On the evidence for species coexistence: a critique of the coexistence program

A major challenge in ecology is to understand how the millions of species on Earth are organized into biological communities. Mechanisms promoting coexistence are one such class of organizing processes, which allow multiple species to persist in the same trophic level of a given web of species interactions. If some mechanism promotes the coexistence of two or more species, each species must be able to increase when it is rare and the others are at their typical abundances; this invasibility criterion is fundamental evidence for species coexistence regardless of the mechanism. In an attempt to evaluate the level of empirical support for coexistence mechanisms in nature, we surveyed the literature for empirical studies of coexistence at a local scale (i.e., species found living together in one place) to determine whether these studies satisfied the invasibility criterion. In our survey, only seven of 323 studies that drew conclusions about species coexistence evaluated invasibility in some way in either observational or experimental studies. In addition, only three other studies evaluated necessary but not sufficient conditions for invasibility (i.e., negative density dependence and a trade-off in performance that influences population regulation). These results indicate that, while species coexistence is a prevalent assumption for why species are able to live together in one place, critical empirical tests of this fundamental assumption of community structure are rarely performed. These tests are central to developing a more robust understanding of the relative contributions of both deterministic and stochastic processes structuring biological communities.     

Heavy metals in fish species from lotic freshwater ecosystem at Afikpo, Nigeria

A study was conducted between March 2006 and February 2007 to assess the concentration of Fe, Zn, Cu, Mn, Pb, and Cr in the gills and muscles of six fish species (Chrysichthys nigrodigitatus, Clarias anguillaris, Tillapia zillii, Mormyrus rume rume, Mormyrus macrophthalmus and Mormyrus tapirus) from lotic freshwater ecosystem at Afikpo South- East Nigeria using Atomic Absorption Spectrophotometer. In all the fish species studied the concentration of metals in the gills was significantly higher than that of the muscles (p<0.05). While the highest concentration (mg l(-1)) of Fe (702.20 +/- 0.04), Zn (34.40 +/- 0.02), Cu (2.10 +/- 0.01), Mn (4.91 +/- 0.08) Pb (0.50 +/- 0.02) and Cr (1.12 +/- 0.07) were recorded in the gills of C. nigrodigitatus, that in the muscles were recorded in T. zillii (443.20 +/- 0.08, 23.30 +/- 0.06, 1.33 +/- 0.06, 3.09 +/- 0.02, 0.31 +/- 0.01 and 0.66 +/- 0.04 for Fe, Zn, Cu, Mn, Pb and Cr respectively). The lowest concentration of all the heavy metals in the gills was recorded in M. tapirus (309.00 +/- 0.07, 16.45 +/- 0.03, 0.92 +/- 0.04, 2.15 +/- 0.04, 0.21 +/- 0.01 and 0.50 +/- 0.06 mg l(-1)for Fe, Zn, Cu Mn, Pb and Cr respectively) while the lowest in the muscles was recorded in C. anguillaris [Fe (186.00 +/- 0.07), Zn (14.20 +/- 0.08), Cu (0.56 +/- 0.03), Mn (1.30 +/- 0.02), Pb (0.10 +/- 0.01) and Cr (0.28 +/- 0.04)]. The order of heavy metals concentrations in both the gills and muscles was Fe>Zn>Mn>Cu>Cr>Pb. While the concentration of Zn, Cu and Pb both in the muscles and gills of all the six fish species studied were within the WHO and FEPA prescribed limits, that of Fe(except in the muscles), Mn and Cr were above the prescribed limits thus indicating contamination of the fish species by these metals. Fe has the highest bio-concentration factor (BCF) in both tissues while the least was Cu. Periodic monitoring of these metals in both fishes and river to ensure safety is advocated.     

