Seed predation by Neotropical rain forest mammals increases diversity in seedling recruitment

Seed dispersal and seedling recruitment (the transition of seeds to seedlings) set the spatiotemporal distribution of new individuals in plant communities. Many terrestrial rain forest mammals consume post-dispersal seeds and seedlings, often inflicting density-dependent mortality. In part because of density-dependent mortality, diversity often increases during seedling recruitment, making it a critical stage for species coexistence. We determined how mammalian predators, adult tree abundance, and seed mass interact to affect seedling recruitment in a western Amazonian rain forest. We used exclosures that were selectively permeable to three size classes of mammals: mice and spiny rats (weighing <1 kg), medium-sized rodents (1-12 kg), and large mammals (20-200 kg). Into each exclosure, we placed seeds of 13 tree species and one canopy liana, which varied by an order of magnitude in adult abundance and seed mass. We followed the fates of the seeds and resulting seedlings for at least 17 months. We assessed the effect of each mammalian size class on seed survival, seedling survival and growth, and the density and diversity of the seedlings that survived to the end of the experiment. Surprisingly, large mammals had no detectable effect at any stage of seedling recruitment. In contrast, small- and medium-sized mammals significantly reduced seed survival, seedling survival, and seedling density. Furthermore, predation by small mammals increased species richness on a per-stem basis. This increase in diversity resulted from their disproportionately intense predation on common species and large-seeded species. Small mammals thereby generated a rare-species advantage in seedling recruitment, the critical ingredient for frequency dependence. Predation by small (and to a lesser extent, medium-sized) mammals on seeds and seedlings significantly increases tree species diversity in tropical forests. This is the first long-term study to dissect the effects of various mammalian predators on the recruitment of a diverse set of tree species.     

澳门松山市政公园物种多样性特征

根据实际情况.将澳门松山市政公园植物群落分为阴香群落、假萍婆群落、假萍婆+石栗群落,调查并分析了各群落的物种多样性特征.结果表明,乔木层、灌木层和草本层的物种多样性水平表现为灌木层较高.草本层与乔木层较低.但总体而言.这些群落的植物种类都较少,物种多样性水平尤其乔木层物种多样性水平低.因此,应加强抚育,可通过生态恢复改善环境条件,为生物多样性的发展创造条件;通过林分改造促进群落物种多样性水平的提高,同时也能丰富森林景观.     

Species dispersal rates alter diversity and ecosystem stability in pond metacommunities

Metacommunity theory suggests that relationships between diversity and ecosystem stability can be determined by the rate of species dispersal among local communities. The predicted relationships, however, may depend upon the relative strength of local environmental processes and disturbance. Here we evaluate the role of dispersal frequency and local predation perturbations in affecting patterns of diversity and stability in pond plankton metacommunities. Pond metacommunities were composed of three mesocosm communities: one of the three communities maintained constant "press" predation from a selective predator, bluegill sunfish (Lepomis macrochirus); the second community maintained "press" conditions without predation; and the third community experienced recurrent "pulsed" predation from bluegill sunfish. The triads of pond communities were connected at either no, low (0.7%/d), or high (20%/d) planktonic dispersal. Richness and composition of zooplankton and stability of plankton biomass and ecosystem productivity were measured at local and regional spatial scales. Dispersal significantly affected diversity such that local and regional biotas at the low dispersal rate maintained the greatest number of species. The unimodal local dispersal-diversity relationship was predator-dependent, however, as selective press predation excluded species regardless of dispersal. Further, there was no effect of dispersal on beta diversity because predation generated local conditions that selected for distinct community assemblages. Spatial and temporal ecosystem stability responded to dispersal frequency but not predation. Low dispersal destabilized the spatial stability of producer biomass but stabilized temporal ecosystem productivity. The results indicate that selective predation can prevent species augmentation from mass effects but has no apparent influence on stability. Dispersal rates, in contrast, can have significant effects on both species diversity and ecosystem stability at multiple spatial scales in metacommunities.     

