Long-term changes in a benthic assemblage associated with artificial reefs

The aim of the study was to evaluate the long-term development of a hard bottom benthic assemblage over a period of 20 years in an area off the mouth of a large river. The artificial reef of Fregene was selected because benthic assemblage data were available for the period 1981–1992. This artificial reef is located in the mid Tyrrhenian Sea, 5 nautical miles north of the two mouths of the Tevere River (Latium, Italy) and 1.5 nautical miles offshore from Fregene (Rome, Italy). The artificial reef was deployed in March 1981 for fisheries enhancement in 10–14 m of water on a sandy-silty seabed. The Tevere River carries suspended materials and a heavy load of organics since it transports Rome’s effluent, resulting in the eutrophic state of area waters. Benthic sampling was conducted in 2001 by SCUBA diving; two standard surfaces of 400 cm² were scraped from the vertical walls of the same uppermost block in four different periods. All organisms were identified and counted. The methodology used is the same as that adopted in the previous periods, so that the 2001 data could be compared with past collected data. The benthic assemblage was analysed by cluster analysis using the Bray-Curtis index and clustered using the group average clustering algorithm. The SIMPER procedure was used to identify those taxa that characterize each station group identified by cluster analysis. Changes in benthic assemblages and hydrological trends of the Tevere River were investigated using the cumulative sum series method. The 20-year development of the benthic community, starting from the new substratum, is composed of different phases characterised by different benthic assemblages. In particular five different phases were distinguished: 1. Pioneer species recruitment (May 1981–June 1981); 2. Mytilus galloprovincialis (mussel) dominance (August 1981–November 1983); 3. M. galloprovincialis regression (July 1984–October 1985); 4. M. galloprovincialis absence (91–92); 5. Bryozoans bioconstruction dominance (2001). The dynamic succession of the observed benthic assemblages exhibited a good relation with the Tevere River flow. The Tevere River flow, and the subsequent sedimentation process, seems to have strongly influenced the benthic assemblage succession of the Fregene artificial reef. Guest editors: G. Relini & J. Ryland Biodiversity in Enclosed Seas and Artificial Marine Habitats     

Trophic niche changes associated with habitat fragmentation in a Neotropical bat species

Habitat fragmentation could alter ecological traits including species trophic habits. Here, we used carbon and nitrogen stable isotope ratios to establish differences in isotopic niche width and food resource use between forest fragments and the continuous forest for the phyllostomid frugivorous bat Artibeus lituratus. Using mist nests, we captured bats from two forest fragments and two sites in continuous forest, and sampled from each individual captured three body tissues with contrasting turnover rates (skin, muscle, and liver). Samples were collected between February and March (austral summer) and between August and September (austral winter). In addition, in each sampling site and season we collected potential food resources (fruits and insects) consumed by our A. lituratus. Our findings indicate that A. lituratus had a predominantly omnivorous diet, with high consumption of insects during summer in forest fragments. The increasing consumption of insects in these fragments seems to have led to a wider isotopic niche, in relation to the continuous forest. Because A. lituratus is typically a seed disperser, changes in trophic habits in the forest fragments from frugivory to insectivory may diminish their role in forest regeneration. Abstract in Portuguese is available with online material.     

高寒草地植物群落关键种对不同放牧家畜组合放牧的响应

如何通过合理的利用方式提高高寒草地管理水平,实现其可持续利用一直是草地生态学领域的研究热点。为明确不同放牧家畜组合下高寒草地植物群落关键种的演替规律及其驱动因素,基于中等放牧强度设置了不同放牧家畜组合放牧样地（牦牛单牧、藏羊单牧、牦牛藏羊1 ： 2混牧、牦牛藏羊1 ： 4混牧、牦牛藏羊1 ： 6混牧）以及围封样地,并在连续放牧处理7年后系统分析了不同放牧家畜组合下植物群落特征与土壤理化性质变化,以期确定最优放牧组合。结果表明：（1）不同放牧家畜组合均会显著降低高寒草地植物盖度,但对其物种丰富度与多样性指数的影响并不显著。（2）牦牛藏羊1 ： 2混牧下植物群落特征与禁牧处理下植物群落特征较为相似,且牦牛藏羊1 ： 2混牧下的植物群落稳定性最强、组织水平最高。（3）牦牛单牧与1 ： 2混牧下关键种为矮生嵩草;藏羊单牧下关键种为天山针茅;1 ： 4混牧下关键种为星毛委陵菜;1 ： 6混牧与围封下关键种为赖草。（4）围封能够显著降低土壤容重,藏羊单牧则会显著增加土壤容重与土壤速效氮、磷含量。（5）土壤速效氮、容重与含水量是驱动不同放牧家畜组合下植物群落关键种演替的重要理化因子。综上所述,中等放牧强度下,牦牛藏羊1 ： 2混牧是青藏高原高寒草地较为理想的利用方式。此外,相较于单纯增加草地物种丰富度或多样性,建立植物群落物种之间的有效关联是提高青藏高原高寒草地管理水平的另一关键途径。     

