Effects of species vs. functional diversity: Understanding the roles of complementarity and competition on ecosystem function in a tropical stream fish assemblage

The positive effects of biodiversity on the functioning of ecosystems are well demonstrated in laboratory microcosms but the precise mechanisms underlying higher ecosystem process rates in natural assemblages are less well understood. We investigated, under semi-natural conditions (field enclosures), the potentially interactive effects of species identity and trophic function (i.e., feeding guild) on consumer growth, using a fish assemblage in a tropical stream. We tested the relative importance of species identity and trophic function on consumer growth by placing 2 fish of either (i) the same species, (ii) different species but of similar trophic function, or (iii) different species of different trophic functions in each of 72 stream enclosures for 48 days and measuring biomass change, individual diet composition and behavior. We predicted that if functional diversity had a larger impact than species diversity, then fish growth would be highest for pairs of species from different functional groups (i.e., those with the highest diet complementarity), intermediate for different species within a guild, and lowest for the same species (those with the lowest complementarity and highest niche overlap), such that functional variation between species in different guilds would exceed functional differences within guilds. Our results show that functional heterogeneity rather than species diversity per se had the greatest impact on food resources used complementarily, leading to higher biomass accrual. Mechanistically, higher growth rates were driven by concomitant increases in resource intake and reductions in antagonistic interactions. Together, these results underscore the importance of functional diversity on macroconsumer production in natural assemblages.     

Effects of longitudinal variations in stream habitat structure on fish abundance: an analysis based on subunit-scale habitat classification

SUMMARY 1. Stream reaches contain assortments of various habitat types that can be defined at different spatial scales, such as channel unit (e.g. pools, riffles) and subunit (patches within channel units). We described longitudinal (upstream–downstream) patterns of stream habitat structure by considering subunits as structural elements, and examined their effects on the abundance of masu salmon ( Oncorhynchus masou ) and rosyface dace ( Leuciscus ezoe ) in a third-order tributary of the Teshio River in northern Hokkaido, Japan.
2. Nine subunit types were determined on the basis of water depth, current velocity and substrate, using 0.5 × 0.5 m grids. Although both masu salmon and rosyface dace used pools as a major habitat, the former preferred a subunit type occurring at pool heads (PH subunit) while the latter preferred a slow-current edge type (SE-2 subunit).
3. Along the course of the stream, slow-edge subunits (SE-1, 2 and 3) increased in frequency downstream while fast-edge subunits (FE-1 and 2) decreased, suggesting a downstream development of slow-current edges. Regression analyses indicated that longitudinal variation in masu salmon abundance was explained by the area of PH, rather than pools. Masu salmon density increased with the area of PH. Rosyface dace abundance was explained by a combination of water depth and the area of SE-2, both effects being positive.
4. Longitudinal variations in the abundance of both species were related to the abundance of their preferred habitat at the subunit scale, rather than channel-unit scale. The results emphasise the importance of fine-scale patchiness when examining stream fish habitats.     

太子河流域鱼类功能群结构与多样性对土地利用类型的响应

研究河岸带土地利用方式对河流生物群落的影响对河岸带管理和河流生态系统修复至关重要。研究了太子河河岸带的土地利用类型(森林用地、森林耕作用地、耕地和城镇建设用地)和鱼类功能群的关系,结果表明:栖息地质量参数在不同土地利用类型内具有显著差异,森林用地区电导率、总溶解固体、淤泥和底质含沙量比例的平均值均较低,分别为(105.05μs/cm、80.38 mg/L、65.00 mL和0%),底质类型以石块为主;耕作区的水深、流量和淤泥的平均值均是最高(186.83 m、80.11 m~3和5333.33 mL),底质类型以沙质和淤泥为主。太子河流域鱼类功能群划分为5种类型(18个亚类),在不同土地利用类型内具有差异显著,森林用地内的鱼食性、石块栖功能群、昆虫食性和黏性卵功能群的比例最高;森林耕作用地内的植食性和底层栖功能群的比例最高;耕地内的沙栖功能群、中下层栖功能群和筑巢产卵功能群的比例最高;城镇建设用地内的淤泥栖功能群、耐污种、中下层栖功能群和杂食性功能群的比例最高。研究显示,栖息地评价得分高、栖境复杂的区域其个体数量较高,而栖息地得分低、底质类型以淤泥为主的区域其个体数量较低。     