Methylmercury, total mercury and total selenium in four common freshwater fish species from Ya-Er Lake, China

Snakehead fish (Ophiocephalus argus cantor), silver carp (Hypophthalmichthys molitrtix), crucian carp (Carassius carassius), and common carp (Cyprinus carpio) are four common freshwater fish species in China. In this study, the level of methylmercury (MeHg), total mercury (T-Hg), and total selenium (T-Se) in muscle samples of these four fish species from Ya-Er Lake, China, were analyzed using atomic fluorescence spectrometry coupled with high-performance liquid chromatography, and inductively coupled plasma mass spectrometry. The concentrations of MeHg in all the fish species were significantly correlated with those of T-Hg. Higher T-Hg and MeHg concentrations had accumulated in the snakehead fish, which is a strongly predatory fish, than in the other three species. The concentration ratios of MeHg and T-Hg in the muscles of these four fish species were almost equal. Conversely, there was negative correlation between the concentrations of T-Hg and T-Se, which implies that there is a competition between these two elements with respect to bioaccumulation. It is noteworthy that of all the muscle samples tested, the level of T-Hg exceeded the maximum allowable limit in fish [0.4 mg kg⁻¹ (w/w) recommended by the World Health Organization] in 38.46% of those of the silver carp, 18.18% of those of the crucian carp, and 100% of those of snakehead fish. These results show that the consumption of contaminated fish is a potential threat to human health and that necessary preventive measures to safeguard public health should be emphasized.     

Effect of cortisol on female freshwater fish Notopterus notopterus

In the present study, effect of cortisol on the ovary of the freshwater fish Notopterus notopterus was studied during four phases of the reproductive cycle. The hormone was injected 60microg/fish for ten days. Cortisol in fish is known to increase the metabolic activity. After the hormone treatment the ovarian somatic index and the hepatosomatic index increases during non breeding phase. The young oocytes transferred into vitellogenic oocytes in all the phases after cortisol treatment. The cortisol induced increased ovarian activity may be due to increase in the metabolic activity through the involvement of hepatic cells specially during non-breeding period providing support for energy production for reproductive activity in the freshwater fish N. notopterus.     

Microbial food partitioning by three species of benthic copepods

Using radioactively labeled bacteria and photoautotrophs in undisturbed sediment cores, we show that three cooccurring species of benthic copepods feed on different microbial food sources in their natural environment. Specifically, Thompsonula hyaenae feeds on photoautotrophs, Halicyclops coulli feeds on bacteria, and Zausodes arenicolus feeds on both photoautotrophs and bacteria. Species of benthic copepods feed differently from one another in the field, and meiofaunal species' distributions could be influenced by distributions of their preferred microbial food.     

Plasticity in circadian activity patterns of mesocarnivores in Southwestern Europe: implications for species coexistence

Limiting similarity theory predicts that competing species must segregate along one or more dimensions of their ecological niche in order to coexist. In predator communities, interspecific interactions are influenced by a diversity of factors; therefore, the behavioural patterns of composing species will differ due to locally adapted interactions. We deployed 32–41 camera-traps in five study areas across the Iberian Peninsula to investigate the temporal relations between mesocarnivores in SW Europe. The selection for a period of the diel cycle and plasticity in activity patterns was evaluated using the Jacobs Selection Index (JSI) and the coefficient of activity overlap (?₁). Furthermore, we investigated whether temporal shifts can facilitate coexistence by reducing activity overlap. Seven species of mesocarnivores were detected and were assigned into one of three behaviourally distinct groups: diurnal (JSI_day?≥?0.8), strictly nocturnal (JSI_night?≥?0.8) or facultative nocturnal species (0.4?≥?JSI_night?>?0.8). Most species exhibited substantial flexibility, which allowed them to locally adapt their foraging strategies (intraspecific ?₁?=?0.70–0.77). Mean Δ₁ from all interspecific pairwise comparisons was negatively correlated with the number of carnivore species with ≥10 detections (r ?0.76, p?=?0.02). Our results suggest that temporal segregation is likely to play an important role in facilitating mesocarnivore coexistence, especially with increasing community complexity, where most species’ activity peaks were asynchronous. These results contribute to understanding the dynamics and behavioural strategies of coexisting mesocarnivores, crucial for forecasting the possible outcomes of conservation or management actions.     

