Respiratory characters of three species of haplochromine cichlids: Implications for use of wetland refugia

Respiratory characters of three east African haplochromine cichlid species that differ in their use of hypoxic wetlands were examined to consider the potential of dissolved oxygen as one factor affecting habitat use. All three species had a large gill surface area, ranging from the 67th ( Pseudocrenilabrus multicolor victoriae ) to 98th ( Astatotilapia velifer ) percentile of the known gill size range for freshwater fishes. Pseudocrenilabrus multicolor victoriae was the most tolerant to hypoxia exhibiting the lowest aquatic surface respiration ( Rs )thresholds and lowest critical oxygen tension of the three species. Astatotilapia velifer had the highest ASR thresholds, gill ventilation rates, and level of surface activity of the three species, indicating a relatively low tolerance to hypoxia. Prognathochromis venator was intermediate in its response to hypoxia. These findings are discussed in light of survivorship and distribution patterns of these species following Nile perch introduction into Lake Nabugabo.     

Effects of Nile perch, Lates niloticus, on functional and specific fish diversity in Uganda's Lake Kyoga system

The introduction of Nile perch, Lates niloticus, to Lake Victoria, East Africa, interacted with eutrophication to cause a reorganization of the lake's food web and the extirpation of many endemic fishes. The Lake Kyoga satellite system lies downstream from Lake Victoria. It encompasses species‐rich lakes where Nile perch are absent or very rare, and low diversity lakes where L. niloticus is abundant. In 1999 we surveyed seven lakes in the Kyoga system using experimental monofilament gill nets (1/4–1 inches variable mesh). At Boston University we assessed δ¹⁵N signatures of epaxial muscle from subsamples of the catch (n = 361). These signatures are often highly correlated with the near‐term mean realized trophic position of an individual organism. A neural network analysis of fish length, species name, trophic level, and lake of origin fish explained 94% of the sample variance in δ¹⁵N. We analysed statistical patterns in these signatures at a number of spatial scales. The relationship between trophic level and δ¹⁵N varied greatly among lakes. Higher diversity perch‐free lakes had greater variance in δ¹⁵N values and fish lengths than lower diversity Nile perch lakes, suggesting an important relationship between species diversity and functional diversity. Against expectations, lake size was negatively correlated with δ¹⁵N. Patterns in stable isotope signatures indicated that Nile perch lakes have shorter food chains than perch‐free lakes. The results throw up two management problems for the Kyoga system. Impacted lakes need to be studied to understand and ameliorate the community‐level effects of Nile perch introduction, whereas the species‐rich nonperch lakes, which harbour a large proportion of the remaining diversity of regionally endemic taxa, are in need of conservation planning.     

Plant species loss decreases arthropod diversity and shifts trophic structure

Plant diversity is predicted to be positively linked to the diversity of herbivores and predators in a foodweb. Yet, the relationship between plant and animal diversity is explained by a variety of competing hypotheses, with mixed empirical results for each hypothesis. We sampled arthropods for over a decade in an experiment that manipulated the number of grassland plant species. We found that herbivore and predator species richness were strongly, positively related to plant species richness, and that these relationships were caused by different mechanisms at herbivore and predator trophic levels. Even more dramatic was the threefold increase, from low- to high-plant species richness, in abundances of predatory and parasitoid arthropods relative to their herbivorous prey. Our results demonstrate that, over the long term, the loss of plant species propagates through food webs, greatly decreasing arthropod species richness, shifting a predator-dominated trophic structure to being herbivore dominated, and likely impacting ecosystem functioning and services.     