Spatio‐temporal scaling of biodiversity and the species–time relationship in a stream fish assemblage

1. The increase of species richness with the area of the habitat sampled, that is the species–area relationship, and its temporal analogue, the species–time relationship (STR), are among the few general laws in ecology with strong conservation implications. However, these two scale‐dependent phenomena have rarely been considered together in biodiversity assessment, especially in freshwater systems. 2. We examined how the spatial scale of sampling influences STRs for a Central‐European stream fish assemblage (second‐order Bernecei stream, Hungary) using field survey data in two simulation‐based experiments. 3. In experiment one, we examined how increasing the number of channel units, such as riffles and pools (13 altogether), and the number of field surveys involved in the analyses (12 sampling occasions during 3 years), influence species richness. Complete nested curves were constructed to quantify how many species one observes in the community on average for a given number of sampling occasions at a given spatial scale. 4. In experiment two, we examined STRs for the Bernecei fish assemblage from a landscape perspective. Here, we evaluated a 10‐year reach level data set (2000–09) for the Bernecei stream and its recipient watercourse (third‐order Kemence stream) to complement results on experiment one and to explore the mechanisms behind the observed patterns in more detail. 5. Experiment one indicated the strong influence of the spatial scale of sampling on the accumulation of species richness, although time clearly had an additional effect. The simulation methodology advocated here helped to estimate the number of species in a diverse combination of spatial and temporal scale and, therefore, to determine how different scale combinations influence sampling sufficiency. 6. Experiment two revealed differences in STRs between the upstream (Bernecei) and downstream (Kemence) sites, with steeper curves for the downstream site. Equations of STR curves were within the range observed in other studies, predominantly from terrestrial systems. Assemblage composition data suggested that extinction–colonisation dynamics of rare, non‐resident (i.e. satellite) species influenced patterns in STRs. 7. Our results highlight that the determination of species richness can benefit from the joint consideration of spatial and temporal scales in biodiversity inventory surveys. Additionally, we reveal how our randomisation‐based methodology may help to quantify the scale dependency of diversity components (α, β, γ) in both space and time, which have critical importance in the applied context.     

Higher taxa vs. functional guilds vs. trophic groups as indicators of soil nematode diversity and community structure

Biodiversity surrogates are often used in ecology to save money and time. One such widely used surrogacy method is the higher taxon approach, also known in the literature as taxonomic sufficiency. This approach has rarely been applied to microscopic organisms like soil microfauna, although there is a lack of detailed taxonomic knowledge regarding the latter. We tested taxonomic sufficiency for genus alpha and beta diversity, as well as for community structure of soil nematodes. We also tested whether the functional classifications of nematodes into functional guilds and trophic groups can serve as efficient indicators of nematode diversity and community structure. We used data from soil nematode communities from five different microhabitats (soil, soil moss, rock moss, low tree trunk moss and high tree trunk moss) in forested and non forested areas at five different stations (differing in elevation and aspect). Our results showed that both the higher taxon approach and the functional surrogacy approach could serve as reliable indicators of alpha and beta diversity. Furthermore, our community analysis of both taxonomic and functional composition identified that the variability at the finer scales, and namely the habitat structure, played the leading role in shaping nematode communities.     