Species diversity reduces parasite infection through cross-generational effects on host abundance

With growing interest in the effects of biodiversity on disease, there is a critical need for studies that empirically identify the mechanisms underlying the diversity-disease relationship. Here, we combined wetland surveys of host community structure with mechanistic experiments involving a multi-host parasite to evaluate competing explanations for the dilution effect. Sampling of 320 wetlands in California indicated that snail host communities were strongly nested, with competent hosts for the trematode Ribeiroia ondatrae predominating in low-richness assemblages and unsuitable hosts increasingly present in more diverse communities. Moreover, competent host density was negatively associated with increases in snail species richness. These patterns in host community assembly support a key prerequisite underlying the dilution effect. Results of multigenerational mesocosm experiments designed to mimic field-observed community assemblages allowed us to evaluate the relative importance of host density and diversity in influencing parasite infection success. Increases in snail species richness (from one to four species) had sharply negative effects on the density of infected hosts (-90% reduction). However, this effect was indirect; competition associated with non-host species led to a 95% reduction in host density (susceptible host regulation), owing primarily to a reduction in host reproduction. Among susceptible hosts, there were no differences in infection prevalence as a function of community structure, indicating a lack of support for a direct effect of diversity on infection (encounter reduction). In monospecific conditions, higher initial host densities increased infection among adult hosts; however, compensatory reproduction in the low-density treatments equalized the final number of infected hosts by the next generation, underscoring the relevance of multigenerational studies in understanding the dilution effect. These findings highlight the role of interspecific competition in mediating the relationship between species richness and parasite infection and emphasize the importance of field-informed experimental research in understanding mechanisms underlying the diversity-disease relationship.     

Species diversity and faunal affinities of meiobenthic Copepoda in the deep sea

This first attempt at measuring species diversity of the numerically abundant marine meiobenthos on a vertical gradient into the deep sea supports the stability-time hypothesis developed with the macrofauna. Diversity, as measured by rarefaction, indicated highly evolved, stable and unique assemblages in the deep sea. Shelf samples were consistently dominated by a few species; below 1000 m many of the species were new and few were duplicated from sample to sample. Rarefaction curves are clearly separable for the three environments, with the vertical component of diversity increasing to abyssal depths. Rarefaction analysis of shelf samples above and below the Cape Hatteras zoogeographical boundary further revealed distinct spilits in the faunal assemblages. Faunal affinity analyses indicate that the deep-sea fauna is not homogenously distributed over wide expanses of the ocean floor and are contrary to previously reported macrofaunal findings.     

茂兰喀斯特原生林不同地球化学环境的植物物种多样性

中国西南岩溶区是中国生物多样性分布中心之一。该区生态系统不断退化,生物多样性大量丧失,不同喀斯特地球化学背景下植被恢复的速度与成林性质存在明显差异。我们在最有代表性之一的贵州茂兰喀斯特森林保护区选取纯灰岩和纯白云岩两种岩性的喀斯特环境,对其元素地球化学特征及其中原生性植物群落的相异性和物种多样性进行了对比,结果发现:①不同岩性区域多样性:乔木层和灌木层为纯灰岩区>纯白云岩区,草本层为纯白云岩区>纯灰岩区,总体上纯灰岩区的高于纯白云岩区的;②各样地间的相似性系数都很低,相同岩性类型内的明显高于不同岩性类型之间的相似度;物种周转率高,且白云岩区高于石灰岩区,种类组成差异明显;③喀斯特区可溶岩地球化学背景通过喀斯特作用导致特殊的地形条件与元素地球化学特征,并引起局部小生境的分异从而影响植物群落特征。     

九龙山南方铁杉群落物种多样性及乔木种种间联结性

采用物种丰富度指数、Shannon-Weiner指数、Simpson指数、Pielou均匀度指数研究南方铁杉(Tsuga tchekiangensis)群落物种多样性.同时在2×2联列表基础上,应用x2检验、联结系数AC、种间共同出现的百分率PC研究南方铁杉群落19个乔木种群的种间联结性.结果表明:(1)物种丰富度的大小均表现为灌木层大于乔木层,乔木层中物种丰富度为乔木中层>乔木下层>乔木上层.(2)乔木层、乔木层上层和乔木层中层的Shannon-Weiner指数、Simpson指数、Pielou均匀度指数在1 300～1 500 m随海拔升高而增大,在1 500～1 620 m随海拔升高而减小,物种随海拔梯度变化呈现"中间高度膨胀"现象;乔木下层和灌木层则表现为相反规律.(3)x2统计度量值表明,在162个种对中,147对联结关系未达到显著水平,种间的独立性较大;共同出现的百分率PC表明多数种对之间联结程度不显著,南方铁杉、猴头杜鹃(Rhododendron simoarum)、木荷(Schima superba)三者联结是比较紧密的;联结系数AC表明,南方铁杉与厚皮香(Temstroemia gymnanthe)、多脉冬青(Ilexpolyneura)、红花油茶(Camellia chekiang-oleosa)、浙江樟(Cinnamomum chekiangense)、青冈栎(Cyclobalanopsis glauc)等种对存在显著负联结,表明这几个树种对南方铁杉的生长存在重要影响,在营造林工林时应重点加于考虑.     

