing complementarity nitrogen (N) uptake patterns in terms of space, time, and chemical form as well as N strategies such as soil N use, symbiotic N fixation, or internal N recycling for different plant species. The relative contribution of different spatial, temporal, and chemical soil N pools to total soil N uptake of plants varied significantly among the investigated plant species, within and across functional groups. This suggests that plants occupy distinct niches with respect to their relative N uptake. However, when the absolute N uptake from the different soil N pools was analyzed, no spatial, temporal, or chemical variability was detected, but plants, and in particular functional groups, differed significantly with respect to their total soil N uptake irrespective of treatment. Consequently, our data suggest that absolute N exploitation on the ecosystem level is determined by species or functional group identity and thus by community composition rather than by complementary biodiversity effects. Across functional groups, total N uptake from the soil was negatively correlated with leaf N concentrations, suggesting that these functional groups follow different N use strategies to meet their N demands. While our findings give no evidence for a biodiversity effect on the quantitative exploitation of different soil N pools, there is evidence for different and complementary N strategies and thus a potentially beneficial effect of functional group diversity on ecosystem functioning.     

Coexistence of three specialist aphids on common milkweed, Asclepias syriaca

Coexistence of host-specific herbivores on plants is believed to be governed by interspecific interactions, but few empirical studies have systematically unraveled these dynamics. We investigated the role of several factors in promoting coexistence among the aphids Aphis nerii, Aphis asclepiadis, and Myzocallis asclepiadis that all specialize on common milkweed (Asclepias syriaca). Competitive exclusion is thought to occur when interspecific competition is stronger than intraspecific competition. Consequently, we investigated whether predators, mutualists, or resource quality affected the strength of intra- vs. interspecific competition among aphids in factorial manipulations of competition with exposure to predation, ants, and variable plant genotypes in three separate experiments. In the predation x competition experiment, predators reduced aphid per capita growth by 66%, but the strength of intra- and interspecific competition did not depend on predators. In the ants x competition experiment, ants reduced per capita growth of A. nerii and M. asclepiadis (neither of which were mutualists with ants) by approximately one-half. In so doing, ants ameliorated the negative effects of these competitors on ant-tended A. asclepiadis by two-thirds, representing a novel benefit of ant-aphid mutualism. Nevertheless, ants alone did not explain the persistence of competitively inferior A. asclepiadis as, even in the presence of ants, interspecific competition remained stronger than intraspecific competition. In the plant genotype x competition experiment, both A. asclepiadis and M. asclepiadis were competitively inferior to A. nerii, with the strength of interspecific competition exceeding that of intraspecific competition by 83% and 23%, respectively. Yet these effects differed among milkweed genotypes, and there were one or more plant genotypes for each aphid species where coexistence was predicted. A synthesis of our results shows that predators play little or no role in preferentially suppressing competitively dominant A. nerii. Nonetheless, A. asclepiadis benefits from ants, and A. asclepiadis and M. asclepiadis may escape competitive exclusion by A. nerii on select milkweed genotypes. Taken as a whole, the coexistence of three host-specific aphid species sharing the same resource was promoted by the dual action of ants as antagonists and mutualists and by genetic diversity in the plant population itself.     

Reproductive interference via interspecific pairing in an amphipod species complex

Many species experience reproductive interference defined as interspecific mating interactions that reduce the fitness of individuals in at least one of the species. We examined reproductive interference among three cryptic amphipod species that co-occur in the vegetated zone of lakes. Because these animals form precopulatory pairs and males commonly have indiscriminant mating behavior, we predicted that (1) there would be interspecific pairing, (2) interspecific pairing would be more common when conspecifics were rare, and (3) interspecific pairing would be more common in species combinations where males are larger than females. Using color phenotypes to distinguish species, a novel discovery reported here, we conducted experiments to evaluate patterns of interspecific pairing. In a no-choice experiment (i.e., amphipods had access to mates of only one species), we observed a low rate (5 %) of interspecific pairing and 80 % of these cases involved males that were larger than females. In a second experiment where individuals had access to conspecific mates at varying relative abundance, only 0.04 % of the pairs observed were between heterospecifics, suggesting that the interspecific pairing observed in the no-choice experiment was an artifact of not having access to conspecific mates. Our results suggest that at least one sex has sufficient species-recognition abilities to circumvent the formation of interspecific pairs; therefore, reproductive interference may be minimal in these species.     