Optimal harvesting of ecologically interdependent fish species

The optimal exploitation of a two-species predator-prey system is considered, using Lotka-Volterra-type equations. Due to the density-dependence of ecological efficiency, both species should be harvested simultaneously over a range of relative prices. Beyond the limits of this price range, either the prey species should be utilized indirectly by harvesting the predator, or the predator should be eliminated to maximize the prey yield. Neglecting harvesting costs, the simultaneous harvest of prey and predators requires that a unit of prey biomass increase in value by being “processed” by predators. Certain results from single-species fishery models are shown not to apply to multispecies models. These are as follows: (i) Optimal regulation of a free access fishery may call for subsidizing instead of taxing the harvest of predator species. (ii) Increasing the discount rate may, at “moderate” levels, imply that the optimal standing stock of biomass increases instead of decreasing. (iii) A rising price or a falling cost per unit fishing effort of a species may raise and not lower the optimal standing stock of that species.     

Effects of sheltering fish on growth of their host corals

Stony corals are the foundation species of tropical reefs, and their structures can harbor a diverse range of mutualist taxa that can confer important benefits, including provision of nutrients. Prominent among the associates of branching coral in the genus Pocillopora are groups of zooplanktivorous damselfishes that take refuge in the coral to avoid their predators. In field and laboratory experiments, we explored the effects of colonies of resident damselfishes on growth of their host corals. Laboratory studies revealed a positive relationship between biomass of fish and output of ammonium. In the field, levels of ammonium were significantly elevated in the water surrounding the branches of Pocillopora occupied by colonies of damselfish, particularly in time periods following active feeding by the fish. Experimental manipulation of the presence of fish on host corals during a month-long field experiment revealed that corals hosting fish grew significantly more than those that lacked fish, and coral growth was positively correlated with the biomass of resident fish. The Pocillopora colonies in the field experiment varied in the degree of openness of their branching structure, and dye studies indicated that this affected their ability to retain waterborne nutrients. Together with biomass of resident fish, colony openness explained 76% of the variation in coral growth rate during the experiment. Corals can exhibit considerable morphological variability, and mutualistic fish respond to colony architecture during habitat selection, with some species preferring more open-branched forms. This makes it likely that corals may face tradeoffs in attracting resident fish and in retaining the nutrients they provide.     

Predicting barrier passage and habitat suitability for migratory fish species

Fish migrate to spawn, feed, seek refuge from predators, and escape harmful environmental conditions. The success of upstream migration is limited by the presence of barriers that can impede the passage of fish. We used a spatially explicit modeling strategy to examine the effects of barriers on passage for 21 native and non-native migratory fish species and the amount of suitable habitat blocked for each species. Spatially derived physical parameter estimates and literature based fish capabilities and tolerances were used to predict fish passage success and habitat suitability. Both the fish passage and the habitat suitability models accurately predicted fish presence above barriers for most common, non-stocked species. The fish passage model predicted that barriers greater than or equal to 6 m block all migratory species. Chinook salmon (Oncorhynchus tshawytscha) was expected to be blocked the least. The habitat suitability model predicted that low gradient streams with intact habitat quality were likely to support the highest number of fish species. The fish passage and habitat suitability models were intended to be used by environmental managers as strategy development tools to prioritize candidate dams for field assessment and make decisions regarding the management of migratory fish populations.     

The methane mussel: roles of symbiont and host in the metabolic utilization of methane

Methane mussels (Bathymodiolus sp., undescribed; personal communication by R. Turner to CRF) were collected in September 1989 and April 1990 from offshore Louisiana in the Gulf of Mexico. These mussels contain endosymbiotic methane-oxidizing bacteria and are capable of utilizing environmental methane as a source of energy and carbon. Oxygen consumption, methane consumption, and carbon dioxide production were measured in mussels with intact symbionts, functionally aposymbiotic mussels, and separated symbiont preparations under controlled oxygen and methane conditions, in order to study the roles of the symbionts and the hosts in methane utilization. The association was found to be very efficient in fixing methane carbon (only 30% of CH₄ consumed is released as CO₂), and to be capable of maximal rates of net carbon uptake of nearly 5 mol g^-1 h^-1. Rates of oxygen and methane consumption were dependent upon oxygen and methane concentrations. Maximal consumption rates were measured at 250 to 300 M O₂ and 200 to 300 M CH₄, under which conditions, oxygen consumption by the gill tissues (containing symbionts) had increased more than 50-fold over rates measured in the absence of methane. A model is proposed for the functioning of the intact association in situ, which shows the symbiosis to be capable of achieving growth rates (net carbon assimilation) in the range of 0.003 to 0.50% per day depending upon oxygen and methane concentrations. Under the conditions measured in the seep environment (200 M O₂, 60 M CH₄), a mussel consuming methane at rates found to be typical (4 to 5 mol g^-1 h^-1) should have a net carbon assimilation rate of about 0.1% per day. We suggest that the effectiveness of this symbiosis arises through integration of the morphological and physiological characteristics inherent to each of the symbiotic partners, rather than from extensive specialization exhibited by other deep-sea chemotrophic associations.     