洞庭湖湿地珍稀濒危鸟类群落组成及多样性

2005年7至2007年4月,采用直接记数法和"GPS"技术定位监测对洞庭湖湿地珍稀濒危保护鸟类资源及其环境进行调查。结果表明:洞庭湖湿地珍稀濒危保护鸟类有218种,隶属于16目46科。其中古北界鸟类占优势,有118种,东洋界鸟类54,广布种鸟类46种。冬候鸟为主体,有118种,留鸟40种,夏侯鸟35种,旅鸟25种。按生态类群分,游禽45种、涉禽69种、陆禽3种、猛禽22种、攀禽11种、鸣禽68种;该湿地有国家Ⅰ级重点保护鸟类7种、Ⅱ级重点保护鸟类43种,国际湿地公约名录中的鸟类37种。洞庭湖湿地珍稀濒危保护鸟类群落Shannon-Wiener多样性指数和Pielou均匀性指数分别为4.489 7和0.576 5。     

Habitat structure affects intraguild predation

Intraguild predation is thought to be ubiquitous in natural food webs. Yet, theory on intraguild predation predicts the intraguild prey to persist only under limited conditions. This gap between theory and empirical observations needs scrutiny. One reason might be that theory has focused on equilibrium dynamics and a limited set of species (usually three) that interact in well-mixed populations in unstructured habitats, and these assumptions will often not hold in natural systems. In this review, we focus on the effects of habitat structure on intraguild predation. Habitat structure could reduce encounter rates between predators and prey and could create refuges for prey. In both cases, habitat structure could reduce the strength of intraguild interactions, thereby facilitating species coexistence. A meta-analysis of studies on manipulation of habitat structure shows that intraguild prey indeed suffer less from intraguild predation in structured habitats. This was further confirmed by a meta-analysis in which studies on intraguild predation were classified according to habitat structure. Intraguild predation reduced densities of the intraguild prey significantly more in habitats with little structure than in habitats rich in structure. The effect of intraguild predation on the shared prey was negative, and not significantly affected by habitat structure. We conclude that habitat structure may increase persistence of the intraguild prey by decreasing the strength of the interaction between intraguild predator and intraguild prey.     

古田山国家自然保护区针阔叶混交林植物物种多样性特征

利用样方法进行野外群落调查.共调查了6个样地,每个样地大小为20m×20 m,研究古田山自然保护区针阔叶混交林植物物种多样性特征.结果表明:古田山针阔混交林的科、属、种组成多样性较高,植物种类丰富.组成复杂.群落的属种多样性>科属多样性>科种多样性.从灌木层、乔木层至草本层物种多样性依次递减,乔木层和灌木层的物种多样性指数明显大于草本层物种多样性指数.     

Ideal free distributions under predation risk

 We examine the trade-off between gathering food and avoiding predation in the context of patch use by a group of animals. Often a forager will have to choose between feeding sites that differ in both energetic gain rate and predation risk. The ideal site will have a high gain rate and low risk of predation. However, intake rate will often decrease when the patch is shared with other foragers and it may be optimal for some individuals to feed elsewhere. Within the framework of ideal free theory, we investigate the distribution of foragers that will equalise individual fitness gains. We focus on a two-patch environment with continuous inputs of food. With reference to existing experimental studies, we examine the effects of risk dilution, food input rates and an animal’s expectations of the future. We identify the effect of total animal numbers when one patch is subject to predation risk and the other is safe. Conditions under which the difference in intake rate in the two patches is constant are identified, as are conditions in which the ratio of animals in the two patches is constant. If current conditions do not alter future expectations an increase in input rates to the patches promotes increased use of the risky patch. Yet, if conditions are assumed to persist indefinitely the opposite effect is seen. When both patches are subject to predation risk, dilution of risk favours more extreme distributions, and may lead to more than one stable distribution. The results of these models are used to critically analyse previous work on the energetic equivalence of risk. This paper is intended to help guide the development of new experimental studies into the energy-risk trade-off. Received: 10 February 1995/Accepted after revision: 1 October 1995     