Increasing plant use of organic nitrogen with elevation is reflected in nitrogen uptake rates and ecosystem delta15N

It is hypothesized that decreasing mean annual temperature and rates of nitrogen (N) cycling causes plants to switch from inorganic to organic forms of N as the primary mode of N nutrition. To test this hypothesis, we conducted field experiments and collected natural-abundance delta15N signatures of foliage, soils, and ectomycorrhizal sporocarps along a steep elevation-climate gradient in the White Mountains, New Hampshire, USA. Here we show that with increasing elevation organic forms of N became the dominant source of N taken up by hardwood and coniferous tree species based on dual-labeled glycine uptake analysis, an important confirmation of an emerging theory for the biogeochemistry of the N cycle. Variation in natural abundance foliar delta15N with elevation was also consistent with increasing organic N uptake, though a simple, mass balance model demonstrated that the uptake of delta15N depleted inorganic N, rather than fractionation upon transfer of N from mycorrhizal fungi, best explains variations in foliar delta15N with elevation.     

Increasing enemy biodiversity strengthens herbivore suppression on two plant species

Concern over biodiversity loss, especially at higher trophic levels, has led to a surge in studies investigating how changes in natural enemy diversity affect community and ecosystem functioning. These studies have found that increasing enemy diversity can strengthen, weaken, and not affect prey suppression, demonstrating that multi-enemy effects on prey are context-dependent. Here we ask how one factor, plant species identity, influences multi-enemy effects on prey. We focused on two plant species of agricultural importance, potato (Solanum tuberosum), and collards (Brassica oleracea L.). These species share a common herbivorous pest, the green peach aphid (Myzus persicae), but vary in structural and chemical traits that affect aphid reproductive rates and which may also influence inter-enemy interactions. In a large-scale field experiment, overall prey exploitation varied dramatically among the plant species, with enemies reducing aphid populations by approximately 94% on potatoes and approximately 62% on collards. Increasing enemy diversity similarly strengthened aphid suppression on both plants, however, and there was no evidence that plant species identity significantly altered the relationship between enemy diversity and prey suppression. Microcosm experiments suggested that, on both collards and potatoes, intraspecific competition among natural enemies exceeded interspecific competition. Enemy species showed consistent and significant differences in where they foraged on the plants, and enemies in the low-diversity treatment tended to spend less time foraging than enemies in the high-diversity treatment. These data suggest that increasing enemy diversity may strengthen aphid suppression because interspecific differences in where enemies forage on the plant allow for greater resource partitioning. Further, these functional benefits of diversity appear to be robust to changes in plant species identity.     

A chemical trait creates a genetic trade-off between intra- and interspecific competitive ability

The importance of non-resource-based mechanisms of competition between plant species has been increasingly recognized, but little is known about how genetic variation and evolutionary changes in the underlying competitive traits might affect species coexistence. I found that genetic variation in sinigrin concentration, a putative allelopathic agent in Brassica nigra, affected the fitness of three heterospecific neighbor species but did not affect neighboring B. nigra individuals. Investment in sinigrin led to a negative genetic correlation between intra- and interspecific competitive ability, which over many generations could provide a strong stabilizing force maintaining both species and genetic diversity in this system.     

Area-specific temporal changes of species composition and species-specific range shifts in rocky-shore mollusks associated with warming Kuroshio Current

Increases of low-latitude species in various sea areas and poleward shifts of ranges of many species have been reported and linked to warming environments. To examine the generality of these trends for mollusks, we conducted 7 quadrat surveys during 1978–2006 on Pacific rocky shores in Japan (26.6–45.0°N). Results showed that the dominance of southern species increased on 11 of the 15 shores in a southern, warming sea area but on only 1 of the 6 shores in a northern area with no warming trend. The latitudes of species ranges increased on average but showed large interspecific variations which showed a weak, positive correlation with interspecific taxonomic distance. The differences of these results from the previously reported trends are discussed in relation to the unique current patterns in our study area and the phylogenetic constraints of species-specific responses to a warming environment.     

Patch size-dependent community recovery after massive disturbance

Massive anthropogenic and climate-related disturbances are now common in ecosystems worldwide, generating widespread die-off and subsequent community recovery dominated by remnant-patch dynamics rather than open-gap dynamics. Whether communities can recover and, if so, which factors mediate recolonization rate and extent remain unresolved. Here we evaluate recolonization dynamics of southern U.S. salt marshes that experienced extensive, drought-induced die-off of the foundation species Spartina alterniflora over the previous decade. Surveys of Georgia (USA) salt marshes showed little seedling recruitment in die-off areas but persistence of Spartina particularly in large, rather than small, remnant patches. Given this natural variation in remnant patch size, we conducted field experiments to test whether key plant-controlling biotic (grazing, plant neighbor presence) and abiotic (water availability) factors differentially impact Spartina recolonization at small and large-patch scales. In the small-patch (< 1 m2) experiment in 2009, removing grazers and plant neighbors prompted dramatically higher expansion and growth of Spartina relative to controls, while adding freshwater to reduce water limitation had little effect. In contrast, large-patch (> 20 m2 borders advanced significantly over the same time period regardless of grazer or neighbor removal. We continued the large-patch experiments in 2010, a year that experienced drought, and also added freshwater or salt to borders to modify ambient drought stress; overall, borders advanced less than the previous year but significantly more where neighbors were removed or freshwater added. Thus, water availability appears to mediate Spartina recovery by fueling large-patch expansion during wet summers and intensifying interspecific competition during drought. Combined, these findings suggest ecosystems can recover from massive disturbance if remnant foundation species' patches are large enough to overcome biotic inhibition and successfully expand during periods of relaxed abiotic stress.     