Impact of copper on haematological profile of freshwater fish, Channa punctatus

The present study was to evaluate heavy metal toxicity stress symptoms in fish blood during long-term exposure of sublethal concentration of copper sulphate (CuSO4). The effects of copper on various heamatological parameters were evaluated exposing Indian fresh water fish, Channa punctatus to a sub-lethal concentration of copper sulphate (0.36 mg l(-1)) for different periods. Exposure of fish to copper showed a significant decrease in the haemoglobin (Hb) content from 10.73 to 6.60%, red blood cells (RBC) from 2.86 to 1.84 x 10(6)/mm3 and packed cell volume (PCV) from 31.00 to 23.33% at the end of 45th day as compared to control. Whereas the white blood cells (WBC) increased from 60.00 to 92.48 x10(3)/mm3, clotting time (CT) from 27.66 to 43.00 second, erythrocyte sedimentation rate from 5.0 to 13.66 mm/hr and mean corpuscular volume from 108.11 to 126.85 cmu, significantly, with increase in exposure periods. Although mean corpuscular haemoglobin concentration (MCHC) and mean corpuscular haemoglobin (MCH) values showed significant increase during 15 and 30 days exposures, both the values were found significantly increased at the end of the experimental period. The differential leucocytes count (DLC) showed a significant increase in the populations of lymphocytes and eosinophils, whereas neutrophils, monocytes and basophiles were found decreased after exposure to copper.     

A neighboring plant species creates associational refuge for consumer and host

Examples of plant-animal and plant-plant associational defenses are common across a variety of systems, yet the potential for them to occur in concert has not been explored. In salt marshes in the Gulf of Mexico, the marsh periwinkle (Littoraria irrorata) is an abundant and conspicuous member of the community, climbing up the stems of marsh plants to remain out of the water at high tide. Though Littoraria are thought to primarily utilize stems of marsh cordgrass Spartina alterniflora as a source of food and refuge, Littoraria were more abundant in mixed assemblages of Spartina and Juncus roemerianus than in Spartina-only areas at the same tidal height. Mesocosm experiments confirmed that Juncus provided a refuge for Littoraria, with predation by Callinectes sapidus (but not Melongena corona) reduced when Juncus was present. However, Littoraria's utilization of Juncus as well as the effectiveness of Juncus as a refuge depended strongly on plant height: when Juncus was experimentally clipped to a shorter height than Spartina, snail abundance on Spartina and snail predation by crabs increased. Interestingly, this plant animal refuge led to a corresponding refuge for Spartina from Littoraria: Spartina plants lost less biomass to snail grazing when growing with Juncus in mesocosm and field experiments, and Spartina plants in natural assemblages were taller when growing with Juncus than when growing alone, even in the presence of abundant snails. This example highlights the potential importance of plant plant and plant-animal associational refuges in competitive plant assemblages.     

Study of carbohydrate metabolism in selected tissues of freshwater mussel, Lamellidens marginalis under copper sulphate toxicity

Sublethal toxicity of copper sulphate on carbohydrate metabolism was studied in selected tissues of freshwater mussel, Lamellidens marginalis. Levels of glycogen and pyruvic acid decreased while lactic acid showed an increase. Activities of LDH, SDH and MDH decreased while G-6-PDH activity increased. There appears to be a shift in the carbohydrate metabolism from aerobic to anaerobic type due to toxicity of copper.     