闽西北地区不同林龄常绿阔叶混交林物种多样性比较

基于野外样方数据,按照林龄与垂直分布层次,分别采用Margalef丰富度指数（Dm）、Pielou均匀度指数（J）与生态优势度指数（C）计算式,对闽西北地区主要优势种为米槠（Castanopsis carlesii）、栲树(Castanopsis fargesii)、甜槠（Cadtanopsis eyrei）、青冈（Quercus glauca）等壳斗科植物为主的天然常绿阔叶混交林群落的物种多样性进行了调查与对比分析。结果表明：（1）不同龄组常绿阔叶混交林乔木层的物种多样性存在显著差异（P<0.05）,以中龄林最为丰富,近熟林与成熟林次之,幼龄林最低,各龄组间的物种分布的均匀度与优势度差异不明显（P﹥0.05）,近熟林J值最大,幼龄林C值最大;（2）不同龄组常绿阔叶混交林灌木层间的Dm、J、C值差异显著（P<0.05）,以中龄林与成熟林内物种较为丰富,近熟林次之,幼龄林最低,物种分布以幼龄林内最为不均;（3）不同龄组常绿阔叶混交林草本层间的Dm、J、C差异显著（P<0.05）,物种多样性以成熟林内最为丰富,幼龄林次之,近熟林内最低,成熟林与幼龄林内草本层物种分布较均匀,中龄林与近熟林内则分布较集中,其物种优势度较高;（4）不同龄组常绿阔叶混交林乔、灌、草层间的物种多样性不同。在幼龄林中,Dm值大小顺序为：乔木层>草本层>灌木层,J值为草本层>乔木层>灌木层。中龄林、近熟林、成熟林的Dm值大小顺序均为：乔木层>灌木层>草本层,J值为灌木层>乔木层>草本层。研究结果可为了解闽西北地区不同龄组常绿阔叶混交林群落结构的物种组成与动态变化特征,探讨常绿阔叶混交林近自然经营及其残次林改造技术,制定其生物多样性的生态保护决策等提供理论依据。     

The role of predation in ecosystem models

Ecosystem models must convert a naturally open system into a theoretically closed one. For phytoplankton-herbivore models the form for the parameterization of predation on the herbivores can have a significant effect on the remainder of the model.     

Feeding under predation hazard: response of the guppy and Hart's rivulus from sites with contrasting predation hazard

Summary Populations of guppies, Poecilia reticulata and Hart's rivulus, Rivulus harti, in Trinidad experience different levels of predation hazard from piscivorous fish. Those from the larger rivers (downstream sites) experience chronically high predation hazard, while those from headwater streams (upstream sites) have few predators. Guppies and Hart's rivulus, collected from downstream and upstream sites, were assayed for their feeding rate in the presence and absence of predators. We defined tenacity as the ratio of the feeding rates in the presence and absence of a predator stimulus. Thus, tenacity expresses the degree to which the forager maintains its feeding rate when a predator stimulus is present. Previous work by Seghers (1973) showed that non-feeding guppies from downstream sites responded more strongly to predators than did guppies from upstream sites. Based on this, we initially hypothesized that fish from downstream sites would show lower tenacities than fish from upstream sites. However, we found the opposite in every case. When confronted with a predator stimulus, guppies and Hart's rivulus from downstream sites fed at consistently greater rates and displayed greater tenacities than did those from upstream sites. These differences were found in experiments using both live and model predators. The results suggest that upstream fish readily trade off feeding for hiding and avoiding predation hazard, a likely response when predators appear infrequently, while downstream fish appear to be selected for boldness and tenacity while foraging under chronically high hazard.     

Fish predation selects for reduced foraging activity

Despite the importance of foraging activity for the growth/predation risk trade-off, studies that demonstrated predator-induced survival selection on foraging activity under semi-natural conditions are relatively rare. Here, we tested for fish-induced selection for reduced foraging activity in two larval Enallagma damselflies using a field enclosure experiment. Fish imposed considerable mortality in both damselfly species and survival selection on foraging activity could be detected in Enallagma geminatum. We did not detect selection in Enallagma hageni, probably because this species already was not eating very much in the absence of fish compared to E. geminatum. Both species responded strongly to the presence of predators by reducing their foraging activity. The documented survival selection on foraging activity was detected despite the already low activity levels in fish lake prey species and despite strong predator-induced plasticity in this trait.     

Linking fruit traits to variation in predispersal vertebrate seed predation, insect seed predation, and pathogen attack