Plant patch structure modifies parasitoid assemblage richness of a specialist herbivore

1. The spatial structure of plant patches has been shown to affect host–parasitoid interactions, but its influence on parasitoid diversity remains largely ignored. Here we tested the prediction that parasitoid species richness of the specialist leafminer Liriomyza commelinae increases in larger and less isolated patches of its host plant Commelina erecta. We also explored whether parasitoid abundance and body size affected the occurrence of parasitoid species in local assemblages. 2. A total of 893 naturally established C. erecta patches were sampled on 18 sites around Córdoba city (Argentina). Also, two experiments were performed by creating patches differing in the number of plants and the distance from a parasitoid source. For these tests, plants were infected with the miner in the laboratory prior to placement in the field. 3. Plant patch size, independently of host abundance, positively affected the number of parasitoid species in both survey observations and experimental data. However, plant patch isolation did not influence parasitoid species richness. 4. The probability of finding rare parasitoid species increased with patch size, whereas occupation of isolated patches was independent of dispersal abilities (body size) of parasitoid species. 5. Overall, the results highlight the importance of considering spatial aspects such as the size of plant patches in the study of parasitoid communities.     

Biogeographical comparison of the arthropod herbivore communities associated with Lepidium draba in its native, expanded and introduced ranges

Aim To examine the composition and structure of the arthropod community on the invasive weed Lepidium draba in its native, expanded and introduced ranges, in order to elucidate the lack of a biotic constraint that may facilitate invasion. Location Europe and western North America. Methods Identical sampling protocols were used to collect data from a total of 35 populations of L. draba in its native (Eastern European), expanded (Western European) and introduced (western US) ranges. A bootstrapping analysis was used to compare herbivore richness, diversity and evenness among the regions. Core species groups (monophages, oligophages and polyphages) on the plant were defined and their abundances and host utilization patterns described. Results Species richness was greatest in the native range, while species diversity and evenness were similar in the native and expanded range, but significantly greater than in the introduced range of L. draba. Specialist herbivore abundance was greater in the native and expanded compared with the introduced range. Oligophagous Brassicaceae‐feeders were equally abundant in all three ranges, and polyphagous herbivore abundance was significantly greater in the introduced range. Overall herbivore abundance was greater in the introduced range. Host utilization was more complete in the two European ranges due to monophagous herbivores that do not exist in the introduced range. Root feeders and gall formers were completely absent from the introduced range, which was dominated by generalist sap‐sucking herbivores. However, one indigenous stem‐mining weevil, Ceutorhynchus americanus, occurred on L. draba in the introduced range. Main conclusions This is, to our knowledge, the first study documenting greater herbivore abundance on an invasive weed in its introduced, compared with its native, range. However, greater abundance does not necessarily translate to greater impact. We argue that, despite the greater total herbivore abundance in the introduced range, differences in the herbivore community structure (specialist vs. generalist herbivory) may contribute to the invasion success of L. draba in the western USA.     

Asexual endophytes and associated alkaloids alter arthropod community structure and increase herbivore abundances on a native grass

Despite their minute biomass, microbial symbionts of plants potentially alter herbivory, diversity and community structure. Infection of grasses by asexual endophytic fungi often decreases herbivore loads and alters arthropod diversity. However, most studies to date have involved agronomic grasses and often consider only infection status (infected vs. uninfected), without explicitly measuring endophyte-produced alkaloids, which vary among endophyte isolates and may impact consumers. We combined field experiments and population surveys to investigate how endophyte infection and associated alkaloids influence abundances, species richness, evenness and guild structure of arthropod communities on a native grass, Achnatherum robustum (sleepygrass). Surprisingly, we found that endophyte-produced alkaloids were associated with increased herbivore abundances and species richness. Our results suggest that, unlike what has been found in agronomic grass systems, high alkaloid levels in native grasses may not protect host grasses from arthropod herbivores, and may instead more negatively affect natural enemies of herbivores.
Ecology Letters (2010) 13: 106–117     

