生态位分化与森林群落物种多样性维持研究展望

一直以来,生态学家和进化生物学家对森林群落物种多样格局及其形成机制持有不同的观点。虽然Robert Ricklefs将进化和生态过程整合的观点已经被群落生态学家广泛接受,但是区域物种进化历史以及局域群落微进化过程是否能够影响群落生态学过程以及这些过程如何影响群落结构和动态还有待商榷。经典的生态位理论同时强调了种间和种内生态位分化对群落多样性维持的影响。但是生态学家普遍认为种间差异足以代表群落内个体间的相互作用关系,并且由于进化过程导致的种内分化往往涉及较长的时间尺度,因此,虽然种内差异是自然选择的重要材料,物种对环境的适应性进化过程所导致的种内分化对群落构建的影响往往被生态学家所忽视。为此,通过回顾种间和个体生态位分化的研究历史,对两类研究分别进行简要阐述,强调在今后的群落生态学研究中需要考虑个体分化对局域群落构建的影响。     

Niche overlap in allotopic and syntopic populations of sexually interacting ground‐hopper species

Abstract There is accumulating evidence that sexual interactions among species (reproductive interference) could have dramatic effects for species’ coexistence. It has been shown that the fitness of individuals can be substantially reduced as a consequence of reproductive interference. This might subsequently lead to displacement of a species (sexual exclusion). On the other hand, some evolutionary and ecological mechanisms might enable species to coexist, such as the divergence of mate recognition systems (reproductive character displacement), habitat partitioning, clumped dispersion patterns or different colonization capabilities. We have previously shown that the two ground‐hopper species Tetrix subulata and Tetrix ceperoi interact sexually in the laboratory as well as in the field. At sites where both species co‐occur niche overlap was high, suggesting that coexistence is maintained by different niche breadths rather than by habitat partitioning. To test the hypothesis that habitat partitioning does not contribute to species’ coexistence, we examined whether allotopic and syntopic populations of these two species differ in niche overlap (competitive release). Our results show that niche overlap is higher in syntopic than in allotopic populations, suggesting that the site‐specific habitat structure (heterogeneity) has a stronger influence on microhabitat utilization than the presence of heterospecifics. Hence, our data do not support the hypothesis that habitat partitioning plays a substantial role for the coexistence of these sexually interacting species.     

Assembly rules of ground-foraging ant assemblages are contingent on disturbance, habitat and spatial scale

Aim A major endeavour of community ecology is documenting non‐random patterns in the composition and body size of coexisting species, and inferring the processes, or assembly rules, that may have given rise to the observed patterns. Such assembly rules include species sorting resulting from interspecific competition, aggregation at patchily distributed resources, and co‐evolutionary dynamics. However, for any given taxon, relatively little is known about how these patterns and processes change through time and vary with habitat type, disturbance history, and spatial scale. Here, we tested for non‐random patterns of species co‐occurrence and body size in assemblages of ground‐foraging ants and asked whether those patterns varied with habitat type, disturbance history, and spatial scale. Location Burned and unburned forests and fens in the Siskiyou Mountains of southern Oregon and northern California, USA. Methods We describe ground‐foraging ant assemblages sampled over two years in two discrete habitat types, namely Darlingtonia fens and upland forests. Half of these sites had been subject to a large‐scale, discrete disturbance – a major fire – in the year prior to our first sample. We used null model analyses to compare observed species co‐occurrence patterns and body‐size distributions in these assemblages with randomly generated assemblages unstructured by competition both within (i.e. at a local spatial scale) and among (i.e. at a regional scale) sites. Results At local spatial scales, species co‐occurrence patterns and body‐size ratios did not differ from randomness. At regional scales, co‐occurrence patterns were random or aggregated, and there was evidence for constant body‐size ratios of forest ants. Although these patterns varied between habitats and years, they did not differ between burned and unburned sites. Main conclusions Our results suggest that the operation of assembly rules depends on spatial scale and habitat type, but that it was not affected by disturbance history from fire.     