Soil fauna responses to invasive alien plants are determined by trophic groups and habitat structure: a global meta‐analysis

Despite increasing frequency of invasions by alien plant species with widespread ecological and economic consequences, it remains unclear how belowground compartments of ecosystems are impacted. In order to synthetize current knowledge and provide future directions for research we performed a meta‐analysis assessing the impact of invasive alien plant species on soil fauna abundance. Compared to previous synthesis on this topic, we included together in our model the trophic group of each soil faunal taxa (from herbivores to predators) and habitat structure, namely open and closed habitats (i.e. grass and shrub dominated areas versus forested areas). In doing so, we highlighted that both moderators strongly interact to determine the response of soil fauna to the presence of invasive alien plants. Soil fauna abundance increase in the presence of invasive species only in closed habitats (+18.2%). This pattern of habitat‐dependent response (positive effect in closed habitats) was only found for primary consumers (i.e. herbivores +29.1% and detritivores +66.7%) within both detritus‐based and live root‐based trophic pathways. Abundances of predators and microbivores did not respond to the presence of IAS irrespective of habitat structure. For several groups, the habitat structure (open or closed) significantly drove their responses to the presence of invasive alien species. In addition, we carefully considered potential sources of bias (e.g. geographic, taxonomic and functional) within the collected data in an attempt to highlight gaps in available knowledge on the subject. Our findings support the conclusions of previous studies on the subject by demonstrating 1) that soil fauna abundance is impacted by biological invasions, 2) that initial habitat structure has a strong influence on the outcome and 3) that responses within the soil fauna differ between trophic levels with a stronger response of primary consumers.     

Habitat structure and invertebrate assemblages on stream stones: A multivariate view from the riffle

Abstract Highly structured habitats, those with complex or heterogeneous physical structure, commonly contain more species than simply structured ones. However, tests of this relationship have been hampered by habitat-specific definitions and measurements of habitat structure and, in stream studies, by insufficient information on spatial variation in faunal abundances. We sampled the fauna of 90 stones collected from an order 4 section of the Taggerty and Steavenson Rivers, southeastern Australia. The spatially nested sampling design encompassed three spatial scales (sites within the same stream order, riffles at the same site and groups of stones within the same riffle) but no temporal replication, as we were only able to sample once. Habitat structure of stones was quantified by measuring the amount and type of epilithon on stones, together with stone shapes, sizes, textures and surface complexities. For the latter two attributes, we used a new method, involving image analysis, which can be used to quantify habitat structure in comparable ways in other systems. Species richness (S) was related to some measures of habitat structure, being higher on stones with a rough texture and weakly related to the amount of epilithon present. Total numbers of individuals (N) were also higher on rough stones, and faunal composition (as quantified by hybrid multidimensional scaling) was related to abundance of epilithon. However, flow environments, as quantified by water velocities and depths, were also related to faunal composition. Riffles varied in both faunal densities and near-bed flow environment, but not in any consistent way. Such variability means that individual riffles cannot be reasonably used to ‘represent’ stream sections or orders, an assumption commonly made in stream studies. Neglect of such issues has also resulted in poor conceptual integration between large- and small-scale studies in stream ecology.     

基于稳定同位素技术的长湖鱼类营养结构研究

本研究应用碳、氮稳定同位素技术分析了2017年秋季（10月）和2018年春季（3月）湖北省荆州长湖鱼类营养结构特征,构建了δ¹³C和δ¹⁵N稳定同位素双位图,并计算了7个相关的量化指标。结果显示：所采集的16种鱼类,δ¹³C均值范围为（-27.7±0.8）‰~（-24.8±0.1）‰,δ¹⁵N的均值范围为（11.4±0.3）‰~（16.6±0.5）‰。营养生态位分析显示,长湖鱼类以偏肉食性鱼类黄颡鱼的营养级最高（3.28±0.2）,草鱼的营养级最低（1.74±0.1）。稳定同位素的量化指标表明,长湖春季鱼类群落核心生态位空间（standard ellipse area, SEA）、生态位总空间（total area, TA）和基础食物来源（δ¹³C range, CR）均高于秋季,两个季度的营养多样性（centrifugal distance, CD）和鱼类群落的整体密度（mean nearest neighbor distance, MNND）相似,但春季的营养长度（δ¹⁵N range, NR）和鱼类群落营养生态位分布范围（standard deviation of nearest neighbor distance, SDNND）值低于秋季。说明春季长湖的生态位总空间（TA上升）和基础食物来源（CR上升）较为丰富,但因鱼类种类的食性相近,其群落的总食物链降低（NR下降）。该研究可为分析拆围后长湖鱼类群落结构特征积累基础数据,也为长湖的渔业管理策略提供依据。     