Fine‐scale variables associated with the presence of native forbs in natural temperate grassland

Broad‐scale threats to floristic diversity in native temperate grasslands are well‐documented and include elevated soil nutrients, changes in disturbance regimes and exotic species. However, fine‐scale variables associated with the presence of native forbs, such as gap size and biomass cover, have received relatively little attention. We conducted a case–control study to determine the relative influence of physical structural dimensions and other fine‐scale variables associated with the presence of native forbs in a modified temperate grassland previously used for domestic grazing. We matched 145 case plots centred on 27 different species of native forbs with 290 control plots not centred on a native forb. For each percentage increase in ground litter cover, dead biomass cover, grass cover or exotic forb cover, or the area of bare ground within 30 cm, the relative odds that a native forb was present vs absent declined by a mean of 10–13%. Living and dead biomass reduces light availability, and the former can also reduce nutrient and water availability. Declines in the presence of native forbs associated with increasing total bare ground may suggest that gap sizes were too small or the soil surface condition too degraded. Our results add to a body of evidence suggesting that native forbs in temperate native grassland are likely to benefit from periodic removal of living and dead grass biomass and a reduction in the cover of exotic forbs.     

Sublethal effects of a rapidly spreading native alga on a key herbivore

1. Multiple anthropogenic stressors are causing a global decline in foundation species, including macrophytes, often resulting in the expansion of functionally different, more stressor‐tolerant macrophytes. Previously subdominant species may experience further positive demographic feedback if they are exposed to weaker plant–herbivore interactions, possibly via decreased palatability or being structurally different from the species they are replacing. However, the consequences of the spread of opportunistic macrophytes for the local distribution and life history of herbivores are unknown.
2. The green alga, Caulerpa filiformis, previously a subdominant macrophyte on low intertidal‐shallow subtidal rock shores, is becoming locally more abundant and has spread into warmer waters across the coast of New South Wales, Australia.
3. In this study, we measured (a) the distribution and abundance of a key consumer, the sea urchin Heliocidaris erythrogramma, across a seascape at sites where C. filiformis has become dominant, (b) performed behavioral field experiments to test the role of habitat selection in determining the local distribution of H. erythrogramma, and (c) consumer experiments to test differential palatability between previously dominant higher quality species like Ecklonia radiata and Sargassum sp. and C. filiformis and the physiological consequences of consuming it.
4. At all sites, urchin densities were positively correlated with distance away from C. filiformis beds, and they actively moved away from beds. Feeding experiments showed that, while urchins consumed C. filiformis, sometimes in equal amounts to higher quality algae, there were strong sublethal consequences associated with C. filiformis consumption, mainly on reproductive potential (gonad size). Specifically, the gonad size of urchins that fed on C. filiformis was equivalent to that in starved urchins. There was also a tendency for urchin mortality to be greater when fed C. filiformis.
5. Overall, strong negative effects on herbivore life‐history traits and potentially their survivorship may establish further positive feedback on C. filiformis abundance that contributes to its spread and may mediate shifts from top‐down to bottom‐up control at locations where C. filiformis has become dominant.

The spread of the previously subdominant Caulerpa filiformis along the coast of NSW, Australia has strong lethal and sublethal effects on a key herbivore population. We performed sea urchin surveys, behavioral experiments, and consumer experiments and found that C aulerpa has strong negative effects on their populations. These results may establish reinforcing feedbacks on the spreading algae and mediate shifts from top‐down to bottom‐up control.     

Functional response in habitat selection and the tradeoffs between foraging niche components in a large herbivore

How herbivore behaviour is influenced by changes in resource levels is central for understanding trophic interactions. We examined whether foraging tradeoffs change with food levels by comparing habitat selection and space use within and between two neighbouring, predator‐free Svalbard reindeer populations. The populations faced different food levels due to contrasting grazing history. Summer resource selection in radiocollared females was assessed by a multi‐dimensional niche approach based on habitat variables obtained from a satellite image (e.g. the normalised difference vegetation index, NDVI) and a digital terrain model. The population at the overgrazed Brøggerhalvøya faced overall lower plant cover, biomass and primary productivity (i.e. lower NDVI) than the population at Sarsøyra. At Brøggerhalvøya, most reindeer selected for productive habitat when choosing home range and patches within the home range. In contrast, habitat selection at Sarsøyra was more affected by abiotic conditions such as moisture, which may influence plant quality. Here, reindeer used patches with even less biomass than the average reindeer at the poorer Brøggerhalvøya. Such a difference in habitat preference with different habitat availability (a functional response in habitat selection) probably reflected increased selection for high‐quality forage at the expense of high forage quantity at Sarsøyra. Accordingly, a negative relationship between habitat productivity and home range size was only present across individuals within Brøggerhalvøya, where forage quantity was the important foraging niche component. Individuals having poor (and large) home ranges apparently could not compensate for this by higher patch selectivity compared to individuals with richer home ranges. The results indicate changes in foraging tradeoffs at contrasting resource levels and that strong interactions occur between habitat selection, space use and the foraging niche structure in the absence of predation.     