Niche differentiation depends on body size in a cichlid fish: a model system of a community structured according to size regularities

1. Communities of different species are often structured according to niche differentiation associated with competitive interactions. We show that similar principles may apply on an ecological time-scale when individuals of a species having a wide size variation compete for resources, using the Lake Tanganyika cichlid Lobochilotes labiatus (5-30 cm). This species has a mouth especially adapted to suck up invertebrates from rock crevices. 2. Individuals defended feeding territories against similar-sized conspecifics, but not against different-sized ones. Thus, territories of similar-sized fish rarely overlapped, but up to a total of seven individuals (of seven size-ranks) had broadly overlapping territories with dissimilar-sized individuals. Comparison with expectation from the null model demonstrated clearly that observed size ratios between adjacent size rank were determined non-randomly regardless of sexual combinations. 3. Larger individuals took larger prey types of larger average size, but more importantly used wider rock crevices from which to suck food than smaller individuals. We calculated pairwise values of Schoener's index of diet overlap C(d) and the values of Levin's index of diet breadth B(d) (prey type and prey size) and the same for the width of the rock crevices used for foraging (C(r) and B(r)). C(d) remained high among all combinations of the seven ranks. In contrast, C(r) declined strongly in combinations of adjacent ranks (to 0.27), and was low or zero among further different size ranks. This shows that fish with overlapping territories divided the food resources largely through foraging site partitioning. Accordingly, B(d) did not depend on the size difference to the nearest two coinhabiting fish, whereas B(r) did. 4. We conclude that this L. labiatus community is structured non-randomly: body size-dependent effects on foraging site usage result in competition with, and territorial exclusion of, similar-sized individuals, but not of dissimilar-sized individuals that were accepted as coinhabitants. Accordingly, mean body size ratios (large/small) between two adjacent ranks were consistently approximately 1.28 [standard deviation (SD) = 0.07, n = 104], while approximately 1.34 from the null model (SD = 0.34, n = 10 400 simulations). We discuss our results as an example of Hutchinson's rule, applied originally to size ratios of different species.     

Non-random interactions within and across guilds shape the potential to coexist in multi-trophic ecological communities

Theory posits that the persistence of species in ecological communities is shaped by their interactions within and across trophic guilds. However, we lack empirical evaluations of how the structure, strength and sign of biotic interactions drive the potential to coexist in diverse multi-trophic communities. Here, we model community feasibility domains, a theoretically informed measure of multi-species coexistence probability, from grassland communities comprising more than 45 species on average from three trophic guilds (plants, pollinators and herbivores). Contrary to our hypothesis, increasing community complexity, measured either as the number of guilds or community richness, did not decrease community feasibility. Rather, we observed that high degrees of species self-regulation and niche partitioning allow for maintaining larger levels of community feasibility and higher species persistence in more diverse communities. Our results show that biotic interactions within and across guilds are not random in nature and both structures significantly contribute to maintaining multi-trophic diversity.     

Niche partitioning among three montane ground‐dwelling pheasant species along multiple ecological dimensions

Pheasants (order Galliformes) are typical ground‐dwelling birds, having a large body size and weak flight abilities. Sympatric pheasants are expected to share narrower niche space and face more extensive interspecific competition. However, little work has been undertaken simultaneously to investigate niche partitioning among sympatric pheasant species across multiple ecological dimensions. We compared microhabitat use, activity pattern and foraging strategy of sympatric Blood Pheasant Ithaginis cruentus, Buff‐throated Partridge Tetraophasis szechenyii and White Eared‐pheasant Crossoptilon crossoptilon on the Qinghai‐Tibet Plateau, China, to identify potential interspecific niche partitioning along different ecological dimensions in the breeding season. We found that the Buff‐throated Partridge differed significantly from the other two species in microhabitat use, and the three species showed different foraging strategies. It is likely that niche partitioning reduced potential interspecific competition, thus facilitating the species’ stable coexistence. Our study provides practical evidence of multidimensional niche theory within sympatric ground‐dwelling pheasant species, emphasizing that species interactions and coexistence within a guild are often not uni‐dimensional. Given global conservation concern for maintaining bird diversity, we recommend further restriction of yak grazing in these species’ habitat.     