Assessment of toxicity of two nitroaromatic compounds in the freshwater fish Cyprinus carpio

This study was conducted to evaluate the toxicological response of p-nitrotoluene and p-nitroaniline to the key fish species, Cyprinus carpio. A freshwater fish bioassay based on the 96 h LC ₅₀ was used to estimate the single and joint toxicity of the two chemicals. The toxicity of p-nitrotoluene was greater than that of p-nitroaniline based on 96 h LC ₅₀ values of 40.74 mg·L^?1 and 48.99 mg·L^?1, respectively. Both compounds had moderate toxicity toward Cyprinus carpio, and this toxicity increased with the exposure duration and concentration. Binary mixtures of the compounds were more toxic than the individual compounds at 96 h, and they acted upon partial addition. When the exposure time was longer, the toxicity increased for mixtures of compounds with the same concentration or toxicity. The results of this study suggest that exposure to a combination of these chemicals would result in a higher environmental risk in aquatic systems than exposure to either compound alone. Further research is needed to investigate the combined effects and sublethal toxicity of p-nitrotoluene and p-nitroaniline, since they are both still used in China.     

Variation partitioning of species data matrices: estimation and comparison of fractions

Establishing relationships between species distributions and environmental characteristics is a major goal in the search for forces driving species distributions. Canonical ordinations such as redundancy analysis and canonical correspondence analysis are invaluable tools for modeling communities through environmental predictors. They provide the means for conducting direct explanatory analysis in which the association among species can be studied according to their common and unique relationships with the environmental variables and other sets of predictors of interest, such as spatial variables. Variation partitioning can then be used to test and determine the likelihood of these sets of predictors in explaining patterns in community structure. Although variation partitioning in canonical analysis is routinely used in ecological analysis, no effort has been reported in the literature to consider appropriate estimators so that comparisons between fractions or, eventually, between different canonical models are meaningful. In this paper, we show that variation partitioning as currently applied in canonical analysis is biased. We present appropriate unbiased estimators. In addition, we outline a statistical test to compare fractions in canonical analysis. The question addressed by the test is whether two fractions of variation are significantly different from each other. Such assessment provides an important step toward attaining an understanding of the factors patterning community structure. The test is shown to have correct Type I. error rates and good power for both redundancy analysis and canonical correspondence analysis.     

Critical thermal maxima of two species of estuarine fish

Critical thermal maxima (CTM) were determined for winter flounder (Pseudopleuronectes americanus, Walbaum) and scup (Stenotomus chrysops, L.), acclimated to ambient water temperature or constant increments above ambient. The acclimation regimes simulated a natural environment by tracking the temperature fluctuations in Point Judith Pond, a small Rhode Island estuary. An elevation in CTM from 30.2° to 35.6°C was observed for scup as the mean acclimation temperature increased from 14.8° to 22.2°C. CTM for flounder increased from 26.4° to 32.4°C over an acclimation range of 4.2° to 23.0°C. The use of CTM studies for predicting the effects of heated effluents is discussed.     

A methodological framework for integrating computational fluid dynamics and ecological models applied to juvenile freshwater mussel dispersal in the Upper Mississippi River

Interdisciplinary research in hydraulics and ecology for river management and restoration must integrate processes that occur over a wide range of spatial and temporal scales, which presents a challenge to ecohydraulics modelers. Computational fluid dynamics (CFD) models are being more widely used to determine flow fields for ecohydraulics applications. In the Upper Mississippi River (UMR), the mussel dynamics model was developed as a tool for management and conservation of freshwater mussels (Unionidae), which are benthic organisms, imperiled in North America, that are inextricably linked with the hydraulics of river flow. We updated the juvenile dispersal component of the mussel dynamics model by using stochastic Lagrangian particle tracking in a three dimensional flow field output from CFD models of reaches in the UMR. We developed a methodological framework to integrate hydrodynamic data with the mussel dynamics model, and we demonstrate the use of the juvenile dispersal model employed within the methodological framework in two reaches of the UMR. The method was used to test the hypothesis that impoundment affects the relationship of some hydraulic parameters with juvenile settling distribution. Simulation results were consistent with this hypothesis, and the relationships of bed shear stress and Froude number with juvenile settling were altered by impoundment most likely through effects on local hydraulics. The methodological framework is robust, integrates Eulerian and Lagrangian reference frameworks, and incorporates processes over a wide range of temporal and spatial scales, from watershed scale hydrologic processes (decades), to reach scale (km) processes that occur over hours or days, and turbulent processes on spatial scales of meter to millimeter and times scales of seconds. The methods are presently being used to assess the impacts of pre- and early post-settlement processes on mussel distributions, including the effects of bed shear stress, and the sensitivity of the location of the host fish when juveniles excyst, on juvenile settling distribution.     