Individual recognition of neighbors by song in a suboscine bird, the alder flycatcher Empidonax alnorum

The ability of territorial males to discriminate between songs of their neighbors and songs of strangers has been demonstrated in 27 species of songbirds. Such experiments test only the ability of a subject to discriminate between two classes of stimuli, familiar (neighbors) and unfamiliar (strangers) songs. Individual recognition of neighbors is a finer, more complex type of discrimination. The ability of territorial males to recognize individual neighbors by song has been documented in 12 species of oscine passerines (Passeriformes, Passeri), but has never been demonstrated in suboscine passerines (Tyranni). We investigated recognition of songs of individual neighbors in a suboscine, the alder flycatcher (Empidonax alnorum). We performed a series of song playback experiments and recorded responses of territorial males to songs of neighbors and songs of strangers broadcast from two locations, the neighbor boundary and an opposite boundary. Subjects responded more aggressively to songs of a neighbor when played from the opposite boundary than when played from the neighbor boundary. They responded with equal aggression to songs of strangers regardless of location of playback. The difference in response to neighbor songs between speaker locations and the lack of a difference in response to stranger songs indicate that territorial males associate a particular song with a particular location (territory), and thus recognize individual neighbors.Communicated by I. Hartley     

Predator detection and dilution as benefits of associations between yellow mongooses and Cape ground squirrels

Associations among organisms are thought to form because the benefits, such as increased foraging efficiency or decreased risk of predation, outweigh any costs, such as resource competition. Though many interspecific associations have been described for closely related mammals, few studies have examined the associations between mammals in different orders. The yellow mongoose (Cynictus pencillata), a carnivore, and the Cape ground squirrel (Xerus inauris), a rodent, co-occur in arid and semi-arid South Africa where they share sleeping burrows, predators, a similar body size, and the capability to emit alarm calls in response to predators. To investigate enhanced predator avoidance as a potential benefit explaining the persistence of this association, we assessed individual mongoose vigilance alone and with squirrels or other mongooses, and with varying interspecific group size, using field observations. We also tested for responses to conspecific and heterospecific alarm calls in both study species using playback experiments. The proportion of time mongoose individuals spent vigilant decreased in the presence of squirrels or other mongooses and was negatively correlated with interspecific group size; a similar pattern was previously shown for conspecific groups of Cape ground squirrels. These results are predicted by both the dilution and collective detection hypotheses. In addition, hetero- and conspecific alarm calls elicited vigilance responses in both species. These results suggest that both species can benefit from the collective detection and dilution arising from their interspecific association and that this interspecific association could be mutualistic.     

House sparrows selectively eject parasitic conspecific eggs and incur very low rejection costs

Most host species of obligate interspecific brood parasites are under strong selection because such parasitism, e.g., that involving evictor nestmates, is highly costly. Egg rejection is one of the most efficient host defences against avian brood parasites. Many hosts have thus evolved egg-recognition ability and rejection behaviour. However, this defensive mechanism has not evolved in most species where only intraspecific brood parasitism occurs, probably because (1) the eggs of conspecific females are very similar in appearance, making egg rejection less likely to emerge, and (2) such parasitism is frequently less costly than interspecific parasitism. Using a captive population of house sparrows (Passer domesticus) with a low breeding density, we here provide new evidence showing that this species actually has a fine capacity to discriminate conspecific eggs and to eject them (44.2% of foreign eggs ejected) while incurring very low rejection costs (4.2% of own eggs ejected). This result contradicts those previously found in high-density house sparrow populations in which very high rejection costs and very high clutch desertion rates were reported, probably as a consequence of intraspecific competition and infanticide provoked by the high breeding density. The house sparrow has only rarely been reported as the host of an interspecific brood parasite, which implies that it is a newly described example of an altricial species in which egg ejection has evolved and is maintained in response to intraspecific brood parasitism.