High resource overlap and small dietary differences are widespread in food-limited warbler (Parulidae) communities

Although both interspecific competition and coexistence mechanisms are central to ecological and evolutionary theory, past empirical studies have generally focused on simple (two-species) communities over short time periods. Experimental tests of these species interactions are challenging in complex study systems. Moreover, several studies of ‘imperfect generalists’, consistent with Liem's Paradox, raise questions about the ability of evolved species differences to partition niche space effectively when resources vary considerably across the annual cycle. Here we used a recently developed theoretical framework to combine past research on population-level processes with observational data on resource use to test for ongoing interspecific competition and understand the nature of resource overlap. We compared species diet overlaps and differences in several distinctive communities centred on a focal species, the American Redstart Setophaga ruticilla replicated both spatially and seasonally, in combination with documentation of population regulation to assess the ability of similar species to partition dietary niche space and limit interspecific competition. Our results document high dietary overlap in most of the communities studied, with only subtle differentiation consistent with known species differences in foraging behaviour and morphology. These findings are largely consistent with species foraging as imperfect generalists. However, in contrast to past studies, the high diet overlaps observed here during times of inferred resource scarcity were driven by low-value prey taxa (e.g. small ants) and did not involve truly ‘private’ resources. All of these factors increase the potential negative impacts of interspecific competition, and limit the ability of these birds to avoid competition if food availability deteriorates further than observed in our study, either seasonally or at longer intervals.     

Habitat disturbance modifies dominance,coexistence, and competitive interactions in tropical ant communities

1. Interspecific competition is a major structuring principle in ecological communities. Despite their prevalence, the outcome of competitive interactions is hard to predict, highly context-dependent, and multiple factors can modulate such interactions. 2. We tested predictions concerning how competitive interactions are modified by anthropogenic habitat disturbance in ground-foraging ant assemblages inhabiting fragmented Inter-Andean tropical dry forests in southwestern Colombia, and investigated ant assemblages recruiting to baits in 10 forest fragments exposed to varying level of human disturbance. 3. Specifically, we evaluated how different components of competitive interactions (patterns of species co-occurrence, resource partitioning, numerical dominance, and interspecific trade-offs between discovery and dominance competition) varied with level of habitat disturbance in a human-dominated ecosystem. 4. Multiple lines of evidence suggest that the role of competitive interactions in structuring ground-foraging ant communities at baits varied with respect to habitat disturbance. As disturbance increased, community structure was more likely to exhibit random co-occurrence patterns, higher levels of monopolization of food resources by dominant ants, and disproportionate dominance of a single species, the little fire ant (Wasmannia auropunctata). At a regional scale, we found evidence for a trade-off between dominance and discovery abilities of the 15 most common species at baits. 5. Together, these results suggest that human disturbance modifies the outcome of competitive interactions in ground-foraging ant assemblages and may promote dominant species that reduce diversity and coexistence in tropical ecosystems.     

Impact of human disturbance on temporal partitioning within carnivore communities