Habitat structure and fish predation: effects on invertebrate colonisation and breakdown of stream leaf packs

1. We investigated the effects of two features of leaf‐pack habitat structure (i.e. mass of a leaf pack and surface area of leaves comprising a leaf pack) and fish predation on colonisation of shredders and leaf breakdown rates in a coldwater stream. Packs were constructed of red maple (Acer rubrum) leaves. 2. A 2 × 3 × 3 factorial experiment was used to manipulate fish predation (exclusion and control cage), leaf‐pack mass (1, 3 and 5 g dry mass) and leaf surface area (small: approx. 17.9 cm², medium: approx. 34.6 cm², large: approx. 65.6 cm²). Exclusion cages had mesh on all sides, whereas control cages lacked mesh on two sides to provide access to fish. 3. Common shredders were Gammarus pseudolimnaeus, Pycnopsyche and Lepidostoma. Shredder biomass per leaf pack increased with the mass of a leaf pack (P < 0.001), but biomass per unit mass of leaf pack did not differ with leaf‐pack mass (P = 0.506). Shredder densities did not respond to the exclusion of fish (P > 0.7) or leaf surface area (P > 0.7), and interactions among treatment factors were not significant (P > 0.2). 4. Breakdown rates were lower for leaf packs comprised of small leaves (P < 0.001) and leaf packs with high mass (P = 0.001). Excluding fish did not significantly affect leaf breakdown rates (P = 0.293), and interactions among treatment factors were not significant (P > 0.3). Breakdown rates were highest when packs consisted of few leaves (i.e. leaf packs with large leaves and low mass) and were colonised by many shredders. 5. Fish predation was not an important factor controlling shredder densities in leaf packs over the spatiotemporal scale of our experiment. Nevertheless, we found shredder colonisation was proportional to leaf‐pack mass and breakdown rates were affected by leaf‐pack size (i.e. number of leaves in a pack). We suspect that fragmentation is the primary mechanism causing the breakdown rates to be dependent on leaf‐pack size.     

Modelling the impacts of two exotic invasive species on a native butterfly: top-down vs. bottom-up effects

1. Exotic invasive species can influence population dynamics of native species through top-down or bottom-up forces. The present study examined separate and interactive effects of multiple exotic species invasions on the native mustard white butterfly, Pieris napi oleracea Harris (Lepidoptera: Pieridae), using a stochastic simulation model. 2. P. n. oleracea populations in North America have decreased regionally since the 1860s. Competition with an exotic congener (P. rapae L.), loss of native host plants and parasitism by the introduced broconid wasp (Cotesia glomerata L.), have been suggested to be independently responsible for its decline. The present study examined these hypotheses, as well as an alternative, invasion by an exotic crucifer, garlic mustard (Alliaria petiolata[Bieb.] Cavara & Grande). 3. A stochastic simulation model of P. n. oleracea population dynamics revealed that decreasing the number of host plants available for oviposition and larval development (i.e. habitat loss), sharply reduced the probability of populations persistence and decreased population size for those that persisted. 4. Simulated invasion by garlic mustard also substantially decreased both probability of persistence (= 0 at approximately 50% cover) and mean population size. Persistence probability never reached zero under any C. glomerata scenarios, even when larval mortality in the second generation due to parasitism was 100%. The impact of garlic mustard was intensified by the addition of C. glomerata parasitism. 5. Results suggest that bottom-up forces, loss of host plants through forest understorey loss and/or garlic mustard invasion are the most important forces driving P. n. oleracea population decline. Parasitism by C. glomerata may interact to reduce P. n. oleracea populations more rapidly, but appears insufficient alone to cause local extinction.     