When invasion may not be harmful: niche relations in a lizard assemblage

Some studies have suggested that non‐native species invasions may threaten local diversity by creating homogenized environments. However, many studies have been based on limited or anecdotal data, and/or have failed to consider the influence of habitat modification together with possible influences of non‐native species on native ones. Hemidactylus mabouia (Squamata, Gekkonidae) likely invaded natural environments in Brazil hundreds of years ago. Yet, little is known about whether it affects native lizard fauna. We tested whether H. mabouia negatively influences native lizard species richness and abundance on a regional scale and locally through niche overlap. We analyzed species abundance and richness of nine lizard assemblages, in five of which H. mabouia occurred. We evaluated niche overlap of species in a lizard assemblage with high H. mabouia abundance through null models. Niche axes included spatial use, temporal activity and diet. Although species abundance did not differ among sites with and without the invasive species, the presence of H. mabouia seems constrained to the richer assemblages sampled. We observed significantly higher niche overlap in spatial (?_obs = 0.63; ?_exp = 0.37; P_obs ≥ P_exp = 0.0002) and trophic axes (?_obs = 0.46; ?_exp = 0.17; P_obs ≥ P_exp < 0.001), but not in activity. When we considered all axes (three‐dimensional niche), there was no overlapping among the lizard species. Our findings did not support the hypothesis that this non‐native species negatively influences other sympatric lizard species.     

Lack of feral livestock interference with native guanaco during the dry season in a South American desert

Analyzing coexistence of exotic and native ungulates in arid areas is important from both a theoretical and a species conservation perspective. We assess the habitat use patterns and possible interference between guanaco (Lama guanicoe) and feral livestock (donkey and cattle) in arid environments of South America. To determine habitat use and niche overlap between exotic and native ungulate species, ten sites with different habitats and six natural waterholes were selected. Plots (20 at each site, ten around each waterhole) were randomly set up and characterized by environmental variables and relative use by cattle, donkey and guanaco through faecal pellet counts. Aggregation, niche breadth and niche overlap of the three herbivores were analyzed at habitat level (mesoscale). A direct redundancy analysis was used to examine the relationships between abundance of herbivore faeces and environmental variables at microscale. Mesoscale analyses showed (i) an extensive use of the area by all three species, with guanaco having the highest niche breadth followed by donkey and cattle and (ii) a large, broad guanaco–donkey and donkey–cattle habitat overlap. However, results at a finer scale showed high spatial aggregation of feral livestock species and an independent use of territory by guanacos. This study is the first to provide information about habitat partitioning between guanacos and feral livestock in the hyper-arid Monte Desert biome and points to an apparent lack of negative effects on the native ungulate.     

Ecological correlates of the non-indigenous parasitoid assemblage associated with a Hawaiian endemic moth

Understanding what ecological factors might predispose indigenous habitats to invasion by invasive species is an important aspect of conservation and invasive species management, particularly when biological control is considered for suppression of the invasive species. This study seeks to identify ecological factors that might play a role in determining the structure of the parasitoid assemblage associated with caterpillars of the endemic Hawaiian moth Udea stellata (Crambidae). Parasitoids were reared from field-collected U. stellata larvae at 18 locations. Fourteen environmental variables were measured at each site. Two multivariate analyses, principal component analysis (PCA) and partial redundancy analysis (RDA), were used to analyze the parasitoid assemblage across a range of habitats varying in environmental characteristics. The PCA analysis showed that the occurrence of some species were highly correlated, and associated with less disturbed sites, whereas other species were associated with sites of medium and high levels of disturbance. The RDA analysis showed that only three of the measured environmental variables (U. stellata density, elevation, and level of habitat disturbance) significantly explained variability in the parasitoid assemblage among sites. There was greater parasitoid species richness associated with U. stellata larvae at higher elevation sites with a lower degree of habitat disturbance by exotic vegetation. The purposely introduced parasitoid species were associated with the non-target moth at sites located at higher elevations with low levels of disturbance. Multivariate analysis has the potential to provide valuable insights into the identification of important environmental factors that mediate parasitoid assemblage structure and level of parasitism on a particular target or non-target species, and therefore facilitate identification of suitable target habitats or susceptible non-target habitats.     