1. Interspecific competition is an important evolutionary force, influencing interactions between species and shaping the composition of biological communities. In mammalian carnivores, to reduce the risks of negative encounters between competitors, species can employ a strategy of temporal partitioning, adapting activity patterns to limit synchronous activity. This strategy of non-human competitor avoidance, however, may be influenced by the expansion of human activities, which has driven wild mammals towards nocturnality.
2. We hypothesise that the disruption of temporal niche partitioning by humans and their activities could increase temporal overlap between carnivores, enhancing interspecific competition.
3. We reviewed the published literature systematically and employed generalised linear models to evaluate quantitatively the relative influence of a range of human, meteorological and ecological variables on coefficients of temporal overlap within mammalian terrestrial carnivore communities (orders Carnivora and Didelphimorphia) on a global scale.
4. None of the models investigated showed evidence of an impact of humans on temporal partitioning between carnivores on a global scale. This illustrates that temporal avoidance of humans and competitors does not always follow a consistent pattern and that its strength may be context-dependent and relative to other dimensions of niche partitioning (spatial and trophic).
5. Similarly, the regulation of activity patterns may be strongly site-specific and may be influenced by a combination of biotic and abiotic characteristics. Temporal avoidance of both humans and competitors by carnivores may take the form of short, reactive responses that do not impact activity patterns in the longer term.
6. Although we did not detect a global disruption of temporal partitioning due to human disturbance, carnivore communities may still experience an increase in interspecific competition in other niche dimensions. Further research would benefit from using controlled experimental designs and investigating multiple dimensions of niche partitioning simultaneously. Finally, we recommend complementing the coefficient of temporal overlap with other metrics of fine-scale spatiotemporal interactions.

     

Diet overlap among three sympatric African annual killifish species Nothobranchius spp. from Mozambique

The diet patterns of three Nothobranchius species (N. furzeri, N. orthonotus and N. rachovii), small, short‐lived annual killifish from temporary pools in African savannah were investigated. Four sites with contrasting fish density and water surface area were sampled in 2008 and 2009 in southern Mozambique. Stomach content analysis showed that all the species examined were generalists, with diets largely based on aquatic invertebrates. The same invertebrate prey categories were consumed by all three species, but their relative proportions varied across species. The largest species, N. orthonotus, showed the most distinct diet and consumed vertebrates (juvenile lungfish Protopterus annectens and larval Amphibia) and a relatively high proportion of Odonata, Coleoptera and Ephemeroptera larvae. The diet of the other two species (N. furzeri and N. rachovii) showed a stronger overlap, did not include vertebrates, but was rich in small crustaceans (Cladocera, Copepoda, Ostracoda and Conchostraca). Mosquito (Diptera) larvae formed only a negligible part of the diet of all the three species.     

Habitat Selection,the Included Niche,and Coexistence in Plant‐Specialist Frogs from Madagascar

Classic niche partitioning suggests that coexistence among asymmetric competitors is facilitated by differential resource use. Coexistence is also possible, however, when a species only has access to resources that are shared with a competitor, providing it is the superior competitor on that resource (the ‘included niche’). To test predictions of these two coexistence mechanisms, we studied habitat selection of two closely related sympatric plant‐specialist frogs from Madagascar (Guibemantis bicalcaratus and Guibemantis punctatus). Both species live and breed only in the water‐filled leaf axils of Pandanus plants, and previous experiments with their tadpoles demonstrated asymmetric competition. In a 3‐yr field study, we: (1) monitored the biotic and abiotic conditions of 348 plants; (2) surveyed these plants for frogs; (3) undertook a mark‐recapture study; and (4) conducted an experiment where we manipulated the amount of detritus in plants. We identified several differences in the conditions of the habitats selected by both species (e.g., plant height, canopy cover over the plant, abundance of heterospecifics). Co‐occurrence was nevertheless common, and G. punctatus persisted almost exclusively in plants it shared with G. bicalcaratus. As predicted by theory, G. punctatus was the superior competitor, at least in the tadpole stage. The inferior competitor (G. bicalcaratus) had access to exclusive resources not available to G. punctatus, by virtue of a faster developmental rate that permitted reproduction in lower quality plants. While there was some evidence of weak niche partitioning, we conclude that coexistence in these plant‐specialist frogs is primarily via an included niche mechanism. Abstract in French is available at http://www.blackwell‐synergy.com/loi/btp .     