生态系统营养结构多样性的测度

生态系统层次上的生物多样性是指生物圈内生境、生物群落和生态过程以及生态系统的多样性,营养结构和营养过程的多样性是其中最重要的一个方面。本文通过宏观营养动力学的理论和方法,对生态系统营养结构进行分析,定义了反映生态系统营养结构多样性的两类指标,即营养物与流通量沿宏观营养链分布特征的多样性(D_Ⅰ),以及所有宏观营养级上营养物与流通量在不同分室上分布特征的多样性(D_Ⅱ)。这两类多样性是度量生态系统营养结构多样性的两个基本测度。     

Environmental variables drive spatial patterns of trophic diversity in mammals

Understanding environmental drivers of species diversity has become increasingly important under climate change. Different trophic groups (predators, omnivores and herbivores) interact with their environments in fundamentally different ways and may therefore be influenced by different environmental drivers. Using random forest models, we identified drivers of terrestrial mammals' total and proportional species richness within trophic groups at a global scale. Precipitation seasonality was the most important predictor of richness for all trophic groups. Richness peaked at intermediate precipitation seasonality, indicating that moderate levels of environmental heterogeneity promote mammal richness. Gross primary production (GPP) was the most important correlate of the relative contribution of each trophic group to total species richness. The strong relationship with GPP demonstrates that basal-level resource availability influences how diversity is structured among trophic groups. Our findings suggest that environmental characteristics that influence resource temporal variability and abundance are important predictors of terrestrial mammal richness at a global scale.     

Divergent ontogenies of trophic morphology in two closely related haplochromine cichlids

Fish develop morphological specializations in their trophic and locomotor systems as a result of varying functional demands in response to environmental pressures at different life stages. These specializations should maximize particular performances in specialists, adapting them to their trophic and habitat niches at each ontogenetic stage. Because differential growth rates of the structural components comprised in the head are likely to be linked to the diet of a fish throughout its development, we investigated the ontogenetic development of two haplochromine cichlid species belonging to different trophic guilds. We employed geometric morphometric techniques to evaluate whether starting from morphologically similar fry they diverge into phenotypes that characterize trophic guilds and locomotor types. Our examination of overall body shape shows that certain specialized morphological features are already present in fry, whereas other traits diverge through ontogeny due to differences in species‐specific allometric variation. Allometric shape variation was found to be more relevant for the biter specialist than for the sucker morphotype. Our results confirm that phenotypic changes during ontogeny can be linked to dietary and habitat shifts in these fish. Furthermore, evidence for an integrated development of trophic and locomotor specializations in morphology was observed. J. Morphol. 276:860–871, 2015. © 2015 Wiley Periodicals, Inc.     

黄海生态系统高营养层次生物群落功能群及其主要种类

根据2000年秋季和2001年春季在黄海的两次大面调查,选取生物量占总生物量90%的生物种类为研究对象,分析了黄海生态系统以及3个生态区(冷水团海域、近岸水域和黄海南部水域)春秋两季高营养层次生物群落的功能群组成及其主要种类.结果表明,黄海生态系统高营养层次生物群落包括6个功能群.按生物量排序为:浮游生物食性功能群、底栖动物食性功能群、鱼食性功能群、虾食性功能群、广食性功能群和虾/鱼食性功能群,各功能群营养级范围分别为3.22～3.35、3.30～3.46、4.04～4.50、3.80～4.00、3.38～3.79和4.01.黄海生态系统的主要功能群为浮游生物食性功能群和底栖动物食性功能群,占总生物量的79.6%;主要种类包括13种:小黄鱼、鳀、细巧仿对虾、银鲳、细点圆趾蟹、带鱼、黑鳃梅童、黄鲫、龙头鱼、双斑蟳、细纹狮子鱼、三疣梭子蟹和凤鲚,约占总生物量的70.6%.从不同季节看,春季黄海不同生态区高营养层次的营养级接近,而秋季差别较大,这主要与生物繁殖和索饵群体组成及摄食习性相关.从不同生态区看,黄海冷水团海域高营养层次生物群落以浮游生物食性功能群为主,受季节变化的影响较小,其高营养层次的营养级接近.黄海近岸水域和黄海南部水域高营养层次生物群落功能群组成受季节的影响较大,秋季的营养级均高于春季的营养级.这表明黄海冷水团海域较近岸水域和南部水域稳定,是黄海的一个典型的生态区域.     