Patterns of functional and taxonomic organization of stream fishes: inferences based on α, β, and γ diversities

The primary objective of this study was to determine whether total biodiversity (γ) is partitioned into within‐community (α) and among‐community (β) components differently for taxonomic and functional organization. I hypothesized that α diversity will contribute more to the functional organization of γ diversity and that β diversity will contribute more to the taxonomic organization of γ diversity. A secondary objective was to determine whether the relationship between taxonomic and functional diversity is scale dependent. Species abundance data was obtained from fisheries surveys conducted by the Texas Parks and Wildlife Dept that focused on least disturbed streams from 11 different ecoregions of Texas, including 62 localities from 18 drainages. Functional and taxonomic organization of assemblages was quantified with two different measures of biodiversity, including richness and the numbers equivalent of Shannon diversity. Scale‐dependent effects on these indices were assessed by multiplicatively partitioning γ into α and β components. The contribution of α and β components to γ diversity differed between functional and taxonomic organization and among different measures of biodiversity. Among‐community components were more influential in structuring the taxonomic organization of stream‐fish assemblages, whereas within‐community components were more important in structuring the functional organization of assemblages. The relationship between taxonomic and functional diversity differed between α and β components and between spatial scales. Indeed, ecological patterns not only change with spatial scale, but how they change is dependent on which aspect of biodiversity is considered.     

山地河流浅滩深潭生境大型底栖动物群落比较研究——以重庆开县东河为例

浅滩和深潭是山地河流中常见的河流生境结构。2011年7月,在重庆开县东河上游双河口-杉木桥河段,选择21个浅滩和深潭,调查大型底栖动物,研究影响不同生境中底栖动物组成、分布和多样性的生态机理。结果表明:调查河段浅滩和深潭中大型底栖动物分别为31种和24种,密度分别为450.62 个/m²和86.24 个/m²,生物量分别为2.88 g/m²和0.55 g/m²。浅滩有指示种11种,即纹石蛾(Hydropsyche sp.)、假蜉(Iron sp.)、假二翅蜉(Pseudocloeon sp.)、舌石蛾(Glossosoma sp.)、高翔蜉(Epeorus sp.1)、背刺蜉(Notacanthurus sp.)、Heterocloeon sp、锯形蜉(Serratella sp.)、朝大蚊(Antocha sp.)、等蜉(Isonychia sp.)、溪颏蜉(Rhithrogena sp.)。深潭指示种仅蜉蝣(Ephemera sp.)和黑大蚊(Hexatoma sp.)两种。刮食者为两类生境的优势功能摄食类群。浅滩中滤食者和刮食者比例显著高于深潭,而收集者和捕食者显著低于深潭。两类生境中大型底栖动物群落结构差异显著。浅滩中大型底栖动物的密度、生物量、丰富度指数、Shannon-Wiener 指数、改进的Shannon-Wiener指数均明显高于深潭。受地貌形态、水力特征和冲淤变化规律影响的生境稳定性和异质性差异是导致大型底栖动物群落差异的主要原因。     

Local and regional assessments of the impacts of plant invaders on vegetation structure and soil properties of Mediterranean islands

Aims Although biological invasions occur throughout the world, and some invaders are widespread in many habitats, few studies on the ecological impact of invaders have examined multiple sites. We tested how the impact of three widespread plant invaders changed depending on the identity of the species and the invaded island. We also tested whether relative species loss was lower in species‐rich communities than in species‐poor ones. Location We conducted floristic surveys and soil analyses in eight Mediterranean Basin islands: Crete and Lesbos (Greece), Sardinia (Italy), Corsica, Bagaud and Porquerolles (France), and Mallorca and Menorca (Spain). Methods We compared native species richness and diversity, proportion of life forms, soil percentage nitrogen, percentage organic carbon, C/N, and soil pH in nearby paired plots of 2 × 2 m: one control and one invaded by either the deciduous tree Ailanthus altissima, the succulent subshrubs Carpobrotus spp. or the annual geophyte Oxalis pes‐caprae, across eight Mediterranean Basin islands. Results On average, the presence of invaders reduced species diversity, Carpobrotus spp. exhibiting the largest impact and Oxalis the least. However, the relative impact was island‐dependent, and was positively but weakly associated with the species richness of the recipient community. Therophytes were the life form that experienced the largest decrease across islands. The effects of invasion on soil properties were very variable. Total N changed (increased) only in plots invaded by Ailanthus, significantly decreasing the C/N ratio. The presence of this tree increased soil pH, whereas the opposite was found in plots invaded by the other two species. Organic C increased in plots invaded by Ailanthus and Carpobrotus species. Main conclusions By conducting an analysis at multiple sites, we found that the three plant invaders had an impact on plant community structure not entirely concordant with changes in soil properties. The impacts depended on the identity of the species and of the invaded island, suggesting that impact of invaders is context‐specific. The impact in terms of species loss was not lower in species‐rich than in species‐poor communities.     