Spatio-temporal patterns of ground-dwelling ant assemblages in a lowland rainforest in southeast Cameroon

We studied the spatio-temporal patterns of ground-dwelling and -foraging ant assemblages in a continuous mixed secondary lowland rainforest on the northern periphery of the Dja Biosphere Reserve in southeast Cameroon. The effect of season and of the interaction of season with vegetation type on ant species density, activity and composition were investigated. We also checked for a possible impact of army ant activity on the ant assemblage. Ants were collected by pitfall trapping in seven vegetation types in each of three seasons during 12 months in 2003 and 2004–2005. Season significantly influenced the ground-dwelling ant assemblage. The late wet season had the lowest ant species density and a different species composition than the early wet and the dry season. Also inter-annual variation in ant species density and composition was detected and could partly be explained by an influence of Dorylus activity. In general, vegetation type and inter-annual variation had a larger influence on the ground-ant assemblage than season. Significant effects of interactions between season, year, vegetation type and army ants on the ant assemblage confirm that also in African rainforests ant assemblages are extremely dynamic and depend on a complex combination of availability of food resources, nest sites and predators. Received 4 October 2006; revised 4 June 2007; accepted 2 July 2007.     

OBSERVATIONS ON THE BREEDING OF THE WOOLLYNECKED STORK

Scott, J. A. 1975. Observations on the breeding of the Woollynecked Stork. Ostrich 46: 201–207.

Little is known about the breeding of the Woollynecked Stork Ciconia episcopus in Africa. This paper discusses breeding, adult and nestling behaviour, nests and sites. Seasonal movements are discussed briefly. Eight nests were studied during 1970 to 1974. At one nest incubation was established at 30 to 31 days and the fledging period 55 to 65 days. No feeding of the young was observed at any time, though one eight hour observation period was undertaken. Few mating displays were seen and none away from the nest.     

重庆特有极小种群植物缙云秋海棠生态位特征研究

该研究通过对重庆缙云山国家级自然保护区的缙云秋海棠(Begonia jinyunensis)及其伴生物种所在群落进行实地调查,并以样地作为一维资源轴,物种重要值作为生态位计测状态指标,采用Levins、Hurlbert、Pianka公式对缙云秋海棠种群的生态位特征进行分析,以阐明该群落物种间对资源的利用和竞争关系,为缙云秋海棠的保护提供依据。结果表明:(1)在群落中缙云秋海棠的Levins和Hurlbert生态位宽度均最大,分别为6.742 40和0.717 80,表明缙云秋海棠对环境资源的利用能力强,生态适应幅度也较大。(2)研究区内缙云秋海棠的伴生物种中生态位宽度(Ba)大于0.3的3种植物为石生楼梯草(Elatostema rupestre)、斜方复叶耳蕨(Arachniodes rhomboidea)、对马耳蕨(Polystichum tsus-simense),其Ba值分别为0.460 00、0.417 04、0.301 60,表明缙云秋海棠的3个主要伴生种在群落中具有较强的竞争力。(3)20个伴生物种中与缙云秋海棠生态位重叠值最高的是石生楼梯草(0.671 4),说明缙云秋海棠与石生楼梯具有相似的环境要求,资源不足时,两者可能存在比较激烈的竞争,其中有12个伴生物种(60%)与缙云秋海棠的生态位重叠值小于0.5,群落中大多数物种与缙云秋海棠的生态位重叠值均较低,说明物种之间对资源的需求不同,环境适应能力相似度低,群落中的种间竞争不激烈。研究认为,缙云秋海棠生态位宽度大,生境适宜生存,种间竞争不激烈,即目前生态位因素不是造成其处于濒危状态的主要原因;导致缙云秋海棠数量少,呈现岛屿状分布的主要原因可能是其生境遭到人为破坏。     

Food utilization of aphidophagous hoverfly larvae (Diptera: Syrphidae, Chamaemyiidae) on herbaceous plants in an urban habitat