Intra‐lake stable isotope ratio variation in selected fish species and their possible carbon sources in Lake Kyoga (Uganda): implications for aquatic food web studies

The stable isotopes of nitrogen (δ¹⁵N) and carbon (δ¹³C) provide powerful tools for quantifying trophic relationships and carbon flow to consumers in food webs; however, the isotopic signatures of organisms vary within a lake. Assessment of carbon and nitrogen isotopic signatures in a suite of plants, invertebrates, and fishes in Lake Kyoga, indicated significant variation between two sites for δ¹³C (paired t = 6.305; df = 14, P < 0.001 and δ¹⁵N paired t = 1.292; df = 14; P < 0.05). The fish fauna in Bukungu was generally more ¹³C enriched (mean δ¹³C = –16.37 ± 1.64‰) than in Iyingo (mean δ¹³C = –20.80 ± 2.41‰) but more δ¹⁵N depleted (mean δ¹⁵N = 5.57 ± 0.71‰) than in Iyingo (mean δ¹⁵N = 6.92 ± 0.83‰). The simultaneous shifts in phytoplankton and consumer signatures confirmed phytoplankton as the major source of carbon for the food chain leading to fish. Limited sampling coverage within lakes may affect lake wide stable isotope signatures, and the same error is transferred into trophic position estimation. Consideration of potential intra‐lake spatial variability in isotope ratios and size is essential in evaluating the spatial and trophic structure of fish assemblages.     

Predator diversity strengthens trophic cascades in kelp forests by modifying herbivore behaviour

Although human-mediated extinctions disproportionately affect higher trophic levels, the ecosystem consequences of declining diversity are best known for plants and herbivores. We combined field surveys and experimental manipulations to examine the consequences of changing predator diversity for trophic cascades in kelp forests. In field surveys we found that predator diversity was negatively correlated with herbivore abundance and positively correlated with kelp abundance. To assess whether this relationship was causal, we manipulated predator richness in kelp mesocosms, and found that decreasing predator richness increased herbivore grazing, leading to a decrease in the biomass of the giant kelp Macrocystis. The presence of different predators caused different herbivores to alter their behaviour by reducing grazing, such that total grazing was lowest at highest predator diversity. Our results suggest that declining predator diversity can have cascading effects on community structure by reducing the abundance of key habitat-providing species.     

The destruction of an endemic species flock: quantitative data on the decline of the haplochromine cichlids of Lake Victoria

Synopsis The Lake Victoria fish fauna included an endemic cichlid flock of more than 300 species. To boost fisheries, Nile perch (Lates sp.) was introduced into the lake in the 1950s. In the early 1980s an explosive increase of this predator was observed. Simultaneously, catches of haplochromines decreased. This paper describes the species composition of haplochromines in a research area in the Mwanza Gulf of Lake Victoria prior to the Nile perch upsurge. The decline of the haplochromines as a group and the decline of the number of species in various habitats in the Mwanza Gulf was monitored between 1979 and 1990. Of the 123+ species originally caught at a series of sampling stations ca. 80 had disappeared from the catches after 1986. In deepwater regions and in sub-littoral regions haplochromine catches decreased to virtually zero after the Nile perch boom. Haplochromines were still caught in the littoral regions where Nile perch densities were lower. However, a considerable decrease of species occurred in these regions too. It is expected that a remnant of the original haplochromine fauna will survive in the littoral region of the lake. Extrapolation of the data of the Mwanza Gulf to the entire lake would imply that approximately 200 of the 300+ endemic haplochromine species have already disappeared, or are threatened with extinction. Although fishing had an impact on the haplochromine stocks, the main cause of their decline was predation by Nile perch. The speed of decline differed between species and appeared to depend on their abundance and size, and on the degree of habitat overlap with Nile perch. Since the Nile perch upsurge, the food web of Lake Victoria has changed considerably and the total yield of the fishery has increased three to four times. Dramatic declines of native species have also been observed in other lakes as a result of the introduction of alien predators. However, such data concern less speciose communities and, in most cases, the actual process of extinction has not been monitored.     

Analysis of trophic relationships in two shallow equatorial lakes Lake Naivasha (Kenya) and Lake Ihema (Rwanda) using a multispecifies trophic model

A multispecifies trophic model called ECOPATH II, which can be used to describe the trophic relationships in aquatic ecosystems on a quantitative basis, is briefly presented. When properly used, it can help to explain the trophic relationships in ecosystems and possible evolution of fishstocks after modifications of the environment (e.g. eutrophication, introduction of a new population and/or a significant increase of the fishing effort), and to compare the trophic structure of several ecosystems. Examples are provided on two shallow lakes: Lake Ihema and Lake Naivasha. They are compared with Lake George which was previously documented.     