The importance of vertebrates, invertebrates, and pathogens for plant communities has long been recognized, but their absolute and relative importance in early recruitment of multiple coexisting tropical plant species has not been quantified. Further, little is known about the relationship of fruit traits to seed mortality due to natural enemies in tropical plants. To investigate the influences of vertebrates, invertebrates, and pathogens on reproduction of seven canopy plant species varying in fruit traits, we quantified reductions in fruit development and seed germination due to vertebrates, invertebrates, and fungal pathogens through experimental removal of these enemies using canopy exclosures, insecticide, and fungicide, respectively. We also measured morphological fruit traits hypothesized to mediate interactions of plants with natural enemies of seeds. Vertebrates, invertebrates, and fungi differentially affected predispersal seed mortality depending on the plant species. Fruit morphology explained some variation among species; species with larger fruit and less physical protection surrounding seeds exhibited greater negative effects of fungi on fruit development and germination and experienced reduced seed survival integrated over fruit development and germination in response to vertebrates. Within species, variation in seed size also contributed to variation in natural enemy effects on seed viability. Further, seedling growth was higher for seeds that developed in vertebrate exclosures for Anacardium excelsum and under the fungicide treatment for Castilla elastica, suggesting that predispersal effects of natural enemies may carry through to the seedling stage. This is the first experimental test of the relative effects of vertebrates, invertebrates, and pathogens on seed survival in the canopy. This study motivates further investigation to determine the generality of our results for plant communities. If there is strong variation in natural enemy attack among species related to differences in fruit morphology, then quantification of fruit traits will aid in predicting the outcomes of interactions between plants and their natural enemies. This is particularly important in tropical forests, where high species diversity makes it logistically impossible to study every plant life history stage of every species.     

Avoiding predation: alarm responses of Caribbean sea urchins to simulated predation on conspecific and heterospecific sea urchins

Alarm responses to the extracts of conspecifics and hetero-specifics were measured for the Caribbean sea urchins Echinometra viridis, E. lucunter, Lytechinus variegatus, L. williamsi, Tripneustes ventricosus, Diadema antillarum, and Eucidaris tribuloides collected along the Caribbean coast (9°3314N; 78°5523W) during October 1984 and July–December 1985. Responses to seawater and extracts of the gnathostomate echinoid Clypeaster sybdepressus were used as controls. The intensity of the response resulting from exposure to sea-urchin extracts was measured by: (1) the percentage of individuals that responded by moving away from the extract and/or towards shelter, and (2) the mean distance moved. Echinometra viridis, E. lucunter, and L. williamsi responded to sea-urchin extracts by moving towards nearby shelter sites. The distance that individuals of each species moved in the first minute following exposure to conspecific extracts was correlated with the distance that species moved from shelter while foraging. L. variegatus and D. antillarum, living in microhabitats not providing protection from predators, responded to extracts of conspecifics and heterospecifics by moving away from the direction of the extract. Eucidaris tribuloides did not exhibit alarm responses to the extracts of con- or hetero-specifics. E. tribuloides secures itself with its stout spines into protected sites within corals. Similarly, L. variegatus living in long, dense seagrass that provided protection from detection by predators, and D. antillarum occupying crevices, showed no alarm responses to extracts of conspecifics. Presumably, in these situations, sea urchins cannot increase their defenses against predation by moving away from an injured neighbor. T. ventricosus showed a weak response to extracts of L. variegatus, but no response to extracts of other species including conspecifics. The reasons for this lack of an alarm response are unclear. For the five species that demonstrated an alarm response to sea-urchin extracts, the intensity of the response varied depending on the type of extract used. L. variegatus, L. williamsi, and D. antillarum responded most strongly to extracts from conspecifics, while Echinometra viridis and E. lucunter responded strongly to extracts from both conspecifics and congeners. The weakest responses were shown to the extracts of T. ventricosus and Eucidaris tribuloides. Habitat overlap, overlap in predators, and phylogenetic relationships did not consistently explain patterns of alarm responses to the extracts of heterospecific sea urchins.     

Uniform predation risk in nature: common,inconspicuous, and a source of error to predation risk experiments

Previous studies indicate that when predation risk is uniform across habitats, foragers concentrate their exploitation in fewer patches. Although uniform predation risk may seem rare in nature, some scenarios might cause it. Testing all scenarios in a single experiment is unfeasible; therefore, we developed a model that points whether concentration of exploitation in specific habitats due to uniform risk requires parameter values similar to what is found in literature. This model was based on Brown’s (Behav Ecol Sociobiol 22:37–47, 1988) fitness function but rescaled to multiple habitats and predators, including uniform risk predators. Deriving function’s maximum allowed comparisons with giving-up density studies. Results showed that uniform predation risk had a u-shaped effect in habitat exploitation, causing a concentration of habitat exploitation at probabilities of survival from 0.2 to 0.8. However, the length of this interval and degree of concentration depended on the value of safety to forager fitness. Heterogeneous, nonuniform, predation risk decreases habitat exploitation where it was higher, therefore suppressing the effect of uniform risk on prey behavior. Time spent in the focal habitat and metabolic costs reduced the detectability of habitat concentration, while total time did not. We also found that uniform risk reduced accuracy of heterogeneous risk measurements. Future studies should aim to control all possible predators, as even the mild ones can induce complex behavior.