We examined food utilization in a community of aphidophagous hoverfly larvae (Diptera: Syrphidae and Chamaemyiidae) in open lands in an urban habitat in central Japan for 3 years. The community consisted of 17 hoverfly species feeding on 20 aphid species occurring on 14 species of dominant herbaceous plants. In terms of larval prey preference, the dominant eight species of hoverfly were categorized into three groups: a polyphagous ‘generalist’ group consisting of four species,Episyrphus balteatus, Betasyrphus serarius, Syrphus vitripennis andSphaerophoria sp.; an oligophagous ‘specialist’ group consisting of three species,Metasyrphus hakiensis, Dideoides latus andParagus hemorrhous; andLeucopis puncticornis, which showed a preference for two aphid species on the plantTorilis scabra. The prey aphids of the second group have behavioral or morphological defense mechanisms that are effective for preventing attacks by generalist hoverflies; two prey aphids are aggressive toward generalist predators and the others are protected by ant-attendance. The specialist hoverflies seem to be adapted to overcome these defense mechanisms. The prey ranges overlapped little between the generalist and the specialist groups, while those within the generalist group overlapped greatly.     

Experimental evidence that the introduced fire ant, Solenopsis invicta, does not competitively suppress co-occurring ants in a disturbed habitat

1. The fire ant, Solenopsis invicta, is a globally distributed invasive ant that is largely restricted to disturbed habitats in its introduced range. For more than half a century, biologists have believed its success results from superior competitive abilities relative to native ant species, as well as an escape from their natural enemies. 2. We used large volumes of hot water to kill fire ant colonies, and only fire ant colonies, on experimental plots in pastures, and found that populations and diversity of co-occurring ants did not subsequently increase. 3. These results are contrary to classical predictions and indicate that S. invicta is not a superior competitor that suppresses native ants, and that the low diversity and abundance of native ants in degraded ecosystems does not result from interaction with fire ants. Instead, other factors such as prior disturbance and recruitment limitation may be the primary limiting factors for native species in these habitats.     

Niche separation and weak interactions in the high tidal zone of saltmarsh‐mangrove mixing communities

1. Saltmarsh‐mangrove ecotones occur at the boundary of the natural geographic distribution of mangroves and salt marshes. Climate warming and species invasion can also drive the formation of saltmarsh‐mangrove mixing communities. How these coastal species live together in a “new” mixed community is important in predicting the dynamic of saltmarsh‐mangrove ecosystems as affected by ongoing climate change or human activities. To date, the understanding of species interactions has been rare on adult species in these ecotones.
2. Two typical coastal wetlands were selected as cases to understand how mangrove and saltmarsh species living together in the ecotones. The leaves of seven species were sampled from these coastal wetlands based on their distribution patterns (living alone or coexisting) in the high tidal zone, and seven commonly used functional traits of these species were analyzed.
3. We found niche separation between saltmarsh and mangrove species, which is probably due to the different adaptive strategies they adopted to deal with intertidal environments.
4. Weak interactions between coexisting species were dominated in the high tidal zone of the two saltmarsh‐mangrove communities, which could be driven by both niche differentiation and neutral theory.
5. Synthesis. Our field study implies a potential opportunity to establish a multispecies community in the high tidal zone of saltmarsh‐mangrove ecotones, where the sediment was characterized by low salinity and high nitrogen.

     

Understanding primate communities: Recent developments and future directions

In 1999, the edited volume Primate Communities presented several studies that examined broad‐scale patterns of primate diversity.¹ Similar studies were being conducted on nonprimate taxa; advances in data availability and statistical approaches were allowing scientists to investigate a variety of new questions and to reexamine classical questions in novel ways. While such studies on nonprimate taxa have continued at a steady pace, they have only crept forward for primate species (Fig. 1 ). In the intervening time, the field of macroecology (Box 1) rapidly developed and has resulted in several books 2 - 4 and the establishment of new research institutes. We suggest that examining primate communities, especially in a macroecological context, is an important line of research for our field to embrace and an area where biological anthropologists can provide major contributions. We review the current state of research, describe new datasets and research tools, and suggest future research directions.     

Intraspecific and interspecific trait variability in tadpole meta‐communities from the Brazilian Atlantic rainforest

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