Ambient and substrate energy influence decomposer diversity differentially across trophic levels

The species-energy hypothesis predicts increasing biodiversity with increasing energy in ecosystems. Proxies for energy availability are often grouped into ambient energy (i.e., solar radiation) and substrate energy (i.e., non-structural carbohydrates or nutritional content). The relative importance of substrate energy is thought to decrease with increasing trophic level from primary consumers to predators, with reciprocal effects of ambient energy. Yet, empirical tests are lacking. We compiled data on 332,557 deadwood-inhabiting beetles of 901 species reared from wood of 49 tree species across Europe. Using host-phylogeny-controlled models, we show that the relative importance of substrate energy versus ambient energy decreases with increasing trophic levels: the diversity of zoophagous and mycetophagous beetles was determined by ambient energy, while non-structural carbohydrate content in woody tissues determined that of xylophagous beetles. Our study thus overall supports the species-energy hypothesis and specifies that the relative importance of ambient temperature increases with increasing trophic level with opposite effects for substrate energy.     

西沙群岛礁栖鱼类物种多样性及其食性特征

通过新的方法(即潜水调查法)调查了西沙群岛浅水区的礁栖鱼类, 并从食性角度分析鱼类群落特征及其对水质环境变化的响应。2006年5-6月, 对西沙群岛6个岛礁(东岛、永兴岛、羚羊礁、金银岛、华光礁和中建岛)共10个站位的礁栖鱼类种类、数量和长度进行了调查, 结合公开资料(FishBase等)确定鱼类食性。共记录到29科71属119种鱼, 其中50种是新记录。至此, 西沙群岛的鱼类总记录已达到717种。从不同站位来看, 永兴岛西的种类数量和Shannon-Wiener多样性指数均最高。多数鱼种分布范围狭小。摄食浮游动物和底栖无脊椎动物的鱼类是优势类群, 而草食性、杂食性和食物链顶端食性的鱼类数量和生物量较低, 反映出西沙群岛珊瑚礁生态系统整体上健康稳定。永兴岛的杂食性鱼类的数量比例和生物量比例均高于其他岛礁, 反映出鱼类群落已对水体富营养化产生响应。     

Ecomorphology of retinal structures in zooplanktivorous haplochromine cichlids (Pisces) from Lake Victoria

Synopsis Before the decline of the species flock of haplochromine cichlids of Lake Victoria due to the Nile perch upsurge, there were many co-existing haploehromine species such as the taxonomically and ecologically well-studied zooplanktivores of the Mwanza Gulf. In spite of the scarcely separated niches of some of these species, no sign of competition for space or food could be demonstrated. As is argued in this paper, optical differentiation could well be an aspect of adaptive radiation of these zooplanktivores, particularly among the highly sympatric species. Our hypothesis is based on the morphological modifications of retinal structures in nine zooplanktivorous species. Interspecific variation was observed in composition, size and density of the photoreceptors and ganglion cells. The analyses included the intraretinal variation and size dependency of some of the structural parameters. The optical functions deduced from retinal structure indicate distinct interspecific differences in sensitivity thresholds and a slight differentiation in visual resolution. These functions correlate poorly with the photic conditions of the species-specific habitats. The optical properties can, on the other hand, be connected with the more subtle differentiation in food items and feeding behaviour among these species. It is our concluding hypothesis, that the optical differentiation among the haplochromine zooplanktivores primarily served resource partitioning by different modes of visual prey detection rather than niche partitioning by habitat.     