Levels of selected heavy metals in commercial fish from five freshwater lakes,Pakistan

The levels of As, Cd, Cr, Cu, Fe, Mn, Ni, Pb, Zn and Hg are estimated by atomic absorption in the muscle of six commercial species of fish belonging to five freshwater lakes of Pakistan. The species included Mastacembelus armatus, Tor putitura, Mystus seenghala, Wallago attu, Catla catla and Labeo rohita. The fish were procured in comparable weight ranges so that a viable comparison of trace metal content could be effected. As, Fe, Pb, Zn and Hg showed elevated levels of 0.006–6.967, 0.933–6.133, 0.060–4.108, 0.978–5.363 and 0.030–3.211 μg/g, wet weight. The relevant statistical parameters, such as standard deviation, standard error, skewness and its t‐value are also reported for establishing the randomness of the distribution in relation to the corresponding data of freshwater lakes of the world and examined in view of species‐specificity and origin.     

Homogenization dynamics and introduction routes of invasive freshwater fish in the Iberian Peninsula.

Nonnative invasive species are one of the main global threats to biodiversity. The understanding of the traits characterizing successful invaders and invasion-prone ecosystems is increasing, but our predictive ability is still limited. Quantitative information on biotic homogenization and particularly its temporal dynamics is even scarcer. We used freshwater fish distribution data in the Iberian Peninsula in four periods (before human intervention, 1991, 1995, and 2001) to assess the temporal dynamics of biotic homogenization among river basins. The percentage of introduced species among fish faunas has increased in recent times (from 41.8% in 1991 to 52.5% in 2001), leading to a clear increase in the similarity of community composition among basins. The mean Jaccard's index increase (a measure of biotic homogenization) from the pristine situation to the present (17.1%) was similar to that for Californian fish but higher than for other studies. However, biotic homogenization was found to be a temporally dynamic process, with finer temporal grain analyses detecting transient stages of biotic differentiation. Introduced species assemblages were spatially structured along a latitudinal gradient in the Iberian Peninsula, with species related to sport fishing being characteristic of northern basins. Although the comparison of fish distributions in the Iberian Peninsula and France showed significant and generalized biotic homogenization, nonnative assemblages of northeastern Iberian basins were more similar to those of France than to those of the rest of the Iberian Peninsula, indicating a main introduction route. Species introduced to the Iberian Peninsula tended to be mainly piscivores or widely introduced species that previously had been introduced to France. Our results indicate that the simultaneous analysis of the spatial distribution of introduced assemblages (excluding native species that reflect other biogeographical patterns) and their specific traits can be an effective tool to detect introduction and invasion routes and to predict future invaders from donor regions.     

Worker size-related task partitioning in the foraging strategy of a seed-harvesting ant species

Messor bouvieri is a seed-harvesting ant species in which workers forage in trails from the nest to a search area. A previous observation of seed transfer events between workers returning to the nest suggested potential task partitioning. In this study, we describe seed transportation and analyze the role of task partitioning in the foraging strategy of this species in terms of seed intake efficiency in relation to costs and benefits based on transport speed and task reliability. We assess the harvesting efficiency of task partitioning by comparing cooperative seed transport (CST) and individual seed transport (IST) events. Our results show task partitioning in the form of a sequence of transfer events among workers going from the search area to the nest. Importantly, and despite the weak worker polymorphism of this species, this sequence involved workers of different sizes, with seeds usually being passed along from smaller to larger workers. In addition, we show that small workers are better at finding seeds (spend less time finding a seed), and large workers are better at transporting them (were faster when walking back to the nest and lost fewer seeds). However, we failed to demonstrate that workers of different sizes are specialized in performing the task in which they excel. Overall, sequential CST in M. bouvieri results in a greater seed intake because seed search time decreases and task reliability increases, compared to IST. The determinants and adaptive benefits of CST are discussed.