Biogeography of Aphodiinae dung beetles based on the regional composition and distribution patterns of genera

Aim  To examine current biogeographical patterns of Aphodiinae dung beetles (Coleoptera: Scarabaeoidea: Scarabaeidae) in order to reveal relationships among regions and their potential impact on the diversification of this group.
Location  Worldwide.
Methods  Information about all Aphodiinae genera was obtained from the literature. An occurrence matrix was built for the six worldwide biogeographical regions, and their faunas were characterized through simple statistics. Regional variations and similarities were further explored using co-occurrence and nestedness analyses, sequential agglomerative, hierarchical and nested clustering (SAHN), and a parsimony analysis of endemicity (PAE). Mantel tests were also employed to assess the relationships between several characteristics of the regions and their faunas.
Results  The Palaearctic and Palaeotropical regions showed the highest total numbers of Aphodiinae genera and the greatest generic endemism. Both these regions and the Oriental also showed higher numbers of genera than would be expected according to their size. Co-occurrence and nestedness analyses confirmed the non-randomness of the distribution of genera. Clustering and PAE showed that the Palaearctic and Oriental regions are the most similar, followed by the Palaeotropical region. Regional dissimilarity in genera composition was related to biological and historical traits, but not to ecoregions.
Main conclusions  A structured geographical pattern for Aphodiinae was confirmed. Land continuity and proximity in the long term could have played a unifying role in regional faunas. We suggest that the different biogeographical regions have acted as either macroevolutionary sources (basically the Palaearctic and the Palaeotropical regions) or sink regions, according to their role as diversification centres. We review the processes and events that could account for current patterns of Aphodiinae diversity.     

Determinants of local spider (Araneidae and Thomisidae) species richness on a regional scale: climate and altitude vs. habitat structure

Abstract.  1. The habitat heterogeneity hypothesis states that the more complex the habitat, the higher the species richness. The present study analyzes the effect of local factors on regional spider ( Araneidae and Thomisidae ) richness. The main objective is to disentangle the relative importance of habitat structure and other environmental variables.
2. Fifteen territorial units of 1 km² were sampled to obtain reliable and comparable inventories of the two spider families. Richness values were modelled using general regression models and a set of climate, topographic and vegetation structure variables. Pure and joint effects were computed via variation partitioning.
3. The results highlight the great importance of vegetation complexity, especially of grass and sub-shrub cover, in determining spider species richness.
4. The maximum temperature is the only climate variable significantly related to species richness, although its effect is combined with that of spatial and vegetation structure variables.
5. These results support the habitat heterogeneity hypothesis, and highlight the importance of taking vegetation complexity into account when managing habitats and where spider conservation is desired.     

The effects of fire on ant communities in north-western Patagonia: the importance of habitat structure and regional context

Abstract. We investigated the effects of recent fires on the native ant communities in two habitats of north-west Patagonia that differ in vegetation structural complexity. Using bait traps, we sampled ants in replicated scrub and steppe areas including paired burned and unburned sites. Fires significantly reduced plant cover and ant diversity only in scrub sites. The drop in diversity was due to (a) a reduction in the abundance of rare species associated with woody vegetation, and (b) an increase in the abundance of the dominant species, which thrive in more xeric microclimatic conditions. Consequently, ant assemblage structure of burned scrub approaches that of steppe sites. Our findings suggest that the effects of disturbances on ant assemblages depends both on habitat characteristics, which in turn determine the extent of the changes induced by the disturbance, and on the regional context of the ant fauna, which in turn determines the ability of the ants to deal with the post-disturbance conditions.