Comparison of size-structured and species-level trophic networks reveals antagonistic effects of temperature on vertical trophic diversity at the population and species level

It is predicted that warmer conditions should lead to a loss of trophic levels, as larger bodied consumers, which occupy higher trophic levels, experience higher metabolic costs at high temperature. Yet, it is unclear whether this prediction is consistent with the effect of warming on the trophic structure of natural systems. Furthermore, effects of temperature at the species level, which arise through a change in species composition, may differ from those at the population level, which arise through a change in population structure. We investigate this by building species-level trophic networks, and size-structured trophic networks, as a proxy for population structure, for 18 648 stream fish communities, from 4 145 234 individual fish samples, across 7024 stream locations in France from 1980 to 2008. We estimated effects of temperature on total trophic diversity (total number of nodes), vertical trophic diversity (mean and maximum trophic level) and distribution of biomass across trophic level (correlation between trophic level and biomass) in these networks. We found a positive effect of temperature on total trophic diversity in both species- and size-structured trophic networks. We found that maximum trophic level and biomass distribution decreased in species-level and size-structured trophic networks, but the mean trophic level decreased only in size-structured trophic networks. These results show that warmer temperatures associate with a lower vertical trophic diversity in size-structured networks, and a higher one in species-level networks. This suggests that vertical trophic diversity is shaped by antagonistic effects of temperature on population structure and on species composition. Our results hence demonstrate that effects of temperature do not only differ across trophic levels, but also across levels of biological organisation, from population to species level, implying complex changes in network structure and functioning with warming.     

Phylogenomics of trophically diverse cichlids disentangles processes driving adaptive radiation and repeated trophic transitions

Cichlid fishes of the tribe Tropheini are a striking case of adaptive radiation, exemplifying multiple trophic transitions between herbivory and carnivory occurring in sympatry with other established cichlid lineages. Tropheini evolved highly specialized eco‐morphologies to exploit similar trophic niches in different ways repeatedly and rapidly. To better understand the evolutionary history and trophic adaptations of this lineage, we generated a dataset of 532 targeted loci from 21 out of the 22 described Tropheini species. We resolved the Tropheini into seven monophyletic genera and discovered one to be polyphyletic. The polyphyletic genus, Petrochromis, represents three convergent origins of the algae grazing trophic specialization. This repeated evolution of grazing may have been facilitated by adaptive introgression as we found evidence for gene flow among algae grazing genera. We also found evidence of gene flow among algae browsing genera, but gene flow was restricted between herbivorous and carnivorous genera. Furthermore, we observed no evidence supporting a hybrid origin of this radiation. Our molecular evolutionary analyses suggest that opsin genes likely evolved in response to selection pressures associated with trophic ecology in the Tropheini. We found surprisingly little evidence of positive selection in coding regions of jaw‐shaping genes in this trophically diverse lineage. This suggests low degrees of freedom for further change in these genes, and possibly a larger role for regulatory variation in driving jaw adaptations. Our study emphasizes Tropheini cichlids as an important model for studying the evolution of trophic specialization and its role in speciation.     

Fish community structure in mesotrophic and eutrophic lakes of southern Finland: the relative abundances of percids and cyprinids along a trophic gradient

In 36 south Finnish lakes, the number of species, as well as the cyprinids:percids ratio, was dependent, not only on total phosphorus (TP), but also on lake size. Total fish biomass and cyprinid biomass increased along the TP gradient, whereas the dependence of percid biomass was less evident. Perch Perca fluviatilis and roach Rutilus rutilus strongly dominated mesotrophic lakes; in eutrophic lakes the proportion of other cyprinids and percids, such as white bream Blicca bjoerkna , bream Abramis brama , pikeperch Stizostedion lucioperca and ruffe Gymnocephalus cernuus , increased. Perch biomass was weakly related to abiotic factors but depended on roach biomass. Lake size and fish species composition are essential factors affecting fish community changes in relation to TP, and may be important as well in regulating the responses of a fish community to biomanipulation.     

Assessing fern diversity: relative species richness and its environmental correlates in Uganda

Techniques for the rapid quantification of tropical biodiversity are of critical importance in deciding where to invest scarce conservation resources. Here we describe a simple survey method for assessing species-level richness of a poorly known plant group, the pteridophytes. We then illustrate the use of a powerful, rarefaction- based technique of controlling for inevitable differences in sampling effort to calculate the relative species richness of our study sites. Lastly, we explore how closely observed patterns of relative species richness of Ugandan forests are correlated with a suite of simple environmental variables. We find that fully 75% of the variance in our estimate of fern diversity can be predicted from just two measures: soil fertility (scored as C/N ratio, itself related to rainfall); and distance from the nearest putative Pleistocene refugium.     