Parasitoid assemblage associated with a North American pine weevil in South Africa

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Use of an exotic host plant reduces viral burden in a native insect herbivore

Incorporation of exotic plants into the diets of native herbivores is a common phenomenon, influencing interactions with natural enemies and providing insight into the tritrophic costs and benefits of dietary expansion. We evaluated how use of an exotic plant, Plantago lanceolata, impacted immune performance, development and susceptibility to pathogen infection in the neotropical herbivore Anartia jatrophae (Lepidoptera: Nymphalidae). Caterpillars were reared on P. lanceolata or a native plant, Bacopa monnieri, and experimentally infected with a pathogenic virus, Junonia coenia densovirus. We found that virus-challenged herbivores exhibited higher survival rates and lower viral burdens when reared on P. lanceolata compared to B. monnieri, though immune performance and development time were largely similar on the two plants. These findings reveal that use of an exotic plant can impact the vulnerability of a native herbivore to pathogen infection, suggesting diet-mediated protection against disease as a potential mechanism facilitating the incorporation of novel resources.     

Transition of a Sambucus nigra L. dominated woody vegetation into grassland by a multi-species herbivore assemblage

We describe and analyse how large herbivores strongly diminished a woody vegetation, dominated by the unpalatable shrub Sambucus nigra L. and changed it into grassland. Density of woody species and cover of vegetation were measured in 1996, 2002 and 2012 in the grazed Oostvaardersplassen. In 2002 and 2012 we also measured density and cover in an ungrazed control site. In 2002 we measured intensity of browsing and bark loss of Sambucus shrubs in the grazed and control sites. In the grazed site the density of Sambucus and Salix spp. declined significantly between 1996 and 2012, and large areas changed into grassland. In the control site the density of Sambucus increased significantly during this period, the density of Salix spp. did not change, and the vegetation consisted of a mixture of woody species and a field layer dominated by tall herbs. In 2002 and 2012 the percentages of dead Sambucus shrubs were significantly higher in the grazed site than in the control site. In 2002 the percentages of twigs browsed and ring barked stems of Sambucus shrubs were significantly higher in the grazed site than in the control site. Our results show that debarking caused mature Sambucus shrubs to die, but that heavy browsing may have helped this process. Our results also point to a significant neighbour effect on the break down of Sambucus, suggesting that Aggregational Resistance and Associational Palatability were both active. Essential conditions for the break down of this woody vegetation were the presence of large herbivores, the low ratio between the areas of summer and winter feeding habitats and the competition amongst herbivores. Browsing may have been responsible for seedling death, as seedlings were found only in the control site and not on the old and newly established grasslands in the grazed site.     

Trophic niche changes associated with the eradication of invasive mammals in an insular lizard: an assessment using isotopes

Invasive species are a major threat to island biodiversity, and their eradications have substantially contributed to the conservation of island endemics. However, the consequences of eradications on the trophic ecology of native taxa are largely unexplored. Here, we used the eradication of invasive black rats Rattus rattus and European rabbits Oryctolagus cuniculus from the Berlenga Island, in the western coast of Portugal, as a whole-ecosystem experiment to investigate the effects of the eradication of invasive mammals on the trophic niche and body dimensions of the island-restricted Berlenga wall lizard Podarcis carbonelli berlengensis over a 2-year period. Our results suggest an expansion of the isotopic niche and an intensification of the sexual dimorphism of the lizard following mammal eradication. Additionally, we found considerable variability in isotopic niche across the island and detected evidence of sex-specific and season-modulated nutritional requirements of this threatened reptile. Our findings support that the eradication of 2 of the planet’s most problematic invasive vertebrates led to changes in the lizard trophic niche and sexual dimorphism in just 2 years. This suggests that the ecological pressures—for example, prey availability and habitat structure—to which lizards are exposed have substantially changed post-eradication. Our study emphasizes the scientific value of island eradications as experiments to address a wide range of ecological questions and adds to the increasing body of evidence supporting substantial conservation gains associated with these restoration interventions.