Previously undocumented diversity and abundance of cryptic species: a phylogenetic analysis of Indo-Pacific Arminidae Rafinesque, 1814 (Mollusca: Nudibranchia) with descriptions of 20 new species of Dermatobranchus

The phylogenetic relationships amongst the Arminidae were analysed based upon morphological characters of 58 presently described species or nudibranchs, including 35 previously described Arminidae and 20 new species of Dermatobranchus. From the literature review and anatomical examinations, 43 characters were considered for 78 taxa. These characters were polarized using Berthella canariensis as the outgroup taxon and the type species of several other genera identified from recent publications. The resulting phylogeny supports the monophyly of Arminidae, Dermatobranchus, Doridina, and Proctonotidae. The paraphyly of the Arminina is further demonstrated in this study. Two previously described, but poorly known, species of Indo-Pacific Armina are redescribed, Armina magnaBaba, 1955 and Armina paucifoliataBaba, 1955. The anatomy and taxonomic status of nine previously described species of Dermatobranchus were examined in this study. The anatomy of Dermatobranchus pustulosus (van Hasselt, 1824) has been overlooked since Bergh (1888) illustrated the radula of van Hasselt's specimen. It is redescribed and its range is extended to several new localities in the western Pacific. Dermatobranchus pulcherrimus Miller & Willan, 1986 is considered here as a new synonym of Dermatobranchus rubidus (Gould, 1852). The following 20 species of Dermatobranchus are new and are described in the present paper: Dermatobranchus albineus sp. nov., Dermatobranchus arminus sp. nov., Dermatobranchus caesitius sp. nov., Dermatobranchus caeruleomaculatus sp. nov., Dermatobranchus cymatilis sp. nov., Dermatobranchus dendonephthyphagus sp. nov., Dermatobranchus diagonalis sp. nov., Dermatobranchus earlei sp. nov., Dermatobranchus fasciatus sp. nov., Dermatobranchus funiculus sp. nov., Dermatobranchus kalyptos sp. nov., Dermatobranchus kokonas sp. nov., Dermatobranchus leoni sp. nov., Dermatobranchus microphallus sp. nov., Dermatobranchus oculus sp. nov., Dermatobranchus phyllodes sp. nov., Dermatobranchus piperoides sp. nov., Dermatobranchus rodmani sp. nov., Dermatobranchus semilunus sp. nov., and Dermatobranchus tuberculatus sp. nov. Eighteen of these new taxa are found in the Indo-Pacific tropics and two are found in temperate South Africa, D. albineus and D. arminus. Unique combinations of morphological characters distinguish these as new species of Dermatobranchus. Several species that are externally similar have radically divergent internal morphology, are members of different clades of Dermatobranchus, and represent cryptic species. Especially important is the radular morphology, which shows remarkable diversity of form, probably related directly to the diversification of feeding of members of this clade on various octocorals.     

Reconstructing the historical trophic status of northwestern Pennsylvania lakes using GIS

From each of 46 watersheds in glaciated northwestern Pennsylvania we estimated phosphorus export (kg P/ha/yr) from weekly or twice-weekly measured stream phosphorus concentrations and measured stream discharges, and determined land covers using GIS. Simple and step-down multiple regression analyses yielded models that explained 24% of the variation in P export using land cover within whole watersheds, and 64% of the variation using land cover within 200 m riparian buffers. We used these models to predict P loading to seven lakes and found that predicted lake [P] was consistent with measured lake [P]. To estimate pre-settlement lake [P] we reapplied the P export models with the assumption that human-impacted land covers were originally forests. Predicted (hindcast) pre-settlement lake [P] indicated that six of the seven lakes were edaphically mesotrophic (10 < [P] < 20 μg/l). Lake remediation targets set on the assumption that area lakes were historically oligotrophic ([P] < 10 μg/l) will be unattainable.